L. Scott Mills

Photo of Mills, L. Scott

L. Scott Mills

Professor, Chancellor's Faculty Excellence Program in Global Environmental Change at NC State University

Website: http://workthatmatters.ncsu.edu/camouflage/

Personal Summary

In July, 2013, Dr. Mills moved to North Carolina State University

Dr. L. Scott Mills was a Professor at the University of Montana (Wildlife Biology Program in the Department of Ecosystem and Conservation Sciences, College of Forestry and Conservation). His research across normally disparate scientific disciplines has led to key advances in applying ecological science to wildlife conservation, including new insights into how genetic variation affects persistence of wild species, new methods for non-invasive abundance estimation and monitoring of population trend, computational measures of the relative importance of various management actions to efficiently affect population growth rate of target species, and direct field measurements of how wildlife respond to climate change and other factors. His research species and systems range from marmots and coyotes in Olympic National Park, to endangered bighorn sheep in the California Sierra Nevada, to fruit bats in the Philippines, to snowshoe hares across North America, to snow leopards and tigers in the Himalayan kingdom of Bhutan.

Early in his career at University of Montana, Dr. Mills was awarded one of the most prestigious awards given by National Science Foundation to junior faculty: A Faculty Early Career Development award.  Since then he has published over 90 scientific articles and has given over 100 professional presentations, including testimony to the U.S. Congress on the role of ethics in conservation science.  

Dr. Mills was a 2009 John Simon Guggenheim Fellow. He has also served on invited committees for the National Science Foundation, National Park Service, National Forest Service, the International Whaling Commission and the National Marine Fisheries Service. He was a member of the Board of Governors for the Society of Conservation Biology, served on the Western Governor’s Association Policy working group on Climate Change Effects on Wildlife, and was a Contributing author to the North America section of the Nobel-Prize winning report from the 2007 International Panel on Climate Change. 

BOOKS - Other books listed under the publications tab

Conservation of Wildlife Populations: Demography, Genetics, and Management (2nd Ed.)

Wiley / Blackwell Publishers
Available December 2012  (at www.wiley.com/buy/0470671491)
[enter code “LS13” at checkout for 20% discount]



PH.D., Biology
University of California, Santa Cruz Ph. D., Biology July 1993
Advisor: Michael Soulé

M.S., Wildlife Ecology
Utah State University, Logan M. S., Wildlife Ecology July 1987
Advisor: Fred Knowlton

B.S., Zoology
North Carolina State University, Raleigh B. S., Zoology May 1983

Courses Taught


Wildlife Population Ecology (WILD 470) (Senior undergrads and beginning graduate students)

Advanced Applied Population Ecology (WILD 570) (Graduate)

General Ecology

Foundations of Wildlife Biology: Techniques and Core Concepts

Frontiers of Conservation Genetics (Graduate)

Conservation Biology (field course at Mountain Lake Biological Station, VA)

Research Interests

My research interests are in the area of applied population ecology. My students and I use population models and genetic tools, coupled with field experiments, to understand population and community-level effects of human stressors. I am also active in developing more efficient and rigorous approaches to population assessment, monitoring, and conservation decision-making.

Currently, much of my work focuses on projects in mountainous landscapes. I am heavily involved in helping to build local capacity for wildlife biology research in the Himalayan Kingdom of Bhutan, where two of my graduate students are using non-invasive genetic sampling and remote cameras to study snow leopards and tigers. We published a book in summer of 2012 providing practical, how-to overviews of wildlife research techniques in mountainous Asian landscapes.  

In mountainous landscapes closer to home, my students and I have studied a number of species ranging from marmots in Olympic National Park confronting invasive coyotes, to Sierra Nevada bighorn sheep whose recovery we have helped guide with population models and genetic analyses, to small mammals along forest edges. I am also continuing my research on snowshoe hares that has been continuous since 1998. Having spent more than a decade understanding hare population dynamics and response to logging (and the powerful role of predation), my primary focus now is on whether hare camouflage can adapt to climate change. Hares change from brown to white seasonally to match their background, and are confronting the single largest signal of climate change in temperate regions: a reduction in duration of snow during winters. My research team is using a multi-disciplinary approach: from climate downscaling, to radiotelemetry and field studies, to gene expression and hormone assays. The goal is to understand how the seasonal coat color change happens, and whether the timing of coat color will be able to shift as the snow season shrinks.

Here are some popular media descriptions of my work:

Snowshoe Hare Chronicles - our research lab blog

The Color of Bunny High Country News, 2/6/2012

Tools that leave wildlife unbothered widen research horizons  New York Times, 3/10/2009

Hare Today, Gone Tomorrow? - The snowshoe hare may become a climate change poster child (VISION - The University of Montana)

The Case of the Climate-Challenged Hare You Tube video - Research by Scott Mills, Professor of Wildlife Population Ecology


Field of Study

Broadly, my field is Wildlife Population Ecology.

But that doesn't capture half the fun! 


Graduate Students

Current graduate students:

  • Julie Betsch (PhD): Time, space, and spillover: Interpreting patterns of emergence to predict disease dynamics at the domestic-wild carnivore interface.
  • Alex Kumar (MS): Effects of forest restoration on snowshoe hare vital rates, behavior and seasonal coat color phenology. 
  • Tshewang Wangchuk (PhD): Monitoring snow leopard populations in remote Himalayan terrain
  • Tshering Tempa (PhD): The Ecology of Tigers in the Intact Mountainous Asian Landscapes of Bhutan
  • Marketa Zimova (MS): Fitness Costs of Seasonal Color Mismatch and Behavioral Plasticity of Snowshoe Hares

Completed graduate students - PhD Degrees

  • Tammy Mildenstein: “Conservation of endangered flying foxes I the Philippines: effects of  anthropogenic disturbance and methods for monitoring.” (2012).
  • Ellen Cheng: “Climate Change, gene flow, and the legendary synchrony of snowshoe hares” (2010).
  • Heather Johnson: “Escaping the extinction vortex: Identifying factors affecting population performance and recovery in endangered Sierra Nevada bighorn sheep” (2010).
  • Sue Cox Griffin: “Demography and ecology of a declining endemic: the Olympic marmot” (2007; co-advised with Mark Taper).
  • John Citta: “Metapopulation ecology using mountain bluebirds as a model system.”  (2005; co-advised with Mark Lindberg).
  • Paul Griffin: “Source-sink dynamics of snowshoe hares in a fragmented landscape.” (2003).
  • Michael Schwartz: “The influence of gene flow and landscape location on genetic variation in two threatened carnivores: the Canada lynx and San Joaquin kit fox.”  (2002)
  • David Tallmon: “Genetic, demographic, and ecological effects of habitat fragmentation.”  (2001; co-advised with Fred Allendorf)

Completed graduate students - M.S. Degrees:

  • Jesse Newby, MS: “”Puma dispersal ecology in the central Rocky Mountains.” (2011).
  • Tshering Tempa, MS (non-thesis): “Teaching Wildlife Biology in Bhutan.”  (2008).
  • Carly Walker: “Landscape structure effects on snowshoe hares in fragmented forests.”  (2005)
  • Julia Witzcuk: “Predation by invasive coyotes on declining endemic Olympic marmots.” (2007).
  • Jenny Woolf: “Effects of thinning and prescribed burning on birds and small mammals.” (2003)
  • Tammy Mildenstein: “Habitat selection of large flying foxes using radiotelemetry: targeting conservation efforts in Subic Bay Philippines.”  (2002)
  • Reesa Yale: “Do highways influence connectivity in small mammals?” (2002)
  • Kevin Lair: "The effects of forest fragmentation and forest edge on Columbian Mouse and Southern Red-Backed vole demography."  (2001)
  • John Citta: “An evaluation of demographic sensitivity analysis models.”  (1998)


  • Rebecca Taylor (2009-2011): Modeling sage grouse population dynamics to inform management decisions.
  • Richard Fredrickson (2007-2010): Evaluating tradeoffs between disease risk and demographic benefits for translocating an endangered bighorn sheep.
  • Cindy Hartway (2006-2010): Sensitivity analysis to inform efficient recovery actions for endangered species on military lands.
  • Matt Kaufmann (2004-2006):  A critical evaluation of the trophic cascade hypothesis for wolves and elk.
  • Karen Hodges (2000-2003): Snowshoe hare population dynamics and response to silvicultural treatments.


Current Senior/Honors Thesis Projects

  • James Goertz: Description of snowshoe hare mortality and survival sites
  • Skyler Suhler: Photos and Phenotypes: Using Camera Traps to Quantify Snowshoe Hare Coat Color Phenology
  • Brandon Davis: Plasticity of snowshoe hare flight response to a potential predation event.
  • Jennifer Hernandez: Analysis of potential cyclic dynamics for southern snowshoe hares.

Past Undergrad Theses and Hall of Fame field and lab assistants
In preparation – 175 or so total students, 20 or so senior and Honor’s thesis projects.


Specialized Skills




  •  John Simon Guggenheim Fellow, 2009 [Lived in Bhutan for 5 months, building local capacity for conducting wildlife science to incorporate into local conservation decision making].
  • Scientific Declaration (Invited) to U.S. District Court on genetic issues associated with delisting Northern Rockies Gray wolves (2008).
  • Invited Member of Western Governor’s Association “Climate Change Working Group” (2008).
  • Invited “Contributor” to North America section, 2007 Intergovernmental Panel on Climate Change Report (IPCC).
  • North American Section Board of Governors, Society for Conservation Biology (2002-04).
  • Certificate of Appreciation Awarded by Cindy S. Swanson, Director of WWFRP (USFS) “For continued support, professionalism, and partnership with Region 1 of the Forest Service as related to the conservation of the Canada lynx”. (Spring 2002)
  • Invited Testimony to U.S. Congress (House Committee on Natural Resources) regarding ethics in conservation science (National Lynx Survey). (March 6, 2002).

State and University-Level

  • One of 3 Professors Invited to Speak in 2009-2010 Provost’s Distinguished Lecturer Series (2009 November). Title of talk:“Wildlife Biology in a Changing World”.
  • Bob Watts Communication Award from the Montana Chapter of The Wildlife Society for my textbook “Conservation of Wildlife Populations.” (Feb. 2009).
  • Keynote Speaker for Academic Convocation, University of Montana. (2006 August)
  • Keynote speaker for University of Montana Undergraduate Research Conference (2006 April)
  • Commencement Speaker, College of Forestry and Conservation (2004 May)
  • School of Forestry Montana Druids Faculty Service Award. Spring 2000.
  • Most Inspirational Faculty Award (University of Montana Silent Sentinel student society, accompanied by $1,000 award from the University). (Spring 1998).

International Experience

Philippine fruit bat work with Tammy Mildenstein (MS and PhD student); 1998-present.
New Zealand collaborative research / teaching exchange.
NSF-supported exchange with Univ. of Porto, Portugal.
Capacity building in Bhutan.
See the 5/6/07 New York Times article on our Program.

Selected Publications


Conservation of Wildlife Populations

Conservation of Wildlife Populations: Demography, Genetics, and Management.  2007 (1st edition) and 2013 (2nd edition). Wiley /Blackwell Press.     




Book Cover


Conservation of Wildlife Populations: Demography, Genetics, and Management (2nd Ed.)

Wiley / Blackwell Publishers
Available December 2012  (at www.wiley.com/buy/0470671491)

[enter code “LS13” at checkout for 20% discount]     



Foresman, K. R., L. S. Mills, and Phurba.  2010. Procedures for Implementing Small Mammal Inventories in Bhutan.  Publication of Ugyen Wangchuck Institute for Conservation and the Environment, Bhutan.




Book Cover



Mills, L. S., Tshering, T., Wangchuk, T.  editors.  Wildlife research techniques in rugged mountainous asian landscapes.  in final preparation for publication in bhutan. (summer 2011 target publication date)








Mills, L. S., M. Zimova, J. Oyler, S. Running, J. Abatzoglou, and P. Lukacs.  2013.  Camouflage mismatch in seasonal coat color due to decreased snow duration.  Proceedings of the National Academy of Sciences 110:7360-7365. [widely covered by media, including Science News, ‘Research Highlights’ of Nature Climate Change, McClatchy Newspaper chain, Science Update].  ** See cover of April 30,2013 Issue of PNAS **

 Witczuk, J., S. Pagacz, L. S. Mills.  In Press.  Disproportionate predation on endemic marmots by invasive coyotes.  Journal of Mammalogy.

Newby, J. R., L. S. Mills, T. K. Ruth, D. H. Pletscher, M. S. Mitchell, H. B. Quigly, K. M. Murphy, and R. DeSimone.  2013.  Human-caused mortality influences spatial population dynamics: pumas in landscapes with varying mortality risks. Biological Conservation. In Press.

Tempa, T., M. Hebblewhite, L. S. Mills, T. Wangchuk, N. Norbu, T. Wangchuk.  2012.  Royal Manas National Park, Bhutan as a wild felid biodiversity hot spot.  Oryx.  In Press.

Hartway, C. and L. S. Mills. 2012. A meta-analysis assessing the effects of common management actions on the nest success of North American birds. Conservation Biology 26:657-666..


Johnson, H. E., L. S. Mills, J. D. Wehausen, T. R. Stephenson, and G. Luikart. 2011. Translating effects of inbreeding depression on component vital rates to overall population growth in endangered bighorn sheep. Conservation Biology 25:1240-1249.


Taylor, R. L., B. L. Walker, D. E. Naugle, and L. S. Mills. 2011. Managing multiple vital rates to maximize greater sage-grouse population growth.   Journal of Wildlife Management 76:336-347.


Walker, C. W., Hodges, K. E., and L. S. Mills. 2011. Influence of stand and landscape features on snowshoe hare densities in fragmented forests. Journal of Mammalogy. 92:561-567.


Hebblewhite, M., M. Musiani, and L. S. Mills. 2010.  Restoration of gene flow among reintroduced Northern Rockies wolf populations.  Molecular Ecology 19:4383-4385.


Johnson, H. R., L. S. Mills, J. Weyhausen, and T. Stevenson. 2010.  Combining ground count, telemetry, and mark-resight data to infer population dynamics in an endangered species. Journal of Applied Ecology.  47:1083-1093.


Johnson, H. R., L. S. Mills, T. Stevenson, and J. Weyhausen. 2010. Population-specific Vital Rate Contributions Influence Management of an Endangered Ungulate. Ecological Applications 20:1753-1765.


Griffin, S. C., M. L. Taper., R. Hoffman, and L. S. Mills. 2010. Ranking Mahalanobis distance models to develop specific predictions of occupancy from presence-only data. Journal of Wildlife Management 74:1112-1121.


Pierson, J. C., L. S. Mills, and D. P. Christian. 2010. Foraging patterns of cavity-nesting birds in fire-suppressed and prescribe-burned ponderosa pine forests in Montana. Open Environmental Sciences 4:49-60.


Humbert, J-Y, L. S. Mills, J. S. Horne, and B. Dennis. 2009. A better  way to estimate population trend. Oikos 118:1487-1498.  PLUS APPENDIX Reviewing exponential trend estimators.


Hodges, K.. E., L. S. Mills, and K. Murphy. 2009. Snowshoe hare distribution and abundance in Yellowstone National Park. Journal of Mammalogy. 90:870-878.


Griffin, P. C., and L. S. Mills. 2009. Sinks without borders: snowshoe hare dynamics in a complex landscape. Oikos 118:1487-1498.


Griffin, S.C., P. C. Griffin, M. L. Taper, and L. S. Mills. 2009. Marmots on the move? Dispersal in a declining mountain mammal. Journal of Mammalogy. 90:686-695.


Witczuk, J., S. Pagacz, and L. S. Mills. 2008. Optimizing methods for monitoring programs: Olympic marmots as a case study. Wildlife Research 35:788-797.


Hodges, K. E. and L. S. Mills. 2008. Designing fecal pellet surveys for snowshoe hares. Forest Ecology and Management. 256:1918-1926.


Harris, N. C., M. Kauffman, and L. S. Mills. 2008. Inferences about ungulate population dynamics derived from age ratios. Journal of Wildlife Management 72:1143-1151.


Griffin,  S. C., M. L. Taper, R. Hoffman, and L. S. Mills. 2008. The case of the missing marmots: are metapopulation dynamics or range-wide declines responsible?  Biological Conservation 141:1293-1309.


Griffin, P. C. and L. S. Mills. 2007. Pre-commercial thinning reduces snowshoe hare abundance in the short term. Journal of Wildlife Management 71:559-564.


Griffin, S. C., T. Valois**, M. L. Taper, and L. S. Mills. 2007. The impact of tourism on Olympic marmot behavior and demography. Conservation Biology 21:1070-1081.


Hard, J.J., Mills, L.S. and Peek, J.M. 2006. Genetic implications of reduced survival of male red deer under harvest. Wildlife Biology 12:349-403.


Griffin, P.C., S. C. *Griffin, C. *Waroquiers, and L. S. Mills.  2005.  Mortality by moonlight: predation risk and the snowshoe hare.  Behavioral Ecology 16:938-944.


Mills, L. S., P. C. *Griffin, K. E. Hodges, K. McKelvery, L. Ruggiero, and T. *Ulizio. 2005.  Pellet count indices compared to mark-recapture estimates for evaluating snowshoe hare density.  Journal of Wildlife Management 69:1053-1062.


Mildenstein, T. L., S. C. Stier, C. E. Nuevo-Diego, and L. S. Mills.  2005.  Habitat selection of endangered and endemic large flying-foxes in Subic Bay, Philippines.  Biological Conservation 126:93-102.


*Schwartz, M. K., and L. S. Mills.  2005.  Gene flow after inbreeding leads to higher survival in deer mice.  Biological Conservation 123:413-420.


*Tallmon, D. A., and L. S. Mills.  2004.  Edge effects and isolation: California red-backed voles revisited.  Conservation Biology 18:1658-1664. 


Leberg, P. L., M. Carloss, L. Dugas, K. L. Pilgrim, L. S. Mills, M. C. Green, and D. S. Scognamillo. 2004.  Recent record of a cougar in Louisiana, with notes on diet, based on analysis of fecal materials.  Southeastern Naturalist 2:653-658.


*Schwartz MK, Mills LS, Ortega Y, Ruggiero L, Allendorf FW. 2003.Landscape Location Affects Genetic Variation of Canada Lynx (Lynx canadensis). Molecular Ecology 12:1807-1816.


*Tallmon, D. A., E. S. Jules, N. J. *Radke, and L. S. Mills.  2003. Of mice and men and trillium: cascading effects of forest fragmentation.  Ecological Applications 13:1193-1203.


Bienen, L., P. C. *Griffin, C. M. Gillin, and L. S. Mills.  2003.  Estimating pregnancy rates and litter size in snowshoe hares using ultrasound.  Wildlife Society Bulletin 31:1066-1072.


*Riddle, A. E., K. L. Pilgrim, L. S. Mills, K. S. McKelvey, L. F. Ruggiero.  2003. Identification of mustelids using mitochondrial DNA and non-invasive sampling.  Conservation Genetics. 4:241-243.


*Funk, W.C., and L. S. Mills.  2003. Potential causes of population declines in forest fragments in an Amazonian frog.  Biological Conservation. 111:205-214.


Griffin, P. C., and L. S. Mills. 2003. “Snowshoe hares in a dynamic managed landscape.”  Pages 438-449 in Editors H. R. Akcakaya, M. A. Burgman, O. Kindvall, C.

Wood, P. Sjogren-Gulve, J. Hatfield, and M. A. McCarthy.  Species Conservation and Management: Case Studies. Oxford University Press.


*Hoekman, S. T., L. S. Mills, D. W. Howerter, J. H. Devries, and I. J. Ball.  2002. Sensitivity analysis of the life cycle of mid-continent mallards.  Journal of Wildlife Management.  66:883-900.


Mills, L. S.  2002.  False samples are not the same as blind controls.  Nature 415:471.


McKelvey, K. S., G. W. McDaniel, L. S. Mills, P. C. *Griffin.  2002.  Effects of plot size and shape on pellet density estimates for snowshoe hare.  Wildlife Society Bulletin 30:751-755. 


*Schwartz, M. K., L. S. Mills, K.S. McKelvey, L.F. Ruggiero, and F. W. Allendorf.  2002.  DNA reveals high dispersal synchronizing the population dynamics of Canada lynx.  Nature 415:520-522.


*Biek, R., W. C. *Funk, B. A. *Maxell, and L. S. Mills. 2002.  What is missing in amphibian decline research: Insights from ecological sensitivity analysis. Conservation Biology.  16:728-734. APPENDIX OF VITAL RATES


*Tallmon, D. A., H. M. *Draheim, L. S. Mills, and F. W. Allendorf. 2002. Insights into recently fragmented vole populations from combined genetic and demographic data.  Molecular Ecology 11:699-709. 


*Biek, R., L. S. Mills, and B. Bury.  2002.  Terrestrial and stream amphibians across clear-cut-forest interfaces in the Siskiyou Mountains, Oregon.  Northwest Science 76:129-140. 


Reed, J. M., L. S. Mills, J. B. Dunning, Jr., E. S. Menges,  K. S. McKelvey, R. Frye, S. R. Beissinger, and M-C Anstett, and P. Miller.  2002.  Emerging issues in population viability analysis.  Invited Paper, Conservation Biology 16:7-19.


Mills, L. S., K. L. Pilgrim, M. K. *Schwartz, and K. McKelvey.  2000.  Identifying lynx and other North American felids based on mtDNA analysis.  Conservation Genetics 1:285-288.


Mills, L. S., D. F. Doak, and M. J. Wisdom.  2000.  Elasticity analysis for conservation decision-making: Reply to Ehrlen et al., Conservation Biology 15:281-283.


Mills, L. S., J. J. *Citta, K. *Lair, M. *Schwartz, D. *Tallmon.  2000. Estimating animal abundance using non-invasive DNA sampling: Promise and Pitfalls.  Ecological Applications 10:283-294.


Wisdom, M. J., L. S. Mills, and D. F. Doak.  2000. Life-stage simulation analysis: estimating vital rate effects on population growth for conservation.  Ecology 81:628-641.


Mills, L. S., D. F. Doak, and M. J. Wisdom.  1999.  The reliability of conservation actions based on sensitivity analysis of matrix models.  Conservation Biology 13:815-829.


Jules, E., E. Frost, D. *Tallmon, and L. S. Mills.  1999.  Ecological consequences of forest fragmentation in the Klamath region.  Natural Areas Journal 19:368-378.


*Citta, J. J., and L. S. Mills.  1999.  What do demographic sensitivity analyses tell us about controlling brown-headed cowbirds?  Studies in Avian Biology 18:121-134.


Soulé, M. E., and L. S. Mills.  1998.  No need to Isolate Genetics.  Science 282:1658-1659.


Wisdom, M. J., and L. S. Mills.  1997.  Using sensitivity analysis to guide population recovery: Prairie chickens as an example.  Journal of Wildlife Management 61:302-312.


Mills, L. S., and F. W. Allendorf.  1996.  The one-migrant-per-generation rule in conservation and management.  Conservation Biology 10:1509-1518.


Morrison, M. L., L. S. Mills, and A. J. Kuenzi.  1996.  Study and management of an isolated, rare population: The Fresno Kangaroo rat.  Wildlife Society Bulletin 24:602-606.


Power, M.E., D. Tilman, J. Estes, B. A. Menge, W. J. Bond, L. S. Mills, G. Daily, J. C. Castilla, J. Lubchenco, and R. T. Paine.  1996.  Challenges in the quest for keystones.  Bioscience 46:609-620.


Mills, L. S., S. G. *Hayes, C. *Baldwin, M. J. *Wisdom, J. *Citta, D. J. *Mattson, and K. *Murphy.  1996.  Factors leading to different viability predictions for a grizzly bear data set.  Conservation Biology 10:863-873.


Mills, L. S.  1995.  Edge effects and isolation: red-backed voles on forest remnants.  Conservation Biology 9:395-403.


Power, M. E., and L. S. Mills.  1995.  The keystone cops meet in Hilo.  Trends in Research in Evolution and Ecology.  10:182-184.


Scott, J. M., T. H. Tear, and L. S. Mills.  1995.  Socioeconomics and the recovery of endangered species: biological assessment in a political world.  Conservation Biology 9:214-216.


Mills, L. S., and P. E. Smouse.  1994.  Demographic consequences of inbreeding in remnant populations.  American Naturalist 144:412-431.


Doak, D., and L. S. Mills.  1994.  A useful role for theory in conservation.  Ecology 75:615-626.


*Clarkson, D. A., and L. S. Mills.  1994.  Ecological factors associated with Hypogeous sporocarps in fragmented forests. Northwest Science 68:259-265.


*Tallmon, D.A., and L. S. Mills.  1994.  Log use and home ranges of California red-backed voles on a forest remnant.  Journal of Mammalogy.  75:97-101.


Mills, L. S., M. S. Soulé, and D. F. Doak.  1993.  The keystone species concept in ecology and conservation.  Bioscience 43:219-224.


Mills, L. S., R. J. Fredrickson, B. B. Moorhead.  1993.  Characteristics of old-growth forests associated with northern spotted owls in Olympic National Park.  Journal of Wildlife Management 57:315-321.


Mills, L. S., and F. F. Knowlton.  1991.  Coyote space use in relation to prey abundance.  Canadian Journal of Zoology 69:1516-1521.


Mills, L. S., and F. F. Knowlton.  1989.  Observer performance in known and blind radio-telemetry accuracy tests.  Journal of Wildlife Management.  52:340-342.




Mills, L. S., and H. E. Johnson. 2013. Wildlife Population Dynamics. In Krausman, P., editor. Wildlife management: Contemporary principles and practices. John Hopkins University Press (In Press).


Mills, L. S., J. M. Scott, K. M. Strickler, and S. A. Temple. 2012. Ecology and management of small populations. Pages 270-292 In Techniques for wildlife investigations and management (ed. N. Silvy). Sixth edition. The Wildlife Society, Bethesda, Maryland, USA.


Kelly, M. J., J. Betsch, C. Wultsch, B. Mesa, and L. S. Mills. 2011. Noninvasive Sampling for carnivores. Pages 47- 69  In Boitani, L. and R. Powell, editors. Carnivore Ecology and Conservation. Oxford University Press, Oxford, UK.


Running, S. W. and L. S. Mills. 2009. Terrestrial Ecosystem Adaptation to Climate Change. Resources for the Future Report for the Climate Policy Program at RFF. 


Mills, L. S. 2008. Crossing Disciplines for Endangered Species. Review for Ecology of Scott, J. M., D. D. Goble, and F.W. Davis, editors. The Endangered Species Act at Thirty: Conserving Biodiversity in Human-Dominated Landscapes (Volume 2). Island Press, Washington, D.C. Ecology 89:592-593.


Mills, L. S. 2006 “Contributing Author” to “Chapter 14:  North America” by Field, C. B., L. D. Mortsch, M. Brklacich, D. L. Forbes, P. Kovaks, J. A. Patz, S. W. Running and M. J. Scott, in: Climate Change 2007: Impacts, Adaptation and Vulnerability. Contribution of Working Group II to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change, M. L. Parry, O. F. Canziani, J. P. Palutikof, P. J. van der Linden and C. E. Hanson, Eds., Cambridge University Press, Cambridge, UK, 6127-652.


Mildenstein, T.L., L.S. Mills, P. Landres, G. Rochefort, J. Scharberl. 2006. Recreation impacts on wildlife: review of recent research and a call for   investigation of population and community level effects. Final report to Mount Rainier National Park VERP Program.


Mills, L. S., J. M. Scott, K. M. Strickler, and S. A. Temple.  2005.  Ecology and management of small populations. Invited Article in Wildlife Techniques Manual.  Wildlife Society. 


Mills, L. S., and M. E. Soulé.  In Press.   A brief history of the role of genetics in conservation.  Invited article (“box”) in Conservation Genetics textbook by F. Allendorf and G. Luikart.


Mills, L. S., M. K. Schwartz, D. A. Tallmon, and K. P. Lair.  2003.  Measuring and interpreting changes in connectivity for mammals in coniferous forests.  Pages 587-613 in C. J. Zabel and R. G. Anthony, editors.  Mammal Community Dynamics: Management and Conservation in the Coniferous Forests of Western North America.  Cambridge University Press, New York, USA.


Griffin, P. C., and L. S. Mills.  2003  “Snowshoe hares in a dynamic managed landscape.”  Pages 438-449 in Editors H. R. Akcakaya, M. A. Burgman, O. Kindvall, C.


Wood, P. Sjogren-Gulve, J. Hatfield,   and M. A. McCarthy.  Species Conservation and Management: Case Studies. Oxford University Press.


Mills, L. S., and M. Lindberg.  2002. “Sensitivity Analysis to Evaluate the Consequences of Conservation Actions.”  Pages 338-366 in S. R. Beissinger and D. R. McCullough, editors.  Population Viability Analysis.  University of Chicago Press.


Mills, L. S.  2002.  “Genetics, Demography, and Viability of Fragmented Populations.” (Book Review)  Quarterly Review of Biology 77:222-223.


Mills, L. S., and D. Tallmon. 1999.  “Genetic issues in forest fragmentation.”  Pages 171-184 In Forest Fragmentation: Wildlife and Management Implications. J.

Rochelle, L. A. Lehmann, and J. Wisniewski, eds.  Brill Publishers (Netherlands).


Dobson, A., K. Ralls, M. Foster, M. E. Soule, D. Simberloff, D. Doak, J. A. Estes, L. S. Mills, D. Mattson, R. Dirzo, H. Arita, S. Ryan, E. A. Norse, R. F. Noss, and D. Johns.  1999.  “Connectivity: maintaining flows in fragmented landscapes.”  Pages 129-171 In Soulé, M. E. and J. Terborgh, editors.  Continental Conservation: Scientific Foundations of Regional Reserve Networks.  Island Press.


Groom, M., D. B. Jensen, R. L. Knight, S. Gatewood, L. Mills, D. Boyd-Heger, L. S. Mills, and M. E. Soulé.  1999.  “Buffer zones: benefits and dangers of compatible stewardship.”  Pages 191-198 In Soulé, M. E. and J. Terborgh, editors.  Continental Conservation: Scientific Foundations of Regional Reserve Networks.  Island Press.


Mills, L. S.  1997.  “Book Review: Population Management for Survival and Recovery: Analytical Methods and Strategies in Small Population Conservation.”  Journal of Wildlife Management 61:251-252.


Mills, L. S.  1996. “Fragmentation of a natural area: Dynamics of isolation for small mammals on forest remnants.” Pages 199-219 In Wright, G., editor.  National Parks and Protected Areas: Their Role in Environmental Protection.  Blackwell Press.


Mills, L.S.  1996.  “Keystone Species.” in Paehlke, R., editor.  Encyclopedia of Conservation and Environmentalism. Garland Publishing Co., New York. 


Mills, L.S. 1995.  “Keystone Species.”  Pages 381-387 in Encyclopedia of Environmental Biology. Academic Press, Inc.


Mills, L.S.  1994.  “Book Review: Principles of Conservation Biology.”  Northwest Science 68:303-304.

Soulé, M.E., and L. S. Mills.  1992.  “Conservation genetics and conservation biology: a troubled marriage.”  Pages 55-69 in Sandlund, O. T., K. Hindar, and A. H. D.


Brown, eds., Conservation of Biodiversity for Sustainable Development.  Scandinavian Univ. Press, Oslo.



  • Hodges, K. E., and L. S. Mills.  2005.  Snowshoe hares in Yellowstone. Yellowstone Science 13:3-6.
  • Yale, R. and L. S. Mills.  2000.  “Do highways fragment small mammal populations?” Proceedings of the International Conference on Ecology and Transportation.
  • Mills, L. S. 1996.  “Cheetah extinction: genetics or extrinsic factors?”  Letter to the editor.  Conservation Biology 10:315.
  • Zager, P., L. S. Mills, W. Wakkinen, and D. Tallmon.  1995.  “Woodland caribou --a conservation dilemma.”  Endangered Species Update.
  • Mills, L. S.  1995. “The use of population ecology approaches to assess re-introduction of endangered Selkirk Caribou.”  Final report, submitted to: a) Idaho Fish and Game; b) Washington Department of Wildlife.
  • Mills, L. S.  1995.  “The role of population ecology in guiding wildlife translocations.”  Final report, submitted to U.S. Forest Service, Missoula, MT.
  • Cassidy, K., E. O. Garton, W. B. Krohn, L. S. Mills, J. M. Scott, and K. Williams.  1994.  “National guidelines for assessment of reliability of GAP vertebrate distributions.”  National Biological Survey.
  • Mills, L. S.  1994.  “Effects of forest fragmentation on small mammals in Southwest Oregon.”  COPE Report [(Coastal Oregon Productivity Enhancement Program), an educational non-technical publication for forest managers.]  7:6-8. 
  • Mills, L. S. and M. Morrison.  1993.  “Final Report: Ecology of the Fresno Kangaroo Rat and associated small mammals at NAS Lemoore”.  Prepared for Lemoore Naval Air Station.
  • Fredrickson, R.J., L.S. Mills, and B.B. Moorhead.  1989.  “Spotted owl surveys in Olympic National Park -- 1988 and 1989.”  Olympic National Park, Port Angeles, WA.
  • Mills, L.S., R.J. Fredrickson, B.B. Moorhead, and D.U. Sharp. 1988. “Spotted owl distribution along elevation and vegetation gradients in Olympic National Park.”  1988 Progress Report.  Olympic National Park, Port Angeles, WA.