https://cpw.cvlcollections.org/files/original/378677b262fc46f11c892f19c12903c3.pdf 37febb164d3f9516af94ea7525a6c6bc PDF Text Text The research in this publication was partially or fully funded by Colorado Parks and Wildlife. Dan Prenzlow, Director, Colorado Parks and Wildlife • Parks and Wildlife Commission: Marvin McDaniel, Chair • Carrie Besnette Hauser, Vice-Chair Marie Haskett, Secretary • Taishya Adams • Betsy Blecha • Charles Garcia • Dallas May • Duke Phillips, IV • Luke B. Schafer • James Jay Tutchton • Eden Vardy �Does a Population of Cougars Exist in Michigan? Authors: Kurta, Allen, SCHWARTZ, MICHAEL K., and ANDERSON, CHARLES R. Source: The American Midland Naturalist, 158(2) : 467-471 Published By: University of Notre Dame URL: https://doi.org/10.1674/00030031(2007)158[467:DAPOCE]2.0.CO;2 BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access titles in the biological, ecological, and environmental sciences published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Complete website, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/terms-of-use. Usage of BioOne Complete content is strictly limited to personal, educational, and non - commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Downloaded From: https://bioone.org/journals/The-American-Midland-Naturalist on 22 Jul 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Colorado Parks and Wildlife �Am. Midl. Nat. 158:467–471 Does a Population of Cougars Exist in Michigan? ABSTRACT.—After analyzing DNA obtained from fecal samples gathered in Michigan, Swanson and Rusz (2006) claimed that 83% of identified scats were from cougars, indicating to them that a population of these large carnivores existed in the state. In this paper, we identify problems with their methodology, suggest that they unreasonably extrapolated their conclusions and point out that their results are improbable, especially in light of no other evidence in the scientific literature suggesting the existence of a population of cougars in Michigan. INTRODUCTION The cougar (Puma concolor) historically was the most widely distributed terrestrial mammal in the New World, other than humans (Biek et al., 2006). During the late 19th and early 20th centuries, North American populations of cougars declined substantially due to human persecution, habitat loss and depletion of prey, especially wild ungulates (Culver et al., 2000; Logan and Sweanor, 2001; Biek et al., 2006). The species became extinct throughout most of the East, although this large felid continued to live in many parts of the West. Since the 1970s, western populations appeared to have increased in size (Logan and Sweanor, 2000; Biek et al., 2006) and recolonization of some historically occupied regions has occurred (Novak et al., 1987; Linnell et al., 2001). In Michigan, most biologists believed that the cougar was extirpated from the Lower Peninsula by 1875 and from the more rugged Upper Peninsula by the early 1900s (Baker, 1983; Kurta, 1995). Nevertheless, there has been a long history of unsubstantiated reports of cougars in the state (Baker, 1983; R. H. Baker, in litt.), with many accounts appearing in the popular media during the last 10–15 y, suggesting to some people that this felid may be recolonizing Michigan or that a persistent relict population exists (Belant et al., 2006). Recently, Swanson and Rusz (2006) published what they believed was strong physical evidence, based on DNA obtained from fecal samples, for a population of at least eight cougars in Michigan between 2001 and 2003. The subspecies of cougar that originally was found in Michigan (Puma concolor couguar) is on the federal list of endangered species, and presence of a population of these animals would have profound management implications for state and federal agencies and how they spend limited conservation resources. More importantly, the claims of Swanson and Rusz have contributed to highly publicized fears for public safety (e.g., Donnelly, 2007). Herein, we describe several concerns that we have regarding the methodology and conclusions of Swanson and Rusz (2006) and address the improbability of their results. CRITIQUE Problems with genetic methodologies.—Swanson and Rusz (2006) identify species from scat using a modification of a method published by Foran et al. (1997). This method attempts to examine a segment of DNA that is quite large (ca. 700 base pairs). Unfortunately, large segments of DNA from noninvasive genetic samples, such as scat, rarely are amplified accurately because of the high level of degradation typically associated with these types of samples (Mills et al., 2000). The method of Foran et al. (1997) also relies on a region of the genome that contains an 80-bp tandem repeat unit (Lopez et al., 1996; Foran et al., 1997). Mills et al. (2000), in contrast, describe a newer method for identifying carnivores that examines a much shorter segment of DNA (ca. 360 base pairs) and uses the 16s rRNA region, a section of the genome without the 80-bp tandem repeat. In fact, the motivation behind developing the 16s-rRNA test in Mills et al. (2000) was failure of the method of Foran et al. (1997) to produce consistent results. In the Wildlife Genetics Laboratory of USDA Forest Service, Rocky Mountain Research Station, amplification of DNA from the large region of the genome containing the tandem repeat unit (i.e., the method of Foran et al., 1997) has a low rate of success and potentially yields misleading results, possibly because the number of repeat units varies in different individuals of the same species (K. Pilgrim, pers. comm.). Reliance on the method of Foran et al. (1997) likely explains the low success rate (4%) that Swanson and Rusz (2006) report for amplifying DNA from fecal samples. 467 Downloaded From: https://bioone.org/journals/The-American-Midland-Naturalist on 22 Jul 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Colorado Parks and Wildlife �468 THE AMERICAN MIDLAND NATURALIST 158(2) Furthermore, Swanson and Rusz (2006) do not properly apply the approach of Foran et al. (1997). The first step in the technique of Foran et al. (1997) is to amplify DNA, and the second step is to cut the DNA with diagnostic restriction enzymes, which is the same approach used by Mills et al. (2000). Both methods also require a panel of multiple restriction enzymes to produce diagnostic results for felids. The technique of Foran et al. (1997) requires use of two restriction enzymes to distinguish among the four felids found in the northern United States: bobcat (Lynx rufus), Canada lynx (L. canadensis), cougar and domestic cat (Felis catus). Swanson and Rusz (2006), however, only used one restriction enzyme (RsaI). Although the original primers used by Foran et al. (1997) would distinguish bobcat from cougar based on RsaI, they would fail to distinguish cougar from lynx (Foran et al., 1997:837–838). To distinguish lynx from cougar, Swanson and Rusz (2006) should have used the restriction enzyme HinfI. Note the absence of a sample from lynx as a control in figure 2 of Swanson and Rusz (2006). It is possible, but unlikely, that the modified primers used by Swanson and Rusz (2006) could separate lynx from cougars, but this never was demonstrated. The authors may have assumed that lynx do not exist in the area or that scat from lynx and cougar can be identified reliably using morphology; if so, they are incorrect. Although lynx are rare in Michigan, there are many verified records of lynx in the Upper Peninsula since the early 1900s (Harger, 1965; Baker, 1983; McKelvey et al., 2000; Beyer et al., 2001), with a confirmed lynx collected by a trapper in the Hiawatha National Forest in 2003 (S. Mighton, pers. comm.). We also question whether it is possible to distinguish reliably scats of lynx from those of cougar based on morphology because suspected cougar scats sent to the Wildlife Genetics Laboratory often are determined to be those of lynx based on either sequencing of DNA or the restriction-digest test of Mills et al. (2000; K. Pilgrim, pers. comm.). We have two other concerns regarding images of gels presented by Swanson and Rusz (2006). First, in theory, the method of Foran et al. (1997), when applied properly, differentiates domestic cats from cougars based on examining the RsaI restriction digest. However, we have found, in practice, that unless positive standards (‘‘knowns’’) for domestic cat and cougar are run simultaneously on a gel, it would be difficult to distinguish these species. Nevertheless, Swanson and Rusz (2006) only used standards from ‘‘three known bobcats, cougars, wolves, and coyotes’’ and not domestic cats. Second, RsaI-restricted samples from bobcats produce only a single band (Foran et al.,1997), yet figure 2b in Swanson and Rusz (2006) clearly shows two lower bands in the same size range as those of cougar. These extra lower bands for the bobcat have various possible causes, including contamination, but their unexplained presence in the gels of Swanson and Rusz (2006) is troubling. Extrapolating beyond supported results.—Swanson and Rusz (2006:370) claim positive identification of cougars from 10 samples gathered at eight sites, and conclude that there were ‘‘at least eight [cougars] in Michigan over the 3 y of this study.’’ They base this statement on the fact that the eight sites were separated by at least 56 km and the assumption that it was unlikely, given limited searching for scats in the field, that ‘‘we could have found scat dropped by the same cougar (dispersing or resident) at any two locations’’ (Swanson and Rusz, 2006:369). However, size of home range for male cougars typically is 200–500 km2, with some ranges as large as 792 km2 (Logan and Sweanor, 2000). In addition, young cougars dispersing from their natal home often travel more than 100 km, and distances over 1000 km have been documented (Logan and Sweanor, 2000, 2001; Thompson and Jenks, 2005). Furthermore, samples that Swanson and Rusz (2006) identified as cougar in Michigan were obtained over 3 y. We contend that distance between samples cannot be used reliably to infer separate identity and minimum number alive, given the time over which the samples were collected and the documented ability of cougars to move hundreds of kilometers. In addition, Swanson and Rusz (2006:369) refer to these eight putative cougars as the ‘‘Michigan population,’’ but there is no evidence that these animals, if they were cougars, were part of a resident population, as opposed to individuals dispersing from other states or recently released captives. There are well-established molecular methods (e.g., analysis of microsatellites) that can determine individual identity from scats produced by cougars (e.g., Ernest et al., 2000; Sinclair et al., 2001; Anderson et al., 2004; McRae et al., 2005; Biek et al., 2006). If Swanson and Rusz (2006) had run microsatellites on their 10 positive scats, then they could have made some inferences as to the minimum number of cougars alive in Michigan. In addition, some of these microsatellites produce allele lengths in a distribution that is species-specific in parts of the cougar’s geographic range (see fig. 2 in Ernest et al., Downloaded From: https://bioone.org/journals/The-American-Midland-Naturalist on 22 Jul 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Colorado Parks and Wildlife �2007 NOTES AND DISCUSSION 469 2000). Thus, using microsatellites could have confirmed the identity of species that Swanson and Rusz (2006) obtained from mitochondrial DNA. Lastly, there are felid-specific tests to determine gender that have been designed for use with scats and hairs (Pilgrim et al., 2005), and these tests could have been used to determine if both males and females were present in the ‘‘population.’’ Overall, without employing such tests and the previously mentioned problems with the methodology that they did use, a more cautious interpretation of the data is warranted. Improbable results.—Swanson and Rusz (2006) succeeded in amplifying DNA from 12 of 297 scats (4% of samples examined), with 10 samples attributed to cougars (83% of identified samples), one to bobcat, and one to a canid. The rate of success for amplifying DNA likely would have been much higher had they employed methods outlined in Mills et al. (2000). If Swanson and Rusz (2006) obtained identifications for 90% of their 297 scats (a rate achieved by McKelvey et al., 2006 in another study of wild felids) and 83% were cougars, Swanson and Rusz (2006) would have identified 223 samples as cougar. The high proportion of cougars among the samples identified by Swanson and Rusz (2006) and the absolute number of cougars that presumably would result if more samples actually had yielded amplified DNA seem improbable—especially without any supporting evidence in the scientific literature that a population occurs in the state. Cougars often are killed by trains, trucks or automobiles (Logan and Sweanor, 2000; Heist et al., 2001), but there is no report in the scientific literature that such a deadly collision ever has occurred in Michigan. Hunting is a significant cause of mortality for cougars, even in states where they are protected (Logan and Sweanor, 2000), yet there is no verifiable record of a cougar being shot in Michigan since the early 1900s (Baker, 1983). Similarly, cougars are caught inadvertently in traps set for other types of mammals (e.g., Wyoming Game and Fish Department, 2006), but there is no record of this happening in Michigan since 1907 (Baker, 1983). Personnel of the Michigan Department of Natural Resources annually perform winter track surveys in various parts of the Upper Peninsula, and tracks of this wideranging species never have been seen (e.g., Earle and Tuovila, 2002). Unverified reports of cougar were rampant near Sleeping Bear Dunes National Lakeshore in the Lower Peninsula, yet no cougars were found despite extensive track surveys, use of remote camera systems, baiting and thorough investigation of reported sightings (Belant et al., 2006). The existence of such consistent disconfirmatory evidence over the past century is difficult to reconcile with the claim of Swanson and Rusz (2006) that a population of cougars exists in Michigan. We admit that detection of cryptic species can occur, especially in remote areas, but we advocate that, upon finding such an odd result, initial tests be confirmed with a second method or in another laboratory. CONCLUSIONS Overall, we are skeptical of the results of Swanson and Rusz (2006). Given the lack of any direct testing of and control lanes for domestic cats and Canada lynx, failure to follow Foran et al.’s (1997) protocol and lack of confirmatory tests, we believe that their claim to have identified multiple samples as being produced by cougars is unsubstantiated. Inferences that there are at least eight individuals and that these individuals are part of a population are pure speculation. Nevertheless, we agree that the authors’ description of a partial sequence from a scat found in Delta County in 2001 (Swanson and Rusz, 2006) appears consistent with haplotype M of cougars (Culver et al., 2000). Thus, all that we would feel comfortable in stating, based on Swanson and Rusz (2006), is that the number of cougars documented in the scientific literature in Michigan since the early 1900s is now one. Whether that sample represents a released pet, a dispersing individual or a planted sample is not something that we can assess. Acknowledgments.—We thank V. Drietz, K. Logan, K. Pilgrim, K. McKelvey and H. Shaw for reviewing the manuscript and R. H. Baker for providing historical insight into the question of cougars in Michigan. LITERATURE CITED ANDERSON, C. R., JR., F. G. LINDZEY AND D. B. MCDONALD. 2004. Genetic structure of cougar populations across the Wyoming Basin: metapopulation or megapopulation. J. Mammal, 85:1207–1214. Downloaded From: https://bioone.org/journals/The-American-Midland-Naturalist on 22 Jul 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Colorado Parks and Wildlife �470 THE AMERICAN MIDLAND NATURALIST 158(2) BAKER, R. H. 1983. Michigan mammals. Michigan State University Press, East Lansing. 642 p. BELANT, J. L., S. E. YANCHO AND K. S. STRUTHERS. 2006. Does the cougar inhabit Sleeping Bear Dunes National Lakeshore? Nat. Areas J., 26:370–375. BEYER, D. E., B. J. ROELL, J. H. HAMMILL AND R. D. EARLE. 2001. Records of Canada lynx, Lynx canadensis, in the Upper Peninsula of Michigan. Can. Field-Nat., 115:234–240. BIEK, R., A. J. DRUMMOND AND M. POSS. 2006. A virus reveals population structure and recent demographic history of its carnivore host. Science, 311:538–541. CULVER, M., W. E. JOHNSON, J. PECON-SLATTERY AND S. J. O’BRIEN. 2000. Genomic ancestry of the American puma (Puma concolor). J. Hered., 91:186–197. DONNELLY, F. X. 2007. Roar of Michigan cougar debate grows louder. The Detroit News, 27 January 2007. http://www.detnews.com/apps/pbcs.dll/article?aid5/20070127/metro/701270374/1003. Accessed 7 February 2007. EARLE, R. D. AND V. R. TUOVILA. 2003. Furbearer winter track survey count for 2002. Mich. Dept. Nat. Res., Wildl. Rept., 3397:1–10. ERNEST, H. B., M. C. PENEDO, P. B. MAY, M. SYVANEN AND W. M. BOYCE. 2000. Molecular tracking of mountain lions in the Yosemite Valley region in California: genetic analysis using microsatellites and fecal DNA. Mol. Ecol., 9:433–441. FORAN, D. R., K. R. CROOKS AND S. C. MINTA. 1997. Species identification from scat: an unambiguous genetic method. Wildl. Soc. Bull., 25:835–839. HARGER, E. M. 1965. The status of the Canada lynx in Michigan. Jack-Pine Warbler, 43:150–153. HEIST, E. J., J. R. BOWLES AND A. WOOLF. 2001. Record of a North American cougar (Puma concolor) from southern Illinois. Trans. Ill. State Acad. Sci., 94:227–229. KURTA, A. 1995. Mammals of the Great Lakes region. University of Michigan Press, Ann Arbor. 376 p. LINNELL, J. D. C., J. E. SWENSON AND R. ANDERSEN. 2001. Predators and people: conservation of large carnivores is possible at high human densities if management policy is favourable. Anim. Conserv., 4:345–349. LOGAN, K. A. AND L. L. SWEANOR. 2000. Puma, p. 347–377. In: S. Demarais and P. Krausman (eds.). Ecology and management of large mammals in North America. Prentice Hall, Englewood Cliffs, New Jersey. 778 p. ——— AND ———. 2001. Desert puma: evolutionary ecology and conservation of an enduring carnivore. Island Press, Washington, D.C. 390 p. LOPEZ, J. V., S. CEVARIO AND S. J. O’BRIEN. 1996. Complete nucleotide sequences of the domestic cat (Felis catus) mitochondrial genome and a transposed mtDNA tandem repeat (Numt) in the nuclear genome. Genomics, 33:229–246. MCKELVEY, K. S., K. B. AUBRY AND Y. K. ORTEGA. 2000. History and distribution of lynx in the contiguous United States, p. 207–259. In: L. F. Ruggiero, K. B. Aubry, S. F. Buskirk, C. J. Krebs, K. S. McKelvey and J. R. Squires (eds.). The ecology and conservation of lynx. University of Colorado Press, Denver. 480 p. ———, J. vON KIENAST, K. B. AUBRY, G. M. KOEHLER, B. T. MALETZKE, J. R. SQUIRES, E. L. LINDQUIST, S. LOCH AND M. K. SCHWARTZ. 2006. DNA analysis of hair and scat collected along snow tracks to document the presence of Canada lynx. Wildl. Soc. Bull., 34:451–455. MCRAE, B. H., P. BEIER, L. E. DEWALD, L. Y. HUYNH AND P. KEIM. 2005. Habitat barriers limit gene flow and illuminate historical events in a wide-ranging carnivore, the American puma. Mol. Ecol., 14:1965–1977. MILLS, L. S., K. L. PILGRIM, M. K. SCHWARTZ AND K. S. MCKELVEY. 2000. Identifying lynx and other North American felids based on mtDNA analysis. Conserv. Gen., 1:258–288. NOVAK, M., J. A. BAKER, M. E. OBBARD AND B. MALLOCH. 1987. Wild furbearer management and conservation in North America. Ontario Ministry of Natural Resources, Toronto, Canada. 1150 p. PILGRIM, K. L., K. S. MCKELVEY, A. E. RIDDLE AND M. K. SCHWARTZ. 2005. Felid sex identification based on noninvasive genetic samples. Mol. Ecol. Notes, 5:60–61. SCHWARTZ, M. K., G. LUIKART AND R. S. WAPLES. 2007. Genetic monitoring of individuals, populations, and species in the wild. Trends Ecol. Evol., 22:25–33. Downloaded From: https://bioone.org/journals/The-American-Midland-Naturalist on 22 Jul 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Colorado Parks and Wildlife �2007 NOTES AND DISCUSSION 471 SINCLAIR, E. A., E. L. SWENSON, M. L. WOLFE, D. C. CHOATE, B. BATES AND K. A. CRANDALL. 2001. Gene flow estimates in Utah’s cougars imply management beyond Utah. Anim. Conserv., 4:257–264. SWANSON, B. J. AND P. J. RUSZ. 2006. Detection and classification of cougars in Michigan using low copy DNA sources. Am. Midl. Nat., 155:363–372. THOMPSON, D. J. AND J. A. JENKS. 2005. Long-distance dispersal by a subadult male cougar (Puma concolor) from the Black Hills, South Dakota. J. Wildl. Manage., 69:818–820. WYOMING GAME AND FISH DEPARTMENT. 2006. Wyoming mountain lion management plan. Wyoming Game and Fish Department, Cheyenne. 45 p. ALLEN KURTA, Department of Biology, Eastern Michigan University, Ypsilanti 48197; MICHAEL K. SCHWARTZ, USDA Forest Service, Rocky Mountain Research Station, 800 E. Beckwith Avenue, Missoula, Montana 59801 and CHARLES R. ANDERSON, JR. Colorado Division of Wildlife, 317 W. Prospect, Ft. Collins 80526. Submitted 9 February 2007; Accepted 11 April 2007. Downloaded From: https://bioone.org/journals/The-American-Midland-Naturalist on 22 Jul 2021 Terms of Use: https://bioone.org/terms-of-use Access provided by Colorado Parks and Wildlife � Dublin Core The Dublin Core metadata element set is common to all Omeka records, including items, files, and collections. For more information see, http://dublincore.org/documents/dces/. Title A name given to the resource Journal Articles Description An account of the resource CPW peer-reviewed journal publications Text A resource consisting primarily of words for reading. Examples include books, letters, dissertations, poems, newspapers, articles, archives of mailing lists. Note that facsimiles or images of texts are still of the genre Text. Dublin Core The Dublin Core metadata element set is common to all Omeka records, including items, files, and collections. For more information see, http://dublincore.org/documents/dces/. Title A name given to the resource Does a population of cougars exist in Michigan? Description An account of the resource <span>After analyzing DNA obtained from fecal samples gathered in Michigan, </span><a class="internal-link" href="https://bioone.org/journals/the-american-midland-naturalist/volume-158/issue-2/0003-0031(2007)158%5B467%3ADAPOCE%5D2.0.CO%3B2/Does-a-Population-of-Cougars-Exist-in-Michigan/10.1674/0003-0031(2007)158%5B467:DAPOCE%5D2.0.CO;2.short#i0003-0031-158-2-467-Swanson1">Swanson and Rusz (2006)</a><span> claimed that 83% of identified scats were from cougars, indicating to them that a population of these large carnivores existed in the state. In this paper, we identify problems with their methodology, suggest that they unreasonably extrapolated their conclusions and point out that their results are improbable, especially in light of no other evidence in the scientific literature suggesting the existence of a population of cougars in Michigan.</span> Bibliographic Citation A bibliographic reference for the resource. Recommended practice is to include sufficient bibliographic detail to identify the resource as unambiguously as possible. Kurta, A., M. K. Schwartz, and C. R. Anderson Jr. 2007. Does a population of cougars exist in Michigan? The American Midland Naturalist 158:467–471. <a href="https://doi.org/10.1674/0003-0031(2007)158%5B467:DAPOCE%5D2.0.CO;2" target="_blank" rel="noreferrer noopener">https://doi.org/10.1674/0003-0031(2007)158[467:DAPOCE]2.0.CO;2</a> Creator An entity primarily responsible for making the resource Kurta, Allen Schwartz, Michael K. Anderson Jr, Charles R. Subject The topic of the resource Cougar Michigan DNA Distribution Extent The size or duration of the resource. 6 pages Date Created Date of creation of the resource. 2007-10-01 Rights Information about rights held in and over the resource <a href="http://rightsstatements.org/vocab/InC-NC/1.0/" target="_blank" rel="noreferrer noopener">In Copyright - Non-Commercial Use Permitted</a> Format The file format, physical medium, or dimensions of the resource application/pdf Language A language of the resource English Is Part Of A related resource in which the described resource is physically or logically included. The American Midland Naturalist Type The nature or genre of the resource Article