Research Interests
My research interests range broadly in evolutionary and behavioral ecology. Special interests include life history theory, mating system theory, and optimal foraging theory. I emphasize experimental approaches to the study of adaptation, particularly with species that display geographical variation in the characteristics of interest. Current research uses birds as study systems, but taxonomic interests include most vertebrates and insects. Current projects include 1) the evolution of sexual dichromatism and cavity-nesting in the Hawaii Akepa, 2) the evolution of reproductive rate and permanent monogamy in tropical House Wrens in lowland central Panama, and 3) evolution of introduced birds in Hawai`i. I am closely associated with researchers in the Hawaiian Evolutionary Biology Program in attempts to use both native and introduced species in Hawai`i as model systems for understanding evolutionary processes.

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Grants and Awards

Australian Research Council, 2004-2005. Ecological Modelling of Infectious Disease for
Conserving Wildlife. (PI, Hamish McCallum; Co-PI, L. Freed)
Environmental Protection Agency, Wildlife Risk Assessment. 2001-2004. Habitat
Degradation and Introduced Diseases Stress the Endangered Hawaii Akepa.
(PI, Rebecca L. Cann, M.Lee Goff, Co-PIs)
University of Hawaii, Educational Improvement Award. 1999. Undergraduate Research
Experience at Hakalau Forest Biological Field Station.
National Science Foundation, 1996-1998. "FSML: Hakalau Forest Biological Field
Station" (Alan Teramura, Co-PI).
U.S. Forest Service, 1996-1997. "What Does an Old Growth Ohia/Koa Forest Look
Like?"
State of Hawaii, Natural Areas Reserve Commission, 1994?1997. "Habitat Use by the
Hawaiian Hoary Bat on the Island of Hawaii."
U.S. Fish and Wildlife Service, 1993?1997. "Comparative Ecological and Genetic
Studies of Native Birds."
U.S. Fish and Wildlife Service, 1993?1994. "Proposal for a Survey of Rota Bridled
White?Eyes in the Commonwealth of the Northern Mariana Islands."
University of Hawaii, Howard Hughes Medical Institute Program for
Undergraduate Research, 1993?1994.
University of Hawaii, Arts and Sciences Advancement Fund for Undergraduate Field
Research Experience, 1993?1994
U.S. Fish and Wildlife Service, 1992?1993. "Hawaii Akepa Artificial Nest Box Study."
U.S. Fish and Wildlife Service, 1992?1993. "Foraging Behavior of the Endangered
Hawaiian Hoary Bat, Lasiurus cinereus semotus."
U.S. Forest Service, 1991?1992. "Artificial Cavities for Preventing Nest?Site Limitation
in the Endangered Hawaii Akepa."
State of Hawaii, Natural Area Reserves Commission, 1991?1992. "Status, Structure, and Health of Native Bird Populations at Hanawi Natural Area Reserve."
University of Hawaii Foundation, 1990. "Conservation of Native Hawaiian Birds."
University of Hawaii Foundation, l990. "Hana Rain Forest Project Fund."
State of Hawaii, Division of Forestry and Wildlife, l990?l99l. "Distribution and
Abundance of the Hawaiian Hoary Bat on the Island of Hawaii."
John D. and Catherine T. MacArthur Foundation, l989?present. "Conservation of
Native Hawaiian Forest Birds and Other Species with a precarious Future."
(Rebecca Cann, Sheila Conant, Co-PIs)
University of Hawaii, Project Development Award, l988?l989. "Social Organization and
Post?Breeding Survival of Native Hawaiian Forest Birds."
University of Hawaii, Fujio Matsuda Fellow, l988?l989. "Social Organization and
Post?Breeding Survival of Native Hawaiian Forest Birds."
University of Hawaii, Educational Improvement Fund Mini?Grant, 1988?1989. "Field
Techniques in Ornithology, Evolutionary Biology, and Conservation Biology."
Research Relations Fund, University of Hawaii, 1987-1988. (Co?PI with Rebecca L.
Cann), l987?l988.
Project Development Award, University of Hawaii, l987?l988.
Research and Training Revolving Fund Award, University of Hawaii, (Co?PI with
Robert C. Fleischer), l986?l987.
University of Hawaii Foundation, l986?l989.
National Science Foundation (Co?PI with Robert C. Fleischer), l986?l988.
Research and Training Revolving Fund Award, University of Hawaii, l985?l986.
National Geographic Society, l985?l986.
National Geographic Society, l984?l985.
Scholarly Studies Program, Smithsonian Institution (Co?PI with Neal G. Smith),
l983.
Hawaiian Evolutionary Biology Program, University of Hawaii, l983.
Frank M. Chapman Memorial Fund, American Museum of Natural History, l98l.
W.C. Allee Award for Best Student Paper, Animal Behavior Society, l98l.
Sigma Xi Grant?in?aid of Research, l98l.
Best Student Paper Award (Runner?up), American Ornithologists' Union, l980.
NSF Doctoral Dissertation Improvement Grant, l978?l980.
N.I.H. National Research Service Award, Neurobehavioral Sciences Program, University
of Iowa, l979?l980.
Navy Achievement Medal, U.S. Navy, l972.

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Publications

Freed, L.A., and Cann, R.L. (in press). On comparing PCR tests for detecting malaria
in birds. Journal of Parasitology:

Freed, L.A., and Lepson, J.K. (in review). Female plumage and fitness in the sexually
dichromatic Hawaii Akepa. Behavioral Ecology:

Hart, P.J., and Freed, L.A. 2003 Structure and dynamics of mixed species flocks
in a Hawaiian rain forest. Auk 120: (in press)

Freed, L.A. 2002. Riding the tiger. (Invited book review). Quarterly Review of
Biology 77: 225.

Shehata, C., Freed, L.A, and Cann, R.L. 2001. Changes in native and introduced
bird populations on Oahu: infectious diseases and species replacement. Pages 264-
273 In Evolution, Ecology, Conservation and Management of Hawaiian Birds: A
Vanishing Avifauna. (M. Scott, S. Conant, C. van Riper, eds). Studies in Avian
Biology No. 22. Cooper Ornithological Society.

Freed, L.A. 2001. Significance of old-growth forest to the Hawaii Akepa. Pages 173-
184 In Evolution, Ecology, Conservation, and Management of Hawaiian Birds: A
Vanishing Avifauna. of the Hawaiian Avifauna (M. Scott, S. Conant, C. van
Riper, eds.). Studies in Avian Biology No. 22. Cooper Ornithological Society.

Freed, L.A. 2000. Female ornamentation, mate choice and sexual selection. Trends in
Ecology and Evolution 15: 471.

Freed, L.A. 1999. Extinction and endangerment of Hawaiian Honeycreepers: a
comparative approach. Pages 137-162 In Genes, Species, and Extinction (L.
Landweber and A. Dobson, eds.). Princeton University Press.

Lepson, J.K., and L.A. Freed. 1997. Akepa (Loxops coccineus). In The Birds of North
America, No. 294 (A.Poole and F. Gill, eds.). The Academy of Natural Sciences,
Philadelphia, PA, and The American Ornithologists' Union, Washington, D.C.

Wingfield, J.C., Hunt, K., Breuner, C., Dunlap, K., Fowler, G.S., Freed, L., and Lepson,
J. 1997. Environmental stress, field endocrinology, and conservation biology. Pages
95-131 In Behavioral Approaches to Conservation in the Wild (J.R. Clemmons and
R. Buchholz, eds.). University Press, Cambridge.

Freed, L.A., Smith, T.B., Carothers, J.H., Lepson, J.K. 1996. Shrinkage is not the most
likely cause of bill change in the iiwi: a rejoinder to Winker. Conservation Biology
10: 659-660.

Cann, R.L., Feldman, R.A., Agullana, L., and Freed, L.A. 1996. A PCR approach to
detection of malaria in Hawaiian birds. Pages 202-213 In Molecular Genetic
Approaches to Conservation Genetics (T. Smith, R. Wayne, eds.). Oxford
University Press.

Lepson, J.K., and L. A. Freed. 1995. Variation in male plumage and behavior of the
Hawaii Akepa. Auk 112: 402-414.

Feldman, R.A., Freed, L.A., and Cann, R.L. 1995. A PCR test for avian malaria in
Hawaiian birds. Molecular Ecology 4: 663-673.

Smith, T.B., Freed, L.A., Lepson, J.K., and Carothers, J.H. 1995. Evolutionary
consequences of extinctions in populations of a Hawaiian Honeycreeper.
Conservation Biology 9: 107?113.

Cann, R.L., Feldman, R.A., Freed, L.A., Lum, J.K., and Reeb, C.A. 1993. Collection
and storage of vertebrate samples for molecular evolutionary studies. Pages 38?51 in
Molecular Evolution: Producing the Biochemical Data (Methods in Enzymology,
volume 224; E.A. Zimmer, T.J. White, and R.L. Cann, eds), Academic Press,
Orlando, FL.

Freed, L.A. 1991. Mate replacement during the non?breeding months in tropical house
wrens. Acta XX Congressus Internationalis Ornithologici:1214?1219.

Freed, L.A. 199l. Selection in a Galapagos finch: value of long?term study, (Invited
book review of Evolutionary Dynamics of a Natural Population, by B. Rosemary
Grant and Peter R. Grant). Evolution 45: l296?l297.

Freed, L.A. 1990. Reproductive sociality. (Invited book review of Cooperative
Breeding in Birds, edited by Peter B. Stacey and Walter D. Koenig). Science 250:
1282?1283.

Freed, L.A. 1990. Invited book review of Conservation Biology in Hawaii (edited by
Charles P. Stone and Danielle B. Stone). Quarterly Review of Biology 65:
105?106.

Freed, L.A., and R.L. Cann. l989. Integrated conservation strategy for Hawaiian forest
birds. Bioscience. 39: 475?476.

Palumbi, S.R., and L.A. Freed. l988. Agonistic interactions in a keystone predatory
starfish. Ecology 69: l624?l627.

Freed, L.A. l988. Forced fledging: investigation of the lengthy nestling period of
tropical house wrens. National Geographic Research 4: 395?407.

Freed, L.A. l988. Demographic and behavioral observations of the Hawaii Akepa.
Elepaio 48:37?39.

Freed, L.A., Conant, S., and R.C. Fleischer. l987. Evolutionary ecology and radiation of Hawaiian passerine birds. Trends in Ecology and Evolution 2: l96?203.

Freed, L.A., Telecky, T.M., Tyler, W.A. III, and M.A. Kjargaard. l987. Nest?site
variability in the Akepa and other forest birds on the island of Hawaii. Elepaio 47:
79?8l.

Freed, L.A. l987. Prospective infanticide and protection of genetic paternity in tropical house wrens. American Naturalist l30: 948?954.

Freed, L.A. l987. The long?term pair bond of tropical house wrens: advantage or
constraint? American Naturalist l30: 507?525.

Freed, L.A. l987. Rufous?and?white wrens kill house wren nestlings during a food
shortage. Condor 89: l95?l97.

Freed, L.A. l986. Usurpatory and opportunistic bigamy in tropical house wrens.
Animal Behaviour 34: l894?l896.

Freed, L.A. l986. Territory takeover and sexually selected infanticide in tropical house
wrens. Behavioral Ecology and Sociobiology l9: l97?206.

Freed, L.A. l985. Journey to adulthood (invited book review of The Growth and
Development of Birds, by R.C. O'Connor). Nature 3l8:5l7.

Freed, L.A. l98l. Breeding biology of House Wrens (Troglodytes aedon): new views of
avian life history phenomena. Ph.D. Dissertation, University of Iowa.

Freed, L.A. l98l. Loss of mass in breeding wrens: stress or adaptation? Ecology 62:
ll79?ll86.

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Symposia and Plenary Sessions Organized

The Role of Behavior in Evolution. Plenary Session for International Ethological
Conference, Honolulu, Hawaii, 1995.

Conflict in Cooperative Systems. Plenary Session for International Ethological
Conference, Honolulu, Hawaii, 1995.

Principles of Restoration and Regeneration. Symposium for Hawaii Conservation
Conference, Honolulu, Hawaii, 1994.

Avian Evolution in Hawaii. Symposium for the Society for the Study of Evolution,
Hilo, Hawaii, l991.

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Invited and Contributed Papers

Molting and breeding strategies in Hawaiian honeycreepers. Third North American
Ornithological Conference, New Orleans, 2002.

Life history characteristics underlie extinction and endangerment in Hawaiian
honeycreepers. Society for Conservation Biology meeting, Hilo, 2001.

Female plumage and fitness select for male preference in the sexually dichromatic Hawaii
Akepa. International Meeting of the Society for Behavioral Ecology, Zurich, 2000
(Jaan Lepson, co-author).

Bright females and fitness: a reason for conserving endangered species is that they can be
extremely interesting. Invited presentation to the President of the National Audubon
Society, Hawaii Audubon Society, Honolulu, HI. 1998.

Non-random community structure and patterns of extinction in Hawaiian honeycreepers
of Metrosideros/Acacia forests. Invited paper for Symposium on Ecology and
Evolution of Hawaiian Birds, Cooper Ornithological Society, Hilo, HI. 1997.
(Rebecca L. Cann, co-author).

Current trends in infectious disease among Hawaiian forest birds. Invited paper for
Symposium on Ecology and Evolution of Hawaiian Birds, Cooper Ornithological
Society, Hilo, HI. 1997 (Co-author with Rebecca L. Cann, Jobriath Rohwer, Eric
VanderWerf, and Cherie Shehata).

Evolution, extinction, and conservation in the Hawaiian Islands. Invited paper for
Symposium on Genes, Species, and Extinction: The Role of DNA in
Conservation. Princeton University. 1996.

Introduction to Hawaii for ethologists. Invited extended introduction to Plenary Session
on the Role of Behavior in Evolution. International Ethological Conference,
Honolulu, Hawaii, 1995.

Regeneration of critical resources for endangered species. Invited paper for
Restoration/Regeneration Symposium, Hawaii Conservation Conference,
Honolulu, 1994.

Trends in the ecology and evolution of Hawaiian passerine birds. Invited paper for
Symposium on Avian Evolution in the Pacific Region. Association for Tropical
Biology, AIBS, Honolulu, 1992.

Hawaiian forest bird breeding biology: it's a tough life. Invited paper for Symposium on
Avian Evolution in Hawaii. Society for the Study of Evolution, Hilo, Hawaii,
l991.

Mate replacement during the non?breeding months in tropical house wrens. Invited
paper for Symposium on Social Behaviour in the Non?breeding Season.
International Ornithological Congress, Christchurch, New Zealand, l990.

Flycatching as a novel foraging behavior in Hawaiian honeycreepers. Animal Behavior
Society, Binghamton, l990.

Kin selection may shape fledging behavior in tropical house wrens. American
Ornithologists' Union, Fayetteville, l988.

Permanent monogamy and sexually selected infanticide in tropical house wrens.
International Ethological Conference, Madison, l987 (invited).

Long?term pair?bonds of tropical house wrens: advantage or constraint? Cooper
Ornithological Society, Davis, l986.

Usurpatory and opportunistic bigamy in tropical house wrens. Animal Behavior Society,
Tucson, l986.

Reproductive rate of tropical house wrens. American Ornithologists' Union, Tempe,
l985.

The unstable permanently monogamous mating system of tropical house wrens. Animal
Behavior Society, Raleigh, l985.

Optimal foraging by parental house wrens: change in decision rules caused by a nest
predator. Societe Quebocoise pour l'Etude Biologique du Comportement,
Montreal, l982.

Male response to female desertion of nestlings in house wrens. Societe Quebecolse pour
l'Etude Biologique du Comportement, Montreal, l982.

Optimization of a behavioral duplex: foraging and protection of young in house wrens.
American Society of Naturalists, Iowa City, l98l.

Optimal foraging by house wrens in the face of conflicting demands. Animal Behavior
Society, Knoxville, l98l.

Weight loss in breeding wrens - stress of adaptation? American Ornithologists' Union,
Fort Collins, l980.

Sibling competition among nestling house wrens. American Society of Zoologists,
Tampa, l979.

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Other Speaking Engagements (Invited and Voluntary)

Seminar, Smithsonian Tropical Research Institute, Panama, 2002.
Seminar, Pacific Ecosystem Research Center, Volcano, Hawaii. 2000.
Seminar, Department of Ecology and Evolutionary Biology, Princeton Unversity, 1996.
Meeting, University of Hawaii Foundation Trustees, 1995.
Seminar, Department of Biology, University of Rome, Tor Vergata, 1993.
Annual Sigma Xi Initiation Dinner, Hawaii Chapter of Sigma Xi, 1992.
Seminar, Department of Botany, University of Hawaii, 1992.
Seminar, Museum of Vertebrate Zoology, University of California at Berkeley, 1989.
Seminar, Institute of Environmental Studies, University of Washington, 1989.
Seminar, Department of Zoology, University of Washington, 1989.
Seminar, Department of Zoology, University of Hawaii, l987.
Meeting, Hawaii Audubon Society, l986.
Seminar, Museum of Vertebrate Zoology, University of California at Berkeley, l986.
Seminar, Department of Zoology, University of Hawaii, l986.
Seminar, Smithsonian Tropical Research Institute, l985.
Meeting, Society for the Study of Pacific Evolutionary Biology, l984.
Lecture, Organization for Tropical Studies (course OTS 83?l), l983.
Seminar, Smithsonian Tropical Research Institute, l983.
Seminar, Department of Neurobiology and Behavior, Cornell University, l982.
Seminar, Department of Biology, McGill University, l982.
Seminar, Department of Zoology, University of Hawaii, l982.
Seminar, Smithsonian Tropical Research Institute, l982.
Meeting, Panama Audubon Society, l982.
Seminar, Department of Zoology, University of Iowa, l98l.
Seminar, Department of Ecology, Ethology, and Evolution; University of Illinois at
Champaign?Urbana, l98l.
Annual Meeting, Iowa Ornithologists' Union, l98l.
Seminar, Department of Biology, Illinois State University, l98l.
Colloquium, Department of Statistics, University of Iowa, l978.

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Professional Societies

American Ornithologists' Union
American Society of Naturalists
Animal Behavior Society
Association of Field Ornithologists
International Society for Behavioral Ecology
Cooper Ornithological Society
Ecological Society of America
Hawaii Audubon Society
Sigma Xi, The Scientific Research Society
Society for Conservation Biology
Society for the Study of Evolution
Wilson Ornithological Society

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Recovery Teams

Hawaii Forest Bird Recovery Team, USFWS, 1994-present

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Community and Professional Service

Reviewer, Environmental Center, University of Hawaii.
Radio interview, Hawaiian forest birds, KTUH, l986.
Invited participant, United States Fish and Wildlife Service, Special Honolulu meeting on
"Conservation of Hawaiian Forest Birds," l987.
Invited speaker, Sierra Club (Hawaii Chapter), l988.
Invited speaker, Hawaii Audubon Society, l988, 1998
Invited workshop leader, Care of the Earth Symposium, The Lutheran Church of
Honolulu, l989.
Planning Committee, Hawaii Conservation Biology Initiative
Reviewer (NSF, National Geographic Society, Condor, Auk, American Naturalist,
Ecology, Journal of Field Ornithology, Molecular Phylogenetics and
Evolution)
Applied Animal Behavior Committee, Animal Behavior Society, 1992?1995.
Director, Organization for Tropical Studies
Member, Animal Species Advisory Commission, State of Hawaii
Member, Hawaii Conservation Forum, 1994.
Organizer, Tester Symposium, University of Hawaii, 1991, 1999

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