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number 14 june1988

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Eloise F. Potter, Acting Editor
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James W. Hardin
Department of Botany
N.C. State University

William M. Palmer
Curator of Lower Vertebrates
N. C. State Museum

David S. Lee
Curator of Birds
N.C. State Museum

Rowland M. Shelley
Curator of Invertebrates
N.C. State Museum

Brimleyana, the Journal of the North Carolina State Museum of Natural
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Subject matter will focus on taxonomy and systematics, ecology, zoo-
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paleontology. Papers will stress the results of original empirical field studies, but
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North Carolina State Museum of Natural Sciences
North Carolina Department of Agriculture
James A. Graham, Commissioner

CODN BRIMD 7
ISSN 0193-4406

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. Culver1

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.

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

Invertebrate Cave Fauna

critically examined and tested against our data in an attempt to gain a
better understanding of the factors that have influenced the present dis-
tribution of cave organisms in the study area. Because there have been
very few detailed studies on the ecology or zoogeography of an entire
regional cave fauna, this study should provide some interesting new
insights in these areas.

The results of the present study will also complement previously
published data on cave faunas of other areas in the eastern United
States and will considerably update the data in earlier papers on the
cave faunas of Tennessee by Barr (1961) and Virginia by Holsinger
(1963a, 1964). The cave faunas of North America have been sampled
extensively in recent years and, as a result, are becoming well docu-
mented. In the eastern United States, regional cave faunas have been
analyzed to one extent or another in papers by Krekeler and Williams
(1966) on Indiana, Barr (1967a) on Kentucky, Peck (1970) on Florida,
Franz and Slifer (1971) on Maryland, Holsinger and Peck (1971) on
Georgia, Holsinger (1976) on Pennsylvania, Holsinger et al. (1976) on
West Virginia, Peck and Lewis (1978) on Illinois, and Hobbs (1981) and
Hobbs and Flynn (1981) on Ohio.

THE STUDY AREA

As shown in Figure 1, the study area encompasses the 26 counties
in western Virginia that contain cavernous limestones, and a part of
eastern Tennessee. The Tennessee portion includes all of the lower
Powell Valley (parts of Campbell, Claiborne, Hancock, and Union
counties); that part of the Clinch Valley extending from the state line
southwest to the confluence of the Clinch and Powell rivers (most of
Hancock and Union counties and parts of Campbell, Grainger, and
Hawkins counties); and the northern periphery of Hawkins and Sullivan
counties, which lies just south of the state line in the Holston Valley.
The study area covers parts of seven major drainage basins (Fig. 2),
detailed discussions of which are included later under “Zoogeography.”

Geographic and Geologic Relationships

Excluding the limestone region of Florida, the major cave and
karst areas of the eastern United States, east of Mississippi River, are
developed in Paleozoic limestones of the Appalachian Valley and Ridge
(or simply “Appalachian Valley”), Appalachian Plateau (or Alleghany
Plateau of some authors), and Interior Low Plateaus physiographic
provinces (see Fig. 3). Although the Appalachian Valley and Ridge and
Appalachian Plateau provinces are usually assigned to a major physio-
graphic division called the Appalachian Highlands, and the Interior
Low Plateaus province is assigned to the Interior Plains division (see

John R. Holsinger and David C. Culver

Hunt 1967), the Interior Low Plateaus is sometimes referred to as
“Appalachian” in the broad sense, because it is closely allied biologically
and geologically with parts of the Appalachian Highlands. References in
this paper to the “greater Appalachian region” include the Interior Low
Plateaus.

For all intents and purposes, the study area as defined above lies
within the Appalachian Valley and Ridge province. Only its western
periphery in southwestern Virginia and east Tennessee encroaches on
the eastern margin of the Appalachian Plateau, where cavernous lime-
stones crop out along the flanks of Cumberland and Stone mountains.
The Appalachian Valley is underlain by folded and faulted bedrock that
varies in geological age from Lower Cambrian to Upper Mississippian,
about half of which is limestone and dolomite. The total number of
caves recorded from the study area through 1980 was 2611, including
2377 in Virginia and 234 in east Tennessee.

Limestones in the Appalachian Valley and Ridge province are
exposed on valley floors and along the sides of low ridges and are gen-
erally restricted to long, linear strike belts. As a result, the principal
orientation of most cave passages is along the regional strike (NE-SW),
trending parallel to the valleys and ridges in which the caves occur.
Strike-oriented belts of cavernous limestone are generally relatively nar-
row and separated from each other by intervening parallel exposures of
non-carbonate, clastic rocks such as sandstones, shales, and quartzites.
Karst topography is generally common on most limestone terranes but
is usually much more prominent in valleys floored by broad exposures
of Middle Cambrian and Middle Ordovician limestones (Holsinger
1975) (see Fig. 4). The overall drainage pattern is trellised, and, with the
exception of a segment of the New River that flows north, most major
streams flow roughly parallel to the strike (see Fig. 2).

South and southwest of the study area, the cave region of the
Appalachian Valley and Ridge extends through eastern Tennessee into
northwestern Georgia and northeastern Alabama. North and west of the
study area, it extends through eastern West Virginia, west-central
Maryland and across south-central Pennsylvania. The Appalachian
Plateau is capped with resistant, non-carbonate elastics of Pennsylvanian
age, but in several places, especially on its eastern and western sides,
cavernous limestones are exposed. On the eastern side significant
exposures of limestones occur in eastern New York (Helderberg Plateau),
southwestern Pennsylvania, western Maryland, parts of eastern West
Virginia, southeastern Kentucky (Pine Mountain), east-central Tennessee
(Grassy Cove, Lookout, and Sequatchie valleys), northwestern Georgia
(Lookout Valley), and northeastern Alabama (Lookout, Sequatchie,
and Wills valleys). Barr (1981a) appropriately termed some of these
disjunct limestone areas “karst islands” to call attention to their

Invertebrate Cave Fauna

Fig. 1. Outline map of the study area in western Virginia and northeastern
Tennessee, showing counties and county seats.

Fig. 2. Map of the study area showing major drainage basins as follows: 1,
Shenandoah; 2, James; 3, Roanoke; 4, New; 5, Holston; 6, Clinch; 7, Powell.
Arrows indicate direction of drainage. Principal mountains and ridges indicated
by hachuring.

John R. Holsinger and David C. Culver

Fig. 3. Part of the eastern United States showing major physiographic
provinces as follows: 1, Coastal Plain; 2, Piedmont; 3, Blue Ridge; 4,
Appalachian Valley and Ridge (= Appalachian Valley); 5, Appalachian Plateau;
6, Interior Low Plateaus.

geographic isolation. Although these karst areas are situated within the
Appalachian Plateau proper, they are geographically close and
geologically similar to belts of cavernous limestone on the western side
of the Appalachian Valley.

In Alabama, Kentucky, and Tennessee, where the Appalachian
Plateau is known locally as the Cumberland Plateau, cavernous
limestones of Mississippian age are exposed along the highly dissected
western margin of the Plateau in all three states. Cavernous areas on the
western side of the plateau, unlike those on the eastern side, are closely

Invertebrate Cave Fauna

allied with those of the adjoining Eastern Highland Rim of the Interior
Low Plateaus.

A few caves are also recorded from the Blue Ridge province south
and southeast of the study area in the higher mountains of southeastern
Tennessee (Barr 1961) and in western North Carolina (Cato Holler, Jr.,
pers. comm.). With the exception of those in Blount County, Tenn.,
however, most of these caves consist of fissure passages in non-carbonate
rocks.

Faunistic Relationships

For the most part, the cave-limited faunas of the Appalachian
Valley and karst islands on the eastern side of the Appalachian Plateau
differ significantly from those of the Interior Low Plateaus and western
margin of the Cumberland Plateau. Since many genera with troglobites
(see definition under “Methods”) are shared by these two major cave
regions, this difference is considerably greater on the species level.
However, differences in the cave faunas of the two regions are less
pronounced in southern Tennessee, northeastern Alabama, and
northwestern Georgia, where only short distances separate cave areas in
the Appalachian Valley, Cumberland Plateau, and Eastern Highland
Rim. Farther north in Kentucky and Virginia- West Virginia, where the
distance between the cave areas of the Interior Low Plateaus-Cumberland
Plateau and Appalachian Valley-eastern Appalachian Plateau is much
greater (see Fig. 3), the faunas are more different.

The ranges of a number of troglobitic species in the study area
extend into adjoining cave areas on the north, south, and west. However,
to the north the number of troglobitic species decreases significantly in
Maryland and Pennsylvania, where the cave-limited fauna is very sparse
(see Franz and Slifer 1971, Holsinger 1976). Even farther north, in the
glaciated cave area of New York, the only known troglobite is the
amphipod crustacean Stygobromus allegheniensis (Holsinger 1967a,
1978).

West of the study area in the adjoining cave areas of eastern West
Virginia, the cave-limited fauna is comparable in diversity to that in
western Virginia, and there is a strong taxonomic affinity among many
troglobitic species throughout much of the two-state area (cf., Holsinger
et al. 1976). There is also a strong affinity between certain cave-limited
species in the study area and those of Pine Mountain, a karst island in
the Appalachian Plateau about 16 km northwest of the study area in
southeastern Kentucky and northwestern Campbell County, Tenn.

Much of the cave-limited fauna in the Appalachian Valley of
eastern Tennessee south and southwest of the study area has not been
documented in the same detail as that of areas north and west of the
study area. But those observations and literature records that are

John R. Holsinger and David C. Culver

Invertebrate Cave Fauna

available (cited under “Review of the Fauna” elsewhere this paper)
suggest a close taxonomic affinity between troglobites of this area and
those of the southern part of the study area. For example, troglobitic
species from the Clinch drainage basin south of the confluence of the
Clinch and Powell rivers are closely related to, or in some cases the
same as, species from that part of the Clinch basin included in the study
area. In contrast to the Clinch basin, the cave-limited fauna of the
adjoining Holston drainage basin in northeastern Tennessee is generally
much less diverse, apparently reflecting the same relationship we have
noted elsewhere in this paper (see “Zoogeography”) between the cave
faunas of these two basins farther north in the study area.

In comparison with northeastern Tennessee (principally the Clinch,
Holston, and Powell basins), the Appalachian Valley of southeastern
Tennessee between Kingston in Roane County and the Georgia-
Tennessee state line near Chattanooga contains fewer and generally
smaller caves (see Barr 1961, Matthews 1971), and the cave-limited
fauna is poorly known. The cave-limited fauna of the Appalachian
Valley and eastern margin of the Appalachian Plateau south of
Chattanooga in northwestern Georgia has been documented in detail,
however (see Holsinger and Peck 1971). This fauna is diverse and shares
affinities with that of both the Appalachian Valley farther north and the
Cumberland Plateau and Eastern Highland Rim in adjacent northern
Alabama and south-central Tennessee.

Cave Vertebrates

The present study is limited to invertebrates, but some pertinent
observations are included on the ecology of the plethodontid salamander
Gyrinophilus porphyriticus (Green), a species that our research has
shown to be a major predator in certain cave-stream communities in
southwestern Virginia and eastern Tennessee (see “Ecology” elsewhere
this paper). Although there are no bona fide troglobitic vertebrates
recorded from the study area, certain populations of G. porphyriticus in
caves of the Clinch and Powell valleys are apparently cave-limited and
dominated by large, pale larvae. The systematics of these populations
warrants further detailed study. Elsewhere in the Appalachian Valley,
just west and south of the study area, several populations of Gyrinophilus
are considered troglobitic and include G. subterraneus Besharse and

Fig. 4. Karst features in the study area: A, entrance to Hugh Young Cave in
large sinkhole, Tazewell County; B, sinkhole topography on Middle Cambrian
limestone in the Clinch Valley, Scott County; C, Maiden Spring, a large
resurgence in the Ward Cove karst, Tazewell County; D, vertical entrance to
Stegers Fissure, Augusta County (courtesy of K. E. Wark and D. G. Whall); E,
sinks of Meadow Creek near Looney, Craig County.

John R. Holsinger and David C. Culver

Holsinger (1977) from General Davis Cave in Greenbrier County,
W.Va., and G. palleucus McCrady {sensu lato ) from several caves and a
temporary spring in Knox, McMinn, and Roane counties, Tenn. (see
Brandon 1965, Simmons 1975).

Troglobitic fishes of the family Amblyopsidae occur on the western
margin of the Cumberland Plateau and in the Interior Low Plateaus
and Ozark Plateaus, but are absent from the Appalachian Valley and
eastern side of the Appalachian Plateau except for the documented
occurrence of Typhlichthys subterraneus Girard in Lookout and Wills
valleys in northeastern Alabama and northwestern Georgia (see Cooper
and lies 1971).

METHODS

Field Work

During the course of our investigation (1961-1980), 450 caves in
Virginia and 53 in eastern Tennessee were explored for biological
specimens. In addition, biological data were obtained from the literature
or from other biologists on approximately 38 caves in Virginia and 9 in
Tennessee not visited by us during the field work. As of 1980, these
totals represented approximately 21% of the recorded caves in the study
area. Caves were visited in all counties in the study area except Clarke
County in northwestern Virginia, which has only four insignificant
caves reported (see Douglas 1964, Holsinger 1975). Virtually all caves
considered “large” (see Holsinger 1975) were checked at least once, and
some of the most complex ones, especially in the Clinch and Powell
valleys, were visited on several separate occasions. In addition to the
field work in Virginia and east Tennessee, biological data were collected
concurrently from 152 caves in adjacent West Virginia, but the results of
this part of the study have been published separately (see Holsinger et
al. 1976).

In most of the caves investigated, sampling for specimens was done
in all potential habitats, including banks of damp clay and silt, decom-
posing organic detritus (e.g., wood, leaves, guano), damp flowstone and
dripstone, pools fed by drips and seeps, and streams (see Fig. 5). With
the exception of specimens obtained from pit-fall traps used for a short
time in four Lee County caves during the summer of 1975, in a special
study by T. C. Kane, and the occasional use of cheese and shrimp baits
on an experimental basis, trapping, baiting, Berlese, and phreatic pump-
ing techniques were not employed in collecting. A majority of records
in this report are based on collections made directly from the substrate,
aided only by small brushes or syringes.

Collecting efforts were focused principally on troglobites and
troglophiles, but selective collections were also made of trogloxenes and
accidentals in order to document their occurrence. Non-cave habitats,

Invertebrate Cave Fauna

Fig. 5. Cave habitats in the study are: A, stream in Gallohan Cave No. 1, Lee
County (courtesy of D. E. Wapinski); B, rimstone pools in Sweet Potato Cave,
Lee County; C, decomposing wood in English Cave, Claiborne County; D, East
Lake in Madisons Saltpetre Cave, Augusta County; E, mud-bottom drip pool in
Molly Wagle Cave, Lee County.

John R. Holsinger and David C. Culver

such as springs and seeps, were sampled occasionally in order to obtain
specimens for comparison with those taken from nearby caves.

The species covered in this study are essentially macroscopic forms
(i.e., generally larger than 1 mm). Microscopic forms (< 1 mm) that are
sometimes reported from cave waters (e.g., protozoans, rotifers; see
Gittleson and Hoover 1970) or from the interstices of sand and gravel
substrates beneath cave streams (e.g., tiny oligochaeates, copepods, ostra-
cods) have not been included. Some preliminary studies, however, on
polluted pools in Banners Corner Cave in Russell County (see Holsinger
1966) and the interstitial habitat beneath a stream in Buis Saltpetre
Cave in Claiborne County (unpubl. data) indicate a potential richness of
subterranean microscopic organisms that would be profitable to investi-
gate in a future study.

Although our study does not cover all of the Clinch Valley in Ten-
nessee (Fig. 2), some pertinent data on the distribution of species
recorded from caves just southwest of the study area in Anderson
County are included. These data add significant details to the picture of
the geographic distribution of species or species groups whose ranges
extend into parts of the Clinch basin outside the study area. Moreover,
except for a few major caves in Hawkins and Sullivan counties, which
lie just south of the Tennessee-Virginia border, our survey does not
cover the Holston Valley in eastern Tennessee (see Fig. 1, 2).

Definition of Terms

Cavernicoles are usually classified ecologically according to their
level of adaptation and degree of restriction to the cave environment.
The commonly accepted system, which is used throughout this paper, is
defined as follows (see also Barr 1963, 1968). (1) Troglobites are obliga-
tory species, which are restricted to caves or similar habitats. Morpho-
logical modifications (specializations) called troglomorphisms usually
distinguish troglobites and may include, among other things, loss or
rudimentation of eyes and pigment, and attenuation of the body,
appendages, or sensory hairs. (2) Troglophiles are facultative species,
which are able to complete their life cycle within a cave but may also
occur in ecologically suitable habitats outside caves. (3) Trogloxenes are
species habitually found in caves or similar cool, dark habitats outside
caves, but they must return periodically to the surface or at least to the
entrance zone of a cave for food. Some species, however, such as certain
cave crickets, may be trogloxenic under one set of circumstances and
troglophilic under another (see Barr 1963). (4) Accidentals are species
that wander, fall, or are washed into caves and generally exist there
temporarily.

Many small aquatic invertebrates (e.g., flatworms, crustaceans)
simultaneously inhabit both caves and subterranean groundwater habitats

Invertebrate Cave Fauna

outside of caves and even outside of karst areas and are sometimes
called phreatobites (see Holsinger 1967a, Barr 1968) or stygobionts.
Because some of these species occur in shallow groundwater (i.e.,
vadose water) above the zone of permanent saturation (i.e., phreatic
water), the less restrictive designation stygobiont , now commonly used
by European workers, is probably preferable to phreatobite. Some
examples of non-cave habitats occupied by stygobionts include springs,
wells, the interstitial media of small gravels either beneath a stream (=
hyporheic or nappes fluviales) or beside a stream (= parafluvial or
nappes phreatiques), small seeps or their outflow above the water table
(= hypotelminorheic or nappes perchees), and outlets of drain tiles
placed beneath cultivated fields with poor natural drainage (for further
details see Henry 1978, Holsinger 1978, Culver 1982).

Edaphobites are obligatory deep-soil species that occasionally occur
in caves. Although frequently blind and weakly pigmented, edaphobites
are usually distinguished from true troglobites by the absence of other
troglomorphisms. This distinction is often a subtle one, however, and is
best made between species in a carefully studied group. Endogean is
used in a rather broad sense to designate species living in deep ground-
litter or soil (i.e., endogean species) or the habitat type itself (i.e.,
endogean habitat).

Cave Nomenclature

Locations and descriptions of most of the caves cited herein
have been published by Barr (1961) and Matthews (1971) for Tennessee
and by Douglas (1964) and Holsinger (1975) for Virginia, or are on file
with the Tennessee Cave Survey or the Virginia Speleological Survey.
Most cave names are now standardized for both states, but a few are
listed in the biological literature under different names and tend to be
confusing. In the following list the currently accepted, standardized
name is followed by the former name in parentheses: Banners Corner
Cave (Big Spring Cave), Russell County; Battlefield Crystal Cave
(Crystal Cave), Shenandoah County; Caney Sinks Cave (Sinks Cave),
Hancock County; Cudjos Cavern (King Solomons Cave in part), Lee
County; Endless Caverns (New Market or Zirkles Cave), Rockingham
County; Fred Bulls Cave (Mark Smiths Cave), Montgomery County;
Giant Caverns (Hopkins Cave), Giles County; Gilley Cave (Elys or
Shalers Cave), Lee County [refers to Ely Cave on p. 294 in Douglas
(1964), not Ely Cave on p. 306]; Grand Caverns (Weyers Cave),
Augusta County; and Wills Cave (Fraleys Cave), Washington County.

It should also be noted that we have retained the original name for
Buck Hill Cave (Rockbridge County), which was recently commercialized
under the name Caverns of Natural Bridge. Cassell Farm Cave No. 1
and 2 in Tazewell County are two separate caves located very close

John R. Holsinger and David C. Culver

together (see Holsinger 1975), but they are frequently not differentiated
as two caves in the older literature. In citing records from the literature
where it was not clear which cave was intended, we have listed the
locality as Cassell Farm Cave(s).

The following caves have been listed in the biological literature but
are unknown to either state’s cave survey by the name given (see also
indication in “Review of the Fauna”): Big Stony Cave, Giles County;
Cave No. 1 and No. 3, Pennington Gap, Lee County; Coopers (or
Parkeys) Cave, Hancock County; Field Cave, Russell County (apparently
not the Fields Cave in Holsinger 1975:240); Hammers Cave, Campbell
County (possibly same as Big Hollow Cave); Mushroom Cave, Page
County (possibly same as Ruffners Cave No. 1); Newman Ridge Cave,
Hancock County (could be any one of several caves in Newman Ridge
near Sneedville, Tenn.); Old Hollins Road Cave, Roanoke County: Old
Joe’s Cave near Wingina, Buckingham County (not in study area): Sikes
Cave, Russell County (apparently not the same as Sykes Cave in
Holsinger 1975:259); and Water Cave (presumably in the Shenandoah
Valley).

REVIEW OF THE FAUNA

Approximately 335 species of invertebrate animals, representing
some 90 families and 173 genera, have been recorded from caves in the
study area. An exact number is meaningless, of course, because many
species are incompletely known taxonomically and some groups have
been collected more intensively than others. Of the known species, 42%
are troglobites (some questionable pending further study); 18% are
troglophiles; 14% are trogloxenes; and the remaining 26% are marginal
trogloxenes and accidentals. The numerical distribution of troglobitic
and troglophilic species by taxonomic order (or subclass for arachnids)
is given in Table 1. Of the 140 troglobitic species, 42 are aquatic and 98
are terrestrial.

In the following list the higher taxa (phyla, classes, orders) are
arranged in generally accepted phylogenetic sequence. The lower taxa
(families, genera, species) are listed alphabetically within their respective
taxonomic groups. Species are arranged in species groups (under genera)
where usage of these groups is well established in the recent literature.
Some of the troglobites listed are only provisionally recognized or just
now in the process of being described (i.e., description in manuscript or
in press) and are therefore designated by upper case letters (viz., sp. A,
sp. B, etc.) under their respective genera or species groups. The
abbreviations TB, TP, TX, and AC designate troglobite, troglophile,
trogloxene, and accidental, respectively. However, as noted in the lists,
the ecological status of some species is questionable or provisional, and

Invertebrate Cave Fauna

Table 1. Frequency distribution by order or subclass of troglobites and
troglophiles in the study area.

Order or Subclass 1

No. of Troglobites 2

No. of Troglophiles 2

Alloeocoela (flatworms)

Tricladida (flatworms)

Lumbriculida (oligochaetes)

Mesogastropoda (snails)

Stylommatophora (snails)

Amphipoda (amphipods)

Isopoda (isopods)

Decapoda (crayfishes)
Pseudoscorpiones

(pseudoscorpions)

Acari (mites)

Opiliones (harvestmen)

Araneae (spiders)

Lithobiomorpha (centipedes)

Spirostreptida (millipeds)

Chordeumatida (millipeds)

Julida (millipeds)

Collembola (springtails)

Diplura (bristletails)

Orthoptera (crickets)

Coleoptera (beetles)

Diptera (flies)

Total number of species

140

1 The arachnid groups Pseudoscorpiones, Acari, Opiliones, and Araneae are
considered subclasses by some workers (see Krantz 1970) and orders by
others (see Barnes 1980).

Includes several species whoes ecological status is presently unclear (see text).

clarification must await additional information on ecology, systematics,
or both.

All known cave records within the study area are listed alpha-
betically by county for each species. Type localities for troglobites are
indicated in parentheses following the cave name when these localities
occur in the study area. Quotation marks and a reference (in parentheses)
to the author who used the name indicate cave localities taken from the
literature and unknown to either the Virginia Speleological Survey or the
Tennessee Cave Survey by the name published. Many of the troglobites
listed are endemic to the study area and, unless indicated otherwise in
the preliminary discussions or under “Comments,” the caves listed

John R. Holsinger and David C. Culver

include all known locality records. A question mark after a cave name
indicates a questionable species record. Additional data on the geo-
graphic distribution or taxonomy of a species are sometimes given
under “Comments,” following the list of cave records.

PHYLUM PLATYHELMINTHES

Among the free-living flatworms (class Turbellaria) found in Virginia
and east Tennessee caves are alloeocoels and tricladids. The former are
restricted to a single, curious species also recorded from single caves in
Kentucky and West Virginia; it is the only alloeocoel reported from
caves (Carpenter 1970a, Holsinger et al. 1976). The other flatworms are
planarians in the genera Sphalloplana and Phagocata.

Cavernicolous flatworms are generally encountered in drip or
stream-fed pools or on the flat surface of rocks in small streams;
population numbers fluctuate greatly. Outside the study area, Sphal-
loplana chandleri is recorded from springs in Davidson County, Tenn.,
and Floyd County, Ind. (Kenk 1977), and is apparently a relatively
widespread stygobiont. In contrast, the troglobites Sphalloplana con-
similis (Fig. 13E) and S. virginiana have narrowly delimited ranges (Fig.
6) and are known only from the caves listed below (see Hyman 1945,
Kenk 1977). The presence of Sphalloplana percoeca in northeastern
Tennessee is highly questionable. In redescribing this species, Kenk
(1977) listed many localities in Alabama, Kentucky, and Tennessee and
indicated that the range might possibly extend into West Virginia and
Georgia. He also pointed out that some of these records, especially
those from Tennessee, need verification.

Phagocata gracilis is recorded from numerous localities (viz., caves,
springs, headwaters of small streams) in the eastern and east-central
United States (Kenk 1970). Although Phagocata subterranea (Hyman
1937) was previously reported from Banners Corner Cave by Holsinger
(1963a, 1964, 1966), it is apparently a subterranean ecophenotype of P.
gracilis and is therefore now considered a synonym of this species by
Kenk (1970). Phagocata morgani , common in the subterranean ground-
waters of the Ward Cove karst in Tazewell County, is recorded from
many springs, small streams, and caves in eastern North America
(Carpenter 1970b).

Order Alloeocoela

Family Prorhynchidae

Geocentrophora cavernicola Carpenter (TB?)

Virginia. — Tazewell Co.: Fallen Rock Cave.
Geocentrophora sp.

Virginia. — Lee Co.: Cliff Cave.

Invertebrate Cave Fauna

a Sphalloplana chandleri
• S. consimilis
■X S. virginiana
□ £. percoeca?

▼ S. spp.

WEST VIRGINIA

VIRGINIA

KENTUCKY /l

0 25 50 km

0 25 50 miles

NORTH CAROLINA

Fig. 6. Distribution of troglobitic planarians ( Sphalloplana ) in the study area.

Order Tricladida

Family Kenkiidae

Sphalloplana ( Speophila ) chandleri Kenk (TB?)

Virginia. — Tazewell Co.: Fallen Rock Cave.

Sphalloplana (Sphalloplana) consimilis Kenk (TB)

Tennessee. — Claiborne Co.: Buis Saltpetre Cave.

Virginia. — Lee Co.: Bowling, Cope, Gallohan No. 1 (type locality),
Gregorys and McClure caves.

Sphalloplana (Sphalloplana) percoeca (?) (Packard) (TB)

Tennessee. — Campbell Co.: Meredith Cave.

Sphalloplana (Speophila) virginiana Hyman (TB)

Virginia. — Rockbridge Co.: Showalters Cave (type locality).
Comments. Previous records from Bland and Lee counties (Holsinger
1963b, 1964) are invalid in light of subsequent revisionary studies
by Kenk (1977).

Sphalloplana spp.

Tennessee. — Claiborne Co.: Chadwells Cave. Union Co.: Oaks
Cave.

John R. Holsinger and David C. Culver

Virginia. — Bland Co.: Newberry-Bane Cave. Frederick Co.: Ogdens
Cave. Lee Co.: Cliff and Smiths Milk caves. Russell Co.: Banners
Corner Cave. Wise Co.: Rocky Hollow Cave.

Comments. — These records are based on juveniles or poorly
preserved specimens of which specific determinations could not
be made.

Family Planariidae

Phagocata gracilis (Haldeman) (TP or TX)

Virginia. — Russell Co.: Banners Corner Cave.

Phagocata morgani (Stevens and Boring) (TP or TX)

Virginia. — Giles Co.: Starnes Cave. Tazewell Co.: Fallen Rock and
Hugh Young caves.

PHYLUM ANNELIDA

All segmented worms recorded from caves in Virginia and east
Tennessee are in the class Oligochaeta and belong to the orders
Branchiobdellida, Haplotaxida, Lumbriculida, and Tubificida. The
records given in the list below are based on either literature references
(e.g., Gates 1959) or selective collecting and by no means represent an
exhaustive survey.

The branchiobdellids occur as epizoites on freshwater crustaceans,
and all species recorded from caves in the study area were taken on the
troglophilic crayfish Cambarus bartonii s. lat. (see Holt 1973). The
occurrence of these species in Appalachian caves is probably largely
accidental, inasmuch as they are generally widespread in epigean habitats
and are transported into caves secondarily by their crayfish hosts.

The haplotaxids include several species of terrestrial and semi-
terrestrial “earthworms” that are probably initially introduced into
caves in mud or silt washed underground by flooding or filtration.
However, many of these species probably exist in caves as trogloxenes,
or even as troglophiles, under certain conditions. All of the haplotaxids
listed below are also reported from caves elsewhere in the eastern
United States (see Gates 1959, Franz and Slifer 1971, Cook 1975,
Holsinger et al. 1976, Peck and Lewis 1978).

Of greater interest zoogeographically are the “thread-like”
lumbriculid worms, of which all three species found to date are
apparently troglobites with narrowly defined ranges. These worms have
been collected from the gravel substrate of small streams, but only after
diligent searching. In comparison with Europe, the North American
cavernicolous lumbriculid fauna is very poorly known (Cook 1975).

Although they have been observed in several study-area caves,
tubificid worms remain poorly known to date. An undetermined genus
and species of the family Enchytraeidae has been collected from the
stream in Fallen Rock Cave in Tazewell County, and Tubifex tubifex

Invertebrate Cave Fauna

Muller (Tubificidae) has been observed in Banners Corner Cave, Russell
County, in pools polluted by sewage (see Holsinger 1966).

Order Branchiobdellida
Family Branchiobdellidae
Ankyrodrilus legacus Holt (AC)

Tennessee. — Hancock Co.: Fairmont School Cave.

Virginia. — Tazewell Co.: Fallen Rock Cave.

Bdellodrilus illuminatus (Moore) (AC)

Tennessee. — Hancock Co.: Cantwell Valley Cave.

Cambarincola fallax Hoffman (TX or AC)

Tennessee. — Hancock Co.: Cantwell Valley and Fairmont School
caves.

Virginia. — Scott Co.: McDavids Cave. Tazewell Co.: Fallen Rock
Cave.

Cambarincola philadephicus (Leidy) (TX or AC)

Tennessee. — Hancock Co.: Fairmont School Cave.

Virginia. — Tazewell Co.: Wagoners Cave.

Cambarincola sp.

Tennessee.— Sullivan Co.: Bristol Caverns.

Oedipodrilus macbaini (Holt) (AC)

Tennessee. — Sullivan Co.: Bristol Caverns.

Xironodrilus formosus Ellis (AC)

Tennessee.— Sullivan Co.: Bristol Caverns.

Xironogiton instabilis (Moore) (AC)

Virginia. — Tazewell Co.: Wagoners Cave.

Order Haplotaxida

Family Lumbricidae

Allolobophora chlorotica (Savigny) (TX)

Virginia. — Rockbridge Co.: Showalters Cave.

Allolobophora turgida Eisen (TX)

Virginia. — Rockbridge Co.: Showalters and Tolleys caves.

Bimastos tumidus (Eisen) (TX)

Tennessee. — Claiborne Co.: English Cave.

Virginia. — Lee Co.: Gilley Cave. Russell Co.: “Field Cave” (Gates,
1959:80).

Dendrobaena rubida (Savigny) (TX)

Tennessee. — Claiborne Co.: English Cave.

Virginia. — Lee Co.: Cudjos Cavern. Russell Co.: Jessie Cave.
Eisenia rosea (Savigny) (TX)

Virginia. — Giles Co.: Clover Hollow and Tawneys caves.

Eiseniella tetraedra (Savigny) (TX)

Virginia. — Russell Co.: Banners Corner Cave. Tazewell Co.: Fallen
Rock Cave.

John R. Holsinger and David C. Culver

Octolasium lactewn (Oerley) (TX)

Virginia. — Bland Co.: Newberry-Bane Cave. Scott Co.: Grigsby
Cave.

Order Lumbriculida

Family Lumbriculidae

Spelaedrilus multiporus Cook (TB)

Virginia. — Russell Co.: Smiths Cave (type locality).

Stylodrilus ( Bythonomus ) beattiei Cook (TB)

Virginia. — Tazewell Co.: Steeles Cave.

Comments. — Also recorded from three caves in southern West
Virginia (Cook 1975).

Genus (?) species (?)

Virginia. — Lee Co.: McClure and Spangler caves.

Comments. — These populations represent an undescribed, troglobitic
species (D. G. Cook, pers. comm.)

PHYLUM MOLLUSCA

Both aquatic and terrestrial snails (class Gastropoda) have been
collected from caves in Virginia and eastern Tennessee, but the former
are far more common in subterranean habitats than are the latter. Aside
from several species of Goniobasis, which are sometimes abundant in
karst springs and occasionally penetrate some distance into cave streams,
aquatic cave snails of the Appalachians are members of the family
Hydrobiidae, and most apparently belong to the genus Fontigens (Fig.
13D). Cavernicolous hydrobiids commonly inhabit the undersides of
flat rocks in small streams with relatively constant flow.

Owing to the fact that the taxonomy of the cave and spring
hydrobiids is based largely on shell morphology (see Hubricht 1976),
which is often highly variable, identities of some of the species listed
below are, in our opinion, questionable. There are a number of
peculiarities that are perplexing about the geographic distribution (Fig.
7) and ecology of these species. For example, Fontigens aldrichi has
been recorded from caves and springs in the Ozarks and Appalachians
and is represented in both regions by eyed, pigmented populations living
principally in springs, and by eyeless, unpigmented populations living
principally in caves (Hubricht 1976, Peck and Lewis 1978). Another
species, F. orolibas, although resticted to the Appalachians, has been
identified from eyed, pigmented populations living in springs in the Blue
Ridge Mountains and from blind, unpigmented populations living in
caves in karst valleys to the west (see Hubricht 1957, 1976). Similarly,
blind, white snails from caves in the Powell Valley of southwestern
Virginia have been tentatively assigned by Hubricht (1976) to

Invertebrate Cave Fauna

Fig. 7. Distribution of aquatic cavernicolous snails ( Fontigens ) in the study
area. Spring localities for F. orolibas in the Blue Ridge Mountains not shown.

F. nickliniana, an epigean species previously recorded from a number
of localities in the eastern United States. However, the recent study of a
population in Unthanks Cave, utilizing internal anatomy in combination
with shell morphology, suggests that one or more undescribed troglobitic
species inhabit caves of the Powell Valley (R. Hershler and F. G.
Thompson, in litt.). Another population in need of additional taxonomic
study and clarification is one from Skyline Caverns tentatively identified
by Morrison (1949, pers. comm.) as an undescribed species of the
European subterranean genus Lartetia. Morrison (1949, pers. comm.)
has taken a different view from that of Hubricht and believes that
Fontigens in the Appalachians represents a complex of closely similar
genera composed collectively of many well-isolated troglobites. Un-
fortunately, his observations are mostly unpublished and thus unavailable
for biogeographic analysis.

Terrestrial cave snails were usually collected from damp, rotting
wood; only a few populations were noted. Seven species in three
families have been recorded to date. Helicodiscus notius specus
(Helicodiscidae), a “somewhat degenerate form” (see Barr 1967a) of the
widespread, primarily epigean H. notius (Hubricht 1962) was originally

John R. Holsinger and David C. Culver

described from Burnet Cave in Barren County, Ky., and has since been
identified from Bristol Caverns in east Tennessee by Hubricht (in litt.).
Helicodiscus inermis, recorded from two caves in west-central Virginia,
is also reported from caves in Alabama, Georgia, and Tennessee, and
from surface localities elsewhere in the eastern and southern United
States (see Hubricht 1964, 1985; Holsinger and Peck 1971).

In the Polygyridae, Mesodon appressus is recorded from Flannery
Cave in Scott County, and this species, in its broadest sense, is also
reported from caves in Kentucky and Tennessee by Barr (1961, 1967a)
and Hubricht (1964). Glyphyalinia specus (Zonitidae), a white, apparently
blind species unknown outside caves and possibly a troglobite, is
recorded from Bristol Caverns in Sullivan County and also from caves
in Alabama, Georgia, Kentucky, middle Tennessee, and possibly West
Virginia (see Hubricht 1965, 1985; Barr 1967a; Holsinger and Peck
1971; Holsinger et al. 1976). Another zonitid, Zonitoides arboreus,
probably a troglophile, is recorded from one cave in the study area and
from many other caves in the east-central and southeastern United
States (see Hubricht 1964, Holsinger and Peck 1971, Peck and Lewis
1978).

Order Mesogastropoda

Family Pleuroceridae

Goniobasis elavaeformis (Lea) (TX or AC)

Tennessee. — Hancock Co.: Cantwell Valley Cave.

Goniobasis simplex (Say) (TX)

Virginia. — Lee Co.: Surgener and Young-Fugate caves. Scott Co.:
Alley and McDavids caves.

Goniobasis sp.

Virginia. — Scott Co.: Speers Ferry Cave.

Family Hydrobiidae

Fontigens aldrichi (Call and Beecher) (?) (TP?)

Virginia. — Bath Co.: Blowing and Butler-Sinking Creek caves.
Frederick Co.: Ogdens Cave.

Comments. — In or near the study area this species is also recorded
from springs in Highland Co., Va., and Washington Co., Md.
(see Hubricht 1976).

Fontigens orolibas Hubricht (TP)

Virginia. — Giles Co.: Smokehole, Starnes, and Tawneys caves.
Tazewell Co.: Hugh Young Cave. Warren Co.: Skyline Caverns.
Fontigens sp. (near nicklinianal ) (TB?)

Virginia. — Lee Co.: Gallohan No. 1, Smiths Milk, and Spangler
caves.

Fontigens spp.

Virginia. — Bath Co.: Witheros Cave. Lee Co.: Unthanks Cave.
Washington Co.: Perkins Cave.

Invertebrate Cave Fauna

Lartetia (?) sp. (TB)

Virginia. — Warren Co.: Skyline Caverns.

Order Stylommatophora

Family Helicodiscidae

Helicodiscus inermis Baker (TP or TX)

Virginia.- — Augusta Co.: Grand Caverns. Bath Co.: Dunns Cave.
Helicodiscus notius specus Hubricht (TP or TX)

Tennessee. — Sullivan Co.: Bristol Caverns.

Family Polygyridae

Mesodon apprcssus (Say) (TX?)

Virginia. — Scott Co.: Flannery Cave.

Mesodon normalis (Pilsbry) (AC?)

Tennessee. — Claiborne Co.: English Cave.

Polygyra albolabris Say (AC?)

Virginia. — Shenandoah Co.: Shenandoah Caverns.

Family Zonitidae

Glyphyalinia specus Hubricht (TB?)

Tennessee. — Sullivan Co.: Bristol Caverns.

Zonitoides arboreus (Say) (TP?)

Virginia. — Rockingham Co.: Endless Caverns.

PHYLUM ARTHROPODA: SUBPHYLUM CRUSTACEA

A significant number of the species recorded from caves in Virginia
and eastern Tennessee are crustaceans and include copepods, amphipods,
isopods, crayfishes, and possibly ostracods. The vast majority, however,
are amphipods and isopods, both of which are frequently well represented
in aquatic cave communities.

Class Copepoda

Cave copepods are very poorly known from the study area and
only a single species has been recorded to date. However, as pointed out
earlier, no attempt was made to sample microscopic cave faunas, and
the lack of data on tiny crustaceans such as copepods and ostracods is
to be expected.

Order Cyclopoida

Family Cyclopidae

Cyclops vernalis Fischer (TX)

Virginia. — Tazewell Co.: Hugh Young Cave.

Comments. — Extremely variable and widespread species sometimes
recorded from caves (e.g., in Georgia, Kentucky, New Mexico,
Texas) (see Barr 1967a, Reddell 1965, Barr and Reddell 1967,
Holsinger and Peck 1971).

John R. Holsinger and David C. Culver

Class Ostracoda

Hobbs (1975) alluded to the presence of the ectocommensal ostracod
Phymocythere phyma (Hobbs and Walton) (Entocytheridae) in Virginia
and West Virginia caves but gave no specific records. Because a
principal host of this species is Cambarus bartonii, a crayfish found in
caves of the study area (see below), the occurrence of this ostracod in
Virginia and east Tennessee caves should be expected. However, to our
knowledge there are no published records.

Class Malacostraca

Malacostracan crustaceans are represented in study-area caves by
three orders: Amphipoda (2 families, 3 genera, 21 species), Isopoda (6
families, 10 genera, 23 species), and Decapoda (1 family, 1 genus, 2
species).

Order Amphipoda

Amphipods are common faunal components of cave waters where
they are usually found among gravels or under small rocks in streams,
on the organically enriched mud substrate of pools fed by drips and or
seeps, and rarely in deep phreatic lakes. A total of 21 species, all in the
suborder Gammaridea, have been recorded, of which 20 are of troglobitic
facies and known only from groundwater biotopes. The troglobitic
species belong to the genera Stygobromus, Bactrurus, and Crangonyx ,
all in the family Crangonyctidae; the single troglophile, Gammarus
minus , is in the family Gammaridae.

A majority of the species (18) have been assigned to Stygobromus , a
large, exclusively subterranean genus that is distributed throughout a
large part of North America (Holsinger 1977, 1978, 1986a, 1986b). Most
species of Stygobromus from the study area have narrowly circumscribed
ranges (Fig. 8, 9), and many are local endemics; three are known only
from their type localities. Only four species listed below occur outside
the study area, and none extends beyond this area for a great distance.
The most common and widespread species is S. mackini, which is
distributed from Monroe County in southern West Virginia (New River
drainage) southwestw'ard to Roane County in eastern Tennessee
(Tennessee River drainage) (Holsinger 1978).

Bactrurus is represented by a single, undescribed (provisionally
recognized) species that is recorded to date from only three caves in the
Pow'ell Valley (Fig. 8). This is one of five or six undescribed species in
the genus (Holsinger 1986b) and the first to be found in the Appalachian
Valley. Three described species are reported from caves and other
groundwater biotopes in the eastern and central United States (see
Holsinger 1972, 1986a, 1986b).

Invertebrate Cave Fauna

Bactrurus sp.

Stygobromus cumberlandus

S. interitus

S. abditus

S. finleyi

S. leensis

S. mackini

KENTUCKY

VIRGINIA

Fig. 8. Distribution of troglobitic amphipods ( Bactrurus and Stygobromus ) in

0 25 50 km

0 25 50 miles

the study area. All localities for S. mackini (including those in Anderson and
Grainger counties, Tenn., and Mercer and Monroe counties, W.Va.) shown
except Berry Cave, Roane County, Tenn. Two symbols in a circle indicate two
species from the same cave.

Stygobromus

emarginatus group ▲ gracilipes group

1 . fergusoni

2. hoffmani

3. morrisoni

4. mundus
ephemerus group

5. ephemerus

6. estesi

7. conradi

8. gracilipes

4 spinosus group

9. pseudospinosus
• ungrouped species

10. baroodyi

11. biggersi

12. stegerorum

WEST VIRGINIA

5^ VIRGINIA

25 ^0 km

50 miles

Fig. 9. Distribution of troglobitic amphipods {Stygobromus) in the study area.
Single localities for S. morrisoni in Hardy and Pendleton counties, W.Va.,
also shown. Two symbols in a circle indicate two species from the same cave.

John R. Holsinger and David C. Culver

Crangonyx antennatus is recorded from numerous caves in the
Powell Valley and the middle and lower parts of the Clinch Valley (Fig.
10, 13B). It is the most common and, after Stygobromus mackini , most
widespread troglobitic amphipod in Virginia and eastern Tennessee. Its
range, which needs further evaluation in view of morphological variation,
extends south of the study area through eastern Tennessee into
northwestern Georgia and northern Alabama and then westward to
south-central Tennessee (see Holsinger 1969a, 1972, 1986a, 1986b). In
addition to caves, C. antennatus has been collected occasionally from
surface springs or seeps, including Spout Spring in Lee County
(Holsinger 1969a).

Gammarus minus is recorded from caves, springs, and small spring-
fed streams, principally in karst regions of the eastern and east-central
United States (see Holsinger and Culver 1970; Holsinger 1969a, 1972;
Stock 1986). In the study area this species is abundant only in caves of
the Ward Cove karst area (upper Clinch drainage) in Tazewell County
(Fig. 10). Here a majority of the populations have developed a trog-
lomorphic facies referred to as Form I in an earlier paper (Holsinger
and Culver 1970). Within the study area, G. minus is more widespread
in springs than in caves and is recorded from the former habitat in the
Tennessee counties of Claiborne, Hancock, and Sullivan, and the Virginia
counties of Alleghany, Bath, Botetourt, Craig, Frederick, Lee, Mont-
gomery, Pulaski, Russell, Scott, Shenandoah, Tazewell, Washington,
and Wythe.

Family Crangonyctidae
Bactrurus sp. (TB)

Tennessee. — Claiborne Co.: Kings Saltpetre and Saur Kraut caves.

Virginia. — Lee Co.: Cumberland Gap Saltpetre Cave.

Crangonyx antennatus Packard (TB)

Tennessee. — Campbell Co.: Meredith Cave. Claiborne Co.: Buis
Saltpetre, Chadwells, English, Hauser Spring, John Lard, Kings
Saltpetre, and Station Creek caves. Grainger Co.: Horseshoe
Cave. Hancock Co.: Cantwell Valley, Fairmont School, and
Subers caves. Hawkins Co.: Pearson Cave. Sullivan Co.: Morrills
Cave. Union Co.: Oaks, Wolf, and Wrights caves.

Virginia. — Lee Co.: Baileys, Bowling, Cave Springs, Cedar Hill,
Chances, Combs No. 1, Cope, Crouse, Cudjos (Cavern),
Cumberland Gap Saltpetre, Frazier, Gallohan No. 1 and 2,
Garretts, Gibson-Frazier, Gilliam, Glen Olingers, Golf Course
No. 1 and 2, Gregorys, Jones Saltpetre, Knapper, Lesters, Litton
No. 1, Lucy Beatty, McClure, Minors Saltpetre, Molly Wagle,
Mount Moriah Pit, Olinger, Roadside No. 1, Seal, Slemp, Smiths
Milk, Spangler, Sweet Potato, Taylor Pit, Thompson, Thompson

Invertebrate Cave Fauna

Fig. 10. Distribution of cavernicolous amphipods ( Crangonyx and
Gammarus) in the study area. Only cave localities shown for G. minus.
Anderson County, Tenn., records for C. antennatus also indicated. Two symbols
in a circle indicate two species from the same cave.

Cedar, Unthanks, Watsons No. 1, and Young-Fugate caves.
Scott Co.: McDavids, Speers Ferry, and Spurlock caves. Wise
Co.: Wildcat Cavern and Wildcat Saltpetre Cave.

Stygobromus (species listed by group as indicated)

cumberlandus group
Stygobromus cumberlandus Holsinger (TB)

Virginia. — Lee Co.: Baileys and Cliff caves. Wise Co.: Wildcat
Saltpetre Cave (type locality).

Comments. — Also recorded from a well at Duffield in Scott County
(Holsinger 1978).

Stygobromus interitus Holsinger (TB)

Virginia. — Craig Co.: New Castle Murder Hole Cave (type locality).

emarginatus group

Stygobromus fergusoni Holsinger (TB)

Virginia. — Montgomery Co.: Old Mill and Slussers Chapel (type
locality) caves.

Stygobromus hoffmani Holsinger (TB)

Virginia. — Alleghany Co.: Lowmoor (type locality) and Me Elwee

caves.

John R. Holsinger and David C. Culver

Stygobromus morrisoni (Holsinger) (TB)

Virginia.— Bath Co.: Witheros Cave (type locality). Highland Co.:
Corbett Cave.

Comments. — Also recorded from single caves in Hardy and
Pendleton counties, W. Va. (Holsinger 1978).

Stygobromus mundus (Holsinger) (TB)

Virginia.— Bath Co.: Witheros Cave (type locality).

Comments. — Also recorded from a tributary to the Cowpasture
River in Alleghany County (see Holsinger 1967a, 1978).

ephemerus group

Stygobromus ephemerus (Holsinger) (TB)

Virginia. — Giles Co.: Canoe and Tawneys (type locality) caves.

Stygobromus estesi (Holsinger) (TB)

Virginia.— Craig Co.: New Castle Murder Hole and Rufe Caldwell
(type locality) caves.

gracilipes group

Stygobromus conradi (Holsinger) (TB)

Virginia.— Bath Co.: Breathing (type locality) and Butler-Sinking
Creek caves.

Stygobromus gracilipes (Holsinger) (TB)

Virginia. — Frederick Co.: Ogdens Cave. Rockingham Co.: Deer
Hole, Endless (Caverns), Massanutten (Caverns), and Three-D
Maze caves. Warren Co.: Skyline Caverns (type locality).
Comments. — Also recorded from caves just north of the study area
in Washington Co., Md.; Franklin Co., Pa.; and Berkeley and
Jefferson counties, W. Va. (Holsinger 1967a, 1978).

mackini group

Stygobromus abditus Holsinger (TB)

Virginia. — Pulaski Co.: James (type locality) and Sam Bells caves.

Stygobromus finleyi Holsinger (TB)

Tennessee. — Claiborne Co.: English Cave (type locality).

Stygobromus leensis Holsinger (TB)

Virginia. — Lee Co.: Gallohan No. 2, Litton No. 1 (type locality),
and Skull caves.

Stygobromus mackini Hubricht (TB)

Tennessee. — Hancock Co.: Cantwell Valley Cave. Hawkins Co.:
Sensabaugh Saltpetre Cave. Union Co.: Lost Creek, Oaks, and
Ridenour Pit caves.

Virginia. — Giles Co.: Ballards, Starnes, and Tawneys caves. Russell
Co.: Banners Corner, Bundys No. 2, Burns, Grays, Jessie,
Johnson, Munsey, Porgie Bundys, “Sikes” (Hubricht 1943:697;
type locality), and Smith Drop caves. Scott Co.: Blair-Collins,
Blowing Hole, Deep Spring, Flannery, Greears Sweet Potato,

Invertebrate Cave Fauna

Grigsby, Hill, Jack, Jackson, Kerns Smoke-Hole, McDavids,
McNew, Moccasin Valley, Natural Tunnel (Cavern), Pond,
Spurlock, Taylor No. 1, Winding Stair, and Wolfe caves. Smyth
Co.: Buchanan Saltpetre, McMullin (?), and Tilson Saltpetre
caves. Tazewell Co.: Cauliflower, Chimney Rock, Crocketts,
Fallen Rock, Glenwood Church, Hugh Young, Lost Mill No. 1,
Steeles, and Ward Cove caves. Washington Co.: Singleton Cave.
Wise Co.: Wildcat Saltpetre Cave.

Comments. — Also recorded from caves in Anderson, Grainger, and
Roane counties, Tenn., and Mercer and Monroe counties, W.
Va., and occasionally from small springs or seeps in Giles,
Tazewell, and Washington counties, Va. (see Holsinger 1978).

spinosus group

Stygobrornus pseudospinosus Holsinger (TB)

Virginia. — Page Co.: Luray Caverns (type locality).

Ungrouped Species

Stygobrornus baroodyi Holsinger (TB)

Virginia. — Rockbridge Co.: Bathers (type locality), Bell, Billy
Williams, Buck Hill, Grahams, and Showalter caves.

Stygobrornus bigger si Holsinger (TB)

Virginia. — Frederick Co.: Johns and Ogdens (type locality) caves.
Comments. — Also recorded from caves just north of the study area
in Washington Co., Md., Franklin Co., Pa., and Jefferson Co.,
W. Va. (Holsinger 1978).

Stygobrornus stegerorurn Holsinger (TB)

Virginia. — Augusta Co.: Madisons Saltpetre (type locality) and
Stegers Fissure caves.

Stygobrornus spp.

Tennessee. — Campbell Co.: Norris Dam Cave.

Virginia. — Craig Co.: New Castle Murder Hole Cave. Washington
Co.: Neals Cave.

Comments. — These populations may represent several undescribed
species, all probably in the rnackini group.

Family Gammaridae

Garnrnarus minus Say (TP)

Tennessee. — Hancock Co.: Cantwell Valley Cave.

Virginia. — Giles Co.: Canoe, Smokehole, and Tawneys caves. Russell
Co.: Smiths Cave. Scott Co.: Alley and Wolfe caves. Tazewell
Co.: Bowens, Cauliflower, Crocketts, Fallen Rock, Gillespie
Water, Hugh Young, Lawson, Lost Mill No. 1 and 3, Quarry,
Rosenbaums Water, and Wagoners caves. Washington Co.:
Hookers Rock Cave.

John R. Holsinger and David C. Culver

Order Isopoda

Isopods are represented in the regional cave fauna by three
suborders: Asellota, Flabellifera, and Oniscoida. All asellotids from
caves in Virginia and eastern Tennessee are in the large, Holarctic,
freshwater family Asellidae. Flabelliferans are represented by a single,
unique member of the predominantly marine family Cirolanidae.
Oniscoids are terrestrial and are represented by the families Arma-
dillidiidae, Ligiidae, Oniscidae, and Trichoniscidae.

Cavernicolous asellids are usually associated with the gravel or
rock substrate of small streams or the mud-bottom substrate of drip/
seep pools. Some species apparently prefer riffle zones, whereas others
are sometimes seen in large concentrations on flowstone surfaces covered
by thin films of moving water (see Culver 1973a, Estes and Holsinger
1982). Two genera, Caecidotea and Lirceus , occur in study-area caves.
The former is represented by 1 1 species, 9 of which are troglobites; the
latter is represented by two species (both troglobites) and possibly
several undescribed (non-troglobitic) ones as well (see Henry et al.
1986).

Although several troglobitic species of Caecidotea have relatively
wide ranges, their distributions generally correspond rather closely to
drainage basins (Fig. 11, 12). Some of the wide-ranging species, such as
C. richardsonae, C. recurvata , and C. pricei , have also been collected
occasionally from subterranean waters outside caves (e.g., seeps, wells).
Five species, viz., C. holsingeri, C. incurva , C. recurvata (Fig. 13C), C.
richardsonae , and C. pricei , have ranges that extend beyond the study
area, whereas C. bowmani , C. henroti , and the undescribed species from
Cliff Cave (Lee County) are local endemics with very restricted ranges.
Of particular interest here is C. bowmani , at present known only from a
drain-tile habitat in Rockbridge County (see Lewis 1980). Although this
species is not recorded from a cave per se, it is of troglobitic facies and,
with careful searching, will possibly be found in caves. Because of this
we have listed it as a troglobite. Moreover, careful reevaluation of
collections from caves in the James River basin previously assigned to
C. vandeli by Fleming (1972) may very well result in their reassignment
to C. bowmani.

Because taxonomic studies of Caecidotea have placed almost
complete emphasis on the morphology of the male second pleopod and
tended toward the “lumping” of species, a careful reevaluation of the
systematics of the Appalachian cave species is warranted. In support of
this view is the recent research by J. J. Lewis (in progress) on the
systematics of subterranean Caecidotea of the east-central United States,
which has revealed additional new species from material previously
assigned to described taxa. Based on geographic distribution and ecology,
we suspect that C. richardsonae , for example, can; with careful

Invertebrate Cave Fauna

NORTH CAROLINA

Fig. 11. Distribution of aquatic troglobitic isopods ( Antrolana and
Caecidotea) in the study area. All localities for C. recurvata (including a single
cave in Washington County, Tenn.) shown except spring in Knox County,
Tenn. Two symbols in a circle indicate two species from the same cave.

•fr Antrolana lira
♦ Caeqdotea bow m a n i
■ C. henroti
□ C. holsingeri

WEST VIRGINIA

* C. incur va

* C. vandeli

• C recurvata

KENTUCKY

VIRGINIA

50 km

50 miles

Fig. 12. Distribution of aquatic troglobitic isopods ( Caecidotea and Lirceus ) in
the study area.

NORTH CAROLINA

Caecidotea pricei
C. richardsonae
C. sp. A
C_ spp.

Lirceus culveri
L. usdagalun

WEST VIRGINIA

KENTUCKY

VIRGINIA

0 25 50 km

0 25 50 miles

John R. Holsinger and David C. Culver

taxonomic analysis, be shown to be a complex of closely related species.
This may also be true of other species,' such as C. holsingeri and C.
pricei.

In addition to the troglobitic species of Caecidotea , C. intermedia
and C. r. racovitzai are both unknown from caves except for the records
cited below from Tazewell and Smyth counties and a record for the
former from southern Illinois (see Lisowski 1979). Caecidotea intermedia
is relatively common in the east-central United States and southeastern
Canada, whereas C. r. racovitzai is relatively common in southeastern
Canada but sparsely distributed in the east-central and northeastern
United States (Williams 1970, Fleming 1972). The Virginia cave
populations warrant further study, especially since they are geograph-
ically and ecologically isolated from other localities documented for
their respective species.

Lirceus is commonly found in springs and occasionally in caves in
eastern North America, but only two troglobitic species have been
recognized to date. Both of these occur in southwestern Virginia, where
their respective ranges (Fig. 12) are greatly delimited as indicated in the
list below. At least one undescribed troglophile inhabits caves and
springs in the Ward Cove karst area of Tazewell County where several
large populations composed of very pale individuals with tiny eyes have
been noted.

The sole member of the family Cirolanidae in the Appalachians is
Antrolana lira, an unusual monotypic form that is restricted to an
isolated groundwater aquifer in Cave Hill in Augusta County (Fig. 1 1,
13 A). This species inhabits lakes of deep phreatic water in two caves
(Bowman 1964, Collins and Holsinger 1981, Botosaneanu et al. 1986). It
is the only freshwater cirolanid in North America north of Texas,
Mexico, and the West Indies, and is therefore of great interest
zoogeographically.

Of the four families of oniscoid isopods, only the Trichoniscidae
contains troglobites. The remainder contain epigean species, some of
which, however, are commonly associated with cave habitats.
Armadillidium vulgare (family Armadillidiidae), one of the so-called
“pill bugs” is a common, widespread epigean species sometimes found
under damp wood in the entrance zone of caves. This species has been
collected from a few Virginia caves.

Ligiidae is represented in study-area caves by Ligidium elrodii , a
species sometimes abundant on wet organic detritus flushed into caves
by flooding. It is recorded from epigean localities in the eastern United
States and southern Canada (Schultz 1970). In addition to the cave
records cited below for Virginia and east Tennessee, it is recorded from
caves in northern Arkansas, southern Illinois, northwestern Georgia,
and southern West Virginia (Schultz 1970, Holsinger and Peck 1971,

Invertebrate Cave Fauna

Fig. 13. Aquatic troglobites from the study area (approximate body lengths in
parentheses): A, isopod, Antrolana lira (16 mm); B, amphipod, Crangonyx
antennatus (14 mm); C, isopod, Caecidotea recurvata (15 mm); D, snail,
Fontigens sp. (3 mm); E, planarian, Sphalloplana comsimilis (14 mm).

John R. Holsinger and David C. Culver

Holsinger et al. 1976, McDaniel and Smith 1976, Peck and Lewis 1978).
Five subspecies have been designated by Schultz (1970), of which
three — leensis , scottensis , and hancockensis — occur in southwestern
Virginia and northeastern Tennessee.

Cylisticus convexus (family Oniscidae), a common epigean species
throughout the United States, has been found in a few Virginia caves
and is also reported from caves elsewhere in the southeastern and south-
central parts of the country (see Schultz 1970, Franz and Slifer 1971,
Holsinger and Peck 1971, Peck and Lewis 1978, Hobbs and Flynn
1981).

Cavernicolous trichoniscid isopods are usually found on damp to
wet, decomposing wood. Six species in four genera are recorded from
caves in the study area. Three of these species are troglobites (Fig. 14):
Amerigoniseus henroti (Fig. 3 IE) from caves in central Lee County
(Holsinger 1967b, Vandel 1977), A. paynei from caves in the Clinch
Valley of eastern Tennessee (Muchmore 1970a), and Miktoniscus r.
racovitzai from caves in the James and Shenandoah river drainage
basins (Vandel 1965a).

Vandel (1977) considered A. paynei Muchmore (1970a) synonymous
with A. nicholasi , a species described earlier by Vandel (1965a) from
Columbia Caverns in middle Tennessee just west of Nashville in Dickson
County. In our opinion, however, the small morphological differences
between this population and those from eastern Tennessee noted by
Muchmore (1970a), combined with the rather wide geographic separation
of the populations, provides a good reason for the recognition of two
separate species.

Outside the study area. Miktoniscus r. racovitzai is reported from
Slacks Cave in Scott County, Ky., by Vandel (1965a); and a second
subspecies, M. r. oklahomensis, was designated by Vandel for a single
cave population in Murray County, Oklahoma. The systematic status of
these populations is questionable, in view of their disjunct distributions,
and should be carefully reevaluated.

The non-troglobitic trichoniscids include: Haplophthalmus danicus ,
recorded from caves and epigean localities throughout a large part of
North America and also found in Europe (see Vandel 1965a. Holsinger
et al. 1976); Miktoniscus medcofl (synonym = M. alabamensis Muchmore;
see Muchmore 1964, Schultz 1976), recorded from many caves in the
southeastern United States; and Trichoniscus pusillus, a common epigean
species occasionally found in caves (see Holsinger et al. 1976).

Suborder Asellota

Family Asellidae

Caecidotea (species listed by groups as indicated)

Invertebrate Cave Fauna

Fig. 14. Distribution of terrestrial troglobitic isopods ( Arnerigonicus and
Miktoniscus ) in the study area. Localities for A. paynei in Anderson County,
Tenn., also shown.

WEST VIRGINIA

(

KENTUCKY /%

VIRGINIA

0__ 25 50 km

0 25 50 miles

NORTH CAROLINA

• Amerigofiiscus henroti
▼ A_ paynei

A Miktoniscus racovitzai

cannula group

Caecidotea bowmani Lewis (TB)

Virginia. — Rockbridge Co.: drain tile near Natural Bridge (type
locality).

Caecidotea henroti (Bresson) (TB)

Virginia. — Giles Co.: Smokehole (type locality) and Tawneys caves.
Pulaski Co.: James Cave.

Caecidotea holsingeri (Steeves) (TB)

Virginia. — Bath Co.: Butler-Sinking Creek Cave.

Comments. — Recorded from numerous caves in West Virginia
(Greenbrier, Monroe, Pocahontas, and Randolph counties) and
from one cave in Garrett Co., Md. (Steeves 1963a, 1969; Holsinger
et al. 1976; Lewis 1980).

Caecidotea incurva (Steeves and Holsinger) (TB)

Virginia. -Smyth Co.: McMullin Cave. Wythe Co.: Groseclose Cave
No. 1.

Comments. — Also recorded from single caves in Blount and Roane
counties, Tenn. (Steeves and Holsinger 1968).

Caecidotea vandeli (Bresson) (TB)

Virginia. — Bath Co.: Blowing Cave. Botetourt Co.: Brough Cave

John R. Holsinger and David C. Culver

No. 2. Giles Co.: New River Cave. Montgomery Co.: Erhart
(type locality), Old Mill, and Slussers Chapel caves. Roanoke
Co.: Goodwins Cave (?).

stygia group

Caecidotea recurvata (Steeves) (TB)

Tennessee. — Campbell Co.: Meredith Cave. Claiborne Co.: Buis
Saltpetre, Chadwells, English, Hauser Spring, Kings Saltpetre,
and Station Creek caves. Hancock Co.: Subers Cave. Union Co.:
Coppock, Lost Creek, Ridenour Pit, Wolf, and Wright caves.

Virginia. — Lee Co.: Baileys, Bowling, Cave Springs, Combs No. 1,
Cope, Crouse, Fisher, Gallohan No. 1 and 2, Gilliam, Golf
Course No. 1 and 2, Kinzer Hollow, Litton No. 1, McClure,
Minors Saltpetre, Molly Wagle, Roadside No. 1, Seal, Skull,
Smiths Milk, Spangler, Sweet Potato, Taylor Pit, T-Bone,
Thompson Cedar, Unthanks (type locality), and Young-Fugate
caves. Russell Co.: Banners Corner, Breeding, Bundys Pearl,
Burns, Daugherty, Grays, Indian, Jessie, Johnson, Munsey, Porgie
Bundys, Seven Springs, Smiths, and “Sikes” (Hubricht 1943:697)
caves. Scott Co.: Blair-Collins, Coley No. 2, Flannery, Jack.
McDavids, Spurlock, and Taylor No. 1 caves. Smyth Co.:
McMullin Cave. Washington Co.: Brass Kettle Hole Cave, Wise
Co.: Hairy Hole, Kelly, Little Kennedy, Parsons, Rocky Hollow,
and Wildcat Saltpetre caves.

Comments. — Also recorded from a spring in Knox County (see
Fleming 1972) and a cave in Washington County, both in eastern
Tennessee. “ Asellus forcipitus n. sp.” recorded from English
Cave by Dearolf (1953), was never described in the literature and
should be regarded as a nomen nudum.

Caecidotea richardsonae Hay (TB)

Tennessee. — Claiborne Co.: Buis Saltpetre, Cline, and Holt caves.
Grainger Co.: Horseshoe Cave. Hancock Co.: Fairmont School
Cave. Hawkins Co.: Sensabaugh Saltpetre Cave.

Virginia. — Lee Co.: Gregory, Olinger, and Smiths Milk caves.
Scott Co.: Blair-Collins, Horton, Moccasin Valley, and Wolfe
caves. Tazewell Co.: Bowens, Fallen Rock, Hugh Young, Lost
Mill No. 3, Rosenbaums Water, and Stonley caves.

Comments. — Also reported from caves and occasionally wells in
central and northeastern Alabama, northwestern Georgia, and
south-central Tennessee (Steeves 1963b, 1969; Fleming 1972).

Ungrouped Species

Caecidotea intermedia (Forbes) (AC?)

Virginia. — Smyth Co.: Interstate-81 Cave.

Invertebrate Cave Fauna

Caecidotea racovitzai racovitzai Williams (TX?)

Virginia. — Tazewell Co.: Lawson and Quarry caves.

Caecidotea pricei Levi (TB)

Virginia. — Augusta Co.: Barterbrook Spring Cave. Frederick Co.:
Ogdens Cave. Page Co.: Will Mauck Cave. Rockbridge Co.:
Bathers, Bell, Billy Williams, Showalters, and Tolleys caves.
Rockingham Co.: Endless Caverns. Shenandoah Co.: Flemings
Cave. Warren Co.: Skyline Caverns. Also: “Water Cave, Va.”
(Dearolf 1953:227).

Comments. — Outside the study area, this species is recorded
from groundwater habitats (mostly caves) in central Maryland,
southern Pennsylvania, and eastern West Virginia (Holsinger and
Steeves 1971, Franz and Slifer 1971, Holsinger 1976, Holsinger et
al. 1976). Within the study area, it is also recorded from three
small springs or seeps in Rockingham County and one spring in
Rockbridge County (see Holsinger and Steeves 1971).

Asellus condei, described by Chappuis (1957) from Ogdens
Cave, is considered a synonym of C. pricei (see Holsinger and
Steeves 1971, Fleming 1973).

Caecidotea sp. A (TB)

Virginia. — Lee Co.: Cliff Cave.

Comments. — Fleming (1972) listed this population as belonging to
Caecidotea scrupulosa Williams, but subsequent examination
indicates that it represents an undescribed troglobitic species.

Caecidotea spp. (TB)

Tennessee. — Hancock Co.: Panther Creek Cave. Union Co.: Oaks
Cave.

Virginia. — Alleghany Co.: Paxtons Cave. Botetourt Co.: Eagle
Rock Cave. Highland Co.: Aqua, Better Forgotten, and Roaring
Springs caves. Lee Co.: Gilley Cave. Scott Co.: Grigsby and
Pond caves. Smyth Co.: Buchanan Saltpetre Cave. Washington
Co.: Neals and Reeds No. 1 caves.

Comments. — Collections from the caves listed above lacked males,
therefore precluding specific determinations.

Lirceus culveri Estes and Holsinger (TB)

Virginia. — Scott Co.: McDavids Cave (type locality).

Lirceus usdagalun Holsinger and Bowman (TB)

Virginia. — Lee Co.: Gallohan No. 1 (type locality), Gallohan No. 2,
Surgener, and Thompson Cedar caves.

Lirceus spp.

Tennessee. — Claiborne Co.: Billingsley and Lower Coonsies Creek
caves. Hancock Co.: Lawsons Cave No. 3.

Virginia. — Lee Co.: Baileys, Olinger, and Young-Fugate
caves. Rockbridge Co.: Tolleys Cave. Scott Co.: Alley, Wolfe,

John R. Holsinger and David C. Culver

Coley No. 2, and Speers Ferry caves. Tazewell Co.: Fallen Rock,
Gillespie Water, and Hugh Young caves. Washington Co.:
Singleton Cave.

Comments. — Several species, some apparently undescribed, are
represented in these collections; none appears to be of troglobitic
facies (see Holsinger and Bowman 1973).

Suborder Flabellifera

Family Cirolanidae

Antrolana lira Bowman (TB)

Virginia. — Augusta Co.: Madisons Saltpetre (type locality) and
Stegers Fissure caves.

Suborder Oniscoidea

Family Armadillidiidae

Armadillidium vulgare (Latreille) (TX or AC)

Virginia. — Alleghany Co.: Lowmoor Cave. Augusta Co.: Madisons
Saltpetre Cave. Bath Co.: Roy Lyle Cave. Page Co.: Foltz Cave
No. 1. Rockbridge Co.: Doll House and Tolleys caves.

Family Ligiidae

Ligidium elrodii (5. lat .) (Packard) (TB)

Tennessee. — Claiborne Co.: Lower Coonsies Creek Cave. Hancock
Co.: Cantwell Valley Cave. Sullivan Co.: Bristol Caverns.
Virginia. — Lee Co.: Bowling and Waltons caves. Scott Co.: Coley
Cave No. 2.

Ligidium sp.

Virginia. — Craig Co.: New Castle Murder Hole Cave. Lee Co.:
Carter Cave.

Comments. — These populations are probably L. elrodii, but the
lack of males precludes specific determination.

Family Oniscidae

Cylisticus convexus (De Greer) (TX)

Virginia. — Botetourt Co.: Thomas Cave. Roanoke Co.: Dixie
Caverns. Smyth Co.: Stones No. 2 and Sugar Grove No. 10
caves.

Family Trichoniscidae

Amerigoniscus henroti Vandel (TB)

Virginia. — Lee Co.: Cope, Gallohan No. 1, Gilley (type locality),
Kinzer Hollow, Smiths Milk, Spangler, Sweet Potato, and
Unthanks caves.

Amerigoniscus paynei (Muchmore) (TB)

Tennessee. — Hancock Co.: Fairmont School Cave. Union Co.:
Lost Creek and Wolf caves.

Invertebrate Cave Fauna

Comments. — This species is also recorded from Hill and Offutts
(type locality) caves just south of the study area in Anderson
County and may also inhabit Melton Hill Cave No. 1 in Roane
County, Tenn. (see Muchmore 1970a).

Haplophthalmus danicus Budde-Lund (TP or TX)

Virginia. — Lee Co.: Ruths Cave. Page Co.: Luray Caverns. Pulaski
Co.: James Cave. Roanoke Co.: Goodwins Cave. Rockbridge
Co.: Showalters Cave. Rockingham Co.: Massanutten Caverns.
Russell Co.: Banners Corner Cave. Tazewell Co.: Wagoners
Cave.

Miktoniscus medcofi (Van Name) (TP or TX)

Virginia.— Alleghany Co.: Lowmoor Cave. Giles Co.: Smokehole
Cave. Rockbridge Co.: Buck Hill Cave.

Miktoniscus racovitzai racovitzai Vandel (TB)

Virginia. — Alleghany Co.: Island Ford and Lowmoor caves.
Botetourt Co.: Peery Saltpetre Cave. Page Co.: Luray Caverns
(type locality). Rockbridge Co.: Buck Hill Cave. Shenandoah
Co.: Shenandoah Caverns.

Miktoniscus spp.

Tennessee. — Campbell Co.: Norris Dam Cave. Sullivan Co.: Bristol
Caverns.

Virginia. — Washington Co.: Hall Bottom Cave No. 1.

Comments. — These populations may be referable to M. medcofi
after further study.

Trichoniscus pusillus Brandt (TX)

Virginia. — Augusta Co.: Staunton Quarry Cave (?). Lee Co.: Cudjos
Cavern. Tazewell Co.: Wagoners Cave. Washington Co.: Hall
Bottom Cave No. 1.

Order Decapoda

The only decapod crustaceans recorded from caves in Virginia and
eastern Tennessee are crayfishes of the family Astacidae. Two species of
the genus Cambarus are known: C. bartonii (s. lat.) and C. dubius (see
also Hobbs et al. 1977, Holthuis 1986). The former is common throughout
much of the eastern United States (see Hobbs 1972, 1974) and is often
found in caves of the Appalachian region, where it is probably a
troglophile. The latter, recorded only once from a cave in the study area,
is apparently an accidental, inasmuch as it is normally found in burrows
and not caves (Hobbs 1974). In caves C. bartonii is usually found in
streams or stream pools. Individuals or whole populations may sometimes
be quite pale, probably reflecting ecophenotypic rather than genetic
changes (see Hobbs and Barr 1960). In addition to the caves listed
below, there are many sight records for C. bartonii , especially from
caves in the Powell and Clinch valleys where relatively large populations

John R. Holsinger and David C. Culver

were sometimes observed in streams. According to Hobbs (1972, 1974)
at least two subspecies inhabit caves of the study area: C. b. cavatus
Hay in the upper Tennessee River drainage of southwestern Virginia
and eastern Tennessee, and C. b. bartonii (Fabricius) elsewhere. The
systematic status of C. b. cavatus , however, is unclear and in need of
further evaluation (H. H. Hobbs, Jr., pers. comm.).

Family Astacidae

Cambarus ( Cambarus ) bartonii (s. lat.) (Fabricius) (TP)

Tennessee. — Hancock Co.: Cantwell Valley and Fairmont School
caves.

Virginia. — Alleghany Co.: Paxtons and Wares caves. Augusta Co.:
Barterbrook Spring Cave. Bath Co.: Roy Lyle Cave. Highland
Co.: Aqua Cave. Lee Co.: Crouse Cave. Rockbridge Co.: Billy
Williams Cave. Russell Co.: Quillens Field Cave. Scott Co.:
Johnson, McDavids, Riggs Chapel, and Wolfe caves. Smyth Co.:
Atwells Tunnel Cave. Tazewell Co.: Fallen Rock, Stonely, and
Wagoners Cave. Warren Co.: Skyline Caverns. Washington Co.:
Hall Bottom Cave No. 1.

Cambarus ( Jugicambarus ) dubius Faxon (AC)

Virginia.— Russell Co.: Jessie Cave.

Cambarus sp.

Virginia. — Montgomery Co.: Fred Bull Cave. Tazewell Co.: Steeles
Cave.

PHYLUM ARTHROPODA: SUBPHYLUM CHELICERATA

All cavernicolous chelicerates are in the class Arachnida, and in the
study area they include pseudoscorpions, acarines (mites and ticks),
harvestmen, and spiders. A considerable number of arachnids are
troglobites, especially spiders and pseudoscorpions.

Subclass Pseudoscorpiones

Although represented by a significant number of species,
pseudoscorpions are generally very rare in a given cave, and a
majority of the species are known only from a few individuals.
Cavernicolous pseudoscorpions are usually found in damp places,
frequently under rocks or small pieces of wood. In caves of Virginia and
eastern Tennessee, they are represented by four families, six genera, and
15 described species. Two species are provisionally recognized but
remain undescribed to date. Most species are troglobitic.

The family Chthoniidae contains the majority of cave species, and
all of these are troglobites in the study area. Kleptochthonius (subgenus
Chamberlinochthonius ) includes 10 species (2 undescribed) (see Malcolm

Invertebrate Cave Fauna

and Chamberlin 1961; Muchmore 1970b, 1974, 1976a), all of which are
rare, extremely localized endemics (Fig. 15) that are morphologically
strongly modified for cave existence (Fig. 19E). Only one species, K.
affinis, has been recorded from more than a single cave to date. All but
two species of Kleptochthonius from the study area occur in caves of
the upper Tennessee drainage in southwestern Virginia and eastern
Tennessee. Five species from this area (viz., K. affinis , K. binoculatus,
_ K . gertschi, K. proximosetus, and K. regulus ) are very closely allied
morphologically and were placed in a proximosetus group by Muchmore
(1976a). Other chthoniids include two species of Apochthonius (see
Muchmore 1963, 1967) and one species of Mundochthonius (see Benedict
and Malcolm 1974), all highly localized in distribution (Fig. 15).

The families Neobisiidae and Syarinidae also include troglobites —
Microcreagris valentinei in the former and Chitrella superba in the
latter. Both species are known only from single caves (Fig. 15) and are
quite rare (Chamberlin 1962, Muchmore 1973). Chitrella cavicola , a
troglophile or trogloxene recorded from Endless Caverns in Rockingham
County, is also reported from several epigean localities in northern
Virginia and a cave in Berkeley County, W.Va. (Muchmore 1973, Holsinger
et al. 1976).

The family Chernetidae is represented by a single species,
Hesperochernes mirabilis (formerly in Pseudozaona, see Muchmore

WEST VIRGINIA

! Apochthonius

1- A. coecus

2- A^ holsinqeri
Chitrella

3- C. superba
Kleptochthonius

4- K. affinis

5- K. anophthalmus

6 - K. binoculatus

7- K- gertschi

8- K. lutzi

9- K. proximosetus

10- K_. regulus

11- tC similis

12- tC sp. A

13- (C sp. B

14- (C spp.
Microcreagris

15- M. valentinei
Mundochthonius

16- M. holsinqeri

KENTUCKY

VIRGINIA

0 25 50 km

0 25 50 miles

Fig. 15. Distribution of troglobitic pseudoscorpions ( Apochthonius , Chitrella ,
Kleptochthonius , Microcreagris , and Mundochthonius) in the study area. Two
symbols in a circle indicate two species from the same cave.

John R. Holsinger and David C. Culver

1981), reported from a cave at Pennington Gap (possibly Gilley Cave).
This species is also recorded from several caves in south-central Kentucky
(Hoff 1958, Barr 1967a), and, although never recorded outside caves,
it is probably not a troglobite (see Chamberlin and Malcolm 1960, Barr
1967a).

Family Chernetidae

Hesperochernes mirabilis (Banks) (TP?)

Virginia. — Lee Co.: “Cave at Pennington Gap” (Banks 1895:4).
Hesperochernes spp.

Virginia. — Bath Co.: Cave Run Pit Cave. Giles Co.: Smokehole
Cave. Highland Co.: Van Devanters Cave.

Family Chthoniidae
Apoehthonius coecus (Packard) (TB)

Virginia. — Augusta Co.: Grand Caverns (type locality) and Madisons
Saltpetre Cave.

Apoehthonius holsingeri Muchmore (TB)

Virginia. — Alleghany Co.: Blue Springs Cave (?). Bath Co.: Cave
Run Pit Cave (type locality).

Comments.— The single specimen from Blue Springs Cave is a
tritonymph, and determination is tentative pending further study
(Muchmore 1976b).

Apoehthonius sp.

Virginia. — Giles Co.: Harris Cave.

Kleptochthonius {Chamber lino chthonius) affinis Muchmore (TB)
Tennessee. — Claiborne Co.: Chadwells (type locality), English and
Jennings caves.

Kleptochthonius (C.) anophthalmus Muchmore (TB)

Virginia. — Bath Co.: Porters Cave (type locality).

Kleptochthonius (C.) binoculatus Muchmore (TB)

Virginia. — Scott Co.: Hill Cave (type locality).

Kleptochthonius (C.) gertschi Malcolm and Chamberlin (TB)

Virginia. — Lee Co.: Gilley Cave (type locality).

Kleptochthonius (C.) lutzi Malcolm and Chamberlin (TB)

Virginia. — Lee Co.: Cudjos Cavern (type locality).

Kleptochthonius (C.) proximosetus Muchmore (TB)

Virginia. — Lee Co.: Gallohan Cave No. 1 (type locality).
Kleptochthonius (C.) regulus Muchmore (TB)

Virginia. — Tazewell Co.: Fallen Rock Cave (type locality).
Kleptochthonius (C.) similis Muchmore (TB)

Virginia. — Lee Co.: Sweet Potato Cave (type locality).
Kleptochthonius (C.) sp. A (TB)

Tennessee. — Hancock Co.: Panther Creek Cave.

Comments. — This population represents an undescribed species
(W. B. Muchmore, in litt.).

Invertebrate Cave Fauna

Kleptochthonius (C.) sp. B (TB)

Virginia. — Augusta Co.: Madisons Saltpetre Cave.

Comments. — This population represents an undescribed species
(see Muchmore 1970b).

Kleptochthonius (C.) spp.

Virginia. — Lee Co.: Elys Moonshine and Molly Wagle caves.
Comments. — Specimens from these caves are juveniles (deutonymphs
or tritonymphs), therefore precluding specific determination.
Mundochthonius holsingeri Benedict and Malcolm (TB)

Virginia. — Shenandoah Co.: Helsley Cave (type locality).

Family Neobisiidae

Microcreagris valentinei Chamberlin (TB)

Virginia. — Lee Co.: Cudjos Cavern (type locality).

Family Syarinidae

Chitrella cavicola (Packard) (TP or TX)

Virginia.— Rockingham Co.: Endless Caverns (type locality).
Chitrella superba Muchmore (TB)

Virginia. — Shenandoah Co.: Maddens Cave (type locality).

Subclass Acari

Although ticks (Ixodida) are occasionally transported into caves by
bats or pack rats, most cavernicolous acarines are mites. Several families
of mites have been recorded from caves in Virginia and east Tennessee,
including Laelapidae and Parasitidae in the order Parasitiformes and
Eupodidae and Rhagidiidae in the order Acariformes (see Holsinger
1965a). Aside from Rhagidiidae, however, the taxonomy and ecology of
cave-associated mites is very poorly known. Rhagidiid mites are relatively
common in caves, and to date four genera and five species have been
recorded from the study area (Fig. 16). In caves these mites are usually
found in mesic areas beneath rocks or decomposing organic detritus.
The family is primarily edaphic; although many species are reported
from caves in the Northern Hemisphere, only a few appear to be bona
fide troglobites (Zacharda 1980). The possession of troglomorphisms
and restriction to caves are criteria used by Zacharda (1980) and
Zacharda and Elliott (1981) to distinguish troglobites from troglophiles
and trogloxenes. In the Virginia-east Tennessee cave-mite fauna, two
species are possibly troglobitic, whereas three are probably troglophilic.

The most common cave mite in Virginia is Robustocheles hilli (Fig.
19c), an apparent troglophile, which is also recorded from many caves
in the eastern and western United States (Zacharda 1985) and from
epigean habitats in Alaska and northern Canada (Zacharda 1980,
Zacharda and Elliott 1981). Poecilophysis weyerensis (formerly Rhagidia
weyerensis ) was originally described from Grand Caverns by Packard

John R. Holsinger and David C. Culver

(1888) and thought to be a troglobite. It was redescribed by Holsinger
(1965b) and subsequently reported from caves in Missouri, New Mexico,
and Mexico by Elliott and Strandtmann (1971). Zacharda (1980) listed
this species from Long Cave in Edmonson County, Ky. (synonym =
Rhagidia cavernarum ) and from epigean localities (scree and moist
ground litter) in Czechoslovakia, but questioned the records given by
Elliott and Strandtmann (1971). Zacharda (1985) gave additional cave
records for this species in the study area (see list below) and also
recorded it from a cave in Monroe County, Tenn.

Three other species of rhagidiid mites have been identified from
caves in the Virginia-east Tennessee area by Zacharda (1985). Two of
these, Foveacheles paralleloseta and Rhagidia varia , are probably
troglobites since they possess some troglomorphisms and are known
only from caves. The third species, Poecilophysis extraneostella , although
known only from caves at present, is not troglomorphic and is probably
a troglophile that eventually will be found outside caves.

Linopodes sp. (possibly motatorius\ see Holsinger 1965a), a member
of the family Eupodidae, was noted occasionally in study-area caves,
but specimens were not collected.

Order Parasitiformes
Suborder Ixodida

Family Ixodidae
Ixodes cookei Pakcard (AC)

Virginia.— Giles Co.: Harris Cave.

Suborder Gamasida

Family Laelapidae
Androlaelaps sp.

Virginia. — Shenandoah Co.: Shenandoah Wild Cave.

Hypoaspis sp.

Virginia.— Shenandoah Co.: Shenandoah Wild Cave.

Family Parasitidae
Eugamasus sp.

Virginia.— Russell Co.: Banners Corner Cave.

Pergamasus sp.

Virginia. — Russell Co.: Banners Corner Cave.

Unidentified gamasid mites are recorded as follows:

Tennessee. — Anderson Co.: Hill Cave. Union Co.: Lost Creek
Cave.

Virginia. — Rockbridge Co.: Showalters Cave. Tazewell Co.: Fallen
Rock and Wagoners caves.

Invertebrate Cave Fauna

■ Foveacheles paralleloseta

♦ Poecilophysis weyerensis
□ P extraneostella

▼ Rhagidia varia

• Robustocheles hilli

WEST VIRGINIA

KENTUCKY

VIRGINIA

Fig. 16. Distribution of cavernicolous mites ( Foveacheles , Poecilophysis ,
Rhagidia , and Robustocheles) in the study area. Two symbols in a circle
indicate two species from the same cave.

50 km

50 miles

Order Acariformes

Family Rhagidiidae

Foveacheles paralleloseta Zacharda (TB?)

Virginia. — Wythe Co.: Sam Six Cave.

Poecilophysis extraneostella Zacharda (TP)

Virginia. — Lee Co.: Bowling Cave.

Comments. — Also recorded from Steeles Cave, Monroe County,
W.Va.

Poecilophysis weyerensis (Packard) (TP)

Tennessee. — Hawkins Co.: Sensabaugh Saltpetre Cave.

Virginia. — Augusta Co.: Grand Caverns (type locality). Rockbridge
Co.: Buck Hill Cave.

Rhagidia varia Zacharda (TB?)

Virginia. — Bath Co.: Butler-Sinking Creek Cave. Pulaski Co.: Sam
Bells Cave. Scott Co.: Hill Cave.

Comments. — Also recorded from single caves in Greenbrier and
Pocahontas counties, W.Va.

Robustocheles hilli (Strandtmann) (TP)

Tennessee. — Hawkins Co.: Pearson Cave.

John R. Holsinger and David C. Culver

Virginia.— Alleghany Co.: Rumbolds Cave. Augusta Co.: Madisons
Saltpetre Cave. Bland Co.: Banes Spring, Hamilton, and Repass
Saltpetre caves. Craig Co.: Loneys Cave. Giles Co.: New River
and Straleys No. 1 caves, Highland Co.: Aqua Cave. Pulaski Co.:
Sam Bells Cave. Roanoke Co.: Goodwins Cave. Rockingham
Co.: Endless Caverns. Scott Co.: Lane and Moccasin Valley
caves. Tazewell Co.: Crocketts, Fallen Rock, Gillespie Water,
and Gully caves. Wise Co.: Kelly Cave.

Undetermined rhagidiid mites:

Tennessee. — Claiborne Co.: Tazewell Saltpetre Cave.

Virginia. — Augusta Co.: Staunton Quarry Cave. Lee Co.: Waltons
Cave. Montgomery Co.: Vickers Road Cave. Scott Co.: Greears
Sweet Potato and Taylor No. 1 caves. Smyth Co.: Tilson Saltpetre
Cave.

Subclass Opiliones

Except for Leiobunum (Phalangiidae), a sporadically common

threshold trogloxene, opilionids (also called phalangids or harvestmen)

are not common in the caves of Virginia and east Tennessee. Only a few

species are recorded, and none is a troglobite. Probably the most

interesting species with respect to cave association is Erebomaster

acanthina (Erebomastridae), a troglophile found in several caves in the

Shenandoah Valley (Fig. 19B). This species is also recorded from caves

in Maryland (under Phalangodes acanthina by Franz and Slifer 1971)

and West Virginia (under Phalangodes flavescens weyerensis by Holsinger

et al. 1976), and from epigean localities in the Piedmont and Blue Ridge

Mountains of North Carolina and Virginia (see Goodnight and Goodnight
1942, Briggs 1969). Packard (1888) described Phalangodes flavescens
var. weyerensis from Grand Caverns, and Hadzi (1935) later described
Cladonychium corii from Endless Caverns. Both the “variety” weyerensis
and C. corii are now considered synonyms of Erebomaster acanthina ,
which was redescribed in detail by Briggs (1969, in litt.).

Erebomaster acanthina may be a troglobite in statu nascendi ,
inasmuch as several populations appear to be cave limited and consist
of individuals with reduced eyes and pigment. Both adults and juveniles
of this species have been observed many times on damp, rotting wood in
Endless Caverns and Madisons Saltpetre Cave.

The range of Phalangodes laciniosa (Phalangodidae), also a
troglophile, extends to the southern end of the study area, where it has
been found once in Norris Dam Cave in Campbell County. To the
south and southwest this species is recorded from caves in northern
Alabama, northern Florida, northwestern Georgia, and other parts of
Tennessee, and occasionally from epigean localities in the same general
region (Goodnight and Goodnight 1960, Barr 1961, Peck 1970, Holsinger
and Peck 1971).

Invertebrate Cave Fauna

Crosbycus goodnighti (Nemastomatidae) is reported from Fountain
Cave in Augusta County. According to W. A. Shear (in litt.), the
description and figure of this species by Roewer (1951) apply to the
juvenile of a European nemastomatid, and the identity of this taxon is
therefore questionable.

Family Erebomastridae

Erebomaster acanthina (Crosby and Bishop) (TP)

Virginia. — Augusta Co.: Fountain, Grand (Caverns), and Madisons
Saltpetre caves. Frederick Co.: Ogdens Cave. Rockingham Co.:
Endless Caverns.

Erebomaster (?) spp.

Virginia. — Alleghany Co.: Paxtons Cave. Bath Co.: Roy Lyle
Cave. Bland Co.: Newberry-Bane Cave.

Family Nemastomatidae

Crosbycus (?) goodnighti (?) Roewer
Virginia. — Augusta Co.: Fountain Cave.

Family Phalangiidae
Leiobunum bicolor (?) (Wood) (TX)

Comments. — One or more species of Leiobunum were seen
sporadically in caves of the study area, but no attempt was made
to collect them systematically.

Family Phalangodidae

Phalangodes ( Bishopella ) laciniosa Crosby and Bishop (TP)

Tennessee. — Campbell Co.: Norris Dam Cave.

Subclass Araneae

The cave spider fauna of Virginia and east Tennessee is quite
diverse and comprises 13 families, 31 genera, and approximately 38
species. However, about one-third of the species recorded are accidentals
or only marginal trogloxenes and contribute very little to the fauna of
most caves. The remaining species are divided roughly equally between
trogloxenes/ trogophiles and troglobites. As noted in the list below,
most of the trogloxenic and trogophilic spiders associated with caves in
the study area are also recorded from caves in other parts of the eastern
and southeastern United States as well as from epigean localities. Many
species are associated with ground litter and similar habitats at the
surface. In caves, spiders occupy a number of microhabitats and are
often found around decomposing wood, in the damp recesses of passage
walls and ceilings, and sometimes beneath rocks near the banks of
streams.

Troglobitic spiders in the study area, as well as in most of the
eastern United States, belong to the families Linyphiidae and Nesticidae.
Five species of linyphiids have been recorded, all of which are presumably

John R. Holsinger and David C. Culver

troglobitic but have wide ranges (Fig. 17) and occur outside of the
Virginia-Tennessee cave region. Anthrobia monmouthia has been found
in several Virginia caves and is also recorded from caves in south-
central Kentucky, middle Tennessee, and southern West Virginia (Barr
1961, 1967a; Holsinger et al. 1976). The Appalachian Valley populations
of Virginia and West Virginia may represent one or more subspecies
and are in need of further study. In the study area Bathyphantes weyeri
is known only from Grand Caverns but is also recorded from caves in
Arkansas, Kentucky, Pennsylvania, West Virginia, and Wisconsin (Ivie
1969). According to W. J. Gertsch (unpublished data) this species was
collected once from an epigean habitat. Islandiana muma, an extremely
rare and poorly known species, was described from Buck Hill Cave in
Rockbridge County but is also reported from a single cave in Colbert
County, Ala. (Ivie 1965).

The most common, widespread linyphiid spider in Virginia and
Tennessee is Phanetta subterranea. It is also common in caves through-
out the eastern United States and ranges from Pennsylvania south to
Georgia and Alabama and west to Illinois and Missouri. Porrhomma
cavernicolum is also widespread in caves of the study area but is
generally not as common as P. subterranea. It is recorded from caves
throughout much of the central and eastern United States and ranges
from Pennsylvania south to Georgia and west to Missouri and
Oklahoma. Linyphiids have the widest ranges of any troglobites in
North America, leading to the speculation that these are morphological
species, each representing several separate gene pools (Barr 1967a,
Holsinger et al. 1976).

With the exception of Eidmannella pallida (formerly Nesticus
pallidus ), a troglophile or trogloxene relatively common in caves
throughout much of the United States and Mexico, cavernicolous
spiders of the family Nesticidae in the study area have relatively
restricted distributions (Fig. 18). Those that are not troglobites are
represented by populations primarily limited to caves. The most common
and widespread nesticid in the Appalachian region is Nesticus carteri, a
troglophile that is sometimes quite abundant in caves of the Powell
Valley (Fig. 19D). It is recorded from numerous caves and a few
epigean localities (ground detritus) in southern Indiana, eastern Kentucky,
eastern Tennessee, southwestern Virginia, and southeastern West Virginia
(Gertsch 1984). In or near the study area this species has been collected
from ground detritus in Dickenson County, Va., and Mercer County,
W.Va. The type locality of N. carteri is Bat Cave (Carter Caves State
Park). Carter County, Ky., and not Mammoth Cave as erroneously
reported by Nicholas (1960) and Holsinger (1963a).

The other species of Nesticus from the study area have more closely
circumscribed ranges and are largely restricted to areas east of the

Invertebrate Cave Fauna

>° -V

a Anthrobia monmouthia

* Bathyphantes weyeri
▼ Islandiana muma

• Phanetta subterranea

□ Porrhomma cavernicolum

WEST VIRGINIA

KENTUCKY

f VIRGINIA

25 50 km

25 50 miles

NORTH CAROLINA

Fig. 17. Distribution of troglobitic spiders ( Anthrobia , Bathyphantes ,
Islandiana , Phanetta , and Porrhomma) in the study area. Two or more symbols
in a circle indicate two or more species from the same cave.

• Nesticus carteri

♦ N. tennesseensis
0 N. holsingeri

□ N. mimus

* N. paynei

* Nesticus spp.

WEST VIRGINIA

KENTUCKY

VIRGINIA

50 km

50 miles

Fig. 18. Distribution of cavernicolous spiders ( Nesticus ) in the study area.
Single cave localities for N. carteri in Greenbrier County, W.Va., and N.
tennesseensis in Grainger County, Tenn., also shown. Epigean localities for N.
carteri and N. tennesseensis not shown.

John R. Holsinger and David C. Culver

Appalachian Plateau. These four species are closely allied morphologi-
cally and are assigned to the tennesseensis group by Gertsch (1984). The
most widespread member of this suite is N. tennesseensis, a probable
troglobite with variation in both eye-pigment reduction and elongation
of the legs. Its range extends from Highland County, Va., southwestward
to Roane County, Tenn. In addition to the localities listed below, it is
recorded from single caves in Grainger and Roane counties, Tenn., and
from ground detritus (epigean) at single localities in Raleigh County,
W.Va., and Giles and Highland counties, Va. (Gertsch 1984). The
remaining three species were recently described by Gertsch (1984) and
inhabit caves in the upper Tennessee drainage in southwestern Virginia
and eastern Tennessee. Nesticus holsingeri , a probable troglobite with
reduced eyes, is known only from caves in Lee and Scott counties.
Nesticus mimus is recorded from two caves in Washington County and
also from epigean habitats at higher elevations in Burke and Watauga
counties in nearby North Carolina. Nesticus paynei has fully developed
eyes but is at present unknown outside caves. This species is also
recorded from caves just south of the study area in Anderson, Carter,
and Knox counties, Tenn.

Probably the most conspicuous cave spider in the Virginia-east
Tennessee area is the orb weaver Meta menardi (Argiopidae), a
trogloxene or troglophile frequently seen near cave entrances. This
species is widespread in caves of the eastern United States and is also
found in western Europe. Sight records are numerous from the study
area, but no attempt was made to collect it systematically.

Family Agelenidae

Calymmaria cavicola (Banks) (TP or TX)

Virginia. — Tazewell Co.: Steeles Cave.

Comments. — Also recorded from caves in Alabama, Georgia,
Illinois, Kentucky, Tennessee, and West Virginia (Barr 1967a,
Holsinger and Peck 1971, Holsinger et al. 1976, Peck and Lewis
1978); widespread in epigean localities.

Cicurina pallida Keyserling (TP or TX)

Virginia. — Augusta Co.: Fountain, Glade, and Madisons Saltpetre
caves. Shenandoah Co.: Hensleys Cave.

Comments. — Also recorded from caves in Illinois and West Virginia
(Holsinger et al. 1976, Peck and Lewis 1978); widespread in the
eastern United States.

Circurina sp.

Virginia. — Rockbridge Co.: Tolleys Cave.

Family Anyphaenidae

Anyphaena sp. (AC)

Virginia. — Roanoke Co.: “Old Hollins Road Cave” (L. M.
Ferguson, in litt.).

Invertebrate Cave Fauna

Fig. 19. Terrestrial cavernicoles from the study area (approximate body lengths
in parentheses): A, salamander (adult), Gyrinophilus porphyriticus (13 cm); B,
opilionid, Erebomaster acathina (3 mm); C, mite, Robustocheles hilli ( 1 mm);
D, spider, Nesticus carteri (4 mm); E, pseudoscorpion, Kleptochthonius sp. (2
mm).

John R. Holsinger and David C. Culver

Aysha sp. (AC)

Virginia. — Roanoke Co.: Newmans Cave.

Family Argiopidae
Araniella displicata (Hentz) (AC)

Virginia. — Roanoke Co.: Newmans Cave.

Leucauge venusta Walckenaer (AC)

Virginia. — Craig Co.: Rufe Caldwell Cave.

Mangora placida Hentz (AC)

Virginia. — Rockbridge Co.: Bell Cave.

Meta menardi (Latreille) (TP or TX)

Comments. — Common throughout the study area; specific records
not documented.

Family Clubionidae

Liocranoides unicolor Keyserling (TP or TX)

Virginia. — Washington Co.: Hall Bottom Cave No. 1.

Comments. — Common in caves in northern Alabama, northwestern
Georgia, and south-central Tennessee (Barr 1961, Holsinger and
Peck 1971).

Liocranoides sp. (TX?)

Virginia. — Washington Co.: Neals Cave.

Comments. — This population probably represents an undescribed
species (W. J. Gertsch, in litt.).

Family Ctenizidae
Antrodiaetus unicolor (Hentz) (AC)

Virginia. — Tazewell Co.: Hugh Young Cave.

Family Dictynidae
Lathys sp. (AC)

Virginia. — Tazewell Co.: Cassell Farm Cave(s).

Family Linyphiidae

Anthrobia monmouthia Tellkampf (TB)

Virginia. — Alleghany Co.: Wares Cave. Bath Co.: Clarks Cave.
Scott Co.: Harris Pit Cave. Smyth Co.: Buchanan Saltpetre
Cave.

Bathyphantes ( Bathyphantes ) albiventris (Banks) (TX or AC)

Virginia. — Lee Co.: Bowling Cave.

Comments. — Reported from epigean localities in the eastern and
northeastern United States (Ivie 1969) and from one cave in
Illinois (Peck and Lewis 1978).

Bathyphantes {Weyerphantes) weyeri (Emerton) (TB?)

Virginia. — Augusta Co.: Grand Caverns (type locality).

Centromerus cornupalpis (Pickard-Cambridge) (TX?)

Virginia. — Montgomery Co.: Erharts Cave.

Invertebrate Cave Fauna

Comments. — Also recorded from caves in Illinois and Missouri (see
Peck and Lewis 1978).

Centromerus latidens (Emerton) (TP or TX)

Virginia. — Lee Co.: Sweet Potato Cave. Shenandoah Co.: Shenan-
doah Wild Cave.

Comments. — Widespread in the central and eastern United States;
recorded from caves in Florida, Illinois, Kentucky, Missouri,
Oklahoma, and possibly Texas (Reddell 1965, Barr 1967a, Peck
1970, Black 1971, Craig 1977, Peck and Lewis 1978).

Centromerus spp.

Tennessee. — Sullivan Co.: Bristol Caverns.

Virginia. — Alleghany Co.: Second Dam and Wares caves. Tazewell
Co.: Lost Mill No. 1 and Steeles caves.

Frontinella communis Hentz (AC)

Virginia. — Rockbridge Co.: Showalters Cave.

Islandiana muma Ivie (TB)

Virginia. — Rockbridge Co.: Buck Hill Cave (type locality)

Islandiana (?) sp.

Virginia. — Montgomery Co.: Vickers Road Cave.

Linyphia marginal a Koch (TX?)

Virginia. — Roanoke Co.: “Old Hollins Road Cave” (L. M.
Ferguson, in litt.).

Comments. — Also recorded from a few caves in Missouri, Okla-
homa, and West Virginia (Black 1971, Holsinger et al. 1976,
Craig 1977).

Meioneta sp. (TX or AC)

Tennessee. — Claiborne Co.: Lower Coonsies Creek Cave.

Microneta sp. (TX or AC)

Virginia. — Alleghany Co.: Wares Cave.

Oreonetides sp. (AC)

Virginia. — Tazewell Co.: Rosenbaums Water Cave.

Phanetta subterranea (Emerton) (TB)

Tennessee. — Campbell Co.: Meredith and Norris Dam caves. Clai-
borne Co.: English and Keck No. 1 caves. Hancock Co.: Cantwell
Valley Cave. Sullivan Co.: Bristol Caverns and Morrill Cave.

Virginia. — Alleghany Co.: Blue Springs, Island Ford, Lowmoor,
and Wares caves. Augusta Co.: Fountain and Madisons Saltpetre
caves. Bath Co.: Boundless, Breathing, Butler-Sinking Creek,
Cave Run Pit, Clarks, Dunns, and Starr Chapel caves. Bland
Co.: Hamilton Cave. Botetourt Co.: Peery Saltpetre and Thomas
caves. Craig Co.: Rufe Caldwell Cave. Frederick Co.: Ogdens
Cave. Giles Co.: Clover Hollow, Harris, New River, Smokehole,
Starnes, Straleys No. 1, and Tawneys caves. Lee Co.: Bowling,
Cudjos (Cavern), Cumberland Gap Saltpetre, Gibson-Frazier,

John R. Holsinger and David C. Culver

Lucy Beatty, Molly Wagle, Olinger and Spangler caves.
Montgomery Co.: Slussers Chapel Cave. Page Co.: Luray
Caverns. Pulaski Co.: Sam Bells Cave. Roanoke Co.: Goodwins,
Hodges No. 1, and Millers Cove caves. Rockingham Co.: Deer
Hole, Massanutten (Caverns), and Stephens caves. Russell Co.:
Banners Corner, Jessie, and Porgie Bundys caves. Scott Co.:
Grigsby, Herron No. 1, Hill, and Kerns No. 1 caves. Smyth Co.:
Beaver Creek Cave. Tazewell Co.: Cassell Farm, Lawson, and
Steeles caves. Warren Co.: Skyline Caverns. Washington Co.:
Hall Bottom No. 1 and Perkins caves. Wise Co.: Kelly and
Wildcat Saltpetre caves. Wythe Co.: Picketts Cave.

Porrhomma cavernicolum (Keyserling) (TB)

Tennessee. — Claiborne Co.: Jennings Cave. Hawkins Co.: Sensa-
baugh Saltpetre Cave.

Virginia. — Augusta Co.: Fountain, Glade, and Madisons Saltpetre
caves. Bath Co.: Clarks, Crossroads, Porters, and Witheros
caves. Bland Co.: Coon and Banes Spring caves. Craig Co.: New
Castle Murder Hole and Rufe Caldwell caves. Frederick Co.:
Beans Cave. Giles Co.: Clover Hollow Cave. Lee Co.: Fisher and
Unthanks caves. Page Co.: Luray Caverns and Ruffners Cave
No. 1. Roanoke Co.: Dixie Caverns. Rockbridge Co.: Bell and
Buck Hill caves. Rockingham Co.: Three-D Maze Cave. Smyth
Co.: Buchanan Saltpetre Cave. Tazewell Co.: Gully and Lawson
caves. Wise Co.: Parsons Cave.

Sciastes sp. (AC)

Virginia. — Lee Co.: Young-Fugate Cave.

Family Lycosidae
Lycosa rabida Walckenaer (AC)

Virginia. — Giles Co.: Tawneys Cave.

Comments. — Also reported from single caves in Oklahoma (Black
1971) and Texas (Reddell 1965).

Pirata sp. (AC)

Virginia. — Lee Co.: Bowling Cave.

Family Nesticidae

Eidmannella pallida (Emerton) (TP or TX)

Virginia. — Alleghany Co.: Walking Cave. Augusta Co.: Fountain,
Glade and Grand (Caverns) caves. Giles Co.: New River Cave.
Lee Co.: Cattle, Gallohan No. 1, Glen Olingers, and Smiths Milk
caves. Page Co.: Luray Caverns and Ruffners Cave No. 1.
Rockbridge Co.: Tolleys Cave. Rockingham Co.: Massanutten
Caverns and Steam Hole Cave. Scott Co.: Ellington and Harris
Pit caves. Washington Co.: Hall Bottom No. 1 and Perkins

caves.

Invertebrate Cave Fauna

Nesticus carteri Emerton (TP)

Tennessee. — Claiborne Co.: Chadwells, English, Keck No. 1, and
Tom Balls caves. Hancock Co.: Subers Cave. Sullivan Co.:
Bristol Caverns. Union Co.: Lost Creek and Oaks caves.

Virginia. — Lee Co.: Cope, Cudjos (Cavern), Frazier, Gilliam, Kinzer
Hollow, McClure, Molly Wagle, Roadside No. 1, Sheep, Skull,
Skylight, Sweet Potato, Taylor Pit, and Thompson caves. Rock-
bridge Co.: Buck Hill and Doll House caves. Scott Co.: Blowing
Hole, Greears Sweet Potato, and Kerns No. 1 caves. Smyth Co.:
Atwells Tunnel and Stones No. 2 caves. Tazewell Co.: Quarry
and Wagoners caves.

Nesticus holsingeri Gertsch (TB)

Virginia. — Lee Co.: Bowling and Gibson No. 1 caves. Scott Co.:
Alley, Blair-Collins, Coley No. 2, Jackson, McDavids, Pond
(type locality), and Taylor No. 1 caves. Wise Co.: Burtons Cave.

Nesticus mimus Gertsch (TP)

Virginia. — Washington Co.: Fritz Breathing and Shiloh School
(type locality) caves.

Nesticus paynei Gertsch (TB?)

Tennessee. — Campbell Co.: “Hammers” (Gertsch 1984:28) and
Norris Dam caves. Hancock Co.: Cantwell Valley Cave. Sullivan
Co.: Morrills Cave. Union Co.: Coppock and Ridenour Pit
caves.

Virginia. — Scott Co.: Wolfe Cave.

Nesticus tennesseensis (Petrunkevitch) (TB?)

Tennessee. — Hawkins Co.: Sensabaugh Saltpetre Cave. Sullivan
Co.: Potters Cave.

Virginia. — Alleghany Co.: Rumbolds Cave. Craig Co.: Fish Hatchery
and Walkthrough caves. Giles Co.: Ballards, Giant (Caverns),
Glenlyn, Harris, Starnes, and Straleys No. 1 caves. Smyth Co.:
Sugar Grove Cave No. 10. Tazewell Co.: Cassell Farm, Chimney
Rock, Fallen Rock, Hugh Young, and Steeles caves.

Nesticus spp.

Tennessee. — Campbell Co.: Easterly Cave. Claiborne Co.: John
Lard and Lower Coonsies Creek caves. Hancock Co.: Fairmont
School and Lawsons No. 3 caves.

Virginia. — Alleghany Co.: Island Ford Cave. Lee Co.: Ely, Fisher,
Indian, Spangler, and Young-Fugate caves. Pulaski Co.: Fifty-
Foot Hell Cave. Russell Co.: Smiths Cave. Scott Co.: Flannery
and Sparks caves. Tazewell Co.: Lost Mill Cave No. 1.

Comments. — Specimens from these caves are juveniles, therefore
precluding specific determination.

Family Pholcidae

Pholcus phalangioides Fuesslin (TX)

John R. Holsinger and David C. Culver

Virginia. — Page Co.: Luray Caverns.

Comments. — Also reported from caves in Tennessee (Barr 1961).

Family Tetragnathidae
Tetragnatha sp. (AC)

Virginia. — Smyth Co.: Stones Cave No. 2.

Family Theridiidae

Achaearanea tepidariorum (Kock) (TP or TX)

Virginia. — Augusta Co.: Barterbrook Spring, Fountain, Grand
(Caverns), and Madisons Saltpetre caves. Page Co.: Will Mauck
Cave. Rockbridge Co.: Bell and Showalters caves.

Comments.— Recorded from caves throughout a large part of the
southeastern and south-central United States (see Black 1971,
Franz and Slifer 1971, Holsinger and Peck 1971, Holsinger et al.
1976, Craig 1977, Peck and Lewis 1978).

Family Thomisidae
Misumenops celer Hentz (AC)

Virginia. — Bath Co.: Porters Cave.

PHYLUM ARTHROPODA: SUBPHYLUM UNIRAMIA
Among the uniramians, the classes Diplopoda (millipeds) and
Insecta are very well represented in the cave fauna of Virginia and east
Tennessee; many troglobites and troglophiles are recorded in each
group. Of significantly less importance in the regional cave fauna is the
class Chilopoda (centipedes), species of which are seldom found in
caves. Only one such species is a possible troglobite. Representatives of
the classes Pauropoda and Symphyla are extremely rare in caves, and
only a single cave record for each group is noted from the study area.
Both pauropods and symphylans are rare, cryptic organisms that live in
soil and leaf mold, and their occurrence in caves is probably accidental.

Class Pauropoda

Genus (?) species (?)

Virginia. — Roanoke Co.: McVitty Cave.

Class Symphyla

Scutigerella sp. (AC)

Virginia. — Lee Co.: Molly Wagle Cave.

Class Chilopoda

Cave records for centipedes are very sparse, and all species but one
are recorded from single caves and are probably accidentals. Nampabius
turbator (Lithobiidae), however, is recorded from two caves in Alleghany
County and possesses reduced eyes and pigment (see Crabill 1952).

Invertebrate Cave Fauna

Although to our knowledge this species has not been found outside
caves to date, its status as a troglobite is uncertain.

Order Geophilomorpha

Family Chilenophilidae
Arctogeophilus umbracticus (McNeill) (AC)

Virginia. — Scott Co.: Coley Cave No. 2. Shenandoah Co.: Pingleys
Cave.

Order Lithobiomorpha

Family Ethypoliidae

Bothropolys multidentatus (Newport) (AC)

Virginia. — Rockbridge Co.: Tolleys Cave.

Family Lithobiidae

Nampabius parienus Chamberlin (TX or AC)

Virginia. — Smyth Co.: Atwells Tunnel Cave.

Nampabius turbator Crabill (TB?)

Virginia. — Alleghany Co.: Island Ford and Lowmoor (type locality)
caves.

Nampabius sp.

Virginia. — Montgomery Co.: Erharts Cave.

Order Scolopendromorpha

Family Cryptopidae

Cryptops hortensis Leach (AC)

Virginia. — Lee Co.: Ruths Cave.

Crytops hyalinus Say (AC)

Virginia. — Tazewell Co. Gully Cave.

Scolopocryptops sexpinosus (Say) (AC)

Tennessee. — Claiborne Co.: English Cave.

Theatops posticus (Say) (AC)

Virginia. — Frederick Co.: Ogdens Cave.

Class Diplopoda

Millipeds are among the most common cavernicoles in Virginia and
east Tennessee and are well represented by a diverse taxonomic
assemblage consisting of 5 orders, 9 families, 12 genera, and 24 described
species. Probably about one-half of the species collected from caves are
undescribed at present. Approximately 25% of the species (including
both described and undescribed forms) are troglobites. Cavernicolous
millipeds are usually found in damp to wet areas associated with
decomposing organic matter (e.g., wood, guano, carcasses)

Clearly the most significant order with respect to the diversity of
cavernicolous species and their affinity for the cave environment is the
Chordeumatida. All of the trogolobitic millipeds in the study area are

John R. Holsinger and David C. Culver

included in this group, which is represented by the families Cleidogoni-
dae, Conotylidae, Striariidae, and Trichopetalidae. The most widespread
genus in caves of the study area is Pseudotremia (Cleidogonidae); it is
found in all major drainage basins except the Shenandoah (Fig. 20, 21).
In study-area caves the genus is represented by 12 described and
approximately 20 undescribed species. Two species, P. nodosa (Fig.
3 ID) and P. cavernarum, have greatly reduced eyes (ocelli) and are
either unpigmented or only lightly so. Both are clearly troglobitic (see
Loomis 1939, Hoffman 1958, Shear 1972). Three other species — P.
deprehendor, P. tuberculata, and P. valga — although known only from
caves at present, are generally pigmented, possess relatively well-
developed ocelli, and are questionable troglobites. The remaining species
(described) have been found in both cave and epigean habitats and are
apparently troglophiles.

Pseudotremia nodosa , originally described from English Cave in
Claiborne County, has been recorded from many caves in the Powell
Valley and, along with morphologically closely allied populations in the
adjacent Clinch Valley, may represent a complex of closely similar
(sibling ?) species (W. A. Shear, in litt.). This species, or complex, is the
most troglomorphic member of the genus in the Virginia-east Tennessee
area. Another species complex in the upper Tennessee basin is
represented by P. fr acta (s. lat.) and P. cocytus. Although Shear (1972)
described P. cottus from cave and epigean habitats in Anderson,
Blount, Knox, Roane, and Sevier counties, Tenn., Hoffman (1981)
pointed out that P.fracta is actually the objective senior synonym of the
species and therefore should take nomenclatural priority. Hoffman
(1981) further divided P. fracta into four subspecies: P. f fracta , P. f.
paynei, P. f ingens, and P. f nantahala. The records listed below for P.
fracta (s. lat.) are based on material determined by W. A. Shear as P.
cottus, but in light of Hoffman’s recent study, they probably should be
assigned to P. f. paynei.

Pseudotremia hobbsi is the most common species of the genus in
west-central Virginia, where it is recorded from a number of caves and a
few epigean localities in the upper James and Roanoke basins; it is also
found in southern West Virginia (Hoffman 1950, Shear 1972, Holsinger
et. al. 1976).

As indicated in the list below, many species of Pseudotremia
remain undescribed. In addition, numerous collections are undetermined,
primarily because they lack mature males. Further, detailed taxonomic
study of the genus is clearly needed to resolve species complexes and
elucidate distributional patterns.

Three troglobitic species of Trichopetalum (Trichopetalidae),
formerly assigned to Zygonopus by Causey (1960a) but reassigned to
the present genus by Shear (1972), occupy caves from the New River

Invertebrate Cave Fauna

♦ Pseudotremia cavemarum
▼ P. deprehendor

• R nodosa IsTTat.

A R tuberculata

* R valga

■ R spp. 1 nodosa complex

KENTUCKY

/IRGINIA

NORTH CAROLINA

25 50 km

25 50 miles

Fig. 20. Distribution of troglobitic millipeds ( Pseudotremia ) in the study area.
Single locality for P. deprehendor in Anderson County, Tenn., also shown. Two
symbols in a circle indicate two species from the same cave.

Pseudotremia armesi

* R cocytus
■ R tracta

♦ P hobbsi
▼ P momus
▲ P princeps

• R sublevis

KENTUCKY

VIRGINIA

0 25 50 km

0 25 50 miles

WEST VIRGINIA

Fig. 21. Distribution of troglophilic millipeds ( Pseudotremia ) in the study

area.

John R. Holsinger and David C. Culver

basin northeastward to the Shenandoah Valley (Fig. 22). The genus has
never been found in caves of the upper Tennessee basin. Troglobitic
species of Trichopetalum are usually much smaller than those of
Pseudotremia, and all individuals lack ocelli and pigment.

Trichopetalum whitei inhabits caves of the Shenandoah Valley and
is also recorded from caves in adjacent Grant and Pendleton counties,
W.Va. (see Holsinger et al. 1976). Trichopetalum weyeriensis ranges
generally south and west of T. whitei and is also recorded from caves in
Greenbrier, Monroe, Pendleton, and Pocahontas counties, W.Va. (see
Holsinger et al. 1976). Trichopetalum packardi occurs to the southwest
of T. weyeriensis and is common in caves of the New River drainage; it
is also recorded from caves in Greenbrier, Mercer, and Monroe counties,
W.Va. (Holsinger et al. 1976).

On the basis of collections made in the early 1960s, N. B. Causey
(in litt.) concluded that some populations of T. weyeriensis showed
evidence of intergradation with both T. packardi and T. whitei in
different parts of West Virginia. Based on these observations, Causey
(1963) suggested that the three species are subspecies of a single, rather
widespread species and not three distinct species as she had indicated
earlier (Causey 1960a). In our judgment, this situation is far from being
as clear-cut as Causey suggested and cannot be properly resolved until
all collections from the Virginia-West Virginia cave region (many of
which have been made since 1963) have been carefully examined and
analyzed in detail.

Other chordeumatids recorded from caves in the study area include
Conotyla venetia (Conotylidae), a possible trogloxene reported from
one cave and two epigean localities in Alleghany County (see Shear
1971); and one or more species of Striaria (Striariidae), of which S.
Columbiana is a possible trogloxene known primarily from epigean
habitats in northwestern Virginia, adjacent Maryland, and the District
of Columbia (Chamberlin and Hoffman 1958). One population of
Striaria from Madisons Saltpetre Cave in Augusta County appears to
be troglomorphic and may represent an undescribed species, but additional
study is needed to determine its status vis-a-vis S. columbiana and other
species in the genus (W. A. Shear, in litt.).

In the order Julida, Ophyiulus pilosus (Julidae), an introduction
from Europe and a probable troglophile. is rather widespread and
occasionally abundant in Virginia caves. It is also recorded from caves
in Maryland, Ohio, and West Virginia (see Franz and Slifer 1971,
Holsinger et al. 1976, Hobbs and Flynn 1981). The order Polydesmida
is poorly represented in study-area caves, and only a few records are
known. Brachydesmus superus (Polydesmidae), either a trogloxene or
accidental, is common in Europe and in cultivated areas of the United
States; it is recorded from single caves in Virginia and West Virginia

Invertebrate Cave Fauna

Fig. 22. Distribution of troglobitic millipeds ( Trichopetalum ) in the study
area.

(see Holsinger et al. 1976). Another polydesmid, Scytonotus granulatus,
probably a trogloxene, is recorded from a single cave in Virginia and
from caves in Maryland, Ohio, and Pennsylvania (Franz and Slifer
1971, Holsinger 1976, Hobbs and Flynn 1981); it is widespread over
most of the eastern half of the United States (Chamberlin and Hoffman
1958).

The order Spirostreptida is represented by two species of Cambala
(Cambalidae). Cambala minor , a troglophile, is widespread in the
southeastern and east-central United States (Shelley 1979) and has been
collected from caves in Virginia and eight other states within its range
(see Loomis 1943, Shear 1969, Holsinger and Peck 1971, Black 1971,
Craig 1975, Holsinger et al. 1976, Peck and Lewis 1978, Hobbs and

John R. Holsinger and David C. Culver

Flynn 1981). Cambala annulata , a probable trogloxene, ranges over a
large part of the southeastern United States (Shelley 1979) and is
recorded from a few caves in Florida, Georgia, Alabama, and Virginia
(see Holsinger and Peck 1971). Abacion magnum (Caspiopetalidae), the
only representative of the order Callipodida in study-area caves, is a
trogloxene recorded from several caves in southwestern Virginia and
northwestern Georgia (see Holsinger and Peck 1971).

Order Polydesmida

Family Polydesmidae
Brachydesmus superus Latzel (TX or AC)

Virginia. — Tazewell Co.: Lawson Cave.

Pseudopolydesmus sp.

Virginia. — Russell Co.: Dickenson Cave.

Polydesmus angustus Latzel (AC)

Virginia. — Page Co.: Ruffners Cave No. 1.

Scytonotus granulatus (Say) (TX?)

Virginia. — Rockbridge Co.: Billy Williams Cave.

Order Spirostreptida

Family Cambalidae

Cambala annulata (Say) (TX?)

Virginia. — Giles Co.: New River Cave. Page Co.: Luray Caverns (?)
Smyth Co.: Sugar Grove Cave No. 10.

Cambala minor Bollman (TP)

Virginia. — Alleghany Co.: Me Elwee Cave. Augusta Co.:, Glade
Cave. Bath Co.: Clarks Cave. Washington Co.: Wills Cave.
Cambala sp.

Virginia. — Augusta Co.: Fountain and Madisons Saltpetre caves.
Giles Co.: Spruce Run Mountain Cave. Roanoke Co.: Millers
Cove Cave.

Order Callipodia

Family Caspiopetalidae
Abacion magnum (Loomis) (TX)

Virginia. — Lee Co.: Cumberland Gap Saltpetre Cave. Russell Co.:
Dickenson Cave. Tazewell Co.: Lost Mill Cave No. 3.

Order Chordeumatida

Family Cleidogonidae

Pseudotremia (species listed by group as indicated)

eburnea group

Pseudotremia nodosa (s. lat.) Loomis (TB)

Tennessee. — Claiborne Co.: Buis Saltpetre, Chadwells, Clines,
English (type locality), Hauser Spring, Keck No. 1, Lower

Invertebrate Cave Fauna

Coonsies Creek, Saur Kraut, and Tazewell Saltpetre caves.
Hancock Co.: Subers Cave. Union Co.: Wolf Cave.

Virginia. — Lee Co.: Cedar Hill, Cope, Crouse, Gallohan No. 1 and
2, Gibson-Frazier, Gilley, Jones Saltpetre, Knapper, Litton No.
1, Lucy Beatty, Molly Wagle, Smith, Spangler, Surgener, Sweet
Potato, Thompson Cedar, Unthanks, and Young-Fugate caves.

Pseudotremia sp. ( nodsa complex) (TB)

Virginia. — Scott Co.: Flannery, Kerns No. 1, and McDavids caves.
Wise Co: Kelly, Wildcat (Cavern) and Wildcat Saltpetre caves.

fracta group

Pseudotremia cocytus Shear (TP)

Tennessee. — Campbell Co.: Norris Dam Cave.

Comments. — Also recorded from two caves and a wooded hillside
(epigean) just south of the study area in Anderson County, Tenn.
(see Shear 1972).

Pseudotremia fracta (5. lat .) Chamberlin (TP)

Tennessee. — Claiborne Co.: Bug Hole No. 1 and John Lard caves.
Union Co.: Lost Creek Cave.

hobbsi group

Pseudotremia cavernarum Cope (TB)

Virginia. — Montgomery Co.: Daves and Erhart (type locality) caves.

Pseudotremia deprehendor Shear (TB?)

Tennessee. — Grainger Co.: Cedar Springs Cave.

Comments. — Also recorded from Feathers Cave (type locality) just
south of the study area in Anderson County, Tenn. (see Shear
1972).

Pseudotremia hobbsi Hoffman (TP)

Virginia. — Alleghany Co.: Arritt Mill Tunnel, Blue Springs,
Chestnut Ridge (type locality), Island Ford, Lowmoor, Rum-
bolds, Second Dam, and Wares caves. Botetourt Co.: Henderson
No. 1 and Thomas caves. Craig Co.: Shires Saltpetre Cave.
Montgomery Co.: Slussers Chapel Cave.

Pseudotremia princeps Loomis (TP)

Virginia. — Highland Co.: Van Devanters Cave.

Comments. — Also recorded from several caves and one epigean
locality just north and west of the study area in Pendleton
County, W.Va. (see Shear 1972, Holsinger et al. 1976).

Pseudotremia sublevis Loomis (TP)

Virginia. — Giles Co.: “Big Stony” (Cope 1869), Clover Hollow,
Smokehole, Spruce Run Mountain, and Tawneys (type locality)
caves.

Comments. — Also recorded from several epigean localities in the
Giles-Montgomery county area (see Loomis 1944, Shear 1972).

John R. Holsinger and David C. Culver

spira group

Pseudotremia valga Loomis (TB?)

Tennessee. — Claiborne Co.: Station Creek Cave.

Virginia. — Lee Co.: Cudjos Cavern (type locality) and Young-
Fugate Cave.

tuberculata group

Pseudotremia armesi {s. lat.) Shear (TP)

Virginia. — Tazewell Co.: Fallen Rock Cave.

Comments. — Also recorded from two caves and one epigean locality
just west of the study area in Mercer County, W.Va. (Shear
1972, Holsinger et al. 1976).

Pseudotremia momus Shear (TP)

Virginia. — Smyth Co.: Atwells Tunnel and Spence (type locality)
caves.

Comments. — Also recorded from an epigean habitat on the crest of
Big Walker Mountain near the Wythe-Bland county line (Shear
1972.

Pseudotremia tuberculata (s. lat.) Loomis (TB?)

Virginia. — Tazewell Co.: Bowens, Cassell Farm (type locality),
Fallen Rock, Lawson, and Stonley caves.

Undescribed and undetermined species
Pseudotremia n. spp. (TB and TP)

In addition to several undescribed probable species in the nodosa
complex listed above, the following cave populations have been
tentatively recognized as undescribed species by either W. A.
Shear or R. L. Hoffman (in litt.). All need further study and are
not counted in our numerical analyses.

1. Ballards Cave, Giles County.

2. Banners Corner and Dickenson caves, Russell County.

3. Blowing Cave, Bath County.

4. Buchanan Saltpetre Cave, Smyth County.

5. Carter Cave, Lee County.

6. Cave School Water Cave, Wythe County.

7. Coley Cave No. 2, Scott County.

8. Crossroads and Porters caves, Bath County.

9. Cumberland Gap Saltpetre Cave, Lee County.

10. Elys Moonshine and Sweet Potato caves, Lee County.

11. Fisher Cave, Lee County.

12. Greears Sweet Potato and Kerns No. 1 caves, Scott County.

13. Little Kennedy Cave, Wise County.

14. Moccasin Valley Cave, Scott County.

15. New Castle Murder Hole and Rufe Caldwell caves, Craig
County (two species).

16. Pearson Cave, Hawkins County.

Invertebrate Cave Fauna

17. Smiths Cave, Russell County; Hugh Young and Steeles caves,
Tazewell County.

18. Starnes Cave, Giles County.

19. Wares Cave, Alleghany County.

Pseudotremia spp.

Tennessee. — Campbell Co.: Easterly, Meredith, and Panther No. 1
caves. Claiborne Co.: Buis Saltpetre and Kings Saltpetre caves.
Grainger Co.: Horseshoe Cave. Hancock Co.: Cantwell Valley,
Fairmont School, and Panther Creek caves. Hawkins Co.: Pear-
son and Sensabaugh Saltpetre caves. Sullivan Co.: Morrill and
Potters caves. Union Co.: Lost Creek and Oaks caves.

Virginia. — Alleghany Co.: Rumbolds, Walking, and Wares caves.
Bath Co.: Clarks and Dunns caves. Bland Co.: Repass Saltpetre
Cave. Craig Co.: Loneys Cave. Giles Co.: Canoe, Giant (Caverns),
New River, and Starnes caves. Highland Co.: Roaring Springs
Cave. Lee Co.: Bowling, Cattle, Davis, Ely, Frazier, Gibson No.
1, Gilley, Gregory, Indian, Kinzer Hollow, McClure, Roadside
No. 1, Ruths, Seals Pit, Skylight, and Smiths Milk caves.
Montgomery Co.: Fred Bulls Cave. Roanoke Co.: Hodges No. 1
and Millers Cove caves. Russell Co.: Jessie, Johnson, and Porgie
Bundy caves. Rockbridge Co.: Doll House Cave. Scott Co.:
Alley, Blair-Collins, Blowing Hole, Bolling, Cox Ram Pump,
Cox Ridge, Grigsby, Harris Pit, Herron No. 1, Hill, Hortons,
Jack, Jackson, Lane, Obeys Creek, Pond, Natural Tunnel
(Cavern), Quillen No. 1, Spurlock, Taylor No. 1, Winding Stair,
and Wolfe caves. Smyth Co.: Tilson Saltpetre Cave. Tazewell
Co.: Barnes Dry, Gillespie Water, Gully, Lawson, Lost Mill No.
3, Rosebaums Water, and Wagoners caves. Washington Co.:
Neals and Perkins caves. Wise Co.: Parsons and Rocky Hollow
caves. Wythe Co.: Pickett Cave.

Comments. — These records are based primarily on collections
containing juveniles and females, of which specific determinations
could not be made.

Family Conotylidae

Conotyla venetia Hoffman (TX?)

Virginia. — Alleghany Co.: Paxtons Cave.

Family Striariidae
Striaria columbiana Cook (TX?)

Virginia. — Warren Co.: Allens Cave.

Striaria sp. A. (TB?)

Virginia. — Augusta Co.: Madisons Saltpetre Cave.

Striaria sp.

Virginia. — Page Co.: Will Mauck Cave. Shenandoah Co.: Hensleys
Cave.

John R. Holsinger and David C. Culver

Family Trichopetalidae

Trichopetalum packardi (, s . lat.) (Causey) (TB)

Virginia. — Bland Co.: Coon, Hamilton, Newberry-Bane, and Repass
Saltpetre caves. Botetourt Co.: Peery Saltpetre Cave. Craig Co.:
Rufe Caldwell Cave. Giles Co.: Canoe, Clover Hollow, Giant
(Caverns), Starnes, Straleys No. 1, and Tawneys caves. Pulaski
Co.: Fifty-Foot Hell and Sam Bells caves. Roanoke Co.: Dixie
Caverns. Wythe Co.: Sam Six Cave.

Trichopetalum weyeriensis (s. lat.) (Causey) (TB)

Virginia. — Augusta Co.: Grand Caverns (type locality) and
Madisons Saltpetre Cave. Bath Co.: Boundless, Breathing,
Butler-Sinking Creek, Porters, and Starr Chapel caves. Rock-
bridge Co.: Billy Williams Cave.

Trichopetalum whitei (s. lat.) (Ryder) (TB)

Virginia. — Augusta Co.: Glade Cave. Page Co.: Luray Caverns
(type locality) and Ruffners Cave No. 1. Rockingham Co.:
Endless (Caverns), Stevens, and Three-D Maze caves. Shenan-
doah Co.: Maddens, Shenandoah (Caverns), and Shenandoah
Wild caves.

Trichopetalum spp.

Virginia. — Alleghany Co.: Blue Spring Cave. Bath Co.: Dunns
Cave. Bland Co.: Banes Spring Cave. Craig Co.: Loneys Cave.
Giles Co.: New River Cave. Highland Co.: Roaring Springs
Cave. Montgomery Co.: Old Mill and Slussers Chapel caves.
Pulaski Co.: James Cave. Roanoke Co.: Goodwins Cave.
Rockbridge Co.: Grahams Cave.

Comments. — These records are based primarily on juveniles and
females, for which specific determinations could not be made.

Order Julida

Family Julidae

Ophyiulus pilosus (Newport) (TP or TX)

Tennessee. — Hawkins Co.: Sensabaugh Saltpetre Cave.

Virginia. — Alleghany Co.: Wares Cave. Giles Co.: Ballards Cave.
Lee Co.: Carter Cave. Montgomery Co.: Erharts Cave. Page Co.:
Ruffners and Will Mauck caves. Rockbridge Co.: Tolleys Cave.
Rockingham Co.: Melrose Cave. Russell Co.: Banners Corner
and Dickenson caves. Washington Co.: Hall Bottom Cave No. 1.
Family Parajulidae
Ptyoiulus sp.

Virginia. — Tazewell Co.: Gully Cave.

Class Insecta

Insects, along with crustaceans, spiders, and millipeds, are the most
common animals in the caves of Virginia and east Tennessee. At least 1 1

Invertebrate Cave Fauna

orders have been documented from study-area caves, but a majority of
the cavernicoles are in the orders Collembola, Diplura, Orthoptera,
Coleoptera, and Diptera. Many troglobites are noted among the
collembolans, diplurans, and coleopterans.

Only a few scattered records exist for representatives of other insect
orders, none of which is commonly found in caves of the study area.
These include the: mayfly order Ephemeroptera; moth and butterfly
order Lepidoptera (e.g., Scoliopteryx libatrix)\ scorpion fly order
Mecoptera (e.g., family Bittacidae); stonefly order Plecoptera (e.g.,
Leuctra decepta ); caddis fly order Trichoptera (e.g., Hydropsyche
deprevata, H. betteni , and Ochrotrichia)\ and bristletail order Thysanura
(e.g., Machiloides ).

Order Collembola

Colembolans or springtails are common and often abundant in
caves where they are frequently seen in and around damp, decaying
organic material. In the study area the order is represented by 5
families, 9 genera, and 26 described species. Three or four species are
troglobites, eight or nine are troglophiles, and the remainder are
trogloxenes and accidentals

Of the five families, Entomobryidae is clearly the most significant
in terms of abundance and diversity. Pseudosinella is represented by
seven described species from caves in the Virginia-east Tennessee area
(Fig. 23, 24) and two, P. hirsuta and P. orba, are troglobites. Outside
the area, the former species is recorded from numerous caves in
northern Alabama, northwestern Georgia, central Kentucky, and middle
Tennessee, and from one epigean locality on Pine Mountain in Campbell
County, Tenn. (Christiansen and Bellinger 1980c). The latter species has
a much narrower range and is restricted to the study area and adjacent
Mercer County in southern West Virginia (see Holsinger et al. 1976).
The other species of this genus reported from study-area caves are
troglophiles and trogloxenes that range over much of the southeastern
United States.

The genus Sinella is representd in study-area caves by four species,
one of which, S. hoffmani (see Wray 1952), is considered a troglobite
(Fig. 23, 24). This species is also recorded from nine counties in eastern
West Virginia (Holsinger et al. 1976) and one in Pennsylvania (K. A.
Christiansen, in litt.). It has been collected three times from surface
habitats, twice in North Carolina and once in West Virginia (Christian-
sen and Bellinger 1980c), but the identity of the North Carolina
specimens is questionable (Christiansen 1982). The lone record of this
species from Tazewell County in the Clinch drainage basin is also
questionable. The other species of Sinella noted from study-area caves
are troglophiles and trogloxenes and have wide ranges outside Virginia
and eastern Tennessee (see Christiansen 1960a).

John R. Holsinger and David C. Culver

Probably the most common and widespread cavernicolous collem-
bolan in the study area is Tomocerus bidentatus (Fig. 31 A), a lightly
pigmented troglophile with small eyes that is also recorded from epigean
and cave habitats in the eastern United States and from two caves in
California (Christiansen 1964a, Christiansen and Bellinger 1980c).
Tomocerus flavescens, also a troglophile (or trogloxene?), is recorded
from caves in many parts of the United States (Christiansen 1964a,
Christiansen and Bellinger 1980c), but it is much less common than T.
bidentatus in the study area (Fig. 25).

The second most significant family in the regional cave collembolan
fauna is Sminthuridae, represented by six species in the genus
Arrhopalites (Fig. 24). Most of these species are troglophiles and
trogloxenes and are recorded from a large part of the United States (see
Christiansen and Bellinger 1981). Arrhopalites clarus, however, is at
present known only from caves and is apparently troglobitic despite its
broad distribution, which includes localities in Arkansas, Missouri,
Virginia, and West Virginia (see Christiansen 1982). The most common
species of the genus in regional caves is A. pygmaeus, a troglophile
uncommon in epigean habitats but recorded from caves throughout a
large part of the southeastern and south-central United States
(Christiansen 1964a, Christiansen and Bellinger 1981). Arrhopalites
ferrugineus (Packard), reported earlier from caves in Virginia by
Holsinger (1963a), is considered a synonym of this species by
Christiansen (1966).

In other families, Folsomia Candida (Isotomidae), a probable
troglophile, is recorded from a few caves in Virginia as well as from
caves over a wide area of the United States (Christiansen and Bellinger
1980b). The families Hypogastruridae and Onychiuridae are represented
in study-area caves by several trogloxene or accidental species, which,
with the exception of Onychiurus ramosus, are based on single cave
records. Onychiurus ramosus is recorded from several caves in Virginia
and one in northeastern Utah (see Peck 1981a); otherwise it is widespread
in epigean habitats over much of the United States (Christiansen and
Bellinger 1980b).

Family Entomobryidae
Entomobrya socia Boren (TX or AC)

Virginia. — Giles Co.: New River Cave.

Pseudosinella aera Christiansen and Bellinger (TP or TX)

Virginia. — Shenandoah Co.: “Cave” (Christiansen and Bellinger
1980c:966).

Comments. — Also recorded from caves in Illinois, Kentucky,
Missouri, Tennessee, and Texas (Christiansen and Bellinger
1980c).

Invertebrate Cave Fauna

Fig. 23. Distribution of troglobitic collembolans (Pseudosinella and Sinella) in
the study area. Single locality for P. orba in Mercer County, W.Va.. also shown.
Two symbols in a circle indicate two species from the same cave.

▼ Pseudosinella hirsuta
• R orba

▲ Sinella hoffmani

WEST VIRGINIA

VIRGINIA

KENTUCKY

■ Arrhopalites clarus
• A. pygmaeus

□ Pseudosinella argentea |

▲ Sinella barri

WEST VIRGINIA

KENTUCKY

VIRGINIA

0 25^ 50 km

50 miles

NORTH CAROLINA

Fig. 24. Distribution of cavernicolous collembolans {Arrhopalites .
Pseudosinella , and Sinella ) in the study area. Two symbols in a circle indicate
two species from the same cave.

John R. Holsinger and David C. Culver

Pseudosinella alba (Packard) (TX)

Virginia. — Shenandoah Co.: Shenandoah Wild Cave.

Comments. — Widely distributed in United States; recorded from a
few caves (Christiansen and Bellinger 1980c).

Pseudosinella argentea Folsom (TP)

Virginia. — Augusta Co.: Grand Caverns and Madisons Saltpetre
Cave. Bland Co.: Newberry-Bane Cave. Highland Co.: Marshall
Cave. Lee Co.: Kinzer Hollow Cave (?). Russell Co.: Fraleys
Cave.

Comments. — Also recorded from caves in Arkansas, Illinois,
Kentucky, Missouri, and Tennessee (Christiansen and Bellinger
1980c).

Pseudosinella collina Wray (TP or TX)

Virginia. — Giles Co.: New River Cave. Pulaski Co.: Sam Bells
Cave.

Comments. — Also recorded from caves in Alabama, Kentucky, and
Tennessee (Christiansen and Bellinger 1980c). All North American
records for P. duodecimpuncata Denis probably should be
referred to this species (Christiansen and Bellinger 1980c).

Pseudosinella hirsuta (Delamare) (TB)

Tennessee. — Campbell Co.: Easterly and Meredith caves.

Virginia. — Lee Co.: Cliff, Cudjos (Cavern), and Skylight caves.

Pseudosinella orba Christiansen (TB)

Tennessee. — Sullivan Co.: Morrill Cave (type locality).

Virginia. — Bland Co.: Hamilton Cave. Craig Co.: Rufe Caldwell
Cave. Giles Co.: Starnes Cave. Lee Co.: Gallohan No. 1, Smith,
and Sweet Potato caves. Pulaski Co.: Sam Bells Cave. Roanoke
Co.: Goodwins Cave (?). Russell Co.: Porgie Bundys Cave. Scott
Co.: Blair-Collins Cave. Smyth Co.: Buchanan Saltpetre, Inter-
state-81, and Tilson Saltpetre caves. Tazewell Co.: Fallen Rock
and Gully caves. Wise Co.: Wildcat Saltpetre Cave.

Pseudosinella sexoculata Schott (TX)

Virginia. — Shenandoah Co.: Battlefield Crystal Cave.

Comments. — Distributed over much of the United States; also
recorded from caves in Iowa, Kentucky, and New Mexico
(Christiansen 1960a,b; Christiansen and Bellinger 1980c).

Pseudosinella spp.

Tennessee. — Hawkins Co.: Pearson Cave.

Virginia. — Tazewell Co.: Cassell Farm and Lawson caves. Wash-
ington Co.: Fritz Breathing Cave.

Comments. — Both the Pearson Cave and Fritz Breathing Cave
populations probably represent undescribed species (K. A.
Christiansen, in litt.).

Sinella barri Christiansen (TP)

Tennessee. — Union Co.: Wolfe Cave (?).

Invertebrate Cave Fauna

• Tomocerus bktontatus
□ T. flavescens

WEST VIRGINIA

KENTUCKY

VIRGINIA

0 25 50 km

Q 25 50 miles

NORTH CAROLINA

Fig. 25.
area.

Distribution of troglophilic collembolans ( Tomocerus ) in the study

Virginia. — Alleghany Co.: Island Ford Cave. Giles Co.: Parsells
Cave. Scott Co.: Herron Cave No. 1. Washington Co.: Vickers
Cave. Wythe Co.: Picketts Cave.

Comments. — Also recorded from caves in Arkansas, Illinois,
Kentucky, Missouri, and Tennessee (Christiansen and Bellinger
1980c).

Sinella caeca Schott (TX)

Virginia. — Frederick Co.: Ogdens Cave. Page Co.: Luray Caverns.
Pulaski Co.: James Cave. Rockbridge Co.: Showalters Cave.
Shenandoah Co.: Shenandoah Caverns.

Comments. — Reported from epigean localities throughout most of
the United States; also recorded from caves in Iowa, Kentucky,
Missouri, Texas, and Wisconsin (Christiansen and Bellinger
1980c).

Sinella curviseta Brook (TX or AC)

Virginia. — Montgomery Co.: Erharts Cave.

Comments. — Reported from epigean localities over much of the
United States; also recorded from a cave in Kentucky (Bonet
1934, Christiansen and Bellinger 1980c).

Sinella hoffmani Wray (TB)

Virginia. — Alleghany Co.: Blue Spring, Island Ford, Lowmoor

John R. Holsinger and David C. Culver

(type locality), Rumbolds, and Wares caves. Bath Co.: Boundless,
Breathing, Butler-Sinking Creek, Crossroads, Dunns, Porters,
Starr Chapel, and Witheros caves. Botetourt Co.: Peery Saltpetre
Cave. Roanoke Co.: Goodwins Cave. Rockbridge Co.: Buck Hill
and Doll House caves. Tazewell Co.: Stonley Cave (?).

Tomocerus bidentatus Folsom (TP)

Tennessee. — Claiborne Co.: English and Lower Coonsies Creek
caves. Hawkins Co.: Sensabaugh Saltpetre Cave. Sullivan Co.:
Bristol Caverns and Morrill Cave. Union Co.: Lost Creek Cave.
Virginia. — Alleghany Co.: Island Ford and Paxtons caves. Augusta
Co.: Glade and Madisons Saltpetre caves. Bath Co.: Roy Lyle
and Porters caves. Botetourt Co.: Peery Saltpetre Cave. Frederick
Co.: Ogdens Cave. Giles Co.: Harris and New River caves.
Highland Co.: Marshalls Cave. Lee Co.: Cudjos (Cavern),
Cumberland Gap Saltpetre, Gallohan No. 1, Kinzer Hollow,
Lucy Beatty, Skylight, and Sweet Potato caves. Montgomery
Co.: Erharts Cave. Page Co.: Foltz Cave No. 1 and Luray
Caverns. Roanoke Co.: Goodwins and Hodges No. 1 caves.
Rockbridge Co.: Showalters and Tolley caves. Rockingham Co.:
Church Mountain, Massanutten (Caverns), Melrose (Caverns),
Round Hill, Steam Hole, and Stephens caves. Russell Co.: Jessie
and Porgie Bundys caves. Scott Co.: Bolling, Hill, Kerns No. 1,
and Lane caves. Shenandoah Co.: Helsley and Shenandoah Wild
caves. Smyth Co.: Atwells Tunnel, Roberts, and Sugar Grove
No. 10 caves. Washington Co.: Hall Bottom No. 1 and Singleton
caves. Wythe Co.: Cave School Water and Sam Six caves.
Tomocerus flavescens (Tullberg) (TP or TX)

Virginia. — Bath Co.: Cave Run Pit Cave. Bland Co.: Banes Spring
Cave.

Family Hypogastruridae
Hypogastrura denticulata (Bagnall) (TX)

Virginia. — Giles Co.: Tawneys Cave.

Comments. — Widespread species complex, occasionally found in
caves (see Christiansen and Bellinger 1980a).

Neanura barberi (Handschin) (TX?)

Virginia. — Augusta Co.: Madisons Saltpetre Cave.

Comments. — Recorded from epigean localities in the eastern and
midwestern United States; occasionally found in caves (Christian-
sen and Bellinger 1980a).

Family Isotomidae
Folsomia Candida Willem (TP)

Tennessee. — Campbell Co.: Meredith Cave.

Virginia. — Augusta Co.: Madisons Saltpetre Cave. Pulaski Co.:

Invertebrate Cave Fauna

Fifty-Foot Hell Cave. Roanoke Co.: Goodwins Cave. Wise Co.:
Wildcat Saltpetre Cave.

Folsomia sp.

Tennessee. — Claiborne Co.: English Cave.

Virginia. — Rockbridge Co.: Showalters Cave.

Family Onychiuridae

Onychiurus magninus Wray (AC)

Virginia. — Roanoke Co.: Goodwins Cave.

Onychiurus ramosus Folsom (TX)

Virginia. — Lee Co.: Sweet Potato Cave. Russell Co.: Bundy Cave
No. 2. Wise Co.: Kelly and Wildcat Saltpetre caves.

Onychiurus reus Christiansen and Bellinger (TX or AC)

Virginia. — Warren Co.: Baldwin Hill Cave(s).

Comments. — Recorded from several epigean localities in the eastern
United States and a cave in Kentucky (Christiansen and Bellinger
1980b).

Family Sminthuridae

Arrhopalites benitus (Folsom) (TX)

Virginia. — Alleghany Co.: Island Ford Cave. Bath Co.: Breathing
Cave.

Comments. — Also recorded from a cave in Greenbrier County,
W.Va. (Holsinger et al. 1976).

Arrhopalites caecus (Tullberg) (TX)

Virginia. — Rockbridge Co.: Showalters Cave.

Comments. — Also recorded from caves in Iowa, Minnesota, and
South Dakota (Christiansen and Bellinger 1981).

Arrhopalites clarus Christiansen (TB?)

Virginia. — Montgomery Co.: Old Mill Cave. Wythe Co.: Sam Six
Cave.

Arrhopalites hirtus Christiansen (TP or TX)

Virginia. — Lee Co.: Gallohan Cave No. 1.

Comments. — Also recorded from caves in Illinois, Iowa, Kentucky,
and Wisconsin (Christiansen and Bellinger 1981).

Arrhopalites pygmaeus (Wankel) (TP)

Tennessee. — Claiborne Co.: Station Creek Cave.

Virginia. — Augusta Co.: Grand Caverns and Madisons Saltpetre
Cave. Lee Co.: Smith and Sweet Potato caves. Rockingham Co.:
Endless Caverns, Scott Co.: Flannery and Greears Sweet Potato
caves. Washington Co.: Wills Cave.

Arrhopalites whiteside Jacot (TX)

Virginia. — Alleghany Co.: Island Ford and Lowmoor caves.

Arrhopalites sp.

Virginia. — Russell Co.: Porgie Bundys Cave.

John R. Holsinger and David C. Culver

Order Diplura

Cavernicolous diplurans (Fig. 3 1C) are represented in the study
area by a single genus, Litocampa (Campodeidae), and six species. Only
one of the species has been described; the remainder were recognized in
a thesis and a dissertation by Ferguson (1974, 1981a), but descriptions
have not been published to date and formal names are not available. All
species of Litocampa (formerly a subgenus of Plusiocampa ) in North
America are troglobites (Ferguson 1981b). The range of Litocampa jn
the study area is restricted to the New and Tennessee drainage basins
(Fig. 26). Cavernicolous diplurans are generally found on damp mud or
silt banks near streams and occasionally on damp to wet surfaces
elsewhere. They are sometimes locally abundant on organically enriched
silt but otherwise usually uncommon in a given cave.

Litocampa cookei inhabits caves of the Powell Valley and parts of
the adjacent Clinch Valley. The species is also recorded from caves in
south-central Kentucky and middle Tennessee, where it is common and
fairly widespread (Ferguson 1974). The other species are endemic to the
Appalachian Valley and eastern side of the Appalachian Plateau and,
with two exceptions, are known only from caves in the study area.
Litocampa sp. A and D have very restricted ranges; the former is found
only in caves of the Ward Cove karst in Tazewell County, and the latter
is known only from a single cave in Hancock County. In comparison, L.
sp. B, C, and E have wider ranges as indicated by the records cited
below.

Litocampa sp. B is recorded from caves in the New River basin
(southeast of Walker and Gap mountains) and parts of the Holston
basin. In addition to three caves in Scott County, L. sp. C has been
found in Angel Cave on Pine Mountain, just west of the study area in
Letcher County, Ky. (Ferguson 1981a). Litocampa sp. E has a
moderately extensive range that covers parts of the New, Holston, and
Clinch basins and includes one cave just outside the study area in
Mercer County, W.Va. (Ferguson 1974, Holsinger et al. 1976).

Family Campodeidae
Litocampa cookei (Packard) (TB)

Tenneessee. — Campbell Co.: Meredith and Norris Dam caves.
Claiborne Co.: Tazewell Saltpetre Cave. Hancock Co.: Panther
Creek and Subers caves.

Virginia. — Lee Co.: Gallohan No. 1, Molly Wagle, Sweet Potato,
and Young-Fugate caves. Scott Co.: Spurlock Cave. Wise Co.:
Little Kennedy, Parsons, and Rocky Hollow caves.

Litocampa sp. A (L. M. Ferguson, in ms.) (TB)

Virginia. — Tazewell Co.: Bowens, Fallen Rock (type locality),
Gillespie Water, and Lost Mill No. 1 and 3 caves.

Invertebrate Cave Fauna

Fig. 26. Distribution of troglobitic diplurans ( Litocampa ) in the study area.

Litocampa sp. B (L. M. Ferguson, in ms.) (TB)

Virginia. — Montgomery Co.: Vickers Road Cave. Pulaski Co.:
Fifty-Foot Hell, James, and Sam Bells caves. Smyth Co.:
Interstate-81 Cave. Washington Co.: Brass Kettle Hole Cave.
Wythe Co.: Speedwell Cave No. 1 (type locality).

Litocampa sp. C (L. M. Ferguson, in ms.) (TB)

Virginia. — Scott Co.: Hill, McDavids (type locality), and Queens
caves.

Litocampa sp. D (L. M. Ferguson, in ms.) (TB)

Tennessee. — Hancock Co.: Panther Creek Cave (type locality).

Litocampa sp. E (L. M. Ferguson, in ms.) (TB)

Virginia. — Bland Co.: Coon, Hamilton, and Newberry-Bane caves.
Giles Co.: Giant Caverns and Starnes Cave (type locality). Russell
Co.: Bundys No. 2 and Grays caves. Scott Co.: Blair-Collins,
Coley No. 2, Lane, and Wolfe caves. Smyth Co.: Beaver Creek,
Buchanan Saltpetre, Hancock, and Tilson Saltpetre caves.
Tazewell Co.: Cassell Farm, Lawson, and Wagoners caves.
Washington Co.: Perkins Cave.

Order Orthoptera

Cave crickets (Rhaphidophoridae) are common in caves of Virginia
and east Tennessee, where they are represented by two genera and at
least five species. Ceuthophilus is yellowish-brown with black bands on
the abdomen, is usually seen near entrances, and rarely, if ever, penetrates
caves for an appreciable distance. Three species have been reported
from study-area caves, but C. gracilipes, a threshold trogloxene, is the
most common and widespread. The range of this species extends from

John R. Holsinger and David C. Culver

the Ozarks eastward throughout much of the Appalachian region and
includes numerous caves (Hubbell 1936, Holsinger and Peck 1971, Peck
and Lewis 1978). No attempt has been made to collect Ceuthophilus
systematically from caves in Virginia and east Tennessee; thus its
occurrence in study-area caves is more common than indicated by the
few records cited below.

In comparison with Ceuthophilus , Euhadenoecus , the other genus
found in regional caves, is light brown in color, lacks conspicuous
banding, and has a more slender body with longer legs. Euhadenoecus
puteanus , a threshold trogloxene like C. gracilipes, is widely distributed
throughout much of the Appalachian region and a part of the Interior
Low Plateaus. It is recorded from numerous caves and epigean localities,
many of these in Virginia and east Tennessee (see Hubbell and Norton
1978). Euhadenoecus fragilis, in contrast to E. puteanus , is a habitual
trogloxene, or a troglophile under some circumstances. It is lightly
pigmented, has attenuated legs (Fig. 32B), and is closely associated with
the cave environment. It breeds in caves and commonly occurs far from
entrance zones. The range of this species (Fig. 27) extends from Bath
County, Va., and southern Randolph County, W.Va., southwestward to
Claiborne County, Tenn., and includes Pine Mountain in southeastern
Kentucky; the majority of locality records are from caves (see Hubbell
and Norton 1978). Although common in the Clinch and Powell valleys,
it is to date unrecorded from the Holston Valley. In addition to the
localities listed below, we have made many unrecorded sightings of E.
fragilis in southwestern Virginia caves.

Family Rhaphidophoridae

Ceuthophilus brevipes Scudder (TX)

Virginia. — Botetourt Co.: Thomas Cave. Giles Co.: Tawneys Cave.
Roanoke Co.: Hodges Cave No. 1. Tazewell Co.: Cassell Farm
and Little Gully caves.

Ceuthophilus gracilipes gracilipes (Haldeman) (TX)

Virginia. — Botetourt Co.: Henderson Cave No. 1. Giles Co.:
Tawneys Cave. Highland Co.: Hamilton Cave. Lee Co.: Waltons
Cave. Montgomery Co.: Fred Bulls Cave. Roanoke Co.: McVitty,
Millers Cove, and New Dixie caves. Rockbridge Co.: Doll House
Cave. Rockingham Co.: Massanutten Caverns. Scott Co.: Queens
and Speers Ferry caves. Tazewell Co.: Cassell Farm and Little
Gully caves.

Comments. — Also recorded from “Old Joe’s Cave,” east of the Blue
Ridge Mountains in Buckingham County, Va. (see Hubbell
1936).

Ceuthophilus pallidipes Walker (TX)

Virginia. — Bland Co.: Hamilton Cave. Highland Co.: Better
Forgotton Cave. Lee Co.: Waltons Cave. Roanoke Co.: Hodges

Invertebrate Cave Fauna

Fig. 27. Distribution of troglophilic crickets ( Euhadenoecus ) in the study area.

Cave No. 1. Rockbridge Co.: Tolleys Cave. Scott Co.: Queens
Cave.

Euhadenocecus fragilis Hubbell (TP or TX)

Tennessee. — Claiborne Co.: Bug Hole No. 1, English, and Saur
Kraut caves. Hancock Co.: Newmans Ridge (Hubbell and Norton
1978:43), Caney Sinks, and Subers caves.

Virginia. — Alleghany Co.: Wares Cave. Giles Co.: Ballards, Links,
Smokehole, Starnes, and Tawneys (type locality) caves. Lee Co.:
Cattle, Cliff, Crouse, Cumberland Gap Saltpetre, Gibson No. 2,
Gibson-Frazier, Gilley, Indian, Kinzer Holow, Molly Wagle,
Roadside No. 1, Smiths Milk, Spangler, Sweet Potato, Thompson
Cedar, Unthanks, Waltons, and Young-Fugate caves; also “small
caves,” Pennington Gap (Hubbell and Norton 1978:43). Roanoke
Co.: Millers Cove Cave. Russell Co.: Banners Corner, Indian,
and Seven Springs caves. Scott Co.: Blowing Hole, Coley No. 1
and 2, Hortons, Queens, and Speers Ferry caves. Tazewell Co.:
Cassell Farm, Glenwood Church, Lawson, Spider, and Wagoners
caves.

Euhadenoecus puteanus (Scudder) (TX)

Tennessee. — Claiborne Co.: Yoakum Cave. Sullivan Co.: Bristol
(Caverns), Morrill, and Potters caves.

Virginia. — Alleghany Co.: Lowmoor and Wares caves. Giles Co.:
New River Cave. Highland Co.: Better Forgotten and Van
Devanters caves. Lee Co.: Gilley Cave (not Billeys Cave or
Baileys Cave as listed by Hubbell and Norton 1978:31). Roanoke

John R. Holsinger and David C. Culver

Co.: Dixie Caverns and McVitty Cave. Rockbridge Co.: Doll
House Cave. Rockingham Co.: “cave” (see Hubbell and Norton
1978:31). Russell Co.: Seven Springs Cave. Smyth Co.: Atwells
Tunnel and Stones No. 2 caves. Tazewell Co.: Cassell Farm
Cave(s). Washington Co.: Hookers Rock Cave.

Euhadenoecus spp.

Virginia. — Bath Co.: Roy Lyle Cave. Giles Co.: Clover Hollow
Cave. Highland Co.: Better Forgotten Cave. Rockbridge Co.:
Billy Williams and Tolleys caves. Rockingham Co.: Gay Hill and
Three-D Maze caves. Russell Co.: Johnson Dry Cave.

Order Coleoptera

Beetles constitute the most diverse group of insects in study-area
caves, where they are represented by 8 families, 36 genera, and more
than 75 species. Biospeleologically, the most important families are
Cantharidae, Carabidae, Leiodidae, Pselaphidae, and Staphylinidae.
Representatives of Cryptophagidae ( Cryptophagus sp.), Dytiscidae
( Hydroporus wickhami), and Scarabaeidae (Ataenuis spretulus and
Aphodius rufipes) were also noted, but only as occasional accidentals.

Most of the cavernicolous beetles in the study area belong to the
Carabidae and the large, predominantly troglobitic genus Pseu-
danophthalmus. Forty-seven species of this genus have been recognized,
36 of which have been described to date (see Barber 1928; Jeannel 1928,
1931, 1949; Valentine 1931, 1932, 1945, 1948; Barr 1960a, 1965, 1981a,
1985). Many closely related species inhabit caves in adjacent areas (e.g.,
West Virginia, eastern Kentucky, southeastern Tennessee). Most of the
species are locally endemic; 26 are known only from a single cave and
13 from a small cluster of caves (Fig. 28, 29, 30). However, a few, like P.
delicatus (Fig. 32A) and P. hoffmani, have significantly wider ranges
with linear extents of approximately 50 and 75 km, respectively. Of the
11 species groups currently recognized from the region by Barr (1981a),
only two are endemic to the study area. The other nine contain species
that also occur outside the area. Both the engelhardti and the hirsutus
groups include species that occur relatively far from the study area in
southeastern Tennessee, northwestern Georgia, and northern Alabama
(see Barr 1981a). One member of the engelhardti group, P. wallacei,
occurs just south of the study area in Anderson County, however. The
gracilis , grandis , hub bar di, and pusio groups contain species that inhabit
caves just west of the study area is eastern West Virginia. Four of the
five species assigned to the hypolithos group by Barr (1981a) occur in
caves on the northwest side of Pine Mountain in southeastern Kentucky,
also just west of the study area. The jonesi group also contains species
that occupy caves in Pine Mountain, one in southeastern Kentucky and
one in Campbell County, Tenn. Another species of this group is found

Invertebrate Cave Fauna

• engelhardti group

deceptivus

engelhardti

holsingeri

rotundatus

sidus

wallacei

sp. A
8 sp. B

Pseudanopthalmus

▲ tennesseensis group
9 paynei

10 pusillus

1 1 unionis

* hypolithos group

12 praetermissus

♦ hubrichti group

13 egberti

14 hubrichti

15 paradoxus

16 quadratus

17 sa net i pauli

18 vicarius ^

19 sp. A

20 sp. D '

KENTUCKY

▼ gracilis group
21 gracilis

VIRGINIA

25 50 km

25 50 miles

— j-T

)

Fig. 28. Distribution of troglobitic beetles ( Pseudanophthalmus ) in the study
area. Localities for three species just south of the study area in Anderson
County, Tenn., also shown.

0 25 50 miles

.OY' 7

Fig. 29. Distribution of troglobitic beetles ( Peudanophthalmus ) in the study

Pseudanophtha I mus

• hirsutus group

1 delicatus

2 hirsutus

3 serious

▼ grandis group
II virginicus

a jonesi

4 cordicollis

5 longiceps

6 pallidus

7 seclusus

8 thomasi

9 sp. A

10 sp. B

KENTUCKY

area.

John R. Holsinger and David C. Culver

in Grassy Cove, a karst island in the Cumberland Plateau some 80 km
southwest of the study area (see Barr 1981a). Three of the four species
of the tennesseensis group are recorded from caves just south of the
study area in Anderson, Knox, and Roane counties (viz., P. paynei , P.
pusillus , and P. tennesseensis).

Although some species of Pseudanophthalmus may be sporadically
abundant in a given cave, most are quite rare; and several species, such
as those of the hubbardi group in the Shenandoah Valley, are known
only from a few specimens collected over a period of many years.
Cavernicolous carabids, especially Pseudanophthalmus , are typically
found in damp to wet areas under rocks or around organic detritus.

The non-troglobitic carabids from regional caves include species of
Atranus , Bembidion, Patrobus, Platynus, Rhadine , Stenolophus, and
Trechus. Perhaps the most common of these is Platynus tenuicollis , a
troglophile recorded from caves in the eastern United States, the
Ozarks, and Texas (Peck and Lewis 1978). Agonum {Platynus) reflexum,
reported from caves in the eastern United States by Barr (1964), is now
considered a synonym of P. tenuicollis (T. C. Barr, Jr., in litt.).
Bembidion and Atranus may also be occasionally abundant. Both B.
lacunarium and B. wingatei are recorded from Virginia caves; the
former is common in caves in the central and eastern United States
(Peck and Lewis 1978); the latter is reported from caves in eastern
Kentucky, Pennsylvania, and West Virginia (Barr 1964, Holsinger et al.
1976). Atranus pubescens, a troglophile recorded from caves in the
central and eastern United States (see Peck and Lewis 1978), is known
from several Virginia caves. Rhadine caudata , a fairly widespread
troglophile in caves in Alabama, Georgia, and Tennessee (Barr 1960b,
1964; Holsinger and Peck 1971), was reported from a single cave in
Virginia by Bolivar and Jeannel (1931). Trechus hydropicus canus,
probably a trbgloxene, is recorded from a single cave in Lee County but
is more common in surface localities at higher elevations in eastern
Kentucky and southwestern Virginia (Barr 1979).

Cavernicolous pselaphid and leiodid beetles are poorly represented
in the study area in contrast to parts of the Cumberland Plateau and
Interior Low Plateaus where they are more diverse and represented by
numerous troglobites (Park 1960, Peck 1973). Only two troglobitic
pselaphids are known from the study area: Arianops jeanneli and Batria-
symmodes greeveri (Fig. 30). Both species are very rare, local endemics
(see Park 1956, 1965; Barr 1974, 1987). The former has been found only
once, despite several diligent searches in the type locality. These species
may be edaphobites and not troglobites, but this remains to be
determined. Other pselaphids include Batriasymmodes monstrosus,
probably accidental, for which Poor Farm Cave in Lee County is the
only documented cave record to date for this widespread epigean species

Invertebrate Cave Fauna

Fig. 30. Distribution of troglobitic beetles ( Arianops , Batriasymmodes, and
Pseudanophthalmus ) in the study area. Single locality for P. potomaca in
Pendleton County, W.Va., also shown.

(see Park 1965, Barr 1987), and Batrisodes globosus, a trogloxene
widespread in eastern North America and recorded from single caves in
Alabama, Georgia, and Virginia (Holsinger and Peck 1971). Priono-
chaeta opaca , a trogloxenic leiodid, is recorded from a single cave in
Virginia but is widespread in eastern North America and reported from
caves elsewhere in the southeastern United States (Peck 1977).

Most of the cavernicolous staphylinid beetles are in three
subfamilies— Aleocharinae, Omaliinae, and Staphylininae. Although
fairly common and sometimes moderately abundant in caves, none is a
troglobite. The systematics of the aleocharines, previously poorly known,
is being revised by J. Klimaszewski and S. B. Peck (Klimaszewski 1984,
Klimaszewski and Peck 1986). Aleochara lucifuga appears to be the
most frequently seen member of the subfamily in regional caves, but
Aloconota insecta and Atheta annexa are also relatively common and
widespread. Outside the study area, all of these aleocharines are found
in a number of cave areas in the southeastern United States.

The omaliine Brathinus nitidus (sometimes placed in the family
Brathinidae) is widespread in eastern North America and reported from
caves in several states (Peck 1975a).

John R. Holsinger and David C. Culver

Several troglophilic species in the genus Quedius (Staphylininae)
inhabit regional caves. Both Q. erythrogaster and Q. spelaeus are
recorded from a number of Virginia caves, are found over much of
North America, and are common in caves elsewhere in the eastern
United States (Smetana 1971, Holsinger and Peck 1971, Peck and Lewis
1978). Quedius mesomelinus is found in caves much less frequently,
although its distribution is Holarctic (Smetana 1971).

The family Cantharidae is represented in study-area caves * by
Cantharis, a genus with one or more undetermined trogloxenic species.
In Virginia and east Tennessee, as well as elsewhere in eastern North
America, only larvae have been found in caves (see Peck 1975b).

Family Cantharidae
Cantharis sp. (TX)

Tennessee. — Claiborne Co.: English Cave.

Virginia. — Pulaski Co.: Sam Bells Cave. Scott. Co.: Greears Sweet
Potato Cave. Washington Co.: Brass Kettle Hole Cave.

Family Carabidae
Atranus pubescens (Dejean) (TP)

Virginia. — Scott Co.: Coley Cave No. 2. Washington Co.: Hall
Bottom Cave No. 1.

Bembidion ( Peryphus ) lacunarium (Zimmermann) (TP)

Virginia. — Smyth Co.: Atwells Tunnel and Stones No. 2 caves.
Bembidion (Amerizus) wingatei (Bland) (TP)

Virginia. — Alleghany Co : Wares Cave. Tazewell Co.: Lawson Cave.
Patrobus longicornis (Say) (TX)

Virginia. — Scott Co.: Coley No. 2 and Sparks caves.

Comments. — Also recorded from caves in Alabama, Illinois,
Kentucky, and Missouri (Barr 1964, Peck and Lewis 1978).
Platynus tenuicollis (LeConte) (TP)

Tennessee. — Claiborne Co.: Bug Hole Cave No. 1.

Virginia. — Giles Co.: Ballards Cave. Montgomery Co.: Old Mill
Cave. Rockbridge Co.: Tolleys Cave. Russell Co.: Banners Corner
Cave. Smyth Co.: Atwells Tunnel Cave. Washington Co.: Hall
Bottom Cave No. 1.

Pseudanophthalmus (species listed by group as indicated)

engelhardti group
Pseudanophthalmus deceptivus Barr (TB)

Virginia. — Lee Co.: Fisher Cave (type locality).

Pseudanophthalmus engelhardti (Barber) (TB)

Tennessee. — Claiborne Co.: English Cave (type locality).
Pseudanophthalmus holsingeri Barr (TB)

Virginia. — Lee Co.: Young-Fugate Cave (type locality).

Invertebrate Cave Fauna

Pseudanophthalmus rotundatus Valentine (TB)

Tennessee. — Claiborne Co.: English Cave (type locality). Hancock
Co.: “Coopers” (Jeannel 1949:82) and Subers caves.

Virginia. — Lee Co.: Elys Moonshine, Smith, and Sweet Potato
caves.

Pseudanophthalmus sidus Barr (TB)

Tennessee. — Campbell Co.: Meredith Cave (type locality).

Pseudanophthalmus sp. A (T. C. Barr, Jr., in ms.) (TB)

Tennessee. — Union Co.: Wolf Cave.

Pseudanophthalmus sp. B (T. C. Barr, Jr., in ms.) (TB)

Tennessee. — Campbell Co.: Valley View Cave.

gracilis group

Pseudanophthalmus gracilis Valentine (TB)

Virginia. — Craig Co.: Rufe Caldwell Cave. Giles Co.: Clover
Hollow, Smokehole, and Tawneys (type locality) caves.

grandis group

Pseudanophthalmus virginicus Barr (TB)

Virginia. — Tazewell Co.: Hugh Young Cave (type locality).
Comments. — This species was originally the type species of the
genus Aphanotrechus but is now assigned to Pseudanophthalmus
(see Barr 1960a, 1981a).

hirsutus group

Pseudanophthalmus delicatus Valentine (TB)

Virginia. — Lee Co.: Baileys, Bowling, Cattle, Gallohan No. 1,
Garrett, Gilley (type locality), Jones Saltpetre, Molly Wagle,
Poor Farm, Seal Pit, Smith, Spangler, and Unthanks caves.

Pseudanophthalmus hirsutus Valentine (TB)

Tennessee. — Claiborne Co.: Powell Mountain Cave.

Virginia. — Lee Co.: Cudjos Caverns (type locality) and Cumberland
Gap Saltpetre Cave.

Pseudanophthalmus sericus Barr (TB)

Virginia. — Scott Co.: Lane Cave (type locality).

hubbardi group

Pseudanophthalmus avernus (Valentine) (TB)

Virginia. — Rockingham Co.: Endless Caverns (type locality).

Pseudanophthalmus hubbardi (Barber) (TB)

Virginia. — Page Co.: Luray Caverns (type locality).

Pseudanophthalmus intersectus Barr (TB)

Virginia. — Bath Co.: Crossroads Cave (type locality).

Pseudanophthalmus limicola (Jeannel) (TB)

Virginia. — Shenandoah Co.: Maddens (type locality), Shenandoah
(Caverns), and Shenandoah Wild caves.

John R. Holsinger and David C. Culver

Pseudanophthalmus parvicollis (Jeannel) (TB)

Virginia. — Shenandoah Co.: Battlefield Crystal Cave (type locality).

Pseudanophthalmus potomaca Valentine (TB)

Virginia. — Highland Co.: Van Devanter Cave.

Comments. — Also recorded from Kenny Simmons Cave (type
locality) in adjoining Pendleton County, W.Va.

Pseudanophthalmus sp. A (T. C. Barr, Jr., in ms.) (TB)

Virginia. — Bath Co.: Breathing and Butler-Sinking Creek caves.

hubrichti group

Pseudanophthalmus egberti Barr (TB)

Virginia. — Giles Co.: Giant Caverns and Starnes Cave (type
locality).

Pseudanophthalmus hubrichti Valentine (TB)

Virginia. — Russell Co.: Daughtery Cave (type locality).

Pseudanophthalmus paradoxus Barr (TB)

Tennessee. — Hawkins Co.: Sensabaugh Saltpetre Cave (type
locality).

Pseudanophthalmus quadratus Barr (TB)

Virginia. — Giles Co.: Straleys Cave No. 1 (type locality).

Pseudanophthalmus sanctipauli Barr (TB)

Virginia. — Russell Co.: Banners Corner Cave (type locality). Scott
Co.: Greears Sweet Potato Cave.

Pseudanophthalmus vicarius Barr (TB)

Virginia. — Tazewell Co.: Bowens, Cauliflower, Fallen Rock, Gully,
Hugh Young (type locality), and Lost Mill No. 3 caves.

Pseudanophthalmus sp. A (T. C. Barr, Jr., in ms.) (TB)

Virginia. — Russell Co.: Banner Cave.

Pseudanophthalmus sp. B (T. C. Barr, Jr., in ms.) (TB)

Virginia. — Russell Co.: Indian Cave.

hypolithos group

Pseudanophthalmus praetermissus Barr (TB)

Virginia. — Scott Co.: Kerns Cave No. 1 (type locality).

jonesi group

Pseudanophthalmus cordicollis Barr (TB)

Virginia. — Wise Co.: Little Kennedy Cave (type locality).

Pseudanophthalmus longiceps Barr (TB)

Tennessee. — Hancock Co.: Panther Creek Cave.

Virginia. — Lee Co.: Fisher Cave (type locality).

Pseudanophthalmus pallidus Barr (TB)

Tennessee. — Claiborne Co.: Buis Saltpetre, Chadwells (type
locality), and English caves.

Pseudanophthalmus seclusus Barr (TB)

Virginia. — Scott Co.: Alley, Cox Ram Pump, Flannery (type
locality), Hill, Kerns No. 1, McDavids, and Pond caves.

Invertebrate Cave Fauna

Pseudanopht halmus thomasi Barr (TB)

Virginia. — Scott Co.: Blair-Collins (type locality) and Coley No. 2
caves.

Pseudanophthalmus sp. A (T. C. Barr, Jr., in ms.) (TB)

Virginia. — Scott Co.: Greears Sweet Potato Cave.

Pseudanophthalmus sp. B (T. C. Barr, Jr., in ms.) (TB)

Tennessee. — Campbell Co.: Valley View’ Cave.

petrunkevitchi group

Pseudanophthalmus hoffmani Barr (TB)

Virginia. — Bland Co.: Coon, Hamilton, Newberry-Bane, and Repass
Saltpetre caves. Smyth Co.: Beaver Creek, Buchanan Saltpetre
(type locality), and Marble caves.

Pseudanophthalmus hortulanus Barr (TB)

Virginia. — Tazewell Co.: Cassell Farm Cave No. 2 (type locality).

Pseudanophthalmus petrunkevitchi Valentine (TB)

Virginia. — Page Co.: Woods Cave. Warren Co.: Skyline Caverns
(type locality).

Pseudanophthalmus sp. A (T. C. Barr, Jr., in ms.) (TB)

Virginia. — Washington Co.: Brass Kettle Hole Cave.

Pseudanophthalmus sp. B (T. C. Barr, Jr., in ms.) (TB)

Virginia. — Wythe Co.: Cave School Water, Pickett, and Sam Six
caves.

Pseudanophthalmus sp. C (T. C. Barr, Jr., in ms.) (TB)

Virginia. — Pulaski Co.: Sam Bells Cave.

pusio group

Pseudanophthalmus nelsoni Barr (TB)

Virginia. — Alleghany Co.: Arritt Mill Tunnel (type locality) and
Blue Springs (?) caves.

Pseudanophthalmus pontis Barr (TB)

Virginia. — Rockbridge Co.: Buck Hill Cave (type locality).

Pseudanophthalmus punctatus Valentine (TB)

Virginia. — Giles Co.: Clover Hollow7, Smokehole, Spruce Run
Mountain, and Tawneys (type locality) caves.

Pseudanophthalmus pusio (Horn) (TB)

Virginia. — Montgomery Co.: Agnew, Aunt Nellies, Erhart (type
locality), Fred Bulls, Old Mill, Slussers Chapel, and Thorn Hill
caves. Roanoke Co.: Goodwins Cave.

Pseudanophthalmus sp. A (T. C. Barr, Jr., in ms.) (TB)

Virginia. — Rockbridge Co.: Showalters Cave.

tennesseensis group

Pseudanophthalmus unionis Barr (TB)

Tennessee. — Union Co.: Wolf (type locality) and Wright caves.

Rhadine caudata (TeConte) (TP)

Virginia. — Roanoke Co.: Dixie Caverns.

John R. Holsinger and David C. Culver

Stenolophus ochropezus (Say) (TX)

Virginia. — Scott Co.: Coley Cave No. 2.

Trechus ( Trechus ) hydropicus canus Barr (TX)

Virginia. — Lee Co.: Bowling Cave.

Other Carabidae

Other species occasionally collected from study-area caves, where
they probably occurred as accidentals, include: Bradycellus sp.,
Harpalus compar , Platynus decens, P. extensicollis, P. gratiosus, P.
punctiforme , P. retractus, Pterostichus lucublandus, and Tachys
( Tachyura ) sp.

Family Leiodidae
Catops sp.

Virginia. — Giles Co.: Spruce Run Mountain Cave.

Nemadus horni (?) Hatch (TX?)

Virginia. — Lee Co.: Gilley and Sweet Potato caves.

Prionochaeta opaca Say (TX)

Virginia. — Smyth Co.: Stones Cave No. 2.

Family Pselaphidae
Arianops jeanneli Park (TB)

Virginia. — Lee Co.: Gilley Cave (type locality).

Batriasymmodes greeveri Park (TB)

Tennessee. — Sullivan Co.: Potters Cave (type locality).
Batriasymmodes monstrosus (LeConte) (AC)

Virginia. — Lee Co.: Poor Farm Cave.

Batrisodes globosus (LeConte) (TX)

Virginia. — Lee Co.: Sweet Potato Cave.

Family Staphylinidae
Aleochara lucifuga (Casey) (TP)

Tennessee. — Claiborne Co.: English Cave.

Virginia. — Bland Co.: Hamilton Cave. Frederick Co.: Ogdens Cave.
Lee Co.: Elys Moonshine, Gilley, Lucy Beatty, Smith, and Sweet
Potato caves. Page Co.: “Mushroom Cave” (Klimaszewski
1984:93). Roanoke Co.: Goodwins Cave. Russell Co.: Dickenson
Cave. Smyth Co.: Roberts and Sugar Grove No. 10 caves.
Tazewell Co.: Cassell Farm Cave(s). Wise Co.: Wildcat Saltpetre
Cave. Wythe Co.: Sam Six Cave.

Aloconota insecta (Thomson) (TP)

Virginia. — Botetourt Co.: Thomas Cave. Lee Co.: Bowling Cave.
Page Co.: Ruffners Cave No. 1. Roanoke Co.: Hodges Cave No.
1. Rockingham Co.: Endless Caverns. Russell Co.: Banners
Corner Cave. Tazewell Co.: Fallen Rock and Gully caves.
Washington Co.: Hall Bottom Cave No. 1.

Invertebrate Cave Fauna

Atheta annexa Casey (TP)

Virginia. — Giles Co.: Giant Caverns. Montgomery Co.: Old Mill
Cave. Roanoke Co.: Goodwins Cave. Shenandoah Co.: Battlefield
Crystal Cave. Smyth Co.: Stones Cave No. 2.

Atheta troglophila Klimaszewski and Peck (TP)

Virginia. — Lee Co.: Lucy Beatty, Smith, and Young-Fugate caves.
Aleocharinae (undetermined genus and species)

Virginia. — Augusta Co.: Glade Cave.

Brathinus nitidus LeConte (TP or TX)

Virginia. — Lee Co.: Bowling Cave. Russell Co.: Banners Corner
Cave. Scott Co.: Coley No. 2 and Flannery caves.

Quedius { Microsaurus ) erythrogaster Mannerheim (TP)

Virginia. — Bath Co.: Porters and Roy Lyle caves. Giles Co.: Giant
(Caverns), Harris and Straleys No. 1 caves. Highland Co.:
Marshalls Cave. Lee Co.: “Cave No. 1 and No. 3, Pennington
Gap” (Smetana, 1971:85), and Indian Cave. Rockbridge Co.:
Doll House Cave. Rockingham Co.: Melrose Cave. Scott Co.:
Sounding Cave. Shenandoah Co.: Hensleys and Shenandoah
Wild caves. Smyth Co.: Buchanan Saltpetre Cave.

Quedius {Microsaurus) mesomelinus (Marsham) (TP or TX)

Virginia. — Pulaski Co.: Sam Bells Cave.

Quedius {Microsaurus) spelaeus Horn (TP)

Virginia.— Bath Co.: Cave Run Pit and Crossroads caves.
Frederick Co.: Ogdens Cave. Giles Co.: Tawneys Cave. Pulaski
Co.: Fifty-Foot Hell Cave. Rockingham Co.: Three-D Maze
Cave.

Quedius sp.

Virginia. — Augusta Co.: Grand Caverns and Madisons Saltpetre
Cave. Lee Co.: Young-Fugate Cave. Page Co.: Luray Caverns.
Rockingham Co.: Massanutten Caverns. Warren Co.: Skyline
Caverns.

Other Staphylinidae

Nine other genera, based largely on single records and presumably
including mostly accidental species, are recorded from study-area
caves as follows: Cratarea, Emplenota, Erichsonius, Homaeotarsus,
Lathrobium , Lesteva (probably L. pallipes LeConte, a common
trogloxene), Megalinus, Philonthus, and Trichophya (probably T.
pilicornis Gyllenhal).

Order Diptera

With the possible exception of one species, there are no troglobitic
dipterans (flies) in the study area. Several species are relatively common
in caves, however, and sometimes make an important contribution to

John R. Holsinger and David C. Culver

the cavernicolous fauna. At least six families are found in regional caves
with some degree of regularity: Heleomyzidae, Mycetophilidae, Phor-
idae, Psychodidae, Sciaridae, and Sphaeroceridae. In addition, seven
other families are sporadically observed, usually in entrance zones; but
they rarely, if ever, contribute significantly to the fauna of a given cave.
These are Calliphoridae, Chironomidae, Culicidae, Dolichopodidae,
Empidae, Streblidae (e.g., Trichobius , an ectoparasite of bats), and
Tipulidae. Although no attempt was made to systematically collect
dipterans from caves, a few collections were made selectively to establish
the identity of the most common species.

The fly most frequently seen in regional caves was Amoebalaria
defessa (Heleomyzidae), a troglophile or trogloxene common in caves
throughout much of the eastern United States (see Gill 1962, Peck and
Lewis 1978) (Fig. 31B). Two other heleomyzids, Aecothea (probably A.
specus Aldrich) and Heleomyza brachyptera (Loew), were observed
occasionally , but specific cave records are unavailable. Heleomyzids are
generally found on damp walls and ceilings, sometimes in large numbers
and usually not far from entrance zones (see also Busacca 1975).

Also relatively common in study-area caves are Megaselia caverni-
cola (Phoridae), a troglophile widespread in the east-central and eastern
United States (see Borgmeier 1965), and members of the Sphaeroceridae,
of which several species are often found in caves of the United States
(see Curran 1965, Stone et al. 1965, Marshall 1985). Although the
sphaerocerid Spelobia tenebrarum is recorded from only two caves in
Lee County, it probably inhabits many other caves in the study area.
This species is recorded from numerous caves in the eastern United
States and has been listed as a troglophile or trogloxene under the name
Leptocera tenebrarum by a number of workers (viz., Barr 1967a,
Holsinger and Peck 1971, Holsinger et al. 1976, Peck and Lewis 1978).
However, in a recent study of cavernicolous sphaerocerids, Marshall
and Peck (1984, 1985) suggest that it may be a troglobite. Both
Megaselia and Spelobia are associated with decaying organic material
(e.g., vegetal matter, feces, and carcasses) in caves.

Both larvae and adults of fungus gnats (families Sciaridae and
Mycetophilidae) are recorded from regional caves. Sciarids are usually
found in and around damp, rotting vegetal debris. Mycetophilid larvae
are sometimes luminescent and build silken webs on dung and damp
clay and under rocks. Peck and Russell (1976) identified the myceto-
philid Macrocera nobilis Johnson from many caves in the southeastern
United States, but none of these records is from the study area.
Although M. nobilis should occur in study-area caves on the basis of its
geographic distribution, most of the larvae seen to date have been
smaller than those of this species and probably represent other genera.

Invertebrate Cave Fauna

Fig. 31. Terrestrial cavernicoles from the study area (approximate body lengths
in parentheses): A, collembolan, Tomocerus bidentatus (3 mm); B, fly,
Amoebalaria defessa (6 mm); C, dipluran, Litocampa sp. (7 mm) (courtesy of
L. M. Ferguson); D, milliped, Pseudotremia nodosa (15 mm); E, terrestrial
isopod, Amerigoniscus henroti (6 mm).

John R. Holsinger and David C. Culver

Family Heleomyzidae

Amoebalaria defessa (Osten-Sacken) (TP or TX)

Tennessee. — Claiborne Co.: Buis Saltpetre and Keck No. 1 caves.

Grainger Co.: Horseshoe Cave. Hancock Co.: Subers Cave.
Virginia. — Bland Co.: Hamilton Cave. Craig Co.: Carpers and
Rufe Caldwell caves. Frederick Co.: Ogdens Cave. Giles Co.:
Ballards Cave. Lee Co.: Gallohan No. 1, Olinger, Roadside No.
1, and Young-Fugate caves. Montgomery Co.: Vickers Road
Cave. Pulaski Co.: Fifty-Foot Hell Cave. Rockbridge Co.:
Showalters Cave. Rockingham Co.: Massanutten Caverns.
Russell Co.: Banners Corner and Campbells Spring caves. Scott
Co.: Blair-Collins and Hill caves. Smyth Co.: Tilson Saltpetre
Cave. Tazewell Co.: Fallen Rock, Hugh Young, and Steeles
caves.

Family Mycetophilidae
Genus (?) species (?)

Tennessee. — Claiborne Co.: Jennings Cave.

Virginia. — Bath Co.: Porters Cave. Lee Co.: Cumberland Gap
Saltpetre Cave. Rockbridge Co.: Doll House Cave.

Family Phoridae

Megaselia cavernicola (Brues) (TP)

Virginia. — Lee Co.: Gallohan No. 1, Molly Wagle, Smith, and
Sweet Potato caves. Page Co.: Luray Caverns. Smyth Co.:
Tilson Saltpetre Cave. Tazewell Co.: Lawson Cave.

Family Psychodidae
Psycho da sp.

Virginia. — Lee Co.: Sweet Potato Cave.

Family Sciaridae

Brady sia luravi (Johannsen) (TX or AC)

Virginia. — Page Co.: Luray Caverns.

Brady sia sp.

Virginia. — Lee Co.: Gallohan No. 1, Molly Wagle, Smith, and
Sweet Potato caves. Smyth Co.: Tilson Saltpetre Cave.

Pnyxia scabiei (Hopkins) (AC)

Virginia. — Page Co.: Luray Caverns.

Sciara (?) sp.

Tennessee. — Claiborne Co.: English Cave.

Virginia. — Tazewell Co.: Lawson Cave.

Family Sphaeroceridae
Leptocera pararoralis (?) Duda (TX?)

Virginia. — Lee Co.: Gallohan No. 1 and Sweet Potato caves.

Invertebrate Cave Fauna

Fig. 32. Terrestrial cavernicoles from the study area (approximate body lengths
in parentheses): A, beetle, Pseudanophthalmus delicatus (4 mm); B, cricket,
Euhadenoecus fragilis (15 mm).

Spelobia semioculata (Richards) (TX?)

Virginia. — Lee Co.: Smith Cave. Shenandoah Co.: Maddens Cave.
Spelobia tenebrarum (Aldrich) (TP?)

Virginia. — Lee Co.: Molly Wagle and Sweet Potato caves.

Spelobia (?) spp.

Virginia. — Page Co.: Luray Caverns. Shenandoah Co.: Shenandoah
Caverns. Smyth Co.: Tilson Saltpetre Cave.

John R. Holsinger and David C. Culver

ECOLOGY

Sources of Food

Aside from darkness, the most striking feature of most caves is the
scarcity of food. Except for a few chemosynthetic autotrophic bacteria
that use iron and sulfur as an electron donor, primary producers are
absent. Thus, in a general sense, cave communities are decomposer
communities. Allochthonous food is brought in by both biological and
physical agents in different amounts, continuously or in pulses, and in
different spatial configurations. These differences affect the kind of
species present, so that it is useful to review them.

In the terrestrial biotope, there are at least five major sources of
food: (1) bat guano, (2) cave cricket eggs and guano, (3) microorganisms,
(4) mammalian feces and dead animals, and (5) plant detritus left by
flooding. A few caves harbor large bat colonies with large guano
concentrations beneath the roosting sites. In this case food is abundant,
and the fauna feeding on guano is quite different from the rest of the
cave fauna (Harris 1970). Caves with large bat populations are rare in
Virginia and east Tennessee and have not been studied with respect to
their invertebrate communities. Small piles of bat guano rarely seem to
have any macroscopic fauna. Perhaps this is because no species are
present that are physiologically equipped to digest bat guano.

A major source of food input comes from the cave crickets in the
genus Euhadenoecus. These crickets regularly leave the cave at night
and feed “opportunistically and omnivorously as a scavenger” (Hubbell
and Norton 1978), eating the vast majority of their food outside the
cave. The females oviposit inside the cave, usually in sandy substrates.
In parts of the Edwards Plateau of Texas and the Interior Low Plateaus
of Kentucky, cave-cricket eggs are the major dietary item for some
species of beetles. This fascinating interaction has been extensively
studied (Culver 1982) because the cricket-beetle interaction comes close
to being a naturally isolated predator-prey pair. This facilitates study of
morphological, behavioral, and demographic characteristics because
selective pressures are relatively simple and clear-cut. We have found no
evidence that cricket eggs form a major part of the diet of any beetles in
Virginia and east-Tennessee caves. We suspect that this interaction is
absent because sandy substrates are rare in Appalachian Valley caves
and Euhadenoecus species oviposit in substrates difficult for beetles to
excavate. Cricket guano, on the other hand, is an important food
source. Some of the most diverse terrestrial communities occur in areas
where cricket guano is spattered on walls and floors. We suspect that in
many caves it is a major source of food, either directly, or indirectly by
serving as a substrate for microflora.

Microorganisms occur on a variety of substrates, including wood,
dung, and plant detritus. At least part of the diet of many terrestrial

Invertebrate Cave Fauna

cave invertebrates is microorganisms (see below). The richest sources of
microorganisms are dung near entrances and decaying arthropod
remains in aphotic passages (Dickson and Kirk 1976). Fungi are more
concentrated and patchily distributed than bacteria, and fungi are also a
more important food source, perhaps because they are concentrated.

Besides serving as a substrate for microfungi, dung and dead
animals are important food in their own right and attract a wide variety
of invertebrates. Peck (1973) has also used human dung as a very
effective bait for cave invertebrates.

Plant detritus may also be an important food source. A layer of
mud and finely divided leaves, often rich in oligochaetes, is deposited in
many caves by slowly receding floodwaters. Such areas often have a rich
fauna. In caves subject to severe, rapid flooding, piles of twigs and
leaves are left behind. On these resource patches is a relatively distinct
fauna that will be described below.

Food in streams is almost entirely allochthonous in origin. Stream
detritus is usually divided into coarse particulate organic matter (CPOM
> 1 mm) and fine particulate organic matter (0.0005 mm < FPOM < 1
mm) (Cummins and Klug 1979). By convention, organic matter smaller
than 0.0005 mm is considered dissolved (DOM). CPOM is a substrate
for microorganisms.

Resource Levels

Although there is a great deal of indirect evidence of food scarcity
in caves, there have been few direct measurements of resource levels in
caves. Many of the physiological and morphological changes associated
with isolation in caves (reviewed by Culver 1982) make sense only in the
context of a relatively stable, food-poor environment. It is obvious to
anyone visiting a cave that at least the standing crop of resources is very
low indeed.

Dickson and Kirk (1976) have provided direct evidence from Old
Mill Cave in Montgomery County. They found that, for the most part,
resource levels were lower in the cave than in forest soil, but there are
exceptions. Dung in the entrance and mud floors with chitin remains
had high plate counts. Thus food is scarce and very patchy. Dickson
and Kirk (1976) also found that fungi are correlated with abundance of
the terrestrial macrofauna whereas bacteria are not. This may help
explain why wet passages have more fauna than dry passages, where
fungi are relatively uncommon.

There remains the question of how much food is actually available
to cave animals. The best comparative study is that of Peck and
Richardson (1976), who compared stomach contents of the “cave
salamander” Eurycea lucifuga Rafinesque from entrance and dark zones
of caves in Tennessee and Alabama. Salamanders collected at the

Table 2. Fauna associated with discrete habitats with abundant resources — dung, wood, and plant detritus. Troglobites
indicated by an asterisk.

John R. Holsinger and David C. Culver

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John R. Holsinger and David C. Culver

entrance, where one would expect food to be more abundant, had 19.6
prey items with a volume of 0.14 ml per stomach. By contrast
salamanders from dark zones had only 3.4 prey items with a volume of
0.05 ml per stomach.

Diets

Our knowledge of the diet of cavernicoles is woefully inadequate.
While food webs are not available for most communities (see Cohen
1978) and most published food webs are fanciful, the problem is
particularly acute for cave communities. Packard (1888) pointed out
nearly a century ago, that “cave animals, even the carnivorous species,
take remarkably little food.” Contemporary ecological theory predicts
that a species faced with scarce resources should increase the range of
foods taken. That is, one would expect little specificity in diet. One
obvious fact about cave animals is that even their surface relatives tend
to be omnivorous.

Many terrestrial cavernicoles feed directly on dead and decaying
organic matter and associated microorganisms. Except for crickets,
millipeds are usually the most abundant terrestial cavernicoles. Millipeds
are frequently found on dead and decaying organic matter, and they
often ingest rotting wood (Shear 1969). Other invertebrates that feed
directly on dead and decaying organic matter include staphylinid beetles,
isopods, and dipterans. Collembolans apparently concentrate on micro-
fungi (Christiansen 1964b).

Carnivores are also very catholic in their diets. Around cave
entrances the orbweb-building spider Meta menardi is often common; it
captures a variety of flying insects, especially Diptera. Cantharid beetle
larvae may also be important predators in entrances (Peck 1975b). In
the dark zones, nesticid and linyphiid spiders construct several small
sheet webs in which they capture a variety of walking invertebrates.
Other web-builders are likely to be in the study area — the larvae of the
fungus gnat Macrocera nobilis (Peck and Russell 1976). These larvae
build webs in which they catch mostly other dipterans, but they also
feed on dead organic matter while constructing their webs (Peck and
Russell 1976). Trechine beetles, which are often common in caves in the
Interior Low Plateaus, are generally uncommon in Virginia and east
Tennessee. In common with the small species of Pseudanopthalmus in
the Interior Low Plateaus, species of this genus in the study area
probably eat collembolans, small oligochaetes in the mud, diplurans,
and small diplopods (Barr 1968, Keith 1975, McKinney 1975). Even less
is known about other invertebrate predators, such as opilionids,
pseudoscorpions, and rhagidiid mites; but they probably feed mainly on
Collembola, spider eggs, and small spiders. Finally, the salamanders,
Eurycea lucifuga and Gyrinophilus prophyriticus eat a wide range of
invertebrates.

Invertebrate Cave Fauna

Most cavernicoles in streams feed directly on detritus and its
associated microorganisms. The epigean amphipod Gammarus pseudo-
limnaeus Bousfield is a facultative shredder (Cummins and Klug 1979),
preferring CPOM, but also using FPOM and DOM. Cave-stream
amphipods are probably similar in this regard. No direct information on
isopods is available, but Estes (1978) suggests that Lirceus usdagalun
tends to eat CPOM whereas Caecidotea recurvata tends to eat DOM.
On the basis of their size, crayfish are probably shredders, and snails
and lumbriculid worms probably ingest DOM.

The diet of cave flatworms is more problematical than that of
crustaceans. Mitchell (1974) has demonstrated that Texas cave flat-
worms ( Sphalloplana sp.) eat injured and moribund amphipods and
crickets. Holsinger (1966), on the other hand, suggested that flatworms
eat tubificid worms. The greatest concentrations of flatworms that we
have observed in cave streams have been in stream pools with no
amphipods or isopods. We suspect that flatworms feed on small
oligochaetes and perhaps on microorganisms.

The primary stream predator is larval Gyrinophilus porphyriticus.
These salamander larvae are voracious feeders on amphipods and
isopods (Culver 1973b, 1985) and appear to be exclusively predaceous.
They are generally limited to caves with high densities of amphipods
and isopods.

The amphipods and isopods occurring in drip pools and in deep
phreatic lakes ingest the organically rich mud. Guts of animals from
these habitats are often filled with mud, as can be seen in the photograph
of the cirolanid isopod Antrolana lira (Fig. 13A). Dickson (1975) shows
that abundance of Crangonyx antennatus in pools is correlated with
abundance of microfungi. A tentative food web for pool habitats in
Banners Corner Cave is shown in Figure 33, based on Holsinger’s (1966)
study.

Habitats

Because of the scarcity and patchiness of resources, terrestrial
cavernicoles are often concentrated on discrete patches of dung, wood,
and plant detritus. Examples of the fauna found in these habitats are
listed in Table 2. The most interesting pattern that emerges from Table
2 is that the frequency of troglobites is lowest on dung (20%), slightly
higher on patches of plant detritus (28%), and much higher on wood
(76%). The difference between wood and the other habitats is highly
significant (G = 9.44 P > 0.99). As Poulson (1978) pointed out,
resources with high caloric value and low residence time, such as dung,
should have a high frequency of vagile troglophiles with relatively high
reproductive rates, compared with long-lasting resources having low
caloric value, such as wood.

Cavernicoles also are found in habitats that are less discrete, where
resources are more or less homogeneously distributed over a larger

John R. Holsinger and David C. Culver

area. The most notable example of this is mud banks, which are often
near streams. The faunas from three such habitats are listed in Table 3.
In Fallen Rock Cave, a layer of finely divided detritus rich in oligochaetes
was present, and in the other two caves there were spatterings of cricket
guano. In all three caves there was a high proportion of troglobites,
ranging from 50% in Tazewell Saltpetre Cave to 73% in Gallohan Cave
No. 1. In these situations, low resource density probably puts trog-
lophiles at a disadvantage.

Most cave streams have an alternating riffle-pool structure (shallows
and deeps) that is characteristic of stony-bottomed streams in general.
Most stream-dwelling cavernicoles prefer riffles for several reasons.
First, water in riffles is well oxygenated. Second, riffles serve as traps
for leaf litter, thus increasing availability of food resources. Species
characteristic of riffles are the isopods Caecidotea and Lirceus, the
amphipod Crangonyx antennatus, and the snails Fontigens. There is a
finer division of the riffle habitat. In general, small individuals tend to
be under small rocks, which are deep in the riffle. In a study of two
caves in southwestern Virginia, Estes (1978) found that the small
Crangonyx antennatus was under small rocks and gravels, but the larger
Lirceus usdagalun and Caecidotea recurvata tended to be under large
rocks and gravels.

A few species are concentrated in steam pools. Flatworms seem to
be more common in pools, where they glide along the surface of the
water, than in riffles; but no quantitative data exist on this point. Larval
Gyrinophilus prophyriticus are concentrated in pools, where it is
relatively easy for them to detect prey movements (Culver 1975).

Drip pools are the habitat of most species of Stygobromus, although
occasionally some (especially Stygobromus mackini) are found in
streams (Holsinger 1978). Crangonyx antennatus also occurs in pools
(Dickson 1977a), where it frequently constructs shallow burrows,
apparently to avoid desiccation during droughts (Holsinger and Dickson
1977). Burrowing behavior may be widespread in Stygobromus as well,
but this point has not been investigated.

Life Histories

Relatively little work has been done on life histories of cavernicoles
in the study area. Nonetheless, because of the importance of the subject,
a brief overview of the problem will be presented. A more comprehensive
treatment is given by Culver (1982). In general three sorts of comparisons
have been made. First, cavernicoles have been compared with epigean
species (Ginet 1960, Rouch 1968). While comparisons have not always
been made with phyletically similar species, these comparisons usually
show striking differences between cave and epigean species. Second,
cavernicoles of different ages in caves have been compared. Age in caves

Invertebrate Cave Fauna

Fig. 33. Food web for Banners Corner Cave, Russell County, Va. The dotted
lines indicate feeding by flatworms on injured and moribund amphipods and
isopods.

Gammorus pulex

'l' Fertilized egg
O Release from marsupium
Eggs in marsupium
if Sexual maturity
if Death

Niphargus orcinus virei

★

■if

YEARS °

3 4 5 6

Fig. 34. Life history comparison of the troglobitic amphipod Niphargus orcinus
virei with the epigean amphipod Gammarus pulex. Modified Irom Ginet ( 1960).

Table 3. Fauna associated with diffuse food supplies. Troglobites indicated by an asterisk.

100

John R. Holsinger and David C. Culver

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Invertebrate Cave Fauna

101

is usually measured by the level of regressive evolution (Poulson 1963).
Third, cavernicoles in different cave habitats have been compared
(Dickson and Holsinger 1981, Estes 1978).

Selection for delayed reproduction, increased longevity, and the
like is frequently called K-selection. Recent models (reviewed by
Charlesworth 1980) show that there is no simple dichotomy between
r-selection and K-selection, but the following characteristics are likely to
„ be selected for in at least some cavernicoles:

1. Delayed maturity

2. Increased longevity

3. Fewer clutches

4. Smaller clutch size

5. Larger eggs

6. Low percentage of mature females ovigerous

7. Sex ratio skewed toward females

Examples of each of these characteristics will be discussed, but their
generality will not.

Ginet’s (1960) study of the amphipod Niphargus orcinus virei
Chevreux illustrates the first three characteristics (Fig. 34). Compared
with Gammarus duebeni Lilljeborg, N. orcinus virei takes four times
longer to mature, lives five times longer, and has only two broods (as
opposed to seven). The most striking example of increased longevity is
the crayfish Orconectes australis australis (Rhoades), which lives a
minimum of 40 years (Cooper 1975). The amphipod Crangonyx
antennatus, which is common in many Virginia and Tennessee caves,

lives at least 6 years in the laboratory. Life spans of terrestrial animals
are apparently shorter. Peck (1975c) found that the leiodid beetle
Ptomaphagus hirtus Tellkampf lived an average of 2 years.

Rouch (1968), in a comprehensive study of cave harpacticoid
copepods in France, found that cavernicolous species had fewer eggs per
unit size of female and that egg diameters were larger than was the case
for low-altitude epigean species. High-altitude epigean harpacticoids,
like cavernicolus species, also had fewer eggs with larger diameters when
compared with low-altitude epigean species.

Many populations of cavernicoles have low percentages of ovigerous
females and mature females. One very clear case is Dickson’s data
(Dickson and Holsinger 1981) on Crangonyx antennatus from two
caves in Lee County, Va. (Fig. 35). These data raise some interesting
evolutionary questions. If ovigerous females are genetically different
from non-ovigerous females, then those genotypes should increase,
resulting in higher frequencies of ovigery. Low ovigery may be
maintained by strong intraspecific competition, with little genetic basis,
which allows only an occasional female to take in enough food to

102

John R. Holsinger and David C. Culver

reproduce. Alternatively, low rates of ovigery may be maintained by
group selection, with populations with excessively high reproductive
rates becoming extinct.

Apparently in response to low population densities, some aquatic
species have evolved to the point where males are quite rare. We suspect
that this is accompanied by some form of parthenogenesis (Culver and
Holsinger 1969), but the genetics have not been studied. The isopod
Lirceus usdagalun has a sex ratio favoring females by at least three to
one (Estes 1978); and in the amphipod Crangonyx antennatus , male
frequency is positively correlated with density (Dickson and Holsinger
1981). However, male rarity is most strikingly developed in the
amphipod genus Stygobromus . Cave populations usually have sex
ratios of the order of 10 females to every male (Culver and Holsinger
1969).

Population Size and Stability

For populations undergoing K-selection, increased efficiency,
reduced clutch size, and the like should result in an increase in the carry-
ing capacity, and thus increase population size. The best comparative
data are from Poulson’s (1963) study of the amblyopsid fishes in caves
in the Interior Low Plateaus. There is an unambiguous increase in pop-
ulation size with increasing morphological adaptation. Considering life-
history characteristics, however, Amblyopsis spelaea De Kay would be
expected to have a high population size because its growth rate is low.
In a less comprehensive study of the isopods Lirceus usdagalun and
Caecidotea recurvata , Culver (1976) found that L. usdagalun had a lower
carrying capacity than did the more specialized C. recurvata.

However, there is considerable doubt that those species with the
longest evolutionary history in caves have the largest population sizes.
Although actual population sizes are determined by interspecific inter-
actions as well as carrying capacity, the intensity of these interactions
should also be under evolutionary control. Culver (1976) showed that
the intensity of competition between any pair of species declined through
evolutionary time but that success in competition did not increase.
Thus, the troglophilic amphipod Gammarus minus (Form I) is much
more common than the troglobitic amphipod Stygobromus mackini and
the isopod Caecidotea richardsonae in the caves of the Ward Cove
karst in Tazewell County. In the milliped genus Pseudotremia, troglo-
philes can be as abundant as troglobites. Comparatively more special-
ized millipeds in the genus Trichopetalum are always less common than
those in Pseudotremia. Many troglobites, such as pseudoscorpions, are
always rare. These observations suggest that rarity per se is advanta-
geous, since the population avoids increases that lead to crashes and
extinction. The most likely mechanism for evolution of rarity is group
selection. This conclusion is supported by the fact that population
extinctions are known to occur (Culver 1970).

% of Amphipods mature

Invertebrate Cave Fauna

103

100

August

November

February

June

Fig. 35. Frequency of males that are mature and frequency of females that are
mature for Crangonyx antennatus. From Dickson and Holsinger (1981).

104

John R. Holsinger and David C. Culver

Interspecific Interactions — General

Compared with other ecological problems, interspecific interactions
among cavernicoles have received considerable attention. Cave
communities offer several advantages to the student of species
interactions. First, cave environments and population sizes are relatively
stable compared with most epigean populations. Thus cave communities
correspond more closely to the assumptions of ecological models than
do most communities. Second, the number of species is so small that all
interactions can be studied. Third, because there are many caves with
very similar environmental conditions, there are many replicates as well
as many “natural experiments” where species composition is different.
The main disadvantage of cave communities is the long generation time
of species, which makes the study of long-term dynamics difficult.

Mutualism

No free-living mutualists have been reported from caves. Mutual-
istic gut endosymbionts may be important, but they have not been
studied. Two ectosymbionts on crayfish are known, branchiodbellid
worms and entocytherid ostracods, both of which occur in the study area.
Hobbs (1975) has studied ostracod symbionts in Iqdiana caves, and his
work shows that cave crayfish may be convenient systems for the study
of symbiosis. The ecosymbiotic entocytherids attach to the exoskeleton
of crayfish and feed on microorganisms and detritus that accumulate on
the host exoskeleton.

The troglobitic crayfish Orconectes inermis Cope has several
advantages for further study. First, more than 90% of the ostracods on
Orconectes inermis inermis belonged to one species, Sagittocythere barri
(Hart and Hobbs), which in turn was rarely found on other crayfish in
Hobbs’ study area. Thus it is essentially a two-species system. Second,
the ectosymbionts are common. More than half of the O. inermis popu-
lations had ostracods. The average number of S. barri per crayfish was
17.2 (Hobbs 1975). One of Hobbs’ most interesting findings was that
there was a significant increase in the number of ostracods with increas-
ing length of the crayfish carapace. Thus the crayfish are like islands
that are colonized by ostracods.

Predation

While predation appears to be more frequent in the terrestrial
fauna than in the aquatic fauna, there have been no ecological or behav-
ioral studies of terrestrial predators in the study area. One aquatic pred-
ator, larval Gyrinophilus porphyriticus, has been extensively studied
(Culver 1973b, 1975, 1985). This work is summarized below.

Gyrinophilus porphyriticus larvae form significant populations in
stream pools in several caves in the Powell Valley of Claiborne and Lee
counties. Their prey are amphipods and isopods. None of the larvae

Invertebrate Cave Fauna

105

reacted to dead amphipods and isopods, but when a live amphipod or
isopod was put in the water, the larva would raise itself on its front legs
and usually its hind legs as well. It then remained motionless until the
prey came within 2 to 4 cm of its snout. Then, with rapid sucking
action, the salamander ate the prey item. Larvae will also attack the
ends of small brushes moved slowly through the water, indicating that
mechano-reception is the primary method of prey detection.

The behavior of the larvae toward prey is remarkably uncompli-
cated. The functional response is linear over a wide range of prey densi-
ties (Fig. 36), a wider range than is normally encountered in the field.
This linearity is apparently due to the very short handling time and the
larvae’s prodigious appetites. One larva ate eight Caecidotea recurvata
in 30 minutes in the laboratory.

Although G. prophyriticus attempts to capture any Caecidotea
recurvata or Crangonyx antennatus that comes close to its snout, it is
about three times as successful at capturing C. recurvata, apparently
because C. antennatus often avoids predation by swimming out of
range. Actual predation rates also depend on the fraction of the popula-
tion accessible to predators. Because nearly all G. porphyriticus larvae
are in pools rather than riffles, and relatively still water aids prey detec-
tion, individuals in riffles and on flowstone are inaccessible to preda-
tors. In McClure Cave in Lee County, it was found that the actual
predation rates (proportional to success rate times the proportion
accessible to predation) did not significantly differ between the two
species (Culver 1975). This is probably coincidental, but it does facilitate
modeling the system. Using standard competition and predation
equations, which is justified in part by the linear functional response, it
was predicted that both the density and frequency of C. antennatus
should increase. Even though it is preyed upon, the reduction in density
of its competitor C. recurvata more than compensates for predation
losses. Field data confirmed these predictions. Frequency of C. antennatus
increased from 0.09 away from larvae, to 0.44 in the immedidate vicinity
of larvae.

Salamander predation also resulted in habitat shifts of the prey.
With salamanders nearby, a greater frequency of both prey species was
found in riffle and flowstone “refuges.” Consequently, when salaman-
ders first invade a cave, a greater frequency of the prey population is
available. Potential predation rates are nearly double those at equili-
brium. Thus, invasion should be easy, but the establishment of a repro-
ducing population should be difficult. This prediction is also confirmed
by the data. Most “populations” are fewer than five individuals.

Competition

Competition is the most extensively studied and probably the most
important interaction in caves. Barr (1967b) and McKinney (1975) have

106

John R. Holsinger and David C. Culver

found it to be important in cave beetles, and Cooper (1975) discussed
the importance of competition among cave crayfish. The most extensive
study of competition has concerned peracarid crustacean communities
in southwest Virginia and east Tennessee caves (Culver 1973a, 1976,
1981, 1982; Dickson 1976, 1977a; Dickson and Holsinger 1981; Estes
1978; Estes and Holsinger 1982). Various combinations of three species
are present: Cacidotea recurvata, Crangonyx antennatus, and Lirceus
usdagalun.

The basis for competition is that, in the stream, amphipods and
isopods need a place to avoid the brunt of the current and a place to
feed on detritus washed into the cave. Because riffles are food-rich and
well oxygenated, amphipods and isopods congregate there, even though
high current velocities increase the risk of being washed out. Although
many stones in a riffle are unoccupied (Culver 1973a), amphipods and
isopods frequently meet because of jostling about by the current and
directed movement toward food. When two individuals meet, one is
almost invariably washed downstream. Some of these individuals die;
all are removed from the riffle.

Since washout is the basis of competition, it was possible to derive
a formula for measuring the competition coefficients and to measure
washout in an artificial stream in the laboratory. Competition coeffi-
cients (cxij) for all pairs of the three species studied were as follows:

C.a.

C.r.

L.u.

Crangonyx antennatus

\ >

0.99

1.32"]

Caecidotea recurvata

0.32

1.29

Lirceus usdagalun

L 1.16

0.49

i J

If these are adequate measures of competition and our reasoning above
has been correct, then the greater the competition, the greater the
microhabitat separation. Crangonyx antennatus and L. usdagalun,
which show the strongest competition, barely coexist in the same cave;
C. recurvata and L. usdagalun occupy different riffles; and C. antenna-
tus and C. recurvata occupy different-sized rocks in the same riffle.

Estes (1978) has closely examined microhabitat niche differences
among the three species and determined that L. usdagalun was almost
always found in current with a velocity/ depth ratio greater than 0.67,
whereas the other two species were found over a greater range of cur-
rent velocities. In addition, relative densities on different rock sizes var-
ied (Table 4). The large C. recurvata was under large rocks, and the
small C. antennatus was in gravel. Lirceus usdagalun was on an
intermediate substrate.

There is evidence that the intensity of competition between a pair
of species declines through evolutionary time. Culver (1976) showed

Invertebrate Cave Fauna

107

8-i

No.

Eaten

• •

-i 1 1

4 8 16

No. Offered

Fig. 36. Functional response of Gyrinophilus porphvriticus larva to
Caecidotea recurvata.

Table 4. Occurrence of Caecidotea recurvata , Lirceus usdagalun , and
Crangonyx antennatus under stones and gravel in areas of strong
current in Gallohan Cave No. 1 (modified from Estes 1978). The
first number is the mean 0.09m~ and the second is the relative
density with the highest density for each species given a value of 100.
Actual stone sizes were not given.

Species/ Individuals

Habitat

Large

Stones

Medium

Stones

Small

Stones

Gravel

C. recurvata 53

1.0(28)

3.63(100)

1.69(47)

2.67(74)

L. usdagalun 412

7.5(28)

21.75(82)

26.62(100)

16.67(63)

C. antennatus 13

0.0(0)

0.50(38)

0.62(47)

1.33(100)

that relative age of the interaction (as measured by the amount of regres-
sive evolution and speciation) and competition where negatively correlated
for Virginia and West Virginia peracarid crustacean communities.

Species Packing and Species Diversity

For most of the best-studied communities, species diversity is
limited by competitive interactions. Culver (1976) showed that competi-
tion theory predicts a maximum of three species. Three-species com-
munities in the Greenbrier Valley in West Virginia and in the Powell

108

John R. Holsinger and David C. Culver

Valley in Virginia were resistant to invasion, but two-species communi-
ties often were not. In the Powell Valley, L. usdagalun successfully
invaded a C. recurvata - C. antennatus community in Gallohan No. 2.
On the other hand, L. usdagalun in Thompson Cedar Cave is resistant
to invasion by L. recurvata, which is limited to a small section of stream
near the entrance. Both of these events are in agreement with the theory.

Salamander predators can have a large effect on species diversity.
In McClure Cave, relative density of C. antennatus increases near G.
porphyriticus larvae, primarily because its competitor C. recurvata is
also preyed upon. Refugia ensure the persistence of both prey. In Sweet
Potato Cave, G. porphyriticus eliminates C. recurvata from rimstone
pools, but C. antennatus escapes predation by burrowing (Holsinger
and Dickson 1977). In pools without salamander larvae, both prey spe-
cies persist.

Flooding has a major impact on both aquatic and terrestrial fauna.
Caves where water slowly recedes following regular flooding often have
a very rich terrestrial fauna as a result of the detritus left by receding
waters. Although no quantitative data are available, caves that have
much detritus often have a diverse fauna (Table 3). On the other hand,
caves that flood have a depauperate aquatic fauna. In a study of caves
of the Greenbrier Valley in West Virginia, Culver (1970) found that
caves that flood have 0.5 amphipod and isopod species (s = 0.7, n = 13),
while caves that do not flood have 2.3 species (s = 1.1, n = 15).

Finally, the island-like nature of caves has two important effects
(Culver 1976). First, the mean number of peracarid crustaceans is less
than the predicted three due to continuing extinctions. Second, the
regional faunal diversity is enhanced because patchiness allows coexist-
ence of competitors.

ZOOGEOGRAPHY
Drainage Basins and Regional Cave Faunas

The Appalachian Valley and Ridge in Virginia and northeastern
Tennessee is drained by seven major river systems or drainage basins as
indicated in Figure 2. Because each basin is well defined geographically
and contains topographically confined karst areas with a unique
assemblage of endemic cave species, we have chosen to treat them as
regional cave faunal units for the purpose of analyzing and discussing
zoogeographical relationships. Although some of the divides and inter-
fluves that separate these basins contain carbonate rocks (limestone and
dolomite) as indicated below, as a rule the major part of each basin is
enclosed by clastic rocks. The faunal units, which correspond to drain-
age basins of the same name, are: (1) Shenandoah, (2) James, (3) Roa-
noke, (4) New, (5) Holston, (6) Clinch, and (7) Powell. A very small
cavernous area in northern Highland County drained by the South

Invertebrate Cave Fauna

109

Branch of the Potomac River actually constitutes an eighth drainage
basin but has been excluded from our analysis because of its insignifi-
cant size in the study area. The cave fauna of this basin was discussed
previously by Holsinger et al. (1976).

In the companion study on the cave invertebrates of West Virginia
(Holsinger et al. 1976), we also divided that area into cave faunal units
that corresponded to major drainage basins and discussed zoogeo-
graphical relationships in a context similar to that of the present paper.
Somewhat similar, but broader, faunal units than those recognized in
the Virginias and northeastern Tennessee were distinguished for the
regional cave faunas of the Interior Low Plateaus by Barr (1967a),
northwestern Georgia by Holsinger and Peck (1971), and Illinois-
southeastern Missouri by Peck and Lewis (1978).

A list of the cave-limited species has been compiled for each basin
or faunal unit (Tables 5-1 1). Although these lists are restricted primarily
to troglobites, we have included a few select troglophiles that our obser-
vations indicate are commonly represented by cave-restricted pop-
ulations.

1. Shenandoah Basin. — This faunal unit includes that part of the
study area drained by the Shenandoah River and its tributaries and
covers approximately 8328 km2 (Fig. 2). It is defined by the Blue Ridge
Mountains on the east, North and Shenandoah mountains on the west,
the Virginia-West Virginia state line on the north, and a drainage divide
(composed partly of carbonate rock) with the James River basin on the
south. Outside the study area, the basin continues for a short distance
through the extreme northeastern corner of West Virginia to a point
where the Shenandoah River joins the Potomac River at Harpers Ferry.

The regional terrain is generally rolling and is significantly punctu-
ated only by Massanutten Mountain, a prominent ridge that partly
divides the basin into two valleys for about half of its length. A total of
396 caves are recorded from the basin in the study area; a majority are
small, and only a few are of major extent. Although the area contains a
fairly extensive exposure of carbonate rock (Cambrian to Devonian),
much of it is dolomite and calcareous shale. As a result, the potential
for extensive cave development has been greatly limited. The regional
cavernicolous fauna contains 25 cave-limited species; 23 are troglobites,
and 14 are endemic to the basin (Table 5).

2. James Basin. — In the study area this faunal unit includes all of
that part of west-central Virginia drained by the James River and its
tributaries and covers approximately 7745 km2 (Fig. 2). It is defined by
the Blue Ridge Mountains on the east, Allegheny and Peters mountains
on the west, drainage divides (with carbonate rocks) with the upper
Potomac drainage system (i.e., South Branch and Shenandoah rivers)
on the north, and drainage divides (also with carbonate rocks) with the

110

John R. Holsinger and David C. Culver

New and Roanoke rivers on the south. The regional terrain varies con-
siderably from one part of the basin to another. In the western two-
thirds it is relatively rugged and characterized by numerous prominent
ridges and narrow valleys, whereas in the eastern third it is of lower
relief and generally rolling.

A total of 431 caves are recorded, including some of the largest in
the study area. There are significant exposures of Silurian and Devo-
nian limestones in the western part of the basin, where caves are often
extensive but frequently localized in isolated belts of limestone that crop
out along the flanks of ridges and in valley floors. Limestones and
dolomites of Cambrian and Ordovician age predominate in the eastern
part of the basin, where caves are less extensive but relatively numerous.
The regional cavernicolous fauna is composed of 32 cave-limited spe-
cies; 31 are troglobites, and 16 are endemic to the basin (Table 6).

3. Roanoke Basin. — This faunal unit is the smallest in the study
area and covers only approximately 1073 km (Fig. 2). It is drained by
the Roanoke River and its tributaries and is defined by the Blue Ridge
Mountains on the east and south, Brush and Catawba mountains in
part on the north, and a drainage divide (with carbonate rock) with the
New River on the west. Although the regional terrain varies considera-
bly, most of the karst topography is moderately rolling and developed
on valley floors and low hills. Ninety-one caves are recorded, and all of
them are developed in Cambrian and Ordovician limestones and dolo-
mites. Because of the extensive exposures of dolomite, most of the caves
are small, although several large ones are excavated in Ordovician
limestone in the valley of the North Fork of the Roanoke River. The
regional cavernicolous fauna consists of only 10 cave-limited species, all
trogobites; three species are endemic to the basin (Table 7).

4. New Basin. — This faunal unit encompasses that part of the
study area drained by the New River and its tributaries and covers
approximately 4087 km (Fig. 2). Unlike other major rivers in the study
area, New River flows generally northward and cuts across the regional
strike instead of flowing parallel to it. The basin is defined by the Iron
Mountains and by Poplar Camp and Macks mountains on the south,
complex drainage divides with the Roanoke and James rivers composed
of several ridges of prominent relief on the east, Peters and East River
mountains and Big Stone Ridge on the north, and complex drainage
divides (partly composed of carbonate rocks) with the Holston and
Clinch rivers on the west. The regional terrain is heterogeneous and
characterized in general by both broad and narrow valleys and a
number of prominent ridges.

A total of 419 caves are recorded, a significant number of which are
extensive. Both caves and karst terranes occur in many parts of the
basin but are especially well developed in large valleys on opposite sides
of Cloyds and Brush mountains, along the western side of Big Walker

Invertebrate Cave Fauna

111

Table 5. List of cave-limited species (see text for definition) in the Shenandoah
basin regional fauna. Species listed in same sequence as in text (cf.,
“Review of the Fauna”). * = endemic species. TB = troglobite or
probable troglobite.

AQUATIC SPECIES

Fontigens orolibas
t *Lartetia sp. (TB)

Stygobromus gracilipes (TB)

*S. pseudospinosus (TB)

S. bigger si (TB)

*S. stegerorum (TB)

Caecidotea pricei (TB)
*Antrolana lira (TB)
TERRESTRIAL SPECIES
Miktoniscus racovitzai (TB)
*Apochthonius coecus (TB)
*Kleptochthonius sp. B (TB)

* Mundoehthonius holsingeri (TB)

TERRESTRIAL SPECIES (continued)

*Chitrella superba (TB)

Erebomaster acanthina
Bathyphantes weyeri (TB)

Phanetta subterranea (TB)

Porrhomma cavernicolum (TB)
*Striaria sp. (TB)

Trichopetalum weyeriensis (TB)

T. whitei (TB)

*Pseudanophthalmus avernus (TB)

*P. hubbardi (TB)

*P. limicola (TB)

*P. parvicollis (TB)

*P. petrunkevitchi (TB)

Summary: Total species = 25 (8 aquatic, 17 terrestrial); endemics = 14.

Table 6. List of cave-limited species (see text for definition) in the James basin
regional fauna. Species listed in same sequence as in text (cf.,
“Review of the Fauna”). * = endemic species. TB = troglobite or
probable troglobite.

AQUATIC SPECIES

*SphaIloplana virginiana (TB)

* Stygobromus interitus (TB)

*S. hoffmani (TB)

S. morrisoni (TB)

*S. mundus (TB)

*S. estesi (TB)

*S. conradi (TB)

*S. baroodyi (TB)

* Caecidotea bowmani (TB)

C. holsingeri (TB)

C. vandeli (TB)

C. pricei (TB)

TERRESTRIAL SPECIES
Miktoniscus racovitzai (TB)
*Apochthonius holsingeri (TB)
*Kleptochthonius anophthalmus (TB)

TERRESTRIAL SPECIES (continued)

Rhagidia varia (TB)

Anthrobia monmouthia (TB)
Islandiana muma (TB)

Phanetta subterranea (TB)

Porrhomma cavernicolum (TB)
Nesticus tennesseensis (TB)
*Nampabius turbator (TB)
Trichopetalum packardi (TB)

T. weyeriensis (TB)

Sinella hoffmani (TB)

Euhadenoecus fragilis
Pseudanophthalmus gracilis (TB)

* Pseudanophthalmus inter sect us (TB)
*P. sp. A ( hubbardi group) (TB)

*P. nelsoni (TB)

*P. pontis (TB)

*P. sp. A [pusio group) (TB)

Summary: Total species = 32 (12 aquatic, 20 terrestrial); endemics = 16.

112

John R. Holsinger and David C. Culver

Mountain and in Burkes Garden. The carbonate rocks are predomi-
nantly limestones and dolomites of Cambrian and Ordovician age,
although a limited exposure of Mississippian limestone crops out in the
extreme northwestern part of the basin. The New River also drains sev-
eral karst areas in adjacent West Virginia, and the cave fauna of these
areas was discussed in some detail in a previous paper (Holsinger et al.
1976). The regional cavernicolous fauna includes 29 cave-limited spe-
cies; 26 are troglobites, and 10 are endemic to the basin (Table 8).

5. Holston Basin. — That part of the Holston basin in the study
area lies almost entirely in Virginia and extends only a few kilometers
into Tennessee (Fig. 2). It is drained by three major tributaries of the
Holston River and covers approximately 3690 km2. This faunal unit is
defined by the Iron Mountains on the southeast, the drainage divide
with the New River on the east, and Clinch Mountain and Moccasin
Ridge on the north and northwest. Outside the study area, the basin
continues southwestward through eastern Tennessee to the vicinity of
Knoxville, where the Holston River joins the French Broad River to
form the Tennessee River. The regional terrain varies from moderately
rugged in areas drained by the North Fork to moderately rolling in the
southern two-thirds of the basin drained by the Middle and South forks
of the Holston River. The basin is bisected in part by Walker Mountain,
which trends southwest and forms a prominent interfluve between the
North Fork and Middle Fork.

Although most of the exposed carbonates are limestones and dolomites
of Cambrian and Ordovician age, limited outcrops of Silurian-Devonian
and Mississippian limestones occur in parts of Scott and Washington
counties. A total of 308 caves are recorded, and a number of them are
large. However, much of the carbonate rock exposed in the southern
part of the basin is dolomite and has limited the development of exten-
sive caves. The regional cavernicolous fauna is composed of 19 cave-
limited species; 18 are troglobites, and only three are endemic to the
basin (Table 9).

6. Clinch Basin. — Most of this basin lies within the study area and
is drained by the Clinch River and its tributaries (Fig. 2). It is defined
by the short drainage divide with New River on the northeast, the east-
ern margin of the Appalachian Plateau and Powell Mountain on the
north and west, and Clinch Mountain on the south except for a short
stretch in Russell and Scott counties where Big Moccasin Creek flows
north of Clinch Mountain before turning south to join the North Fork
of the Holston River south of Gate City, Va. As defined in the present
study, this faunal unit ends in Campbell County and in the vicinity of
Norris Dam about 10 km south of where the Clinch River is joined by
the Powell River; it covers approximately 4048 km2. Beyond the study
area, however, the basin extends southwestward for approximately 60

Invertebrate Cave Fauna

113

Table 7. List of cave-limited species (see text for definition) in the Roanoke
Drainage basin regional fauna. Species listed in same sequence as in
text (cf., “Review of the Fauna”). * - endemic species. TB = troglobite
or probable troglobite.

AQUATIC SPECIES TERRESTRIAL SPECIES

*Stygobromus fergusoni (TB) Phanetta subterranea (TB)

Caecidotea vandeli (TB) Porrhomma cavernicolum (TB)

* Pseudotremia cavernarum (TB)
Trichopetalum packardi (TB)
Pseudosinella orba (TB)

Sinella hoffmani (TB)
Arrhopalites clarus (TB)

* Pseudanophthalmus pusio (TB)

Summary: Total species - 10 (2 aquatic, 8 terrestrial); endemics = 3.

Table 8. List of cave-limited species (see text for definition) in the New basin
regional fauna. Species listed in same sequence as in text (cf.,
“Review of the Fauna”). * = endemic species. TB = troglobite
or probable troglobite.

AQUATIC SPECIES

Fontigens orolibas
*Stygobromus ephemerus (TB)
*S. abditus (TB)

S. mackini (TB)

*Caecidotea henroti (TB)

C. incurva (TB)

C. vandeli (TB)

C. richardsonae (TB)
TERRESTRIAL SPECIES
* Foveacheles paralleloseta (TB)
Rhagidia varia (TB)

Phanetta subterranea (TB)
Porrhomma cavernicolum (TB)
Nesticus carteri
Nesticus tennesseensis (TB)

TERRESTRIAL SPECIES (continued)

Pseudotremia tuberculata (TB)
Trichopetalum packardi (TB)
Pseudosinella orba (TB)

Arrhopalites clarus (TB)

Litocampa sp. B (TB)

L. sp. E (TB)

Euhadenoecus fragilis
Pseudanophthalmus gracilis (TB)

*P. egberti (TB)

*P. quadratus (TB)

P. hoffmani (TB)

*P. hortulanus (TB)

*P. sp. B ( petrunkevitchi group) (TB)
*P. sp. C ( petrunkevitchi group) (TB)
*P. punctatus (TB)

Summary: Total species = 29 (8 aquatic, 21 terrestrial); endemics = 10.

114

John R. Holsinger and David C. Culver

Table 9. List of cave-limited species (see text for definition) in the Holston
basin regional fauna. Species listed in same sequence as in text (cf.,
“Review of the Fauna”). * = endemic species. TB = troglobite
or probable troglobite.

AQUATIC SPECIES

Crangonyx antennatus (TB)
Stygobromus mackini (TB)
Caecidotea incurva (TB)

C. recurvata (TB)

C. richardsonae (TB)
TERRESTRIAL SPECIES
Glyphyalinia specus (TB)
Anthrobia monmouthia (TB)
Phanetta subterranea (TB)
Porrhomma cavernicolum (TB)

TERRESTRIAL SPECIES (continued)

Nesticus carteri
N. tennesseensis (TB)

N. pavnei (TB)

Pseudosinella orba (TB)

Litocampa sp. B (TB)

L. sp. E (TB)

* Pseudanophthalmus paradoxus (TB)

P. hoffmani (TB)

*P. sp. A {petrunkevitchi group) (TB)

* Batriasymmodes greeveri (TB)

Summary: Total species = 19 (5 aquatic, 14 terrestrial); endemics - 3.

km to the vicinity of Kingston in Roane County, Tenn., where the
Clinch River joins the Tennessee River. The regional terrain is highly
variable but is generally rugged and characterized by prominent ridges
and relatively narrow valleys. However, in several places, especially in
the northeastern half, broad coves with karsted limestone floors are
formed between mountains, two good examples being Ward Cove in
Tazewell County and Rye Cove in Scott County.

Numerous belts of carbonate rock, ranging in age from Cambrian
to Mississippian, are exposed in the basin. A total of 537 caves are
recorded, many of which are extensive and most of which are excavated
in Cambrian, Ordovician, and Mississippian limestones. The regional
cavernicolous fauna contains 51 cave-limited species; 47 are troglobites,
and 24 are endemic to the basin (Table 10).

7. Powell Basin. — This basin lies completely within the study area
and is drained by the Powell River and its tributaries (Fig. 2). It covers
approximately 2278 km2 and is defined by the eastern margin of the
Appalachian Plateau on the northeast, north, and west, and by Powell
Mountain on the south except in Claiborne and Union counties where
the interfluve with the Clinch drainage is Wallen Ridge and a series of
low, dolomitic ridges that extend across the Central Peninsula between
the arms of Norris Lake. The Powell basin is essentially one large valley
with generally rolling terrain of moderately low relief. However, Wallen
Ridge on the eastern side of the Valley is a topographic feature.

Extensive exposures of Cambrian, Ordovician, and Mississippian
limestones and dolomites occur throughout the basin. The middle of the
valley is floored by several broad belts of Ordovician limestone, whereas
a significant belt of Mississippian limestone crops out along the front of

Invertebrate Cave Fauna

115

Table 10. List of cave-limited species (see text for definition) in the Clinch
basin regional fauna. Species listed in same sequence as in text (cf.,
“Review of the Fauna”). * = endemic species. TB = troglobite or
probable troglobite.

AQUATIC SPECIES

Geocentrophora cavernicola (TB)
Sphalloplana chandleri (TB)
*Spelaedrilus multiporus (TB)
Stylodrilus beattiei (TB)

Fontigens orolibas
Crangonvx antennatus (TB)
Stygobromus cumberlandus (TB)
Stygobromus mackini (TB)
Gammarus minus (Form I)
Caecidotea recurvata (TB)

C. richardsonae (TB)

*Lirceus culveri (TB)
TERRESTRIAL SPECIES
*Amerigoniscus pavnei (TB)

* Kleptochthonius binoculatus (TB)
*K. regulus (TB)

*K. sp. A (TB)

Rhagidia v aria- (TB)

Anthrobia monmouthia (TB)
Phanetta subterranea (TB)
Porrhomma cavernicolum (TB)
Nesticus carteri
N. tennesseensis (TB)

N. ho (singer i (TB)

N. paynei (TB)

Pseudotremia nodosa (TB)

TERRESTRIAL SPECIES (continued)

Pseudotremia tubereulata (TB)

*P. deprehendor (TB)

Pseudosinella orba (TB)

P. hirsuta (TB)

Sinella hoffmani (TB)

Litocampa cookei (TB)

*L. sp. A (TB)

L. sp. C (TB)

*L. sp. D (TB)

L. sp. E (TB)

Euhadenoecus fragilis
* Pseudanophthalmus deeeptivus (TB)

* P. sp. A ( enge/hardti group) (TB)

*P. viearius (TB)

*P. serieus (TB)

*P. hubrichti (TB)

*P. sanctipauli (TB)

*P. sp. A ( hubrichti group) (TB)

*P. sp. B ( hubrichti group) (TB)

*P. praetermissus (TB)

*P. longieeps (TB)

*P. sec/usus (TB)

*P. thomasi (TB)

*P. sp. A (jonesi group) (TB)

*P. unionis (TB)

*P. virginicus (TB)

Summary: Total species = 51 (12 aquatic, 39 terrestrial); endemics = 24.

the Appalachian Plateau on the western side of the valley. A total of
394 caves are recorded, many of which are extensive and among the
largest in the study area. Karst terrane is especially well developed, and
the center of the valley, from Jonesville, Va., to Tazewell, Tenn., is the larg-
est continuous karst corridor in the study area. The regional cavernico-
lous fauna contains 44 cave-limited species; 41 are troglobites, and 27
are endemic to the basin (Table 1 1).

Patterns of Distribution

We have recognized three general types of distributional patterns
exhibited by cave-limited species of the study area. Ranges may be (1)
very widespread, (2) trans-Appalachian, or (3) Appalachian Valley. Most

116

John R. Holsinger and David C. Culver

of the species exhibit the Appalachian Valley pattern, whereas
only 13 are trans-Appalachian. Four are very widespread.

The four very widespread species are the linyphiid spiders Bathy-
phantes weyeri, Phanetta subterranea, and Porrhomma cavernicolum,
and the collembolan Arrhopalites clarus. The spiders are recorded from
caves throughout much of the eastern United States and apparently
show little morphological variation (W. J. Gertsch, in litt.). The range
of A. clarus, on the other hand, is disjunct, with cave populations re-
stricted to the Ozark region and the Appalachians (Christiansen 1982).

Trans-Appalachian species are recorded from caves on both sides
of the Appalachian Plateau — in the Appalachian Valley and Ridge and
eastern margin of the Appalachian Plateau on the east and Interior Low
Plateaus and western margin of the Appalachian Plateau on the west.
The taxonomic status of many of these species is unclear, and, as a
result, their geographic distributions are questionable and in need of
further study. The 13 trans- Appalachian species are the flatworms Geo-
centrophora cavernicola, Sphalloplana chandleri, and S. percoeca; the
terrestrial snail Glyphyalinia specus\ the amphipod Crangonyx anten-
natus\ the aquatic isopod Caecidotea richardsonae\ the terrestrial isopod
Miktoniscus racovitzav, the pseudoscorpion Hesperochernes mirabilis,
the spiders Anthrobia monmouthia, Islandiana muma, and Nesticus
carteri\ the collembolan Pseudosinella hirsuta ; and the dipluran
Litocampa cookei.

Appalachian Valley species are limited to the Appalachian Valley
and Ridge province and closely associated karst islands on the eastern
side of the Appalachian Plateau. There are basically two categories of
Appalachian Valley species with respect to range: (a) species usually
with moderately extensive ranges that inhabit caves in two or more of
the faunal units in the study area and sometimes occur outside the
study area, and (b) species known only from a single faunal unit in the
study area. Species in the first category are the lumbriculid worm
Stylodrilus beattiei; the aquatic snail Fontigens orolibas (s. lat.)\ the
amphipods Gammarus minus (Form I), Stygobromus biggersi, S.
cumberlandus, S. gracilipes, S. mackini, and S. morrisoni\ the aquatic
isopods Caecidotea incurva, C. holsingeri, C. pricei, C. recurvata, and
C. vandelv, the mite Rhagidia varia\ the phalangid Erebomaster
acanthina; the spiders Nesticus tennesseensis, N. holsingeri , and N.
paynei ; the millipeds Pseudotremia nodosa, P. tuberculata, Tricho-
petalum packardi, T. weyeriensis, and T. whiter, the collembolans
Pseudosinella orba and Sinella hoffmani\ the diplurans Litocampa spp.
B, C, and E; the cricket Euhadenoecus fragilis ; and the beetles
Pseudanophthalmus gracilis and P. hoffmani.

The second category comprises 97 species, or 66% of the 146 cave-
limited species in the study area. Except for the beetle Pseudanoph-
thalmus potomaca (northern Highland County and southern Pendleton
County, W.Va.), these species are indicated by an asterisk in the lists

Invertebrate Cave Fauna

117

1 able 11. List of cave-limited species (see text for definition) in the Powell
basin regional fauna. Species listed in same sequence as in text (cf.,
“Review of the Fauna”). * - endemic species. TB = troglobite or
probable troglobite.

AQUATIC SPECIES

*Sphalloplana consimilis (TB)

S. percoeca (?) (TB)
*Lumbriculid (sp.) (TB)

* Fontigens sp. (TB)

* Bactrurus sp. (TB)

Crangonyx antennatus (TB)
Stygobromus cumberlandus (TB)
*S. finleyi (TB)

*S. leensis (TB)

S. mackini (TB)

Caecidotea recurvata (TB)

C. richardsonae (TB)

*C. sp. A (TB)

*Lirceus usdagalun (TB)
TERRESTRIAL SPECIES
*Amerigonsicus henroti (TB)

* Kleptochthonius affinis (TB)

*K. gertschi (TB)

*K. lutzi (TB)

*K. proximosetus (TB)

* K. similis (TB)

* Microcreagris valentinei (TB)
Hesperochernes tnirabilis

TERRESTRIAL SPECIES (continued)

Phanetta subterranea (TB)

Porrhomma cavernicolum (TB)
Nesticus carteri
N. holsingeri (TB)

N. paynei (TB)

* Pseudotremia valga (TB)

P. nodosa (TB)

Pseudosine/la hirsuta (TB)

P. orba (TB)

Litocampa cookei (TB)

Euhadenoecus fragilis
* Pseudanophthalmus engelhardti (TB)
*P. holsingeri (TB)

*P. rotundatus (TB)

*P. sidus (TB)

*P. sp. B ( engelhardti group) (TB)

*P. de lie at us (TB)

*P. hirsutus (TB)

*P. cordicollis (TB)

*P. pallidus (TB)

*P. sp. B (jonesi group) (TB)

*Arianops jeanneli (TB)

Summary: Total species = 44 ( 14 aquatic, 30 terrestrial); endemics - 27.

given in Tables 5 through 1 1 and need not be enumerated here. They are
distributed numerically by taxon as follows: flatworms (2), lumbriculid
worms (2), aquatic snails (2), amphipods (14), aquatic isopods (6), ter-
restrial isopods (2), pseudoscorpions (15), mites (1), centipedes (1), mil-
lipeds (4), diplurans (2), and beetles (46). An analysis of the ranges of
these species indicates that 53 are recorded from single caves, 15 from
two or rarely three caves within 5 km of each other, and 29 from two or
more (usually more) caves located some distance apart. The percent of
endemic species in each basin is: Shenandoah (0.56), James (0.50), Roa-
noke (0.30), New (0.34), Holston (0.16), Clinch (0.47), and Powell
(0.61).

These data give a clear picture of the degree of endemism among
cave-limited species in the study area. The highest percentage of regional
faunal endemics is found among the beetles (principally Pseudanoph-
thalmus) and pseudoscorpions (principally Kleptochthonius), where, in

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John R. Holsinger and David C. Culver

both taxa, almost all species are restricted to a single faunal unit, and
a majority of species in both groups are single-cave isolates. There is
also a high percentage of regional faunal endemics among lumbriculid
worms, snails, amphipods, and isopods; and more than half of the spe-
cies in each of these groups are recorded from a single faunal unit. On
the whole, comparatively fewer endemics are noted among flatworms,
mites, millipeds, and diplurans; and no endemics are recorded for spiders
and collembolans.

Although the cave-limited fauna contains many highly localized
endemics, there are a number of species — especially the few, select troglo-
philes, troglobitic spiders, and collembolans, and some of the troglobitic
amphipods, isopods, millipeds, and diplurans — that have relatively exten-
sive ranges within the study area and are apparently good dispersers.
These species are found in two or more of the seven faunal units, and at
least two species, the highly vagile spiders Phanetta subterranea and
Porrhomma cavernicolum , occur in all seven.

The extent to which drainage basins or faunal units share cave-
limited species is indicated by the data compiled in Table 12. Theoreti-
cally, dispersal between basins could take place by some of the more
vagile troglobites through caves and solution channels developed in
parts of drainage divides and interfluves composed of carbonate rock,
and through endogean habitats (e.g., deep ground litter and shallow
underground compartments) and groundwater habitats (e.g., interstitial,
hypotelminorheic) outside karst areas. At least three of the troglo-
philes we have included in the cave-limited fauna ( Euhadenoecus fragi-
lis , Erebomaster acanthina , and Nesticus carteri ) should be able to
undergo limited dispersal through ecologically suitable epigean habitats.
The frequency of species exchange between basins would be influenced
by proximity of the basins as well as by the geological structure and
geographic extent of drainage divides. The farther removed two basins
are from each other, the fewer species they would be expected to share,
and, by the same analogy, the closer they are, the more species they
would be expected to share. Moreover, long common divides or inter-
fluves, especially those containing carbonate rock, would be expected to
facilitate more dispersal than do short divides or divides composed
entirely of non-carbonate, clastic rock.

Our data (Table 12) strongly support these assumptions and indi-
cate clearly that, with few exceptions, adjacent basins have more species
in common than far removed ones, and, furthermore, that basins on
opposite sides of divides or interfluves containing carbonate rock gener-
ally share more species than those separated by divides composed
entirely of non-carbonate rock.

Diversity-Area Relationships

In the course of the present investigation it became increasingly
obvious that some drainage basins in the study area had signifi-

Invertebrate Cave Fauna

119

Table 12. Number of cave-liriiited species shared by drainage basins. Data
based on species listed in Tables 5 through 1 1.

Drainage

Basin

Shenandoah

James

Roanoke

•

New

Holston

Clinch

Powell

Shenandoah

James

Roanoke

New

1 1

Holston

Clinch

Powell

cantly more cave-limited species and ecologically more complex cave
communities than others, and, that with few exceptions, cave species
diversity and ecological complexity increased in those areas with large
numbers of caves and extensive karst development. To gain a better
understanding of this apparent relationship and to translate it into
quantitative terms, we compiled the data shown in Table 13. As these
data indicate, there are major differences among the seven drainage bas-
ins with respect to area, number of cave-limited species, and number of
recorded caves. In order to demonstrate the relationship between these
variables concisely, we calculated both species density and cave density
per unit of area for each basin (Table 13). We then plotted these values
and found a strong linear relationship between species density and cave
density in the different basins (Fig. 37). There is a progressive increase
in density of species per unit of area from the Shenandoah basin, with
the lowest cave density, to the Powell basin, with the highest cave den-
sity. The slope of the regression line in Figure 37 is highly significant (b
= 0.13, p < .001) and 92% of the variation can be explained by variation
in cave density.

Cave density alone probably does not determine the number of
cave-limited species in a given area or basin. Other factors, which are
more difficult to quantify, but probably equally important in determin-
ing the species diversity of a faunal unit, include both the amount and
continuity of exposed, cavernous limestone and the degree of karst
development. Cave density, however, which can be easily calculated for
an area where the caves are well documented, is an excellent indication
of the extent of cavernous limestone and also often reflects the extent of
karst development.

120

John R. Holsinger and David C. Culver

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Invertebrate Cave Fauna

121

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Fig. 37. Relationship of species density to cave density in the seven drainage
basins of the study area. Regression line calculated by least squares method.
Drainage basins: S = Shenandoah; J = James; R = Roanoke; N = New; H =
Holston; C = Clinch; P = Powell.

The geological structure of an area, then, appears to play a signifi-
cant role in determining the diversity of cave species. There are appar-
ently several complex, interrelated reasons for this. First, large areas of
continuously exposed, cavernous limestone would increase the oppor-
tunities for invasion of subterranean habitats by surface ancestors of
cavernicoles; expand the potential for dispersal by hypogean species fol-
lowing cave colonization; increase potential habitat space in the form of
cave passages and solution channels, thus allowing additional coloniza-
tions and, subsequently, the development of complex communities as
the number of species increased; and increase the accessibility of caves
to trogloxenes and troglophiles, which periodically introduce food
underground. Second, well-developed karst terranes, characterized in
the Appalachians by numerous sinkholes, blind valleys, sinking streams,
bare limestone outcrops, and springs, would provide accessible avenues
for the invasion of subterranean habitats by surface ancestors, greatly
facilitate the movement of organic nutrients into subterranean channels,
and increase the hydrological complexity of subterranean groundwater
systems.

Regional terrains and the potential for cave and karst development
in the seven drainage basins have already been characterized briefly, and

122

John R. Holsinger and David C. Culver

our assumption that geological structure directly affects cave species
diversity and ecological complexity appears to be supported by the dif-
ferences noted. Clearly, those basins with limited, discontinuous expo-
sures of cavernous limestone and restricted karst terranes, such as the
Shenandoah, James, and Holston, have significantly fewer cave-limited
species per unit of area than those basins with extensive, continuous
exposures of cavernous limestone and well-developed karst terranes,
such as the Clinch and Powell. Those basins intermediate in these geo-
logical parameters, the Roanoke and New, fall somewhere between the
two extremes of species diversity. Direct field observations tend to rein-
force our assumption, namely that, by and large, we found more species
and larger populations in the caves of the Clinch and Powell basins than
anywhere else in the study area.

The numbers of genera and families represented in the cave fauna
of a basin should provide a further indication of taxonomic diversity
and ecological complexity. In order to check this, we compiled the
numbers of genera and families with cave-limited species in the seven
drainage basins of the study area (Table 14). As the data indicate, the
numbers of both genera and families are highest for the Clinch and
Powell basins, again emphasizing that the greatest faunistic diversity
occurs in areas with the most extensive, continuous exposure of
cavernous limestone.

Barr (1967b, 1968) compared the cave systems of the Appalachian
Valley and Ridge province with those of the Mississippian limestone
plateaus in the Interior Low Plateaus region (i.e., Mitchell Plain in
southern Indiana, Pennyroyal Plateau in Kentucky, and the Cumber-
land Plateau margin in Kentucky, central Tennessee, and northern Ala-
bama) and concluded that they were ecologically very different. Ecolog-
ical differences were attributed to differences in geological structure.
The Paleozoic limestones (Cambrian to Mississippian) of the Appala-
chian Valley and Ridge are faulted and folded and exposed in long, linear
anticlinal valleys that are separated by synclinal ridges of clastic rocks.
In comparison, the Mississippian limestones of the Interior Low Pla-
teaus are relatively undisturbed and exposed over broad areas. Because
of this, dispersal of troglobites in the Appalachian Valley and Ridge
would be restricted, whereas dispersal would be enhanced in the Missis-
sippian plateaus. This comparison was based primarily on troglobitic
trechine beetles (especially Pseudanophthalmus ), which are represented
by many species in both regions. But as Barr (1967b, 1968) has pointed
out, the implications apparently apply to many other groups of caverni-
coles as well. On a broad scale, the troglobitic fauna of the Mississip-
pian plateau is more diverse than that of the Appalachian Valley.

Although our comparison of drainage basins in the study area of
Virginia and northeastern Tennessee focused on relatively small cave

Invertebrate Cave Fauna

123

Table 14. Frequency distribution by drainage basin of genera and families with
cave-limited species.

Drainage Basin

Number of Genera

Number of Families

Shenandoah

James

Roanoke

New

Holston

Clinch

Powell

regions within a single physiographic province and considered trechine
beetles as well as all other groups of cave-limited species, our findings
are essentially the same as those of Barr. The results of both of these
studies corroborate the hypothesis that in a geographic region where
climatic conditions are historically similar, areas with extensive, contin-
uous exposures of cavernous limestone will harbor more diverse troglo-
bite faunas than areas with limited, discontinuous exposures of limestone.

The data from two other studies, both on Appalachian cave faunas,
also tend to corroborate this hypothesis. In a study of the invertebrate
cave fauna of West Virginia (Holsinger et al. 1976), we documented a
significantly richer troglobitic fauna in the Greenbrier Valley than in
any other limestone region of that state. Like the Clinch and Powell
valleys in Virginia, the Greenbrier Valley contains extensive, continuous
exposures of cavernous limestone and a well-developed karst terrane;
cave density is very high. In other major cave regions of West Virginia
(e.g., the Monongahela and Potomac basins), where limestone expo-
sures and karst terranes are more restricted and cave density is lower,
the troglobitic faunas are correspondingly less diverse. In the other
study, the troglobitic fauna of Pennsylvania was compared on a broad,
regional scale with that of the Virginias (Holsinger 1976). In Pennsylva-
nia, limestone areas are mostly very narrowly delimited, and caves are
typically very small. The troglobitic fauna is exceedingly sparse (only 15
species) and contains largely aquatic species, some of which are stygobi-
onts. In contrast, the troglobitic fauna of the Virginias, where cavernous
limestone areas are generally much more extensive, is significantly
richer. Although, admittedly, the proximity of the cave region of Penn-
sylvania to Pleistocene glaciation probably has had something to do
with its impoverished troglobitic fauna (Holsinger 1976), the effect of
geological structure has probably been of equal or greater significance
(Holsinger 1976).

124

John R. Holsinger and David C. Culver

Karst Areas as Islands

Several authors (e.g., Barr 1968, Culver et al. 1973) have explored
the potential analogy between caves and islands. Of special interest is
whether the number of species in a cave or a karst region is determined
by an equilibrium of immigration and extinction rates when applied to
individual caves or parts of caves. The time scale is ecological in the
sense that populations rather than species are becoming extinct. Craw-
ford (1981) and Culver (1982) have critically reviewed the validity of the
cave-island analogy in ecological time. On a larger geographic scale, the
number of species in a karst region may be determined by a balance
between the rate of isolation of species in caves and the rate of extinc-
tion of cave species. The time scale for this process is evolutionary
rather than ecological.

As Simberloff (1976) points out, there has been an uncritical accep-
tance of island biogeography theory, and attempts to test the hypothesis
are few. The best tests of the hypothesis involve direct observations of
immigrations and extinctions, but such verification is clearly not possi-
ble for evolutionary time scales. Therefore we must fall back on an
analysis of area effect. It is often assumed that there is a one-to-one
correspondence between island biogeography theory and a value of z in
the following equation:

S=CAZ (1)

where S is species numbers, A is area, and C and z are fitted constants.
Although processes other than an equilibrium between immigration and
extinction can result in a z-value near 0.26 (Connor and McCoy 1979),
the validity of the equilibrium model does not require a z-value of 0.26
(Culver 1982).

Analysis of area effect can provide some useful clues about the pro-
cesses that determine species numbers. First, the absence of an area
effect would indicate that area was incorrectly measured or that
some other variable and some other process is more important. For
example, terrestrial cave species numbers might be determined by avail-
ability of suitable epigean ancestors for which elevation might be more
important than area. Second, if the island analogy holds, then equation
(1), sometimes called the power function, should be a better fit than the
untransformed linear model:

S=C + zA (2)

Equation (2) represents a model for area effect where species numbers
are controlled by passive sampling from the species pool (Connor and
McCoy 1979), and does not involve a balance between immigration and
extinction. Third, if the power function is the best fit, then the larger the
exponent z, the longer the time required to reach equilibrium (Culver et
al. 1973). A large z-value indicates that it is unlikely the system is in
equilibrium. What follows is a preliminary analysis, with extensive
analysis in preparation by the authors.

Invertebrate Cave Fauna

125

The basic units of analysis are the seven drainage basins defined
above. Area was estimated in three ways, each with somewhat different
biological interpretations and with its share of technical problems. The
first measurement is the total area of the drainage basin. While rela-
tively accurate, the measurement combines karst and non-karst areas. If
most terrestrial cave species occur in shallow underground compart-
ments (Juberthie and Delay 1981, and see elsewhere this paper) in karst
and non-karst areas as well as caves, and if most aquatic cave species
occur in interstitial habitats as well as caves (Henry 1978, and elsewhere
this paper), then total area of the drainage basin is the appropriate mea-
surement of area. The second measurement is that of the area underlain
by soluble carbonate rock in each drainage basin. This should measure
the area in which caves potentially occur. The measurement itself was
obtained by finding the percentage of limestone on a series of randomly
chosen USGS topographic quadrangles, according to Douglas’s (1964)
mapping of exposed carbonate rock. The problem is that not all of the
exposed carbonates are equally likely to have caves, because, as shown
by Douglas, they include both limestone and dolomite, and the latter
usually has significantly fewer and smaller caves depending on its com-
position (see Holsinger 1975). The third measurement estimates the area
underlain by caves by the total number of caves for each drainage basin.
This measurement avoids the problem of differential cave development
in different limestone strata, but adds the problem that the number of
known caves is correlated with sampling intensity.

We have limited our analysis to terrestrial troglobites that are
endemic to a particular drainage basin for several reasons. First and
most important, the terrestrial troglobite endemics are assumed to be a
relatively homogeneous group with respect to their time of isolation in
caves. This allows us to formulate and test the following hypothesis that
terrestrial species were isolated in caves during the series of Pleistocene
interglacials (immigrations) with extinctions also occurring. If the endemic
terrestrial fauna is a more or less perfect record of these events, then the
power series model should be a better fit than the linear model, and the
exponent z of equation (1) should be around 0.26. Alternative hy-
potheses of special interest are three. If the number of endemic terrestrial
troglobites reflects sampling intensity, then the linear model should pro-
vide a better fit. If a significant proportion of the terrestrial species
arose not directly from epigean ancestors, but from subterranean colo-
nization by other troglobites, then species numbers should depend on the
amount of fragmentation of the limestone. Finally, if significant extinc-
tions have occurred since the Pleistocene, the coefficient of area effect z,
will be much greater than 0.26.

The second reason for limiting analysis to terrestrial troglobites
endemic to a basin is that non-endemic terrestrial species either can
move between basins, unlike the endemics, or are actually several

126

John R. Holsinger and David C. Culver

unrecognized sibling species. In any case, they complicate interpretation
of the results. The third reason for limiting the analysis is that aquatic
troglobites frequently inhabit caves and interstitial or non-cave karstic
waters simultaneously.

The lists of basin endemic terrestrial species are given in Tables 5
through 11, and various area measurements for the drainage basins are
given in Table 15. The primary basis for choosing between regression
models is whether the residuals after regression show any systematic
pattern (see Sugihara 1981). However, the residuals of neither equation
(1) nor equation (2) show any systematic bias, which is not surprising
given the small number of points. A secondary criterion, how much of
the variance in the dependent variable is explained by the independent
variable, can be used to tentatively decide between alternatives (see
Connor and McCoy 1979).

The results are given in Table 16, and given the small number of
data points, the results are remarkably consistent. For both the log and
linear models, number of caves (or log of the number of caves)
explained more of the variance than either drainage area or limestone
area. The log model consistently gave a better fit than the linear model.
Finally, the best fit was provided by the log model using number of
caves as the independent variable, with C = 0.007 and z = 1.20. The large
z-value indicates extinctions have been occurring since the Pleistocene,
but that the basic island analogy holds. The lack of significant correlation
between the log of species and a measure of limestone fragmentation,
namely percent of area covered by limestone, indicates that speciation
resulting from underground movement is unimportant. Finally, we must
stress the tentative nature of our conclusions, especially because
correlations among independent variables have not been thoroughly
explored.

Origin, Evolution, and Dispersal

Because there are fundamental differences between aquatic and
terrestrial cave species with respect to modes of origin, habitats, and
dispersal, we will discuss them under separate headings.

Aquatic Species. — Basically two different patterns have been noted
for aquatic troglobites in the study area with regard to their origin. It
should be noted that these patterns are perceived as general trends only
and are not intended to be rigid categories. The first pattern is
exemplified by species that appear to have evolved directly from
preadapted epigean ancestors. Morphological and physiological changes
have developed concurrently with colonization of subterranean waters.
These species belong to genera that are simultaneously represented by
eyed, pigmented surface species, some of which are not far removed
taxonomically (or genetically?) from subterranean forms. Taxa fitting
this pattern are hydrobiid snails ( Fontigens ), some crangonyctid
amphipods ( Crangonyx ), and asellid isopods ( Caecidotea and Lirceus).

Invertebrate Cave Fauna

127

Table 15. Summary of data on area used in regression analysis.

Drainage

Basin

Area(km2)

% Limestone

Limestone

Area(km2)

Number

of Caves*

Shenandoah

8328

3847

396

James

7745

2943

431

Roanoke

1073

386

New

4087

1349

419

Holston

3690

2288

308

Clinch

4048

2210

537

Powell

2278

981

394

♦Includes all caves recorded from the study area through 1980.

Table 16.

Comparison of regression equations using data in Tables 5 through

and Table ]

15. Abbreviations for independent variables: DA =

drainage area;

LA = limestone

area; NC = Number of caves.

Endemic terrestrial species (dependent variable) is abbreviated ET.

Percent

Dependent

Independent

Variance

Variable

Explained

0.0

N.S.

0.6

N.S.

52.0

>0.95

In ET

In DA

17.3

N.S.

In ET

In LA

17.7

N.S.

In ET

In NC

63.9

>0.95

The second pattern is exemplified by species that do not appear to
have evolved directly from epigean ancestors but instead were probably
derived through lineages from ancestors already living in subterranean
groundwater habitats. These species have no known surface congeners
and belong to phylogenetically very old groups. Taxa corresponding to
this pattern are crangonyctid amphipods ( Bactrurus and Stygobromus),
cirolanid isopods (Antrolana), and possibly planarians ( Sphalloplana ).

Alloeocoelid and lumbriculid worms are still too poorly known
taxonomically and ecologically in North American subterranean waters
to identify them with either of the two patterns (see Carpenter 1970a,
Cook 1977). Common troglophiles, such as Phagocata spp., Fontigens
orolibas, Gammarus minus, and Cambarus bartonii, are found in both
epigean and hypogean waters, and the cave populations probably
represent recent invasions of subterranean habitats. However, as indi-
cated below, F. orolibas and G. minus (Form I) could be special cases.

128

John R. Holsinger and David C. Culver

It is difficult to speculate on a time of origin for aquatic troglobites.
Factors that might have been responsible for invasion and colonization
of subterranean waters by putative ancestors undoubtedly revolve around
a complex of interacting, interrelated biological and geological pro-
cesses (see also Barr and Holsinger 1985). Changes in stream gradients
and flow patterns and diversion of surface streams into underground
channels by subterranean stream piracy in karst areas have been sug-
gested as possible factors (Barr 1968, Holsinger et al. 1976, Culver
1982). All of these geological processes have continued over millions of
years of erosional history in the Appalachians (see Hack 1969), and
none is easily identifiable with a given time period. If combined biologi-
cal and geological-area cladograms can be developed for some of the
taxa and areas in question, they might prove useful in approximating
vicariant events that led to the isolation of ancestral populations in
groundwater habitats.

One intuitively obvious avenue for the underground invasion of
aquatic organisms in karst areas would be through springs whose waters
are the continuation to the surface of cave streams. Many aquatic
trogloxenes and troglophiles (viz., species of Phagocata, Fontigens, Goni-
obasis , Gammarus , Lirceus , and Cambarus) inhabit both springs and
cave streams, and populations are occasionally continuous from a
spring well upstream into an adjoining cave. Moreover, some troglobitic
species (viz., in Fontigens , Crangonyx, Caecidotea , and Lirceus ) are
closely allied taxonomically with congeneric epigean species living in
surface springs, suggesting close genetic affinities between surface and
cave forms. The geographic isolation of a population in an underground
stream could occur if the spring fed by this stream was eliminated by
erosion or lowering of base level.

The origin of troglobitic planarians of the genus Sphalloplana is
somewhat obscure. Of the four species in the study area, two — S. con-
similis and S. virginiana — are isolated in widely separated karst areas
and have been assigned to different subgenera (see Kenk 1977). The
taxonomic position of S. pereoeca is unclear as mentioned earlier, and
its range may extend far outside the study area. Sphalloplana ehanderli ,
which is widespread but known only from three disjunct localities, is
probably a “morphological” species. Except for one species with eyes
and dark pigmentation from a spring in central Japan (see Mitchell
1968), the anatomy of which has not been studied (Kenk 1977), all other
members of Sphalloplana are eyeless, unpigmented forms restricted to
subterranean groundwater habitats, and most of them inhabit caves in
North America (Kenk 1977, Kawakatsu and Mitchell 1981).

Despite unresolved taxonomic problems in the genus Fontigens , it
is obvious from similarities in shell structure that the eyeless, unpig-
mented cave populations are closely allied morphologically with the

Invertebrate Cave Fauna

129

eyed, pigmented spring populations; the former probably evolved directly
from the latter. Spring populations of the Fontigens orolibas “complex”
in the Blue Ridge Mountains are presumably physically well isolated
from cave populations in the Appalachian Valley at present, and
because there is no evidence for much dispersal mobility in these tiny
snails, we must assume that this isolation has prevailed for a long period
of time. Presumably, however, at one time in the past these populations
were more or less contiguous.

The isopod family Asellidae is probably a very ancient freshwater
group dating back perhaps to the Mesozoic (Birstein 1964). There are
many genera and numerous subterranean species. Some of the troglo-
bitic species of Caecidotea undoubtedly have been in subterranean
groundwaters for a long period of time, but based on the fact that there
are a number of epigean species in the genus, of which some are appar-
ently not far removed taxonomically from hypogean species, we suspect
that the majority of troglobitic species have evolved directly from sur-
face ancestors. The genus also contains a small number of troglophiles
or trogloxenes, and some of the stygobionts from the east-central Uni-
ted States sometimes have vestigial eyes and light pigmentation (Lewis
and Bowman 1981). In the Greenbrier Valley of West Virginia just west
of the study area, Caecidotea scrupulosa (Williams), a typical epigean
species outside karst areas, has apparently recently invaded caves,
and populations show varying degrees of eye and pigment loss, some-
times corresponding to the distance these animals live inside caves
(Steeves 1969).

The occurrence in subterranean waters of Lirceus is probably more
recent than that of Caecidotea , inasmuch as species of the former are
much less common in caves and only two of the 15 described species in
this genus are troglobites. These two troglobites, which are closely
related sister species that occupy very delimited ranges in different karst
areas on opposite sides of Powell Mountain in southwestern Virginia,
do not appear highly specialized morphologically for a cave existence or
far removed traxonomically from epigean congeners (Holsinger and
Bowman 1973, Estes and Holsinger 1976). In the Ward Cove karst of
Tazewell County, an undescribed species of Lirceus with tiny eyes
inhabits several cave streams and their combined resurgence at Maiden
Spring. Specimens from the spring population are pigmented, whereas
those from the caves are not.

Following colonization of subterranean waters, asellid isopods can
apparently disperse over relatively broad areas through groundwater
habitats outside of caves and even karst areas. Of the 1 1 troglobites in
the study area, six are found in more than one drainage basin, and only
four are known from a single karst area. Caecidotea pricei for example,
although usually found in caves, has been collected several times in the

130

John R. Holsinger and David C. Culver

Shenandoah Valley from small springs and seeps in alluvium underlain
by Martinsburg shale (Holsinger and Steeves 1971). The relatively long
range of this species, i.e., Rockbridge County, Va., northeastward to
southeastern Pennsylvania, can probably be attributed to its ecological
flexibility to live in both interstitial and hypotelminorheic habitats
between karst areas. Several other troglobitic isopods in the study area
have also been collected from non-cave groundwater habitats.

The occurrence of the cirolanid isopod Antrolana lira in a single,
isolated subterranean groundwater aquifer in the Shenandoah Valley is
one of the most intriguing zoogeographic problems in North American
biospeleology. This unique, monotypic form is the only subterranean
freshwater cirolanid isopod found in North America north of Texas,
Mexico, and the West Indies, and is probably a very old relict. The
family Cirolanidae is predominantly marine and only a small number of
species live in freshwater, all of which, except for two poorly known
forms from Africa and one from Cuba, are eyeless, unpigmented species
obligatory to subterranean groundwaters. A total of 17 species in 10
genera have been described to date from subterranean waters in the
Western Hemisphere, and several additional forms from the Bahamas,
Grand Cayman Island, and Haiti are being described. Other genera
and species have been reported from groundwater habitats in southern
Europe and the Mediterranean region.

With the notable exception of A. lira , troglobitic cirolanids live in
areas that are either presently near coastal marine zones or were
exposed to shallow marine transgressions in the Cretaceous or Tertiary.
Because a majority of the cirolanids are marine and the troglobitic species
live either in close proximity to the sea or in old marine embayment
areas, many workers have hypothesized that the subterranean fresh-
water species were derived directly from marine ancestors during the
recession of seawater from limestone regions (Bowman 1964, Vandel
1965b, Cole and Minckley 1966, Carpenter 1981, Contreras-Balderas
and Purata-Velarde 1982, and others). The presence of troglobitic ciro-
lanids in the saline water of a small cave on San Salvador Island in the
Bahamas (see Carpenter 1981) and in the brackish water of a limestone
tunnel on the island of Aruba (see Botosaneanu and Stock 1979) offers
additional support for this hypothesis. Since these troglobitic cirolanids
live in brackish or saline water, they may well represent ecological tran-
sition stages in the evolution of subterranean freshwater forms from
preadapted marine ancestors (Carpenter 1981).

If the prevailing hypothesis for the origin of troglobitic cirolanids is
applied to A. lira , then this species would have to be regarded as the
derivative of an ancient lineage dating back to the Paleozoic, when what
is now the Appalachian Valley was last subjected to marine transgres-
sions (Collins and Holsinger 1981). The evidence, however, argues

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strongly against a Paleozoic origin. Although the oldest known isopods
are recorded from fossils of Pennsylvanian age, these early forms were
phreatoicideans and not flabelliferans (Schram 1974, 1977). Based on
fossil evidence, flabelliferan isopods did not appear until the Triassic
(Schram 1974). Furthermore, the Appalachians did not develop into
their present form until periods of extensive uplifting, folding, and fault-
ing occurred in late Paleozoic and early Mesozoic times. “Stable”
groundwater habitats almost certainly could not have existed in this
region until post-Triassic times.

It appears more probable that freshwater cirolanids ancestral to A.
lira were derived from marine forms in the Late Cretaceous or early
Tertiary when marine embayments existed on the coastal plain of
Virginia approximately 100 km east of the Appalachian Valley. Their
invasion of freshwater habitats would have been followed by their
migration west into karst areas west of the Blue Ridge. Bowman (1964)
has also suggested the possibility of an origin along the Atlantic coast
with subsequent dispersal to the west. But he pointed out that this mode
of origin would have required a freshwater epigean progenitor, in
contrast to other troglobitic cirolanids, which are believed to have
descended directly from marine ancestors.

Whether its ancestral stock was epigean or hypogean cannot be
determined, but in view of the evidence given above, it is doubtful that
A. lira was derived directly from a marine ancestor. It should be noted
further that, despite the extreme rarity of epigean freshwater cirolanids,
at least one bona fide freshwater species, Saharolana seurati Monod, is
recorded from a spring basin in southern Tunisia (see Monod 1930,
Vandel 1965b). The reduced eyes and association with a groundwater
outlet of this species suggest that it is preadapted to a subterranean
existence. A similar stage may have occurred during the evolutionary
history of A. lira.

The troglobitic amphipod fauna has probably originated directly
both from epigean ancestors (e.g., Crangonyx ) and from ancestral
lineages already living in subterranean waters (e.g., Bactrurus and
Stygobromus). The family Crangonyctidae, which contains all of the
troglobitic amphipods in the study area, is widespread over the Holarctic
region and, like Asellidae, is presumably an ancient freshwater group
dating back to the Mesozoic (Holsinger 1977, 1978, 1986a, 1986b).

With the exception of one species from Florida and two or three
from Europe, troglobitic species of Crangonyx are not far removed
taxonomically from surface congeners and do not appear to be as highly
specialized for a subterranean existence as species of Bactrurus and
Stygobromus (see Holsinger 1969a, 1977; Culver 1976; Dickson and
Holsinger 1981). Of the 22 described species in the genus, eight are
troglobitic (or phreatobitic) and two are troglophilic. Most of the

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troglobites in North America, including C. antennatus from the study
area, are represented by some populations with vestigial eyes, but the
presence or absence of eyes may vary both within and between
populations.

Recent studies by Dickson (1976, 1977a, 1977b, 1979) and Dickson
and Holsinger (1981) on the ecology of C. antennatus in Lee County,
where the species is very common in caves, have revealed what are
apparently two microgeographic races corresponding to habitat types.
One race is found in mud-bottom drip pools, and the other in small
gravel-bottom streams. Dickson has found small differences in behavior,
ecology, and morphology between the races. In addition to ecological
studies, six populations from the same area were genetically analyzed by
electrophoresis (Dickson et al. 1979), but allozyme allele frequencies
were determined at only two polymorphic loci. This study revealed a
high degree of allele frequency heterogeneity among the populations
and indicated a tendency for stream and pool amphipods to cluster in
populations distinct from one another (see Table 1 in Dickson et al.
1979).

Despite some apparent isolation between the two habitat types and
the microgeographic heterogeneity among populations, additional
observations on C. antennatus indicate that at least a limited amount of
dispersal can occur between cave populations. One of us (Holsinger
1969a, 1978) has shown that this species may also inhabit perched
groundwater above cave passages and occasionally enter caves from this
habitat in dripping vadose water. The ecological flexibility of C.
antennatus that allows it to inhabit simultaneously several different
types of subterranean groundwater habitats undoubtedly accounts in
part for its large range, which extends far southwest of the study area.
The fact that C. antennatus does not appear to be far removed
taxonomically from surface congeners and is not a highly specialized
troglobite indicates to us that it has evolved directly from an epigean
ancestor in fairly recent times.

In contrast to Crangonyx, Bactrurus and Stygobromus are ex-
clusively subterranean groups in which all known species are eyeless and
unpigmented, and apparently highly specialized stygobionts. Species of
these genera inhabit a wide variety of groundwater biotopes: cave
streams, pools and phreatic lakes; interstitial media; small springs and
seeps; wells; drain tiles; and, rarely, even Pleistocene relict lakes. Some
species are restricted primarily to caves per se, where they are often
associated with small streams; others simultaneously inhabit caves
(usually drip or seep pools) and groundwater habitats outside caves; and
even others inhabit groundwater habitats outside karst areas and are
never found in caves (Holsinger 1967a, 1969b, 1972, 1977, 1978; Culver
1982). Although Bactrurus and Stygobromus are apparently closely

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133

allied morphologically, the latter has a much wider geographic dis-
tribution and contains many more species (Holsinger 1977).

Bactrurus and Stygobromus are exclusively of subterranean facies
and occupy virtually every conceivable type of groundwater habitat.
Moreover, Stygobromus occurs throughout a large part of North
America north of Mexico and is represented by numerous species (more
than 160 counting undescribed forms). These facts strongly imply that
these amphipods are very old stygobionts that have inhabited ground-
waters for a long period of time. The invasion of cave waters, especially
small drip and seep pools, appears to be a dynamic, ongoing process
that has occurred in the past and is continuing at present (Holsinger
1978, Culver 1982). The original colonization of subterranean ground-
waters by various crangonyctid amphipods probably occurred long
before the present generation of cave habitats was available (Culver
1982). Our observations (Holsinger 1978, Culver 1982) indicate that
cave drip-pools are not usually the primary habitats of many small
species of Stygobromus. These cave habitats are only secondary or
marginal biotopes that are periodically populated by animals from
interstitial groundwaters outside caves per se. Some good examples in
the study area of small species recorded only from drip/seep pools
include S. cumberlandus, S. ephemerus, S. estesi, S. finleyi, S. hoffmani ,
S. leensis, and S. pseudospinosus. Other species in the study area appear
to be permanent members of the cavernicole fauna and include S.
baroodyi, S. conradi, S. gracilipes, S. mackini, S. morrisoni, and
Bactrurus sp. These species are usually comparatively large in size and
inhabit small streams, although some, especially V. mackini, are
commonly found in drip pools as well. One cavernicolous species in the
study area, S', stegerorum, is unique, however. It is known only from
deep lakes of phreatic water in two caves in Cave Hill in Augusta
County, where it is associated with the cirolanid isopod Antrolana lira.

Another amphipod, Gammarus minus (Gammaridae), although not
considered a troglobite, is of considerable interest. In the Appalachian
Valley this species is apparently represented by three morphological
forms (see Holsinger and Culver 1970). Form III has relatively short
antennae, well-developed eyes, and dark pigmentation; it inhabits surface
springs throughout the range of the species. Form II has relatively long
antennae, slightly reduced eyes, and weak pigmentation (variable); it
inhabits caves over a broad geographic area. Form I has relatively long
antennae, greatly reduced eyes, and weak pigmentation; it is known
only from caves in the Ward Cove karst of Tazewell County, Va., and
the Great Savannah karst of Greenbrier County, W.Va. These two areas
are situated in different drainage basins and are separated
geographically by a distance of 122 km and several prominent mountains.
Form II occurs in many caves in the Virginias but is rarely as common

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as Form I in a given cave. These two forms are never found together in
the same cave, although they sometimes inhabit caves no more than 5 to
10 km apart.

In a previous study (Holsinger and Culver 1970), in which the
morphological variation of this species was carefully analyzed, we
concluded that the three forms probably represented different eco-
phenotypes of a single, highly variable species. However, we were
unable to give a satisfactory explanation for the presence of Form I in
only two isolated karst areas, except to suggest that it might represent a
convergent ecotype that occurs only under special environmental
conditions in the presence of proper genetic variants. Both of these karst
areas are similar in that they contain large, integrated subterranean
drainage systems. On the other hand, Form II populations are also
sometimes found in caves that are components of extensive subterranean
drainage systems and, in some instances, not far removed geographically
from caves with Form I populations.

Genetic studies on G. minus , principally in the Greenbrier Valley of
West Virginia, by Hetrick (1975), Hetrick and Gooch (1981), and Gooch
and Hetrick (1979), in which allozyme allele frequencies were determined
at three polymorphic loci, tend to support the ecophenotype concept.
The results of these studies indicate that there is generally a greater
genetic distance between populations of the same ecophenotype in
different geographic areas than between populations of different
ecophenotypes in the same small, defined geographic area, and that
most of the sharper discontinuities coincide closely with potential
barriers to dispersal, such as streams, stream and karst drainage divides,
and stratigraphic changes.

Whether the troglomorphic populations in Ward Cove and the
Great Savannah are already incipient troglobitic species, are on their
way to becoming separate species, or simply represent the extreme
expression of a highly plastic phenotype is debatable and cannot be
resolved on the basis of the information presently available. Further
study is clearly needed.

Terrestrial Species. — The origin of terrestrial troglobites has perhaps
been more direct and has involved fewer variables than that of aquatic
troglobites. Most workers agree that the troglobitic terrestrial fauna of
the north temperate region (viz., North America, Europe, and Japan)
has been derived from preadapted, epigean ancestors that occupied
moss, ground litter, and deep-soil habitats of humid forest floors (see
Vandel 1965b, Barr 1968, Peck and Lewis 1978, Peck 1981b, Culver
1982, Barr and Holsinger 1985). Moreover, the invasion and colonization
of caves by terrestrial invertebrates has probably been, and still is, an
ongoing process, involving the dynamics of taxon cycles and pulses (see
also Peck 1980, 1981b). With few exceptions, all terrestrial troglobites in

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135

the study area belong to higher taxa that are simultaneously well
represented by cryptozoic, epigean species living in the cool, moist litter
microhabitats of montane forests of the Appalachians. Furthermore, the
greater Appalachian region, because of its rich diversity of both habitats
and biota, has been suggested as the site of origin for much of the
ancestral stock of the terrestrial troglobite fauna of the entire eastern
United States (Peck and Lewis 1978).

The widely accepted model for the origin of terrestrial troglobites
in temperate regions of the Northern Hemisphere is based on climatic
fluctuations in the Pleistocene (Barr 1967a, 1968, 1973, 1985; Poulson
and White 1969; Peck 1981b; Culver 1982). According to this hypothesis,
during periods of glacial maxima, the cold, moist areas lying south of
glaciation, such as the southern Appalachians, would have provided a
suitable environment for the extensive distribution of a cryophilic
endogean fauna. Both caves and ecologically suitable surface habitats
would have been colonized by this fauna. During interglacial periods,
when the regional climate became warmer and drier, many elements of
this fauna would have become extinct at the surface, especially at low
elevations, but other elements would have survived in caves and at high
elevations in cool-mesic forests. The extinction of surface populations at
low elevations during interglacials would have resulted in the genetic
isolation of founder populations in caves, because migration and gene
exchange between epigean and hypogean populations would have been
eliminated in many karst areas. Ultimately this series of events would
have led to the evolution of troglobitic species, depending on the length
of time of physical isolation underground and whether or not certain
populations subsequently reinvaded suitable surface habitats during
succeeding glacial advances (c.f., the taxon cycle of Peck 1980).
Eventually, however, isolation was completed for many cave populations.
Since the onset of the Pleistocene, there has probably been a sequence
of invasions and colonizations of caves by preadapted, troglophile
ancestors and concomitant extirpations of closely related epigean
populations. The detailed evolutionary history of any troglobitic group,
however, must be relatively complex, because, as Peck (1980) suggests,
many groups have probably passed through a taxon cycle that first
involved isolation of populations in caves, followed by expansion into
epigean habitats and, then, ultimately by isolation again in caves during
a succeeding interglacial.

One of us (Culver 1982) recently reviewed the evidence in favor of
the Pleistocene climatic-effect paradigm and concluded that, although
indirect, the evidence supporting the hypothesis was strong. Nonetheless,
this hypothesis has recently been questioned by several workers. Based
on studies of the newly discovered troglobitic fauna of Hawaiian lava
caves, Howarth (1980, 1981) suggested that troglobitic organisms have

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evolved, in the tropical Hawaiian Islands at least, through adaptive
shifts of preadapted ancestors into newly opened niches and not by
isolation of troglophilic ancestors during climatic shifts in the Pleis-
tocene. Although Howarth’s theory is based on data from Hawaiian
lava tubes, which are generally much younger geologically and contain a
significantly different food supply than limestone caves in temperate
regions, he believes that his theory can be extended to explain to a large
extent the evolution of terrestrial troglobites in temperate regions,
where, as he points out, a complex geological history and glaciations
have obscured the early history and obfuscated the previous distribution
and evolution of troglobites. In Hawaii, cave populations may be larger
than those of their surface relatives because the colonizable subterranean
habitat is much larger in area than the rain forest or new lava substrate
habitats on the surface (Howarth 1980). Howarth also suggests that an
analogous situation may exist in limestone karst areas as well, but the
data from our present study neither confirm nor refute this.

In her research on cave spiders in the temperate region of southern
Europe, Deeleman-Reinhold (1981) concluded that physical properties
of the subterranean environment and the present areal climate have
been the principal factors in the evolution of the high diversity of
troglobitic species in the southwestern Yugoslavian karst. Her conclusion
also raises a serious question about the effect of past climates on the
evolution of terrestrial troglobites, specifically in temperate karst areas.

One of the arguments made in the past in support of the Pleistocene
climatic-effect theory was that terrestrial troglobites are far more
abundant in the temperate zone than in the tropics (Barr 1968). It was
previously assumed that terrestrial troglobites were extremely rare in
tropical areas (see Vandel 1965b, Barr 1968, Mitchell 1969), but the
recent discovery of rich terrestrial troglobitic faunas in Hawaii (Howarth
1972), Jamaica (Peck 1975d), and the lowlands of Mexico and Central
America (Reddell 1981) has proven otherwise. Because the effects of
climatic fluctuations in the Pleistocene were probably different at low
elevations in the tropics than in temperate zones, the Pleistocene
climatic-effect model is questionable for areas outside temperate karst
regions and high elevations in the tropics; thus, other explanations for
the origin of terrestrial troglobites in the lowland tropics may be
warranted. One such explanation is Howarth’s adaptive-shift hypothesis,
discussed above.

The adaptive-shift theory is attractive because it can be applied to
all parts of the world. Therefore, it eliminates the need for different
paradigms for different regions and is applicable to both terrestrial and
aquatic troglobites. It is not necessarily an allopatric model, however,
but infers a kind of parapatric speciation in which new troglobitic
species can arise in the absence of complete physical isolation between
epigean ancestral populations and hypogean founder populations.

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137

Barr (1965, 1967a, 1967c, 1968, 1973, 1981a, 1985), on the other
hand, based on his studies on the geographic distribution and ecology of
troglobitic trechine beetles, has made a convincing case for the
Pleistocene climatic-effect model. Barr (1967b, 1968) has also made a
strong argument for the allopatric speciation process in the evolution of
troglobites and has suggested that after isolation of a founder population
in a cave or series of interconnected caves, following the extinction of
epigean ancestors, the newly isolated cave colony will pass through a
period of lowered genetic variability (genetic bottleneck). Moreover, if
the colony survives, an extensive genetic reorganization will result in a
reconstructed epigenotype, the end point of which is a well-adapted
troglobite. Genetic, studies by Sbordoni et al. (1981) on cave crickets in
southern Europe tend to support the bottleneck effect in the evolution
of cave species, but the degree to which an epigenotype is reconstructed
in the evolution of a troglobite remains unclear. It is entirely possible,
however, that genetic differences between troglobites and epigean
congeners have been overstressed, despite the prominent regressive
features that develop almost universally in highly specialized cavernicoles.

It is beyond the scope of this paper to debate the pros and cons of
allopatric versus parapatric speciation. But it should be pointed out
that, whereas allopatric speciation is still favored over parapatric
speciation for most groups of organisms, a rather strong case has been
made for parapatric speciation (sensu Bush 1975, Endler 1977) in some
groups under certain conditions, and it cannot be ruled out as a possible
mode of evolution for some troglobites.

Trechine beetles of the genus Pseudanophthalmus are taxonomically
the most numerous and thoroughly studied terrestrial troglobites in the
study area and therefore provide good data for zoogeographic analyses.
According to Barr (1981a, 1981b), ancestors of troglobitic species
presently living in caves of both the Appalachian Valley and Ridge and
the Interior Low Plateaus probably originated in upland forests of the
Appalachian Plateau and spread out under periglacial climates. Caves
were colonized at the beginning of interglacial periods. The ancestors
were probably edaphobites already strongly preadapted for a cave
existence. An earlier hypothesis by Jeannel (1949) suggested that
ancestors spread out from an interglacial refugium in the Unaka
Mountains along the Tennessee-North Carolina border, but Barr has
made a more convincing case for an Appalachian Plateau center of
distribution. Barr’s theory is based principally on dissimilarities of
species on opposite sides of the Appalachian Plateau, the increased
richness of species closer to the plateau front in the Appalachian Valley,
and the occurrence of a single edaphobitic species ( Pseudanophthalmus
sylvaticus ) in the Plateau and not in the Unakas. The Unaka hypothesis
was not discarded altogether by Barr, however, since, as he points out,
distributions of the engelhardti and petrunkevitchi groups are not

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incompatible with an origin in the higher mountains on the eastern side
of the Appalachian Valley.

The presence of vestigial eyes in some species of Pseudanophthalmus
viz., members of the petrunkevitchi group and P. vicarius ) suggests a
fairly recent invasion of caves by some species (Barr 1965) if the degree
of eye reduction is a crude measure of the length of time a species has
lived in a cave. On the other hand, the level of intrageneric diversity and
the occurrence of many distinct species groups suggest the possibility
that Pseudanophthalmus is much older than the Pleistocene (see Barr
1981a). Given this background, one might postulate that the colonization
of caves by species of Pseudanophthalmus has taken place over a long
period of time through a succession of independent invasions. The
occurrence of many distinct species groups, some of which broadly
overlap geographically in southwestern Virginia and eastern Tennessee,
suggests several independent colonizations of caves by ancestral stocks.
How closely these colonizations might have coincided with the beginning
of Pleistocene interglacials is difficult to determine, however.

The presence of P. sylvaticus in a non-cave habitat is of zoogeo-
graphic interest because this species is the only non-troglobitic
Pseudanophthalmus recorded from North America. Barr (1967c, 1969)
believes it is probably a periglacial relict that survived in the ecologically
suitable habitat of a cold mountain forest during one of the interglacials
when many of its congeners either were extirpated on the surface by a
progressively warmer and drier climate or survived by colonizing caves
at low elevations. Pseudanophthalmus sylvaticus was collected and
described by Barr (1967c) from an endogean habitat in the Yew
Mountains, approximately 36 km west of the study area on the eastern
margin of the Appalachian Plateau in Pocahontas County, W.Va. This
species, which has rudimentation of both eyes and pigment, is an
edaphobite, presumably closely similar to putative preadapted ancestors
of troglobitic members of the genus. It is not far removed taxonomically
from some of the present cave forms living in limestone areas just to the
east.

Although Barr (1967c) suggested that the discovery of P. sylvaticus
supports the Pleistocene climatic-effect theory, we believe that it could
also support Howarth’s adaptive-shift theory. For example, if preadapted
species of Pseudanophthalmus colonized caves in response to newly
opened niches, it is unlikely that all members of the genus would have
gone underground. Those left behind on the surface could have persisted
in ecologically suitable habitats like that of P. sylvaticus in the Yew
Mountains. In reality, neither hypothesis is falsified by the discovery of
P. sylvaticus , since both predict the occurrence of preadapted epigean
congeners in groups with troglobitic species.

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139

With the exception of P. nelsoni, for which there is an apparent
identity problem (see Barr 1965:45), the ranges of all species of
Pseudanophthalmus in the study area are restricted to continuous
exposures of cavernous limestone. As already pointed out, most of the
species are known from single caves or small clusters of caves, although
a few, like P. delicatus, P. gracilis , P. hoffmani, and P. rotundatus, have
significantly larger ranges. Closely delimited ranges that coincide with
continuous exposures of limestone strongly suggest that the dispersal of
these beetles is limited to caves, solution channels, and other openings
in carbonate bedrock. The two troglobitic pselaphid beetles from the
study area also have highly restricted ranges, and each is known only
from a single cave. These species were probably derived directly from
edaphobitic ancestors in relatively recent times. Epigean congeners live
in damp, deciduous-forest floors or in deep soil; in the genus Arianops,
both epigean and hypogean species are eyeless (see Park 1960, 1965;
Barr 1974).

Troglobitic pseudoscorpions, like troglobitic trechine and pselaphid
beetles, also have narrowly circumscribed ranges, and most are known
only from single cave localities. Subterranean dispersal is apparently
highly restricted and limited to continuous belts of cavernous limestone.
Chamberlin and Malcolm (1960) concluded that the pseudoscorpion
cave fauna is derived from epigean (endogean) forms in the same
general geographic area. The highly localized distribution of the
cavernicolous species tends to support their conclusion.

Muchmore (1981) has pointed out that all of the troglobitic species
of Kleptochthonius (29 described species in the subgenus Chamberlin-
ochthonius) are restricted to the southeastern cave region in Kentucky,
Tennessee, Virginia, West Virginia, and southern Indiana, and that
troglobitic species in other chthoniid genera (viz., Apochthonius and
Mundochthonius in the study area) occur on the periphery of the range
of cavernicolous Kleptochthonius with little or no overlap. This suggests
the possibility of competitive exclusion of other cavernicolous pseudo-
scorpions by the strongly troglomorphic species of Kleptochthonius.
The range of Kleptochthonius ( Chamber lino chthonius ), like that of
Pseudanophthalmus , forms a trans-Appalachian distributional track,
extending from the Interior Low Plateaus on the west to the Appalachian
Valley and Ridge on the east. By comparison, troglobitic species of
Apochthonius and Microcreagris are very widely scattered (Muchmore
1981), but troglobitic Chitrella and Mundochthonius are rare and
represented by only a few species (Malcolm and Chamberlin 1960,
Muchmore 1973, Benedict and Malcolm 1974).

With few exceptions, the ranges of other troglobitic arthropods
(e.g., isopods, mites, spiders, millipeds, collembolans, and diplurans) in

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the study area do not show the same high degrees of restriction to
isolated, continuous belts of limestone as do those of beetles and
pseudoscorpions. Populations of many of these species are found in
caves developed in discontinuous exposures of limestone physically
separated by clastic rocks. Assuming, however, that some gene exchange
takes place between cave populations of the same species in different
karst areas, then limited dispersal through areas composed of non-
calcareous rock must occur.

Recent discoveries of troglobites in non-calcareous caves and
artificial mine adits in Japan by Ueno (1977) and in shallow underground
compartments in Europe by Juberthie and Delay (1981) indicate how
subterranean dispersal may occur outside caves per se. Ueno found
troglobitic beetles, isopods, millipeds, and spiders in natural cavities and
artificial mines excavated in fissured, non-calcareous rocks in Japan. In
the Alps, Pyrenees, and Carpathians, Juberthie and his colleagues
discovered troglobitic beetles, millipeds, isopods, and spiders in a distinct
habitat type they named the shallow underground compartment (S.U.C.).
Most of these species had been recorded previously from nearby caves.
According to Juberthie and Delay (1981), the S.U.C. exists under the
last layer of soil in mountainous areas and consists of cracks and
fissures in the mantle rock. These cracks and fissures are in turn
connected to caves and/or deep cracks that represent the deep
underground compartment. In non-limestone areas, the S.U.C. was
usually identified in schists; in limestone areas it was commonly
associated with screes or talus.

The observations by Ueno (1977) and Juberthie and Delay (1981)
are good evidence that many troglobites inhabit shallow fissures and
crevices near the surface in non-cavernous areas. Although not yet
specifically identified, similar conditions probably exist in the Appala-
chians. We do have good evidence, however, that some terrestrial
troglobites in the study area occur outside caves and are therefore able
to move between caves situated in different exposures of limestone.
Both spiders ( Nesticus tennesseensis ) and collembolans {Pseudo sinella
hirsuta and Sinella hoffmani ) have been collected from deep ground-
litter habitats in forested areas on mountainsides outside limestone
terranes (Barr 1967c, Christiansen and Bellinger 1980c, Gertsch 1984,
and elsewhere this paper). It will not be surprising if other troglobitic
species are eventually found in similar habitats, either in deep ground
litter or under conditions analogous to those described by Ueno (1977)
and Juberthie and Delay (1981).

Of the three troglobitic trichoniscid isopods in the Virginia-east
Tennessee area, Miktoniscus racovitzai is fairly widely distributed,
whereas Amerigoniscus henroti and A. paynei have relatively limited
ranges. The range of A. henroti is restricted to caves in a continuous

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141

exposure of cavernous limestone in the Powell Valley of Lee County
(see Holsinger 1967b). In the adjacent Clinch Valley, A. paynei, a
probable sister species, inhabits caves that are developed in several
separate exposures of limestone.

Miktoniscus racovitzai (5. lat.), the only eyeless, troglobitic member
of its genus, is closely allied morphologically with epigean congeners in
the eastern United States (see Vandel 1950). It is probably a relatively
recent derivative of a widespread, preadapted troglophile ancestor. In
comparison, Amerigoniscus comprises 10 eyeless, unpigmented species,
of which nine are troglobites and seven are recorded from single
localities. The widespread, highly disjunct distribution of the species in
this genus (viz., three from northwestern Georgia, one from south-
central Oklahoma, two from Oregon, one from middle Tennessee, one
from northwestern Texas, and two from the study area; see Vandel
1965a and 1977, Schultz 1982), combined with the fact that all are of
troglobitic facies, suggests that members of this genus are old, isolated,
subterranean relicts of a formerly widespread surface fauna. With the
exception of A. rothi (Vandel) from an endogean habitat (under rocks
and moss in a dense forest; see Vandel 1953) in Curry County, Oregon,
no other epigean congener is known.

Recent studies by Zacharda (1980, 1985) indicate that a majority of
the cavernicolous rhagidiid mites in the North American and European
faunas are troglophiles and that only a few species have well-developed
troglomorphisms and are restricted to caves. Because the family
Rhagidiidae is predominantly edaphic and some of the edaphobites
occur in caves, it is reasonable to assume that the troglobites are
relatively recent derivatives of soil-dwelling forms. Of the two species
considered troglobitic in the study area, one ( Foveacheles paralleloseta )
is known only from a single cave, whereas the other ( Rhagidia viria ) has
a much broader distribution and is recorded from caves in several
drainage basins.

None of the eight spiders considered troglobitic in the study area
has a range that is limited to a single, continuous exposure of limestone.
As already mentioned, the ranges of the troglobitic linyphiid spiders are
among the most extensive of all troglobites in North America. Several
explanations for these broad ranges have been suggested (Holsinger
1963a, Barr 1967a, Holsinger et al. 1976, and elsewhere this paper), but
until the genetics of the species are studied, nothing definitive can be
said. However, the presence of eyes (although variable) and some
pigment in many populations, combined with the wide ranges, strongly
indicates that these species are recently evolved troglobites.

Compared with linyphiids, nesticid spiders have much smaller
ranges. Troglobitic nesticids show varying levels of eye and pigment
reduction and appendage attenuation. Gertsch (1984) has pointed out

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John R. Holsinger and David C. Culver

that Nesticus tennesseensis has more reduced eyes, lighter pigmentation,
and longer legs in the southern part of its range than in the northern
part. In addition, this species has been collected occasionally from deep
ground litter outside caves in the northern part of its range. Of the
three troglobitic nesticids in the study area, Nesticus holsingeri has the
most reduced eyes and pigmentation and also the most limited range.
Reductions in eye structure, pigment, and geographic distribution may
be positively correlated with a relatively advanced level of cave
specialization.

In the eastern United States Nesticus is represented by 24 species
that inhabit both caves and the rich ground litter of mesic forest floors,
largely in the southern Appalachian region (Gertsch 1984). Troglobitic
nesticids have apparently evolved from troglophilic ancestors as the
latter became progressively more restricted to caves. The moderately
widespread troglophile Nesticus earteri may very well be a good example
of a troglobite in statu nascendi in a portion of its range, inasmuch as it
is represented by numerous cavernicole populations, some of which are
large and feed and reproduce in caves.

Millipeds constitute one of the most significant groups of
cavernicoles in the eastern United States, but unfortunately they remain
one of the most underworked taxonomically. Probably more than 50%
of the species known from caves are undescribed, which makes
zoogeographic analysis difficult. The genus Pseudotremia is represented
by many species that inhabit both caves and epigean habitats in parts of
the Appalachian Valley, Appalachian Plateau, and Interior Low
Plateaus. The range of the genus forms a distributional track across the
Appalachian Plateau similar to that of Pseudanophthalmus and
Kleptochthonius ( Chamberlinochthonius ). Of the 34 species of Pseudo-
tremia recognized by Shear (1972), 15 are obvious troglobites, 7 are
questionable troglobites, 8 are troglophiles, and 4 are apparently strictly
epigean. Ranges of both the troglobites and the troglophiles are generally
localized; but without further taxonomic refinements, it cannot be
determined how closely the geographic distributions of troglobites
coincide with isolated exposures of limestone.

In the study area, the most highly specialized troglobitic pseudo-
trimiids are in the nodosa complex, a group of closely allied species that
are unpigmented and have greatly reduced eyes (ca. 10-11 ocelli per
eye). They are common in caves in the Clinch and Powell valleys.
Presumably, colonization of caves by members of this complex predates
that of the less specialized troglobites, which are pigmented and have
more ocelli. In caves of the Clinch and Powell valleys, it is not
uncommon to find both pigmented and unpigmented species in the
same cave, but the latter (species of the P. nodosa complex) are usually
more abundant and often occur at greater distances from cave entrances.

Invertebrate Cave Fauna

143

The milliped genus Trichopetalum has a much broader geographic
distribution and fewer species (ca. 15) than Pseudotremia (see Shear
1972). Troglobites are unpigmented and completely eyeless. Scoterpes is
closely related to Trichopetalum and contains perhaps 30 troglobitic
species (many undescribed) that inhabit caves to the south and west of
the study area in Alabama, Georgia, Illinois, Kentucky, Missouri, and
Tennessee (Causey 1960b; Shear 1969, 1972). Trichopetalum contains
five troglobites and three troglophiles. Four of the troglobites occur in
the Appalachians of Virginia and West Virginia, and one is known from
northern Alabama (Causey 1960a, Shear 1972). The troglophiles occur
in Alabama, Kentucky, Maryland, and Oklahoma (see Causey 1967,
1969; Shear 1972), and all species possess eyes and pigment.

The three species of Trichopetalum in the study area are apparently
very closely related genetically, as suggested by the possibility of
hybridization between some of the populations (see Causey 1963 and
elsewhere this paper). This possibility, combined with the contiguous
distribution and closely similar morphologies of the species, suggests a
relatively recent common ancestor, possibly involving a moderately
widespread humicolous epigean form that invaded caves over parts of
western Virginia and eastern West Virginia. A fourth species, T. krekeleri
(Causey), from caves in Randolph and Tucker counties, W.Va., is
distinct, but it was probably derived from the same ancestor.

A majority of the North American troglobitic collembolans are in
the Entomobryinae genera Pseudosinella and Sinella. Three troglobites
(viz., P. hirsuta, P. orba, and S. hoffmani ) and a number of
troglophiles/ trogloxenes occur in the study area. In two recent papers
on the zoogeography of eastern North American cave collembolans,
Christiansen (1981, 1982) assigned caves in the Appalachian Valley and
Interior Low Plateaus region to a category he called “heartland caves.”
Two other categories in the eastern United States were designated
“glaciated area caves” and “non-glaciated non-heartland caves.” As
might be expected, the most highly specialized troglobitic collembolans
(based on degree of troglomorphy) generally occur in heartland caves.
In an earlier paper, Christiansen (1961) recognized two types of
characteristics in cave species: cave-dependent and cave-independent
characters. Using cave-dependent characters as a basis to measure
evolutionary changes leading to an increase in troglomorphy, he devised
a seven-step evolutionary scale for the cave Entomobryinae, with step 7
representing the highest level of adaptation. On this scale, Pseudosinella
hirsuta was considered to be in step 5; P. orba and Sinella hoffmani
were in step 6.

Troglobitic collembolans in the study area have relatively extensive
ranges, and none is restricted to caves in a single exposure of limestone.
Troglobites, however, have much more compact ranges than troglophiles.

144

John R. Holsinger and David C. Culver

In the Entomobryinae, Christiansen (1981, 1982) postulated several
evolutionary lineages for each genus. In Sinella , 5. hoffmani is placed in
a lineage with the troglophile S. barri. In Pseudosinella , P. hirsuta, P.
orba , and P. argentea (a troglophile) are each placed in a separate
lineage.

To account for the extensive range of P. hirsuta (parts of Alabama,
Georgia, Kentucky, Tennessee, and Virginia), Christiansen and Culver
(1968) and Christiansen (1982) postulated a process they termed “parallel
speciation,” which was envisioned as having resulted from the invasion
of caves by an ancestor over a wide geographic area, followed by
independent parallel development of the same morphology in separate,
physically isolated lineages. The end products would resemble each
other so precisely in behavior, morphology, and ecology that they could
be called the same species. According to this interpretation, P. hirsuta
would have to be regarded as a complex of several biological (sibling?)
species. Populations in the study area, although apparently morpholo-
gically indistinguishable from those farther west in Kentucky and middle
Tennessee, are probably genetically distinct. Christiansen (1982)
interprets the rather wide and discontinuous range of S. hoffmani as
either the result of parallel speciation or representative of the vestiges of
a previously continuous range. However, both P. hirsuta and S. hoffmani
have been found outside caves on rare occasions, and the possibility
that their wide ranges have resulted in part from dispersal between karst
areas through shallow underground compartments or similar endogean
habitats cannot be dismissed.

According to Christiansen (1982), the evolution of P. orba , which
has a more limited geographic distribution than either P. hirsuta or S.
hoffmani, has probably resulted from the single invasion of caves by a
putative ancestor and subsequent subsurface dispersal to the present
limits of its range. In the genus Arrhopalites, Christiansen (1982)
suggests that the troglomorphic A clarus, recorded from caves in both
the Ozarks and the Appalachian Valley, might be the product of parallel
speciation at least twice from a common, widespread ancestor.

The dipluran genus Litocampa contains 32 species worldwide; a
majority (20) inhabit caves in the United States, and all are troglobites
(Ferguson 1981a, 1981b). The wide geographic distribution of the genus
(viz., parts of Africa, Europe, and North and South America), combined
with retention of certain characters judged to be primitive for the order,
has led Ferguson (1981a, 1981b) to suggest that its origin may predate
the breakup of the supercontinent Pangaea in the Mesozoic. The
absence of epigean congeners anywhere in North America suggests that
troglobitic species of Litocampa are probably relatively old cavernicoles.
Moreover, based on the richness of species and number of endemics in
the southern Appalachian region, Ferguson has suggested this area as a

Invertebrate Cave Fauna

145

probable center of distribution for species of Litocampa in the United
States. Of the 20 species currently recognized from the United States, 17
occur in the greater Appalachian region, and their combined ranges
form a distributional track from the Interior Low Plateaus to the
Appalachian Valley and Ridge (see Ferguson 1981a: Fig. 43).

Of the six species of Litocampa in the study area, only two have
narrowly circumscribed ranges that coincide with isolated exposures of
limestone. The other species have wider ranges, although all except L.
cookei, whose range extends as far west as central Kentucky and middle
Tennessee, have relatively localized ranges confined to the study area or
its periphery. The extensive distribution of L. cookei, the largest of any
troglobitic dipluran in North America, is puzzling. Its distribution is not
contiguous, however, but occurs in five disjunct clusters (Ferguson
1981a: Fig. 43). Ferguson (1981a) has studied the morphology of this
species in detail and has concluded that it may represent a complex of
allopatric sibling species.

Two final points should be made with respect to the origin of
terrestrial troglobites. (1) The geographic distributions of four genera —
viz., Kleptochthonius (Chamber lino chthonius), Litocampa, Pseuda-
nophthalmus, and Pseudotremia — represented collectively by numerous
troglobites in the study area are nearly congruent and together form a
strong generalized distributional track that extends across the Ap-
palachian Plateau. The importance of generalized tracks in biogeographic
analysis has been reviewed by Wiley (1981). Such tracks may be used to
estimate the range of ancestral species in monophyletic groups with
similar distributions.

The possibility suggested by Barr (1981a) that ancestors of
troglobitic species of Pseudanophthalmus originated in the forest floors
of the Appalachian Plateau in late Cenozoic times, with subsequent
thrusts into limestone areas on either side, was discussed above. The
coincident distributions of Kleptochthonius (Chamber lino chthonius),
Litocampa, and Pseudotremia suggest a similar place of origin and
center of distribution for ancestors of troglobites in these groups as well.
Shear (1972) alluded to the possibility that Pseudotremia originated in
the southern Appalachian Mountains through evolution from a proto-
Pseudotremia stock in the Cenozoic. Similarly, Ferguson (1981a)
suggested that the southern Appalachians might have been the center of
distribution for North American species of Litocampa. The generalized
track formed by these taxa tends to support these ideas and points to
the central and southern parts of the Appalachian Plateau as an
important geographic center for the distribution of ancestors of terrestrial
troglobites in the Appalachian Valley, Interior Low Plateaus, and
limestone areas on the eastern and western sides of the Appalachian
Plateau.

146

John R. Holsinger and David C. Culver

(2) Some common caves species in the study area, such as the
spider Meta menardi, the cricket Euhadenoecus fragilis, and the dipterans
Amoebalaria defessa and Megaselia cavernicola, are apparently well
adapted in their present roles as troglophiles or (habitual) trogloxenes
and show no evidence of evolving into troglobites. Other troglophiles,
however, such as the harvestman Erebomaster acanthina, the spider
Nesticus carteri, and certain species of the milliped Pseudotremia, show
good evidence of becoming bona fide troglobites, and, as such, they are
probably troglobites in statu nascendi.

ACKNOWLEDGMENTS. — We are extremely grateful to the many
individuals who have assisted us in various phases of the field work or
furnished us with useful data from their own collections. In particular
we thank R. A. Baroody, T. C. Barr, Jr., J. H. Carpenter, J. M. Beck,
III, L. G. Conrad, J. E. Cooper, G. D. Corbett, J. Cox, G. W. Dickson,
J. E. Estes, L. M. Ferguson, D. L. Finley, S. W. Hetrick, R. L.
Hoffman, L. R. Hubricht, T. C. Kane, C. H. Krekeler, P. C. Lucas,
G. H. Marland, T. G. Marsh, the late D. R. Martin, the late J. P. E.
Morrison, S. B. Peck, R. D. Powers, Jr., R. M. Norton, J. E. Tichenor,
V. M. (Tipton) Dalton, R. L. Wallace, and R. E. Whittemore.

Many systematists aided in the determination of specimens, and
their expertise and helpful comments are appreciated. We thank: T. C.
Barr, Jr., beetles; T. E. Bowman, isopods; T. S. Briggs, opilionids; J. H.
Carpenter, flatworms; the late N. B. Causey, milllipeds; K. A.
Christiansen, collembolans; D. G. Cook, oligochaetes; R. E. Crabill, Jr.,
centipedes; D. A. Crossley, Jr., mites; W. R. Elliott, mites; L. M.
Ferguson, diplurans; L. E. Fleming, isopods; B. A. Foote, dipterans;
R. J. Gagne, dipterans; G. E. Gates, oligochaetes; W. J. Gertsch,
spiders; C. J. Goodnight, opilionids; A. B. Gurney, crickets; L. J.
Herman, Jr., beetles; R. Hershler, snails; H. H. Hobbs, Jr., crayfishes;
R. L. Hoffman, millipeds; P. C. Holt, branchiobdellids; T. H. Hubbell,
crickets; L. R. Hubricht, snails; L. H. Hyman, flatworms; R. Kenk,
flatworms; J. J. Lewis, isopods; D. R. Malcolm, pseudoscorpions; S. A.
Marashall, dipterans; A. C. Mickelbacher, symphylans; the late J. P. E.
Morrison, snails; W. B. Muchmore, pseudoscorpions; S. B. Peck,
beetles; G. A. Schultz, isopods; W. A. Shear, millipeds and opilionids;
D. E. Sonenshine, ticks; H. R. Steeves, III, isopods; G. C. Steyskal,
dipterans; F. C. Thompson, dipterans; F. G. Thompson, snails; the late
A. Vandel, isopods; W. C. Welbourn, mites; W. W. Wirth, dipterans;
D. L. Wray, collembolans; H. C. Yeatman, copepods; and M. Zackarda,
mites.

We are indebted to numerous landowners in Virginia and eastern
Tennessee who so willingly cooperated with us in allowing access to

Invertebrate Cave Fauna

147

their caves. Many owners went out of their way to assist with the field
work. We are also grateful to the managements of the privately operated
commercial caves in Virginia and to officials of Cumberland Gap
National Historical Park, Natural Tunnel State Park, and the Upper
Valley Regional Park Authority for giving us free access to work in the
caves under their jurisdiction. J. R. Jordan, Jr., and W. H. Redman of
the Regional Heritage Program of the Tennessee Valley Authority
(TV A) are thanked for their encouragement and support of our research
in southwestern Virginia and eastern Tennessee. D. L. Miller-Carson, of
the Old Dominion University Center for Instructional Development,
and B. A. McBride, of the TVA Division of Natural Resources Services,
are thanked for their assistance with preparation of the maps. Certain
parts of this paper have benefited from discussions and/or the exchange
of information with T. C. Barr, Jr., E. M. Benedict, K. A. Christiansen,
L. M. Ferguson, F. G. Howarth, T. C. Kane, J. J. Lewis, S. B. Peck,
T. L. Poulson, J. R. Reddell, F. D. Stone, and many others.

This study was supported in part by grants from the Research
Advisory Committee of the National Speleological Society, the National
Science Foundation (GB-42332), and Old Dominion University (Faculty
Summer Research Grant in 1979) and by travel funds from the
Department of Invertebrate Zoology of the Smithsonian Institution
(summer of 1972) to J. R. Holsinger; by grants from the National
Science Foundation and Northwestern University to D. C. Culver; and
by a personal service contract from the Tennessee Valley Authority (TV
51940A) in 1979 to both of us. This paper is the final contribution of the
Cave Biogeography of the Central Appalachians Study Group, a former
research project of the National Speleological Society.

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Accepted 16 February 1987

163

NEW EDITOR

Frank J. Radovsky, Curator of Research and Collections at the
North Carolina State Museum of Natural Sciences, has been named
Editor of Brimleyana, effective 15 January 1988. Eloise F. Potter, who
has been Acting Editor of the journal since the resignation of John E.
Cooper, will continue in that capacity through the publication of
Brimleyana No. 15 and will serve as Managing Editor under Dr.
Radovsky.

During the transition period, Radovsky will conclude his editorship
of the Journal of Medical Entomology, which is published by the
Entomological Society of America. In December 1987 he was named to
a 5-year term on the Editorial Board of that journal. He has served as
an Associate Editor of the Annual Review of Entomology for 10 years,
and he was recently appointed to another 5-year term. Radovsky is on
the Executive Committee of the Acarological Society of America, and
he formerly served on the Board of Directors of the Association of
Systematics Collections (1982-1985) and as Executive Secretary of the
International Congress of Acarology (1971-1978).

Prior to coming to the North Carolina State Museum of Natural
Sciences, Radovsky was on the staff of the Bishop Museum in Honolulu,
Hawaii (1969-1986), where he was Assistant Director (1977-1985) and
holder of the L. A. Bishop Distinguished Chair of Zoology (1984-1986).
From 1986 to 1987, he was Visiting Professor of Entomology at Oregon
State University, Corvallis.

Radovsky received an A.B. degree in Zoology from the University
of Colorado, Boulder, and M.S. and Ph.D. degrees in Parasitology
from the University of California, Berkeley. His current research interests
include the systematics and ecology of mites, ticks, and fleas.

DATE OF MAILING

Brimleyana No. 13 was mailed on 16 July 1987.

164

ENDANGERED, THREATENED, AND
RARE FAUNA OF NORTH CAROLINA

PART I.

A RE-EVALUATION OF THE MAMMALS
Edited by Mary Kay Clark

This book is a report prepared by a committee appointed in 1985
by the North Carolina State Museum of Natural Sciences to re-evaluate
the list of mammals presented in Endangered and Threatened Plants
and Animals of North Carolina (John E. Cooper, Sarah S. Robinson,
and John B. Funderburg, editors. N.C. State Mus. Nat. Hist., Raleigh,
1977), which is now out of print. Committee members were Mary Kay
Clark, David A. Adams, William F. Adams, Carl W. Betsill, John B.
Funderburg, Roger A. Powell, Wm. David Webster, and Peter D.
Weigh The report treats 21 species listed in the following status
categories: Endangered (5), Threatened (1), Vulnerable (6), and
Undetermined (9). Most species accounts discuss the animal’s physical
characteristics, range, habitat, life history and ecology, special sig-
nificance, and status (including the rationale for the evaluation and
recommendations for protection) and provide a range map and an
illustration of the animal’s external characters. Ruth Brunstetter and
Renaldo Kuhler illustrated the book. An introductory section contributed
by Ms. Clark discusses the changes in status that occurred in the decade
between 1975 and 1985. It also mentions efforts to protect marine
mammals and includes a checklist of the cetaceans known from North
Carolina.

1987 52 pages Softbound ISBN 0-917134-14-1

Price: $5 postpaid. North Carolina residents add 5% sales tax. Please make
checks payable in U.S. currency to NCDA Museum Extension Fund.

Send order to: ETR MAMMALS, N.C. State Museum of Natural Sciences,

P.O. Box 27647, Raleigh, NC 27611.

INFORMATION FOR CONTRIBUTORS

Submit original and two copies of manuscripts to Editor, Brimleyana, North Carolina
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Preparation of manuscript. Adhere generally to the Council of Biology Editors Style
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Where appropriate, the title will indicate at least two higher categories to which taxa
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Woodall, W. Robert, Jr., and J. B. Wallace, 1972. The benthic fauna in four small
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for proofreading.

BRIMLEYANA NO. 14, JUNE 1988

CONTENTS

The Invertebrate Cave Fauna of Virginia and a Part
of Eastern Tennessee: Zoogeography and Ecology.
John R. Holsinger and David C. Culver

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