78 
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 
