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Journal of Cave and Karst Studies December 2020 Volume 82, Number 4 JOURNAL OF ISSN 1090-6924 A Publication of the National CAVE AND KARST Speleological Society STUDIES DEDICATED TO THE ADVANCEMENT OF SCIENCE, EDUCATION, EXPLORATION, AND CONSERVATION Published By BOARD OF EDITORS The National Speleological Society Anthropology George Crothers http://caves.org/pub/journal University of Kentucky Lexington, KY Office george.crothers@utk.edu 6001 Pulaski Pike NW Huntsville, AL 35810 USA Conservation-Life Sciences Julian J. Lewis & Salisa L. Lewis Tel:256-852-1300 Lewis & Associates, LLC. nss@caves.org Borden, IN lewisbioconsult@aol.com Editor-in-Chief Earth Sciences Benjamin Schwartz Malcolm S. Field Texas State University National Center of Environmental San Marcos, TX Assessment (8623P) bs37@txstate.edu Office of Research and Development U.S. Environmental Protection Agency Leslie A. North 1200 Pennsylvania Avenue NW Western Kentucky University Bowling Green, KY Washington, DC 20460-0001 leslie.north@wku.edu 703-347-8601 Voice 703-347-8692 Fax field.malcolm@epa.gov Mario Parise University Aldo Moro Production Editor Bari, Italy mario.parise@uniba.it Scott A. Engel Knoxville, TN Carol Wicks 225-281-3914 Louisiana State University saecaver@gmail.com Baton Rouge, LA cwicks@lsu.edu Exploration Paul Burger National Park Service Eagle River, Alaska paul_burger@nps.gov Microbiology Kathleen H. Lavoie State University of New York Plattsburgh, NY lavoiekh@plattsburgh.edu Paleontology Greg McDonald National Park Service Fort Collins, CO The Journal of Cave and Karst Studies , ISSN 1090-6924, CPM greg_mcdonald@nps.gov Number #40065056, is a multi-disciplinary, refereed journal pub- lished four times a year by the National Speleological Society. The Journal is available by open access on its website, or check Social Sciences the website for current print subscription rates. Back issues are Joseph C. Douglas available from the NSS office. Volunteer State Community College Gallatin, TN POSTMASTER: send address changes to the National Speleo- 615-230-3241 logical Society Office listed above. joe.douglas@volstate.edu The Journal of Cave and Karst Studies is covered by the follow- Book Reviews ing ISI Thomson Services Science Citation Index Expanded, ISI Arthur N. Palmer & Margaret V Palmer Alerting Services, and Current Contents/Physical, Chemical, and State University of New York Earth Sciences. Oneonta, NY palmeran@oneonta.edu Copyright © 2020 Front cover: The pseudoscorpion Kleptochthonius griseoanus. by the National Speleological Society, Inc. See Lewis et al. this issue. Cato Holler Jr., Jonathan D. Mays, and Matthew L. Niemiller. The fauna of caves and other subterranean habitats of North Carolina, USA. Journal of Cave and Karst Studies, v. 82, no. 4, p. 221-260. DOI:10.4311/2019LSC0133 THE FAUNA OF CAVES AND OTHER SUBTERRANEAN HABITATS OF NORTH CAROLINA, USA Cato Holler Jr.1, Jonathan D. Mays2, and Matthew L. Niemiller3,C Abstract Over 1,500 caves have been documented in North Carolina, however, cave fauna in the Blue Ridge Mountains and Piedmont regions of North Carolina have been overlooked historically compared to the cave-rich karst terrains in the Appalachian Valley and Ridge and Interior Low Plateau to the west. Here, we provide the first comprehensive faunal list of caves and other subterranean habitats in the state based on over 40 years of periodic surveys and compilation of literature, biodiversity databases, and museum records. We report 475 occurrences from 127 caves, springs, and wells in 29 counties, representing 5 phyla, 17 classes, 43 orders, 90 families, 124 genera, and at least 164 species. Vertebrate fauna comprised 32 species, including 4 fishes, 9 salamanders, 1 lizard, 4 snakes, 2 birds, and 12 mammals (8 bats). Diverse invertebrate groups included spiders (11 families and 18 genera), springtails (7 families and 9 genera), segmented worms (3 families and 8 genera), and snails (6 families and 9 genera). At least 25 taxa are troglobites/ stygobites (cave obligates), including 5 species of cave flatworms, 5 cave springtails, and 5 cave amphipods. Most troglobitic/stygobitic fauna documented in this study are endemic to North Carolina. Counties with the greatest cave biodiversity include Rutherford, McDowell, Swain, Henderson, Polk, and Avery counties. Over 20 species documented are of conservation concern, including 14 troglobites and 3 federally-listed bats. Although not as diverse as adjacent states, caves and other subterranean habitats in North Carolina support a diverse community of invertebrates and ver- tebrates. Our review serves as a base line for future cave biological surveys in the state and highlights the importance of subterranean habitats for North Carolina biodiversity. Introduction Caves and associated subterranean habitats are home to a unique and taxonomically diverse assemblage of inver- tebrate and vertebrate organisms in North America. In addition to the more than 1,350 species that are obligate inhab- itants of terrestrial and aquatic subterranean habitats in the United States (i.e., troglobites and stygobites) (Niemiller et al., 2019), hundreds of other species use caves on an occasional to semi-permanent basis. Most of this subterranean diversity is known from the ten major karst biogeographic regions defined in the United States that are associated with carbonate exposures (Culver et al., 2003; Hobbs, 2012). Several additional smaller karst regions exist, but these regions have received comparatively little attention from biospeleologists. The fauna of caves and associated subterranean habitats in the Blue Ridge Mountains and Piedmont physiographic provinces of North Carolina has been grossly understudied compared to subterranean fauna of the cave-rich Appala- chian and Interior Low Plateau karst to the west of the Appalachian Mountains in Tennessee, Alabama, Georgia, and Virginia. In contrast to most cave systems in the Appalachians and Interior Low Plateau karst regions, cave systems in North Carolina are predominately non-solutional (i.e., pseudo-karst) and granite-gneiss talus and fissure caves, in par- ticular. A few troglobites and stygobites are known from North Carolina, including some that are endemic to the state, such as the Carolina Groundwater Amphipod (Stygobromus carolinensis) and Blowing Springs Cave Springtail (Pseu- dosinella flatua). The bat fauna from several caves and mines has been characterized (Boynton et al., 1992), particularly in association to winter hibernation surveys in recent years (e.g., North Carolina Wildlife Resources Commission, 2017b). However, a comprehensive, annotated list of the subterranean fauna of North Carolina has never been published. In early 1972, after the Flittermouse Grotto of the National Speleological Society was chartered, its members initi- ated the North Carolina Cave Survey (NCCS) to track and catalog caves in the state. In conjunction with this statewide cave inventory, and with the suggestion and encouragement of the late Dr. John Cooper, former Director of Research and Collections at the North Carolina State Museum of Natural Sciences, a survey of North Carolina’s little-known cave fauna also began. Here we report new distributional records of subterranean fauna documented during periodic sur- veys of caves between 1970 and 2018 conducted by the lead author and maintained by the NCCS. We also compiled occurrence records from caves and other subterranean habitats (e.g., wells, springs, and seeps) for invertebrate and vertebrate fauna of North Carolina from several sources, including published and unpublished literature, databases, and museum collections. We include these data to generate the first comprehensive faunal list from caves in the state. 1 Director, North Carolina Cave Survey, PO Box 100, Old Fort, North Carolina 28762; hollers2@frontier.com 2 Fish and Wildlife Research Institute, Florida Fish and Wildlife Conservation Commission, 1105 SW Williston Rd., Gainesville, Florida, 32601; jonathan.mays@myfwc.com 3 Department of Biological Sciences, The University of Alabama in Huntsville, Huntsville, Alabama 35899; cavemander17@gmail.com, matthew. niemiller@uah.edu CCorresponding author Journal of Cave and Karst Studies, December 2020 • 221 Holler Jr., Mays, and Niemiller Materials and Methods STUDY REGION AND GEOLOGICAL SETTING North Carolina lies in the southeastern United States bordered on the east by the Atlantic Ocean, on the south by South Carolina and Georgia, on the west by Tennessee, and on the north by Virginia. The state is divided into three physiographic regions: i) the Blue Ridge Mountains, composed of assorted igneous, metamorphic, volcanic, and sedi- mentary rocks of widely ranging ages; ii) the Piedmont Plateau consisting primarily of metamorphic and igneous rocks, and iii) the extensive Coastal Plain with its sands, clays, phosphates, and limestone, making up 45% of the state. As of October 2018, the North Carolina Cave Survey (NCCS) has 1,582 caves on record (Figure 1). Some states limit the term “cave” to a specific length measure; for example, several state cave surveys require that a passage be at least 50 feet (15.2m) in length to be considered a cave. Karst regions and solution caves are not common in North Car- olina, thus the NCCS recognizes the term as any naturally occurring opening within the earth capable of being entered by a human. True karst (solution) caves are limited to a few areas of the state. Madison County has three cave-bearing geological formations: Shady Dolomite, the Honaker Limestone, and a dolomite member of
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