The Invertebrate Cave Fauna of Virginia 
and a Part of Eastern Tennessee: 
Zoogeography and Ecology 
John R. Holsinger 
Department of Biological Sciences, 
Old Dominion University, Norfolk, Virginia 23529 
AND 
David C. Culver 1 
Department of Ecology and Evolutionary Biology, 
Northwestern University, Evanston, Illinois 60201 
ABSTRACT . — Collections of macroscopic invertebrate animals and 
ecological data were made from approximately 500 caves in the 
Appalachian Valley and Ridge province of Virginia and eastern Ten- 
nessee from 1961 to 1980. The study area comprised 26 counties in 
western Virginia and all or parts of seven counties in northeastern 
Tennessee. Approximately 335 species of invertebrates were recorded 
from the caves, including 140 troglobites (obligatory cavernicoles) and 
61 troglophiles (facultative cavernicoles). The troglobites are numeri- 
cally distributed as follows: flatworms (5), oligochaetes (3), snails (3), 
amphipods (20), isopods (15), pseudoscorpions (15), mites (2), spiders 
(8), centipedes (1), millipeds (9), collembolans (4), diplurans (6), and 
beetles (49). Basic ecological characteristics of cave species are consi- 
dered, including habitats, trophic relationships, life histories, and spe- 
cies interactions. 
Seven regional cave faunas, which coincide with major drainage 
basins, are recognized: (1) Shenandoah, (2) James, (3) Roanoke, (4) 
New, (5) Holston, (6) Clinch, and (7) Powell. Drainage basins that 
contain extensive exposures of cavernous limestone, such as the Clinch 
and Powell, have a greater diversity of cave-limited species than those 
with limited exposures of limestone. There is a strong linear relation- 
ship between cave species density and cave density, and an “area 
effect” exists among the endemic terrestrial troglobites. Aquatic tro- 
globites are apparently derived both indirectly through ancestral line- 
ages living in subterranean water prior to the present generation of 
caves and directly from surface ancestors. Terrestrial troglobites are 
apparently derived from preadapted surface ancestors living on cool, 
moist forest floors in the Appalachian Mountains. Invasion and colo- 
nization of caves by terrestrial organisms might have occurred in 
response to changing climates during the Pleistocene. Many troglobites 
are highly localized endemics that are restricted to only one or a few 
caves, whereas others have much broader ranges. 
1 Present address: Department of Biology, American University, 4400 
Massachusetts Avenue, N.W., Washington, D.C. 20016. 
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