58 
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 
