Appalachian Pseudanophthalmus 39 
Pseudanophthalmus Jeannel 
ENGELHARDTI COMPLEX 
Integuments generally pubescent, although dorsum of head and prono- 
tum disc subglabrous in a few species. Elytral microsculpture always 
more or less transverse, usually a little confused, often obsolescent on 
central disc, rarely forming distinct meshes; elytra not pruinose. Anterior 
discal puncture at or near level of 4th umbilicate puncture. Apical recur- 
rent groove elongate, oblique or subparallel, usually bisinuate (except 
alabamae group) and usually connected to apex of 3rd stria with or 
without a crosier. Humeri setulose, rarely serrulate. Mentum tooth short, 
usually emarginate. Mesosternum simply declivous. Profemur unmodi- 
fied. Last abdominal sternite of males with apical margin entire. Transfer 
apparatus consisting of an elongate, canoe-shaped right piece (sometimes 
with dorsal keel) partly enfolding a smaller, usually shorter, elongate- 
triangular, spinulose left piece. 
Distribution: — The species of the engelhardti complex occur primar- 
ily in the Appalachian Valley and Ridge Province from the New River 
valley in Giles County, Virginia, southwest through east Tennessee, 
northwest Georgia, and north Alabama, and also in the Tennessee River 
valley in west-central Tennessee as far north as Decatur and Perry coun- 
ties. They also inhabit karst “islands” near the eastern edge of the Alle- 
gheny Plateau — Grassy Cove, Tennessee; Pine Mountain, Tennessee 
and Kentucky; Lookout Valley, Tennessee and Georgia; Wills Valley, 
Alabama; and Sequatchie Valley, Tennessee and Alabama. Southwest of 
Tazewell County, Virginia, all known species of cave trechines in the 
Appalachian valley or in karst islands at the edge of the Allegheny Pla- 
teau belong to the engelhardti complex. Limited overlap (but not sympa- 
try in the same caves) occurs with species of the petrunkevitchi group (all 
species with circular, well-formed eye rudiment), which are known from 
caves in Washington, Smyth, Wythe, Pulaski, Tazewell (Burkes Garden), 
and Bland counties, Virginia. Ranges of engelhardti complex species {P. 
quadratus, P. egberti) and pusio and gracilis group species in Giles 
County are mutually exclusive (see Barr 1965). 
Pleistocene Refugia 
Cave species of Pseudanophthalmus are presumably descendants of 
edaphobitic ancestors which became extinct outside of caves during 
warmer, drier interglacials (Jeannel 1949; Barr 1967a). Jeannel (1949) 
proposed a Unaka refugium hypothesis: ancestral beetle stocks spread 
over the Appalachian valley and Interior Low Plateaus during glacial 
maxima from an interglacial refugium in the higher mountains along the 
Tennessee-North Carolina border. A pulse theory of cave colonization 
