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April 2003 JOURNAL OF Volume 65 Number 1 ISSN 1090-6924 A Publication of the National CAVE AND KARST Speleological Society STUDIES Journal of Cave and Karst Studies Editor Louise D. Hose of the National Speleological Society National Cave and Karst Research Institute Volume 65 Number 1 April 2003 1400 University Drive Carlsbad, NM 88220 505-234-5561 or 887-2759 Voice CONTENTS 505-887-3051 FAX [email protected] The Cave-inhabiting rove beetles of the United States Production Editor (Coleoptera; Staphylinidae; excluding Aleocharinae and Pselaphinae): James A. Pisarowicz Diversity and distributions Wind Cave National Park Stewart B. Peck and Margaret K. Thayer 3 Hot Springs, SD 57747 605-673-5582 A conservation focused inventory of subterranean invertebrates [email protected] of the southwest Illinois Karst BOARD OF EDITORS Julian J. Lewis, Philip Moss, Diane Tecic, and Matthew E. Nelson 9 Anthropology Patty Jo Watson Mycological survey of Río Camuy Caves Park, Puerto Rico Department of Anthropology Ángel M. Nieves-Rivera 22 Washington University St. Louis, MO 63130 [email protected] Morphometric and spatial distribution prarmeters of karstic depressions, Lower Suwannee River Basin, Florida Conservation Can Denizman 29 Julian J. Lewis J. Lewis & Associates, Biological Consulting 217 West Carter Avenue Arrhopalites (Collembola: Arrhopalitidae) in U.S. caves Clarksville, IN 47129 with the description of seven new species 812-283-6120 Douglas Zeppelini and Kenneth Christiansen 36 [email protected] Earth Sciences-Journal Index Gypsum wedging and cavern breakdown: Studies in the Ira D. Sasowsky Mammoth Cave System Kentucky Department of Geology William B. White and Elizabeth L. White 43 University of Akron Akron, OH 44325-4101 330-972-5389 The Kuka'iau Cave, Mauna Kea, Hawaii, Created by water erosion: [email protected] A new Hawaiian cave type Stephan Kempe and Marlin Spike Werner 53 Exploration Andrea Futrell 987 Dow Creek Road Raw sewarge and sold waste dumps in lave tube caves of Hawaii Island Pembroke, VA 24136 William R. Halliday 68 540-626-5349 [email protected] Pa'auhau Civil Defense Cave on Mauna Kea, Hawaii: Life Sciences A lava tube modified by water erosion Steve Taylor Stephan Kempe, Ingo Bauer, and Horst-Volker Henschel 76 Center for Biodiversity Illinois Natural History Survey 607 East Peabody Drive (MC-652) Champaign, IL 61820-6970 Cave Science News 86 217-333-5702 [email protected] Editorial The National Cave and Karst Research Institute: Social Sciences American speleology’s remarkable opportunity Marion O. Smith P.O. Box 8276 Penelope Boston 87 University of Tennessee Station Knoxville, TN 37996 National Cave and Karst Research Act of 1998 88 Book Reviews Ernst H. Kastning P.O. Box 1048 Radford, VA 24141-0048 The Journal of Cave and Karst Studies (ISSN 1090-6924, CPM Number #40065056) is a multi-disciplinary, refereed [email protected] journal published three times a year by the National Speleological Society, 2813 Cave Avenue, Huntsville, Alabama 35810-4431 USA; (256) 852-1300; FAX (256) 851-9241, e-mail: [email protected]; World Wide Web: Proofreader http://www.caves.org/pub/journal/. The annual subscription fee, worldwide, by surface mail, is $18 US. Airmail deliv- Donald G. Davis ery outside the United States of both the NSS News and the Journal of Cave and Karst Studies is available for an addi- tional fee of $40 (total $58); The Journal of Cave and Karst Studies is not available alone by airmail. Back issues and cumulative indices are available from the NSS office. POSTMASTER: send address changes to the Journal of Cave JOURNAL ADVISORY BOARD and Karst Studies, 2813 Cave Avenue, Huntsville, Alabama 35810-4431 USA. David Ashley Rane Curl Malcolm Field Andrew Flurkey Copyright © 2003 by the National Speleological Society, Inc. Printed on recycled paper by American Web, 4040 Dahlia Street, Denver, Colorado 80216 USA John Ganter Donald MacFarlane Diana Northup Art Palmer Front cover: Main entrance of Clara Cave, Río Camuy Cave Park. See Nieves-Rivera, p. 22. William White Stewart B. Peck and Margaret K. Thayer - The Cave-inhabiting rove beetles of the United States (Coleoptera; Staphylinidae; excluding Aleocharinae and Pselaphinae): Diversity and distributions. Journal of Cave and Karst Studies, v. 65, n. 1, p. 3-8. THE CAVE-INHABITING ROVE BEETLES OF THE UNITED STATES (COLEOPTERA; STAPHYLINIDAE; EXCLUDING ALEOCHARINAE AND PSELAPHINAE): DIVERSITY AND DISTRIBUTIONS STEWART B. PECK Department of Biology, Carleton University, Ottawa, ON, K1S 5B6 CANADA [email protected] MARGARET K. THAYER Department of Zoology, Field Museum of Natural History, Chicago IL 60605-2496 USA A taxonomic listing is given for new records of 66 species of staphylinid beetles (excluding Aleocharinae and Pselaphinae) collected in caves in the contiguous United States. Most species are judged to be either accidentals or infrequent troglophilic inhabitants of caves. Nine species are classed as frequent troglophiles. When added to the 6 frequent troglophile species of aleocharine staphylinids, this yields a total of 15 species of staphylinid beetles (excluding Pselaphinae) frequently found in US cave ecosys- tems. No troglobitic species are known from US caves. Troglobitic staphylinids (excluding Pselaphinae) elsewhere in the world are few (some 30 species). They are briefly considered and discussed. Worldwide, troglobitic staphylinids are taxonomically, geographically, and geologically concentrated in the Canary Islands (in volcanic lava tube caves) and in nearby Spain and northwestern Africa. Worldwide, caves are most frequently and abundantly (Poggi et al. 1998). Some 53 species of them in 8 genera are occupied by beetles in the families Carabidae, Leiodidae, and considered to be troglobites in the USA (Peck 1998; Chandler Staphylinidae. The first two of these families contain a great & Reddell 2001). We are not aware of any cave records for many cave-specialized and cave-restricted (troglobitic) genera scaphidiines, which are strictly mycophagous as far as known and species in many parts of the world, mostly in Europe (Leschen & Löbl 1995), and they seem unlikely to be cave- (Casale et al. 1998; Giachino et al. 1998). In contrast, very few associated. cave-specialized or cave-restricted (troglobitic) staphylinid Staphylinid beetles occur frequently and commonly in beetles are known. Jeannel and Jarrige (1949) summarized caves in the United States. The published records of staphylin- data on about 150 species of Staphylinidae from over 1000 ids from US caves are many and scattered in numerous region- caves worldwide and considered very few of these species to al and state cave faunal reports. These have been summarized be troglobites. Recent reviews by Bordoni and Oromi (1998) by Roth (2001). After many years of faunal surveys of many and Outerelo et al. (1998) list only 30 troglobitic species caves by many people, no troglobitic species (outside worldwide, and most of these are from Europe and North Pselaphinae) have been found in the USA. It now seems most Africa. None are from the USA. likely that none will be found in the future. Most staphylinid beetles are distinctive in their appearance, Identification of species in this very large beetle family is with an exceptionally elongate and flexible body form and difficult and time consuming, even for the specialist, and since very short elytra (front wing covers) over their hind wings troglobitic species are not known from the US, staphylinid (Figs. 1–6). Their English common name, “rove beetle,” beetles have often received little or no specific attention in sur- comes from their behavior of rapidly running about in many veys of cave insect diversity. For these reasons, we have a very directions. Some species in the family are eyeless and without scattered and incomplete knowledge of which species of flight wings, but other than the few cave species, these are staphylinid beetles are frequent inhabitants of US caves and mostly small to minute litter- or soil-inhabiting species. This components of cave ecosystems. beetle family is one of the world’s largest, with some 4100 Klimaszewski and Peck (1986) presented a comprehensive species recorded in the USA and Canada, and over 47,000 systematic report on the cave-inhabiting aleocharine staphylin- species known to science worldwide (Newton et al. 2000). ids of the USA. The aim of this paper is to complement that Recently, the traditional beetle families Scaphidiidae and report and the literature summary of Roth (2001). We present Pselaphidae have been combined with the family a systematic summary of new records of authoritatively iden- Staphylinidae, and each is now treated as a subfamily (Kasule tified species of Staphylinidae (excluding Aleocharinae and 1966; Newton & Thayer 1995). These two groups are not Pselaphinae) from cave localities throughout the USA. This included in the following discussions, which are restricted to documentation of the species diversity and distribution of the subfamilies traditionally placed in the Staphylinidae. these staphylinid beetles helps to complete an understanding of Pselaphines are known from many caves, especially in the this previously poorly known part of the fauna of US caves. USA (Chandler 1992; Chandler & Reddell 2001) and Europe Copyright © 2003 by The National Speleological Society Journal of Cave and Karst Studies, April 2003 • 3 CAVE-INHABITING ROVE BEETLES OF THE UNITED STATES Figure 1. Brathinus nitidus. Figure 2. Geodromicus brunneus. Figure 3. Lesteva pal-