The fauna of caves and other subter-ranean habitats of North Carolina, USA

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 vertebrates. 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 invertebrate and vertebrate organisms in North America. In addition to the more than 1,350 species that are obligate inhabitants 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 Appalachian 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 particular. 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 (Pseudosinella 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 initiated 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 surveys 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.