2 
John R. Holsinger and David C. Culver 
Areas in the Appalachians of western Virginia and eastern Tennessee 
underlain by carbonate rocks contain numerous caves inhabited by 
an interesting diversity of cavernicolous organisms. The caves in these 
areas have been investigated extensively for many years, resulting in the 
accumulation of a significant body of information on many important 
aspects of biology, geology, and hydrology. Biologists have long been 
interested in the cave faunas of the Appalachian region. Probably one 
of the first biologists to visit caves in Virginia was E. D. Cope, who 
collected beetles and millipeds from caves in Giles and Montgomery 
counties (see Horn 1868, Cope 1869). A. S. Packard (1881, 1888) visited 
four caves in Virginia in 1874 and 1880 and collected specimens from 
Grand Caverns (then called Weyers Cave) and Fountain Cave (mistak- 
enly called Madisons Cave by Packard) in Augusta County, Endless 
Caverns (then called New Market Cave) in Rockingham County, and 
Luray Caverns in Page County (see also Emerton 1875, Ryder 1881). 
After a long lull, caves in the study area were visited in the 1920s 
and early 1930s by American biologists H. S. Barber (1928) and J. M. 
Valentine (1931, 1932) and by European biologists C. Bolivar and Rene 
Jeannel (see Berland 1931, Bolivar and Jeannel 1931, Chopard 1931, 
Jeannel 1931). In the late 1930s and early 1940s, specimens were col- 
lected from caves in the area by Kenneth Dearolf (1953; see also Loomis 
1939), Leslie Hubricht (1943), J. P. E. Morrison (1949), and others (see 
Fowler 1942, 1943, 1944, 1946). In 1946, the French biologist Henri 
Henrot visited 11 caves in Virginia and four in northeastern Tennessee. 
He collected many specimens, some of which were subsequently described 
as new species (see Henrot 1949, Jeannel 1949, Vandel 1950, Bresson 
1955). Bruno Conde, another French biologist, visited several caves in 
Virginia in 1956 (see Chappuis 1957). In 1958, T. C. Barr, Jr., visited 24 
caves in Tennessee and 37 in Virginia and made many important biolog- 
ical collections (Barr 1959). The present study was initiated in 1961, 
when J. R. Holsinger began a detailed survey of the Virginia-West Vir- 
ginia cave fauna (see Holsinger 1962, 1963a, 1963b, 1964). The survey 
was joined later by D. C. Culver, and in the early 1970s it was extended 
to include parts of eastern Tennessee. During our field work, which 
extended through 1980, collections of biological specimens and ecologi- 
cal observations were made in approximately 500 caves in Virginia and 
northeastern Tennessee. 
In the present paper, we have prepared an annotated listing of all 
invertebrate species from caves in the study area (defined below), using 
the data collected during our field work and supplemented by informa- 
tion from the literature. We have also discussed observations on the 
ecology of cavernicolous species and presented a detailed zoogeographi- 
cal analysis of the cave-limited fauna of the study area. Prevailing hypo- 
theses on the ecology and zoogeography of invertebrate cave faunas are 
