Appalachian Pseudanophthalmus 41 
and elsewhere) demonstrated a close relationship between the Eurasian 
genus Trechoblemus (eyes and wings usually present) and Pseudanoph- 
thalmus and Neaphaenops (I would add Nelsonites), postulating a Euro- 
pean introduction ancestral to these cave beetles. In my judgment Lasio- 
trechus discus, from eastern Canada and northeastern United States, 
belongs within the same phyletic series. Trechoblemus was recently 
reported from Oregon (Barr 1972). 
The notion of a Unaka refugium should not be totally discarded: the 
distribution of the engelhardti group (one of seven groups in the engel- 
hardti complex) is compatible with the hypothesis of successive pulses 
from a Unaka refugium, and the distribution of the petrunkevitchi group 
(Virginia; not treated in detail in this paper) suggests an ancestral refu- 
gium in the Blue Ridge. But new and more complete distributional data 
for cave trechines in eastern United States, including Pseudanophthal- 
mus, strongly support the hypothesis that the Allegheny Plateau played a 
major role as a refugium during Pleistocene time and perhaps even ear- 
lier. At least two other introductions must be postulated to account for a) 
Darlingtonea and Ameroduvalius, of unknown affinities within the Tre- 
chini; and b) Xenotrechus, whose two Missouri species are unquestiona- 
bly very close to Chaetoduv alius, a genus of the Carpathians and Tran- 
sylvanian Alps. 
KEY TO SPECIES GROUPS OF THE ENGELHARDTI COMPLEX 
1. Apex of aedeagus without constriction before apex (dorsal view) and 
without hatchet shape or sharp ventral cusp (lateral view) 2 
Apex of aedeagus arrow-shaped in dorsal view because of subapical 
constriction and/or hatchet-shaped with sharp ventral cusp in lateral 
view (Figs. 5-11) engelhardti group 
2(1). Pronotum disc with 1-4 long setae each side in addition to normal 
pubescence 3 
Pronotum disc pubescent but without longer setae 5 
3(2). Small, slender, usually depressed species (3. 1-4.5 mm) 4 
Larger (4.4-6. 2 mm), slender, often convex species; aedeagal apex pro- 
duced and slender or (Rye Cove, VA) button-shaped (Figs. 48-54) 
jonesi group (new group) 
4(3). Aedeagus with apex slender, moderately produced and attenuate, with 
very finely truncate knob in lateral view (Figs. 12-15) 
tennesseensis group (new group) 
Aedeagus with apex feebly produced and bluntly rounded at tip, median 
lobe weakly arcuate, basal bulb rather small, left copulatory piece very 
small (Figs. 20-25) hirsutus group 
5(2). Aedeagus with apex slender and considerably produced and attenuate . .6 
Aedeagus with apex briefly produced, either bluntly rounded or finely 
truncate, in some species also deflexed (Figs. 32-33) 
hubrichti group (new group) 
