Fish and Wildlife Population Ecology  - Dr. Edward O. Garton




Wlf 541:  Advanced Population Biology

Fall 2010 - 3 Credits

MWF 10:30-11:20 - CNR 203

Instructors: E. O. Garton, CNR 104E, 885-7426, (
                  Brian Kennedy, CNR 103C, 885-5171, (
                  Dan Schill, IDFG, Boise and Nampa, 334-3791, (

Office Hours: Oz Garton: MF 9:30-10:30 a.m.
                      TTh 3:30-4:30 p.m. Other times by arrangement

Have you read the classic papers that form the foundation for modern theories of population ecology? Participants in this class will have the opportunity to read and discuss some of the most significant and interesting papers on population ecology, published in the scientific literature historically and within the last decade. We will begin with a brief review on the scientific method where we'll extend the classic Popperian paradigm to incorporate complex issues such as pseudoreplication, multicausal systems, quasi-experimentation and the growing use of information theoretic tools. We will read a series of classic papers on theoretical topics and experimental concepts that have continued to be important such as niche, competition, habitat selection, natality, mortality, dispersal and life-history strategies before exploring the genetic structure of populations. Next we will focus on density-dependence, -independence, -vagueness, population cycles, eruptions, chaos, regulation, and viability. A major section will explore fundamental and optimization approaches to stability of predator-prey interactions while evaluating these concepts in the context of modern harvest and management practices. The final sections will explore efforts at integration and application across large spatial scales for hierarchically structured metapopulations and those population processes projected to experience impacts due to climate and landscape change.


The primary goal of this course is to develop an understanding of the major concepts of population dynamics. We will do this by reading and discussing a selection of classical papers in population biology. The discussions are directed toward analyzing and criticizing the concepts proposed and only secondarily toward criticizing the papers themselves. I hope that this will stimulate your interest in the literature in the field of population ecology.


This course is conducted as a seminar in which the students and instructor lead discussions of important papers selected from the literature in population biology. At appropriate times the instructor will provide brief comments to help students understand the mathematical aspects of significance or the general subject matter being discussed. Each student should prepare for a discussion on a particular paper by reading the paper carefully. If you are in charge of a discussion on a topic you should find a half-dozen other recent papers on the topic which support or refute theory presented in the selected paper. Hand out a brief bibliography at the start of the class and then spend about one third of the period reviewing the selected paper for that day along with the supporting papers. During the remaining time the student in charge should direct discussion of the paper into a critical examination of the paper's implications and significance by raising a series of questions for discussion. Try to avoid excessive criticism of the validity of the paper but use it as a spring-board for better ideas.

A copy of each paper is available from a web site for the class. Each student in the class is responsible for reading the papers before the presentation. 


A short mid-term and final examination will be given in this course. Each will consist of two take-home questions requiring the integration of the concepts covered. Answers will be limited to two pages (500 words) per question. These two exams will make up one-half of the grade, the other half being based on the students' paper presentations.

 Download PAPERS --


8/25 EOG Romesburg, H. C. 1981. Wildlife science: gaining reliable knowledge. J. Wildl. Manage. 45:293-313.

8/25 EOG Quinn, J. F., and A. E. Dunham. 1983. On hypothesis testing in ecology and evolution. Amer. Natur. 122:602-617.

8/25 EOG Hurlbert, S. H. 1984. Pseudoreplication and the design of ecological field experiments. Ecol. Monogr. 54:187-211.

8/27 SKIP Cook, T. D. and D. T. Campbell. 1979. Causal inference and the language of experimentation. Pages 1-36 In Cook, T. D., and D. T. Campbell. Quasi-experimentation: Design and analysis issues for field settings. Houghton Mifflin Co., Boston, Mass. 405 p.

8/27 Brian Hobbs, N. T. and R. Hilborn.  2006.  Alternatives to hypothesis testing in ecology: a guide to self teaching.  Ecological Applications 16:5-19.

8/27  Brian Sinclair, A. R. E. 1991. Science and the practice of wildlife management. J. Wildl. Manage. 55:767-773.

9/1  GiffJames, F. C. and C. E. McCulloch. 1985. Data analysis and the design of experiments in ornithology. Pages 1-63 In R. F. Johnston, ed. Current Ornithology, Vol. 2, Plenum Press, New York, N. Y.

9/1  Giff  Kuhn, Thomas S.  1970.  The Structure of Scientific Revolutions (2nd ed.). Chapter 2 - The route to normal science pp. 10-22.  University of Chicago Press, Chicago, IL.

9/1  Giff  Burnham, K. P. and D. R. Anderson.  2002.  Model Selection and Multimodel Inference: A Practical Information-Theoretic Approach.  2nd ed. Chap. 8 Summary, pp. 437-454.  Springer, New York, NY.

9/ Dan  Johnson, Douglas H.  1999.  The insignificance of statistical significance testing.  Journal of Wildlife Management 63:763-772.  

9/3  Dan Garton, E. O., J. Horne, J Aycrigg, and J. Ratti.  2010. Research and experimental design.  Preprint of pages ??-?? In Nova Silvy (editor). Techniques for Wildlife Investigations and Management.  The Wildlife Society, Bethesda, MD.


9/8    Jeff  Hutchinson, G. Evelyn.  Concluding remarks.  Cold Spring Harbor Symposium Quantitative Biology 22:415-427.

9/8     Jeff Whittaker, R. H., S. A. Levin, and R. B. Root. 1973. Niche, habitat and ecotope. Amer. Natur. 107:321-328.

9/10    Sam Root, R. B. 1967. The niche exploitation of the Blue-Gray Gnatcatcher. Ecol. Monogr. 37:317-319, 331-349.

9/10   Sam Ayala, F. J. 1968. Genotype, environment and population numbers. Science 162:1453-1459.


9/13          Gause, G. F. 1934.  The Struggle for Existence.  Williams and Wilkins/Reprinted by Dover, New York, NY.  pp 44-58, 90-113.

9/13 __   _ Ayala, F. J.  1969. Experimental invalidation of the principle of competitive exclusion.  Nature 224:1076-1079.

9/13 __   _ Roughgarden, J. 1983. Competition and theory in community ecology. Amer. Natur. 122:583-601.

9/13          Simberloff, D. 1983. Competition theory, hypothesis testing, and other community ecological buzzwords. Amer. Natur. 122:626-635.

9/13  _   __  Tilman, D. 1982. Resource Competition and Community Structure. Monographs in Population Biology No. 17 ( p 43-96 only) Princeton Univ. Press, Princeton , NJ.

9/15           Heller, H. C. 1971. Altitudinal zonation of chipmunks (Eutamias): Interspecific aggression. Ecology 52:312-319.

Skip            Strong, D. R., Jr. 1983. Natural variability and the manifold mechanisms of ecological communities. Amer. Natur. 122:636-660.

9/15            MacArthur, R. H. 1958. Population ecology of some warblers of northeastern coniferous forests. Ecology 39:599-612, 617-619.

9/17 __    _ Odum, H. T. and W. C. Allee.  1954.  A note on the stable point of populations showing both intraspecific cooperation and disoperation.  Ecology 35: 95-97..

9/17           Noon, B. R. 1981. The distribution of an avian guild along a temperate elevational gradient: The importance and expression of competition. Ecol. Monogr. 51(1):105-124.

9/17            Connell, J. H. 1983. On the prevalence and relative importance of interspecific competition: evidence from field experiments. Amer. Natur. 122:661-696.


9/20             Fretwell, S. D. & Lucas, H. L., Jr. 1970. On territorial behavior and other factors influencing habitat distribution in birds. I. Theoretical Development. Acta Biotheoretica 19: 16–36.

9/20               Van Horne, B.  Density as a misleading indicator of habitat quality.  Journal of Wildlife Management 47:893-901.

9/20               Rosenzweig, M. L. 1991. Habitat selection and population interactions: The search for mechanism. Amer. Natur. 137:S5-S28.

9/20             Pulliam, H. R. and B. J. Danielson. 1991. Sources, sinks, and habitat selection: A landscape perspective on population dynamics. Amer. Natur. 137:S50-S66.


9/22            Cody, M. 1966. A general theory of clutch size. Evolution 20:174-184.

9/22             Murray, B. G., Jr. 1991. Sir Isaac Newton and the evolution of clutch size in birds: A defense of the hypothetico-deductive method in ecology and evolutionary biology. Pages 143-180 In J. L. Casti and A. Karlqvist (eds.) Beyond belief: Randomness, predition and explanation in science. CRC Press, Boca Raton. 334p.

9/22             Murdoch, W.W.  1966.  Population stability and life history phenomena.  Amer. Nat. 100:5-11.

9/24             Murphy, G. 1968.  Pattern in life history and the environment.  American Naturalist 102:391-403.

9/24             Charnov, E.L., & W.M. Schaffer.  1973.  Life history consequences of natural selection: Cole's result revisited. Amer. Nat. 107:791-793.

9/27             Luckinbill, L.S.  1984.  An experimental analysis of a life history theory.  Ecology 65:1170-1184.

9/27______Gross, M. R.   R. M. Coleman and R. M. McDowall.  1988.  Aquatic productivity and the evolution of diadromous fish migration.  Science 239: 1291-1293.

9/27______Gross, M. R.  1996.  Alternative reproductive strategies and tactics: diversity within sexes.  TREE 11(2):92-97.

9/27 _____ Reznick, D., M. J. Bryant, and F. Bashey.  2002.  r-and K-selection revisited: the role of population regulation in life-history evolution.  Ecology 83:1509-1520.


9/29              Caughley, G. 1966. Mortality patterns in mammals. Ecology 47:906-918.

9/29           Botkin, D. B., and R. S. Miller. 1974. Mortality rates and survival of birds. Amer. Natur. 108:181-192.

9/29            Conover, D. O., and S. B. Munch.  2002.  Sustaining fisheries yields over evolutionary time scales.  Science 297: 94-96.


10/1            Wright, S. 1940. Breeding structure of populations in relation to speciation. Amer. Natur.74:232-248.

10/1             Nelson, M. E. and L. D. Mech. 1987. Demes within a Northeastern Minnesota deer population. pp 27-40 in B. D. Chepko-Sade and Z. T. Halpin, eds. Mammalian Dispersal Patterns: The Effects of Social Structure on Population Genetics. Univ. Chicago Press, Chicago, IL.

10/1            Lande, R. 1988. Genetics and demography in biological conservation. Science 241:1455-1460.


10/4       Solomon, M. E. 1958. Meaning of density-dependence and related terms in population dynamics. Nature 181:1778-1780.

10/4         Andrewartha, H. G. 1961. Introduction to the study of animal populations. Univ. of Chicago Press, Chicago, IL. pp 5-10 only.

Skip             Strong, D. R. 1984. Density vague ecology and liberal population regulation in insects. pp. 313-327 in Price, P. W., C. N. Slobodchikoff, and W. S. Gaud, (eds.). A new ecology: Novel approaches to interactive systems. 515 pp.

10/4             Ostfeld, R. S. and C. D. Canham. 1995. Density-dependent processes in meadow voles: an experimental approach. Ecology 76: 521-532.


10/6            Chitty, D. 1967. The natural selection of self-regulatory behavior in animal populations. Proc. Ecol. Soc. of Australia 2:51-78.

10/6           Christian, J. J., and D. E. Davis. 1964. Endocrines, behavior and population. Science 146:1550-1560.

10/6              Keith, L. B., J. R. Cary, O. J. Rongstad, and M. C. Brittingham. 1984. Demography and ecology of a declining snowshoe hare population. Wildl. Monogr. No. 90. 43pp.

10/8            Krebs, C. J., S. Boutin, R. Boonstra, A. R. E. Sinclair, J. N. M. Smith, R. R. Dale, K. Martin, and R. Turkington.  1995.  Impact of food and predation on the snowshoe hare cycle.  Science 269: 112-115.

10/8           Krebs, C. J. 1970. Genetic and behavioral studies on fluctuating vole populations. Proc. Adv. Study Inst. Dynamics Numbers Popul. (Oosterbeek), pp. 243-256.

10/8           Caughley, G. 1970. Erruption of ungulate populations, with emphasis on Himalayan thar in New Zealand. Ecology. 51(1):53-72.

10/11          May, R. M. 1974. Biological populations with non-overlapping generations: stable points, stable cycles, and chaos. Science 186:645-647.

10/11            Tilman, D. and D. Wedin. 1991. Oscillations and chaos in the dynamics of a perennial grass. Nature 353: 653-655.

10/13            Hanski, I., P. Turchin, E. Korpimaki, and H. Henttonen. 1993. Population oscillations of boreal rodents: regulation by mustelid predators leads to chaos. Nature 364:232-235.

10/13             Ritchie, M. E. 1992. Chaotic dynamics in food-limited populations: implications for wildlife management. p 139-146 in D. R. McCullough and R. H. Barrett (eds.) Wildlife 20001: Populations. Elsevier.



10/15           Lack, D. 1954. The natural regulation of animal numbers. Chapter 13. pp. 141-153. Food as a limiting factor in birds. Clarendon Press, Oxford.

10/15             Wynn-Edwards, V. C. 1962. Animal dispersion in relation to social behaviors. Chapter 1, pp. 1-22. An outline of the principle of animal dispersion.

10/18            Watson, A. and R. Moss. 1969. Dominance, spacing behavior and aggression in relation to population limitation in vertebrates, pp. 167-218 in Watson, A. (ed.). Animal populations in relation to their food resources. Blackwell Sci. Pub., Oxford.

10/18 _______ Pulliam, H.R.  1988.  Sources, sinks and population regulation.  American Naturalist 132:652-661.


10/20          Shaffer, M. L. 1981. Minimum population sizes for species conservation. BioScience 31(2):131-134.

____            Shaffer, M. L. 1985. The metapopulation and species conservation: The special case of the northern spotted owl. pp. 86-99 in Gutiérrez, R. J., and A. B. Carey (eds.). Ecology and management of the spotted owl in the Pacific Northwest. USDA Forest Service, General Tech. Report PNW-185. Portland, OR. 119pp.

_____ Lande, R.  1993.  Risks of population extinction from demographic and environmental stochasticity and random catastrophes.  American Naturalist 142: 911-927.

10/20 Shaffer, M. L., L. H. Watchman, W. J. Snape, III and I. K. Latchis.  2002.  Population viability analysis and conservation policy.  pp. 127-141 in S. r. Beissinger and D. R. McCullough (eds).  Population Viability Analysis. University of Chicago Press, Chicago, IL. 

10/22               Harris, R. B., and F. W. Allendorf. 1988. Genetically effective population size of large mammals: an assessment of estimators. Conservation Biology 3:181-191.

10/22           Senner, J. W. 1980. Inbreeding depression and the survival of zoo populations. pp. 209-224 in Soule, M. E., and B. A. Wilcox (eds.). Conservation biology: An evolutionary-ecological perspective. Sinauer Associates, Sunderland, MA. 395pp.

10/22          Chepko-Sade, B. D., and W. M. Shields. 1987. The effects of dispersal and social structure on effective population size. pp.287-321 in Chepko-Sade, B. D., and Z. T. Halpin, (eds.). Mammalian dispersal patterns. The effects of social structure on population genetics. University of Chicago Press, Chicago, IL. 324pp.


10/25 ______ Lotka,A. J.  1924.  pp. 77-79,88-94 in Elements of Physical Biology: reprinted in 1956 as Elements of Mathematical Biology, Dover, New York, NY. 

Functional Response

10/25           Holling, C. S. 1959. Some characteristics of simple types of predation and parasitism. Canad. Entom. 91:385-398.

10/25            Holling, C. S. 1959. The components of predation as revealed by a study of small mammal predation of the European Pine Sawfly. Canad. Entom. 90:293-320.

10/25 _____ Vucetich, J. A., R. O. Peterson, and C. L. Schaefer.  2002.  The effect of prey and predator densities on wolf predation.  Ecology 83:3003-3013.

 Search Images

10/27          Heinrich, B. and S. L. collins.  1983.  Caterpillar damage, and the game of hide-and-seek with birds.  Ecology 64:592-602.

10/27           Gibb, J. A. 1962. L. Tinbergen's hypothesis of the role of specific search images. Ibis 104:106-111.

Optimal Diet

10/29            Pyke, G. H., H. R. Pulliam, and E. L. Charnov. 1977. Optimal foraging: a selective review of theory and tests. Quart. Rev. Biol. 52(2):137-154.

10/29            Sephens, D. W. and J. R. Krebs. 1986. Testing foraging models. Chapter 9 In Foraging Theory. Princeton Univ. Press, Princeton, NJ.


11/1           Anderson, D. R. 1975. Optimal exploitation strategies for an animal population in a Markovian environment: A theory and an example. Ecology 56:1281-1297.

11/1            Peterman, R. M. 1977. A simple mechanism that causes collapsing stability regions in exploited salmonid populations. J. Fisheries Research Board of Canada 34:1130-1142.

11/1            Ludwig, D., R. Hilborn, and C. Walters. 1993. Uncertainty, resource exploitation, and conservation: lessons from history. Science 260:17, 36.





11/3             Levins, R. 1970. Extinction. Pages 77-107 in M. Gerstenhaber, ed., Some Mathematical Questions in Biology. American Mathematical Society, Providence, RI.

11/3           Hanski, I. and M. Gilpin. 1991. Metapopulation dynamics: brief history and conceptual domain. Biological Journal of the Linnean Society 42:3-16.

11/3             Harrison, S. 1991. Local extinction in a metapopulation context: an empirical evaluation. Biological Journal of the Linnean Society 42:73-88.

11/5            Hansson, L. 1991. Dispersal and connectivity in metapopulations. Biological Journal of the Linnean Society 42:89-103.

11/5             Fritz, R. S. 1979. Consequences of insular population structure: distribution and extinction of spruce grouse populations. Oecologia 42:57-65.

11/8            Gilpin, M. 1991. The genetic effective size of a metapopulation. Biological Journal of the Linnean Society 42:165-175.

11/8             Pulliam, H. R. 1996. Sources and sinks: empirical evidence and population consequences. pp 45-69 in O. E. Rhodes, Jr., R. K. Chesser, and M.H. Smith, eds. Population Dynamics in Ecological Space and Time. Univ. Chicago Press., Chicago, IL.

11/10           Wiens, J. A. 1996. Wildlife in patchy environments: metapopulations, mosaics, and management. pp53-84 in D. R. McCullough, ed. Metapopulations and Wildlife Conservation. Island Press, Covelo, CA.

11/10              Watkinson, A. R., and W. J. Sutherland.  1995.  Sources, sinks and pseudo-sinks.  Journal of Animal Ecology 64:126-130.

11/10             Boughton, D. A.  1999.  Empirical evidence for complex source-sink dynamics with alternative states in a butterfly metapopulation.  Ecology 80: 2727-2739.

11/10 ______ Parmesan, C.  2006.  Ecological and evolutionary responses to recent climate change.  Ann. Rev. Ecol. Syst. 37:637-669.


11/12              Bunnell, F. L. and D. E. N. Tait. 1981. Population dynamics of bears - implications. pp. 75-88 in Fowler, C. W., and T. D. Smith (eds.). Dynamics of large mammal populations. John Wiley and Sons, New York. 477pp.

11/12            Fowler, C. W. 1981. Comparative population dynamics in large mammals. pp. 437-455 in Fowler, C. W. and T. D. Smith (eds.). Dynamics of large mammal populations. John Wiley and Sons, New York. 477pp.

11/15             Tanner, J. T. 1975. The stability and the intrinsic growth rates of prey and predator populations. Ecology 56:855-867.

11/15            Walters, C. J., M. Stocker, and G. C. Haber. 1981. Simulation and optimization models for a wolf-ungulate system. pp. 317-337 in Fowler, C. W. and T. D. Smith (eds.). Dynamics of large mammal populations. John Wiley and Sons, New York. 477pp.

11/17            Price, P. W. 1984. Alternative paradigms in community ecology. pp.353-383 in Price, P. W., C. N. Slobodchikoff, and W. S. Gaud, (eds.). A new ecology: Novel approaches to interactive systems. 515pp.

11/17            Moss, R. and A. Watson. 2001. Population cycles in birds of the grouse family (Tetraonidae).  Advances in Ecological Research 32:53-111.

11/19             Weins, J. A. 1984. Resource systems, populations, and communities. pp.397-436 in Price, P. W., C. N. Slobodchikoff, and W. S. Gaud, (eds.). A new ecology: Novel approaches to interactive systems. 515pp.


  11/19             Hall, C. A. S. and J. W. Day, Jr. 1977.  Systems and models: Terms and basic principles.  Pp.5-36 in Hall, C. A. S. and J. W. Day, Jr. (eds.). Ecosystem Modeling in Theory and Practice: An Introduction with Case Histories.  John Wiley and Sons, New York, NY.684pp.

11/29             Kingsland, S. E. 1985. Modelling Nature: Episodes in the History of Population Ecology. Chapter 8: The eclipse of history. Pp. 176-205.

11/29            Starfield, A. M. and A. L. Bleloch. 1986. Building Models for Conservation and Wildlife Management.  MacMillan Publ. Co. New York, NY. Pp. 17-36

12/           Walters, C. 1992. Trends in applied ecological modelling. p 117-122 in D. R. McCullough and R. H. Barrett (eds.) Wildlife 20001: Populations.  Elsevier.

12/1             Caswell, H.  1997.Matrix models for population analysis.  Pp. 19-58 in S. Tuljapurkar and H. Caswell (eds.).  Structured-population Models in Marine, Terrestrial, and Freshwater Systems.  Chapman & Hall, New York, NY.  

12/3            Turchin, P.  1998.  Modeling movement.  Chapter 3, Pp. 33-73 in P. Turchin, Quantitative Analysis of Movement: Measuring and Modeling Population Redistribution in Animals and Plants.  Sinauer Assoc., Sunderland, MA.

12/6            Boyce, M. S.  1992. Wolf recovery for Yellowstone National Park: A simulation model. Pp 123-138 in D. R. McCullough and R. H. Barrett (eds.) Wildlife 20001: Populations.  Elsevier.

12/6            Sibly, R. M., and J. Hone.  2003.  Population growth rate and its determinants: an overview.  Pp. 11-40 In R. M Sibly, J. Hone, and T. H. Clutton-brock (eds.).  Wildlife Population Growth Rates.  Cambridge University Press, Cambridge, UK.

12/8           Dennis, B., P. M. Munholland and J. M. Scott.  1991.  Estimation of growth and extinction parameters for endangered species.  Ecology 61:115-143

12/8             Lele, S., M. L. Taper, and S. Gage. 1992. Statistical Analysis of population dynamics in space and time using estimating functions.  Ecology 79:1489-1502

  12/10            Pugesek, B. H.  2003.  Concepts of structural equation modeling in biological research.  Pages 40-57 In B. H. Pugesek, A. Tomer, and A. Von Eye (editors).  Structural Equation Modeling, Cambridge Univ. Press, Cambridge, UK.