BANISTERIA A JOURNAL DEVOTED TO THE NATURAL HISTORY OF VIRGINIA Bundy Cave, Russell County, Virginia This special issue of Banisteria consists of a lead article on the history of biospeleology in Virginia, followed by a review of the 168 obligate cave-dwelling invertebrate species that inhabit the Commonwealth. Number 42 ISSN 1066-0712 2013 BANISTERIA A JOURNAL DEVOTED TO THE NATURAL HISTORY OF VIRGINIA ISSN 1066-0712 Published by the Virginia Natural History Society The Virginia Natural History Society (VNHS) is a nonprofit organization dedicated to the dissemination of scientific information on all aspects of natural history in the Commonwealth of Virginia, including botany, zoology, ecology, archeology, anthropology, paleontology, geology, geography, and climatology. Membership in VNHS includes a subscription to Banisteria. Annual dues are $20.00 (per calendar year); library subscriptions to Banisteria are $40.00. Subscribers/members outside the United States should add $3.00 for additional postage. Checks or money orders should be made payable to the Virginia Natural History Society (credits cards cannot be accepted). Membership dues and inquires should be directed to the Secretary-Treasurer (address, page 2); correspondence regarding Banisteria to the Editor. Banisteria is a peer-reviewed journal. The Editor will consider manuscripts on any aspect of natural history in Virginia or neighboring states if the information concerns a species native to Virginia or the topic is directly related to regional natural history (as defined above). Book reviews, biographies, and historical accounts of relevance to natural history in Virginia also are suitable for publication in Banisteria. For additional information regarding the VNHS, including other membership categories, field events, symposia, representative papers from past issues of Banisteria, and instructions for prospective authors, consult our website at: http://virginianaturalhistorysociety.com/ Editorial Staff: Banisteria Editor Steven M. Roble Virginia Department of Conservation and Recreation Division of Natural Heritage 600 E. Main Street, 24" Floor Richmond, Virginia 23219 Associate Editors Joseph C. Mitchell, Mitchell Ecological Research Service, LLC P.O. Box 2520, High Springs, Florida 32655 Alfred G. Wheeler, Jr., Department of Entomology Clemson University, Clemson, South Carolina 29634 Thomas F. Wieboldt, Department of Biology Virginia Polytechnic Institute & State University, Blacksburg, Virginia 24061 Banisteria No. 41 was published on 23 August 2013. Cover: Lucie L. Ferguson standing on a slope in front of the enormous entrance to Bundy Cave No. 1, Russell Co., Virginia. Photo by Lynn M. Ferguson, April 1968. Back cover: Top: Lynn M. Ferguson at entrance of Bundy Cave No. 1, Russell County, VA. Photo by Lucie L. Ferguson, April 1968. Bottom, left: Renowned biospeleologist and amphipod systematist John R. Holsinger exploring a cave stream. He has been the leading proponent for the inventory, documentation, and conservation of Virginia caves and their biota during the past half century, and is the lead author of both articles in this special issue of Banisteria. Photo by Lynn M. Ferguson, July 1991. Bottom, right: Lirceus culveri Estes & Holsinger, a troglobiotic isopod crustacean from on and around gravels in a stream in McDavids Cave, Scott Co., Virginia. The species was named in honor of renowned biospeleologist David C. Culver, second author of the feature article in this journal. This female specimen (length 5.9 mm) was photographed by Christopher S. Hobson. BANISTERIA A JOURNAL DEVOTED TO THE NATURAL HISTORY OF VIRGINIA Number 42, 2013 Contributed Papers A Brief History of Biospeleology in Virginia Pelarh Kt IOUS BCL P15 re gceeserernscseohoa ae aks iyi nahediongrribdt dibiehd ss MNeiy Va Reatesaerer htt opptecadhaaaniiwidiedtpeeeesttsdowenand see magia The Invertebrate Cave Fauna of Virginia John R. Holsinger, David C. Culver, David A. Hubbard, Jr., William D. Orndorff, ANC CNTISHOPHEr S- AAODSO Mes savccvccrcormnunnndbactstansadgelanennnepopenivnnenk el de onleovielllsccvyeedusinannnthhiSda hele labled soso caahekenekisas dace Miscellanea FEED OTS recreaitets chun natey desta tenpinaenntaneRaxans edna nace idilyens eee ease Mitr t lglbtepis i teevd beds pash apat latency radireretie duct sng Med tglitats ADHOUNCEMENIS.© 6.051, oA eecct scat yrs scnsceanceuuh cactteeetecnas Loven tecedun mise th hago St nessont yes seausearcyuab deeb teagan ny iasesce tien Ns Virginia Natural History Society Officers, 2013 President Todd S. Fredericksen School of Natural Science and Mathematics Ferrum College Ferrum, Virginia 24088 tfredericksen @ ferrum.edu (term expires December, 2014) Vice President Michael Lachance Virginia Cooperative Extension P.O. Box 298 Lovingston, Virginia 22949 lachance @ vt.edu (term expires December, 2014) Secretary-Treasurer William A. Shear Department of Biology Hampden-Sydney College Hampden-Sydney, Virginia 23943 wshear @hsc.edu (term expires December, 2016) Councilors Lisa D. Williams, Annandale (term expires December, 2013) Richard S. Groover, Mechanicsville (term expires December, 2014) Nancy Moncrief, Martinsville (term expires December, 2016) Honorary Councilors Michael Kosztarab Joseph C. Mitchell Webmaster John White vhs.webmaster @ verizon.net Banisteria, Editor Steven M. Roble steve.roble @ dcr. virginia.gov Banisteria, Number 42, pages 3-8 © 2013 Virginia Natural History Society A Brief History of Biospeleology in Virginia John R. Holsinger Department of Biological Sciences Old Dominion University Norfolk, Virginia 23529 ABSTRACT The study of cave biology or biospeleology in Virginia had its beginning in the second part of the 19th century with two papers published by E. D. Cope followed by a monograph in 1888 by A. S. Packard. Packard’s extensive work resulted from his observations on caves in Kentucky, Indiana, and Virginia. Following a period of inactivity, biological study of North American caves was resumed in the 1930s with investigations by both American and European workers, presumably driven by the rapidly increasing interest in evolutionary biology. Explanations for the loss of eyes, pigment, and attenuation of appendages in cave animals were especially interesting. Beginning in the 1950s and continuing until the present time, both extensive and intensive investigations of caves and their biotas have been carried out and many new species of insects and crustaceans have been discovered and described. One of the most remarkable has been the discovery of the marine relict isopod Antrolana lira from Madison Saltpetre Cave. Concurrent with a rapidly developing interest in both the biology and geology of caves, a “formal” Biological Survey of Virginia Caves was initiated in the early 1960s and resulted in a publication in 1988 titled “The invertebrate cave fauna of Virginia and a part of eastern Tennessee: zoogeography and ecology.” Subsequent to this publication, collecting has continued on a reduced scale paralleled by ecological studies by graduate students. Long-term studies on ecology, biological monitoring, and conservation of specific caves continue as well. Key words: biospeleology, Edward Drinker Cope, A. S. Packard, Madison’s Cave, troglomorphic, Burkes Garden, Thomas C. Barr, Jr., Biological Survey of Virginia Caves, Virginia Natural Heritage Program. The study of cave biology or biospeleology had its beginning in Virginia in approximately 1869 with a paper published by Edward Drinker Cope, who recorded beetles and millipeds he collected from caves two years earlier in Giles and Montgomery counties (Cope, 1869; Grady, 1987). This early acknow- ledgement of living organisms in caves is one of the first scientific documentations of subterranean animal life in North America and is probably the first listing of cave-adapted animals in Virginia. A few years later, Cope (1872) described the subterranean amphipod crustacean genus Stygobromus based on specimens collected in Mammoth Cave, Kentucky. Subsequently, many species of this genus have been discovered and described from Virginia caves as well as elsewhere in North America and Asia (e.g., Holsinger, 1967, 1978, 2009). It is interesting that Cope, who is usually thought of as primarily a paleontologist, also made these early collections and descriptions of arthropods. Nevertheless, Cope’s significant work was soon overshadowed by the extensive biospeleological research in Virginia and other parts of eastern North America by A. S. Packard (1888), who published an important monograph titled “The cave fauna of North America, with remarks on the anatomy of the brain and origin of the blind species” in Memoirs of the National Academy of Sciences. Notably, Packard’s principal observations and collections were based primarily on his visits to the relatively well known and easily accessible “show” caves in Virginia, Kentucky, and Indiana at a time when literally nothing was known about thousands of other caves that would be discovered and explored in North America in the years to follow. In Virginia, these early known “show” caves were all in the Shenandoah Valley and included Fountain, Grand (formerly Weyer’s), and Madison’s in “Cave Hill” near the town of Grottoes; Endless or Zirkles Cave near New Market; and Luray Cave (now Luray Caverns) near the town of Luray. Elsewhere, Packard visited and _ collected specimens from Wyandotte Cave in southern Indiana, Mammoth Cave in central Kentucky, and a few others. Prior to 4 BANISTERIA publication of Packard’s extensive volume, literally nothing was known about the numbers or kinds of organisms that lived in the subterranean ecosystems of North American caves. It is interesting that prior to the early works of Cope and Packard at least one cave in Virginia was visited by both George Washington and Thomas Jefferson in the early part of the 19" century. Although neither was a biologist, both were apparently interested in natural history, and their visits to a cave later called Madison’s Cave (more recently Madison Saltpetre Cave) probably marked the beginning of speleology in Virginia. Moreover, Jefferson’s “eye draught” map of Madison’s Cave is one of the first maps of a cave in America (see Halliday, 1968). Also of historical interest is the fact that Packard mistakenly referred to Fountain Cave as nearby Madison’s Cave in his 1888 monograph. It is unfortunate that Packard did not visit Madison’s Cave. This cave is clearly the most biologically significant of the three Cave Hill caves and data from there would have enhanced Packard’s work. Following Packard’s extensive publication in 1888, little was written about the exploration of caves, their geological origins, or biological contents during the next 20 years. This apparent lack of interest in cave science seems to have extended well into the early part of the 20" century and is surprising in view of the excitement that one would have expected from the earlier papers of Cope and Packard. However, all of this changed in the 1920s and early 1930s. During this period, American coleopterists S. Barber and J. M. Valentine and European biospeleologists C. Boliver and René Jeannel made extensive collections of invertebrates (primarily insects) from Virginia caves and published their findings in a series of papers that followed their visits. This new awakening of biospeleological activity appears to have occurred simultaneously with a renewed interest in cave exploration and documentation. Following the lag of speleological activity in the early 20" century, the 1930s also witnessed a merger of taxonomic and genetic thinking and development of the so-called “synthetic theory” in evolutionary and systematic biology (see Mayr, 1963). It followed that blind, unpigmented animals, often with attenuated append- ages, living in caves demanded attention and an explanation for their loss of eyes and pigment and development of other “troglomorphic” characters common to cave-adapted organisms. This newly developing emphasis on evolutionary biology in the 1930s clearly had a significant effect on the renewed interest in cave animals, not just in Virginia, but elsewhere in America as well. NO. 42, 2013 From the late 1930s well into the 1950s, in concert with the developing interest in evolutionary biology and exploration and documentation of caves, a number of both American and European biologists collected specimens from caves in Virginia and adjoining states. These workers and their taxonomic group(s) of interest included Kenneth Dearolf (general collecting), Leslie Hubricht (Fig. 1; crustaceans and snails), J. P. E. Morrison (snails), Henri Henrot (crustaceans), and Bruno Condé (diplurans). These collections resulted not only in descriptions of many interesting new species but provided significant new information on the ecology and biogeography of cave-adapted animals. Probably the most remarkable Virginia cave biology story to emerge from the 1950s involved a collecting trip to southwestern Virginia by Bruno Condé from Nancy, France. Condé came to Washington, D.C. in 1955 to attend an international zoological meeting. While in close proximity to the Appalachian cave region, Condé planned to use this occasion to visit Lawson Cave in southwestern Virginia from which he had seen specimens of an interesting, undescribed dipluran insect that he hoped to describe and study in more detail. Because this cave was in Virginia, Condé believed it would be easy for him to travel to it and collect additional specimens during his visit to the U.S. Little did he know that Wytheville, the only town of any size “close” to Lawson Cave in Burkes Garden, was approximately 250 miles from D.C. in “distant” southwestern Virginia. Determined to visit this cave, Fig. 1. Leslie Hubricht, an early collector of aquatic cave and spring organisms in Virginia and elsewhere in eastern North America. HOLSINGER: HISTORY OF BIOSPELEOLOGY 5 Condé took what must have been at the least a 12-hour trip by Greyhound bus from Washington, D.C. to Wytheville. Once in Wytheville he was told that Burkes Garden was in a remote, high elevation valley in the mountains some 20 miles to the northwest. The only road to Burkes Garden from Wytheville was an unimproved, gravel-bedded Forest Service road. Amazingly, Condé was able to talk a cab driver into taking him to Burkes Garden. Once at the cave, he paid the cabbie to wait while he collected diplurans. Apparently everything went as planned and Condé found numerous diplurans on a mud bank in the cave. He returned to Wytheville and took a bus back to D.C. In telling this fantastic story to a group of colleagues, at a meeting of subterranean biologists many years later in Moulis, France, Condé never said how long these bus rides lasted or how he found Lawson Cave once he reached Burkes Garden. However, in those days there were no interstate highways and the road across the mountain to Burkes Garden wasn’t much more than a narrow, winding, unimproved graveled pathway with hairpin curves and switchbacks. One can only imagine what the locals thought when they encountered this vivacious French zoologist who had traveled all the way to Tazewell County from Europe to collect tiny, white, eyeless insects from a cave in remote Burkes Garden. Whatever the obstacles, Condé returned to France with a nice collection of insects and a great story to tell his colleagues. Following the renewed biological interest in caves in the 1930s and 1940s, Thomas C. Barr, Jr. (Fig. 2), a biospeleologist from Tennessee, made an extensive collecting trip through western Virginia in 1958, where he visited 37 caves. Although Barr’s primary goal was to collect trechine cave beetles in the genus Pseudanophthalmus, one of his most important discoveries was finding a rare, marine relict cirolanid isopod in a deep lake in Madison Saltpetre Cave in Augusta County. This new, biogeographically significant isopod was subsequently described by Thomas C. Bowman, a curator of Crustacea at the Smithsonian Institution, and named Antrolana lira (Bowman, 1964). Originally believed to be extremely rare and restricted to a single subterranean aquifer, A. lira was placed on the Federal Threatened Species List. Although now known to be more common and widespread than first thought, it is nevertheless one of the most significant cave-adapted animals in eastern North America (Holsinger et al., 1994), and its protection as a “threatened species” is_ clearly warranted. In the early 1960s, in concert with the renewed interest in cave exploration and documentation and the potential for finding undescribed new species as well as Fig. 2. Dr. Thomas C. Barr, Jr. collecting cave beetles from an Appalachian cave. Fig. 3. Drs. John R. Holsinger (left) and John E. Cooper (right) in front of Showalter’s Cave, Rockbridge Co., VA ona collecting trip in May 1961 that marked the beginning of the “Biological Survey of Virginia Caves” project. a need for additional data on the systematics and ecology of cave organisms, an extensive biological survey of caves in Virginia was initiated by the author. This newly developed project named the “Biological Survey of Virginia Caves” was first assisted by John E. Cooper (Fig. 3) and later joined by David C. Culver (Fig. 4). It was subsequently expanded to include the contiguous karst areas of upper eastern Tennessee. The results of this study, which were based on collections and observations in approximately 500 caves in Virginia and northeastern Tennessee, were published in 6 BANISTERIA enn ee, A cee Fig. 4. Dr. David C. Culver collecting “cave-like” crustaceans from a groundwater seep in the Piedmont of eastern Virginia just west of the Appalachian cave region. a monograph titled “The invertebrate cave fauna of Virginia and a part of eastern Tennessee: zoogeography and ecology” by Holsinger & Culver (1988). It is perhaps of interest that publication of this monograph was on the 100 year anniversary of Packard’s earlier monograph. Approximately 335 species of invertebrate animals, representing some 90 families and 173 genera, were recorded from caves in the study area. A companion to the Biological Survey of Virginia Caves was developed later to include the adjacent karst region in neighboring West Virginia. The West Virginia Cave Survey data were published in Bulletin 7 of the West Virginia Speleological Survey by J. R. Holsinger, R. Baroody, and D. C. Culver (1976). A revised and updated edition of this monograph was published recently as Bulletin 16 of the West Virginia Speleological Survey (Fong et al., 2007). A similar revision and update of the invertebrate cave fauna of Virginia by J. R. Holsinger, D. C. Culver, D. A. Hubbard, Jr., W. D. Orndorff, and C. S. Hobson has been completed and appears as the next paper in this issue of Banisteria. Subsequent to the publication by Holsinger & Culver (1988), collecting in Virginia caves has continued on a reduced scale, consisting primarily of obtaining specimens from caves newly discovered since 1988 and returning to previously studied caves for additional specimens and ecological observations. However, many new records, including additional undescribed species, were obtained during the 1990s by David A. Hubbard, Jr. (Fig. 5), who visited many caves while simultaneously collecting geological data for a state “Karst Map” being developed by the Virginia Division of Mineral Resources. More _ recently, biological studies of caves have concentrated on gathering pertinent data on the ecology and NO. 42, 2013 biogeography of specific taxa in some of the state’s most important biologically significant caves. Paralleling these studies has been a number of important conservation efforts as well. These include acquisition and/or long-term protection of biologically significant caves by the Virginia Natural Heritage Program and the Cave Conservancy of the Virginias (CCV), and the development of protective measures for karst drainage basins associated with large cave systems. Beginning in the early to middle 1970s, there have been a number of important studies by former graduate students (Fig. 6) working on the systematics and/or ecology of specific taxa or on the ecology of the fauna of a selected cave. The results of much of this important research is published and readers are referred to the bibliography in Holsinger & Culver (1988) for a complete listing of the published research papers based on projects completed in the 1970s and 1980s. Briefly, these projects included research on the ecology of the amphipod Gammarus minus by Steven W. Hetrick; population ecology of Antrolana lira in Madison Saltpetre Cave and nearby Stegers Fissure by T. Lynn Collins; a series of papers on the ecology of the amphipod crustacean Crangonyx antennatus in Lee County by Gary W. Dickson; ecology of the isopod crustacean Lirceus usdagalun in Thompson Cedar Cave combined with assistance on the description of the sister species Lirceus culveri in Rye Cove by James A. Estes; systematics and biogeography of dipluran insects in the family Campodeidae by Lynn M. Ferguson and additional biological collecting; and taxonomy of Fig. 5. David A. Hubbard, Jr., collected extensively in Virginia caves during the 1990s (photo courtesy of Lynn M. Ferguson). HOLSINGER: HISTORY OF BIOSPELEOLOGY 7 Fig. 6. Three former Ecological Science graduate students from Old Dominion University collecting crustaceans from a cave stream in Lee County. asellid isopods in the genus Caecidotea by Julian J. Lewis and Lawrence E. Fleming. The most recent graduate student research was carried out by Ben Hutchins on the genetic structure of the Madison Cave Isopod under the direction of Daniel W. Fong and David B. Carlini at American University (Hutchins et al., 2010). In addition to the flourish of graduate student research during the period described above, Robert Hershler, a curator of Mollusca at the Smithsonian Institution, described Holsingeria unthanksensis, a rare new hydrobiid snail from a stream in Unthanks Cave in Lee County (Hershler, 1989). In the last few years, personnel of the Virginia Department of Conservation and Recreation’s Division of Natural Heritage have also become involved in long- term, biological monitoring of selected caves, with special emphasis on the conservation and protection of the Federally Threatened Madison Cave Isopod and the Federally Endangered Lee County Cave Isopod, Lirceus usdagalun. A recent study, involving personnel from both local universities and the state’s Natural Heritage Program is underway to precisely determine the relationship between deep groundwater aquifers and the geographic distribution of the Madison Cave Isopod in the Shenandoah Valley karst region of western Virginia and eastern West Virginia. In addition, efforts are being made by personnel of the Heritage Program’s Karst Protection group and the Virginia Department of Game and Inland Fisheries (VDGIF) to record and monitor the distribution and detrimental effects of White Nose Syndrome (WNS) on the state’s bat populations. Although not affected to date by WNS, the lone maternity colony in Virginia of the Federally Endangered Virginia Big-eared Bat (Corynorhinus townsendii virginianus) 1n a cave in Burkes Garden is routinely monitored by VDGIF personnel. What is in the future for cave biology research in Virginia? Many new caves have been discovered in recent years and many more remain to be found. In addition, new passages in previously known caves continue to be discovered. Currently, based on the Virginia Speleological Survey database, the number of recorded caves in the state is approximately 3,000, excluding tight fissures, crevices, and most rock shelters. It is of further interest that five of these caves are greater than 10 miles in length, 17 are greater than 5 miles in length, and many more are longer than 1 mile. Biological exploration of newly discovered caves and/or passages in previously known caves have the potential for discovery of new species and perhaps new genera, and range extensions for previously known species. In addition, discovery of new taxa can lead to important new studies on their evolution, ecology, and biogeography. ACKNOWLEDGMENTS I am grateful to the following colleagues who were kind enough to read an earlier draft of this paper and provide additional information and suggestions for improvement: John E. Cooper, David C. Culver, Daniel W. Fong, Julian J. Lewis, the late Thomas C. Barr, Jr., Lynn M. Ferguson, and Wil Orndorff. Mike Futrell provided current data from the Virginia Speleological Survey on the number of caves in the state. In addition, I thank the many taxonomists who have provided identifications and/or descriptions of taxa from Virginia caves over the years. Their contributions have added immensely to our knowledge of Virginia’s significant cave fauna. A near complete list of these workers can be found under “Acknowledgments” on page 146 in Holsinger & Culver (1988). LITERATURE CITED Bowman, T. E. 1964. Antrolana lira, a new genus and species of troglobitic cirolanid isopod from Madison Cave, Virginia. International Journal of Speleology 1: 229-236. Cope, E. D. 1869. Synopsis of the extinct Mammalia of the cave formations in the United States, with observations on some Myriapoda found in and near the same, and on some extinct mammals of the caves of Anguilla, W. I., and of other localities. Proceedings of the American Philosophical Society 11: 171-192. 8 BANISTERIA Cope, E. D. 1872. On the Wyandotte Cave and its fauna. American Naturalist 6: 406-422. Fong, D. W., D. C. Culver, H. H. Hobbs, III, & T. Pipan. 2007. The Invertebrate Cave Fauna of West Virginia. Second Edition. West Virginia Speleological Survey Bulletin No. 16. 163 pp. Grady, F. 1987. Edward Drinker Cope’s contributions to speleology. Journal of Spelean History 21: 37-38. Halliday, W. R. (ed). 1968. The grottoes of the Shenandoah. Journal of Spelean History 1: 20-43. Hershler, R. 1989. Holsingeria unthanksensis, a new genus and species of aquatic cavesnail from eastern North America. Malacological Review 22: 93- 100. Holsinger, J. R. 1967. Systematics, speciation, and distribution of the subterranean amphipod genus Stygonectes (Gammaridae). United States National Museum Bulletin No. 259. 176 pp. Holsinger, J. R. 1978. Systematics of the subterranean amphipod genus Stygobromus (Crangonyctidae), Part II: species of the eastern United States. Smithsonian Contributions to Zoology No. 266. 144 pp. Holsinger, J. R. 2009. Three new species of the subterranean amphipod crustacean genus Stygobromus (Crangonyctidae) from the District of Columbia, Maryland, and Virginia. Pp. 261-276 In S. M. Roble & J. C. Mitchell (eds.), A Lifetime of NO. 42, 2013 Contributions to Myriapodology and the Natural History of Virginia: A Festschrift in Honor of Richard L. Hoffman’s 80" Birthday. Virginia Museum of Natural History Special Publication No. 16, Martinsville, VA. Holsinger, J. R., R. Baroody, & D. C. Culver. 1976. The Invertebrate Cave Fauna of West Virginia. West Virginia Speleological Survey Bulletin No. 7. 82 pp. Holsinger, J. R., & D. C. Culver. 1988. The invertebrate cave fauna of Virginia and a part of eastern Tennessee: zoogeography and ecology. Brimleyana 14: 1-162. Holsinger, J. R., D. A. Hubbard, Jr., & T. E. Bowman. 1994. Biogeographic and ecological implications of newly discovered populations of the stygobiont isopod crustacean Antrolana lira Bowman _ (Cirolanidae). Journal of Natural History 28: 1047-1058. Hutchins, B., D. W. Fong, & D. B. Carlini. 2010. Genetic population structure of the Madison Cave isopod, Antrolana lira (Cymothoida: Cirolanidae) in the Shenandoah Valley of the eastern United States. Journal of Crustacean Biology 30: 312-322. Mayr, E. 1963. Animal Species and Evolution. Belknap Press of Harvard University Press, Cambridge, MA. 797 pp. Packard, A. S. 1888. The cave fauna of North America with remarks on the anatomy of the brain and origin of the blind species. Memoirs of the National Academy of Sciences 4: 1-156. Exterior and interior views of the entrance to Surgener Cave in Lee County, one of Virginia’s most biologically significant caves. Photos by John R. Holsinger. Banisteria, Number 42, pages 9-56 © 2013 Virginia Natural History Society The Invertebrate Cave Fauna of Virginia John R. Holsinger Department of Biological Sciences Old Dominion University Norfolk, Virginia 23529 David C. Culver Department of Environmental Science American University 4400 Massachusetts Avenue NW Washington, DC 20016 David A. Hubbard, Jr. Virginia Speleological Survey 40 Woodlake Drive Charlottesville, Virginia 22901 William D. Orndorff Virginia Department of Conservation and Recreation Division of Natural Heritage Karst Program 8 Radford Street, Suite 102 Christiansburg, Virginia 24073 Christopher S. Hobson Virginia Department of Conservation and Recreation Division of Natural Heritage 600 East Main Street, 24 Floor Richmond, Virginia 23219 ABSTRACT The obligate cave-dwelling invertebrate fauna of Virginia is reviewed, with the taxonomic status and distribution of each species and subspecies summarized. There are a total of 121 terrestrial (troglobiotic) and 47 aquatic (stygobiotic) species and subspecies, to which can be added 17 stygobiotic species known from Coastal Plain and Piedmont non-cave groundwater habitats, and published elsewhere (Culver et al., 2012a). Richest terrestrial groups are Coleoptera, Collembola, and Diplopoda. The richest aquatic group is Amphipoda. A number of undescribed species are known and the facultative cave-dwelling species are yet to be summarized. Key words: Appalachians, biogeography, biospeleology, caves, springs, stygobionts, subterranean, troglobionts. 10 BANISTERIA INTRODUCTION The cave fauna of Virginia, most particularly the obligate cave-dwelling fauna, has been studied and described for over 100 years. The first obligate cave- dwelling species described from a Virginia cave was a beetle, Pseudanophthalmus pusio, described by Horn in 1868. As of 2012, 168 obligate subterranean species have been described from Virginia caves and associated habitats (Table 1), and another 17 species described from other subterranean habitats (Culver et al., 2012a). The present study grew out of the realization that the last compilation of the Virginia cave fauna by Holsinger & Culver (1988) was seriously out of date, and that in the intervening years extensive collections had been made in Virginia caves, especially by David Hubbard, and that many new species have been described, especially by William Shear and Kenneth Christiansen. In addition, the establishment of the Virginia Natural Heritage Program in 1986, and its extensive records of subterranean fauna, meant that it would be relatively easy to assemble information on the subterranean fauna of Virginia. Early on, we recognized that the subterranean fauna of seepage springs and shallow wells of the Coastal Plain and Piedmont, while sharing many morphological features with cave- dwelling species, especially loss of eyes and pigment and elongation of appendages (see Culver et al., 2010), was geographically distinct, and thus best treated separately. This shallow subterranean fauna, together with that of Maryland and the District of Columbia, was summarized in a separate publication (Culver et al., 2012a). The present study enumerates the described obligate subterranean fauna of caves and associated habitats in Virginia, together with a list of all known localities, a distribution map, and some comments on the systematic status of the species and subspecies. Table 1. Number of described troglobionts and stygobionts known from Virginia at various times. The number in parentheses is the additional number of stygobionts known from shallow subterranean habitats in the Coastal Plain and Piedmont (Culver et al., 2012a). Reference Troglobionts Stygobionts Packard (1888) 9 0 Holsinger (1963) 38 wi Holsinger & Culver (1988) 67 35 Present study 121 47(17) NO. 42, 2013 METHODS AND MATERIALS We assembled all published records, all records from the Virginia Natural Heritage Program database, and supplemented this with our own unpublished records. The final list was reconciled with the Virginia Natural Heritage Program database. All records are based on voucher specimens deposited in various museums and research collections around the world. No unconfirmed sight records are included. Locations were obtained from the same sources, and supplemented by information supplied by the Virginia Speleological Survey. In a very few cases, especially old records involving springs, it was not possible to determine an exact location. All locations have an error of less than 1 km. Distribution maps were created using ArcMap 10. In order to emphasize not only where a species was found but also where it was not found, gray dots on all maps indicate all terrestrial or aquatic sampling sites, depending on the species. A map of the counties where caves were sampled is shown in Fig. 1. The most difficult decision was selecting which species to include in this review. Holsinger & Culver (1988) treated both obligate and non-obligate cave- dwelling species, including both described and undescribed species. We decided to include only described, obligate cave-dwelling species in this paper for several reasons. The first is a practical one—the number of species has grown to such an extent that even the limitation to described obligate cave-dwellers is a formidable one. Second, records on species that are not obligate cave-dwellers are very incomplete, and very few new records have been added since the Holsinger & Culver report. This is not to imply that these species are unimportant. Many are major components of both aquatic and _ terrestrial communities, and in the case of crickets (the genera Ceuthophilus and Euhadenoecus), they are a major source of organic carbon and nutrients as a result of their regular trips in and out of caves. The third reason has to do with limitation to described species. Undescribed species range from ones that have been well studied, with detailed drawings, but are not yet published to those that are not adequately characterized or described. Even in the case of the cave fauna where endemism is the rule, not all potential undescribed species turn out to be new species upon further study. About one-third, on closer examination, prove to be minor variants of existing species (Culver et al., 2012b). Our criterion for inclusion of a described species in this list is that at least 90 percent of the records for the species were from _ subterranean habitats. This distributional definition corresponds to the original HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 11 Ae ® Sample sites se | wate Sek in . @ = : Norfolk ¥ fl \ Fig. 1. Distribution of sampling sites in Virginia. More than 95 percent of these records are from caves; the rest are from springs and other groundwater-associated habitats. ecological classification of Schiner (1854) and others (see Sket, 2008), with troglobionts (terrestrial) and stygobionts (aquatic) the terms for _ obligate subterranean dwellers. Most troglobionts and stygobionts have reduced or absent eyes and pigment, and these criteria (collectively termed troglomorphy by Christiansen [1962]) have sometimes been used to determine which species are troglobionts and stygobionts. Operationally, this often works but there are several problems. First, it is often difficult to determine how much eye and pigment reduction is necessary to call a species troglomorphic. Second, troglomorphy is based on the idea of strong convergence of subterranean species, but this is being challenged by the discovery of divergence of stygobionts and troglobionts within a community (FiSer et al., 2012). Pipan & Culver (2012) point out that most cave communities have non-troglomorphic species, and that many other subterranean habitats have troglomorphic species. As Daniel Fong (pers. comm.) points out, troglomorphy is a hypothesis, not an invariant law. In those cases where non-troglomorphic species have been included, we have indicated such. It may well be that some or most of these species will prove to be troglophiles and stygophiles rather than troglobionts and stygobionts, but until they have been regularly found outside of caves, we have included them. RESULTS A total of 121 troglobiotic and 47 stygobiotic species and subspecies have been found in Virginia caves. In 1988, the comparable numbers were 67 and 35 (Table 1). Comparable numbers are not available for most states, but in neighboring West Virginia there are 63 troglobionts and 33 stygobionts (updated from Fong et al., 2007). In Arkansas, there are 27 troglobionts and 28 stygobionts (Graening et al., 2011). The taxonomic distribution of the troglobionts and stygobionts is enumerated in Table 2. Among stygobionts, amphipods with 51 percent (24) of the species and subspecies, are the most diverse. The only other order with more than ten percent of the known stygobionts is Isopoda, with 13 species. Three orders dominate the terrestrial fauna, each with 23 or more species — chordeumatid millipeds, Collembola, and Coleoptera (Table 2). Only one other order, Pseudoscorpiones with 13 species, has as much as 10 percent of the fauna. 12 BANISTERIA NO. 42, 2013 Table 2. Taxonomic position of Virginia stygobionts and troglobionts, excluding species from the Piedmont and Coastal Plain. Phylum Class Order Platyhelminthes Turbellaria Lecithoepetheliata Tricladida Annelida Clitellata Lumbriculida Mollusca Gastropoda Mesogastropoda Arthropoda Chelicerata Acari Araneae Pseudoscorpiones Diplopoda Chordeumatida Chilopoda Lithobiomorpha Hexapoda Collembola Diplura Insecta Coleoptera Diptera Malacostraca Amphipoda Isopoda Number of Terrestrial Species and Subspecies Number of Aquatic Species and Subspecies RN We 24 13 4 ANNOTATED LIST OF SPECIES PHYLUM PLATYHELMINTHES Class Turbellaria Order Lecithoepetheliata Family Prorhynchidae Geocentrophora cavernicola Carpenter 1970 Type Locality: John Rogers Cave, Jackson County, Kentucky Virginia Record (Fig. 2): Tazewell Co.: Fallen Rock Cave. Remarks: This is the only species in the genus known from subterranean sites in North America. However, two other species of Geocentrophora are recorded from subterranean waters in Europe (Schwank, 1986). G. cavernicola is also known from Blowhole Cave, Pendleton Co., West Virginia (Fong et al., 2007). Order Tricladida Family Kenkiidae Sphalloplana chandleri Kenk 1977 Type Locality: Stokes Lane spring, Davidson County, Tennessee Virginia Record (Fig. 3): Tazewell Co.: Fallen Rock Cave. Remarks: This planarian is also known from a cave spring in Tennessee, as well as a spring in Floyd Co., Indiana. The type locality spring issues from a cave (Kenk, 1977). Sphalloplana consimilis Kenk 1977 Type Locality: Gallohan No. 1 Cave, Lee County, Virginia Other Virginia Records (Fig. 3): Lee Co.: Bowling Cave, Cope Cave, Gregorys Cave, Hamblin School Cave, McClure Cave. Remarks: The occurrence of this species is sporadic but it is common where found. It is also known from Claiborne County, Tennessee, and is endemic to the Powell River watershed in Virginia and Tennessee. Sphalloplana virginiana Hyman 1945 Type Locality: Showalter Cave, Rockbridge County, Virginia Other Virginia Records: Known only from the type locality (Fig. 3). Remarks: The species is apparently endemic to a single cave. Holsinger & Culver (1988) noted that previous records from Bland and Lee counties (Holsinger, 1963) are invalid in light of subsequent revisionary studies by Kenk (1977). HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA Kilometers Fig. 2. Distribution of Geocentrophora cavernicola in Virginia. Gray dots are all the aquatic sampling sites. # Sphalloplana chandler A Sphalloplana consimilis gw Sphalloplana virginiana Big Stone Gap Owed, tnt FX Fig. 3. Distribution of Sphalloplana chandleri, S. consimilis, and S. virginiana in Virginia. Gray dots are all the aquatic sampling sites. 150 kilometers * Spelaedrilus multiporis A Stylodrilus beattiei “a Big Stone Gap pg AS- Fig. 4. Distribution of Spelaedrilus multiporus and Stylodrilus beattiei in Virginia. Gray dots are all the aquatic sampling sites. Kilometers 13 14 BANISTERIA PHYLUM ANNELIDA Class Clitellata Order Lumbriculida Family Lumbriculidae Spelaedrilus multiporus Cook 1975 Type Locality: Smiths Cave, Russell County, Virginia Other Virginia Records: Known only from the type locality (Fig. 4). Remarks: This is the only species in the genus. Stylodrilus beattiei Cook 1975 Type Locality: Tub Cave, Pocahontas County, West Virginia Virginia Record (Fig. 4): Tazewell Co.: Steeles Cave. Remarks: This worm is also known from Court Street Cave and Friars Hole Cave in Greenbrier Co., West Virginia (Fong et al., 2007). PHYLUM MOLLUSCA Class Gastropoda Order Mesogastropoda Family Hydrobiidae Fontigens bottimeri (Walker 1925) Type Locality: Glenn Echo, Montgomery County, Maryland Virginia Records (Fig. 5): Frederick Co.: Ogdens Cave, seep 0.8 km S of Hayfield. Remarks: Found in both caves and small springs in the Potomac and Shenandoah River basins in northern Virginia, Maryland, and District of Columbia (Hershler et al., 1990), this snail is listed as a State Endangered Species in Virginia. Its distribution in the rest of the state is shown in Culver et al. (2012a). It ranges in size from | to 3 mm. Fontigens morrisoni Hershler, Holsinger & Hubricht 1990 Type Locality: A small, spring-fed brook SW of Mustoe, Highland County, Virginia Other Virginia Records (Fig. 5): Bath Co.: Blowing Cave; Highland Co.: Aqua Cave, Butler-Sinking Creek Cave, Vandevander Cave, spring 2.1 km N of Mustoe. Remarks: A small species (1.5-2.5 mm) that can be found in both springs and cave habitats within the upper James River watershed. NO. 42, 2013 Fontigens tartarea Hubricht 1963 Type Locality: Organ Cave, Greenbrier County, West Virginia Virginia Records (Fig. 5): Bath Co.: Crossroads Cave; Highland Co.: Secret Anthodite Cave. Remarks: One of the smallest freshwater snails (1.7-2 mm) in eastern North America, it was discovered in Virginia for the first time in 1994. Holsingeria unthanksensis Hershler 1989 Type Locality: Unthanks Cave, Lee County, Virginia Other Virginia Records (Fig. 5): Lee Co.: Burial Cave, Gibson No. | Cave, Jones Saltpetre Cave, Tater Cave. Remarks: This local endemic is found among small submerged rocks in small streams in five caves confined to a portion of the Powell River basin in Lee County, Virginia. It is listed as a State Endangered Species. PHYLUM ARTHROPODA Class Chelicerata Order Acari Family Rhagidiidae Poecilophysis extraneostella Zacharda 1985 Type Locality: Bowling Cave, Lee County, Virginia Other Virginia Records: Known in Virginia only from the type locality (Fig. 6). Remarks: Although this mite is known only from caves and hence by definition is a troglobiont, it shows little if any modification for subterranean life. It is also known from four caves in West Virginia (Fong et al., 2007). Poecilophysis weyerensis (Packard 1888) Type Locality: Grand Caverns, Augusta County, Virginia Other Virginia Record (Fig. 6): Rockbridge Co.: Buck Hill Cave. Remarks: Originally described by Packard (1888) from Grand Caverns, it was redescribed by Holsinger (1965) and subsequently reported from caves in Missouri, New Mexico, and Mexico by Elliott & Strandtmann (1971). Zacharda (1980) listed this species (synonym = Rhagidia cavernarum) from Long Cave in Edmonson County, Kentucky, and from scree habitats in the Czech Republic, but questioned the records given by Elliott & Strandtmann (1971). Zacharda (1985) gave an additional record from Buck Hill Cave as well as Sensabaught Saltpetre Cave in Hawkins Co., Tennessee. As with P. extraneostella, it shows little HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 15 morphological modification for cave life. Rhagidia varia Zacharda 1985 Type Locality: Butler-Sinking Creek Cave, County, Virginia Other Virginia Records (Fig. 6): Bath Co.: Starr Chapel Disappointment Cave; Scott Co.: Hill Cave; Wythe Co.: Sam Six Cave. Remarks: This predaceous mite is often found in leaf litter in caves. It presumably preys upon other mites and especially Collembola. It is also known from caves in West Virginia (Zacharda, 1985). Bath Traegaardhia paralleloseta (Zacharda 1985) Type Locality: Sam Six Cave, Wythe County, Virginia Other Virginia Records: Known only from the type locality (Fig. 6). Remarks: According to Zacharda et al. (2010), T. paralleloseta shows extensive morphological modification for cave life, such as elaborated sensory structures (solenidia) on the tibia and elsewhere. It is known from one specimen! Order Araneae Family Linyphiidae Anthrobia coylei Miller 2005a Type Locality: Benedicts Cave, Greenbrier County, West Virginia Virginia Records (Fig. 7): Alleghany Co.: Wares Cave; Bath Co.: Clarks Cave. Remarks: Formerly assigned to A. monmouthia, Miller (2005a) recognized these records and a number of additional records from Greenbrier, Mercer, Monroe, and Pocahontas counties in West Virginia (Fong et al., 2007) as being a distinct species. Anthrobia monmouthia Tellkampf 1844 Type Locality: Mammoth Cave, Edmonson County, Kentucky Virginia Records (Fig. 7): Scott Co.: Harris Pit Cave, McNew Cave; Smyth Co.: Buchanon Saltpetre Cave. Remarks: This widespread troglobiotic spider is also found in many caves in Kentucky and Tennessee. Bathyphantes weyeri (Emerton 1875) Type Locality: Grand Caverns, Virginia Other Virginia Records: Known in Virginia only from the type locality (Fig. 7). Remarks: This species is widely distributed in caves Augusta County, from Wisconsin and Arkansas east to Pennsylvania, Virginia, and West Virginia (Ivie, 1969), but it is found in relatively few caves in any region. Paquin & Dupérré (2003) report this species from surface localities in Canada. It may be that the species is troglobiotic in southern parts of its range, such as Virginia, and troglophilic in northern parts of its range, but this conjecture needs further field studies for confirmation. As with other Linyphiidae, it is a predator, probably of mites and Collembola. Tslandiana muma Ivie 1965 Type Locality: Buck Hill Cave, Rockbridge County, Virginia Other Virginia Records: Known in Virginia only from the type locality (Fig. 7). Remarks: This is a very rare species with an enigmatic distribution. In addition to the type locality, it is only known from one cave in Colbert County, Alabama (Ivie, 1965). Oreonetides beattyi Paquin, Dupérré, Buckle & Lewis 2009 Type Locality: Smiths Folly Cave, Lawrence County, Indiana Virginia Records (Fig. 7): Montgomery Co.: Vicker Road Cave; Tazewell Co.: Rosenbaums Water Cave. Remarks: This species is also known from caves in Indiana, Maryland, Pennsylvania, and Tennessee. There is variation in the amount of eye reduction, but it is reduced in all populations (Paquin et al., 2009). Phanetta subterranea (Emerton 1875) Type Locality: Wyandotte Cave, Crawford Co., Indiana Virginia Records (Fig. 8): Alleghany Co.: Blue Spring Cave, Fudges Cave, Island Ford Cave, Lowmoor Cave, Wares Cave; Augusta Co.: Fountain Cave, Madison Saltpetre Cave; Bath Co.: Boundless Cave, Breathing Cave, Butler-Sinking Creek Cave, Cave Run Pit Cave, Clarks Cave, Dunns Cave, Russells Reserve Cave, Starr Chapel Saltpetre Cave; Bland Co.: Hamilton Cave; Botetourt Co.: Peerys Saltpetre Cave, Thomas Cave; Craig Co.: Rufe Caldwell Cave; Frederick Co.: Ogdens Cave; Giles Co.: Clover Hollow Cave, Giant Caverns Cave, Harris Cave, New River Cave, Salamander Cave, Smokehole Cave, Starnes Cave, Straleys No. 1 Cave, Tawneys Cave; Highland Co.: Vandevander Cave; Lee Co.: Bowling Cave, Cudjo’s Cavern (part of the Gap Cave System), Cumberland Gap Saltpetre Cave (part of the Gap Cave System), Gibson-Frazier Cave, Lucy Beatty Cave, Molly Wagle Cave, Olinger Cave, Spangler Cave; Montgomery Co.: Slussers Chapel Cave; Page Co.: Luray Caverns; 16 BANISTERIA # Fontigens bottimeri a Fontigens mortisoni g Fontigens tartarea @ Holsingera unthanksensis M Big Stone Gap a IKilometers Fig. 5. Distribution of Fontigens bottimeri, F. morrisoni, F. tartarea, and Holsingeria unthanksensis in Virginia. Gray dots are all the aquatic sampling sites. Winchester * Poecilophysis extraneostella ; : A Poecilophysis weyerensis Harrisonburg g Rhagidia varia ? @ Jraegaardhia paralleloseta Big Stone Gap oe GS 150 es: ae rere hilo meters Fig. 6. Distribution of Poecilophysis extraneostella, P. weyerensis, Rhagidia varia, and Traegaardhia paralleloseta in Virginia. Gray dots are all the terrestrial sampling sites. Winchester # Anthrobia coylei a Anthrobia monmouthia a Bathyphantes weyen @ /slandiana muma w Oreonetides beattyi Big Stone Gap at et GORGE, 5 * Fig. 7. Distribution of Anthrobia coylei, A. monmouthia, Bathyphantes weyeri, Islandiana muma, and Oreonetides beattyi in Virginia. Gray dots are all the terrestrial sampling sites. 150 eee hKilometers NO. 42, 2013 HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 17 Pulaski Co.: Sam Bells Cave; Roanoke Co.: Goodwins Cave, Hodges No. 1 Cave, Millers Cove Cave; Rockingham Co.: Deer Drop Cave; Massanutten Caverns; Melrose Caverns, Stephens Cave; Russell Co.: Banners Corner Cave, Concrete Tank Cave, Daugherty Cave, Densmore Hill Cave, Dorton Spring Cave, Echo Lake Cave, Jessie Cave, Litton Cave, Mountain View Cave, Porgie Bundys Cave, Stewart Monks Cave, Tumbez Cave; Scott Co.: Carlinbrook Cave, Cave Hollow Cave, Grigsby Cave, Herron No. 1 Cave, Hill Cave, Jessee Branch Cave, Kerns No. | Cave, Sounding Cave, Valley Creek Pirate Cave, Winding Stair Cave; Smyth Co.: Beaver Creek Cave; Tazewell Co.: Cassell Farm Cave No. 2, Chimney Rock Cave, Higginbotham No. 1 Cave, Lawson Cave, Steeles Cave; Warren Co.: Skyline Caverns; Washington Co.: Hall Bottom No. 1 Cave, Perkins Cave, Red Wolf Cave; Wise Co.: Kelly Cave, Wildcat Saltpetre Cave; Wythe Co.: Picketts Cave. Remarks: This is the most common and widespread troglobiotic spider in Virginia caves. It is also common in caves throughout the eastern United States and ranges from Pennsylvania south to Georgia and Alabama and west to Missouri and Illinois. Porrhomma cavernicola (Keyserling 1886) Type Locality: Reynolds Cave, Barren County, Kentucky Virginia Records (Fig. 9): Augusta Co.: Fountain Cave, Glade Cave, Madison Saltpetre Cave; Bath Co.: Clarks Cave, Crossroads Cave, Porters Cave, Witheros Cave; Bland Co.: Banes Spring Cave, Coon Cave; Craig Co.: New Castle Murder Hole, Rufe Caldwell Cave; Frederick Co.: Beans Cave; Giles Co.: Clover Hollow Cave; Lee Co.: Fisher Cave, Unthanks Cave; Montgomery Co.: Aunt Nellies Hole Cave; Page Co.: Luray Caverns, Ruffners No. 1 Cave; Roanoke Co.: Dixie Caverns; Rockbridge Co.: Bell Cave, Rock Hill Cave; Rockingham Co.: 3-D Maze Cave; Russell Co.: Cartop Cave, Maggie Baker Cave; Scott Co.: Abrams Cave, Little Duck Cave, Queens Cave; Smyth Co.: Buchanan Saltpetre Cave; Tazewell Co.: Gully Cave, Lawson Cave; Washington Co.: Robinson Cave; Wise Co.: Parsons Cave; Wythe Co.: Sam Six Cave. Remarks: This species, like P. subterranea, occurs over much of the eastern United States. It ranges from Pennsylvania south to Georgia and west to Missouri and Oklahoma. These two species have among the largest ranges of any North American troglobiont or stygobiont. This large range would seem to indicate that it is either a species complex or is not really a troglobiont, and thus able to disperse through and survive in surface habitats. However, it is practically unknown from surface habitats and Miller (2005b) found no morphological differences among populations in different parts of its range. It is possible that this taxon may encompass cryptic species detectable only from DNA sequencing. Family Nesticidae Nesticus holsingeri Gertsch 1984 Type Locality: Pond Cave, Scott County, Virginia Other Virginia Records (Fig. 10): Lee Co.: Bowling Cave, Gibson No. 1 Cave; Montgomery Co.: Eve Einstein Cave; Pulaski Co.: Collier Cave; Russell Co.: Austins Hilltop Cave, Carpenters Cave, Concrete Tank Cave, Ferguson Cave, Stewart Monks Cave; Scott Co.: Alley Cave, Blair-Collins Cave, Coley No. 2 Cave, Jackson Cave, McDavids Cave, Pond Cave, Taylor No. 1 Cave; Washington Co.: Thomas No. 1 Cave; Wise Co.: Burton Cave. Remarks: This species is known only from caves in southwestern Virginia. Eyes are present but reduced. Nesticus paynei Gertsch 1984 Type Locality: Reeders Cave, Tennessee Virginia Record (Fig. 11): Scott Co.: Wolfe Cave. Remarks: This species is common in caves in eastern Tennessee. Although it has well developed eyes, it is known only from caves (Gertsch, 1984). Anderson County, Nesticus tennesseensis (Petrunkevitch 1925) Type Locality: Indian Cave, Grainger Tennessee Virginia Records (Fig. 11): Alleghany Co.: Rumbolds Cave; Craig Co.: Fish Hatchery Cave, Walkthrough Cave; Giles Co.: Ballards Cave, Doe Mountain Cave, Giant Caverns Cave, Glenlyn Cave, Harris Cave, Starnes Cave, Straleys No. 1 Cave, Sugar Run Cave System; Highland Co.: Locust Cave; Lee Co.: Cattle Cave; Russell Co.: Broken Formation Cave, Dorton Spring Cave; Scott Co.: Booher Cave; Smyth Co.: Sugar Grove No. 10 Cave; Tazewell Co.: Cassell Farm No. 2 Cave, Chimney Rock Cave, Fallen Rock Cave, Hugh Young Cave, Steeles Cave; Wise Co.: Hairy Hole Cave. Remarks: In addition to the localities listed above, this species is known from caves in West Virginia and Tennessee. According to Gertsch (1984), it is morphologically variable and is sometimes found in surface habitats in Virginia and West Virginia, but not in Tennessee. However, nearly all records are from caves and therefore we consider it a troglobiont. It shows variable amounts of eye and pigment reduction. County, 18 BANISTERIA + Phanetta subterranea 150 a Tkilometers Fig. 8. Distribution of Phanetta subterranea in Virginia. Gray dots are all the terrestrial sampling sites. Winchester “8 Harrisonburg + = Porrhomma cavernicola Ccee———Kilometers Fig. 9. Distribution of Porrhomma cavernicola in Virginia. Gray dots are all the terrestrial sampling sites. Winchester *# Nesticus holsingeri Cer Kilometers Fig. 10. Distribution of Nesticus holsingeri in Virginia. Gray dots are all the terrestrial sampling sites. NO. 42, 2013 HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 19 a # Nesticus payne/ A. Nesticus tennesseensis SY: oka 150 —*“——X—T Kilometers Big Stone Gap Fig. 11. Distribution of Nesticus paynei and N. tennesseensis in Virginia. Gray dots are all the terrestrial sampling sites. # Hesperochernes mirabilis Harrisonburg A Kleptochthonius anophthalmus B Apochthonius coecus kilometers Fig. 12. Distribution of Apochthonius coecus, Hesperochernes mirabilis, and Kleptochthonius anophthalmus in Virginia. Gray dots are all the terrestrial sampling sites. # Kleptochthonius gertschi Harrisonburg A Apochthonius holsingeri g Kleptochthonius similis 150 Kilometers Fig. 13. Distribution of Apochthonius holsingeri, Kleptochthonius gertschi, and K. similis in Virginia. Gray dots are all the terrestrial sampling sites. 20 BANISTERIA Order Pseudoscorpiones Family Chernetidae Hesperochernes mirabilis (Banks 1895) Type Locality: “Cave at Pennington Gap”, Lee County, Virginia Other Virginia Records (Fig. 12): Bath Co.: Cave Run Pit Cave; Giles Co.: Smokehole Cave; Highland Co.: Vandevander Cave; Lee Co.: Gilley Cave, Secret Cave; Roanoke Co.: Goodwins Cave; Rockingham Co.: Eggleston Cave; Russell Co.: Boy Scout Cave, Ferguson Hollow Cave; Scott Co.: Basil Duncan Cave, Big Spiders in a Littlke Maze Cave, Canyon to Nowhere Cave, Creek No. 1 Cave, Darty Cave, McCulley Cave, W.R. Combs Cave; Washington Co.: Davenport Cave, Three Chambers Cave. Remarks: This species is also known from caves throughout the southeastern U.S. (Holsinger & Culver, 1988; Niemiller & Zigler, 2013). It appears to be morphologically unmodified for subterranean life, but it has not been found outside of caves. Family Chthoniidae Apochthonius coecus (Packard 1884) Type Locality: Grand Caverns, Virginia Other Virginia Record (Fig. 12): Augusta Co.: Madison Saltpetre Cave. Remarks: Chthontidae, including all the species listed below, are typically found in damp places, under rocks or wood. Augusta County, Apochthonius holsingeri Muchmore 1967 Type Locality: Cave Run Pit Cave, Bath County, Virginia Other Virginia Record (Fig. 13): Alleghany Co.: Blue Spring Cave. Kleptochthonius anophthalmus Muchmore 1970 Type Locality: Porters Cave, Bath County, Virginia Other Virginia Records: Known only from the type locality (Fig. 12). Kleptochthonius binoculatus Muchmore 1974 Type Locality: Hill Cave, Scott County, Virginia Other Virginia Records (Fig. 14): Scott Co.: Dancing Rock Cave, Jack Hartstock Cave, Jimmys Suck Hole Cave. Remarks: For a pseudoscorpion, it has a_ large distribution, being known from four caves! NO. 42, 2013 Kleptochthonius gertschi Malcolm & Chamberlin 1961 Type Locality: Gilley Cave, Lee County, Virginia Other Virginia Records: Known only from the type locality (Fig. 13). Kleptochonthius lutzi Malcolm & Chamberlin 1961 Type Locality: Cudjo’s Cavern (part of the Gap Cave System), Lee County, Virginia Other Virginia Records: Known only from the type locality (Fig. 14). Kleptochthonius proximosetus Muchmore 1976 Type Locality: Gallohan No. 1 Cave, Lee County, Virginia Other Virginia Records: Known only from the type locality (Fig. 15). Kleptochthonius regulus Muchmore 1970 Type Locality: Fallen Rock Cave, Tazewell County, Virginia Other Virginia Record (Fig. Higginbotham No. 1 Cave. 15): Tazewell Co.: Kleptochthonius similis Muchmore 1976 Type Locality: Sweet Potato Cave, Tazewell County, Virginia Other Virginia Records: Known only from the type locality (Fig. 13). Mundochthonius holsingeri Benedict & Malcolm 1974 Type Locality: Helsley Cave, Shenandoah County, Virginia Other Virginia Records: Known only from the type locality (Fig. 14). Family Neobisiidae Lissocreagris valentinei (Chamberlin 1962) Type Locality: Cudjo’s Cavern (part of the Gap Cave System), Lee County, Virginia Other Virginia Record (Fig. 15): Lee Co.: Indian Cave. Family Syanaridae Chitrella superba Muchmore 1973 Type Locality: Maddens Cave, Shenandoah County, Virginia Other Virginia Records: Known only from the type locality (Fig. 15). Remarks: This rare species is less morphologically modified than most of the species in the family Chthoniidae. HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA Winchester * Kleptochthonius binoculatus Harrisonburg é Mundochthonits holsingen g Kleptochthonius lutzi 150 eT Kilometers Fig. 14. Distribution of Kleptochthonius binoculatus, K. lutzi, and Mundochthonius holsingeri in Virginia. Gray dots are all the terrestrial sampling sites. Winchester * Kleptochthonius regulus A Kleptochthonius proximosetus gi Lissocreagris valentinei @ Chitrella superba ———————————TKilometers Fig. 15. Distribution of Chitrella superba, Kleptochthonius proximosetus, K. regulus, and Lissocreagris valentinei in Virginia. Gray dots are all the terrestrial sampling sites. Winchester * Pseudotremia contorta Harrisonburg a Pseudotremia cavermarum g Pseudotremia cerberus Big Stone Gap 150 kilometers Fig. 16. Distribution of Pseudotremia cavernarum, P. cerberus, and P. contorta in Virginia. Gray dots are all the terrestrial sampling sites. 21 22 BANISTERIA Class Diplopoda Order Chordeumatida Family Cleidogonidae A total of 19 troglobiotic Pseudotremia millipeds have been found in Virginia caves, 16 of them endemic to the state. Pseudotremia cavernarum Cope 1869 Type Locality: Erhart Cave, Montgomery County Other Virginia Records (Fig. 16): Montgomery Co.: Aunt Nellies Hole, Daves Cave, Heartbeat Cave, Unnamed Cave A. Remarks: Listed as State Endangered in Virginia, the type locality was destroyed by quarrying. Its range is limited to approximately 40 km? in a developing suburban setting. Pseudotremia cerberus Shear 2011 Type Locality: Wares Cave, Virginia Other Virginia Records: Known only from the type locality (Fig. 16). Alleghany County, Pseudotremia contorta Shear 2011 Type Locality: Mountain Grove Saltpetre Cave, Bath County, Virginia Other Virginia Records (Fig. 16): Bath Co.: Little Mountain Cave. Remarks: Considered an epigean or somewhat troglophilic species by Shear (2011), it is currently known only from caves. Pseudotremia culveri Shear 2011 Type Locality: Fisher Cave, Lee County, Virginia Other Virginia Records: Known only from the type locality (Fig. 17). Remarks: Occurs in the middle of the range of the more widespread P. nodosa. Pseudotremia deprehendor Shear 1972 Type Locality: Feathers Cave, Anderson County, Tennessee Virginia Record (Fig. 17): Wise Co.: Hairy Hole. Remarks: Known from a few caves in eastern Tennessee. Pseudotremia fergusoni Shear 2011 Type Locality: Stones No. 1 Cave, Smyth County, Virginia Other Virginia Records: Known only from the type locality (Fig. 17). NO. 42, 2013 Pseudotremia hubbardi Shear 2011 Type Locality: Crackers Neck Saltpetre Cave, Wise County, Virginia Other Virginia Records (Fig. 18): Wise Co.: Kelly Cave, Little Kennedy Cave, Omega Cave System, Parsons Cave, Ridge Cave, Wildcat Saltpetre Cave. Remarks: Known only from caves near the eastern end of the Powell Valley. Pseudotremia inexpectata Shear 2011 Type Locality: Davault Cave, Lee County, Virginia Other Virginia Records: Known only from the type locality (Fig. 18). Remarks: Occurs in the midst of the range of the more widespread P. nodosa. Pseudotremia jaculohamatum Shear 2011 Type Locality: Walker Mountain Saltpetre Cave, Washington County, Virginia Other Virginia Records (Fig. 17): none confirmed. Remarks: Specimens from three other Washington County caves (Clarke Cave, Millard Cave, and Three Chambers Cave) are likely P. jaculohamatum, but confirmation is not possible due to curatorial problems (Shear, 2011). Pseudotremia johnholsingeri Shear 2011 Type Locality: Carter Cave, Lee County, Virginia Other Virginia Records: Known only from the type locality (Fig. 19). Pseudotremia loomisi Shear 2011 Type Locality: Clarks Cave, Bath County, Virginia Other Virginia Records: Known only from the type locality (Fig. 18). Remarks: According to Shear (2011), this species appears to be a troglophile rather than a true troglobiont, but it is known only from a cave so we have included it. Pseudotremia nodosa Loomis 1939 Type Locality: English Cave, Tennessee Virginia Records (Fig. 20): Lee Co.: Cavin Cave, Cedar Hill Cave, Cope Cave, Crouse Cave, Gallohan No. 1 Cave, Gallohan No. 2 Cave, Gibson-Frazier Cave, Gilley Cave, Indian Burial Cave, Jones Saltpetre Cave, Knapper Cave, Litton No. 1 Cave, Lucy Beatty Cave, Longs Cave, Molly Waggle Cave, Smith Cave, Spangler Cave, Surgener Cave, Sweet Potato Cave, Thompson Cedar Cave, Unthanks Cave, Young-Fugate Cave. Remarks: This species is widespread Tennessee and Lee County, Virginia. Claiborne County, in eastern HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 23 Winchester * Pseudotremia jaculohamatum a Pseudotremia fergusoni g Pseudotremia deprehendor @ Pseudotremia culveri 150 Ci itometers Fig. 17. Distribution of Pseudotremia culveri, P. deprehendor, P. fergusoni, and P. jaculohamatum in Virginia. Gray dots are all the terrestrial sampling sites. ey, Winchester + Pseudotremia hubbardi ps Harrisonburg A Pseudotremia inexpectata mg Pseudotremia loamisi S fa Roanoke 150 Kilometers Fig. 18. Distribution of Pseudotremia hubbardi, P. inexpectata, and P. loomisi in Virginia. Gray dots are all the terrestrial sampling sites. Winchester + Pseudotremia ryensis Harrisonburg a Pseudotremia orndorffi g Pseudotremia joiinholsingen Big Stone Ga eg OO Fig. 19. Distribution of Pseudotremia johnholsingeri, P. orndorffi, and P. ryensis in Virginia. Gray dots are all the terrestrial sampling sites. 150 kilometers 24 BANISTERIA Pseudotremia orndorffi Shear 2011 Type Locality: Sugar Run Cave System, Giles County, Virginia Other Virginia Records (Fig. 19): Craig Co.: Rufe Caldwell Cave; Giles Co.: Starnes Cave. Pseudotremia peponocranium Shear 2011 Type Locality: Giant Caverns, Giles County, Virginia Other Virginia Record (Fig. 20): Giles Co.: Scooters Boneyard Cave. Pseudotremia piscator Shear 2011 Type Locality: Fisher Cave, Lee County, Virginia Other Virginia Records: Known only from the type locality (Fig. 20). Remarks: Relative to other cave-inhabiting Pseudotremia, this species has large eyes (21 contiguous ocelli; Shear, 2011). Perhaps, it will prove to be a troglophile. Pseudotremia ryensis Shear 2011 Type Locality: Franklin Cave, Virginia Other Virginia Records (Fig. 19): Scott Co.: Flannery Cave, McDavids Cave, Riggs Chapel Cave, Thatcher Swallet Cave. Remarks: The known distribution is limited to the Rye Cove area of Scott County. Scott County, Pseudotremia salifodina Shear 2011 Type Locality: Jones Saltpetre Cave, Lee County, Virginia Other Records: Known only from the type locality (Fig. 2A Remarks: This species shows little if any modification for subterranean life (Shear, 2011), but it is known only from a cave. Pseudotremia tuberculata Loomis 1939 Type Locality: Cassell Farm Cave No. 2, Tazewell County, Virginia Other Virginia Records (Fig. 21): Tazewell Co.: Bowens Cave, Chimney Rock Cave, Fallen Rock Cave, Lawson Cave, Stonley Cave. Remarks: Known only from caves, but it retains eyes and pigment. Pseudotremia valga Loomis 1943 Type Locality: Cudjo’s Cavern (part of the Gap Cave System), Lee County, Virginia Other Virginia Record (Fig. 21): Lee Co.: Young- Fugate Cave. Remarks: Also known from Station Creek Cave in bordering Claiborne County, Tennessee. NO. 42, 2013 Family Trichopetalidae Zygonopus packardi Causey 1960 Type Locality: Pattons Cave, Monroe County, West Virginia Virginia Records (Fig. 22): Augusta Co.: Blue Hole Cave; Bland Co.: Coon Cave, Hamilton Cave, Newberry-Bane Cave, Repass Saltpetre Cave; Botetourt Co.: Peerys Saltpetre Cave; Craig Co.: Loneys Cave, Rufe Caldwell Cave; Giles Co.: Canoe Cave, Clover Hollow Cave, Doe Mountain Cave, Echols Cave, Giant Caverns, New River Cave, Salamander Cave, Starnes Cave, Straleys No. | Cave, Tawneys Cave; Montgomery Co.: Old Mill Cave, Slussers Chapel Cave; Page Co.: Luray Caverns; Pulaski Co.: Colliers Cave, Fifty-foot Hell Cave, James Cave, Sam Bells Cave; Roanoke Co.: Dixie Caverns; Wythe Co.: Early Cave, Gardners Cave, Sam Six Cave. Remarks: This is the most widespread of the Zygonopus species. It occurs syntopically with Z. whitei in Luray Caverns. The genus was revised by Shear (2010). Zygonopus weyeriensis Causey 1960 Type Locality: Grand Caverns, Virginia Other Virginia Records (Fig. 22): Augusta Co.: Madison Saltpetre Cave; Bath Co.: Boundless Cave, Breathing Cave, Butler-Sinking Creek Cave, Porters Cave, Russells Reserve Cave, Starr Chapel Saltpetre Cave; Rockbridge Co.: Billy Williams Cave, Grahams Cave. Remarks: The entrance to Billy Williams Cave was covered during construction of Interstate 81. Augusta County, Zygonopus whitei Causey 1960 Type Locality: Glade Cave, Augusta County Virginia Other Virginia Records (Fig. 22): Page Co.: Luray Caverns, Ruffner No. 1 Cave; Rockingham Co.: 3-D Maze Cave, Bakers Cave, Cedar Hill Cave, Endless Caverns, Melrose Caverns, New Market Cave, Orebaugh Cave, Ponderosa Fissure, Stephens Cave; Shenandoah Co.: Maddens Cave, Shenandoah Caverns, Shenandoah Wild Cave. Family Zosteractinidae Ameractis satis Causey 1959 Type Locality: Indian Cave, White County, Tennessee Virginia Records (Fig. 23): Russell Co.: Ferguson Hollow Cave; Washington Co.: Millard Cave. Remarks: This species occurs in Alabama, Georgia, North Carolina, Tennessee, and Virginia and is the most widely distributed troglobiotic milliped in North America (Shear & Hubbard, 1998). HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 25 Winchester + Pseudcotremia nodosa “Harrisonburg a Pseudotremia peponocranium g $FPseudotremia piscator kilometers Fig. 20. Distribution of Pseudotremia nodosa, P. peponocranium, and P. piscator in Virginia. Gray dots are all the terrestrial sampling sites. ace) Winchester fie PAR. £ # Pseudotremia valga A Pseudoiremia tuberculata mg Pseudotremia salifodina Harrisonburg Big Stone Gap 150 CK ilometers Fig. 21. Distribution of Pseudotremia salifodina, P. tuberculata, and P. valga in Virginia. Gray dots are all the terrestrial sampling sites. Zygonopus packardi Harrisonburg A Zygonopus weyernensis Bg Zygonopus whitei a ’ Ags tS 150 CI Kilometers Fig. 22. Distribution of Zygonopus packardi, Z. weyeriensis, and Z. whitei in Virginia. Gray dots are all the terrestrial sampling sites. 26 BANISTERIA Class Chilopoda Order Lithobiomorpha Family Lithobiidae Nampabius turbator Crabill 1952 Type Locality: Lowmoor Cave, Alleghany County, Virginia Other Virginia Record (Fig. 24): Alleghany Co.: Island Ford Cave. Class Hexapoda Order Collembola Family Entomobryidae Pseudosinella bona Christiansen & Bellinger 1996a Type Locality: Goodwins Cave, Roanoke County, Virginia Other Virginia Records (Fig. 25): Washington Co.: Lowes Cave; Wythe Co.: Chockedfullofit Cave, Williams Cave. Pseudosinella erehwon Christiansen & Bellinger 1996a Type Locality: Canyon to Nowhere Cave, Scott County, Virginia Other Virginia Records (Fig. 26): Lee Co.: Coffin Cave; Russell Co.: Stewart Monks Cave; Washington Co.: Clarke Cave; Wythe Co.: Connor Valley Cave. Pseudosinella extra Christiansen & Bellinger 1996a Type Locality: Jack Cave, Scott County, Virginia Other Virginia Records (Fig. 25): Scott Co.: Basil Duncan Cave, Lane Caves; Wythe Co.: Brown Cave. Pseudosinella flatua Christiansen & Bellinger 1996a Type Locality: Blowing Springs Cave, Swain County, North Carolina Virginia Records (Fig. 25): Russell Co.: Broken Formation Cave; Washington Co.: Leonard Cave, Walker Mountain Saltpetre Cave. Pseudosinella gisini virginia Christiansen & Bellinger 1996a Type Locality: Spangler Cave, Lee County, Virginia Other Virginia Records (Fig. 25): Lee Co.: Indian Burial Cave; Russell Co.: Daugherty Cave. Remarks: One of three subspecies of Pseudosinella gisini, with P. g. gisini in the Greenbrier Valley of West Virginia and P. g. carolina from western North Carolina. NO. 42, 2013 Pseudosinella granda Christiansen & Bellinger 1996a Type Locality: Grand Caverns, Augusta County, Virginia Other Virginia Records (Fig. 27): Scott Co.: Darty Cave, John W. Fugate Cave; Washington Co.: Whispering Breeze Cave; Wise Co.: Burtons Cave, Huff caves; Wythe Co.: Lone Ash No. 2 Cave. Pseudosinella hirsuta Delamare Deboutteville 1949 Type Locality: Tennessee Caverns (Crystal Caves), Hamilton County, Tennessee Virginia Records (Fig. 27): Lee Co.: Cliff Cave, Cudjo’s Cavern (part of the Gap Cave System), Skylight Cave, Young-Fugate Cave. Remarks: A widely distributed troglobiotic springtail in the central and southeastern U.S. Pseudosinella ops Christiansen & Bellinger 1998 Type Locality: Herrons Echo Hall Cave, Scott County, Virginia Other Virginia Records (Fig. 26): Alleghany Co.: Island Ford Cave; Montgomery Co.: Adams Cave; Russell Co.: Buffalo Hollow Hill Cave; Washington Co.: Leonard and Thomas No. | Cave. Pseudosinella orba Christiansen 1960a Type Locality: Morrill Cave, Sullivan County, Tennessee Virginia Records (Fig. 28): Bath Co.: Starr Chapel Saltpetre Cave; Bland Co.: Hamilton Cave; Botetourt Co.: Catawba Murder Hole; Craig Co.: Rufe Caldwell Cave; Giles Co.: Starnes Cave, Sugar Run Cave System; Lee Co.: Cavin Cave, Gallohan No. 1 Cave, Long Cave, Smith Cave, Sweet Potato Cave; Pulaski Co.: Colliers Cave, Sam Bells Cave; Roanoke Co.: Smiths No. 1 Cave; Russell Co.: Austins Hilltop Cave, Concrete Tank Cave, Mountain View Cave, Porgie Bundys Cave, Stewart Monks Cave; Scott Co.: Blair- Collins Cave, Littlke Duck Cave, Sikes Cave; Smyth Co.: Buchanan Saltpetre Cave, Dead Air Cave, Interstate-81 Cave, Little Saltpetre Cave, Tilson Saltpetre Cave; Tazewell Co.: Fallen Rock Cave, Gully Cave, Higginbotham No. 1 Cave; Washington Co.: Fraleys Cave; Wise Co.: Wildcat Saltpetre Cave; Wythe Co.: Brown Cave, Collins Cove Cave, Macs Cave. Remarks: Also known from caves in North Carolina, Tennessee, and West Virginia. Pseudosinella testa Christiansen & Bellinger 1980 Type Locality: Neely Farm Cave, Mercer County, West Virginia Virginia Record (Fig. 27): Scott Co.: Canyon to Nowhere Cave. Remarks: Also known from caves in Arkansas. HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 27 # Ameractis satis Kilometers Fig. 23. Distribution of Ameractis satis in Virginia. Gray dots are all the terrestrial sampling sites. * Nampabius turbator Fig. 24. Distribution of Nampabius turbator in Virginia. Gray dots are all the terrestrial sampling sites. # Pseudosinella bona A Pseudosinella extra gw Pseudosinella gisini virginia @ Pseudosinella flatua ————————Nilometers Fig. 25. Distribution of Pseudosinella bona, P. extra, P. flatua, and P. gisini virginia in Virginia. Gray dots are all the terrestrial sampling sites. 28 BANISTERIA Winchester Harrisonburg Pseudosinella erehwon a Pseudosinella ops M Big Stone Gap 150 Ce Kilometers Fig. 26. Distribution of Pseudosinella erehwon and P. ops in Virginia. Gray dots are all the terrestrial sampling sites. Winchester + Pseudosinella granda ; @ Pseudosinella hirsuta Harrisonburg gw Pseudosinella vespera A Pseudosinella testa N Big St — iometers Fig. 27. Distribution of Pseudosinella granda, P. hirsuta, P. testa, and P. vespera in Virginia. Gray dots are all the terrestrial sampling sites. Winchester + Pseudosinella orba A Sinella barri Big ee aa Fig. 28. Distribution of Pseudosinella orba and Sinella barri in Virginia. Gray dots are all the terrestrial sampling sites. 150 eee NKilometers NO. 42, 2013 HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 29 Pseudosinella vespera Christiansen & Bellinger 1996a Type Locality: Bat Cave, Rutherford County, North Carolina Virginia Record (Fig. 27): Bland Co.: Cedar Cliff Saltpetre Cave. Remarks: Known from one other North Carolina cave. Sinella barri Christiansen 1960b Type Locality: Inman Cave, Perry County, Tennessee Virginia Records (Fig. 28): Alleghany Co.: Arritt Mill Tunnel Cave; Giles Co.: Echols Cave, Parsells Cave; Scott Co.: Herrons Echo Hall Cave, McCulley Cave, Wininger Cave; Warren Co.: Bee Tree Cave; Washington Co.: Fraleys Cave, Vickers Cave; Wythe Co.: Picketts Cave. Remarks: Widespread through Eastern and Midwestern states, it was originally listed by Holsinger & Culver (1988) as a troglophile, but because more than 90 percent of its localities are in caves, we list it here as a troglobiont. Sinella hoffmani Wray 1952 Type Locality: Lowmoor Cave, Alleghany County, Virginia Other Virginia Records (Fig. 29): Alleghany Co.: Blowhole Cave, Blue Spring Cave, Canterbury Cave, Island Ford Cave, Matthews No. 2 Cave, Rumbolds Cave, Walking Cave, Wares Cave; Bath Co.: Blowing Cave, Boundless Cave, Breathing Cave, Butler-Sinking Creek Cave, Cave Run Pit, Bicentennial Cave, Clarks Cave, Crossroads Cave, Dunns Cave, Little Mountain Cave, Little Starr Chapel Cave, Mays Mountain Saltpetre Cave, Porters Cave, Serpent Cave, Starr Chapel Saltpetre Cave, Witheros Cave; Botetourt Co.: Peerys Saltpetre Cave; Highland Co.: Cabin Cave, Helictite Cave, Hiner Cave, Secret Anthodite Cave, Sugar Maple Cave, Vandevander Cave, Water Sinks Cave; Lee Co.: Glen Olingers Cave; Roanoke Co.: Goodwins Cave; Rockbridge Co.: Buck Hill Cave, Doll House Cave, Turkey Hill Cave; Rockingham Co.: Devil’s Hole, Raindrops a Dripping Cave; Russell Co.: Cascade Pit; Scott Co.: M.J. Mann Cave, Cave Hollow Cave; Tazewell Co.: Stonley Cave; Washington Co.: Three Chambers Cave; Wise Co.: Bloomer Cave; Wythe Co.: Connor Valley Cave. Remarks: Widespread in the central and southern Appalachians. Family Hypogastruridae Schaefferia hubbardi Thibaud 1995 Type Locality: Hiner Cave, Highland County, Virginia Other Virginia Records (Fig. 30): Pulaski Co.: Maze Cave; Russell Co.: Family Secret Cave; Scott Co.: Greears Sweet Potato Cave. Schaefferia valentini Thibaud 1996 Type Localities: Grigsby and Herrons Echo Hall caves, Scott County, Virginia Other Virginia Records (Fig. 30): Giles Co.: Echols Cave; Pulaski Co.: Colliers Cave, Maze Cave; Russell Co.: Amos Cave, Densmore Hill Cave, Sentinel Oak Cave, Smiths Drop Cave; Scott Co.: Little Duck Cave, M.J. Mann Cave, Prince Albert in a Cave, Queens Cave, Sikes Cave; Washington Co.: Robinson Cave; Wythe Co.: Blue Grotto Cave, Brown Cave, Collins Cove Cave, Early Cave, Sam Six Cave. Family Neanuridae Paleonura petebellingeri Palacios-Vargas & Benito 2007 Type Locality: Skyline Caverns, Warren County, Virginia Other Records: Known only from the type locality (Fig. 31). Speleonura kenchristianseni Palacios-Vargas & Benito 2007 Type Locality: Porters Cave, Bath County, Virginia Other Records: Known only from the type locality (Fig. 31). Remarks: This is the only species currently known in its genus. Family Oncopoduridae Oncopodura hubbardi Christiansen & Bellinger 1996a Type Locality: Reasors Cave, Lee County, Virginia Other Virginia Records (Fig. 32): Lee Co.: Burton Cave, Cattle Cave, Spangler Cave; Scott Co.: Carters Pit, Jessee Branch Cave. Remarks: Known only from the Upper Tennessee River basin in southwestern Virginia. Family Sminthuridae Pygmarrhopalites caedus (Christiansen & Bellinger 1996b) Type Locality: Catawba Murder Hole, County, Virginia Other Virginia Records (Fig. 32): Bath Co.: Little Starr Chapel Cave, Starr Chapel Saltpetre Cave. Remarks: Relatively unmodified for subterranean life, it is known only from caves in the Alleghany Highlands of Virginia. Botetourt 30 BANISTERIA Winchester + Sinella hoffmani Roanoke Pe Big Stone Gap op Gb 150 ae. 7 oo iitometers Fig. 29. Distribution of Sinella hoffmani in Virginia. Gray dots are all the terrestrial sampling sites. + Schaefferia valentini A Schaefferia hubbardi Kilometers Fig. 30. Distribution of Schaefferia hubbardi and S. valentini in Virginia. Gray dots are all the terrestrial sampling sites. * Paleonura petebellingeri A Speleonura kenchristianseni eee Iilometers Fig. 31. Distribution of Paleonura petebellingeri and Speleonura kenchristianseni in Virginia. Gray dots are all the terrestrial sampling sites. NO. 42, 2013 HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 31 Pygmarrhopalites carolynae (Christiansen & Bellinger 1996b) Type Locality: Wildcat Saltpetre Cave, Wise County, Virginia Other Virginia Records (Fig. 33): Alleghany Co.: Matthews No. 2 Cave; Augusta Co.: Staunton Quarry Cave; Bath Co.: Butler-Sinking Creek Cave, Little Starr Chapel Cave, Starr Chapel Saltpetre Cave; Botetourt Co.: Peerys Saltpetre Cave; Highland Co.: Marshalls Cave, Water Sinks Cave; Lee Co.: Spangler Cave; Russell Co.: Tumbez Cave; Shenandoah Co.: Crystal Caverns at Hupps Hill; Wythe Co.: Lone Ashe No. 2 Cave. Remarks: This species is highly ~ variable morphologically, and is also known from caves in Indiana. Pygmarrhopalites clarus (Christiansen 1966) Type Locality: Rockhouse Cave, Barry County, Missouri Virginia Records (Fig. 34): Bath Co.: Starr Chapel Saltpetre Cave; Montgomery Co.: Old Mill Cave; Wythe Co.: Campbell Cave, Gardners Cave, Sam Six Cave. Remarks: A strongly troglomorphic species, widespread through Eastern and Midwestern states. Pygmarrhopalites commorus (Christiansen & Bellinger 1996b) Type Locality: Stay High Cave, Giles County, Virginia Other Virginia Records (Fig. 34): Lee Co.: Burton Cave; Scott Co.: Queens Cave; Wise Co.: Big Kennedy Cave, Little Kennedy Cave. Remarks: Also known from a cave in Pocahontas Co., West Virginia. Pygmarrhopalites lacuna (Christiansen & Bellinger 1996b) Type Locality: Paxtons Cave, Alleghany County, Virginia Other Virginia Record (Fig. 32): Wise Co.: Kelly Cave. Pygmarrhopalites marshalli (Christiansen & Bellinger 1996b) Type Locality: Wininger Cave, Scott County, Virginia Other Virginia Records (Fig. 33): Giles Co.: Echols Cave; Highland Co.: Helictite Cave; Russell Co.: Burns Cave, Concrete Tank Cave, Daugherty Cave, Dorton Springs Cave, Mountain View Cave, Phreatic to a Fault Cave, Sentinel Oak Cave, Stewart Monks Cave; Scott Co.: Canyon to Nowhere Cave, Dancing Rock Cave, Darty Cave, Hartsock Cave, Jack Hartsock Cave, Jessee Branch Cave, Jim Robinson Cave, Sounding Cave; Smyth Co.: Little Saltpetre Cave; Tazewell Co.: Chimney Rock Cave; Washington Co.: Fraleys Cave; Wythe Co.: Minnie Corvins Cave. Remarks: Also known from caves in northeastern Tennessee. Pygmarrhopalites obtusus (Zeppelini & Christiansen 2003) Type Locality: Bakers Cave, Rockingham County, Virginia Other Virginia Record (Fig. 33): Rockingham Co.: Orebaugh Cave. Pygmarrhopalites pavo (Christiansen & Bellinger 1996b) Type Locality: Turkey Hill Cave, Rockbridge County, Virginia Other Virginia Records (Fig. 35): Rockbridge Co.: Ringbat Pothole Cave; Scott Co.: Bolling Cave, Wininger Cave. Remarks: Also known from caves in Tennessee and West Virginia. Pygmarrhopalites sacer (Christiansen & Bellinger 1996b) Type Locality: Little Starr Chapel Cave, Bath County, Virginia Other Virginia Records (Fig. 34): Bath Co.: Butler- Sinking Creek Cave, Barberry Cave, Starr Chapel Saltpetre Cave; Pulaski Co.: Colliers Cave; Tazewell Co.: Hugh Young Cave. Remarks: Also known from caves in Monroe County, West Virginia. Pygmarrhopalites sextus (Zeppelini & Christiansen 2003) Type Locality: Sam Six Cave, Wythe County, Virginia Other Virginia Record (Fig. 35): Wythe Co.: Irvin Cave. Remarks: Known only from one Virginia county. Pygmarrhopalites silvus (Christiansen & Bellinger 1996b) Type Locality: Woods-Terry Cave, Highland County, Virginia Other Virginia Records (Fig. 35): Bath Co.: Clarks Cave; Highland Co.: Hiner Cave. Order Diplura Family Campodeidae Litocampa cookei (Packard 1871) Type Locality: Mammoth Cave, Edmonson County, Kentucky 52 BANISTERIA Winchester * Oncopodura hubbardi Harrisonburg é& Pygmarrhopalites lacuna gm Pygmarrhopalites caedus Big Stone Gap Fig. 32. Distribution of Oncopodura hubbardi, Pygmarrhopalites caedus, and P. lacuna in Virginia. Gray dots are all the terrestrial sampling sites. 150 kilometers Winchester + Pygimarrhopalites marshalli Harrisonburg a Pygmarmopalites carolynae ga Pygmarrhopalites obtusus Big Stone Gap— fz 150 4. Kilometers Fig. 33. Distribution of Pygmarrhopalites carolynae, P. marshalli, and P. obtusus in Virginia. Gray dots are all the terrestrial sampling sites. Winchester # Pygmarrhopalites commons Harrisonburg a Pygmarrhopalites sacer g Pygmarrhopalites clarus Big Stone Gap— on a Fig. 34. Distribution of Pygmarrhopalites clarus, P. commorus, and P. sacer in Virginia. Gray dots are all the terrestrial sampling sites. 150 Kilometers NO. 42, 2013 HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 33 Virginia Records (Fig. 36): Lee Co.: Cave Road Cave, Gallohan No. | Cave, Garretts Cave, Jones Saltpetre Cave, Long Cave, Molly Wagle Cave, Mud Cave, Sweet Potato Cave, Young-Fugate Cave; Scott Co.: Cave Hollow Cave, Spurlock Cave; Wise Co.: Big Kennedy Cave, Crackers Neck Saltpetre Cave, Hairy Hole, Littlke Kennedy Cave, Parsons Cave, Rocky Hollow Cave. Remarks: There are a number of undescribed species in the genus. This species is also recorded from caves in east and middle Tennessee and _ south-central Kentucky. Litocampa fieldingae (Condé 1949) Type Locality: McClungs Cave, Greenbrier County, West Virginia Virginia Records (Fig. 36): Highland Co.: Aqua Cave. Remarks: Occurs in a number of caves in Monroe, Greenbrier, and Pocahontas counties, West Virginia (Fong et al., 2007). Litocampa hoffmani Ferguson 2009 Type Locality: Speedwell Cave No. 1, Wythe County, Virginia Other Virginia Records (Fig. 36): Montgomery Co.: Vicker Road Cave; Smyth Co.: Dead Air Cave, Interstate-81 Cave, Overbay Hill Cave, Roberts Cave, Ward Saltpetre Cave; Washington Co.: Brass Kettle Hole, Ira Miller Cave, Maddens Pyramid Cave; Wythe Co.: Groseclose No. 1 Cave. Litocampa pucketti Condé & Bareth 1996 Type Locality: Lawson Cave, Tazewell County, Virginia Other Virginia Records (Fig. 36): Bland Co.: Hamilton Cave; Giles Co.: Harris Cave; Tazewell Co.: Cassell Farm No. 2 Cave. Class Insecta Order Coleoptera Family Carabidae A total of 31 species of Pseudanophthalmus have been found in Virginia caves, 29 of them endemic to the state. Pseudanophthalmus avernus (Valentine 1945) Type Locality: Endless Caverns, Rockingham County, Virginia Other Virginia Records: Known only from the type locality (Fig. 37). Remarks: This beetle is typically seen on sediment banks along the stream in the non-commercial sections of this popular commercial cave. Pseudanophthalmus cordicollis Barr 1981 Type Locality: Little Kennedy Cave, Wise County, Virginia Other Virginia Records (Fig. 37): Wise Co.: Franklins Pit, Omega Cave System, Wildcat Saltpetre Cave. Pseudanophthalmus deceptivus Barr 1981 Type Locality: Fisher Cave, Lee County, Virginia Other Virginia Records: Known only from the type locality (Fig. 37). Pseudanophthalmus delicatus Valentine 1932 Type Locality: Gilley Cave, Lee County, Virginia Other Virginia Records (Fig. 38): Lee Co.: Baileys Cave, Bowling Cave, Burial Cave, Cattle Cave, Gallohan No. 1 Cave, Garretts Cave, Gibson No. 1 Cave, Jones Saltpetre Cave, Litton No. 1 Cave, McCurry Indian Cave, Molly Wagle Cave, Poor Farm Cave, Seal Pit, Smith Cave, Spangler Cave, Unthanks Cave. Remarks: This is one of the most widespread species of Pseudanophthalmus in the Commonwealth, yet it is confined to a single county. Pseudanophthalmus egberti Barr 1965 Type Locality: Starnes Cave, Giles County, Virginia Other Virginia Record (Fig. 37): Giles Co.: Giant Caverns. Pseudanophthalmus gracilis Valentine 1931 Type Locality: Tawneys Cave, Giles County, Virginia Other Virginia Records (Fig. 38): Craig Co.: Rufe Caldwell Cave; Giles Co.: Clover Hollow Cave, Smokehole Cave, Stay High Cave. Pseudanophthalmus hirsutus Valentine 1931 Type Locality: Cudjo’s Cavern (part of the Gap Cave System), Lee County, Virginia Other Virginia Record (Fig. 39): Lee Co.: Cumberland Gap Saltpetre Cave (part of the Gap Cave System). Remarks: Both of these caves are now part of the Cumberland Gap Cave System. Pseudanophthalmus hoffmani Barr 1965 Type Locality: Buchanan Saltpetre Cave, County, Virginia Other Virginia Records (Fig. 38): Bland Co.: Byrds Water Cave, Coon Cave, Hamilton Cave, Newberry- Bane Cave, Repass Saltpetre Cave; Giles Co.: Salamander Cave; Smyth Co.: Beaver Creek Cave, Smyth 34 BANISTERIA Winchester * Pygmarrhopalites pavo & Pygmarrhopalites sextus gw Pygmarmopalites silvus 150 kilometers Fig. 35. Distribution of Pygmarrhopalties pavo, P. sextus, and P. silvus in Virginia. Gray dots are all the terrestrial sampling sites. Winchester + Litocampa hoffmani A Litocampa cookei ga Litocampa fieldingae @ Litocampa pucketti Kilometers Fig. 36. Distribution of Litocampa cookei, L. fieldingae, L. hoffmani, and L. pucketti in Virginia. Gray dots are all the terrestrial sampling sites. by Winchester 78 Harrisonburg * Pseudanophthalmus cordicollis A Pseudanophthalmus egberti mg Pseudanophthalmus avernus @ Pseudanophthalmus deceptivus a Le, —————————aiKiometers Fig. 37. Distribution of Pseudanophthalmus avernus, P. cordicollis, P. deceptivus, and P. egberti in Virginia. Gray dots are all the terrestrial sampling sites. NO. 42, 2013 HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 35 Winchester + Pseudanophthalmus delicatus Harrisonburg A Pseudanophthalmus hoffmani g Pseudanophthalmus gracilis ———————————rilometers Fig. 38. Distribution of Pseudanophthalmus delicatus, P. gracilis, and P. hoffmani in Virginia. Gray dots are all the terrestrial sampling sites. Winchester # Pseudanophthalmus hirsutus A Pseudanophthalmus hortulanus g Pseudanophthalmus hubbardi @ Pseudanophthalmus hubrichti Kilometers Fig. 39. Distribution of Pseudanophthalmus hirsutus, P. hortulanus, P. hubbardi, and P. hubrichti in Virginia. Gray dots are all the terrestrial sampling sites. Winchester + Pseudanophthalmus limicola A Pseudanophthalmus intersectus gm Pseudanophthalmus holsingeri @ Pseudanophthalmus nelsoni Kilometers Fig. 40. Distribution of Pseudanophthalmus holsingeri, P. intersectus, P. limicola, and P. nelsoni in Virginia. Gray dots are all the terrestrial sampling sites. 36 BANISTERIA Marble Cave, Walkers Cave; Washington Co.: Quarry No. | Cave. Remarks: Range extensions into both Giles and Washington counties as well as the new Byrds Water Cave record in Bland County supplement records in Holsinger & Culver (1988) and Barr (2004). Pseudanophthalmus holsingeri Barr 1965 Type Locality: Young-Fugate Cave, Lee County, Virginia Other Virginia Records: Known only from the type locality (Fig. 40). Remarks: This species is listed as State Endangered. Pseudanophthalmus hortulanus Barr 1965 Type Locality: Cassell Farm No. 2 Cave, Tazewell County, Virginia Other Virginia Records: Known only from the type locality (Fig. 39). Pseudanophthalmus hubbardi (Barber 1928) Type Locality: Luray Caverns, Page County, Virginia Other Virginia Records: Known only from the type locality (Fig. 39). Remarks: Barr (2004) erroneously reported that this species had “not been retaken for several decades, perhaps because of commercialization of the cave.” In fact, Hubbard collected both male and _ female specimens, with and without bait, near Broaddus Lake in Luray Caverns in February 1996. Pseudanophthalmus hubrichti Valentine 1948 Type Locality: Daugherty Cave, Russell County, Virginia Other Virginia Records: Known only from the type locality (Fig. 39). Pseudanophthalmus intersectus Barr 1965 Type Locality: Crossroads Cave, Bath County, Virginia Other Virginia Record (Fig. 40): Bath Co.: Williams Cave. Pseudanophthalmus limicola Jeannel 1931 Type Locality: Maddens Cave, Shenandoah County, Virginia Other Virginia Records (Fig. 40): Rockingham Co.: Bakers Cave; Shenandoah Co.: Shenandoah Caverns, Shenandoah Wild Cave. Pseudanophthalmus longiceps Barr 1981 Type Locality: Fisher Cave, Lee County, Virginia Other Virginia Records: Known only from the type locality (Fig. 41). NO. 42, 2013 Pseudanophthalmus nelsoni Barr 1965 Type Locality: Arritt Mill Tunnel Cave, Alleghany County, Virginia Other Virginia Record (Fig. 40): Alleghany Co.: Blue Springs Cave. Pseudanophthalmus parvicollis Jeannel 1931 Type Locality: Battlefield Crystal Cave, Shenandoah County, Virginia Other Virginia Record (Fig. 41): Frederick Co.: Ogdens Cave. Remarks: Repeated attempts by Hubbard to find this species at its type locality, even with bait, met with failure in March 1992, June 1998, November 1998, and February 1999. This commercial cave is presently known as Crystal Caverns at Hupps Hill. Pseudanophthalmus petrunkevitchi Valentine 1945 Type Locality: Skyline Caverns, Warren County, Virginia Other Virginia Records (Fig. 41): Page Co.: Woods Cave; Warren Co.: Allens Cave. Pseudanophthalmus pontis Barr 1965 Type Locality: Buck Hill Cave, Rockbridge County, Virginia Other Virginia Records: Known only from the type locality (Fig. 41). Remarks: Repeated attempts to find this species at its type locality, even with bait, have met with failure beginning in November 1992. Environmental changes accompanying the commercialization of this cave (Natural Bridge Caverns) are thought to have adversely affected and possibly extirpated this species. Pseudanophthalmus potomaca Valentine 1932 Type Locality: Kenny Simmons Cave, Pendleton County, West Virginia Virginia Records (Fig. 42): Bath Co.: Little Mountain Cave, Starr Chapel Saltpetre Cave; Highland Co.: Vandevander Cave. Remarks: The range extension into Bath County supplements records in Holsinger & Culver (1988) and Barr (2004). Pseudanophthalmus praetermissus Barr 1981 Type Locality: Kerns No. 1 Cave, Scott County, Virginia Other Virginia Records: Known only from the type locality (Fig. 42). Pseudanophthalmus punctatus Valentine 1931 Type Locality: Tawneys Cave, Giles County, Virginia Other Virginia Records (Fig. 42): Giles Co.: Clover HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 37 Winchester * Pseudanophthalmus parvicollis a Pseudanophthalmus petrunkevitchi g Pseudanophthalmus longiceps @ Pseudanophthalmus pontis Big Stone Gap Oat 7 150 KKitometers Fig. 41. Distribution of Pseudanophthalmus longiceps, P. parvicollis, P. petrunkevitchi, and P. pontis in Virginia. Gray dots are all the terrestrial sampling sites. Winchester @ Pseudanophthalmus potomaca + Pseudanophthalmus pusio a Pseudanophthalmus punctatus g_ Pseudanophthalmus praetermissus 150 ——————KKilometers Fig. 42. Distribution of Pseudanophthalmus potomaca, P. praetermissus, P. punctatus, and P. pusio in Virginia. Gray dots are all the terrestrial sampling sites. Winchester # Pseudanophthalmus rotundatus A Pseudanophthalmus sanctipauli g_ Pseudanophthalmus sericus @ Pseudanophthalmus quadratus 150 —————————————KKitometers Fig. 43. Distribution of Pseudanophthalmus quadratus, P. rotundatus, P. sanctipauli, and P. sericus in Virginia. Gray dots are all the terrestrial sampling sites. 38 BANISTERIA Hollow Cave, Smokehole Cave, Spruce Run Mountain Cave, Stay High Cave. Remarks: The Stay High Cave record of this species supplements its reporting in Holsinger & Culver (1988) and Barr (2004). Pseudanophthalmus pusio (Horn 1868) Type Locality: Erhart Cave, Montgomery County, Virginia Other Virginia Records (Fig. 42): Montgomery Co.: Agnews Cave, Aunt Nellies Hole, Fred Bulls Cave, Mill Creek Cave, Old Mill Cave, Slussers Chapel Cave, Thorn Hill Cave; Roanoke Co.: Goodwins Cave, New Dixie Cave, Smith No. 1 Cave. Remarks: The type locality was destroyed by quarrying (Barr, 2004). Pseudanophthalmus quadratus Barr 1965 Type Locality: Straleys No. 1 Cave, Giles County, Virginia Other Virginia Record (Fig. 43): Giles Co.: Sugar Run Cave System. Remarks: The Sugar Run Cave System record of this species supplements its reporting in Holsinger & Culver (1988). Pseudanophthalmus rotundatus Valentine 1932 Type Locality: English Cave, Claiborne County, Tennessee Virginia Records (Fig. 43): Lee Co.: Elys Moonshine Cave, Smith Cave, Sweet Potato Cave. Pseudanophthalmus sanctipauli Barr 1981 Type Locality: Banners Corner Cave, Russell County, Virginia Other Virginia Record (Fig. 43): Scott Co.: Greears Sweet Potato Cave. Pseudanophthalmus seclusus Barr 1981 Type Locality: Flannery Cave, Scott County, Virginia Other Virginia Records (Fig. 44): Scott Co.: Alley Cave, Cox Ram Pump Cave, Hill Cave, Kerns No. 1 Cave, McDavids Cave, Pond Cave, Queens Cave. Pseudanophthalmus sericus Barr 1981 Type Locality: Lane Cave, Scott County, Virginia Other Virginia Records: Known only from the type locality (Fig. 43). Pseudanophthalmus thomasi Barr 1981 Type Locality: Blair-Collins Cave, Virginia Other Virginia Record (Fig. 45): Scott Co.: Coley No. 2 Cave. Scott County, NO. 42, 2013 Remarks: This species is listed as State Endangered. Pseudanophthalmus vicarius Barr 1965 Type Locality: Hugh Young Cave, Tazewell County, Virginia Other Virginia Records (Fig. 44): Tazewell Co.: Bowens Cave, Cauliflower Cave, Fallen Rock Cave, Gully Cave, Lost Mill No. 3 Cave, Martins Cave. Pseudanophthalmus virginicus (Barr 1960) Type Locality: Hugh Young Cave, Tazewell County, Virginia Other Virginia Records: Known only from the type locality (Fig. 45). Remarks: This species was originally the type species of the genus Aphanotrechus but is now assigned to Pseudanophthalmus. Family Pselaphidae Arianops jeanneli Park 1956 Type Locality: Gilley Cave, Lee County, Virginia Other Virginia Records: Known only from the type locality (Fig. 46). Order Diptera Family Sphaeroceridae Spelobia tenebrarum (Aldrich 1897) Type Locality: Wyandotte Cave, Crawford County, Indiana Virginia Records (Fig. 47): Lee Co.: Molly Wagle Cave, Sweet Potato Cave. Remarks: This fly, often associated with dung, is widespread in caves throughout the eastern United States. It has not been routinely collected, and is undoubtedly much more common and widespread in Virginia than the two records indicate. Class Malacostraca Order Amphipoda Family Crangonyctidae Bactrurus angulus Koenemann & Holsinger 2001 Type Locality: Saur Kraut Cave, Claiborne County, Tennessee Virginia Record (Fig. 48): Lee Co.: Cumberland Gap Saltpetre Cave (part of the Gap Cave System) Remarks: This large, rare species is also recorded from Kings Saltpetre Cave, Claiborne Co., Tennessee. HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 39 Winchester + Pseudanophthaimus seclusus a Pseudanophthalmus vicarius 150 Ce hKitometers Big Stone Gap om Eten Fig. 44. Distribution of Pseudanophthalmus seclusus and P. vicarius in Virginia. Gray dots are all the terrestrial sampling sites. *% Pseudanophthalmus thomasi A Pseudanophthalmus virginicus a hKKilometers Fig. 45. Distribution of Pseudanophthalmus thomasi and P. virginicus in Virginia. Gray dots are all the terrestrial sampling sites. * Arianops jeanneli Big Stone Ga ye Fig. 46. Distribution of Arianops jeanneli in Virginia. Gray dots are all the terrestrial sampling sites. 150 Kilometers 40 BANISTERIA * Spelobia tenebrarum Big Stone Ga gO : Fig. 47. Distribution of Spelobia tenebrarum in Virginia. Gray dots are all the terrestrial sampling sites. 150 CC TKilometers ™§ Bactrurnis angulus é& Gammarus minus tenuipes Big Stone Gap Ses SOE Fig. 48. Distribution of Bactrurus angulus and Gammarus minus var. tenuipes in Virginia. Gray dots are all the aquatic sampling sites. 150 _ Kilometers + Crangonyx antennatus kilometers Fig. 49. Distribution of Crangonyx antennatus in Virginia. Gray dots are all the aquatic sampling sites. NO. 42, 2013 HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 4] Crangonyx antennatus Packard 1881 Type Locality: Nickajack Cave, Tennessee Virginia Records (Fig. 49): Lee Co.: Baileys Cave, Barney Cave, Bowling Cave, Burial Cave, Cave Springs Cave, Cavin Cave, Cedar Hill Cave, Chances Cave, Combs No. 1 Cave, Cope Cave, Crouse Cave, Cudjo’s Cavern (part of the Gap Cave System), Cumberland Saltpetre Cave (part of the Gap Cave System), Flannery Bridge Springs, Frazier Cave, Gallohan No. 1 Cave, Gallohan No. 2 Cave, Garretts Cave, Gibson No. 1 Cave, Gibson-Frazier Cave, Gilliam Cave, Glen Olingers Cave, Golf Course No. 1 Cave, Golf Course No. 2 Cave, Gregorys Cave, Hamblin School Cave, Indian Burial Cave, Jones Saltpetre Cave, Knapper Cave, Lesters Cave, Litton No. 1 Cave, Long Cave, Lucy Beatty Cave, McClure Cave, McCurry Indian Cave, Minors Saltpetre Cave, Molly Wagle Cave, Mount Moriah Pit, Olinger Cave, Reasors Cave, Roadside No. 1 Cave, Seal Cave, Secret Cave, shallow spring in “Cedars”, small stream near Sims Spring, Sims Spring Cave, Slemp Cave, Smiths Milk Cave, Spangler Cave, Spout Spring, spring at entrance to Gallohan Cave No. 1, Sweet Potato Cave, Taylor Pit, Thompson Cave, Thompson Cedar Cave, Unthanks Cave, Watsons No. 1 Cave, Young-Fugate Cave; Scott Co.: Carters Pit Cave, Daniel Boone Caverns, Daniel Boones Spring, McDavids Cave, Pack Rat Cave, Speers Ferry Cave, Spurlock Cave; Washington Co.: Debusk Natural Caverns; Wise Co.: Wildcat Cavern Cave, Wildcat Saltpetre Cave. Remarks: This widespread amphipod is also recorded from caves from eastern Tennessee to northern Alabama. Marion County, A total of 21 Stygobromus species are found in the study area, 20 from caves, and 16 endemic to the state. Stygobromus abditus Holsinger 1978 Type Locality: James Cave, Pulaski County, Virginia Other Virginia Records (Fig. 50): Giles Co.: New River Cave; Montgomery Co.: Eve Einstein Cave; Pulaski Co.: Grindstead Cave, Sam Bells Cave, spring seep/trout pond at New River; Smyth Co.: Hillside Hideaway Cave; Washington Co.: Dolingers Cave, Ira Miller Cave, Mennick Cave, Red Wolf Cave; Wythe Co.: Bertha Cave, Catron Cave, Pierce Mill Cave, Watsons Cave. Remarks: This species is relatively widespread and common where found. Stygobromus baroodyi Holsinger 1978 Type Locality: Bathers Cave, Rockbridge County, Virginia Other Virginia Records (Fig. 50): Rockbridge Co.: Bell Cave, Billy Williams Cave, Hottingers Cave, Limekiln Cave, Buck Hill Cave, Grahams Cave, Showalters Cave, Turkey Hill Cave. Remarks: This species is relatively common and known to date only from Rockbridge County caves. Stygobromus biggersi Holsinger 1978 Type Locality: Ogdens Cave, Frederick County, Virginia Other Virginia Records (Fig. 50): Clarke Co.: well on farm near Boyce; Frederick Co.: Johns Cave, monitoring well and test well near Stephens City; Loudoun Co.: Rust Cave No. 1; Warren Co.: Brother Daves Cave. Remarks: Other localities, north and northeast of Virginia, include sites (caves and wells) in Jefferson and Berkeley counties, West Virginia, Washington Co., Maryland, and Cumberland and Franklin counties, Pennsylvania. Stygobromus conradi (Holsinger 1967) Type Locality: Breathing Cave, Virginia Other Virginia Records (Fig. 51): Bath Co.: Barberry Cave, Bobcat Cave, Butler Cave, Mountain Grove Cave, Serpent Cave, Starr Chapel Cave; Highland Co.: Aqua Cave, Bullpasture Mountainside Cave, Huppmans Saltpetre Cave, Huppman Jr. Cave, Marshalls Cave, Helictite Cave, Water Sinks Cave. Bath County, Stygobromus cumberlandus Holsinger 1978 Type Locality: Wildcat Saltpetre Cave, Wise County, Virginia Other Virginia Records (Fig. 51): Lee Co.: Baileys Cave, Cliff Cave, Indian Cave; Scott Co.: well near Duffield; Wise Co.: Kelly Cave, Parsons Cave, Rocky Hollow Cave. Remarks: This species is relatively widespread and recorded from caves and a water well in the 3-county area of Lee, Scott, and Wise counties in far southwestern Virginia. Stygobromus ephemerus (Holsinger 1969) Type Locality: Tawneys Cave, Giles County, Virginia Other Virginia Records (Fig. 51): Giles Co.: Canoe Cave, Stay High Cave. Remarks: As presently known, this species has a very limited distribution, and the majority of collections to date have been from Tawneys Cave. Stygobromus estesi Holsinger 1978 Type Locality: Rufe Caldwell Cave, Craig County, Virginia 42 BANISTERIA # Stygobromus abclitus A Stygobromus baroodyi BH Stygobromus biggersi Big Stone Gap 150 Kilometers Fig. 50. Distribution of Stygobromus abditus, S. baroodyi, and S. biggersi in Virginia. Gray dots are all the aquatic sampling sites. + Stygobromus ephemerus A Stygobromus estesi gw Stygobromus conradi @ Stygobromus cumberlandus BS o3 Big Stone Gap Es. 150 eee hKilometers Fig. 51. Distribution of Stygobromus conradi, S. cumberlandus, S. ephemerus, and S. estesi in Virginia. Gray dots are all the aquatic sampling sites. # Stygobromus gracilipes A Stygobromus finleyi @ Stygobromus fergusoni @ Stygobromus hoffmani CC TKilometers Fig. 52. Distribution of Stygobromus fergusoni, S. finleyi, S. gracilipes, and S. hoffmani in Virginia. Gray dots are all the aquatic sampling sites. NO. 42, 2013 HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 43 Other Virginia Records (Fig. 51): Alleghany Co.: Arritt Mill Tunnel Cave, Paxtons Cave; Craig Co.: Newcastle Murder Hole, Shires Saltpetre Cave; Giles Co.: spring on tributary of Johns Creek; Montgomery Co.: spring in Poverty Hollow and seeps at old mine drainage near McCoy. Remarks: This species is recorded from a variety of groundwater habitats, including caves, springs, seeps, and mine drainage. Stygobromus fergusoni Holsinger 1978 Type Locality: Slussers Chapel Cave, Montgomery County, Virginia Other Virginia Records (Fig. 52): Botetourt Co.: Karls Pit Cave; Montgomery Co.: Old Mill Cave. Remarks: This species is recorded primarily from seep pools in three caves. Stygobromus finleyi Holsinger 1978 Type Locality: English Cave, Tennessee Virginia Records (Fig. 52): Lee Co.: Cavin Cave, Indian Burial Cave, Jones Saltpetre Cave, Longs Cave. Remarks: This species is recorded from a few caves in the Powell Valley of southwestern Virginia and northeastern Tennessee where it is usually found in drip pools. Claiborne County, Stygobromus gracilipes (Holsinger 1967) Type Locality: Skyline Caverns, Warren County, Virginia Other Virginia Records (Fig. 52): Clarke Co.: well on John Staelin farm and well on Greenwood farm; Frederick Co.: Ogdens Cave; Page Co.: Woods Cave; Rockingham Co.: Church Mountain Cave, Deer Hole (cave), Devil’s Hole (cave), Endless Caverns, Massanutten Caverns, 3-D Maze Cave; Shenandoah Co.: Crystal Cave; Warren Co.: Allens Cave, Brother Daves Cave, Front Royal Caverns. Remarks: This species is also recorded from caves and wells north and northeast of Virginia in Berkeley and Jefferson counties, West Virginia; a spring in Washington Co., Maryland; and Needy Cave in Franklin Co., Pennsylvania. Stygobromus hoffmani Holsinger 1978 Type Locality: Lowmoor Cave, Alleghany County, Virginia Other Virginia Records (Fig. 52): Alleghany Co.: McElwee Cave; Craig Co.: small spring with culvert under road near Bald Mountain; Rockbridge Co.: seep in Goshen Wildlife Management Area. Remarks: This species has been collected from a variety of groundwater habitats in the three Virginia counties listed. Stygobromus hubbardi Holsinger 2009 Type Locality: Battlefield Crystal Cave (Crystal Caverns at Hupps Hill), Shenandoah County, Virginia Other Virginia Record (Fig. 53): Shenandoah Co.: Keyhole Cave. Remarks: This tiny, rare species (ca. 1.7-2.3 mm in length) has been collected to date from tiny drip pools in the two caves listed above. Stygobromus interitus Holsinger 1978 Type Locality: New Castle Murder Hole Cave, Craig County, Virginia Other Virginia Records: Known only from the type locality (Fig. 53). Remarks: A very rare species known only on the basis of two female specimens. Stygobromus leensis Holsinger 1978 Type Locality: Litton No. 1 Cave, Lee County, Virginia Other Virginia Records (Fig. 53): Lee Co.: Gallohan No. 2 Cave, Skull Cave. Remarks: To date, this species is recorded from drip/seep pools in three caves in central Lee County. Stygobromus mackini Hubricht 1943 Type Locality: Sikes Cave, Russell County, Virginia Other Virginia Records (Fig. 54): Bland Co.: Becky Knob Cave, Buddy Penleys Cave, Cedar Cliff Saltpetre Cave, Gollehorn Cave, Hamilton Cave, Newberry- Banes Cave, Repass Saltpetre Cave, Spring Hollow Cave; Giles Co.: Ballards Cave, Birthday Cave, Colby Cave, Doe Mountain Cave, Giant Caverns, New River Cave, Salamander Cave, spring at Bear Cliff near Mountain Lake, spring along small tributary to Johns Creek, Spruce Run Mountain Cave, Starnes Cave, Stay High Cave, Straleys Cave, Tawneys Cave; Lee Co.: Cave Road Cave, Fisher Cave; Russell Co.: Austines Hilltop Cave, Banners Corner Cave, Bundys No. 2 Cave, Burns Handline Cave, Bush Handline Cave, Concrete Tank Cave, Densmore Hill Cave, Dickenson Cave, Dorton Spring Cave, Grays Cave, Jessie Cave, Johnson Cave, Munsey Cave, Porgie Bundys Cave, Riverbend Cave, Rocky Fork Cave, Sikes Cave, Smiths Drop Cave, Trooper Said Cave; Scott Co.: Abrams Cave, Big Spiders in a Little Maze Cave, Blair-Collins Cave, Blowing Hole Cave, Bolling Cave, Cave Hollow Cave, Contact Cave, Deep Spring Cave, Fields Cave, Flannery Cave, Fugate Cave, Greears Sweet Potato Cave, Grigsby Cave, Hill Cave, Hillman Cemetery Pit Cave, Jack Cave, Jackson Cave, Jessee Branch Cave, Kerns Smoke-Hole Cave, Lane Cave, Little Duck Cave, 44 BANISTERIA McDavids Cave, McNew Cave, Moccasin Valley Cave, Natural Tunnel Cave, Pack Rat Cassel Cave, Pond Cave, Queens Cave, Rhoton Milk and Butter Cave, Speers Ferry Cave, Spurlock Cave, Taylor No. 1 Cave, Taylor No. 2 Cave, Winding Stair Cave, Wolfe Cave; Smyth Co.: Big Sink Cave, Buchanan Saltpetre Cave, Hancock Cave, Harmon Cave, Little Saltpetre Cave, McMullin Cave, Tilson Saltpetre Cave, Walkers Cave, Worleys Cave; Tazewell Co.: Cauliflower Cave, Chimney Rock Cave, Corkscrew Cave, Crocketts Cave, Fallen Rock Cave, Glenwood Church Cave, Higginbothams No. | Cave, Hugh Young Cave, Little River Cave, Lost Mill No. 1 Cave, Martins Cave, small spring on east slope of Beartown Mountain, Steeles Cave, Ward Cove Cave; Washington Co.: Ball Battery Cave, covered spring | mile NW of Court House, Singleton Cave, Substation Cave, Thunder Cave; Wise Co.: Big Kelly Cave, Blowing Cave, Omega Cave System, Parsons Cave, Wildcat Saltpetre Cave. Remarks: Stygobromus mackini 1s clearly the most common and widespread stygobiotic species in Virginia. It is also recorded from seven caves in neighboring West Virginia, including five in Mercer County and two in Monroe County (Fong et al., 2007), as well as nine caves in seven counties of upper East Tennessee (Holsinger, 1978; Holsinger & Culver, 1988; J.R. Holsinger, unpub. data). In addition, this species or a morphologically very closely similar species is recorded from a cave in Charlestown in Clark Co., Indiana, and at least five caves in eastern Kentucky. However, the populations in Indiana and Kentucky need further study before their taxonomic relationship with S. mackini s. str. can be determined. Stygobromus mausi Holsinger 2009 Type Locality: Round Hill Cave, Rockingham County, Virginia Other Virginia Records: Known only from the type locality (Fig. 53). Remarks: To date, this tiny, rare species is known only from nine specimens collected from a single drip pool in the type locality. Stygobromus morrisoni (Holsinger 1967) Type Locality: Witheros Cave, Bath County, Virginia Other Virginia Records (Fig. 54): Bath Co.: Crossroads Cave, Clarks Cave; Highland Co.: Corbett Cave, Secret Anthodite Cave. Remarks: The range of this species extends north- northeast into West Virginia where it has been collected from Dyers Cave in Hardy Co. and Kenny Simmons Cave in Pendleton Co. (Fong et al., 2007). NO. 42, 2013 Stygobromus mundus (Holsinger 1967) Type Locality: Witheros Cave, Bath County, Virginia Other Virginia Records (Fig. 55): Alleghany Co.: tributary to Cowpasture River; Bath Co.: Porters Cave, Russells Reserve Cave, Williams Cave. Remarks: This species is fairly common in small streams but is more often collected from drip pools. As noted above, it was also collected once from a surface stream, presumably flushed out of a nearby subterranean habitat. Stygobromus pseudospinosus Holsinger 1978 Type Locality: Luray Caverns, Page County, Virginia Other Virginia Records: Known only from the type locality (Fig. 55). Remarks: This species is known only from drip pools in Luray Caverns. Both S. pseudospinosus and S. spinosus are morphologically closely similar and may well be conspecific despite being considered separate species by Holsinger (1978). A resolution of this question awaits a DNA analysis. Stygobromus spinosus (Hubricht & Mackin 1940) Type Locality: spring near Hawksbill Mountain on Skyline Drive, Madison County, Virginia Other Virginia Records (Fig. 55): Albemarle Co.: spring in Saddle Hollow; spring 2.1 km SE of Sawmill Ridge Overlook in Shenandoah National Park; Clarke Co.: seep at Reservoir Hollow on Appalachian Trail; Fauquier Co.: seeps in Wildcat Hollow, G. Richard Thompson Wildlife Management Area; Greene Co.: spring at Swift Run Gap, Skyline Drive, Shenandoah National Park; spring near Pinefield Shelter, Shenandoah National Park; Nelson Co.: Old Afton Railroad Tunnel; Page Co.: 3.2 km S of Big Meadows, Skyline Drive; Rappahannock Co.: spring at Indian Run Shelter, Shenandoah National Park; Warren Co.: seep in woods near Blue Mountain; spring near Browntown Valley Overlook, Shenandoah National Park. Remarks: This is widely distributed in small springs and seeps, and is the only Stygobromus species in this study not to be found in caves. The habitat is under- collected relative to caves, and there are likely many more populations. Together with S. obrutus (see Culver et al., 2012a), it is outside the range of all other described Stygobromus. Stygobromus stegerorum Holsinger 1978 Type Locality: Madison Saltpetre Cave, County, Virginia Other Virginia Records (Fig. 55): Augusta Co.: Blue Hole Cave, Stegers Fissure. Augusta HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA # = Stygobromus hubbardi A Stygobromus interitus HM Stygobromus leensis @ Stygobromus mausi Kilometers Fig. 53. Distribution of Stygobromus hubbardi, S. interitus, S. leensis, and S. mausi in Virginia. Gray dots are all the aquatic sampling sites. # Stygobromus morrisoni A Stygobromus mackini CC TKilometers Fig. 54. Distribution of Stygobromus mackini and S. morrisoni in Virginia. Gray dots are all the aquatic sampling sites. Stygobromus mundus Stygobromus pseudospinosus Harrisonburg Stygobromus spinosus Stygobromus stegerorum i a ao, CC TKilometers Fig. 55. Distribution of Stygobromus mundus, S. pseudospinosus, S. spinosus, and S. stegerorum in Virginia. Gray dots are all the aquatic sampling sites. 45 46 BANISTERIA Remarks: To date, this species has been recorded from deep, phreatic lakes in the caves listed above. All of these lakes are also inhabited by the troglomorphic cirolanid isopod Antrolana lira Bowman. This species is listed as State Endangered. Family Gammaridae Gammarus minus var. tenuipes Shoemaker 1940 Type Locality: Organ Cave, Greenbrier County, West Virginia Virginia Records (Fig. 48): Tazewell Co.: Fallen Rock Cave, Gillespie Water Cave, Hugh Young Cave. Remarks: See Fong et al. (2007) for the details of a study of this morphologically similar variety in a karst basin of Greenbrier and Monroe counties in southern West Virginia. It is assumed that the variety G. minus var. tenuipes has evolved at least twice and independently under similar selection pressures, resulting in a morphologically similar form “tenuipes” in two completely separate karst basins: one in southern West Virginia and the other in Tazewell Co., Virginia. Order Isopoda Family Asellidae Caecidotea bowmani Lewis 1980 Type Locality: Natural Bridge Park, drain on trail, Rockbridge County, Virginia Other Virginia Record (Fig. 56): Botetourt Co.: Purgatory Cave. Remarks: The Purgatory Cave record is an addition to the Natural Bridge Park record noted by Lewis (2009). This isopod occupies both shallow subterranean habitats and caves. Caecidotea cumberlandensis Lewis 2000 Type Locality: Indian Cave, Lee County, Virginia Other Virginia Record (Fig. 56): Lee Co.: Cliff Cave. Remarks: Known only from two neighboring caves in extreme southwestern Virginia. Caecidotea henroti (Bresson 1955) Type Locality: Smokehole Cave, Giles County, Virginia Other Virginia Records (Fig. 56): Giles Co.: Tawneys Cave; Nelson Co.: Old Afton Railroad Tunnel; Pulaski Co.: James Cave. Remarks: Known from four sites, abandoned railroad tunnel. including an Caecidotea holsingeri (Steeves 1963) Type Locality: Organ Cave, Greenbrier County, West NO. 42, 2013 Virginia Virginia Records (Fig. 56): Alleghany Co.: Paxtons Cave; Bath Co.: Barberry Cave, Breathing Cave, Butler-Sinking Creek Cave, Cave Run Pit Cave, Clarks Cave; Giles Co.: Spring on small tributary of Johns Creek; Highland Co.: Aqua Cave, Helictite Cave, Sweet Anthodite Cave. Remarks: This is a widespread species found in caves and occasionally surface streams in karst areas in parts of Virginia, West Virginia, and Maryland (Fong et al., 2007). Caecidotea incurva (Steeves & Holsinger 1968) Type Locality: Berry Cave, Roane County, Tennessee Virginia Records (Fig. 57): Smyth Co.: McMullin Cave; Washington Co.: Dolingers Cave; Wythe Co.: Bowles Spring Cave, Deep Spring Cave, Early Cave, Gardners Cave, Groseclose Cave No. 1, Lone Ash No. 2 Cave, Mockleys Cave. Remarks: This species also occurs in Blount County, Tennessee. Caecidotea mausi Lewis 2009 Type Locality: Brother Daves Cave, Warren County, Virginia Other Virginia Records: Known in Virginia only from the type locality (Fig. 58). Remarks: This species is also known from several sites in Maryland. Caecidotea pricei Levi 1949 Type Locality: Refton Cave, Pennsylvania Virginia Records (Fig. 57): Augusta Co.: Barterbrook Spring Cave, Glade Cave; Bath Co.: Butler-Sinking Creek Cave; Frederick Co.: Ogdens Cave; Loudoun Co.: Rust Cave; Page Co.: Will Mauck Cave; Rockbridge Co.: Bathers Cave, Bell Cave, Billy Williams Cave, Hottinger Cave, Showalters Cave, Tolleys Cave; Rockingham Co.: Ann Millers Cave, Cedar Hill Cave, Endless Cave, Massanutten Caverns, Mauzy Spring, Stonewall’s Spring; Shenandoah Co.: Flemings Cave; Warren Co.: Skyline Caverns, Water Hole Cave. Remarks: This species is at the southernmost portion of its range in Virginia. It is also known from West Virginia, Maryland, and Pennsylvania. Lancaster County, Caecidotea recurvata (Steeves 1963) Type Locality: Unthanks Cave, Lee County, Virginia Other Virginia Records (Fig. 58): Lee Co.: Baileys Cave, Barney Cave, Bowling Cave, Burial Cave, Cattle Cave, Cave Springs Cave, Combs No. | Cave, Cope Cave, Crouse Cave, Earlys Cave, Fisher Cave, HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 47 Caecidotea bowmani Caecidotea henroti Caecidotea cumberlandensis Caecidotea holsingeri ee oe eee NKilometers Fig. 56. Distribution of Caecidotea bowmani, C. cumberlandensis, C. henroti, and C. holsingeri in Virginia. Gray dots are all the aquatic sampling sites. §- Caecidotea incurva A Caecidotea pricei eee Kilometers Fig. 57. Distribution of Caecidotea incurva and C. pricei in Virginia. Gray dots are all the aquatic sampling sites. + Caecidotea recurvata A Caecidotea mausi Kilometers Fig. 58. Distribution of Caecidotea mausi and C. recurvata in Virginia. Gray dots are all the aquatic sampling sites. 48 BANISTERIA Gallohan No. 1 Cave, Gallohan No. 2 Cave, Gibson No. 1 Cave, Gilley Cave, Gilliam Cave, Golf Course No. 1 Cave, Golf Course No. 2 Cave, Hamblin School Cave, Kinzer Hollow Cave, Knapper Cave, Litton No. | Cave, Long Cave, McClure Cave, McCurry Indian Cave, Minors Saltpetre Cave, Molly Wagle Cave, Olinger Cave, Reasors Cave, Roadside No. 1 Cave, Seal Cave, Secret Cave, Skull Cave, Smiths Milk Cave, Spangler Cave, Sweet Potato Cave, Tater Cave, Taylor Pit Cave, T-Bone Cave, Thompson Cedar Cave, Young-Fugate Cave; Russell Co.: Banners Corner Cave, Big Spring Cave, Breeding Cave, Bundys No. 2 Cave, Bundys Pearl Cave, Burns Cave, Concrete Tank Cave, Daugherty Cave, Dickerson Cave, Dorton Spring Cave, Grays Cave, Indian Cave, Jessie Cave, Johnson Cave, Munsey Cave, Phreatic To A Fault Cave, Porgie Bundys Cave, Seven Springs Cave, Smiths Cave, Stewart Monks Cave, Sykes Cave; Scott Co.: Blair- Collins Cave, Bob Dean Cave, Coley No. 2 Cave, Contact Cave, Flannery Cave, Jack Cave, Jessee Branch Cave, Lane Cave, McDavids Cave, Rhoton Milk and Butter Cave, Spurlock Cave, Sweet Potato Cave, Taylor No. 1 Cave, Taylor No. 2 Cave, Thomas Smith Cave, Triplet Wells Cave; Smyth Co.: McMullin Cave; Washington Co.: Brass Kettle Hole, Cave Spring Cave, David Debusk Cave, Debusk Mill No. 1 Cave, Fraleys Cave, Thunder Cave; Wise Co.: Bloomer Cave, Burtons Cave, Hairy Hole Cave, Kelly Cave, Little Kennedy Cave, Parsons Cave, Rocky Hollow Cave, Wildcat Saltpetre Cave. Remarks: Found in caves and karst springs in Virginia and Tennessee. This species can be found in drip pools and streams within caves. Caecidotea richardsonae Hay 1901 Type Locality: Nickajack Cave, Tennessee Virginia Records (Fig. 59): Lee Co.: Gregorys Cave, Olinger Cave, Smiths Milk Cave; Russell Co.: Cascade Pit Cave; Scott Co.: Blair-Collins Cave, Hortons Cave, Moccasin Valley Cave, Speers Ferry Cave, Wolfe Cave; Tazewell Co.: Bowens Cave, Fallen Rock Cave, Hugh Young Cave, Little River Cave, Lost Mill No. 3 Cave, Rosenbaums Water Cave, Stonley Cave; Washington Co.: Walker Mountain Saltpetre Cave. Remarks: This species co-occurs with C. recurvata at a few caves, and shares a similar range. Marion County, Caecidotea vandeli (Bresson 1955) Type Locality: Erhart Cave, Montgomery County, Virginia Other Virginia Records (Fig. 59): Alleghany Co.: Falling Spring Creek Cave; Bath Co.: Blowing Cave; Botetourt Co.: Brough Cave No. 2; Craig Co.: Rufe NO. 42, 2013 Caldwell Cave; Giles Co.: New River Cave, Spruce Run Mountain Cave, Stay High Cave; Montgomery Co.: Aunt Nellies Hole Cave, Old Mill Cave, Slussers Chapel Cave; Roanoke Co.: Goodwins Cave; Shenandoah Co.: Barb Cave; roadside seeps along Short Mountain. Remarks: The type locality was destroyed by quarrying. This species also occurs in Maryland, where it has been recorded from several springs and a mine (Lewis et al., 2011). Lirceus culveri Estes & Holsinger 1976 Type Locality: McDavids Cave, Virginia Other Virginia Records (Fig. 60): Scott Co.: Carters Pit Cave, Fugate Cave, Jackson Cave. Remarks: This species is apparently endemic to a single karst spring basin in the Rye Cove area of Scott County. It is typically found on and among small to medium-sized rocks in shallow portions of streams. Scott County, Lirceus usdagalun Holsinger & Bowman 1973 Type Locality: Gallohan No. 1 Cave, Lee County, Virginia Other Virginia Records (Fig. 60): Lee Co.: Flannery Bridge Springs, Gallohan No. 2 Cave, Golf Course No. 2 Cave, Masons Cave, Sims Creek Springs, Surgener Cave, Thompson Cedar Cave. Remarks: A Virginia endemic, this species is listed as Endangered by the U.S. Fish and Wildlife Service. It is restricted to a handful of shallow cave systems in the Cedars area of Lee County. Family Cirolanidae Antrolana lira Bowman 1964 Type Locality: Madison Saltpetre Cave, County, Virginia Other Virginia Records (Fig. 61): Augusta Co.: Blue Hole Cave, Stegers Fissure; Clarke Co.: well at Meade Church; Rockbridge Co.: Limekiln Cave; Rockingham Co.: 3-D Maze Cave, Devils Hole Cave, Linville Quarry No. 3 Cave, Massanutten Caverns; Warren Co.: Brother Daves Cave, Front Royal Caverns Cave, Power Plant Pit Cave. Remarks: For thirty years, this unique species was known only from a single isolated, groundwater aquifer (Holsinger & Culver, 1988). It is now known from deep groundwater aquifers in the Shenandoah Valley of Virginia and West Virginia (Holsinger et al., 1994; Fong et al., 2007), and these populations exhibit considerable genetic variation (Hutchins et al., 2010). Globally, stygobiotic cirolanids are found in areas with a past connection to marine waters. The presence of Augusta HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA * Caecidotea richardsonae A Caecidotea vandeli Big Stone Gap 150 Kilometers Fig. 59. Distribution of Caecidotea richardsonae and C. vandeli in Virginia. Gray dots are all the aquatic sampling sites. # = Lirceus culveri A Lirceus usdagalun Big Stone Ga nee Fig. 60. Distribution of Lirceus culveri and L. usdagalun in Virginia. Gray dots are all the aquatic sampling sites. 150 Kilometers * Antrolana lira kilometers Fig. 61. Distribution of Antrolana lira in Virginia. Gray dots are all the aquatic sampling sites. 49 50 BANISTERIA + Amerigoniscus henroti A Miktoniscus racovitzai racovitzai g Ligidium elrodii scottensis @ Ligidium elrodii leensis NO. 42, 2013 kilometers Fig. 62. Distribution of Amerigoniscus henroti, Ligidium elrodii leensis, L. e. scottensis, and Miktoniscus racovitzai racovitzai in Virginia. Gray dots are all the terrestrial sampling sites. A. lira in the Shenandoah Valley implies that such a past connection existed here. It is listed as Threatened by the U.S. Fish and Wildlife Service. Family Ligiidae Ligidium elrodii leensis Schultz 1970 Type Locality: Bowling Cave, Lee County, Virginia Other Virginia Records: Known only from the type locality (Fig. 62). Remarks: L. elrodii is a wide-ranging, surface-dwelling isopod in the eastern U.S. and Canada. This subspecies is slightly troglomorphic and lives on wet organic debris. Ligidium elrodii scottensis Schultz 1970 Type Locality: Coley Cave No. 2, Scott County, Virginia Other Virginia Records: Known only from the type locality (Fig. 62) Remarks: This subspecies is slightly troglomorphic and lives on wet organic debris. Family Trichoniscidae Amerigoniscus henroti Vandel 1950 Type Locality: Gilley Cave, Lee County, Virginia Other Virginia Records (Fig. 62): Lee Co.: Cope Cave, Earlys Cave, Gallohan No. 1 Cave, Kinzer Hollow Cave, Smiths Milk Cave, Spangler Cave, Sweet Potato Cave, Unthanks Cave. Miktoniscus racovitzai racovitzai Vandel 1965 Type Locality: Luray Caverns, Page County, Virginia Other Virginia Records (Fig. 62): Alleghany Co.: Island Ford Cave, Lowmoor Cave; Bath Co.: Russells Reserve Cave; Botetourt Co.: Karls Pit, Peerys Saltpetre Cave; Rockbridge Co.: Buck Hill Cave; Shenandoah Co.: Shenandoah Caverns; Warren Co.: Allens Cave, Brother Daves Cave. Remarks: This subspecies was also reported from Slacks Cave in Scott Co., Kentucky (Vandel, 1965). A second subspecies, M. r. oklahomensis, is found in one cave in Oklahoma. Given the great distances separating these populations, they may be separate species. DISCUSSION There are many ways, lengthy though it is, that this list of species is incomplete. The limitation of the list to only troglobionts, even when morphologically unmodified species limited to subterranean habitats are included, does not encompass all species with reproducing populations in caves. Graening et al. (2011) listed over 1,300 species found in caves in Arkansas and Oklahoma, yet less than ten percent of these are troglobionts and stygobionts. While many of the species listed by these authors are accidentals or temporary visitors, there are a number of ecologically important species in Arkansas and Oklahoma caves that are troglophiles and stygophiles. The same holds for Virginia caves. Holsinger & Culver (1988) discussed some of these, but there needs to be an up-to-date HOLSINGER ET AL.: INVERTEBRATE CAVE FAUNA OF VIRGINIA 51 treatment. There are also undescribed and/or undiscovered stygobionts and troglobionts in Virginia. The number of undiscovered species can be estimated if multiple samples of the same habitat are available (e.g., Schneider & Culver, 2004), but this technique has not been applied to the Virginia cave fauna. What is known is that there are a number of undescribed species that have been collected. This is particularly true for beetles in the genus Pseudanophthalmus (Barr, 2004) and diplurans in the genus Litocampa, where undescribed species have been known since Holsinger & Culver’s (1988) monograph. Some of the microfauna and some habitats (especially epikarst) have hardly been collected at all, but likely are relatively rich in species (see Pipan & Culver [2005] for an example from a West Virginia cave). The topics of biogeography and ecology of subterranean animals have not been covered in this report, but we hope that the data presented here will provide a stimulus for analytical studies in these fields. Little work has been done on phylogeography (e.g., Hutchins et al., 2010) and a quantitative analysis of species richness patterns (see Zagmajster et al. [2010] for some techniques) is also desirable. Finally, we note that much of the Virginia cave fauna may be at risk of extinction, both because of the restricted ranges of nearly all species (many are single cave endemics) and because relatively few caves, their surface area footprints, and their recharge areas are protected. The bright spots are both NGOs such as The Nature Conservancy, Virginia Speleological Survey, Cave Conservancy of the Virginias, Appalachian Cave Conservancy (formerly Perkins Cave Conservancy), Butler Cave Conservation Society, West Virginia Cave Conservancy (which owns caves in Virginia as well), and the Mid-Atlantic Karst Conservancy, and government agencies, especially the Virginia Department of Conservation and Recreation’s Natural Heritage Program and the Virginia Department of Game and Inland Fisheries. All have been active in protecting Virginia caves. The U.S. Fish and Wildlife Service continues to fund numerous projects in Virginia related to the study of listed troglobiotic and stygobiotic species, the implementation of best management practices on private lands to protect these species, and the acquisition of critical habitat for long-term protection. In addition, there is a Virginia Cave Board appointed by the governor which acts as an information source and provides guidance with respect to cave and karst protection. The Virginia Cave Protection Act of 1979 established the Cave Board, and also specifies criminal penalties for cave vandalism, sale of cave formations, and unpermitted scientific collections, while shielding landowners allowing access to caves. from liability when ACKNOWLEDGEMENTS We are grateful to many colleagues who have assisted us in recent years in various phases of the present study, including advice, new descriptions, species determinations, and useful data from their own collections. An older listing pertaining to a previous study can be found in Holsinger & Culver (1988). We apologize to anyone who might have been inadvertently omitted: Thomas C. Barr, Jr. (late), Thomas E. Bowman (late), Kenneth A. Christiansen, Bruno Condé (late), Lynn M. Ferguson, Daniel W. Fong, Robert Hershler, Roman Kenk (late), Julian J. Lewis, Jeremy A. Miller, William B. Muchmore, José Palacios- Vargas, William B. Shear, Marc Thibaud, Miloslav Zacharda, and Douglas Zeppelini. The Cave Conservancy of the Virginias provided a grant to support the preparation of this monograph. We are also indebted to Steven M. Roble for his editorial advice and suggestions which have greatly improved this work. Jerry Lewis provided a thorough and helpful review. Florita Gunasekara and Kristina Hsu assisted with the preparation of the maps. Mike Futtrell of the Virginia Speleological Survey provided numerous cave locations and was essential in resolving discrepancies in reporting cave locations. We also thank the Virginia Speleological Survey for permission to use cave location information in this study. None of this work would have been possible without the help of cavers and cave owners in Virginia. They are the unsung heroes of this study. LITERATURE CITED Aldrich, J. M. 1897. A collection of Diptera from Indiana caves. Annual Report of the Indiana Department of Geology and Natural Resources 21: 186- 190. Banks, N. 1895. Notes on the Pseudoscorpionida. 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Notes on the amphipod Gammarus minus Say and description of a new variety, Gammarus minus var. tenuipes. Journal of the Washington Academy of Science 30: 388-391. Sket, B. 2008. Can we agree on an _ ecological classification of subterranean animals? Journal of Natural History 42: 1549-1563. Steeves, H. R., III. 1963. Two new troglobitic asellids from West Virginia. American Midland Naturalist 70: 462-465. Steeves, H. R., III, & J. R. Holsinger. 1968. Biology of three new _ species of troglobitic asellids from Tennessee. American Midland Naturalist 80: 75- 83. Tellkampf, T. A. 1844. Beschreibung einiger neuer in der Mammuth-Hohle in Kentucky aufgefundener Gattungen von Gliederthieren. Mullers Archiv Anatomie und Physiologie 4: 308-312. Thibaud, J.M. 1995. Une nouvelle espece d’ Schaefferia dun grotte des Etats Unis d’Amerique (Collembola, Hypogastruridae). Revue Francaise d'Entomologie 17: 77-78. Thibaud, J. M. 1996. Une nouvelle espece dTyphlogastrura de deux grottes des Etats Unis d’Amerique (Collembola, Hypogastruridae). Revue Francaise d'Entomologie 18: 11-12. Valentine, J. M. 1931. New cavernicole Carabidae of the subfamily Trechinae Jeannel. Journal of the Elisha Mitchell Scientific Society 46: 247-258. Valentine, J. M. 1932. Horologion, a new genus of cave beetles. Annals of the Entomological Society of America 25: 1-9. Valentine, J. M. 1937. Anophthalmid beetles (fam. Carabidae) from Tennessee caves. Journal of the Elisha Mitchell Scientific Society 53: 93-100. NO. 42, 2013 Valentine, J. M. 1945. Speciation and raciation in Pseudanophthalmus (cavernicolous Carabidae). Trans- actions of the Connecticut Academy of Arts and Sciences 36: 631-672. Valentine, J. M. 1948. New anophthalmid beetles from the Appalachian region. Geological Survey of Alabama Museum Papers 27: 1-20. Vandel, A. 1950. Campagne spéléologique do Bolivar et R. Jeannel dans |l’Amérique du Nord (1928). Isopodes terrestres recuellis par C. Bolivar et R. Jeannel (1928) et le Dr. Henrot (1946). Archives de Zoologie Experimentale et Generale 87: 183-210. Vandel, A. 1965. Les Trichoniscidae cavernicoles (Isopoda terrestria; Crustacea) de |’ Amerique du Nord. Annales de Spéléologie 20: 347-389. Walker, B. 1925. New operculates. Nautilus 39: 5-8. species of fresh-water Wray, D. L. 1952. Some new North American Collembola. Bulletin of the Brooklyn Entomological Society 47: 95-106. Zacharda, M. 1980. Soil mites of the family Rhagidiidae (Actinedida: Euopdoidea). Acta Universitatis Carolinae — Biologica 1978: 489-795. Zacharda, M. 1985. New Rhagidiidae (Acarina: Prostigmata) from caves of the U.S.A. Véstnik Ceskoslovenské Spole¢nosti Zoologické 49: 67-80. Zacharda, M., D. Fong, H. H. Hobbs III, E. Piva, M. E. Slay, & S. J. Taylor. 2010. A review of the genus Traegaardhia (Acari, Prostigmata, Rhagidiidae) with descriptions of new species and a key to species. Zootaxa 2474: 1-64. Zagmajster, M., D. C. Culver, M. C. Christman, & B. Sket. 2010. Evaluating the sampling bias in pattern of subterranean species richness — combining approaches. Biodiversity and Conservation 19: 3035-3048. Zeppelini, D., & K. A. Christiansen. 2003. Arrhopalites (Collembola: Arrhopalitidae) in United States caves with the description of seven new species. Journal of Cave and Karst Studies 65: 36-42. MISCELLANEA 57 Miscellanea Reports 1. President’s Report The Executive Committee of the Virginia Natural History Society consists of the Executive Council (three Councilors) in addition to the President, Vice- President, Secretary-Treasurer, Banisteria Editor, Webmaster, and the last three Presidents. Please consider running for an open Councilor position with the VNHS. The term of appointment for this position is four years (2014-2017). We thank Lisa Williams for serving as Councilor for the past four years. If you are interested in running for this position please contact me at tfredericksen @ferrum.edu. This issue of Banisteria is devoted to cave biology, a topic of special importance to the natural history of Virginia. Please consider our journal as an outlet for research papers and field notes. Volumes are published in the spring and fall. The Master Naturalist program is thriving in Virginia. If our members are interacting with local chapters, please tell them about the Virginia Natural History Society. Respectfully submitted Todd Fredericksen, President Virginia Natural History Society 2. Secretary-Treasurer’s Report As of November 4, 2013, our treasury balance is $10,340. Membership stands at 113, of which 17 are institutions. Respectfully submitted, William A. Shear, Secretary/Treasurer 3. Webmaster’s Report VNHS website traffic from June to October 2013 is summarized in the following table: 365 302 840 347 791 Protat [= | azar [eae 34723 | 47008 | The Banisteria webpage (http://virginianaturalhisto. ipage.com/banisteria/banisteria.htm) contains the table of contents of all past issues of the journal as well as pdf files of selected articles. More articles will be added in the coming year. The following were the top five, most frequently viewed articles or pages for the months of June to October 2013. October 2013 1. Banisteria #32 — Potomac Gorge Bioblitz (Part 4). 2. Banisteria #35 — John Abbot, the English Naturalist- Artist, in Virginia - John V. Calhoun & David W. Johnston 3. Banisteria Main Page 4. Banisteria #21 — Common and _ Occasional Bryophytes of the Virginia Piedmont - David A. Breil 5. Banisteria #28 — List of the Sawflies (Hymenoptera: Symphyta) of Virginia - David R. Smith September 2013 1. Banisteria #5 -— Birds and Natural History Observations in the Life of William Byrd II of Westover - David W. Johnston 2. Banisteria #32 — Potomac Gorge Bioblitz (Part 5). 3. Banisteria #32 — Potomac Gorge Bioblitz (Part 3). 4. Banisteria #21 — Common and _ Occasional Bryophytes of the Virginia Piedmont - David A. Breil 5. Banisteria #28 — List of the Sawflies (Hymenoptera: Symphyta) of Virginia - David R. Smith August 2013 1. Banisteria #29 — Obituary: Roger H. de Rageot - Joseph C. Mitchell & David Liebman; My Favorite Story about Roger Rageot - Franklin J. Tobey, II 2. Banisteria #32 — Potomac Gorge Bioblitz (Part 5). 3. Banisteria #3 — Timber Rattlesnakes (Crotalus horridus) in Prince William Forest Park: Released Captives or Native Population? - Joseph C. Mitchell 58 BANISTERIA 4. Banisteria #21 — Common and _ Occasional Bryophytes of the Virginia Piedmont - David A. Breil 5. Banisteria Main Page July 2013 1. Banisteria #32 — Potomac Gorge Bioblitz (Part 2). 2. Banisteria #21 — Common and _ Occasional Bryophytes of the Virginia Piedmont - David A. Breil 3. Banisteria #32 — Potomac Gorge Bioblitz (Part 5). 4. Banisteria #32 — Potomac Gorge Bioblitz (Part 4). 5. Banisteria Main Page June 2013 1. Banisteria #21 — Common and _ Occasional Bryophytes of the Virginia Piedmont - David A. Breil 2. Banisteria #29 — Obituary: Roger H. de Rageot - Joseph C. Mitchell & David Liebman; My Favorite Story about Roger Rageot - Franklin J. Tobey, II 3. Banisteria #32 — Potomac Gorge Bioblitz (Part 5). 4. Banisteria #33 — The Gulf Fritillary (Agraulis vanillae): Breeding in Richmond, Virginia - Allen Belden, Jr. 5. Banisteria Main Page The url: http://virginianaturalhistorysociety.com is our current website address. Respectfully submitted, John White, VNHS Webmaster NO. 42, 2013 4. Editor’s Report This issue of Banisteria features a significant paper summarizing current knowledge of the obligate cave- inhabiting invertebrate fauna of Virginia, one of the richest in North America, as well as a historical review of past survey and research efforts on this fauna. I thank Lynn Ferguson for his assistance with the cover photographs and design. Manuscripts that are currently in review will comprise the first issue of 2014 and help to get the publication schedule back on track. I would like to thank the following people for serving as peer reviewers during the past two years (* = reviewed more than one manuscript): Steve Ahlstedt, Robert Barney, Yves Bousquet*, Jan Ciegler, John Cooper, Bob Davidson, Ralph Eckerlin, Art Evans*, Henri Goulet, Gary Graves*, Sergio Harding, Tom Henry*, Dick Highton, Eric Hilton, Richard Hoffman’, Paul Johnson, Kerry Katovitch, Leroy Koch, Bob Leberman, Jerry Lewis, Roger Mann, Joe Mitchell’, Bob Mulvihill, Brent Opell, John Orr, John Pagels, Stewart Peck*, Bob Rose, Nathan Schiff, Don Schwab*, Bill Shear*, Rowland Shelley*, Charlie Staines*, Warren Steiner*, Mark Swingle, Brian Taber, Keith Thomson, Steve Tilley, Linda Weir, Al Wheeler*, Jim White, Tom Wieboldt*, Bill Williams, and Mike Wilson. Their efforts were important in maintaining the quality of Banisteria. I apologize if any names were inadvertently omitted from this list. Respectfully submitted, Steve Roble, Editor, Banisteria Announcements 1. Recently published Daniel C. Dourson. 2013. Land Snails of the Great Smoky Mountains National Park and Southern Appalachians. Goatslug Publications, Bakersville, NC. 343 pp. Virginia Natural History Society http://virginianaturalhistorysociety.com/ General Information The Virginia Natural History Society (VNHS) was formed in 1992 to bring together persons interested in the natural history of the Commonwealth of Virginia. The VNHS defines natural history in a broad sense, from the study of plants, animals, and other organisms to the geology and ecology of the state, to the natural history of the native people who inhabit it. The goals of the VNHS are to promote research on the natural history of Virginia, educate the citizens of the Commonwealth on natural history topics, and to encourage the conservation of natural resources. Dissemination of natural history information occurs through publication of the journal Banisteria, named for John Banister (1650-1692) who was the first university- trained naturalist to work in Virginia. The first issue was published in 1992, and the journal is published twice per year in spring and fall. Articles cover a wide array of subjects, and prospective authors are encouraged to submit manuscripts on any aspect of natural history in Virginia; papers may pertain to Virginia or regional archaeology, anthropology, botany, ecology, zoology, paleontology, geology, geography, or climatology. Book reviews, biographies, obituaries, and historical accounts of relevance to natural history in Virginia also are welcomed. Manuscripts are peer- reviewed for suitability and edited for inclusion in the journal. Page charges ($20/page) are waived if the sole or first author is a VNHS member. All authors must pay $75/page if they desire color printing of figures. The society’s website contains detailed instructions for authors and the titles, abstracts or full PDF versions of articles from past Banisteria issues. Memberships The VNHS is open to anyone with an interest in natural history and welcomes participation by all members in society activities and efforts to promote education and conservation. Membership includes a subscription to Banisteria and invitations to periodic symposia and field events. Annual dues for members are $20 (per calendar year); library subscriptions are $40 per year. Checks or money orders (credit cards are not accepted) should be sent to the Secretary/Treasurer, who also has back issues of Banisteria available for sale. The VNHS is a tax-exempt, nonprofit, society under Section 501(C)3 of the IRS. We welcome donations to support our mission in Virginia. Virginia Natural History Society Application for Membership ANNUAL DUES AND SUBSCRIPTIONS TO BANISTERIA (memberships and subscriptions are by calendar year; subscribers/members outside the United States should add $3.00 for additional postage) L] $500.00 Life (not annual) $300.00 Benefactor $100.00 Patron $50.00 Supporting $40.00 Institutional $25.00 Family $20.00 Regular $5.00 Student (see below) L L L L L L L L I have added a contribution of $___ to my membership dues. The special student rate is applicable only when accompanied by the following certification signed by a faculty advisor. Institution Advisor Make checks or money orders payable to: Virginia Natural History Society Send membership form and dues to: Dr. William Shear, Secretary-Treasurer Virginia Natural History Society Box 96 Hampden-Sydney, VA 23943