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Wildlife Genetics

Wildlife Genetics
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The Wildlife Ecology Program of the Rocky Mountain Research Station, with its partners, recognized the contemporary role that molecular genetics plays in the management of wildlife. Thus, it initiated efforts in wildlife genetics in 1998 with the University of Montana and the Northern Region of the USFS. Currently the RMRS Conservation Genetics Program, of which the Wildlife Genetics Laboratory is central, focuses on conducting research that is highly integrated with USFS management needs across the country. The Wildlife Genetics Laboratory works with states, tribes, universities and private groups answering state-of-the-art questions in wildlife genetics, and providing answers to pressing wildlife management needs.

Projects

  • Lynx - Bobcat Hybridization
  • Detecting Allelic Dropout in Non-Invasive Genetic Samples
  • Landscape Genetics
  • Determining the Distribution of Fisher in the Rocky Mountains
  • Estimating Gene Flow of Fisher in the Rocky Mountains
  • Species / Individual Identification From Scat, Hair, and Urine
  • Ancient DNA of Wolverine in California
  • Mountain Lion Relatedness in Yellowstone National Park
  • Wolverine Relatedness in the Sawtooth Mountain Range
  • Otter Genetic Structure in Idaho and Montana

Fisher Survey

Determining Fisher Distribution in the Rocky Mountains

The USFWS has established that fisher in the Pacific Coast States are “warranted, but precluded” from listing under the Endangered Species Act (2004). In the Rocky Mountains fisher habitat may be more contiguous than in the Pacific Coast States, but the presence / absence of fisher on forest lands and their level of connectivity to other fisher populations remains largely unknown. We are currently working with multiple agencies and organizations to detect fisher in the Rocky Mountains.

Objectives

This work has a primary objective and two secondary objectives:

  • To delineate the geographic range of fisher in the Rocky Mountains. Specifically we are interested in detecting all the populations, and determining these populations' boundaries
  • To determine which Rocky Mountain fisher populations have native genes and which fisher populations are comprised of reintroduced individuals
  • To index the abundance of fisher (e.g., minimum number of individuals alive) in each population through the use of DNA

To meet these goals we have established a standardized survey. The accompanying grid and GIS layers can also be downloaded (54MB).

Fisher article in Horizon Air Magazine

Fascinating Fishers, from the July 2006 issue of Horizon Air magazine, is posted here with the written permission of the publisher, Paradigm Communicatons Group, Seattle, WA, Copyright 2006. All rights reserved by the publisher. No printing, Web posting, duplication or distribution by any means, manual or electronic, may be done without the express written permission of the publisher.

Partners

This project is a partnership between Forest Service Research (Rocky Mountain Research Station, Pacific Northwest Research Station, Pacific Southwest Research Station), Coeur D’Alaine Tribes, Potlach Timber Company, Forest Capital Partners, The Northern Region (Region 1 of the US Forest Service), Clearwater and Panhandle National Forests, University of Idaho, Idaho Fish and Game, Great Burn Study Group, and Montana Fish Wildlife and Parks.

Software

KERNEL: A GIS utility

image generated by the software

Moving windows are commonly used in GIS. An important use is to create buffers around features. Buffering can be done using focal functions, but in many cases you want a focal function whose value declines with distance from the feature. ESRI's ArcGIS allows these functions to be applied through the creation of a kernel file. What Kernel does is provide an easy graphical approach to create kernel files. Importantly, it will automatically scale these files so that the function's area is constant regardless of its shape.

Download

Kernel is free software for the MS-Windows operating system. The software was developed by Kevin McKelvey. The download file is in ZIP format and includes documentation. If you download this software please contact Kevin with your name and affiliation.

DROPOUT: Genotyping Error Software

Genotyping error, often associated with low-quantity/quality DNA samples, is an important issue when using genetic tags to estimate abundance using capture-mark-recapture (CMR). DROPOUT, a MS-Windows program, identifies both loci and samples that likely contain errors affecting CMR estimates. DROPOUT uses a 'bimodal test', that enumerates the number of loci different between each pair of samples, and a 'difference in capture history test' (DCH) to determine those loci producing the most errors. Importantly, the DCH test allows one to determine that a data set is error-free.

Download

DROPOUT is free software for the MS-Windows operating system. The software was developed by Kevin McKelvey. The download file is in ZIP format and includes documentation. If you download this software please contact Kevin with your name and affiliation.

Contained within DROPOUT are several programs that compute capture/mark/recapture statistics on genetic data. You can download them separately.

Related Publications

McKelvey, K. and Schwartz, M.K. (2005). DROPOUT: a program to identify problem loci and samples for noninvasive genetic samples in a capture-mark-recapture framework. Molecular Ecology Notes 5: 716-718.

Schwartz, M.K., Cushman, S.A., McKelvey, K., Hayden, J., and Engkjer, C. (2006). Detecting genotyping errors and describing American black bear movement in northern Idaho. Ursus 17(2):138-148.

McKelvey, K. and Schwartz, M.K. (2004). Providing reliable and accurate genetic mark-recapture in a cost-effective manner. Journal of Wildlife Management 68: 453-456.

McKelvey, K. and Schwartz, M.K. (2004). Genetic errors associated with population estimation using non-invasive molecular tagging: problems and new solutions. Journal of Wildlife Management 68: 439-448.

In the News

People

Engkjer, Cory L    Biological Science Technician    406-542-3255
McKelvey, Kevin S    Research Ecologist    406-542-4163
Pilgrim, Kristine L    Laboratory Supervisor    406-542-3255
Schwartz, Michael    Research Ecologist    406-542-4161

Selected Publications

Hansen, Michael M.; Olivieri, Isabelle; Waller, Donald M.; Nielsen, Einar E.; Allendorf, F. W.; Schwartz, M. K.; Baker, C. S.; Gregovich, D. P.; Jackson, J. A.; Kendall, K. C.; Laikre, L.; McKelvey, K.; Neel, M. C.; Ryman, N.; Short Bull, R.; Stetz, J. B.; Tallmon, D. A.; Vojta, C. D.; Waples, R. S. 2012. Monitoring adaptive genetic responses to environmental change. Molecular Ecology. doi: 10.1111/j.1365-294X.2011.05463.x.

Pilgrim, Kristine L.; Zielinski, William J.; Schlexer, Fredrick V.; Schwartz, Michael K. 2012. Development of a reliable method for determining sex for a primitive rodent, the Point Arena mountain beaver (Aplodontia rufa nigra). Conservation Genetics Resources. doi: 10.1007/s12686-012-9686-6.

Zielinski, W.J., F.Z. Schlexer, S.A. Parks, KL. Pilgrim and M.K. Schwartz 2012. Conserv Genet DOI 10.1007/s10592-012-0387-1.

Russell, Robin E.; Royle, J. Andrew; Desimone, Richard; Schwartz, Michael K.; Edwards, Victoria L.; Pilgrim, Kristy P.; McKelvey, Kevin S. 2012. Estimating abundance of mountain lions from unstructured spatial sampling. The Journal of Wildlife Management. doi: 10.1002/jwmg.412.

Campbell, N. R.; Amish, S. J.; Prichard, V. L.; McKelvey, K. M.; Young, M. K.; Schwartz, M. K.; Garza, J. C.; Luikart, G.; Narum, S. R. 2012. Development and evaluation of 200 novel SNP assays for population genetic studies of westslope cutthroat trout and genetic identification of related taxa. Molecular Ecology Resources. doi: 10.1111/j.1755-0998.2012.03161.x.

Short Bull, R. A.; Cushman, Samuel; Mace, R.; Chilton, T.; Kendall, K. C.; Landguth, E. L.; Schwartz, Michael; McKelvey, Kevin; Allendorf, Fred W.; Luikart, G. 2011. Why replication is important in landscape genetics: American black bear in the Rocky Mountains. Molecular Ecology. 20: 1092-1107.

Magoun, Audrey J.; Long, Clinton D.; Schwartz, Michael K.; Pilgrim, Kristine L.; Lowell, Richard E.; Valkenburg, Patrick. 2011. Integrating motion-detection cameras and hair snags for wolverine identification. The Journal of Wildlife Management. 75(3): 731-739.

Hare, M.P., L. Nunney, M.K. Schwartz, D. E. Ruzzante, M. Burford, R.S. Waples, K. Ruegg, and F. Palstra. 2011. Understanding and Estimating Effective Population Size for Practical Application in Marine Species Management. Conservation Biology, Volume 25, No. 3, 438-449.

Marucco, Francesca; Boitani, Luigi; Pletscher, Daniel H.; Schwartz, Michael K. 2011. Bridging the gaps between non-invasive genetic sampling and population parameter estimation. European Journal of Wildlife Research. 57:1-13.

Luikart, Gordon; Ryman, Nils; Tallmon, David A.; Schwartz, Michael K.; Allendorf, Fred W. 2010. Estimation of census and effective population sizes: the increasing usefulness of DNA-based approaches. Conservation Genetics. 11: 355-373.

Schwartz, Michael K.; Luikart, Gordon; McKelvey, Kevin S.; Cushman, Samuel A. 2010. Landscape genomics: A brief perspective [Chapter 9]. In: Cushman, S. A.; Huettmann, F., eds. Spatial complexity, informatics, and wildlife conservation. New York: Springer. p. 165-174.

McKelvey, Kevin S.; Cushman, Samuel A.; Schwartz, Michael K. 2009. Landscape genetics [Chapter 17]. In: Cushman, S. A.; Huettmann, F., eds. Spatial complexity, informatics, and wildlife conservation. New York: Springer. p. 313-328.

Landguth, E. L.; Cushman, S. A; Schwartz, M. K.; McKelvey, K. S.; Murphy, M.; Luikart, G. 2010. Quantifying the lag time to detect barriers in landscape genetics. Molecular Ecology. 19: 4179-4191.

Tallmon, David A.; Gregovich, Dave; Waples, Robin S.; Baker, C. Scott; Jackson, Jennifer; Taylor, Barbara L.; Archer, Eric; Martien, Karen K.; Allendorf, Fred W.; Schwartz, Michael K. 2010. When are genetic methods useful for estimating contemporary abundance and detecting population trends?. Molecular Ecology Resources. 10: 684-692.

Laikre, Linda; Schwartz, Michael K.; Waples, Robin S.; Ryman, Nils; Allendorf, F. W.; Baker, C. S.; Gregovich, D. P.; Hansen, M. M.; Jackson, J. A.; Kendall, K. C.; McKelvey, K.; Neel, M. C.; Olivieri, I.; Short Bull, R.; Stetz, J. B.; Tallmon, D. A.; Vojta, C. D.; Waller, D. M. 2010. Compromising genetic diversity in the wild: Unmonitored large-scale release of plants and animals. Trends in Ecology and Evolution. 25: 520-529.

Laikre, Linda; Allendorf, Fred W.; Aroner, Laurel C.; Baker, C. Scott; Gregovich, David P.; Hansen, Michael M.; Jackson, Jennifer A.; Kendall, Katherine C.; Mckelvey, Kevin; Neel, Maile C.; Olivieri, Isabelle; Ryman, Nils; Schwartz, Michael K.; Bull, Ruth Short; Stetz, Jeffrey B.; Tallmon, David A.; Taylor, Barbara L.; Vojta, Christina D.; Waller, Donald M.; Waples, Robin S. 2009. Neglect of genetic diversity in implementation of the Convention on Biological Diversity. Conservation Biology. 24(1): 86-88.

Brinkman, Todd J.; Person, David K.; Schwartz, Michael K.; Pilgrim, Kristine L.; Colson, Kevin E.; Hundertmark, Kris J. 2010. Individual identification of Sitka black-tailed deer (Odocoileus hemionus sitkensis) using DNA from fecal pellets. Conservation Genetics Resources. 2: 115-118.

Mullen, Lindy B.; Woods, H. Arthur; Schwartz, Michael K.; Sepulveda, Adam J.; Lowe, Winsor H. 2010. Scale-dependent genetic structure of the Idaho giant salamander (Dicamptodon aterrimus) in stream networks. Molecular Ecology. 19: 898-909.

Schwartz, M.K., and J.A. Vucetich. 2009. Molecules and beyond: assessing the distinctness of the Great Lakes wolf. Molecular Ecology 18:2307-2309. Molecular Ecology 18:2307-2309.

Moriarty, Katie M.; Zielinski, William J.; Gonzales, Armand G; Dawson, Todd E.; Boatner, Kristie M.; Wilson, Craig A; Schlexer, Frederick V.; Pilgrim, Kristine L.; Copeland, Jeffrey P.; Schwartz, Michael K. 2009. Wolverine confirmation in California after nearly a century: native or long-distance immigrant. Northwest Science 83(2): 154-162.

Schwartz, Michael K.; Copeland, Jeffrey P.; Anderson, Neil J.; Squires, John R.; Inman, Robert M.; Mckelvey, Kevin S.; Pilgrim, Kristy L.; Waits, Lisette P.; Cushman, Samuel A. 2009. Wolverine gene flow across a narrow climatic niche. Ecology. 90(11): 3222-3232.

Schwartz, M. K. 2009. Uniting ecological and genetic data for the conservation of wild ibex. Animal Conservation. 12: 103-104.

Marucco, Francesca; Pletscher, Daniel H.; Boitani, Luigi; Schwartz, Michael K.; Pilgrim, Kristy L.; Lebreton, Jean-Dominique. 2009. Wolf survival and population trend using non-invasive capture-recapture techniques in the Western Alps. Journal of Applied Ecology. doi: 10.1111/j.1365-2664.2009.01696.x_ 2009.

Beja-Pereira, Albano; Oliveira, Rita; Alves, Paulo C.; Schwartz, Michael K.; Luikart, Gordon. 2009. Advancing ecological understandings through technological transformations in noninvasive genetics. Molecular Ecology Resources. 9: 1279-1301.

Cushman, S.A., K.S. McKelvey, and M.K. Schwartz. 2009. Evaluating habitat connectivity and mapping of corridors between Yellowstone National Park and the Canadian Border with landscape genetics and least cost path analysis. Conservation Biology 23:368-376.

McKelvey, Kevin S.; Aubry, Keith B.; Schwartz, Michael K. 2008. Using anecdotal occurrence data for rare or elusive species: the illusion of reality and a call for evidentiary standards. BioScience. 58(6): 549-555.

Homyack, Jessica A.; Vashon, Jennifer H.; Libby, Cade; Lindquist, Edward L.; Loch, Steve; McAlpine, Donald F.; Pilgrim, Kristine L.; Schwartz, Michael K. 2008. Canada lynx-bobcat (Lynx canadensis x L. rufus) hybrids at the southern periphery of lynx range in Maine, Minnesota and New Brunswick. American Midland Naturalist. 159: 504-508.

Luikart, G., K. Pilgrim, J. Visty, V. Ezenwa, and M.K. Schwartz. 2008. Candidate gene microsatellite variation is associated with parasitism in wild bighorn sheep. Biology Letters 4:228-231.

Cushman, Samuel; McKelvey, Kevin; Schwartz, Michael. 2008. Case study 6.1: DNA survey for fisher in northern Idaho. In: Long, R. A.; MacKay, P.; Ray, J. C.; Zielinski, W. J., editors. Noninvasive survey methods for North American carnivores. Washington, D. C.: Island Press. p. 173-174.

Kendall, Katherine C.; McKelvey, Kevin S. 2008. Chapter 6: Hair collection. In: Long, R. A.; MacKay, P.; Ray, J. C.; Zielinski, W. J., editors. Noninvasive survey methods for North American carnivores. Washington, D. C.: Island Press. p. 135-176.

Schwartz, Michael K.; Monfort, Steven L. 2008. Genetic and endocrine tools for carnivore surveys. In: Long, Robert; MacKay, Paula; Ray, Justina; Zielinski, William, editors. Noninvasive survey methods for North American carnivores. Washington, D.C.: Island Press. p. 228-250.

Schwartz, Michael K.; McKelvey, Kevin S. 2008. Why sampling scheme matters: the effect of sampling scheme on landscape genetic results. Conservation Genetics. DOI 10.1007/s10592-008-9622-1.

Schwartz MK. 2007. Ancient DNA confirms native rocky mountain fisher Martes pennanti avoided early 20th century extinction. Journal of Mammalogy 88:921-925.

Jordan, Mark J. , J . Mark Higley, Sean M. Matthews, Olin E. Rhodes, Michael K. Schwartz, Reginald H. Barrett and Per J. Palsbll. 2007. Development of 22 new microsatellite loci for fishers (Martes pennanti) with variability results from across their range. Molecular Ecology Notes.

Schwartz MK, Luikart G, and Waples RS. 2007. Genetic monitoring as a promising tool for conservation and management. Trends in Ecology and Evolution 22: 25-33.

Schwartz, Michael K.; Aubry, Keith B.; McKelvey, Kevin S.; Pilgrim, Kristine L.; Copeland, Jeffrey P.; Squires, John R.; Inman, Robert M.; Wisely, Samantha M.; Ruggiero, Leonard F. 2007. Inferring geographic isolation of wolverines in California using historical DNA. Journal of Wildlife Management. 71(7): 2170-2179.

Compton, B., K. McGarigal, S.A. Cushman and L. Gamble. 2007. A resistant kernel model of connectivity for vernal pool breeding amphibians. Conservation Biology.

McKelvey, Kevin S.; von Kienast, Jeffrey; Aubry, Keith B.; Koehler, Gary M.; Maletzke, Bejamin T.; Squires, John R.; Lindquist, Edward L.; Loch, Steve; Schwartz, Michael K. 2006. DNA analysis of hair and scat collected along snow tracks to document the presence of Canada Lynx. Wildlife Society Bulletin. 34(2): 451-455.

Todd J. Ulizio, John R. Squires, Daniel H. Pletscher, Michael K. Schwartz, James J. Claar, and Leonard F. Ruggiero. 2006. The Efficacy of Obtaining Genetic-Based Identifications from Putative Wolverine Snow Tracks. Wildlife Society Bulletin 34:1326-1334.

Vinkey, Ray S.; Schwartz, Michael K.; McKelvey, Kevin S.; Foresman, Kerry R.; Pilgrim, Kristine L.; Giddings, Brian J.; Lofroth, Eric C. 2006. When reintroductions are augmentations: the genetic legacy of the fisher (Martes pennanti) in Montana. Journal of Mammalogy. 87(2): 265-271.

Zielinski, William J.; Schlexer, Fredrick V.; Pilgrim, Kristine L.; Schwartz, Michael K. 2006. The efficacy of wire and glue hair snares in identifying mesocarnivores. Wildlife Society Bulletin 34(4): 1152-1161.

Schwartz, Michael K.; Cushman, Samuel A.; McKelvey, Kevin S.; Hayden, Jim; Engkjer, Cory. 2006. Detecting genotyping errors and describing black bear movement in northern Idaho. Ursus. 17(20): 138-148.

Pilgrim, Kristine L.; Zielinski, William J.; Mazurek, Mary J.; Schlexer, Frederick V.; Schwartz, Michael K. 2006. Development and characterization of microsatellite markers in the Point Arena mountain beaver Aplodontia rufa nigra. Molecular Ecology Notes. 6(3): 800-802.

Roman Biek, Naomi Akamine, Michael K. Schwartz, Toni K. Ruth, Kerry M. Murphy and Mary Poss. 2006. Genetic consequences of sex-biased dispersal in a solitary carnivore: Yellowstone cougars. Biol. Lett. (2006) 2, 312-315.

Cushman, Samuel A.; McKelvey, Kevin S.; Hayden, Jim; Schwartz, Michael K. 2006. Gene flow in complex landscapes: Testing multiple hypotheses with causal modeling. American Naturalist 168(4): 486-499.

Schwartz, Michael K. 2005. Guidelines on the use of molecular genetics in reintroduction programs. In: The EU LIFE-Nature Projects to guidelines for the reintroduction of threatened species; Caramanico Terme, Italy; March 21-22, 2005. p. 51-58.

S. S. Loew, D. F. Williams, K. Ralls, K. Pilgrim and R. C. Fleischer. 2005. Population structure and genetic variation in the endangered Giant Kangaroo Rat (Dipodomys ingens). Conservation Genetics 6(4) 485-654.

Holthausen, Richard; Czaplewski, Raymond L.; DeLorenzo, Don; Hayward, Greg; Kessler, Winifred B.; Manley, Pat; McKelvey, Kevin S.; Powell, Douglas S.; Ruggiero, Leonard F.; Schwartz, Michael K.; Van Horne, Bea; Vojta, Christina D. 2005. Strategies for monitoring terrestrial animals and habitats. Gen. Tech. Rep. RMRS-GTR-161. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 34 p.

McKelvey KS and Schwartz MK. 2005. DROPOUT: a program to identify problem loci and samples for noninvasive genetic sampling in a capture-mark-recapture framework. Molecular Ecology Notes 5: 716-718.

Michael K. Schwartz, Katherine Ralls, Dan F. Williams, Brian L. Cypher, Kristine L. Pilgrim and Robert C. Fleischer. 2005. Gene flow among San Joaquin kit fox populations in a severely changed ecosystem. Conservation Genetics 6: 25-37.

Michael K. Schwartz L. Scott Mills. 2005. Gene flow after inbreeding leads to higher survival in deer mice. Biological Conservation 123 413-420.

K. L. Pilgrim, K . S. McKelvey, A . E. Riddle And M. K. Schwartz. 2005. Felid sex identification based on noninvasive genetic samples. Molecular Ecology Notes. 5, 60-61.

McKelvey, K. S. and M. K. Schwartz. 2004. Genetic Errors Associated with Population Estimation Using Non-Invasive Molecular Tagging: Problems and New Solutions. J. Wildl. Manage. 68(3):439-448.

Paul L. Leberg, Michael R. Carloss, Larry J. Dugas, Kristine L. Pilgrim, L. Scott Mills, M. Clay Green, and Daniel Scognamillo. 2004. Recent Record of a Cougar (Puma concolor) in Louisiana, with Notes on Diet, Based on Analysis of Fecal Materials. Southeastern Naturalist 3(4):653-658.

Michael K. Schwartz, Kristine L. Pilgrim, Kevin S. McKelvey, Edward L. Lindquist, James J. Claar, Steve Loch and Leonard F. Ruggiero. 2004. Hybridization between Canada lynx and bobcats: Genetic results and management implications. Conservation Genetics 5: 349-355.

Neil J. Gemmell, Michael K. Schwartz, and Bruce C. Robertson. 2004. Moa were many. Proc Biol Sci. 2004 Dec 7;271 Suppl 6:S430-2.

M. K. Schwartz, L . S . Mills, Y. Ortega, L. F. Ruggiero and F. W. Allendorf. 2003. Landscape location affects genetic variation of Canada lynx (Lynx canadensis). Molecular Ecology (2003) 12, 1807-1816.

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

Mills LS, Schwartz MK, Tallmon DA, and Lair KP. 2003. Measuring and interpreting changes in connectivity for mammals in coniferous forests. In: Zabel C, Anthony RG (eds.) Mammal community dynamics in Western coniferous forests: management and conservation in the new millennium. Island Press, Washington D.C., pp 587-613.

Manel, Stephanie; Schwartz, Michael K.; Luikart, Gordon; Taberlet, Pierre. 2003. Landscape genetics: combining landscape ecology and population genetics. Trends in Ecology and Evolution. 18(4): 189-197.

Michael K. Schwartz, L. Scott Mills, Kevin S. McKelvey. 2002. DNA reveals high dispersal synchronizing the population dynamics of Canada lynx. Nature Vol 415, 520-522.

P. F. Kukuk, S . H. Forbes, R. Zahorchack, A. Riddle And K. Pilgrim. 2002. Highly polymorphic microsatellite markers developed for the social halictine bee Lasioglossum (Chilalictus) hemichalceum. Molecular Ecology Notes 2 529-530.

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(3):285-288.

Spruell, P., K. L. Pilgrim, B. A. Greene, C. Habicht, K. L. Knudsen, K. R. Lindner, G. K. Sage, J. E. Seeb, and F. W. Allendorf. 1999. Inheritance of nuclear DNA markers in gynogenetic haploid pink salmon. Journal of Heredity 90:289-296.

Schwartz MK, Tallmon DA, and Luikart G. 1999. Using genetics to estimate the size of wild populations: many methods, much potential, uncertain utility. Animal Conservation 2: 320-322.

Michael K. Schwartz, David A. Tallmon and Gordon Luikart. 1998. Review of DNA-based census and effective population size estimators. Animal Conservation (1998) 1, 293-299.

Pilgrim, K. P., S. H. Forbes and D. K. Boyd. 1998. Testing for wolf-coyote hybridization in the Rocky Mountains using mitochondrial DNA. Journal of Wildlife Management 62(2):683-689.