NORTH CAROLINA MUSEUM of NATURAL SCIENCES FOURTH COLLOQUIUM ON CONSERVATION OF MAMMALS IN THE SOUTHEASTERN UNITED STATES Edited by BRIAN R. CHAPMAN and JOSHUA LAERM Occasional Papers of the North Carolina Museum of Natural Sciences and the North Carolina Biological Survey Number 12 Fall, 2000 NORTH CAROLINA STATE MUSEUM OF NATURAL SCIENCES FOUNDED 1879 MISSION The purpose of the North Carolina State Museum of Natural Sciences is to enhance the public’s understanding and appreciation of the natural environment in ways that emphasize the biodiversity of North Carolina and the southeastern United States and relate the region to the natural world as a whole. NORTH CAROLINA BIOLOGICAL SURVEY FOUNDED 1976 Member of The Consortium of State Biological Surveys MISSION The mission of the North Carolina Biological Survey is to acquire, organize, preserve, and interpret the natural his¬ tory collections and data associated with the state’s biological resources housed at the North Carolina State Museum of Natural Sciences. OCCASIONAL PAPERS OF THE NORTH CAROLINA MUSEUM OF NATURAL SCIENCES AND THE NORTH CAROLINA BIOLOGICAL SURVEY Occasional Papers publishes manuscripts of 100 or more manuscript pages plus figures (no color) and tables. Suitable topics include geology, paleontology, and systematics and natural history topics in the fields of ichthyology, herpetology, ornithology, and mammalogy, or regarding non-insect terrestrial or aquatic invertebrates. In general, subject matter should be compatible with the mission of the NC Museum of Natural Sciences. Suitability should be determined prior to submitting manuscripts to the managing editor for scientific publications [(919) 733-7450 x701]. Prior to publication, all manuscripts must be considered acceptable by a peer-review board comprised of not more than three external reviewers. Stephen D. Busack, Ph.D. Director, Research and Collections Managing Editor, Scientific Publications ([email protected]) © 2000 North Carolina State Museum of Natural Sciences ISBN 0-917134-17-0 NORTH CAROLINA MUSEUM Of NATURAL SCIENCES Fourth Colloquium on Conservation of Mammals in the Southeastern United States Edited by BRIAN R. CHAPMAN and JOSHUA LAERM Occasional Papers of the North Carolina Museum of Natural Sciences and the North Carolina Biological Survey Number 12 Fall, 2000 Forward Mammals have been the focus of intensive scientific representing different agencies or institutions to meet, get inquiry and much is known of the life-history attributes acquainted, and exchange information and ideas concern¬ of many species. However, recent environmental concerns ing mammalian research and conservation priorities. and new legislation requiring environmental assessments The 1994 Colloquium was held at the University of have brought demands for more detailed information on Georgia and was sponsored jointly by the Institute of the systematics, population status, and habitat require¬ Ecology and the Daniel B. Warned School of Forest ments of mammals. It will take the concerted efforts of Resources. The scientific program consisted of 19 con¬ professional mammalogists from federal and state agen¬ tributed papers and two posters. The presentations pri¬ cies, academic institutions, and the private sector to marily focused on two areas—the status of selected taxa establish priorities and meet the mammalian research and (especially of protected or little-known species), and conservation needs that society requires. aspects of life history. This symposium volume is dedi¬ The “Colloquium” is a loosely organized group of cated to the men and women who initiated and fostered professional scientists from the southeastern United communication and research on mammals in the south¬ States that has interest in conservation and management eastern United States, to those who have recently added of mammals in the region. The group first met at the enthusiasm and skill to the crusade, and to those who will University of Memphis in 1991 and has met annually contribute to future progress in the conservation of mam¬ since then. The Colloquium exists to provide individuals mal populations and habitats. Brian R. Chapman Acknowledgments This publication represents the results of a colloqui¬ We thank the 20 anonymous referees who provided um convened in Athens, Georgia, in February 1994, as excellent reviews of the manuscripts. We express our the Fourth Colloquium on the Conservation of Mammals appreciation to J. F. Boone who provided critical editorial in the Southeastern United States. This colloquium was advice on several manuscripts. We also thank the many made possible by funding and logistical support provided authors who responded well to advice and criticisms of by the Institute of Ecology and the Daniel B. Wamell their work and who waited patiently for their papers to School of Forest Resources of the University of Georgia. appear in print. The Editors: Brian R. Chapman, Joshua Faerm. Fourth Colloquium on Conservation of Mammals in the Southeastern United States Edited by Brian R. Chapman and Joshua Laerm Contents Colloquium on conservation of mammals in the southeastern United States.1 Michael L. Kennedy Conservation status of terrestrial mammals of the southeastern United States,.4 Joshua Laerm, W. Mark Ford, and Brian R. Chapman Analysis of distribution and habitat associations of Sorex hoyi winnemana in the southern Appalachians.17 Joshua Laerm, W. Mark Ford, Michael A. Menzel, and Timothy S. McCoy Radiotelemetric assessment of movement patterns of the gray bat (Myotis grisescens).27 at Guntersville Reservoir, Alabama, David P. Thomas and Troy L. Best Distribution and status of lasiurine bats in Georgia.40 Brian R. Chapman, Joshua Laerm, and Sandra S. Chapman Distribution and abundance of bats in caves and mines of northeastern Mississippi.45 Troy L. Best and Katherine G. Caesar Body size, reproductive biology, and sex ratio of a year-round colony of Eptesicus fuscus fuscus and.50 Tadarida brasiliensis cynocephala in eastern Alabama, Travis H. Henry, Troy L. Best, and Clayton D. Hilton Gastrointestinal helminth parasites of bats in Alabama, Clayton D. Hilton and Troy L. Best.57 Distribution and status of LeConte’s free-tailed bat (Tadarida brasiliensis cynocephala) in Alabama.67 W. Mark Kiser Ecology and conservation of a frontier population of the round-tailed muskrat (Neofiber alleni).74 Bradley J. Bergstrom, Tim Farley, Harvey L. Hill, Jr., and Tip Hon Population dynamics of the beach mouse (Peromyscus polionotus trissyllepsis): a simulation study.83 to examine extinction probabilities, Mahesh Sankaran, Nicholas R. Holler, and Michael C. Wooten Kennedy—Colloquium on Conservation of Mammals Colloquium on Conservation of Mammals in the Southeastern United States Michael L. Kennedy Department of Biology, The University of Memphis, Memphis, TN 38152 The Colloquium on Conservation of Mammals in the sponsored by the Edward J. Meeman Biological Station, Southeastern United States stemmed from an interest to College of Arts and Sciences, and Office of Public foster communications and research on mammals by pro¬ Service at The University of Memphis. The meeting was fessional biologists working in the Southeast. Earliest dis¬ hosted by M. L. Kennedy, and, for this first meeting, cussions of such a conference were held informally by T. papers and participants (39 attended) were invited. L. Best, G. A. Heidt, M. L. Kennedy, P. K. Kennedy, and Following an official welcome by Associate Dean H. D. V. R. McDaniel during the 1980s. The number of profes¬ Black, J. K. Jones, Jr., spoke during the Plenary Session. sional mammalogists and biologists conducting research His talk was entitled “Concerns for Mammalian and either charged or interested in the conservation of Biodiversity and Bioconservation in the South-central mammalian resources in the southeastern states grew States.” G. A. Heidt, V. R. McDaniel, and J. P. Nelson, Jr., significantly from the 1950s and 1960s to the 1980s. served as chairpersons for three paper sessions. Paper ses¬ There was a feeling among the early organizers of the sions were followed by a panel discussion entitled Colloquium of a need for better communication among “Priorities in Mammalian Bioconservation in the South- biologists working in this region. A desire of the early central States.” Panel members included V. R. McDaniel organizers was to bring individuals from academic insti¬ (Chair), J. K. Jones, Jr., T. L. Best, and G. A. Heidt. The tutions, state and federal agencies, and companies in the meeting concluded with a wrap up and discussion of private sector together to discuss conservation issues of future plans. Participants voiced a “successful meeting” significance to people in the region, as well as to estab¬ and voted to make the meeting an annual event. Informal lish needs and priorities (if possible) for scientific study. discussions tended to indicate that the best time for the A unique feature of this meeting, from the start, was a annual meeting would be in February. It was decided that gathering of professionals from different disciplines with the second meeting should be in northern Alabama (to be mutual interests in mammals. There was feeling among hosted by T. L. Best, Auburn University). the early organizers that communication among agencies, The Second Colloquium on Conservation of as well as among individuals, was a key to success relat¬ Mammals in the South-central United States was held at ing to mammalian conservation in the Southeast. The Guntersville State Park in northern Alabama on 20-21 original region of interest was limited to the south-central February 1992. The meeting was sponsored by the states (Alabama, Arkansas, Kentucky, Louisiana, Department of Zoology and Wildlife Science at Auburn Mississippi, Missouri, Tennessee), and, initially, the University. A pre-meeting social during the evening of the loosely structured organization was referred to as the 20th was well attended. Forty-eight participants attended Colloquium on Conservation of Mammals in the South- the meeting on the 21st. J. F. Pritchett (Head, Department central States. At the 1994 meeting, the geographic region of Zoology and Wildlife Science) extended the official was expanded to include all of the southeastern United welcome. The Plenary Session was a paper entitled States and the organization named appropriately “Problems in the Conservation of Small Mammals: Beach (Colloquium on Conservation of Mammals in the Mice as an Example” presented by N. R. Holler. A panel Southeastern United States). A brief history of the discussion on “Problems and Prospects for Conservation Colloquium is presented in the following paragraphs. of Mammals in the South-central United States” fol¬ On 17 May 1991, the First Colloquium on lowed. Members of the panel included M. L. Kennedy Conservation of Mammals in the South-central United (Chair), R. K. Chesser, N. R. Holler, R. Jordan, V. R. States was held at The University of Memphis' Edward J. McDaniel, and M. C. Wooten. The remainder of the for¬ Meeman Biological Station in Shelby Co., Tennessee. mal program included one poster and 12 paper presenta¬ The meeting focused on conservation biology and was tions. This was another very successful meeting. The Kennedy—Colloquium on Conservation of Mammals interaction of individuals from different institutions and States was held at the Executive Inn in Cookeville, agencies was very high. It was decided that the third Tennessee. The meeting was sponsored by the meeting would be in Arkansas (to be hosted by G. A. Department of Biology at Tennessee Tech University. Heidt, University of Arkansas at Little Rock, and V. R. Following the informal reception on the 24th, M. J. McDaniel, Arkansas State University). A very successful Harvey extended an official welcome on the morning of field trip (led by T. L. Best) to Blowing Wind Cave fol¬ the 25th. Three paper sessions (chaired by B. Cushing, J. lowed the meeting. R Nelson, and M. J. Lacki, respectively) followed. On 25-26 February 1993, the Third Colloquium on Seventeen papers and two posters were presented. The Conservation of Mammals in the South-central United first paper entitled “Conservation Mammalogy in and States was held at the Ozark Folk Center State Park, Around Tennessee” by M. J. Harvey got the day off to an Mountain View, Arkansas. The meeting was co-sponsored excellent start. The day ended with a discussion session by the Department of Biology at the University of relating to conservation of mammals in the southeastern Arkansas Little Rock and Department of Biological United States; business relating to the journal issue and Sciences at Arkansas State University. Despite snowy the next annual meeting were discussed. This was another weather, the pre-meeting social on the evening of the highly successful meeting. M. J. Lacki invited the group 25th was well patronized. Forty-three participants attend¬ to hold the Sixth Colloquium in Kentucky. ed the meeting on the 26th. Technical sessions were Since the 1995 meeting in Cookeville, the Colloquium chaired by T. L. Best, J. Laerm, C. R. Tumlison, and J. P. on Conservation of Mammals in the Southeastern United Nelson. Twenty papers were presented. At the conclusion States has held four additional meetings. Locations and of the paper sessions, a roundtable discussion was held hosts were as follows: 1996 (sixth meeting)—Kentucky among all participants regarding conservation issues of Leadership Center, Somerset, Kentucky (M. J. Lacki, concern. Future direction and plans for the Colloquium University of Kentucky); 1997 (seventh meeting)— also were discussed. A field trip to Blanchard Springs YMCA Blue Ridge Assembly, Black Mountain, North Caverns was cancelled due to bad weather. Participants Carolina (M. K. Clark, North Carolina State Museum of believed that the meeting had been very successful and Natural Science); 1998 (eighth meeting)—Arlington decided that the fourth meeting would be held in Georgia Hotel, Hot Springs, Arkansas (D. A. Saugey, Ouachita (to be hosted by J. Laerm and B. Chapman). National Forest, United States Forest Service); 1999 The Fourth Colloquium on Conservation of (ninth meeting)—Holiday Inn, Wytheville, Virginia (R. Mammals in the Southeastern United States was held on Reynolds, Virginia Department of Game and Inland the campus of the University of Georgia in Athens on 25- Fisheries). The ninth meeting was sponsored by Virginia 26 February 1994. Hosts arranged an outstanding recep¬ Department of Game and Inland Fisheries, Ferrum tion for participants on the evening of the 25th at the College, Westvaco, George Washington and Jefferson Museum of Natural History. Food was bountiful, bever¬ national forests, and Virginia Department of Conservation ages copious, atmosphere biological, and interaction and Recreation, Division of Natural Heritage. among participants excellent. On the morning of the 26th, Beginning in 1996, the Colloquium has met jointly J. Laerm extended an official welcome to the group. with the Southeastern Bat Diversity Network. Format of Three technical sessions followed (chaired by J. Laerm, the meeting has included a day focused on topics relating B. Chapman, and M. L. Kennedy). Nineteen oral and two to bats and another on conservation of mammals in the poster presentations were presented. A roundtable discus¬ region. This joint meeting seemed natural because many sion included remarks relating to a number of conserva¬ individuals associated with the Colloquium also were tion issues. A decision was made to publish a journal interested in bats occurring in the southeastern United issue that would contain selected papers presented at the States. In 1999, an ad hoc Allegheny Woodrat Recovery Fourth Colloquium, as well as papers solicited from other Team met concurrently with the Colloquium and the researchers in the southeastern United States. About 80 Southeastern Bat Diversity Network. participants attended the meeting. M. J. Harvey invited The Colloquium and the field of conservation suf¬ the group to meet in Tennessee in 1995. fered a great loss in 1997 with the death of Dr. Joshua On 24-25 February 1995, the Fifth Colloquium on Laerm (University of Georgia). The 1998 meeting in Conservation of Mammals in the Southeastern United Hot Springs was special in that it was dedicated to the Kennedy—Colloquium on Conservation of Mammals memory of Dr. Laerm. The meeting was outstanding in a regional scale in addition to state levels. Individuals that ail regards. have participated in the meetings and shared information An assessment of the Colloquium after 9 years would with others in the region are to be commended. Because seem to indicate that the goals of the early organizers are of the many concerns relating to conservation of mam¬ being met. Each meeting has been somewhat larger than mals in the region, meetings like the Colloquium will the previous one in attendance, and communications play a major role in identifying problems and fostering among individuals working in the Southeast have been solutions. Overall, the future of the Colloquium appears improved, especially among academic institutions and to be very bright. Planning for the 10th annual meeting state and federal agencies. An annual newsletter prepared (to be held in Alabama and hosted by T. L. Best, Auburn by J. P. Nelson at Bethel College contributed in the most University) is in progress, and it promises to be another positive manner toward promoting exchange of informa¬ outstanding meeting. All individuals with an interest in tion during the first 5 years of the Colloquium. Most mammalian conservation in the southeastern United encouraging has been the discussions among participants States are invited to attend the annual meeting and partic¬ of the Colloquium relating to mammalian conservation on ipate in all parts of the program. 3 Laerm et al.—Conservation Status of Mammals Conservation Status of Terrestrial Mammals of the Southeastern United States Joshua Laerm, W. Mark Ford, and Brian R. Chapman Museum of Natural History and Institute of Ecology, University of Georgia, Athens, GA 30602 (JL) United States Department of Agriculture Forest Service, Northeastern Research Station, Parsons, VW 26287 (WMF) Daniel B. Warned School of Forest Resources, University of Georgia, Athens, GA 30602 (BRC) We reviewed the conservation status of terrestrial mammals of the southeastern United States based on taxa listed as endangered, threatened, rate, or of special concern under federal, state, and Natural Heritage Program guidelines. Of the 101 species of terrestrial mammals that occur in the southeastern United States, 78 (77.2%) are viewed as sensitive from a conservation perspective. Of the 276 currently recognized taxa (monotypic species plus subspecies of polytypic species), 137 (49.6%) known to occur in the region are viewed as sensitive. Sixteen species (15.6% of the regional species richness), including 23 taxa (8.3% of the total taxonomic richness), currently are listed as threatened or endan¬ gered under federal guidelines. Two taxa are candidates for listing under federal guidelines. At the state level, 62 species (61.4%) comprising 80 taxa (or 32.2%) officially are listed under respective state guidelines as endangered, threatened, rare, or of special concern. The Heritage Inventory Program lists of the respective states include an additional 16 species (including 42 taxa) beyond those listed under various federal and official state guidelines. Of the 21 families of mam¬ mals in the region, all but five have one or more sensitive taxa. For all families, the mean number of sensitive taxa rela¬ tive to total species richness is 68.7%. Current knowledge of taxonomy, distribution, and life-history attributes of many sensitive taxa in the southeastern United States is too poor to adequately assess their conservation status. Pursuant to the Endangered Species Act of 1973, taxa and their status with regard to federal, state, and several agencies of the United States Government have Heritage Inventory lists. A comprehensive review of the been authorized to compile lists of species, subspecies, or status of each sensitive mammalian taxon in the region distinct population segments of species that are either clearly is beyond the scope of this paper. Our intent is to endangered or threatened, as defined by law, following focus attention of professional biologists and representa¬ appropriate review (Bean, 1993; Wilcove et al., 1993). In tives of various state and federal agencies on those taxa addition, the respective states have initiated their own that are viewed as sensitive in the region. We hope that lists and protective regulations of taxa regarded as endan¬ identification of these taxa on a regional basis will foster gered, threatened, rare, or of special concern. greater understanding of problems associated with listed Furthermore, within most state governments, Heritage species and enhance opportunities for support of more Inventory programs have developed another set of species thorough studies. categories and lists to meet their own conservation imper¬ MATERIALS AND METHODS atives. The emphasis on, and concern for, listed species and subspecies has become the focus around which many The southeastern United States was defined as the conservation efforts have revolved (Lazell, 1993). Southern Region of the United States Department of A primary focus of this colloquium, and others in the Agriculture Forest Service, and includes Virginia, North series, was to provide regionally focused biologists a per¬ Carolina, South Carolina, Georgia, Florida, Alabama, spective from which research and education regarding Mississippi, Louisiana, Tennessee, Kentucky, Arkansas, conservation concerns of mammals in the region can be and the eastern portions of Texas and Oklahoma. We initiated. To identify various taxa of terrestrial mammals reviewed standard references (Hall, 1981; Jones et al., in the southeastern United States sensitive from a conser¬ 1992; Wilson and Reeder, 1993) and current literature to vation perspective, we undertook a review of the regional produce a list of mammalian species that occur in the 4 Laerm et al.—Conservation Status of Mammals region. We included all extant native species and intro¬ with respect to federal, state, and Heritage Inventory lists. duced species with established regional populations Of the 101 species that occur regionally, 78 (77.2%) (Rattus nor\’egicus, R. rattus, Mus musculus, Myocastor occur on one or another of the lists of sensitive species. coypus, Sus scrofa). We did not include taxa known to be Furthermore, of the 276 individual taxa, 128 (46.4%) are extirpated (e.g., Canis lupus, Bos bison, Census elaphus). considered sensitive. We determined total taxonomic richness through Sixteen species, including 23 taxa, currently are list¬ direct counts of monotypic species, plus currently recog¬ ed as threatened or endangered under federal guidelines nized subspecies of polytypic species. We were aware of (United States Fish and Wildlife Service, 1993). These controversy regarding taxonomy of numerous popula¬ represent 15.6% of the species richness and 8.3% of the tions. To resolve these controversies we used standard total mammalian taxonomic richness in the region, references (Hall, 1981; Jones et al., 1992; Wilson and respectively. Three taxa are listed as threatened and the Reeder, 1993) and current literature. Departures from remainder are listed as endangered. In addition, two taxa these authorities are discussed in Appendix I. are current candidates for listing (United States Fish and We compiled a faunal list of those terrestrial mam¬ Wildlife Service, 1996). All federally listed endangered malian species and subspecies that were considered to be or threatened taxa also are listed in the respective states sensitive from a conservation perspective. These includ¬ in which populations occur. Of the 23 federally listed ed: 1) taxa listed as endangered, threatened, or candidates taxa, three (Myotis grisescens, M. sodalis, and Canis for listing under United States Fish and Wildlife Service rufus) are listed throughout the range of the species. The guidelines prior to suspension of formal candidate- remaining 20 listed taxa are subspecies or populations of species designation (United States Fish and Wildlife more widely ranging species, and only designated sub¬ Service, 1993, 1996); 2) taxa listed as endangered, threat¬ species or populations are listed. ened, rare, or of special concern under respective official Sixty-two species (61.4% of the species richness), state guidelines; 3) taxa monitored or tracked under vari¬ including 87 taxa (or 32.3% of the total richness), are ous Heritage Inventory programs in the respective states. listed under respective state guidelines as endangered, Official state-listed taxa and those on Heritage Inventory threatened, rare, of special concern, or deemed in need of lists were obtained from the respective states. management. The Heritage Inventory programs of the We did not intend to provide a comprehensive evalu¬ respective states list an additional 16 species, including ation of the status of each sensitive taxon identified in our 42 taxa beyond those listed under various federal and analysis. New federal listings and rulings are issued fre¬ official state guidelines. quently. State lists are updated and changed on a yearly In the southeastern United States, 21 families of basis. Heritage Inventory lists are subject to continuous mammals occur, and all but five (Didelphidae, change. However, we believe that errors of inclusion or Dasypodidae, Castoridae, Myocastoridae, Suidae) have exclusion would not significantly alter our results qualita¬ one or more sensitive taxa (Table 2). In the remaining 16 tively or quantitatively. families, percentage of sensitive species relative to total Because respective state and Heritage Inventory lists species richness ranges from a low of 50% (Canidae) do not always specify subspecific affinities of species to a high of 100% (Talpidae, Molossidae, Leporidae, considered sensitive, we have attempted to refer popula¬ Geomyidae, Heteromyidae, Dipodidae, Canidae, tions in that state to particular subspecies, where possible, Procyonidae, Cervidae). In 10 families (essentially one- based on published range maps. We do not believe that half of the total family richness in the region), every this exaggerates the numbers of sensitive taxa. Rather, species has one or more subspecies listed as sensitive. For this permits a clearer view of the problem with respect to all families, mean number of sensitive species relative to total regional taxonomic richness. total species richness is 68.7%. Percentage of sensitive taxa per family relative to RESULTS total taxonomic richness ranges from a low of 21% in the In the southeastern United States, there are 101 Procyonidae to a high of 100% in the Molossidae, species of terrestrial mammals, including 276 individual Dipodidae, and Ursidae. We note that those families with taxa. These are listed in Table 1. We included only those 100% of taxa listed as sensitive are among the least spe- subspecies listed as sensitive and indicate their status ciose in the region. However, the proportion of sensitive 5