Draft:Robert K. Colwell

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Robert Knight Colwell (born 9 October 1943 in Denver, Colorado) is an American evolutionary ecologist, biogeographer, biodiversity scientist, and tropical biologist.

Early life and education

Colwell is the younger of two children of Robert Pulliam Colwell,[i] [ii] a public-school principal and rancher, and Eleanor Knight Colwell,[iii] a concert flutist and amateur naturalist. He attended public schools in Denver, but his interest in biology was kindled by weekends and summers on the family cattle ranch in the mountains, collecting and preparing butterflies and moths under his mother's tutelage.

In 1965, Colwell received a Bachelor of Arts (AB) degree from Harvard University.[iv] In 1966, he worked as curatorial assistant in ethnobotany under Richard Evans Schultes at Harvard. In 1969, he received his Ph.D. in Ecology from the University of Michigan, under the supervision of Lawrence B. Slobodkin. [v] He was a Ford Foundation Fellow at the University of Chicago from 1969 to 1970, with Monte Lloyd[vi] and Richard Levins.

Career

Colwell served on the faculty of the Department of Zoology at the University of California, Berkeley from 1970 to 1989. In 1989, he joined the faculty of the Department of Ecology and Evolutionary Biology[vii] at the University of Connecticut, where he was named a Board of Trustees Distinguished Professor in 2001.[viii] After retirement from teaching in 2014, he became a Distinguished Research Professor at Connecticut. Since 2010, he has been an International Collaborator at the Center for Macroecology, Evolution and Climate at University of Copenhagen.[ix] He also holds positions as Curator Adjoint of Entomology and Zoology[x] at the University of Colorado Museum of Natural History, and Pesquisador Visitante Especial[xi] at the Universidade Federal de Goiás, Brazil.

Other professional work

Colwell served on the Board of Reviewing Editors of Science from 2002 to 2005. He has been an Editor-in-Chief for Reviews and Syntheses[xii] for Ecography since 2014 and has served in various editorial roles for other journals. He was Associate Editor for the Encyclopedia of Biodiversity, 1999-2012.[xiii]

In the 1980s and 1990s, Colwell was active in advising public agencies—including the US Environmental Protection Agency (EPA), National Institutes of Health (NIH), and Department of Agriculture (USDA), as well as the Organization for Economic Co-operation and Development (OECD)—and publishing on the environmental, ecological, and evolutionary risks of genetically engineered (transgenic) organisms.[xiv]

Research

The linking theme of Colwell's work is biological diversity—from coexistence and coevolution of species within contemporary communities to the origin, diversification, and evolutionary adaptation of species on geographical spatial scales and geological timescales. He has contributed to advances in process-based simulation of biogeography, global change biogeography, biodiversity statistics, biodiversity inventory, and biodiversity informatics. In the field, he studies complex interactions and coevolution among tropical species.[xv] Examples below:

• Biogeographical null models. With co-author David Winkler, in 1984, Colwell published the first biogeographical null model to include simulated evolutionary diversification and stochastic dispersal.[xvi] Many subsequent process-based biogeographical models trace their roots to this work.[xvii]
• The mid-domain effect (MDE). In 1994, with co-author George Hurtt, Colwell showed that species diversity inevitably peaks in the middle of any bounded geographical domain (such as elevational transects from sea to mountaintop), if species ranges are shuffled at random.[xviii] Colwell and Lees later called this phenomenon the mid-domain effect.[xix] MDE has proved to be both illuminating and controversial.
• Biodiversity statistics. With co-author Jonathan Coddington, in 1994, Colwell reviewed existing statistical methods for estimating species richness from sampling data.[xx] His frequent collaboration with Anne Chao and Nicholas Gotelli subsequently yielded more than 25 publications on biodiversity statistics.[xxi] As of 2025, Colwell's stand-alone freeware application EstimateS: Statistical estimation of species richness and shared species from samples, had been cited more than 12,000 times.[xxii]
• Biodiversity informatics. Between 1996 and 2015, Colwell was co-director (with John Longino) and of Project ALAS, a 15-year tropical arthropod diversity project in Costa Rica.[xxiii] In his role as data manager for the project, Colwell developed the software Biota: The Biodiversity Database Manager,[xxiv] later published by Sinauer Associates and widely used by similar inventory projects and museums.
• Climate-driven range shifts. In 2008, Colwell and colleagues published the first simulation of the upslope elevational range shifts of tropical species to be expected with climate warming, based on field data for more than 1900 species of plants and animals,[xxv] and predicted lowland biotic attrition in the equatorial tropics[xxvi]
• Hummingbird flower mites. In 1973, Colwell published the first field study on the lives of hummingbird flower mites, a diverse group of species that feed and breed in the flowers of hummingbird-pollinated plants, compete with hummingbirds for nectar, and disperse between plants in the nostrils of the birds.[xxvii] Work on this system has contributed to knowledge of sex ratio evolution,[xxviii] sexual selection for host fidelity,[xxix] host-finding,[xxx] and community ecology.[xxxi]

Honors and awards

In 1998, Colwell was elected President of the American Society of Naturalists[xxxii] and received the President's Award in 2001 for the best paper in The American Naturalist the previous year[xxxiii]. From 1994 to 1995 he was Vice President of the Ecological Society of America,[xxxiv] received the ESA Distinguished Service Award in 1999,[xxxv] and was elected Fellow of the Society in 2012.[xxxvi] He is an elected Fellow of the American Association for the Advancement of Science,[xxxvii] the California Academy of Sciences,[xxxviii] and the Connecticut Academy of Science and Engineering.[xxxix] In 2011, he was inducted into the American Academy of Arts and Sciences. ¹⁵ In 2018, he was named a Clarivate Highly Cited Researcher in the Cross-Field category.[xl]

The mite species Tropicoseius (Rhinoseius) colwelli Hunter 1972,[xli] the katydid species Acantheremus colwelli Naskrecki 1997 (Colwell's horned katydid),[xlii] and the ant species Lenomyrmex colwelli Longino 2006[xliii] were named to honor Colwell.

Marriages and children

Colwell was married to the writer Mary Mackey from 1965 to 1972, to the geneticist Mary-Claire King from 1973 to 1983 (one child), and since 1985 to the tropical forest ecologist Robin Chazdon (two children).

Selected publications

Chao, A., N. J. Gotelli, T. Hsieh, E. L. Sander, K. Ma, R. K. Colwell, and A. M. Ellison. 2014. Rarefaction and extrapolation with Hill numbers: a framework for sampling and estimation in species diversity studies. Ecological Monographs 84:45-67. < https://www.robertkcolwell.org/publications/6842.pdf>

Chao, A., R. L. Chazdon, R. K. Colwell, and T. J. Shen. 2005. A new statistical approach for assessing similarity of species composition with incidence and abundance data. Ecology Letters 8:148-159. https://www.robertkcolwell.org/publications/7597.pdf

Chazdon, R. L., R. K. Colwell, J. S. Denslow, and M. R. Guariguata. 1998. Statistical methods for estimating species richness of woody regeneration in primary and secondary rain forests of NE Costa Rica. Pages 285-309 in F. Dallmeier and J. A. Comiskey, editors. Forest biodiversity research, monitoring and modeling: Conceptual background and Old World case studies. Parthenon Publishing, Paris. https://www.robertkcolwell.org/publications/7536.pdf

Colwell, R. K. 2013. EstimateS: Statistical estimation of species richness and shared species from samples. Version 9 and earlier. User's Guide and application. <https://www.robertkcolwell.org/pages/1407>

Colwell, R. K., A. Chao, N. J. Gotelli, S.-Y. Lin, C. X. Mao, R. L. Chazdon, and J. T. Longino. 2012. Models and estimators linking individual-based and sample-based rarefaction, extrapolation and comparison of assemblages. Journal of Plant Ecology 5:3-21. https://www.robertkcolwell.org/publications/6855.pdf

Colwell, R. K., and D. C. Lees. 2000. The mid-domain effect: geometric constraints on the geography of species richness. Trends in Ecology & Evolution 15:70-76. https://www.robertkcolwell.org/publications/7535.pdf

Colwell, R. K., and D. J. Futuyma. 1971. On the measurement of niche breadth and overlap. Ecology 52:567-576. https://www.robertkcolwell.org/publications/7646.pdf

Colwell, R. K., and D. W. Winkler. 1984. A null model for null models in biogeography. Pages 344-359 in D. R. Strong, Jr., D. Simberloff, L. G. Abele, and A. B. Thistle, editors. Ecological communities: Conceptual issues and the evidence. Princeton University Press, Princeton, N. J. https://www.robertkcolwell.org/publications/15859.pdf

Colwell, R. K., and J. A. Coddington. 1994. Estimating terrestrial biodiversity through extrapolation. Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences 345:101-118. https://www.robertkcolwell.org/publications/7087.pdf

Colwell, R. K., C. X. Mao, and J. Chang. 2004. Interpolating, extrapolating, and comparing incidence‐based species accumulation curves. Ecology 85:2717-2727. https://www.robertkcolwell.org/publications/7532.pdf

Colwell, R. K., G. Brehm, C. L. Cardelús, A. C. Gilman, and J. T. Longino. 2008. Global warming, elevational range shifts, and lowland biotic attrition in the wet tropics. Science 322:258-261. https://www.robertkcolwell.org/publications/7542.pdf

Gotelli, N. J., and R. K. Colwell. 2001. Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness. Ecology Letters 4:379-391. https://www.robertkcolwell.org/publications/7086.pdf

Gotelli, N. J., and R. K. Colwell. 2011. Estimating species richness. Pages 39-54 in A. E. Magurran and B. J. McGill, editors. Frontiers in measuring biodiversity. Oxford University Press, New York. https://www.robertkcolwell.org/publications/7091.pdf

Kremen, C., R. Colwell, T. Erwin, D. Murphy, R. a. Noss, and M. Sanjayan. 1993. Terrestrial arthropod assemblages: their use in conservation planning. Conservation Biology:796-808. https://www.robertkcolwell.org/publications/7533.pdf

Martiny, J. B. H., B. J. Bohannan, J. H. Brown, R. K. Colwell, J. A. Fuhrman, J. L. Green, M. C. Horner-Devine, M. Kane, J. A. Krumins, and C. R. Kuske. 2006. Microbial biogeography: putting microorganisms on the map. Nature Reviews Microbiology 4:102-112. https://www.robertkcolwell.org/publications/7534.pdf

Pecl, G. T., M. B. Araújo, J. D. Bell, J. Blanchard, T. C. Bonebrake, I.-C. Chen, T. D. Clark, R. K. Colwell, F. Danielsen, B. Evengård., et al. 2017. Biodiversity redistribution under climate change: Impacts on ecosystems and human well-being. Science 355:eaai9214. <https://www.robertkcolwell.org/publications/6829.pdf>

Sunday, J. M., A. E. Bates, M. R. Kearney, R. K. Colwell, N. K. Dulvy, J. T. Longino, and R. B. Huey. 2014. Thermal-safety margins and the necessity of thermoregulatory behavior across latitude and elevation. Proceedings of the National Academy of Sciences 111:5610-5615. <https://www.robertkcolwell.org/publications/6847.pdf>

References

[i] Anonymous. 2004. Colwell, Robert Pulliam (Obituary). Denver Post. October 12, 2004, Page C-11.

[ii] Martin, C. 2004. A Colorado Life—East High principal innovative. Denver Post. October 14, 2004, Page C-4.

[iii] Anonymous. 2006. Colwell, Eleanor Knight (Obituary). Denver Post. March 5, 2006, Page 6-06.

[iv] 5 Record of degrees: <https://community.alumni.harvard.edu/person/4994237671>

[vi] Monte Lloyd http://chronicle.uchicago.edu/001116/lloyd-obituary.shtml

[vii]Department of Ecology and Evolutionary Biology at the University of Connecticut https://eeb.uconn.edu

[viii] Board of Trustees Distinguished Professor < https://provost.uconn.edu/awards-honors/the-board-of-trustees-distinguished-professor-award/>

[ix] International Collaborator at CMEC < https://macroecology.ku.dk/people/>

[x] University of Colorado Museum of Natural History https://www.colorado.edu/cumuseum/dr-robert-colwell

[xi] Pesquisador Visitante Especial at the Universidade Federal de Goiás <https://ufg.br/n/112080-ufg-tem-dois-professores-entre-os-mais-influentes-do-mundo>

[xii] Ecography Editor < https://www.ecography.org/about-journal/editorial-board>

[xiii] Encyclopedia of Biodiversity < https://shop.elsevier.com/books/encyclopedia-of-biodiversity/levin/978-0-08-091782-5>

[xiv] Colwell, R. K. 1989. Natural and unnatural history: Biological diversity and genetic engineering. Pages 1-40 in W. R. Shea and B. Sitter, editors. Scientists and their responsibility. Watson International Publishing, Canton, MA.

Tiedje, J. M., R. K. Colwell, Y. L. Grossman, R. E. Hodson, R. E. Lenski, R. N. Mack, and P. J. Regal. 1989. The planned introduction of genetically engineered organisms: ecological considerations and recommendations. Ecology 70:298-315.

[xv] American Academy of Arts and Sciences biography.  https://www.amacad.org/person/robert-k-colwell

[xvi] Colwell, R. K., and D. W. Winkler. 1984. A null model for null models in biogeography. Pages 344-359 in D. R. Strong, Jr., D. Simberloff, L. G. Abele, and A. B. Thistle, editors. Ecological communities: Conceptual issues and the evidence. Princeton University Press, Princeton, N. J.

[xvii] Gotelli, N. J. 2014. Commentary on "A Null Model for Null Models in Biogeography (1984), R. K. Colwell and D. W. Winkler". Pages 760-761 in F. A. Smith, J. L. Gittleman, and J. H. Brown, editors. Foundations of Macroecology. University of Chicago Press.

[xviii] Colwell, R. K., and G. C. Hurtt. 1994. Nonbiological gradients in species richness and a spurious Rapoport effect. American Naturalist 144:570-595.

[xix] Colwell, R. K., and D. C. Lees. 2000. The mid-domain effect: geometric constraints on the geography of species richness. Trends in Ecology and Evolution 15:70-76.

[xx] Colwell, R. K., and D. C. Lees. 2000. The mid-domain effect: geometric constraints on the geography of species richness. Trends in Ecology and Evolution 15:70-76.

[xxi] Google Scholar https://scholar.google.com/citations?user=i2OF2g0AAAAJ&hl=en&oi=ao

[xxii] Colwell, R. K. 2012 EstimateS: statistical estimation of species richness and shared species from samples. Version 9. User's Guide and application published at: http://purl.oclc.org/estimates.

Colwell, R. K., and J. E. Elsensohn. 2014. EstimateS turns 20: statistical estimation of species richness and shared species from samples, with non‐parametric extrapolation. Ecography 37:609-613.

[xxiii] Longino, J. T., and R. K. Colwell. 1997. Biodiversity assessment using structured inventory: Capturing the ant fauna of a lowland tropical rainforest. Ecological Applications 7:1263-1277. <https://ants.biology.utah.edu/ALAS/>

[xxiv] Colwell, R. K. 2004. Biota 2: The biodiversity database manager. (Manual and database management software.). Sinauer Associates, Sunderland, MA.

[xxv] Colwell, R. K., G. Brehm, C. Cardelús, A. C. Gilman, and J. T. Longino. 2008. Global warming, elevational range shifts, and lowland biotic attrition in the wet tropics. Science 322:258-261.

[xxvi] Colwell, R. K., and K. J. Feeley. 2024. Still little evidence of poleward range shifts in the tropics, but lowland biotic attrition may be underway. Biotropica n/a:e13358

[xxvii] Colwell, R. K. 1973. Competition and coexistence in a simple tropical community. The American Naturalist 107:737-760.

[xxviii] Wilson, D. S., and R. K. Colwell. 1981. Evolution of sex ratio in structured demes. Evolution 35:882-897.

[xxix] Colwell, R. K. 1986. Population structure and sexual selection for host fidelity in the speciation of hummingbird flower mites. Evolutionary processes and theory. Karlin, S, and E. Nevo, Eds.:475-495.

Colwell, R. K. 1986. Community biology and sexual selection: Lessons from hummingbird flower mites. Pages 406-424 in J. Diamond and T. J. Case, editors. Ecological communities. Harper and Row, New York.

[xxx] Heyneman, A. J., R. K. Colwell, S. Naeem, D. S. Dobkin, and B. Hallet. 1991. Host plant discrimination:  Experiments with hummingbird flower mites. Pages 455-485 in P. W. Price, T. M. Lewinsohn, G. W. Fernandes, and W. W. Benson, editors. Plant-animal interactions:  Evolutionary ecology in tropical and temperate regions. John Wiley and Sons, New York.

[xxxi] Colwell, R. K., and S. Naeem. 1994. Life history patterns of hummingbird flower mites in relation to host phenology and morphology. Pages 23-44 in M. A. Houck, editor. Mites:  Ecological and evolutionary analyses of life history patterns. Chapman and Hall, New York.

[xxxii] Past Presidents of the American Society of Naturalists. https://www.amnat.org/about/history/past-ec.html

[xxxiii] ASN Presidential Award. https://www.amnat.org/awards.html#President

[xxxiv] Vice President of the Ecological Society of America. https://esa.org/history/2014/02/vice-presidents/

[xxxv] ESA Distinguished Service Award. https://esa.org/wp-content/uploads/sites/94/2022/02/service1999.pdf

[xxxvi] ESA Fellow. https://esa.org/about/awards/fellows-program/esa-fellows/

[xxxvii] AAAS Fellow. https://www.aaas.org/fellows/historic

[xxxviii] California Academy of Science Fellow. https://www.calacademy.org/scientists/academy-fellows#CE

[xxxix] Connecticut Academy of Science and Engineering Fellow. https://casemembers.wildapricot.org/Sys/PublicProfile/5990064/1086086

[xl] Clarivate Highly-Cited. https://clarivate.com/highly-cited-researchers/

[xli] Hunter, P. E. (1972) New Rhinoseius species (Mesostigmata: Ascidae) from Costa Rican hummingbirds. Journal of the Georgia Entomological Society, 7, 26 - 35.

[xlii] Naskrecki, P. 1997. A revision of the neotropical genus Acantheremus Karny, 1907 (Orthoptera: Tettigoniidae: Copiphorinae). Transactions of the American Entomological Society:137-161.

[xliii] Longino, J.T. 2006. New species and nomenclatural changes for the Costa Rican ant fauna (Hymenoptera: Formicidae). Myrmecologische Nachrichten. 8:131143. https://www.antwiki.org/wiki/Lenomyrmex_colwelli

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