Institute of Arctic Biology
311 Irving, University of Alaska
Fairbanks, Alaska 99775-7000 USA

Research Interests

           My group works on the evolution and ecology of plant-insect interactions, with special focus on pollination ecology and insect herbivory. We use a variety of approaches including field and lab experiments, chemical analysis of plant secretions, and comparative ecology. In particular, we have attempted to gain a long-term (macroevolutionary) perspective on plant-insect (co-)evolution by conducting phylogeny-based studies of the evolution of traits and of pollination relationships. This involves integrating ecological field studies with phylogenies estimated from morphological and molecular data. We also work on quantitative genetics and development of floral traits involved in attracting, rewarding, and manipulating pollinators.

          Our arctic/subarctic research program has focused primarily on the effects of local variation in microclimate on the composition and function of plant and insect communities. For example, the flora, insect fauna, and pollination ecology is often completely different on nearby hill slopes that face north vs. south. This research includes understanding insect thermal ecology, weather effects on pollination, and latitudinal patterns in beta-diversity (species turnover across microhabitats). We have related interests in Beringian paleoenvironments, and the distribution, ecology, and phylogeography of Beringian plants and insects.

          Another research program, in collaboration with Bruce Baldwin (UC Berkeley), Christa Mulder (UAF), and Susan Kalisz (University of Pittsburgh), examines the evolution of plant mating systems and such floral traits as corolla size, pollen/ovule ratio, and timing of self pollination, in a molecular phylogenetic context. We are employing the sister genera Collinsia and Tonella (Scrophulariaceae) as our study system. Preliminary data suggest that mating-system evolution in this group has been driven at least partly by pollinator dependability, as influenced by the weather during the flowering season.

          Our research on the above topics has been conducted in Alaska, Yukon Territory, the Russian Far East, California, Norway, and Svalbard. Research on plant-insect interactions has also taken us to Madagascar and various parts of Africa, South America, and Meso-America. Our work in the tropics has recently expanded from studies of plants and insects to an ongoing conservation-biology project examining the biological consequences and social causes of tropical-forest fragmentation.

Refereed Publications

• Armbruster, W. S., and D. G. Rogers.  2004.  Does pollen competition reduce the cost of inbreeding?  American J. Botany (in press).
• Fenster, C. B., W. S. Armbruster, P. Wilson, M. R. Dudash, and J. T. Thomson.  2004.  Pollination syndromes and floral specialization. Annual Review of Ecology and Systematics 35: 375-403. 
• Pélabon, C., M. L. Carlson, T. F. Hansen, and W. S. Armbruster. 2004.  Variational and genetic properties of developmental stability in Dalechampia scandens.  Evolution 58: 504-514.
• Pélabon C., T. F. Hansen, M. L. Carlson, N. G. Yoccoz, and W. S. Armbruster.  2004.  Consequences of inter-population crosses on developmental stability and canalization of floral traits in Dalechampia scandens (Euphorbiaceae).  J. Evolutionary Biology 17: 19-32.
• Wilson, P. M. C. Castellanos, J. N. Hogue. J. D. Thomson, and W. S. Armbruster.  2004.  A multivariate search for pollination syndromes among penstemons.  Oikos 104: 345-361.
• Hansen, T. F., C. Pélabon, W. S. Armbruster, and M. L. Carlson. 2003.  Evolvability and genetic constraint in Dalechampia blossoms: Components of variance and measures of evolvability.  J. Evolutionary Biology 16: 754-766.
• Hansen, T. F., W. S. Armbruster, M. L. Carlson, and C. Pélabon. 2003. Evolvability and genetic constraint in Dalechampia blossoms: Genetic correlations and conditional evolvability. J. Experimental Zoology. Molecular and Developmental Evolution 296B: 23-39.
• Baldwin, B. G., and W. S. Armbruster.  2002.  Collinsia antonina Hardham is evolutionarily distinct from C. parryi A. Gray (Scrophulariaceae, sensu lato).  Madroño 49: 295-297.
• Armbruster, W. S., E. M. Debevec, and M. F. Willson. 2002. The evolution of syncarpy in angiosperms: theoretical and phylogenetic analyses of the effects of carpel fusion on offspring quantity and quality. Journal of Evolutionary Biology 15: 657-672.
• Armbruster, W. S. 2002. Can indirect selection and genetic context contribute to trait diversification? A transition-probability study of blossom-color evolution in two genera. Journal of Evolutionary Biology 15: 468-486.
• Armbruster, W. S., C. P. H. Mulder, B. G. Baldwin, S. Kalisz, B. Wessa, and H. Nute. 2002. Comparative analysis of late floral development and mating-system evolution in tribe Collinsieae (Scrophulariaceae, s.l.). Amer. J. Botany 89: 37-49.
• Armbruster, W. S. 2001. Evolution of floral form: Electrostatic forces, pollination, and adaptive compromise. New Phytologist 152: 181-183.
• Hansen, T., W. S. Armbruster, and L. Antonsen. 2000. Comparative analysis of character displacement and spatial adaptations as illustrated by the evolution of Dalechampia blossoms. American Naturalist 156: S17-S34.
• Armbruster, W. S., C. B. Fenster, and M. R. Dudash. 2000. Pollination "principles" revisited: Specialization, pollination syndromes, and the evolution of flowers. Det Norske Videnskapsacademi I. Matematisk Naturvidenskapelige Klasse Skrifter, Ny Serie 39: 139-148.
• Bishop, J. A., and W. S. Armbruster. 1999. The relationship between body size and thermoregulatory ability in bees of interior Alaska. Functional Ecology 13: 711-724.
• Armbruster, W. S., V. S. Di Stilio, J. D. Tuxill, T. C. Flores, and J. L. Velasquez Runk. 1999. Covariance and decoupling of floral and vegetative traits in nine neotropical plants: A reevaluation of Berg's correlation-pleiades concept. American J. Botany 86: 39-55.
• Armbruster, W. S., and B. G. Baldwin. 1998. Switch from specialized to generalized pollination. Nature 394: 632.
• Armbruster, W. S. 1997. Exaptations link the evolution of plant-herbivore and plant-pollinator interactions: a phylogenetic inquiry. Ecology 78:1661-1674.
• Strauss, S. Y., and W. S. Armbruster. 1997. Linking herbivory and pollination - new perspectives on plant and animal ecology and evolution.. Ecology 78:1617-1618.
• Armbruster, W. S. , J.J. Howard, T. P. Clausen, E. Debevec, J. Loquvam, M Matsuki, B. Cerendolo, and F. Andel. 1997. Do biochemical exaptations link evolution of plant defense and pollination systems? -- Historical hypotheses and experimental test with Dalechampia vines. American Naturalist 149:461-484.
• Gillespie, L. and W. S. Armbruster. 1997. A contribution to the Guianan flora: Dalechampia, Haematostemon, Omphalia, Pera, Plukenetia, and Tragia (Euphorbiaceae) with notes on subfamily Acalyphoideae. Smithsonian Contributions in Botany 86:1-48.
• Armbruster, W.S., and K.E. Schwaegerle. 1996. Causes of covariation of phenotypic traits among populations. J. Evolutionary Biology 9:261-276.
• Armbruster, W.S. 1996 A cladistic analysis and revision of Dalechampia sections Rhopalostylis and Brevicolumnae. Syst. Bot. 21:209-235.
• Armbruster, W.S., P. Martin, J. Kidd, R. Stafford, and D.G. Rogers. 1995. Reproductive significance of indirect pollen-tube growth in Dalechampia (Euphorbiaceae). Am. J. Bot. 82:51-56.
• Armbruster, W.S., M.E. Edwards, and E.M. Debevec. 1994. Character displacement generates assemblage structure of Western Australian triggerplants (Stylidium). Ecology 75:315-329.
• Armbruster, W.S. 1994. Early evolution of Dalechampia (Euphorbiaceae): Insights from phylogeny, biogeography, and comparative ecology. Annals of Missouri Botanical Garden 81:302-306.
• Armbruster, W.S., and E.E. Berg. 1994. Thermal ecology of male euglossine bees (Apidae: Euglossini) in a tropical wet forest: fragrance foraging in relation to operative temperatures. Biotropica 26:50-60.
• Lloyd, A.H., W.S. Armbruster, and M.E. Edwards. 1994. Ecology of a steppe-tundra gradient in interior Alaska. J. Veg. Sci. 5:897-912.
• Armbruster, W.S. 1993. Evolution of plant pollination systems: hypotheses and tests with the neotropical vine Dalechampia. Evolution 47:1480-1505.
• Armbruster, W.S. 1993. Within-habitat heterogeneity in baiting samples of male euglossine bees: possible causes and implications. Biotropica 24:122-128.
• Armbruster, W.S., M.E. Edwards, J.F. Hines, R.L.A. Mahunnah, and P. Munyenyembe. 1993. Evolution and pollination of Madagascan and African Dalechampia (Euphorbiaceae). National Geographic Research and Exploration 9:430-444.
• Armbruster, W.S. 1992. Phylogeny and the evolution of plant-animal interactions. BioScience 42:12-20.
• Armbruster, W.S., and K. E. Steiner. 1992. Pollination ecology of four Dalechampia species (Euphorbiaceae) in northern Natal, South Africa. American J. Botany 79:306-313.
• Armbruster, W.S., A.L. Herzig, and T.P. Clausen. 1992. Pollination of two sympatric species of Dalechampia (Euphorbiaceae) in Suriname by male euglossine bees. Amer. J. Bot. 79:1374-1381.
• Armbruster, W.S., and A. D. McGuire. 1991. Experimental assessment of reproductive interactions between sympatric Aster and Erigeron (Asteraceae) in interior Alaska. Amer. J. Bot. 78:1449-1457.
• McGuire, A.D., and W.S. Armbruster. 1991. An experimental test for reproductive interactions between two sequentially blooming Saxifraga species. Amer. J. Bot. 78: 214-219.
• Webster, G.L. and W.S. Armbruster. 1991. A synopsis of the neotropical species of Dalechampia. Bot. J. Linnean Soc. London 105:137-177.
• Wesser, S.D., and W.S. Armbruster. 1991. Controls over species distribution across a forest-steppe transition: a causal model and experimental test. Ecological Monographs 61:323-342.
• Edwards, M.E., J. Dawe, and W.S. Armbruster. 1991. Pollen size of Betula in northern Alaska and interpretation of late Quaternary vegetation records. Can. J. Botany 69:1666-1672.
• Armbruster, W.S. 1991. Multilevel analyses of morphometric data from natural plant populations: insights into ontogenetic, genetic, and selective correlations in Dalechampia scandens. Evolution 45:1229-1244.
• Armbruster, W.S. 1990. Estimating and testing the shapes of adaptive surfaces: the morphology and pollination of Dalechampia blossoms. Am. Nat. 135:14-31.
• Armbruster, W.S., and K.D. McCormick. 1990. Diel foraging patterns of male euglossine bees: ecological causes and evolutionary response by plants. Biotropica 22:160-171.
• Armbruster, W.S. 1989. Three new species of Dalechampia from Venezuelan Guayana . Brittonia 41:44-52.
• Keller, C.S., and W.S. Armbruster. 1989. Pollination of Hyptis capitata in Panama by eumenid wasps. Biotropica 21:190-192.
• Edwards, M.E., and W.S. Armbruster. 1989. A steppe-tundra transition on Kathul Mountain, Alaska. Arctic and Alpine Research 21:296-304.
• Armbruster, W.S., C.S. Keller, and M. Matsuki, and T.P. Clausen. 1989. Pollination of Dalechampia magnoliifolia (Euphorbiaceae) by male euglossine bees (Apidae: Euglossini). Amer. J. Bot. 76:1279-1285.
• Armbruster, W.S., and D.A. Guinn. 1989. The solitary bee fauna of interior and arctic Alaska: flower associations, habitat use, and phenology. J. Kansas Entomol. Soc. 62:468-483.
• Armbruster, W.S. 1988. A new species, section, and synposis of Dalechampia (Euphorbiaceae) from Costa Rica. Syst. Bot. 13:303-312.
• Armbruster, W.S. 1988. Multilevel comparative analysis of morphology, function, and evolution of Dalechampia blossoms. Ecology 69: 1746-1761.
• Armbruster, W.S., and W. Mziray. 1987. Pollination and herbivore ecology of an African Dalechampia (Euphorbiaceae): comparisons with New World species. Biotropica 19:64-73.
• Armbruster, W.S. 1986. Reproductive interactions between sympatric Dalechampia species: are natural assemblages "random" or organized? Ecology 67:522-533.
• Whitten, W.M., N.H. Williams, W.S. Armbruster, M.A. Battiste, L. Strekowski, and N. Lindquist. 1986. Carvone oxide: an example of convergent evolution in euglossine-pollinated plants. Syst. Bot. 11:222-228.
• Armbruster, W.S. 1985. Patterns of character divergence and the evolution of reproductive ecotypes of Dalechampia scandens (Euphorbiaceae). Evolution 39:733-752.
• Armbruster, W.S. and A.L. Herzig. 1984. Partitioning and sharing of pollinators by four sympatric species of Dalechampia (Euphorbiaceae) in Panama. Ann. Missouri Bot. Gard. 71:1-16.
• Armbruster, W.S. 1984. Two new species of Dalechampia (Euphorbiaceae) from Mesoamerica. Syst. Bot. 9:272-278.
• Armbruster, W.S. 1984. The role of resin in angiosperm pollination: ecological and chemical considerations. Amer. J. Bot. 71:1149-1160.
• Webster, G.L., and W.S. Armbruster. 1982. An unusual new species of Dalechampia (Euphorbiaceae) from Surinam. Syst. Bot. 7:484-488.
• Armbruster, W.S. 1982. Seed production and dispersal in Dalechampia (Euphorbiaceae): divergent patterns and ecological consequences. Amer. J. Bot. 69:1429-1440.
• Armbruster, W.S., and G.L. Webster. 1982. Divergent pollination systems in sympatric species of South American Dalechampia (Euphorbiaceae). Amer. Midl. Nat. 108:325-337.
• Armbruster, W.S., and G.L. Webster. 1981. Sistemas de polinizacao de duas especies simpatricas de Dalechampia (Euphorbiaceae) no Amazonas, Brasil. Acta Amazonica 11:13-17.
• Armbruster, W.S., and G.L. Webster. 1979. Pollination of two species of Dalechampia (Euphorbiaceae) in Mexico by euglossine bees. Biotropica 11:278-283.
• Webster, G.L., and W.S. Armbruster. 1979. A new species of euglossine-pollinated Dalechampia (Euphorbiaceae) from Mexico. Brittonia 31:352-357.

Chapters in Books

• Armbruster, W.S., D. Rae, and M. E. Edwards.  2004.  Topographic complexity and biotic response to climate change: Variance is as important as the mean.  In, J. B. Ørbæk (ed.), Arctic-Alpine Ecosystems and People in a Changing Environment. Springer Verlag (in press).
• Armbruster, W. S.  2004.  Evolutionary and ecological perspectives on specialization: from the arctic to the tropics.  In, N. Waser and J. Ollerton (eds.), Specialization and Generalization in Plant-Pollinator Interactions, University of Chicago Press, Chicago (in press). 
• Armbruster, W. S., C. Pélabon, T. F. Hansen, and C. P. H. Mulder.  2004.  Floral integration and modularity: Distinguishing complex adaptations from genetic constraints.  Pp. 23-49 in, M. Pigliucci and K. A. Preston (eds.), The Evolutionary Biology of Complex Phenotypes, Oxford University Press, Oxford, UK. 
• Ratsirarson, J., M. Edwards, G. Olsson, S. Armbruster, I. Pareliussen, B. Reau.  2003.  Forest restoration and biodiversity conservation in the central highlands: the case of the Réserve Spéciale d'Ambohitantely.  Pp. 1476-1480 in, S.J. Goodman and J. Benstead (eds.), The Natural History of Madagascar, University of Chicago Press, Chicago, USA.
• Armbruster, W. S. and B. G. Baldwin.  2003.  Pollination and evolution of euphorb vines in Madagascar.  Pp. 391-393 in, S. Goodman and J. Benstead (eds.), The Natural History of Madagascar.  University of Chicago Press, Chicago, USA.
• Armbruster, W. S.  2002.  Dalechampia.  In S.A. Mori, G. Cremers, C. A. Gracie, J.-J. de Granville, S. V. Heald, M. Hoff, and J. D. Mitchell (eds.), Guide to the Vascular Plants of Central French Guiana. Part 2. Dicotyledons.  Mem. New York Bot. Gard. 76 (2).
• Edwards, M., S. Armbruster, I. Kalland, J. Klein, G. Olsson, I. Pareliussen, J. Ratsirarson, B. Reau,.  2002.  Biodiversity conservation and rural society in Highland Madagascar - worlds apart?  In, Interdisciplinary Research at the Norwegian University of Science and Technology. Universitets forlag, Oslo.
• Armbruster, W. S., and G. Rosenqvist.  2001.  Mutualisms between free-living species.  In, Encyclopedia of Life Sciences.  Macmillan Reference Ltd., London (web and hardcopy editions).
• Armbruster, W. S. 2002. Evolution and pollination of euphorb vines in Madagascar. In S. Goodman and J. Benstead, eds., Natural History of Madagascar. Univ. Chicago Press, Chicago (in press).
• Armbruster, W. S., and G. Rosenqvist. 2002. Mutualisms between free-living species. In, Encyclopedia of Life Sciences. Macmillan Reference Ltd, London (in press).
• Armbruster, W. S. 2002. Dalechampia, In Flora of Central French Guiana. New York Botanical Gardens (in press).
• Armbruster, W.S. 1999. Dalechampia. Pp. 133-139 in, J. A. Steyermark, P E. Berry, K. Yatskievych, B. K. Holst (eds.), Flora of the Venezuelan Guayana. Missouri Botanical Garden Press.
• Armbruster, W. S. 1996. Exaptation, adaptation, and homoplasy: Evolution of ecological traits in Dalechampia. Pp. 227-243 in, M. J. Sanderson and L. Hufford (eds.), Homoplasy: The recurrence of similarity in evolution. Academic Press, New York.
• Armbruster, W.S. 1996. Evolution of floral morphology and function: an integrated approach to adaptation, constraint, and compromise in Dalechampia (Euphorbiaceae). Pp. 241-272 in, D. Lloyd and S. Barrett (eds), Floral biology. Chapman & Hall, New York.
• Armbruster, W. S. 1995. Plant-animal interactions. Pp. 93-102 in Encyclopedia of Environmental Biology, Academic Press, New York.
• Wesser, S. D., W. S. Armbruster, E. M. Debevec, and M. E. Edwards. 1994. Subarctic steppe distribution through space and time: a microclimate modeling approach. Pp 63-69, in R. H. Meehan, V. Sergienka, and G. Weller (eds.), Bridges of science between North America and the Russian Far East. Arctic Division, AAAS, Fairbanks, AK.
• Armbruster, W.S. 1993. Evolution and ecology of animal-pollinated flowers. Pp. 166-167 in McGraw-Hill Yearbook of Science and Technology, 1994. McGraw-Hill, New York.
• Armbruster, W.S. 1983. Dalechampia scandens, in D.H. Janzen (ed.), Costa Rican Natural History, p. 230-233, Univ. of Chicago Press, Chicago, IL.

Manuscripts Submitted and In Preparation

• Hansen, T. F., C. Pelabon, W. S. Armbruster, and M. Carlson. Evolvability and genetic constraint in Dalechampia blossoms: I. Components of variance and measures of evolvability. Evolution (submitted).
• Hansen, T. F., W. S. Armbruster, C. Pelabon, and M. Carlson. Evolvability and genetic constraint in Dalechampia blossoms: II. Genetic correlations and conditional evolvability. Evolution (submitted).
• Armbruster, W. S. and H. R. Nute. Covariance and decoupling of floral and vegetative traits in response to environmental variation imposed on Dalechampia scandens (Euphorbiaceae). (in prep.)

Graduate Students

• Anthony D. McGuire. Ph.D. 1989. Dissertation title: The organization of flowering times of the insect-pollinated flora of south-facing bluffs in interior Alaska: evaluation of the reproductive interference hypothesis.
• Sara D. Wesser. M.S. 1989. Thesis title: Controls over species distribution across a forest-steppe transition in interior Alaska: a correlative model and experimental tests.
• David A. Guinn. M.S. 1991. Thesis title: Distribution and flower use of anthophilious insects interior Alaska, with special reference to solitary bees (Hymenoptera: Apoidea).
• Jon A. Bishop. M.S. 1992. Thesis title: The relationship between size and thermoregulatory capability in social and solitary bees of interior Alaska.
• Andrea Lloyd. M.S. 1993. Thesis title: Ecology of a steppe-tundra ecotone in interior Alaska: modern dynamics and paleoecological interpretations.
• Carl Roland. M.S. 1996. Thesis title: Biogeography and ecology of subarctic steppe in Alaska and the Russian Far East.
• Denise Gobeille Rogers. M.S. 1997. Thesis title: Dynamics of pollen competition in Dalechampia.
• Matthew Carlson. Ph.D. student. Dissertation topic: Mating-system evolution and inbreeding depression in Mimulus (Scrophulariaceae).
• Liv Antonsen. M.S. student. Thesis topic: Phenotypic selection analysis of defense and attraction functions of Dalechampia floral bracts.
• Trond Einar Brobakk. M.S. 2000. Thesis title: Thermal ecology of flower-visiting insects in Norway: activity levels on flowers in relation to operative temperature.
• Ane Moe. M.S. 2000. Thesis title: Patterns of insect and plant distribution in relation to radiation loads and microclimate on Spitsbergen.
• Monica Lundberg. M.S. 2000. Thesis title: Effects of tropical forest fragmentation on tree regeneration in upland Madagascar.
• Suzanne Hansen. M.S. student. Thesis topic: Effects of slope and aspect on forest loss to fire: A GIS study of deforestation in upland Madagascar.
• David Rae. Doctoral student. Dissertation topic: Patterns of beta diversity (turnover) of plant and insect species in response to radiation loads and microclimate at three latitudes.
• Ingar Pareliussen. Doctor student. Dissertation topic: Environmental limits on tree establishment in anthropogenic grasslands of upland Madagascar: implications for reforestation.

Major Research Grants

• US National Science Foundation (NSF) (2004; collaborative with S. Kalisz (Pittsburgh) and B. G. Baldwin (Berkeley)).  Unraveling the dynamics of mating-system evolution in the tribe Collinsieae (Scrophulariaceae).
• Norwegian Research Council (NFR) Research Grant. (2001). Evolution of mating systems in tribe Collinsieae (Scrophulariaceae).
• NFR Equipment Grant. (1999; Arne Strøm and Charles Fenster , co-PI's): Automated DNA sequencing facility for population genetics and biotechnology
• NFR Research Grant (1998; co-PI with Thomas Hansen): Molecular genetic analysis of relationships among Dalechampia scandens population and a comparative test of the pollination character displacement hypothesis.
• NFR Research Grant (1997): Latitudinal patterns in beta diversity (species turnover) of insect and plant species in response to variation in topography, radiation load, and microclimate.
• NFR Research Grant (1997; collaboration with T. Hansen): Using comparative methods to study adaptation and genetic constraint in the evolution of floral morphology and pollination in Dalechampia scandens populations.
• NFR Research Grant (1997; collaboration with M. E. Edwards and G. A. Olsson). Forest fragmentation and biodiversity conservation in upland Madagascar.
• US National Science Foundation (NSF) Research Grant (1997; WSA, PI; J. Thomson, A. Wolfe, and P. Wilson collaborating investigators): Collaborative research: Evolution of floral function: A test of pollen presentation theory.
• NFR Research Grant (1996): Thermal ecology of flower-visiting insects: bee biogeography and pollination.
• NSF Research Grant (1994; WSA, PI; B. Baldwin collaborating investigator): Collaborative research: Phylogenetic analysis of Old World Dalechampia using DNA and morphological data.
• National Geographic Field Research Grant (1992; WSA, PI; M.E. Edwards, co-PI): Ecology and biogeography of azonal steppe in Beringia: comparisons of Asian and North American Elements.
• NSF Research Grant (1990; WSA, PI; T.P. Clausen, co-PI): Chemical ecology of resin secretion in Dalechampia (Euphorbiaceae): Radical evolutionary shifts in the functions of triterpenes?.
• NSF Research Grant (1990; WSA, PI): Systematics and phylogeny of Madagascan and African Dalechampia (Euphorbiaceae).
• National Geographic Field Research Grant (1990; WSA, PI): Systematics and ecology of Madagascan Dalechampia: insights into evolutionary and biogeographic history.
• NSF Biological Centers Program Grant (1988; WSA, PI; F.S. Chapin, J.P. Bryant, J.F. Fox, D.F. Murray, and K.E. Schwaegerle, co-PI's): Plant Growth Facilities for Experimental Botany at the University of Alaska Fairbanks.
• NSF Research Grant BSR-8516864 (1985; WSA, PI): Reproductive Interactions and Plant Community Organization on South-Facing Bluffs in Interior Alaska.
• NSF Research Grant BSR-8509031 (1985; WSA, PI; T.P. Clausen, co-PI): An Analysis of Phylogenetic Relationships within Dalechampia and a Provisional Delimitation of Sections.

Education

• B.A. in Biology (1972); College of Creative Studies, University of California, Santa Barbara; 1969-1972.
• M.S. in Evolutionary Ecology (1977); Ecology Graduate Group, University of California, Davis; 1975-1977.
• Ph.D. in Evolutionary Ecology (1981); Ecology Graduate Group, University of California, Davis; 1977-1980.

Professional Experience

• Professor of Biodiversity, Department of Botany, Norwegian University of Science and Technology, N-7491 Trondheim, Norway (1996 - present).
• Chairman, Department of Botany, Norwegian University of Science and Technology, N-7491 Trondheim, Norway (1999 - 2000)
• Program Director, Ecology Program, National Science Foundation, Arlington, VA (1993 - 1994).
• Deputy Director, Institute of Arctic Biology, University of Alaska Fairbanks (1992 - 1993).
• Professor of Biology, Department of Biology & Wildlife and Institute of Arctic Biology, University of Alaska Fairbanks, Fairbanks AK 99775-0180 (1992 - 1997).
• Associate Professor of Botany, Department of Biology & Wildlife and Institute of Arctic Biology, University of Alaska Fairbanks (1987 - 1992).
• Assistant Professor of Botany, Department of Biology, Fisheries & Wildlife and Institute of Arctic Biology, University of Alaska Fairbanks (1980 - 1987).
• Lecturer, Department of Botany, University of California, Davis (Spring 1980).

Membership in Professional Societies

• Society for the Study of Evolution
• Ecological Society of America
• Botanical Society of America
• American Society of Plant Taxonomists
• Association for Tropical Biology

Field Activities