Department of Zoology and Physiology

Program in Ecology

Evolutionary Biology

Biodiversity Institute

Contact information:

Office: Berry Center 149
Lab: Biological Sciences 439

Mailing Address:
Department of Zoology & Physiology, 3166
University of Wyoming
1000 E. University Ave.
Laramie, WY 82071

Office Ph: (307) 399-4785
Fax: (307) 766-5625


Curriculum Vitae

Craig W. Benkman
Department of Zoology and Physiology
University of Wyoming
Laramie, WY 82071-3166
(307) 766-2978
FAX (307) 766-5625
E-mail: cbenkman (at)

Academic Jobs

2004-present Professor and Robert B. Berry Distinguished Chair in Ecology, University of Wyoming.
2004 Professor of Biology, New Mexico State University.
1999-2004 Associate Professor of Biology, New Mexico State University.
1993-1999 Assistant Professor of Biology, New Mexico State University.
1993 Lecturer, University of California at Santa Cruz.


1985 Ph.D. Biology; State University of New York at Albany.
Dissertation title: The foraging ecology of crossbills in eastern North America. 185 pp. Advisor: Dr. H. Ronald Pulliam

1981 M.S. Biology; Northern Arizona University, Flagstaff, Arizona.
Thesis title: Co-adaptations of red squirrels and Clark's Nutcrackers with limber pine. 95 pp. Advisor: Dr. Russell P. Balda

1978 B.A. Biology; University of California at Berkeley

Postdoctoral Research Experience

1990-91 NSERC International Fellow at the University of British Columbia. Advisor: Dr. Dolph Schluter

1989-90 NSF NATO Postdoctoral Fellow at the University of British Columbia. Advisor: Dr. Dolph Schluter

1987-89 NSF Postdoctoral Research Fellow at Princeton University. Advisor: Dr. Peter R. Grant


2014 E. O. Wilson Naturalist Award from American Society of Naturalists

2002 Fellow, American Ornithologists’ Union

1991 Elective Member, American Ornithologists' Union

1981 A. Brazier Howell Award, best student paper, Annual Meeting of Cooper Ornithological Society

Post-doctoral Fellows in my Laboratory

2007-2008 Dr. Anna Chalfoun, Postdoctoral Fellow funded by the National Park Service

2003-2005 Dr. Eduardo T. Mezquida, Postdoctoral Fellow funded by Spanish Ministry of Education and Science

2002-2003 Dr. Pim Edelaar, NWO Talent Fellow from The Netherlands

Courses Taught

Contemporary Problems in Biology
Natural History of Life
Avian Ecology/Ornithology
Evolutionary Ecology

Seminars on topics including: Adaptive Radiations, Behavioral Ecology, Coevolution, Darwin and the Galápagos, Granivory, Ornithology, and Speciation.

Participated in a four-day National Science Foundation sponsored “FIRST” workshop on inquiry-based teaching at the Southwest Research Station, Portal, Arizona in 1999.

Grants (partial list)

2005 NSF, Division of Environmental Biology/Ecology, Two-year grant ($106,189)

2003 NSF, Division of Environmental Biology/Ecology, Five-year grant ($354,510)

2002 NSF, Division of Environmental Biology/Population Biology. Five-year grant ($480,000)

2000 Grant from the National Geographic Society ($11,500)

1997 NSF, Division of Environmental Biology/Ecological Studies. Two-year grant ($100,000)

1990 NSERC International Fellowship ($28,000 Canadian)

1989 NATO Postdoctoral Fellowship in Science and Engineering ($26,463)

1986 NSF Post-doctoral Research Fellowship in Environmental Biology ($52,800)


Porter, C. K., and C. W. Benkman. 2017. Assessing the potential contributions of reduced immigrant viability and fecundity to reproductive isolation. American Naturalist 189:580-591.

Benkman, C. W. 2017. Matching habitat choice in nomadic crossbills appears most pronounced when food is most limiting. Evolution 71:778-785.

Benkman, C. W. 2016. The natural history of the South Hills crossbill in relation to its impending extinction. American Naturalist 188:589-601.

Parchman, T. L., C. A. Buerkle, V. Soria-Carrasco, and C. W. Benkman. 2016. Genome divergence and diversification within a geographic mosaic of coevolution. Molecular Ecology 25:5705-5718.

Benkman, C. W., S. Jech, and M. V. Talluto. 2016. From the ground up: Biotic and abiotic features that set the course from genes to ecosystems. Ecology and Evolution 6:7032-7038.

Benkman, C. W., and E. T. Mezquida. 2015. Phenotypic selection exerted by a seed predator is replicated in space, time, and among prey species. American Naturalist 186:682-691.

Talluto, M. V., and C. W. Benkman. 2014. Conflicting selection from fire and seed predation drives fine-scaled phenotypic variation in a widespread North American conifer. Proceedings of the National Academy of Sciences USA 111:9543-9548.

Mezquida, E. T., and C. W. Benkman. 2014. Causes of variation in biotic interaction strength and phenotypic selection along an altitudinal gradient. Evolution 68:1710-1721.

Benkman, C. W. 2013. Biotic interaction strength and the intensity of selection. Ecology Letters 16:1054-1560.

Benkman, C. W., and T. L. Parchman. 2013. When directional selection reduces geographic variation in traits mediating species interactions. Ecology and Evolution 3:961-970.

Talluto, M. V., and C. W. Benkman. 2013. Landscape-scale eco-evolutionary dynamics: selection by seed predators and fire determine a major reproductive strategy. Ecology 94:1307-1316.

Benkman, C. W., J. W. Smith, M. Maier, L. Hansen, and M. V. Talluto. 2013. Consistency and variation in phenotypic selection exerted by a community of seed predators. Evolution 67:157-169.

Smith, J. W., S. M. Sjoberg, M. C. Mueller, and C. W. Benkman. 2012. Assortative flocking in crossbills and implications for ecological speciation. Proceedings of the Royal Society of London Series B 279:4223-4229.

Parchman, T. L., Z. Gompert, J. Mudge, F. D. Schilkey, C. W. Benkman, and C. A. Buerkle. 2012. Genome-wide association genetics of an adaptive trait in lodgepole pine. Molecular Ecology 21:2991-3005.

Benkman, C. W. 2012. White-winged Crossbill (Loxia leucoptera), The Birds of North America Online (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology.

Benkman, C. W., A. M. Siepielski, and J. W. Smith. 2012. Consequences of trait evolution in a multi-species system. Pages 278-292 in Interaction Richness and Complexity: Ecological and Evolutionary Aspects of Trait-Mediated Indirect Interactions. T. Ohgushi, O. Schmitz, and R. Holt, eds. Cambridge University Press.

Benkman, C. W., T. Fetz, and M. V. Talluto. 2012. Variable resource availability when resource replenishment is constant: the coupling of predators and prey. Auk 129:115-123.

Santisteban, L., C. W. Benkman, T. Fetz, and J. W. Smith. 2012. Survival and population size of a resident bird species are declining as temperature increases. Journal of Animal Ecology 81:352–363.

Myczko, Ł., and C. W. Benkman. 2011. Great Spotted Woodpeckers Dendrocopos major exert multiple forms of phenotypic selection on Scots pine Pinus sylvestris. Journal of Avian Biology 42:429-433.

Benkman, C. W., and A. M. Siepielski. 2011. Sources and sinks in the
evolution and persistence of mutualisms. Pages 82-98 in Sources, Sinks, and Sustainability. J. Liu, V. Hull, A. Morzillo, and J. Wiens, eds. Cambridge University Press.

Parchman, T. L., C. W. Benkman, B. Jenkins, and C. A. Buerkle. 2011. Low levels of population genetic structure in lodgepole pine across a geographic mosaic of coevolution. American Journal of Botany 98:669-679.

Benkman C. W., T. L. Parchman, and E. T. Mezquida. 2010. Patterns of coevolution in the adaptive radiation of crossbills. Annals of the New York Academy of Sciences 1206:1-16.

Berry, R. B., C. W. Benkman, A. Muela, Y. Seminario, and M. Curti. 2010. Isolation and decline of an Orange-breasted Falcon population. Condor 112:479-489.

Parchman, T. L., K. S. Geist, J. A. Grahnen, C. W. Benkman, and C. A. Buerkle. 2010. Transcriptome sequencing in an ecologically important tree species: assembly, annotation, and marker discovery. BMC Genomics 11:180.

Mezquida, E. T., and C. W. Benkman. 2010. Habitat area and structure affect the impact of seed predators and the potential for coevolutionary arms races. Ecology 91:802-814.

Benkman, C. W. 2010. Diversifying coevolution between crossbills and conifers. Evolution: Education and Outreach 3:47-53.

Siepielski, A. M., and C. W. Benkman. 2010. Conflicting selection from an antagonist and a mutualist enhances phenotypic variation in a plant. Evolution 64:1120-1128.

Benkman, C. W., and T. L. Parchman. 2009. Coevolution between crossbills and black pine: the importance of competitors, forest area, and resource stability. Journal of Evolutionary Biology 22:942-953.

Garcia, R., A. M. Siepielski, and C. W. Benkman. 2009. Cone and seed trait variation in whitebark pine Pinus albicaulis (Pinaceae) and the potential for phenotypic selection. American Journal of Botany 96:1050-1054.

Snowberg, L. K., and C. W. Benkman. 2009. Mate choice based on a key ecological performance trait. Journal of Evolutionary Biology 22:762-769.

Benkman, C. W., J. W. Smith, P. C. Keenan, T. L. Parchman, and L. Santisteban. 2009. A new species of red crossbill (Fringillidae: Loxia) from Idaho. Condor 111:169-176.

Siepielski, A. M., and C. W. Benkman. 2008. A seed predator drives the evolution of a seed dispersal mutualism. Proceedings of the Royal Society of London Series B 275:1917-1925.

Siepielski, A. M., and C. W. Benkman. 2008. Seed predation and selection exerted by a seed predator influence subalpine tree densities. Ecology 89:2960-2966.

Keenan, P. C., and C. W. Benkman. 2008. Call imitation and call modification in Red Crossbills. Condor 110:93-101.

Benkman, C. W., T. L. Parchman, and A. M. Siepielski. 2008. The geographic mosaic of coevolution and its conservation significance. Pages 225-238 in Conservation biology: evolution in action. S. P. Carroll and C. W. Fox, eds. Oxford University Press.

Parchman, T. L., and C. W. Benkman. 2008. The geographic selection mosaic for ponderosa pine and crossbills: a tale of two squirrels. Evolution 62:348-360.

Benkman, C. W., A. M. Siepielski, and T. L. Parchman. 2008. The local introduction of strongly interacting species and the loss of geographic variation in species and species interactions. Molecular Ecology 17:395-404.

Benkman, C. W. 2007. Red crossbill types in Colorado: their ecology, evolution and distribution. Colorado Birds 41:153-163.

Parchman, T. L., C. W. Benkman, and E. T. Mezquida. 2007. Coevolution between Hispaniolan crossbills and pine: Does more time allow for greater phenotypic escalation at lower latitude? Evolution 61:2142-2153.

Siepielski, A. M., and C. W. Benkman. 2007. Extreme environmental variation sharpens selection that drives the evolution of a mutualism. Proceedings of the Royal Society of London Series B 274:1799-1805.

Snowberg, L. K., and C. W. Benkman. 2007. The role of marker traits in the assortative mating within red crossbills, Loxia curvirostra complex. Journal of Evolutionary Biology 20:1924-1932.

Siepielski, A. M., and C. W. Benkman. 2007. Selection by a pre-dispersal seed predator constrains the evolution of avian seed dispersal in pines. Functional Ecology 21:611-618.

Smith, J. W., and C. W. Benkman. 2007. A coevolutionary arms race causes ecological speciation in crossbills. American Naturalist 169:455-465.

Siepielski, A. M., and C. W. Benkman. 2007. Convergent patterns in the selection mosaic for two North American bird-dispersed pines. Ecological Monographs 77:203-220.

Mezquida, E. T., S. J. Slater, and C. W. Benkman. 2006. Sage-grouse and indirect interactions: potential implications of coyote control on sage-grouse populations. Condor 108:747-759.

Parchman, T. L., C. W. Benkman, and S. C. Britch. 2006. Patterns of genetic variation in the adaptive radiation of New World crossbills (Aves: Loxia). Molecular Ecology 15:1873-1887.

Edelaar, P., and C. W. Benkman. 2006. Replicated population divergence caused by localised coevolution? A test of three hypotheses in the red crossbill-lodgepole pine system. Journal of Evolutionary Biology 19:1651-1659.

Benkman, C. W., J. S. Colquitt, W. R. Gould, T. Fetz, P. C. Keenan, and L. Santisteban. 2005. Can selection by an ectoparasite drive a population of red crossbills from its adaptive peak? Evolution 59:2025-2032.

Siepielski, A. M., and C. W. Benkman. 2005. A role for habitat area in the geographic mosaic of coevolution between red crossbills and lodgepole pine. Journal of Evolutionary Biology 18:1042-1049.

Mezquida, E. T., and C. W. Benkman. 2005. The geographic selection mosaic for squirrels, crossbills and Aleppo pine. Journal of Evolutionary Biology 18:348-357.

Benkman, C. W., and A. M. Siepielski. 2004. A keystone selective agent? Pine squirrels and the frequency of serotiny in lodgepole pine. Ecology 85:2082-2087.

Siepielski, A. M., and C. W. Benkman. 2004. Interactions among moths, crossbills, squirrels and lodgepole pine in a geographic selection mosaic. Evolution 58:95-101.

Benkman, C. W., T. L. Parchman, A. Favis, and A. M. Siepielski. 2003. Reciprocal selection causes a coevolutionary arms race between crossbills and lodgepole pine. American Naturalist 162: 182-194.

Benkman, C. W. 2003. Divergent selection drives the adaptive radiation of crossbills. Evolution 57:1176-1181.

Parchman, T. L., and C. W. Benkman. 2002. Diversifying coevolution between crossbills and black spruce on Newfoundland. Evolution 56:1663-1672.

Hulme, P., and C. W. Benkman. 2002. Granivory. Pages 132-154 in Plant-animal interactions: an evolutionary approach. C. Herrera and O. Pellmyr, eds. Blackwell Scientific Publications, New York.

Benkman, C. W., W. C. Holimon, and J. W. Smith. 2001. The influence of a competitor on the geographic mosaic of coevolution between crossbills and lodgepole pine. Evolution 55:282-294.

Bardwell, E., C. W. Benkman, and W. R. Gould. 2001. Adaptive geographic variation in Western Scrub-jays. Ecology 82:2617-2627.

Benkman, C. W. 1999. The selection mosaic and diversifying coevolution between crossbills and lodgepole pine. American Naturalist 154:S75-S91.

Levey, D. J., and C. W. Benkman. 1999. Fruit-seed disperser interactions: timely insights from a long-term perspective. Trends in Ecology and Evolution 14:41-43.

Coffey, K., C. W. Benkman, and B. G. Milligan. 1999. The adaptive significance of spines on pine cones. Ecology 80:1221-1229

Smith, J. W., C. W. Benkman, and K. Coffey. 1999. The use and mis-use of public information by foraging red crossbills. Behavioral Ecology 10:54-62.

Valone, T. J., and C. W. Benkman. 1999. Public information as a mechanism favoring social aggregation: a brief review of empirical evidence. In: Adams, N. & Slotow, R. (eds), Proceedings of the 22nd International Ornithological Congress. Durban, University of Natal: 1328-1336.

Holimon, W. C., C. W. Benkman, and M. F. Willson. 1998. The importance of mature conifers to red crossbills in southeast Alaska. Forest Ecology and Management 102:167-172.

Benkman, C. W. 1997. Feeding behavior, flock-size dynamics, and variation in sexual selection in crossbills. Auk 114:163-178.

Benkman, C. W., and R. E. Miller. 1996. Morphological evolution in response to fluctuating selection. Evolution 50:2499-2504.

Benkman, C. W. 1996. Are the ratios of bill crossing morphs in crossbills the result of frequency-dependent selection? Evolutionary Ecology 10:119-126.

Benkman, C. W. 1995. The impact of tree squirrels (Tamiasciurus) on limber pine seed dispersal adaptations. Evolution 49:585-592.

Benkman, C. W. 1995. Wind dispersal capacity of pine seeds, with comments on the evolution of different seed dispersal modes in pines. Oikos 73: 221-224.

Hill, G. E., and C. W. Benkman. 1995. Exceptional response to dietary carotenoid supplementation by female Red Crossbills. Wilson Bulletin 107:555-557.

Benkman, C. W. 1994. Comments on the ecology and status of the Hispaniolan crossbill (Loxia leucoptera megaplaga), with recommendations for its conservation. Caribbean Journal of Science 30:250-254.

Benkman, C. W. 1993. Logging, conifers, and the conservation of crossbills. Conservation Biology 7:473-479.

Benkman, C. W. 1993. Adaptation to single resources and the evolution of crossbill (Loxia) diversity. Ecological Monographs 63:305-325.

Benkman, C. W. 1993. The evolution, ecology, and decline of the Red Crossbill of Newfoundland. American Birds 47:225-229.

Benkman, C. W. 1992. White-winged Crossbills. In A. Poole, P. Stettenheim, and F. B. Gill, editors. The Birds of North America. The Academy of Natural Sciences of Philadelphia. Philadelphia, PA.

Benkman, C. W. 1992. A crossbill's twist of fate. Natural History 101(12):38-43.

Benkman, C. W. 1991. Predation, seed size partitioning, and the evolution of body size in seed-eating finches. Evolutionary Ecology 5:118-127.

Benkman, C. W. and A. K. Lindholm. 1991. The advantages and evolution of a morphological novelty. Nature 349:519-520.

Benkman, C. W. 1990. Foraging rates and the timing of crossbill reproduction. Auk 107:376-386.

Benkman, C. W. 1989. On the evolution and ecology of island populations of crossbills. Evolution 43:1324-1330.

Benkman, C. W. 1989. Intake rate maximization and the foraging behaviour of crossbills. Ornis Scandinavica 20:65-68.

Benkman, C. W. 1989. On the foraging behavior of the Kauai Akepa. 'Elepaio 49:29-30.

Benkman, C. W. 1989. Breeding opportunities, foraging rates, and parental care in White-winged Crossbills. Auk 106:483-485.

Benkman, C. W. 1988. Why White-winged Crossbills do not defend feeding territories. Auk 105:370-371.

Benkman, C. W., and H. R. Pulliam. 1988. The comparative feeding rates of North American sparrows and finches. Ecology 69:1195-1199.

Benkman, C. W. 1988. A 3:1 ratio of mandible crossing direction in White-winged Crossbills. Auk 105:578-579.

Benkman, C. W. 1988. Flock size, food dispersion, and the feeding behavior of crossbills. Behavioural Ecology and Sociobiology 23:167-175.

Benkman, C. W. 1988. Seed handling efficiency, bill structure, and the cost of bill specialization for crossbills. Auk 105:715-719.

Benkman, C. W. 1988. On the advantages of crossed mandibles: an experimental approach. Ibis 130:288-293.

Benkman, C. W. 1987. Crossbill foraging behavior, bill structure, and patterns of food profitability. Wilson Bulletin 99:351-368.

Benkman, C. W. 1987. Food profitability and the foraging ecology of crossbills. Ecological Monographs 57:251-267.

Benkman, C. W., R. P. Balda, and C. C. Smith. 1984. Adaptations for seed dispersal and the compromises due to seed predation in limber pine. Ecology 65:632-642.

Book Reviews

Siepielski, A. M., T. L. Parchman, and C. W. Benkman. 2002. Review of Evolutionary Ecology: Concepts and Case Studies. Animal Behaviour 64:323-324.

Benkman, C. W. 1998. Review of Made for Each Other: A symbiosis of birds and pines. Condor 100:190-191.

Invited Seminars in past 10 years

2016 University of Arizona; Symposium on Birds in Coupled Human and Natural Systems at North American Ornithological Conference

2015 University of Nevada at Reno

2014 University of Arkansas; University of Tennessee; Symposium at AOS/COS Meeting on Conservation Ecology of Montane Birds

2013 Symposium on Implications of Pulsed Resources on Forest Birds at the European Ornithological Conference in Norwich, England; Saturday U lecture in Sheridan, WY

2012 Estación Biológica de Doñana; Lund University; Northern Colorado University

2011 Plenary speaker at annual meeting of the American Ornithologists' Union; Casper College

2010 University of Nebraska; Plenary speaker at annual meetings of the Colorado Field Ornithologists and the Association of Field Ornithologists

2009 Pennsylvania State University; St. Louis University; Casper College; University of Washington; Plenary speaker at annual meeting of the Western Field Ornithologists; Plenary Speaker for Symposium on “The Phenotype-Fitness Map Re-visited: Agents of Selection and the Importance of Ecology in Evolutionary Studies”, European Society of Evolutionary Biology, Italy

2008 University of Montana, Iowa State University, University of Colorado at Boulder

2007 Dartmouth College, University of Aberdeen, Leiden University, University of North Carolina at Chapel Hill (invited by graduate students), Museu Ciencies Naturals (Barcelona), Black Hills State University

Professional Societies

American Association for the Advancement of Science
American Ornithologists' Union
American Society of Naturalists
Cooper Ornithological Society
Ecological Society of America
Society for the Study of Evolution

Professional Service

Editor of Natural History Miscellany, The American Naturalist (2009- 2011)
Associate Editor for The American Naturalist (2006-2008)
Associate Editor for Functional Ecology (2006-2008)
Associate Editor for Evolution (2002-2005)
Board of Directors, Cooper Ornithological Society (2010-)
Board of Directors, Wyoming Audubon (2005-)
President, New Mexico Ornithological Society (2000-2004)
Member of Scientific Advisory Committee (2001-2003), Southwestern Research Station, Portal, Arizona (American Museum of Natural History)
Panel member, National Science Foundation

ad hoc Reviewer for (in last five years)

National Science Foundation
National Geographic Society
Natural Environment Research Council
Academy of Sciences of the Czech Republic
National Research Foundation, South Africa
Swiss National Science Foundation

Acta Oecologia, American Naturalist, Ardeola, Auk, Austral Ecology, Behavioral Ecology, Biological Journal of the Linnean Society of London, Biology Letters, BMC Evolutionary Biology, Canadian Journal of Zoology, Condor, Diversity and Distributions, Ecography, Ecology, Ecology Letters, Ecoscience, Ethology, Ecology & Evolution, European Journal of Wildlife Research, Evolution, Evolutionary Ecology, Forest Ecology & Management, Forestry, Ibis, Journal of Avian Biology, Journal of Ecology, Journal of Evolutionary Biology, Journal of Zoology, Molecular Ecology, Nature, New Phytologist, Oecologia, Oikos, Ornitologia Neotropical, Proceedings of the National Academy of Sciences, Proceedings of the Royal Society of London, Science, Trends in Ecology & Evolution, Wilson Journal of Ornithology