Curriculum Vitae

Joan's Photo

Postdoctoral Students (Primary Advisor)
Dr. Joan Herbers 1978-1979                 Dr. David King 1979-1981
Dr. Yoh Iwasa 1981-1983   Dr. Elizabeth Denley 1982-1984
Dr. Steven Gaines 1982-1987   Dr. Stephen Brown 1984-1986
Dr. Sheryl Horowitz 1985-1986   Dr. Hugh Possingham 1987-1988
Dr. Terrence Farrell 1987-1989   Dr. Lloyd Goldwasser 1988-1990
Dr. Douglas Stoner 1989-1991   Dr. Timothy Pennington 1989-1991
Dr. Yehoshua Shkedy 1992-1995   Dr. Jef Huisman 1997-1998
Dr. Erol Akçay 2008-2009  

Ph.D. Students (Primary Advisor)
Dr. Ronald Yoshiyama 1972-1977                 Dr. David Heckel 1975-1980
Dr. Stephen Pacala 1978-1982   Dr. John D. Rummel 1979-1984
Dr. Kenneth Naganuma 1982-1989   Dr. Sally Blower 1982-1987
Dr. John McLaughlin 1984-1990   Dr. Kristina Miller 1987-1992
Dr. Susan Alexander 1988-1993   Dr. Roman Dial 1988-1992
Dr. Brian Grantham 1991-1997   Dr. Sharoni Shafir 1991-1995
Dr. Eric Bjorkstedt 1992-1997   Dr. Sean Connolly 1994-1998
Dr. Henrique Pereira 1997-2002   Dr. Paul Armsworth 1999-2003
Dr. Lauren Buckley 2001-2005   Dr. Erol Akçay 2004-2008
Dr. Priya Iyer 2005-2009   Dr. Henri Folse 2006-2011


  1. 2015, Roughgarden, J., Ram-2050: A Ramayana Epic for the Future. Kauai Institute. Kapaa HI. (Fiction)

  2. 2009, Roughgarden, J., The Genial Gene: Deconstructing Darwinian Selfishness. University of California Press. Berkeley CA. French translation, 2012, published by Le Seuil, Paris.

  3. 2006, Roughgarden, J., Evolution and Christian Faith: Reflections of an Evolutionary Biologist. Island Press. Washington DC.

  4. 2004, Roughgarden, J., Evolution's Rainbow: Diversity, Gender, and Sexuality in Nature and People. University of California Press. Berkeley CA. Awarded 2005 Stonewall Prize for nonfiction from American Library Association. Portuguese translation, 2005, published by editoria Planta, Brazil. Korean translation, 2010, published by Puriwa Ipari Publishing Co., Seoul, Korea. Tenth Anniversary Edition, 2014, University of California Press, Berkeley CA.

  5. 1998, Roughgarden, J., Primer of Ecological Theory. Prentice Hall, Upper Saddle River, New Jersey.

  6. 1995, Roughgarden, J., Anolis Lizards of the Caribbean: Ecology, Evolution, and Plate Tectonics. Oxford University Press, Oxford UK.

  7. 1989, Roughgarden, J., R. May, and S. Levin, Eds., Perspectives in Ecological Theory. Princeton University Press, Princeton, New Jersey.

  8. 1987, Ehrlich, P., and J. Roughgarden, The Science of Ecology. Macmillan Publishing Company, New York, New York.

  9. 1979, Roughgarden, J., Theory of Population Genetics and Evolutionary Ecology: An Introduction. Macmillan Publishing Company, New York. (Reprinted 1987 by Macmillan, and in 1996 by Prentice Hall).

Selected Articles

  1. 2020, Roughgarden, J., Holobiont evolution: Population genetic theory for the hologenome. BioRxiv doi:

  2. 2020, Roughgarden, J., Holobiont evolution: Mathematical model with vertical vs. horizontal microbiome transmission. Philos Theor Pract Biol 12:2  doi:10.3998/ptpbio.16039257.0012.002

  3. 2018, Roughgarden, J., Scott F. Gilbert, Eugene Rosenberg, Ilana Zilber-Rosenberg and Elisabeth A. Lloyd, Holobionts as units of selection and a model of their population dynamics and evolution. Biological Theory 13(1):44-65 (

  4. 2017, Roughgarden, J. Model of holobiont population dynamics and evolution: A preliminary sketch. Pp 325-350 in: Snait Gissis, Ehud Lamm, and Ayelet Shavit (eds.) Landscapes of Collectivity in the Life Sciences , Vienna Series in Theoretical Biology, The MIT Press.

  5. 2017, Roughgarden, J. Incentivizing biological cooperation: Direct benefits and the economic theory of the firm. Pp. 227-244 in: Snait Gissis, Ehud Lamm, and Ayelet Shavit (eds.) Landscapes of Collectivity in the Life Sciences , Vienna Series in Theoretical Biology, The MIT Press.

  6. 2017, Roughgarden, J. A theoretician’s perspective on collectivity. Pp. 49-52 in: Snait Gissis, Ehud Lamm, and Ayelet Shavit (eds.) Landscapes of Collectivity in the Life Sciences , Vienna Series in Theoretical Biology, The MIT Press.

  7. 2017, Roughgarden, J. The scientist dreamer: Epilogue. In: Harman, Oren and Michael Dietrich, (eds). Dreamers, Visionaries, and Revolutionaries in the Life Sciences. University of Chicago Press. (in press)

  8. 2017, Roughgarden, J. The gender binary in nature, across human cultures, and in the Bible. Pp. 413-440 in: Gerhard Schreiber (ed.), Transsexualität in Theologie und Neurowissenschaften. Ergebnisse, Kontroversen, Perspektiven (De Gruyter) [In: Gerhard Schreiber (ed.), Transsexualism in Theology and Neuroscience. Results, Controversies, Perspectives (De Gruyter)].

  9. 2017, Roughgarden, J. Homosexuality and evolution: A critical appraisal. Pp. 495-516 (Chapter 30) in: Tibayrenc, Michel and Francisco Ayala (eds). On Human Nature: Biology, Psychology, Ethics, Politics, and Religion, Academic Press, Elsevier, London.

  10. 2015, Roughgarden J, Adkins-Regan E, Akçay E, Crawford JC, Gadagkar R, Griffith SC, Hinde CA, Hoquet T, O’Connor C, Prokop ZM, Prum RO, Shafir S, Snow SS, Taylor D, Van Cleve J, Weisberg M. Sexual selection studies: a NESCent catalyst meeting. PeerJ PrePrints 3:e680v3

  11. 2015, Roughgarden, J. Sexual selection: Is anything left? In: Hoquet, T. (Ed) Current Perspectives on Sexual Selection: What's Left After Darwin? Springer Verlag. Pp. 85-102.

  12. 2014, Roughgarden, J. and Song, Z. Incentives in the family I: The family firm, an evolutionary/economic theory for parent-offspring Relations. Pp. 220-239 in: Narvaez, D., Valentino, K., Fuentes, A., McKenna, J., & Gray, P. (Eds.) Ancestral Landscapes in Human Evolution: Culture, Childrearing and Social Wellbeing. New York: Oxford University Press.

  13. 2013, Akçay, E., Roughgarden, J., Fearon, J., Ferejohn, A. and B. Weingast. Biological institutions: the political science of animal cooperation. Social Science Research Network

  14. 2012, Roughgarden, J. Individual based models in ecology: An evaluation, or how not to ruin a good thing. In: [2012] Philosophy of Science Assoc. 23rd Biennial Mtg (San Diego, CA)

  15. 2012. Roughgarden J. A principal-agent theory of courtship feeding, nuptial gifts and divorce. Preprint available at

  16. 2012, Roughgarden, J. The social selection alternative to sexual selection. Phil. Trans. Roy. Soc. B. 367:2294-2303.

  17. 2012, Roughgarden, J. Teamwork, pleasure and bargaining in animal social behaviour J. Evol. Biol. 25:1454-1462, doi: 10.1111/j.1420-9101.2012.02505.x

  18. 2012, Folse III, H. and Roughgarden. J. Direct benefits of genetic mosaicism and intraorganismal selection: Modeling coevolution between a long-lived tree and a short-lived herbivore. Evolution 66-4: 1091-1113.

  19. 2011, Roughgarden, J and Iyer, P. Contact, not conflict causes the evolution of anisogamy. pp 96-110, In: The Evolution of Anisogamy: A Fundamental Phenomenon Underlying Sexual Selection. Togashi, T. and Cox, P. (eds.) Cambridge University Press, Cambridge.

  20. 2011, Akçay, E., and Roughgarden, J. The evolution of payoff matrices: providing incentives to cooperate. Proc. R. Soc. B 278:2198-2206, DOI: 10.1098/rspb.2010.2105.

  21. 2011, Armsworth, P.R., Block, B., Eagle, J., and Roughgarden, J. The role of discounting and dynamics in determining the economic efficiency of time-area closures. Theoretical Ecology 4:513-526, DOI 10.1007/s12080-010-0093-x published online October 2010.

  22. 2011, Milam, E., Millstein, R., Potochnik, A., and Roughgarden. J. Sex and sensibility: the role of social selection. Book Symposium. Metascience 20:253-277. DOI 10.1007/s11016-010-9464-6

  23. 2010, Folse III, H. and Roughgarden, J. What is an individual organism? A multilevel selection perspective. Quart. Rev. Biol. 85:447-472.

  24. 2010, Roughgarden, J. The responsibility of theology to science. Tikkun Nov/Dec: 37-39.

  25. 2010, Jung, P. and J. Roughgarden. Gender in heaven: The story of the Ethiopian eunuch in light of evolutionary biology. Pp. 224-240. In: Jung, P. and Vigen, A. (eds.) God, Science, Sex, Gender. University of Illinois Press, Urbana, Illinois.

  26. 2010, Roughgarden, J. Evolutionary biology and sexual diversity. Pp. 89-104. In: Jung, P. and Vigen, A. (eds.) God, Science, Sex, Gender. University of Illinois Press, Urbana, Illinois.

  27. 2010, Armsworth, P.R., Block, B.A., Eagle, J. and Roughgarden, J. The economic efficiency of a time-area closure to protect spawning bluefin tuna. Journal of Applied Ecology, 47:36-46.

  28. 2010, Roughgarden, J. and Akçay, E. Final response: sexual selection needs an alternative. Animal Behaviour. 79(3):e18-e23.

  29. 2010, Roughgarden, J. and Akçay, E. Do we need a Sexual Selection 2.0? Animal Behaviour 79(3):e1-e4.

  30. 2009, Akçay, E, and J. Roughgarden. The perfect family: Decision making in biparental care. PloS ONE 4:e7345 1-10.

  31. 2009, Akçay, E., Van Cleve, J., Feldman, M. W. and Roughgarden, J. A theory for the evolution of other-regard integrating proximate and ultimate perspectives. Proc. Natl. Acad. Sci. USA 106:19061-6.

  32. 2009, Roughgarden, J. Is there a general theory of community ecology? Biology and Philosophy 24:521-529.

  33. 2009, Iyer, P. and J. Roughgarden. Alternation of haploid and diploid generations: Evolution by gamete amplification. Evolutionary Ecology Research 11:57-77.

  34. 2008, Iyer, P. and J. Roughgarden. Dioecy as a specialization promoting sperm delivery. Evolutionary Ecology Research 10:867-892.

  35. 2008, Iyer, P. and J. Roughgarden. Gametic conflict versus contact in the evolution of anisogamy. Theoretical Population Biology 73:461-472.

  36. 2008, Roughgarden, J. Social selection vs. sexual selection: Comparison of hypotheses. In: Daniel Lee Kleinman, Karen Cloud-Hansen, Christina Matta, and Jo Handelsman, eds. Controversies in Science and Technology Volume II: From Climate to Chromosomes. New Rochelle, NY: Mary Ann Liebert, Inc. pp. 421-463.

  37. 2008, Armsworth P. and J. Roughgarden. The structure of clines with fitness dependent dispersal American Naturalist 172:648-657.

  38. 2007, Akçay, E, and J. Roughgarden. Extra-pair parentage: a new theory based on transactions in a cooperative game. Evolutionary Ecology Research. 9:1223-1243.

  39. 2007, Akçay, E, and J. Roughgarden. Extra-pair paternity in birds: Review of the genetic benefits. Evolutionary Ecology Research. 9:855-868.

  40. 2007, Akçay, E., and J. Roughgarden. Negotiation of mutualism: Rhizobia and legumes. Proc. Royal Society B. 274:25-32.

  41. 2007, Roughgarden J. Challenging Darwin's theory of sexual selection. Daedalus. 136(2):1-14

  42. 2006, Akçay, E. and J. Roughgarden. Neither should competition be assumed. Trends in Ecology and Evolution. 22:4

  43. 2006, Roughgarden J., M. Oishi, and E. Akçay. Reproductive social behavior: Cooperative games to replace sexual selection. Science. 311:965-969.

  44. 2006, Roughgarden J. Foreward. In: J. Kritzer and P. Sale (eds.), Marine Metapopulations. Elsevier, Burlington, MA. Pp. xvii--xx

  45. 2006, Buckley, L. and J. Roughgarden. Climate, competition, and the coexistence of island lizards. Functional Ecology 20:315-322.

  46. 2006, Buckley, L. and J. Roughgarden. A hump-shaped density-area relationship for island lizards. Oikos 113:243-250.

  47. 2005, Roughgarden. J. The myth of sexual selection. California Wild 53(3):18-23

  48. 2005, Buckley, L. and J. Roughgarden. Lizard habitat partitioning on islands: the interaction of local and landscape scales. Journal of Biogeography 32(12):2113-2121.

  49. 2005, Buckley, L. and J. Roughgarden. Effect of species interactions on landscape abundance patterns. Journal of Animal Ecology 74:1182-1194.

  50. 2005, Armsworth, P. and J. Roughgarden. Disturbance induces the contrasting evolution of reinforcement and dispersiveness in directed and random movers. Evolution 59(10):2083-2096.

  51. 2005, Armsworth, P. and J. Roughgarden. The impact of directed versus random movement on population dynamics and biodiversity patterns. American Naturalist 165:449-465.

  52. 2004, Pereira, H., G. Daily, and J. Roughgarden. A framework for assessing the relative vulnerability of species to land-use change. Ecological Applications. 14(3):730-742.

  53. 2004, Dial, R. and J. Roughgarden. Physical transport, heterogeneity, and interactions involving canopy anoles. In: Lowman, M. and H. B. Rinker. (eds.) Forest Canopies. Elsevier. Amsterdam. Pp. 270-296.

  54. 2004, Roughgarden, J. The in-crowd. New Scientist 181:36-39.

  55. 2004, Roughgarden, J. Evolution and the embodiment of gender. GLQ: A Journal of Lesbian and Gay Studies. 10:287-291.

  56. 2004, Gerber, L., O. Reichman, and J. Roughgarden. Food hoarding: Future value in optimal foraging decisions. Ecological Modelling. 175:77-85.

  57. 2003, Hughes, T., A. Baird, D. Bellwood, M. Card, S. Connolly, C. Folke, R. Grosberg, O. Hoegh-Guldberg, J. Jackson, J. Kleypas, J. Lough, P. Marshall, M. Nyström, S. Palumbi, J. Pandolfi, B. Rosen, and J. Roughgarden. Climate change, human impacts, and the resilience of coral reefs. Science 301:929-933.

  58. 2003, Armsworth, P. and J. Roughgarden. The economic value of ecological stability. Proc. Nat. Acad. (USA) 100:7147-7151.

  59. 2003, Pereira, H., A. Bergman, and J. Roughgarden. Socially stable territories: The negotiation of space by interacting foragers. American Naturalist 161:143-152.

  60. 2002, Bjorkstedt, E., L. Rosenfeld, B. Grantham, Y. Shkedy, and J. Roughgarden. Distributions of larval rockfish (Sebastes spp.) across nearshore fronts in a coastal upwelling region. Marine Ecology Progress Series 242:215-228.

  61. 2002, Balmford, A., A. Bruner, P. Cooper, R. Costanza, S. Farber, R. Green, M. Jenkins, P. Jefferiss, V. Jessamy, J. Madden, K. Munro, N. Myers, S. Naeem, J. Paavola, M. Rayment, S. Rosendo, J. Roughgarden, K. Trumper, and K. Turner. Economic reasons for conserving wild nature. Science 297:950-953.

  62. 2002, Pereira, H., S. Loarie, and J. Roughgarden. Monogamy, polygyny and interspecific interactions in the lizards Anolis gingivinus and Anolis pogus. Caribbean Journal of Science, 38:8-12.

  63. 2002, Bascompte, J. H. Possingham, and J. Roughgarden. Patchy populations in stochastic environments: Critical number of patches for persistence. American Naturalist 159:128-137.

  64. 2001, Connolly, S., B. A. Menge and J. Roughgarden. A latitudinal gradient in recruitment of intertidal invertebrates in the Northeast Pacific Ocean. Ecology, 82:1799-1813.

  65. 2001, Roughgarden, J. and P. Armsworth. Managing ecosystem services. In: Press, M. N. Huntly, and S. Levin, (eds.) Ecology: Achievement and Challenge, Blackwell Science, pp. 337-356.

  66. 2001, P. Armsworth and J. Roughgarden. An invitation to ecological economics, Trends in Ecology and Evolution 16:229-234.

  67. 2001, Roughgarden, J. Guide to diplomatic relations with economists. Bull. Ecol. Soc. America 82:85-88.

  68. 2000, Hughes, J. and J. Roughgarden. Species diversity and biomass stability. American Naturalist, 155:618-627.

  69. 2000, Boggs, C. and J. Roughgarden. Biodiversity: result of speciation and extinction. Chapter 17 in G. Ernst, ed., Earth Systems: Processes and Issues, Cambridge University Press. Pp 255-266.

  70. 2000, Boggs, C. and J. Roughgarden. Evolution: adaptation and environmental change. Chapter 18 in G. Ernst, ed., Earth Systems: Processes and Issues, Cambridge University Press. Pp. 267-277.

  71. 1999, Connolly, S. and J. Roughgarden. Theory of marine communities: competition, predation and recruitment-dependent interaction strength. Ecological Monographs, 69:277-296.

  72. 1999, Connolly, S. and J. Roughgarden. Increased recruitment of northeast Pacific barnacles during the 1997 El Niño. Limnol. Oceanogr. 44:466-469.

  73. 1999, S. Murray, R. F. Ambrose, J. A. Bohnsack, L. W. Botsford, M. H. Carr, G. E. Davis, P. K. Dayton, D. Gotshall, D. R. Gunderson, M. A. Hixon, J. Lubchenco, M. Mangel, A. MacCall, D. A. McArdle, J. C. Ogden, J. Roughgarden, R. M. Starr, M. J. Tegner, and M. M. Yoklavich. No-take reserve networks: sustaining fishery populations and marine ecosystems. Fisheries Nov. 1999 pp. 11-25.

  74. 1998, Connolly, S. and J. Roughgarden. A range extension for the volcano barnacle, Tetraclita rubescens. California Fish and Game 84:182-183.

  75. 1998, Hughes, J. and J. Roughgarden. Aggregate community properties and the strength of species' interactions. Proc. Nat. Acad. Sci. (USA), 95:6837-6842.

  76. 1998, Dial, R., and J. Roughgarden. Theory of marine communities: the intermediate disturbance hypothesis, Ecology 79:1412-1424.

  77. 1998, Shafir, S. and J. Roughgarden. Testing predictions of foraging theory for a sit-and-wait forager, Anolis gingivinus. Behavioral Ecology 9:74-84.

  78. 1998, Connolly, S. and J. Roughgarden, A latitudinal gradient in northeast Pacific intertidal community structure: evidence for an oceanographically based synthesis of marine community theory. American Naturalist, 151:311-326.

  79. 1998, Roughgarden, J., How to manage fisheries. Ecological Applications, 8(1):S160-S164.

  80. 1998, Roughgarden, J., Production functions from ecological populations: a survey with emphasis on spatially explicit models. In: Tilman, D. and P. Kareiva, (eds.) Spatial Ecology: The Role of Space in Population Dynamics and Interspecific Interactions, Princeton University Press, pp. 296-317.

  81. 1997, Shkedy, Y., and J. Roughgarden, Barnacle recruitment and population dynamics predicted from coastal upwelling. Oikos 80(3): 487-498.

  82. 1997, Bjorkstedt E., and J. Roughgarden, Larval transport and coastal upwelling: an application of HF radar in ecological research. Oceanography 10(2): 64-67.

  83. 1997, Brown, G. and J. Roughgarden, A metapopulation model with private property and a common pool. Ecological Economics, 22:65-71.

  84. 1997, Goldwasser, L. and J. Roughgarden, Sampling effects and the estimation of food web properties. Ecology, 78:41-54.

  85. 1996, Dial, R., and J. Roughgarden, Natural history observations of Anolisomyia rufianalis (Diptera: Sarcophagidae) infesting Anolis lizards in a rain forest canopy. Environ. Entomol. 25:13:25-1328.

  86. 1996, Alexander, S.E. and J. Roughgarden, Larval transport and population dynamic of intertidal barnacles: a coupled benthic/oceanic model. Ecological Monographs, 66:259-275.

  87. 1996, Roughgarden, J. and F. Smith, Why fisheries collapse and what to do about it. Proc. Nat. Acad. Sci., (USA), 93:5078-5083

  88. 1996, Shafir, S. and J. Roughgarden, The effect of memory length on the individual fitness of a lizard. In Belew, R.K. and M. Mitchell (Eds.), Adaptive Individuals in Evolving Populations: Models and Algorithms, Santa Fe Institute Studies in the Science of Complexity, Addison Wesley, Reading, MA., pp. 173-182.

  89. 1996, Roughgarden, J., A. Bergman, S. Shafir and C. Taylor, Adaptive computation in ecology and evolution: a guide for future research. In Belew, R.K. and M. Mitchell (Eds.), Adaptive Individuals in Evolving Populations: Models and Algorithms, Santa Fe Institute Studies in the Science of Complexity, Addison Wesley, Reading, MA., pp. 25-32.

  90. 1996, Roughgarden, J., Ecology. In: Stephen H. Schneider (ed) Encyclopedia of Climate and Weather. (1st Ed.) Pp. 268-269.

  91. 1995, Rosenfeld, L., T. Anderson, G. Hatcher, J. Roughgarden, and Y. Shkedy, Upwelling fronts and barnacle recruitment in central California. Technical Report 95-19, Monterey Bay Aquarium Research Institute. Moss Landing, CA

  92. 1995, Roughgarden, J., Can economics protect biodiversity? Pp. 149-156 in: T. Swanson, Ed., The Economics and Ecology of Biodiversity Decline. Cambridge University Press.

  93. 1995, Holling, C.S., D.W. Schindler, B.W. Walker and J. Roughgarden, Biodiversity in the functioning of ecosystems: an ecological synthesis. In Perings, C., C.G. Maler, C. Folke, C.S. Holling, and B.O. Jansson (Eds.), Biodiversity Loss: Economic and Ecological Issues, Cambridge, Cambridge University Press, pp. 44-83.

  94. 1995, Brown, G. and J. Roughgarden, An ecological economy: notes on harvest and growth. In Perrings, C., K.G. Maler, C. Folke, C.S. Holling and B.O. Jansson (Eds.), Biodiversity Loss: Ecological and Economic Issues, Cambridge, Cambridge University Press, pp. 150-189. (Reprinted in: J. Barkley Rosser, Jr., 2004. Complexity in Economics, Volume III. The International Library of Critical Writings in Economics 174. Edward Elgar, Cheltenham, UK, pp. 383-423.)

  95. 1995, Roughgarden, J., Vertebrate patterns on islands. In Vitousek, P.M., L.L. Loope, and H. Andersen (Eds.), Islands: Biological Diversity and Ecosystem Function, Springer, Germany, pp. 51-56.

  96. 1995, Dial, R., and J. Roughgarden, Experimental removal of insectivores from rain forest canopy: direct and indirect effects. Ecology, 76 (6) pp. 1821-1834.

  97. 1994, Shafir, S., and J. Roughgarden, The effect of memory length on the foraging behavior of a lizard. In Cliff, D., P. Husbands, J.A. Meyer, and S.W. Wilson (Eds.), From Animals to Animats 3: Proceedings of the Third International Conference on Simulation of Adaptive Behavior, MIT Press, Cambridge, MA. pp. 221-225.

  98. 1994, Shafir, S., and J. Roughgarden, Instrumental discrimination conditioning of Anolis cristatellus in the field with food as a reward. Caribbean Journal of Science, 30 (3-4): 228-233.

  99. 1994, Dial, R. and J. Roughgarden, Notes on the absolute abundance of canopy anoles: Anolis cuvieri, Anolis stratulus, and Anolis evermanni (Lacertilia: polychridea) in the Luquillo Forest, Puerto Rico. Caribbean Journal of Science, 30 (3-4): 278-279.

  100. 1993, Fernandez, D., C. Teague, J. Vesecky, and J. Roughgarden, Upwelling and relaxation events inferred from high-frequency radar measurements of coastal surface currents. Proceedings of the IGARSS '93 Conference -- IEEE International Geoscience and Remote Sensing Symposium, 4:1931-1933.

  101. 1993, Miller, K., and J. Roughgarden, Descriptions of the larvae of Tetraclita rubescens and Megabalanus californicus with a comparison of the common barnacle larvae of the central California coast. Journal of Crustacean Biology, pp. 579-600.

  102. 1993, Shkedy, Y., D. Fernandez, C. Teague, J. Vesecky, and J. Roughgarden, Detecting upwelling along the central coast of California during an El-Niño year using HF-radar. Continental Shelf Research, 15 (7): 803-814.

  103. 1993, Roughgarden, J., H. Possingham, S. Tuljapurker and M. Wilks, Population cycling in space-limited organisms subject to density-dependent predation. The American Naturalist, 143 (4): 563-582.

  104. 1993, McLaughlin, J., and J. Roughgarden, Species interactions in space. Chapter 8 in Species Diversity in Ecological Communities: Historical and Geographical Perspectives, Ed. by R. E. Ricklefs and D. Schluter, pp. 89-98.

  105. 1993, Roughgarden, J., Pennington and S. Alexander, Dynamics of the rocky intertidal zone with remarks on generalization in ecology. Philosophical Transactions of the Royal Society London B., 341:449-454.

  106. 1993, Goldwasser, L., and J. Roughgarden, Construction and analysis of a large Caribbean food web. Ecology, 74:4, pp. 1216-1233.

  107. 1992, Dobson, A., S. Pacala, J. Roughgarden, E. Carper, and E. Harris., The Parasites of Anolis lizards in the northern Lesser Antilles. Oecologia, 91:1, pp. 110-117.

  108. 1992, Roughgarden, J., Comments on the paper by Losos: Character displacement versus taxon loop. Copeia, 2:288-295.

  109. 1992, Koza, J., J. Rice, and J. Roughgarden., Evolution of food-foraging strategies for the Caribbean Anolis lizard using genetic programming. Adaptive Behavior, 1:2, pp. 171-199.

  110. 1991, Roughgarden, J., S. Running, and P. Matson, What does remote sensing do for ecology? Ecology, 72:6, pp. 1918-1922.

  111. 1991, Miller, K., P. Jones, and J. Roughgarden, Monoclonal antibodies as species-specific probes in oceanographic research: Examples with intertidal barnacle larvae. Molecular Marine Biology and Biotechnology, 1:35-47.

  112. 1991, McLaughlin, J., and J. Roughgarden, Predation across spatial scales in heterogeneous environments. Theoretical Population Biology, 40:2, pp. 149-172.

  113. 1991, Farrell, T., D. Bracher, and J. Roughgarden, Cross-shelf transport causes recruitment to intertidal populations in central California. Limology and Oceanography, 36:279-288.

  114. 1991, McLaughlin, J., and J. Roughgarden, Pattern and stability in predator communities: How diffusion in spatially variable environments affects the Lotka-Volterra model. Theoretical Population Biology, 40:148-172.

  115. 1991, Roughgarden, J., J. Pennington, D. Stoner, S. Alexander, and K. Miller, Collisions of upwelling fronts with the intertidal zone: The cause of recruitment pulses in barnacle populations of central California. Acta Oecologica, 12:35-51.

  116. 1991, Roughgarden, J., The evolution of sex. American Naturalist, 138(4) pp. 934-985.

  117. 1990, Possingham, H., and J. Roughgarden, Spatial population dynamics of an organism with a complex life cycle. Ecology, 71:973-985.

  118. 1990, Roughgarden, J., Origin of the Eastern Caribbean: Data from reptiles and amphibians. In D. K. Larue and G. Draper (Eds.), Trans. 12th Caribbean Geological Conference, St. Croix, USVI, Miami Geological Survey, pp. 10-26.

  119. 1990, Naganuma, K., and J. Roughgarden, Optimal body size in Lesser Antillean Anolis lizards-A mechanistic approach. Ecological Monographs, 60:239-256.

  120. 1990, Brown, J., and J. Roughgarden, Ecology for a changing earth. Bulletin of the Ecological Society of America, 71:173-188.

  121. 1989, Roughgarden, J., The structure and assembly of communities. In Roughgarden, R. M. May, and S. A. Levin (Eds.) Perspectives in Ecological Theory, Princeton University Press, pp. 203-226.

  122. 1989, Roughgarden, J., Viewpoint: The United States needs an ecological survey. Bioscience, 39:5.

  123. 1989, McLaughlin, J., and J. Roughgarden, Avian predation on Anolis lizards in the northeastern Caribbean: An inter-island contrast. Ecology, 70:617-628.

  124. 1989, Brown, J., and J. Roughgarden, U. S. ecologists address global change. Trends in Ecology and Evolution, 4:255.

  125. 1989, Blower, S., and J. Roughgarden, Parasites detect host spatial pattern and density: A field experimental analysis. Oecologia, 78:138-141.

  126. 1989, Miller, K., S. Blower, D. Hedgecock, and J. Roughgarden, Comparison of larval and adult stages of Chthamalus dalli and Chthamalus fissus (Cirripedia: Thoracica). Journal Crustacean Biology, 9:242-256.

  127. 1989, Blower, S., and J. Roughgarden, Population dynamics and parasitic castration: A mathematical model. American Naturalist, 134:848-858.

  128. 1988, Roughgarden, J., and S. Pacala, Taxon cycle among Anolis lizard populations: Review of evidence. In Daniel Otte and John Endler (Eds.), Speciation and its Consequences, Sinauer Associates, Inc., pp. 403-432.

  129. 1988, Blower, S., and J. Roughgarden, Parasitic castration: host species preferences, size-selectivity and spatial heterogeneity. Oecologia, 75:512-515.

  130. 1988, Roughgarden, J., S. Gaines, and H. Possingham, Recruitment dynamics in complex life cycles. Science, 241:1460-1466.

  131. 1988, Roughgarden, J., The evolution of marine life cycles. In M. W. Feldman (Ed.), Mathematical Evolutionary Theory, Princeton University Press, pp. 270-300.

  132. 1987, Roughgarden, J., Community coevolution: A comment. Evolution, 41:5,1130-1134.

  133. 1987, Gaines, S., and J. Roughgarden, Fish in offshore kelp forests affect recruitment to intertidal barnacle populations. Science, 235:479-481.

  134. 1987, Roughgarden, J., S. Gaines, and S. Pacala, Supply side ecology: The role of physical transport processes. In J. H. R. Gee and P. S. Giller (Eds.), Organization of Communities, Past and Present, The 27th Symposium of the The British Ecological Society, Blackwell Scientific Publications, pp. 491-518.

  135. 1987, Blower, S., and J. Roughgarden, Population dynamics and parasitic castration: Test of a model. American Naturalist, 134:848-858

  136. 1986, Roughgarden, J., and Y. Iwasa, Dynamics of a metapopulation with space-limited subpopulations. Theoretical Population Biology, 29:235-261.

  137. 1986, Roughgarden, J., and J. Diamond, Overview: The role of species interactions in community ecology. In J. Diamond and T. Case (Eds.), Community Ecology, Harper & Row, New York, pp. 333-344.

  138. 1986, Roughgarden, J., A Comparison of food-limited and space-limited animal competition communities. In J. Diamond and T. Case (Eds.), Community Ecology, Harper & Row, New York, pp. 492-516.

  139. 1986, Roughgarden, J., Predicting invasions and rates of spread. In Harold Mooney and James A. Drake (Eds.), Ecology of Biological Invasions of North America and Hawaii, Ecological Studies, Springer-Verlag, 58:179-188.

  140. 1986, Iwasa, Y., and J. Roughgarden, Interspecific competition among metapopulations with space-limited subpopulations. Theoretical Population Biology, 30:260-214.

  141. 1986, Roughgarden, J., T. Schoener, and T. Fenchel, The body-size-prey-size hypothesis: A defense. Ecology, 67:260-261.

  142. 1986, Roughgarden, J., Models of population processes for plants. In L. J. Gross and R. M. Miura (Eds.), Some Mathematical Questions in Biology--Plant Biology. Lectures on Mathematics in the Life Sciences, American Mathematical Society, Providence, R.I., 18:235-267.

  143. 1985, Brown, S., and J. Roughgarden, Growth, morphology, and laboratory culture of Balanus glandula larvae (Cirripedia:Thoracica). J. Crustacean Biology, 5:574-590.

  144. 1985, Rummel, J., and J. Roughgarden., Effects of reduced perch-height separation on competition between two Anolis lizards. Ecology, 66:2 pp. 430-444.

  145. 1985, Roughgarden, J., Y. Iwasa, and C. Baxter, Demographic theory for an open marine population with space-limited recruitment. Ecology, 66:54-57.

  146. 1985, Gaines, S., and J. Roughgarden, Larval settlement rate: A leading determinant of structure in an ecological community of the marine intertidal zone. Proc. Nat. Acad. Sci., (USA), 82:3707-3711.

  147. 1985, Gaines, S., S. Brown, and J. Roughgarden, Spatial variation in larval concentrations as a cause of spatial variation in settlement for the barnacle, Balanus glandula. Oecologia, 67:267-272.

  148. 1985, Iwasa, Y., and J. Roughgarden, Evolution in a metapopulation with space-limited subpopulation. IMA Journal of Mathematics Applied in Medicine & Biology, 2:93-107.

  149. 1985, Pacala, S., and J. Roughgarden, Population experiments with the Anolis lizards of St. Maarten and St. Eustatius. Ecology, 66:129-141.

  150. 1985, Rummel, J., and J. Roughgarden, A theory of faunal buildup for competition communities. Evolution, 39:1009-1033.

  151. 1985, Holsinger, K., and J. Roughgarden, A model for the dynamics of an annual plant population. Theoretical Population Biology, 28:288-313.

  152. 1984, Iwasa, Y., and J. Roughgarden, Shoot/root balance of plants: Optimal growth of a system with many vegetative organs. Theoretical Population Biology, 25:78-105.

  153. 1984, Pacala, S., and J. Roughgarden, Control of arthropod abundance by Anolis lizards on St. Eustatius (Neth. Antilles). Oecologia, 64:160-162.

  154. 1984, Roughgarden, J., S. Pacala, and J. Rummel, Strong present-day competition between the Anolis lizard populations of St. Maarten (Neth. Antilles). In B. Shorrocks (Ed.), Evolutionary Ecology, The 23rd Symposium of the British Ecological Society, Blackwell Scientific Publications, pp. 203-220.

  155. 1984, Roughgarden, J. and N. Chiariello, Storage allocation in seasonal races of an annual plant: Optimal versus actual allocation. Ecology, 65:1290-1301.

  156. 1984, Roughgarden, J., S. Gaines, and Y. Iwasa, Dynamics and evolution of marine populations with pelagic larval dispersal. In R. M. May (Ed.), Exploitation of Marine Communities. Dahlem Konferenzen, Berlin. Springer-Verlag, pp. 111-128.

  157. 1983, Roughgarden, J., D. Heckel, and E. Fuentes, Coevolutionary theory and the biogeography and community structure of Anolis. In R. Huey, E. Pianka and T. Schoener (Eds.), Lizard Ecology: Studies of a Model Organism. Harvard University Press, pp. 371-410.

  158. 1983, Roughgarden, J., J. Rummel, and S. Pacala, Experimental evidence of strong present-day competition between Anolis populations of the Anguilla Bank, A preliminary report. In A. Rhodin and K. Miyata, (Eds.), Advances in Herpetology and Evolutionary Biology: Essays in Honor of Ernest Williams. Museum of Comparative Zoology, Harvard University Press, pp. 499-506.

  159. 1983, Adolph, S., and J. Roughgarden, Foraging by Passerine birds and Anolis lizards on St. Eustatius (Neth. Antilles): Implications for interclass competition and predation. Oecologia, 56:313-317.

  160. 1983, Pacala, S., J. Rummel, and J. Roughgarden, A technique for enclosing Anolis lizard populations under field conditions. J. Herpetology, 17:94-97.

  161. 1983, King, D., and J. Roughgarden, Energy allocation patterns of the California grassland annuals, Plantago erecta and Clarkia rubicunda. Ecology, 64:16-24.

  162. 1983, Roughgarden, J., The theory of coevolution. In D. Futuyma and M. Slatkin (Eds.), Coevolution, Sinauer Associates, pp. 33-64.

  163. 1983, Roughgarden, J., Coevolution between competitors. In D. Futuyma, and M. Slatkin (Eds.), Coevolution, Sinauer Associates, pp. 383-403.

  164. 1983, Roughgarden, J., Competition and theory in community ecology. American Naturalist, 122:583-601.

  165. 1983, Rummel, J., and J. Roughgarden, Some differences between invasion-structured and coevolution-structured competitive communities: A preliminary theoretical analysis. Oikos, 41:477-486.

  166. 1982, Pacala, S., and J. Roughgarden, Spatial heterogeneity and interspecific competition. Theoretical Population Biology, 21:92-113.

  167. 1982, King, D., and J. Roughgarden, Graded allocation between vegetative and reproductive growth for annual plants in growing seasons of random length. Theoretical Population Biology, 22:1-16.

  168. 1982, King, D., and J. Roughgarden, Multiple switches between vegetative reproductive growth in annual plants. Theoretical Population Biology, 21:194-204.

  169. 1982, Shigesada, N., and J. Roughgarden, The role of rapid dispersal in the population dynamics of competition. Theoretical Population Biology, 21:353-372.

  170. 1982, Pacala, S., and J. Roughgarden, The evolution of resource partitioning in a multidimensional resource space. Theoretical Population Biology, 22:127-145.

  171. 1982, Pacala, S., and J. Roughgarden, Resource partitioning and interspecific competition in two two-species insular Anolis lizard communities. Science, 217:444-446.

  172. 1982, Roughgarden, J., W. Porter and D. Heckel, Resource partitioning of space and its relationship to body temperature in Anolis lizard populations. Oecologia, 50:256-264.

  173. 1981, Anderson, G., A. Ehrlich, P. Ehrlich, J. Roughgarden, B. Russell, and F. Talbot, The community structure of coral reef fishes. American Naturalist, 117:476-495.

  174. 1981, Roughgarden, J., W. Porter, and D. Heckel, Resource partitioning of space and its relationship to body temperature in Anolis lizard populations. Oecologia, 50:256-264.

  175. 1980, Heckel, D., and J. Roughgarden, A species near its equilibrium size in a fluctuating environment can evolve a lower intrinsic rate of increase. Proc. Nat. Acad. Sci., 77:7497-7500.

  176. 1979, Heckel, D., and J. Roughgarden, A technique for estimating the size of lizard populations. Ecology, 60:966-975.

  177. 1979, Roughgarden, J., Coevolution in ecological systems III. Coadaptation and equilibrium population size. In P. F. Brussard (Ed.), Ecological Genetics: The Interface, Springer-Verlag, New York, pp. 27-50.

  178. 1979, Roughgarden, J., A local concept of structural homology for ecological communities with examples from simple communities of West Indian Anolis lizards. In U. Halbach and J. Jacobs (Eds.), Population Ecology, Fortschritte der Zoology. Symposium Mainz, May 1978. Band 25, Heft 2/3 149-158.

  179. 1978, Roughgarden, J., Influence of competition on patchiness in a random environment. Theoretical Population Biology, 14:185-203.

  180. 1977, Roughgarden, J., Patchiness in the spatial distribution of a population caused by stochastic fluctuations in resources. Oikos, 29:52-59.

  181. 1977, Roughgarden, J., Coevolution in ecological systems: Results from ``loop analysis" for purely density-dependent coevolution. In F. B. Christiansen and T. M. Fenchel (Eds.), Measuring Selection in Natural Populations, Vol. 19 in Lecture Notes in Biomathematics. Springer-Verlag, New York, pp. 499-517.

  182. 1977, Roughgarden, J., and E. Fuentes, The environmental determinants of size in solitary populations of West Indian Anolis lizards. Oikos, 29:44-51.

  183. 1977, Yoshiyama, R., and J. Roughgarden, Species packing in two dimensions. American Naturalist, 111:107-121.

  184. 1976, Roughgarden, J., Resource partitioning among competing species: A coevolutionary approach. Theoretical Population Biology, 9:388-424.

  185. 1976, Roughgarden, J., A simple model for population dynamics in stochastic environments. American Naturalist, 109:713-736.

  186. 1975, Roughgarden, J., Populations dynamics in a stochastic environment: Spectral theory for the linearized N-species Lotka-Volterra competition equations. Theoretical Population Biology, 7:1-12.

  187. 1975, Feldman, M., and J. Roughgarden, A population's stationary distribution and chance of extinction in a stochastic environment with remarks on the theory of species packing. Theoretical Population Biology, 7:197-207.

  188. 1975, Roughgarden, J., and M. Feldman, Species packing and predation pressure. Ecology, 56:489-492.

  189. 1975, Roughgarden, J., Evolution of marine symbiosis - a simple cost-benefit model. Ecology, 56:1201-1208.

  190. 1974, Roughgarden, J., The fundamental and realized niche of a solitary population. American Naturalist, 108:232-235.

  191. 1974, Roughgarden, J., Species packing and the competition function with illustrations from coral reef fish. Theoretical Population Biology, 5:163-186.

  192. 1974, Roughgarden, J., Niche width: Biogeographic patterns among Anolis lizard populations. American Naturalist, 108:429-442.

  193. 1974, Roughgarden, J., Population dynamics in a spatially varying environment: How population size "tracks" spatial variation in carrying capacity. American Naturalist, 108:649-664.

  194. 1972, Roughgarden, J., Evolution of niche width American Naturalist, 106:683-718.

  195. 1971, Roughgarden, J., Density-dependent natural selection Ecology, 52:453-468.

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