JL

^FIELDIANA
Zoology

Published by Field Museum of Natural History

Volume 62, No. 4 November 26, 1973

Cave Beetles of the Genus Pseudanophthalmus

(Coleoptera, Carabidae) from the Kentucky

Bluegrass and Vicinity

Carl H. Krekeler

Valparaiso University, Valparaiso, Ind.

The first true cave carabids known from America, blind, depig-
mented forms known as anophthalmids, were found in Mammoth
Cave in Kentucky (Erichson, 1844). Between their discovery and
1957 about ten additional forms of anophthalmids were described
from Kentucky by Motschoulsky (1862), Horn (1868, 1883), Garman
(1892), and Jeannel (1931, 1949).

In the same years, however, almost 50 species and subspecies of
anophthalmids were described from Tennessee, Virginia, and West
Virginia where collecting had been more intensive. All but a few
of these forms were described between 1928 and 1952 by Jeannel and
Valentine. Suspecting (correctly, as it turned out) that a rich
Kentucky anophthalmid fauna remained to be found, we spent about
five weeks of the summer of 1957 searching for anophthalmids in
central Kentucky. Specimens collected at that time constitute the
majority of the specimens on which this paper is based. At about
the same time, Dr. Thomas C. Barr, Jr. became interested in the
anophthalmid fauna of Kentucky and has subsequently described
(1959, 1962a) several Kentucky forms. In order to avoid duplication
of effort, Dr. Barr and I have agreed that each of us will focus his
attention on certain geographic areas and/or species groups. Hence
his papers in which Kentucky anophthalmids are considered have
dealt with anophthalmids of the Mammoth Cave area and the

Library of Congress Catalog Card Number: 73-79267 ™c L'&KAr<Y OF THE

US ISSN 0015-0754 J/\N 7 ]$7A

Publication 1173 35 UNIVlksi I y OF ILLINOIS

AT URBAN/ CI """T^PN

M"** ftS>»

36 FIELDIANA: ZOOLOGY, VOLUME 62

Pennyroyal Plateau and the menetriesi, pubescens, audax, and robustus
groups. Considered in this paper are other species groups and species
from the Kentucky Bluegrass and vicinity.

I am indeed grateful to Dr. Thomas C. Barr, Jr., for his co-opera-
tion in the arrangement described above, for his generous exchange
of specimens and ideas with me, for his patience while this work
was in progress, and for his helpful criticism of this manuscript. I am
also most grateful to John Rittman who assisted me so capably in
the 1957 collecting and to Signe Heide who prepared the distribution
map. I acknowledge with appreciation the assistance of the National
Science Foundation whose grant G-3888 supported the 1957 collect-
ing and whose grant G-5637 supported the analysis of the material.

The holotypes and allotypes of all the new species described here-
in are deposited in the collection of Field Museum of Natural His-
tory (FMNH). Except for Pseudaywphthalmus puteanus and P.
conditus, paratypes are in both my collection and that of Thomas
C. Barr, Jr. The single paratype of puteanus is in the collection of
Dr. Barr and the single paratype of conditus is in my collection.

The measurements (in mm.) used are those proposed and de-
scribed by Valentine (1932, 1945) and used by me (1958) previously.
The abbreviations used are: TL — total length; HL — head length;
HW — head width; PL — pronotum length; PW — pronotum width;
EL— elytron length; EW— elytra width; DE— depth; ANT— an-
tenna length; AED — aedeagus length. I have included the extended
mandibles in the head length, and in total length, as does Valentine;
Barr's recent measurements do not include the mandibles. Most of
the indices which Valentine proposed as being useful can be deter-
mined from the measurements given if they are desired. A useful
index that cannot be determined from the measurements, the
chaetotaxial index (abbreviated "c"), is given with the measure-
ments. It should be noted that though the drawings of the aedeagi
are dorsal views of the organ as it lies in situ, it is convenient and
customary to refer to this as a lateral view and hence to the convex
portion as dorsal, the concave as ventral, that surface toward the
viewer as left, that away from him as right.

All but one species of the anophthalmids of the Kentucky Blue-
grass and vicinity may be assigned to four species groups. In all the
species two discal setae are found, with the first of them typically
at the level of the 4th humeral papilla, and the recurrent stria joins
either the 3rd or the 5th elytral stria. A key by means of which the

KREKELER: CAVE BEETLES 37

four groups and the species not assigned to a group may be dis-
tinguished is provided below. First it is appropriate that comments
be made concerning some of the key characters.

The mesosternum in anophthalmids typically slopes gently, or
follows a broad convex curve, to an anterior narrowing of the meso-
thorax. Barr (1966) has noted, however, that in some central
Kentucky forms this portion of the mesosternum is distinctly
angulate and that in many cases there is a tubercle at this angle.

In most anophthalmids the posterior margin of the last (6th)
abdominal sternite of the male is gently arched without interruption.
Barr (1967a) has pointed out that in males of the gracilis and
inexpectatus groups there is a shallow but distinct indentation in
this margin. A similar, though usually not as distinct an indentation,
is often found in males of the horni group.

Another character of considerable utility in distinguishing be-
tween groups is the nature of the microsculpture, especially that of
the elytra. The surface of anophthalmids is generally alutaceous
(leathery) in appearance. Some forms, however, when viewed under
relatively low magnification have a matte surface. Barr (pers.
comm.) has termed this "prumose-" The difference in microsculp-
ture is most distinctive under the higher magnifications of a dissect-
ing microscope. The sculpturing of the alutaceous surface is trans-
verse and irregular. The pruinose surface, in contrast, is composed
of minute uniform-sized polygons, each with a minute hair (micro-
trichia?) .

Key to the Species Groups, and Species Not Assigned to a Group, of

Pseudanophthalmus From the Kentucky Bluegrass and Vicinity

1 Inner margin of femora of anterior legs distinctly angulate about one-
third the distance from their base barri group

1' Inner margin of profemora broadly rounded 2

2 (1') Humeri of elytra with distinct or feeble serrations (because the reflex ed

margin sometimes extends vertically these serrations may be visible only
when the specimen is examined from a lateral angle) 3

2' Humeri of elytra lacking serrations 4

3 (2) Aedeagus with single copulatory piece; last (6th) abdominal sternite of

male sometimes feebly indented horni group

3' Aedeagus with two long, slender copulatory pieces; last (6th) abdominal

sternite of male shallowly , but distinctly, indented . inexpectatus group

4 (2') Forms from the southeast portion of the Kentucky Bluegrass and the

adjacent Cumberland Plateau with two copulatory pieces which basally
are two concentric cylinders rittmani group

38 FIELDIANA: ZOOLOGY, VOLUME 62

4' Form from the southwestern portion of the Kentucky Bluegrass with a
single copulatory piece P. conditus n. sp.

HORNI GROUP

Length 3.3-5.3 mm. Microsculpture alutaceous or pruinose. Labrum with
low, broad, weakly to well-defined median tooth. Posterior angles of pronotum
sharply rectangular to somewhat acute with straight or slightly concave lateral
margins. Mesosternum distinctly angled except in ohioensis, exoticus, krameri,
and pholeter. Humeri of elytra distinctly serrate; striae fairly distinct on disc,
becoming indistinct laterally (except in exoticus) ; 1st discal seta just posterior to
level of 4th marginal humeral papilla (except in exoticus); recurved portion of
apical groove weakly impressed and so variable (either curving medially to the
3rd stria or curving laterally or extending subparallel to the elytral suture and
interpretable as joining either the 3rd or 5th stria) that it is of no diagnostic value.

Aedeagus moderately to extremely elongate; ranging in shape from those flat-
tened ventrally and weakly arched dorsally with a distinct basal flexure to those
arched both ventrally and dorsally and lacking a distinct basal flexure; apex
moderately to very extensively produced; transfer apparatus short to moderately
long, consisting of a single element which in lateral view tapers from a thick base
to a pointed tip. Parameres moderately long, bearing 4 or 5 setae.

The only species of anophthalmid heretofore described from the
Bluegrass is Pseud anophthalmias homi after which this group is
named. Jeannel grouped this species at one time (1931) with pusio
and at another (1949) with robustus. Valentine (1932) considered
this species as forming a distinct group related to the menetriesi
group. Barr (1959), like Valentine, distinguished between the homi
and robustus groups and having compared the forms recently retains
this distinction (Barr, pers. comm.). I concur with Valentine and
Barr in considering this a distinct group.

The homi group species are similar in external appearances to
those of the inexpectatus group. But species of the homi group can
be distinguished from those of the inexpectatus group on the basis
of the following external characteristics. 1) The posterior margin of
the last abdominal sternite of males of the inexpectatus group is
typically rather clearly indented; there is often an indentation here
in males of the homi group also, but this indentation is rather shal-
low and poorly defined. 2) In caves where they coexist (Reid's=
Picadome, Swope, Arnold, Dix Dam, Clifton, Robinson, and Hay-

Figs. 1-4. Aedeagi of Pseudanophthalmns species of homi group. 1. P. homi
abditus, n. ssp. 2. P. solivagus, n. sp. 3. P. tenebrosus, n. sp. 4. P. desertus deser-
tus, n. sp. and ssp.

0.5 mm

39

40 FIELDIANA: ZOOLOGY, VOLUME 62

den) the horni group form is larger than the inexpectatus group form,
though the range of size is not sharply discontinuous in the first two
caves listed. 3) In caves where they coexist the horni group forms
have a distinctly angulate mesosternum in contrast to the gradually
sloping mesosternum of the inexpectatus group. It should be noted,
however, that certain forms of the horni group, namely ohioensis,
exoticus, krameri, and pholeter, do not have a distinctly angulate
mesosternum.

It is, however, on the basis of the aedeagal characteristics, and
specifically those of the copulatory pieces, that the horni and inex-
pectatus groups are most sharply distinguished. The copulatory
pieces of species of the inexpectatus group are in the form of two long
subparallel blades which regularly protrude through the aperture
of the median lobe. In species of the horni group there is a single
copulatory piece which in lateral view tapers from a thick base to a
pointed tip. When viewed dorsally or ventrally (cf. fig. 5a), this
single element is seen to be a blunt-tipped elongate triangular ex-
tension of a flattened cylinder; denticles are often found at its tip.
The membranous internal sac sheaths the copulatory piece for much
of its length. Ventrally the membranous sac extends toward the
produced tip of the median lobe and forms a distinct chitinized
' 'floor" within the U-shaped tube which extends beyond the copula-
tory piece. In certain preparations and or when viewed at a particu-
lar angle this chitinized membrane may appear to be a second
copulatory piece. In no case, however, is it detached from the
median lobe so as to extend through the aperture in association with
the copulatory piece which lies dorsal to it.

Forms of the horni group are found in greatest concentration in
the Kentucky River basin of the Bluegrass. They are scattered also
in the Licking River drainage of the Bluegrass and north of the
Ohio River in southeastern Indiana and southwestern Ohio. One
species tentatively assigned to this group is found in the Cumberland
Plateau immediately to the east of the Bluegrass.

Key to the Species and Subspecies of the horni Group

1 Body, especially elytra, with alutaceous microsculpture 2

1 ' Body, especially elytra, with pruinose microsculpture 12

2 (1) Apex of aedeagus directed downward 3

2' Apex of aedeagus directed upward 4

3 (2) Apex of aedeagus broadly rounded (fig. 5) chthonius n. sp.

3' Apex of aedeagus blunt (fig. 7) exoticus n. sp.

7(5)

7'

8 (7')

8'

9 (8')

9'

10 (7)

10'

KREKELER: CAVE BEETLES 41

4 (2') Apex of aedeagus curved gently upward (fig. 6) ohioensis n. sp.

4 ' Apex of aedeagus recurved or with tip extending upward at an angle . 5

5 (4') Apex of aedeagus with tip extending upward at an angle (figs. 1, 2,

3, 9) 7

5' Apex of aedeagus recurved (fig. 4) desertus n. sp. 6

6 (5') Smaller body (3.87-4.84 mm.) and aedeagus (0.91-1.17 mm.)

desertus n. sp.

6' Larger body (4.93-5.30 mm.) and aedeagus (1.29 mm.)

desertus major n. ssp.

Apex of aedeagus much produced beyond the aperture (figs. 2, 3, 9) . 10

Apex of aedeagus only slightly produced beyond the aperture (fig. 1).

horni (Garman) 8

Aedeagus less than 0.8 mm. long horni s. str.

Aedeagus more than 0.8 mm. long 9

Aedeagus 0.85-0.96 mm. long horni abditus n. ssp.

Aedeagus more than 1.0 mm. long horni caecus n. ssp.

Apex of aedeagus distinctly swollen beyond the aperture (figs. 2, 9) . 11

Apex of aedeagus not distinctly swollen beyond the aperture (fig. 3).

tenebrosus n. sp.

11 (10) Swollen portion of produced apex of aedeagus very long (fig. 9);

aedeagus more than 1.35 mm. long; copulatory piece more than 0.3
length of aedeagus elongatus n. sp.

11 ' Swollen portion of produced apex of aedeagus moderately long (fig. 2) ;
aedeagus less than 1.2 mm. long; copulatory piece less than 0.3 length
of aedeagus solivagus n. sp.

12 (1') Smaller body (3.40-4.07 mm.) and aedeagus (1.11-1.17 mm.)

krameri n. sp.

12' Larger body (3.94-4.94 mm.) and aedeagus (1.55 mm.). pholeter n. sp.

Pseudanophthalmus horni horni (Garman), new status

Anophthalmus horni Garman, 1892, p. 241. Type locality: caves and quarry
fissures within the corporate limits of Lexington, Fayette Co., Ky.
Barber, 1928, p. 196.

Pseudanophthalmus horni: Jeannel, 1928; p. 128, figs. 1,372-1,376; Jeannel,
1931, p. 456, figs. 69-73; Valentine, 1932, p. 276, pi. 23, fig. 4.

Pseudanophthalmus horni horni: Jeannel, 1949, p. 49, fig. 27. Type locality:
as above, though designated by Jeannel as a "small cavern on the site
of the University of Kentucky."

Pseudanophthalmus horni garmani: Jeannel, 1949, p. 49, fig. 37. Type locality:
Reid's Cave (one mile west of Lexington), Fayette Co., Ky. (Mus. Nat.
Hist. Nat., Paris). (NEW SYNONYMY.)

42 FIELDIANA: ZOOLOGY, VOLUME 62

Pseudanophthalmus horni minor: Jeannel, 1949, p. 49, fig. 36. Type locality.
Phelp's Cave (five miles southwest of Lexington), Fayette Co., Ky. (coll:
Henrot; cotype Mus. Nat. Hist. Nat., Paris). (NEW SYNONOMY.)

Though specimens of P. horni were set aside by Garman as types
they were not available to Jeannel or Valentine and have, in fact,
only recently been located (Barr, pers. comm., May 9, 1968). Par-
tially as a result of this there is, unfortunately, a good deal of confu-
sion about this species in the literature. Jeannel's 1928 figures of
horni, labelled simply "from Lexington," must be of a Reid's Cave
specimen (where he collected two specimens in 1928) : they are
identical to those in his 1931 paper identified more specifically as
being from Reid's Cave. In 1949 Jeannel described three subspecies
of horni: 1) horni s. str., on the basis of a specimen in his possession
collected by Garman which he acknowledged receiving from the
U. S. National Museum; 2) horni garmani on the basis of two speci-
mens he had collected from Reid's Cave and one collected by Henrot
in Picadome Cave; 3) horni minor, on the basis of 15 specimens from
Phelp's Cave. In the meantime, Barr and his associates (pers.
comm.) have concluded that "Reid's Cave" and "Picadome Cave"
are two names applied to the same cave. Finally, when Garman's
types were located the holotype was identified as coming from
Reid's Cave. There is no question, therefore, that garmani must be
placed into synonymy. As a matter of fact, the characters cited by
Jeannel as diagnostic of subspecies minor — smaller size, less deeply
insinuated pronotal margin, and less recurved aedeagal apex — are
variations not restricted to the Phelp's Cave population. Hence
minor, too, is placed into synonymy.

We were able to collect only two specimens from Reid's (= Pica-
dome) Cave in 1957. Fortunately, both were males. When I re-
moved the genitalia I discovered that there were two quite distinct
forms: 1) that figured by Jeannel (1928, 1931, 1949) and Valentine
(1932) for horni; 2) that described below for umbratilis. Fortu-
nately, the specimens which Jeannel and Valentine used for their
dissections and figures were of the form of the recently located holo-
type which Barr (pers. comm.) confirms as being the "robust local
species with prominent mesosternal spine."

The forms of P. horni are most readily distinguished from other
species of the group by the shape of the apex of the aedeagus: its tip
extends upward at a rather distinct angle but it is produced only
slightly beyond the aperture. The nominate subspecies is the
smallest form of the species, being 3.72 ± 0.04 mm. in length. The

KREKELER: CAVE BEETLES 43

aedeagus is likewise small, the seven measured ranging from 0.77-
0.78 mm. There is a great amount of variation in the chaetotaxial
index (range 0.56-1.13) and a comparatively high mean chaetotaxial
index (0.73) . There are several long hairs anteriorly on the pronotum
on either side of the midline among the short pubescence.

This form has been collected from several caves in the vicinity of
Lexington, Kentucky: Reid's (=Picadome) Cave (type locality),
immediately north of Picadome Elementary School, 2 miles south-
west of Lexington courthouse, Fayette Co., Ky., July 1, 1957
(C. H. K. and J. Rittman); Phelp's Cave, 5 miles west-southwest of
Lexington courthouse, Fayette Co., Ky., Oct. 8, 1961 (T. C. Barr);
Russell Cave, 6 miles north-northeast of Lexington courthouse,
Fayette Co., Ky., Nov. 16, 1963 (J. Holsinger), Aug. 5, 1964 (S.
Peck and W. Andrews) ; Church Cave, 2.5 miles west-northwest of
Georgetown, Scott Co., Ky., Sept., 1964 (R. McAdams and W.
Andrews) . Included in the measurements are five specimens in the
U. S. National Museum labelled as being collected in Lexington
by Hubbard and Schwarz on Oct. 9, 1892. These caves are located
in the Inner Bluegrass in the drainage basins of North Elkhorn
Creek and South Elkhorn Creek (including its Town Branch).
Also found in Reid's Cave, as noted above, is P. umbratilis, a species
of the inexpectatus group.

Pseudanophthalmus horni caecus, new subspecies

Type series. — Holotype, allotype, and 22 paratypes, from Clifton
Cave, 0.6 mile east-southeast of Clifton, Woodford Co., Ky., June
24, 1963 (T. C. Barr).

Holotype male (FMNH). TL 4.37, HL 1.17, HW 0.80, PL 0.80,
PW 1.00, EL 2.40, EW 1.57, DE 0.97, ANT 2.73, AED 1.01, c 0.58.

Allotype female (FMNH). TL 3.74, HL 1.00, HW 0.67, PL
0.70, PW 0.83, EL 2.03, EW 1.23, DE 0.80, ANT 2.23, c 0.67.

P. h. caecus, though not significantly larger in size (3.80 ± 0.08
mm.) than horni s. str., has a distinctly longer aedeagus (the three
measured range from 1.01-1.05 mm.) ; as a result the genital index is
slightly higher (on the order of 0.24 as compared to 0.21). In this
form also there is great variation in the chaetotaxial index (range
0.53-1.04) and a relatively high mean chaetotaxial index (0.78).

Clifton Cave is very near the Kentucky River several miles up-
stream from Frankfort. Occupying the same cave is P. umbratilis
of the inexpectatus group, a form noticeably smaller in size.

44 FIELDIANA: ZOOLOGY, VOLUME 62

Pseudanophthalmus horni abditus, new subspecies

Type series. — Holotype, allotype, and 29 paratypes, from Swope
Cave, 4.5 miles north of Versailles, Woodford, Co., Ky., holotype,
allotype, and 23 paratypes, July 3, 1957 (C. H. K. and J. Rittman),
6 paratypes Sept. 17, 1961 (T. C. Barr); 4 paratypes from Weber
#2 Cave, 3 miles southeast of Versailles, Woodford Co., Ky., Aug.
10, 1963 (T. C. Barr) ; 4 paratypes from Meece Cave, 4.5 miles north-
northwest of Nicholasville, Jessamine Co., Ky., April 7, 1964 (R.
McAdams and W. Andrews).

Holotype male (FMNH). Fig. 1. TL 4.10, HL 1.05, HW 0.70,
PL 0.80, PW 0.95, EL 2.25, EW 1.40, DE 0.80, ANT 2.40, AED
0.88, c 0.63.

Allotype female (FMNH). TL 3.80, HL 0.95, HW 0.65, PL
0.75, PW 0.85, EL 2.10, EW 1.25, DE 0.80, ANT 2.20, c 0.74.

P. h. abditus is the largest of the three subspecies of horni here
recognized (3.93 ± 0.04 mm.); but its aedeagus is smaller (the 11
measured range from 0.85-0.96 mm., average 0.91 mm.) than that
of h. caecus. The variation in the chaetotaxial index (range 0.54-
0.77) is much less, and the mean chaetotaxial index (0.66) is lower
than in the other forms.

The caves from which this form has been taken are in the vicinity
of Versailles. Swope Cave, north of Versailles, is near Lee Branch
which drains into South Elkhorn Creek. Weber #2 Cave and Meece
Cave, southeast of Versailles, are in the karst area lying between
Clear Creek and South Elkhorn Creek. Living with h. abditus in
Swope Cave is P. umbratilis, a species of the inexpectatus group;
there is no appreciable size difference between these two forms.

Pseudanophthalmus solivagus, new species

Type series. — Holotype, allotype, and 62 paratypes from Weber
Cave, 2 miles northwest of Nonesuch, Woodford Co., Ky., July 2,
1957 (C. H. K. and J. Rittman) ; 39 paratypes from Nonesuch Cave,
2.5 miles east-northeast of Nonesuch, Woodford Co., Ky., Oct. 8,
1961 (T. C. Barr); 5 paratypes from Britton Cave, 4.5 miles south-
west of Troy, Woodford Co., Ky., Nov. 4, 1962 (T. C. Barr); 1
paratype from Keene Cave, 1 mile east of Keene, Woodford Co.,
Ky., July 16, 1965 (W. Andrews and N. Hornback).

Holotype male (FMNH). Fig. 2. TL 4.35, HL 1.20, HW 0.75,
PL 0.80, PW 0.95, EL 2.35, EW 1.55, DE 0.90, ANT 2.80, AED
1.08, c 0.68.

0.5mm

Figs. 5-8. Aedeagi of Pseudanophthalmus species of horni group. 5. P. chthon-
ius, n. sp.; 5a, transfer apparatus, ventral aspect. 6. P. ohioensis, n. sp. 7. P.
exoticus, n. sp. 8. P. krameri, n. sp.

45

46 FIELDIANA: ZOOLOGY, VOLUME 62

Allotype female (FMNH). TL 4.30, HL 1.20, HW 0.75, PL
0.80, PW 0.95, EL 2.30, EW 1.50, DE 0.85, ANT 2.60, c 0.74.

P. solivagus, horni, elongatus, and tenebrosus seem to form a
natural subgroup of the horni group. In general form, color, and
pubescence these species are so similar as to be indistinguishable.
The pronotum and elytra are covered with a short and sparse
pubescence, and in most specimens (but not all) there are several
long hairs anteriorly on the pronotum on either side of the midline.
Moreover, the aedeagi of these species are similar in that the pro-
duced apex has a tip which extends upward at a rather distinct angle
and the parameres bear four setae.

P. solivagus, with a length of 4.14 + 0.02 mm., is distinctly
smaller in size than elongatus and tenebrosus; it can also be dis-
tinguished from these species by aedeagal shape as described below.
On the other hand, solivagus is larger than any of the horni subspecies.
Also the aedeagus of solivagus is longer than that of any of the horni
subspecies: the nine aedeagi measured range from 1.03-1.19 mm.,
and average 1.12 mm., in length. P. solivagus is distinguished most
readily from horni by the shape of the aedeagus: the produced tip
of solivagus is distinctly swollen beyond the copulatory piece, while
in horni there is no such swelling of the produced tip.

P. solivagus was found in the greatest abundance of all the
species reported in this paper. In Weber's Cave approximately 30
specimens were taken in the flat crawlway which forms its entrance;
the remainder were collected with the diminished effort which en-
sued when we found that a water channel opened into virgin cave.
The caves in which solivagus is found are north and east of the
deeply entrenched Kentucky River where, shortly after being joined
by the Dix River, it changes its generally westward course to a
generally northern one. Weber's Cave stream flows into Clear Creek,
a tributary of the Kentucky River; Nonesuch Cave lies along the
East Fork of Clear Creek; and Keene Cave is in the upland a short
distance north of the intermittent headwaters of the East Fork of
Clear Creek. Britton Cave is in the upland above a bend in the
Kentucky River, more than 200 ft. higher in altitude than, and less
than a mile north and less than a mile southeast of, the river.

Pseudanophthalmus elongatus, new species

Type series. — Holotype, allotype, and 54 paratypes from Old
Fort Cave, 1.2 miles northeast of Harrodsburg, Mercer Co., Ky.,

KREKELER: CAVE BEETLES 47

July 2, 1957 (C. H. K. and J. Rittman); 2 paratypes from Dix Dam
Cave, 2 miles northwest of Buena Vista, Garrard Co., Ky., July 2,
1957 (C. H. K. and J. Rittman); 3 paratypes from Robinson Cave,
4.5 miles west-northwest of Lancaster, Garrard Co., Ky., Aug. 16,
1957 (C. H. K. and J. Rittman); 8 paratypes from Arnold Cave, 1.2
miles west-northwest of Bryantsville, Garrard Co., Ky., July 2,
1957 (C. H. K. and J. Rittman).

Holotype male (FMNH). Fig. 9. TL 4.65, HL 1.30, HW 0.75,
PL 0.80, PW 1.00, EL 2.55, EW 1.50, DE 1.10, ANT 2.75, AED
1.38, c. 0.64.

Allotype female. TL 4.75, HL 1.30, HW 0.80, PL 0.90, PW 1.10,
EL 2.55, EW 1.60, DE 0.95, ANT 3.05, c 0.58.

P. elongatus, with a length of 4.71 ± 0.02 mm., and P. tenebrosus,
with a length of 4.75 + 0.05 mm., are the largest species of the horni
group. The aedeagus of elongatus is by far the longest in the group;
the seven measured range from 1.35-1.43 mm., and average 1.38
mm., in length. As in horni, solivagus, and tenebrosus, the apex of
the aedeagus of elongatus is produced and has a tip which extends
upward at a distinct angle. As in solivagus, but unlike in horni and
tenebrosus, the apex is clearly swollen beyond the position of the
copulatory piece. The apex of elongatus is much more produced,
however, than that of solivagus; moreover, the copulatory piece is
approximately one-third the length of the aedeagus in elongatus (the
seven measured range from 0.31-0.37, average 0.35) as compared
to less than three-tenths the length of the aedeagus in solivagus (the
eight measured range from 0.25-0.30, average 0.27).

Old Fort Cave lies in a karst plain about 6 miles southwest of the
Kentucky River at the southwest margin of the Inner Bluegrass.
Whether the waters of this cave drain north and east via Shaker
Creek into the Kentucky River or west via Town Creek into Salt
River is not known. Dix Dam Cave, Arnold Cave, and Robinson
Cave lie immediately east of Dix River, a major tributary of the
Kentucky River. In these last three caves is found also P. umbratilis,
a species of the inexpectatus group, which is clearly smaller in size
than elongatus. The three Robinson Cave specimens listed as para-
types of elongatus are all females hence cannot be assigned to this
species with certainty. They are, however, clearly of the horni
group, are in geographic proximity to the other populations of
elongatus, and are of the size of eloyigatus.

48

FIELDIANA: ZOOLOGY, VOLUME 62

Figs. 9-11. Aedeagi of Pscudatwphthahniis species of horni and ritimani
groups. 9. P. elongatiis n. sp. 10. P. pholeter n. sp. 11. P. riUmani n. sp.

Pseudanophthalmus tenebrosus, new species

Type series. — Holotype, allotype, and 11 paratypes from Stevens
Creek Cave, 0.85 mile east-southeast of Orville, Henry Co.. Ky.,
July 9, 1966 (T. C. Barr and R. Norton).

Holotype male (FMNH). Fig. 3. TL 4.87, HL 1.30, HW 0.80.
PL 0.87. PW 1.00, EL 2.70, EW 1.63, DE 1.10, ANT 3.00. AED 1.12.
c. 0.52.

Allotype female (FMNH). TL 4.90, HL 1.23, HW 0.87. PL
0.90, PW 1.03. EL 2.77, EW 1.73. DE 1.23, ANT 3.10, c 0.51.

KREKELER: CAVE BEETLES 49

Although the body length of tenebrosus and elongatus is com-
parable, their aedeagi are quite different in length. The three aedeagi
of tenebrosus measured average 1.16 mm. in length (ranging from
1.12-1.19 mm.) as compared to the average 1.38 mm. length of
elongatus. As a result there is, of course, a difference in genital
index: that of tenebrosus which averages 0.24 (range 0.23-0.24) is
lower than that of elongatus which averages 0.29 (range 0.28-0.30).
The apex of the aedeagus of tenebrosus, like that of elongatus but
unlike that of horni, is extensively produced beyond the copulatory
piece. The produced portion of the aedeagus is, however, like that of
horni but unlike that of elongatus, not clearly swollen.

Stevens Creek Cave is about 20 miles north of Clifton Cave and
even farther from the other caves in which are found the other forms
of its subgroup (horni, solivagus, and elongatus) . It lies near Stevens
Creek less than a mile from where it enters the Kentucky River.

Pseudanophthalmus desertus desertus, new species and sub-
species
Type series. — Holotype, allotype, and 8 paratypes from Jones
Cave, 0.8 mile northeast of Locust Grove, Clark Co., Ky., June
17, 1963 (T. C. Barr); 5 paratypes from Price Cave, 1.8 miles east-
northeast of Eminence, Henry Co., Ky., July 9, 1966 (T. C. Barr
and R. Norton); 2 paratypes from Slack's Cave, 3 miles west-
southwest of Georgetown, Scott Co., Ky., Jan. 18, 1963 (T. C.
Barr); 1 paratype from Hayden Cave, 0.5 mile south-southeast of
Gratz, Owen Co., Ky., July 9, 1966 (R. Norton).

Holotype male (FMNH). Fig. 4. TL 4.03, HL 1.03, HW 0.73,
PL 0.70, PW 0.90, EL 2.30, EW 1.40, DE 0.83, ANT 2.47, AED
0.93, c 0.64.

Allotype female (FMNH). TL 4.53, HL 1.30, HW 0.73, PL
0.83, PW 0.90, EL 2.40, EW 1.53, DE 1.07, ANT 3.00, c 0.70.

The size of P. desertus desertus (4.39 + 0.08 mm.) is intermediate
in the size range of the horni group. There is a considerable range of
variation both in body length (from 3.87-4.84 mm.) and in aedeagus
length (from 0.91-1.17 mm., average 1.01 mm., in the nine specimens
measured), and more than one form may be represented in this
sample. I am, however, unable to distinguish any such forms on the
basis of the small number of specimens available. This species is
distinguished from horni, solivagus, elongatus, and tenebrosus by the
fact that its pronotum and elytra are covered with abundant and

30 FIELDIANA: ZOOLOGY, VOLUME 62

medium length hairs rather than short and sparse pubescence; also
the longer hairs often found anteriorly on the pronotum of these
species are not found in desertus. The apical groove of d. desertus is
rather distinctly impressed and clearly joins the 3rd stria. The
chaetotaxial index of this form ranges from 0.52-0.77, averages
0.64. Most distinctive of this species is the apex of the aedeagus:
it forms a distinct hook with the tip pointing back toward the base
of the aedeagus. The transfer apparatus is clearly of the horni type.

P. d. desertus is found, apparently in small populations, in several
widely scattered caves. Price. Hayden, and Slack's caves are north
and west of Lexington, the former two about 40 miles distant, the
latter about 10 miles. Jones Cave, on the other hand, is about 10
miles southeast of Lexington. All four caves are in the Kentucky
River drainage basin. Found also in Hayden Cave is P. umbratilis
of the inexpectatus group, a form which is clearly smaller in size than
d. desertus.

Pseudanophthalmus desertus major, new subspecies

Type series. — Holotype. allotype, and one male paratype, from
Beaver Cave. 3 miles northeast of Oddville. Harrison Co., Ky.,July
17. 1966 (T. C. Barn.

Holotype male (FMNH). TL 4.97, HL 1.30, HW 0.83, PL 0.90,
PW 1.13. EL 2.77. EW 1.70. DE 1.10. AXT 3.24. AED 1.29. c 0.56.

Allotype female (FMNH). TL 5.30. HL 1.40, HW 0.90, PL
0.97. PW 1.17. EL 2.93. EW 1.80. DE 1.17. AXT 3.30, c 0.48.

P. d. major is noticeably larger (range 4.93-5.30 mm., mean
5.04 ± 0.10 mm. ) than desertus s. str. In addition the single aedeagus
measured, at 1.29 mm., is considerably larger than any of desertus
s. str. P. d. major also has somewhat more distinct lateral striae on
the elytra and has a smaller chaetotaxial index (range 0.48-0.56,
mean 0.52). The apical groove joins either the 3rd or the 5th stria.

Beaver Cave, about 35 miles northeast of Lexington, is 30 miles
distant from the nearest cave (Slack's Cave) from which the nomi-
nate subspecies has been taken. It lies near Beaver Creek, a tribu-
tary of Licking River.

Pseudanophthalmus chthonius, new species

Type series. — Holotype, allotype, and 28 paratypes from Wilson
Cave 1.5 miles northwest of Kent, Jefferson Co., Ind., holotype,
allotype, and 22 paratypes. June 9, 1957 (C. H. K. and J. Rittman),

KREKELER: CAVE BEETLES 51

6 paratypes, Aug. 9, 1964 (T. C. Barr); 22 paratypes from Morris
Cave, 4.5 miles south-southwest of Kent, Jefferson Co., Ind., Aug. 3,
1959 (C. H. K. and W. Bloom); 2 paratypes from Lowry Cave, 0.5
mile east of Commiskey, Jennings Co., Ind., June 9, 1957 (C. H. K.
and J. Rittman) ; 1 paratype from Indian Cave, 0.5 mile southwest
of Charlestown, Clark Co., Ind., Aug. 2, 1959 (C. H. K. and W.
Bloom); 1 paratype from Peyton Beechwood Cave, 3 miles south-
west of Charlestown, Clark Co., Ind., Aug. 2, 1959 (C. H. K. and
W. Bloom).

Holotype male (FMNH). Figs. 5, 5a. TL 4.65, HL 1.20, HW
0.80, PL 0.85, PW 1.00, EL 2.60, EW 1.55, DE 0.90, ANT 2.55,
AED 0.85, c 0.63.

Allotype female (FMNH). TL 4.45, HL 1.10, HW 0.80, PL
0.85, PW 1.05, EL 2.50, EW 1.55, DE 0.95, ANT 2.70, c 0.56.

P. chthonius, with a length of 4.31 ± 0.02 mm., is intermediate in
the size range of the horni group. The aedeagus, however, is short
(the six measured range from 0.77-0.86 mm., average 0.82 mm.),
hence the genital index is low (range 0.18-0.20, mean 0.18). The
mesosternum is distinctly angled. The pronotum and elytra are
covered with a short and sparse pubescence. Most, but not all,
specimens have several longer hairs anteriorly on the pronotum on
either side of the midline. The pronotal angles are somewhat acute
by virtue of the fact that the posterior margin is rather deeply con-
cave as it joins the rather prominent basal angle.

The tip of the aedeagus is most distinctive: it is club shaped and
broadly rounded. The parameres bear 4 or 5 setae. The general
form of the copulatory piece is like that of the other species of the
horni group. The copulatory piece is approximately four-tenths
the length of the aedeagus (from 0.39-0.44, mean 0.42, in the
six measured), distinctly longer proportionately than in any other
species of the group. The portion of the membranous sac which
forms the floor of the produced apical tube of the aedeagus is well
chitinized and may appear to be a second copulatory piece. It re-
mains attached to the tube, however, rather than extending freely
through the aperture as the copulatory piece does, hence is not
interpreted as a copulatory piece. Small spurs are also visible at the
base of the copulatory piece of this species. On careful examination
these spurs are seen to be rather heavily chitinized portions of the
base of the flattened cylinder from which the copulatory piece in this
group typically arises.

52 FIELDIANA: ZOOLOGY, VOLUME 62

The caves from which this species is known are in southeastern
Indiana, across the Ohio River from where the forms described
above are found. Indian Cave and Peyton Beechwood Cave lie near
Pleasant Run, a tributary of Silver Creek which after a short distance
flows into the Ohio River. In these caves P. chthonius is the rare
associate of P. barri described below. The other caves are in the
drainage basin of the Muscatatuck River which drains by a long
route (via the White and Wabash rivers) into the Ohio River. All
five caves in which P. chthonius is found are in rocks covered with
glacial drift of Kansan and Illinoian ages. All previously described
anophthalmids have come from non-glaciated areas, and the sig-
nificance of this and other exceptions reported below in this paper
will be discussed later. The rocks in which these caves are found
correspond to those of the Bluegrass section of the Interior Low
Plateaus province of Kentucky. However, southeastern Indiana,
where glacial rather than bedrock controlled topography predomi-
nates, is classified by physiographers (Thornbury, 1965) as the Till
Plains section of the Central Lowland province. For convenience
the boundary between these sections northeast of Louisville is
placed at the Ohio River.

Pseudanophthalmus ohioensis, new species

Type series. — Holotype, allotype, and 5 paratypes from Freeland
Cave, 6 miles southeast of Peebles, Adams Co., Ohio, holotype, allo-
type, and 2 paratypes, June 7, 1957 (C. H. K. and J. Rittman), 3
paratypes, March 13, 1960 (F. Kramer and W. Menrath).

Holotype male (FMNH). Fig. 6. TL 4.10, HL 1.00, HW 0.70,
PL 0.75, PW 0.90, EL 2.35, EW 1.45, DE 1.00, ANT 2.35, AED 0.78,
c 0.55.

Allotype female (FMNH). TL 4.30, HL 1.20, HW 0.80, PL 0.80,
PW 0.95, EL 2.30, EW 1.45, DE 0.90, ANT 2.45, c 0.62.

P. ohioensis, like chthonius, is of intermediate length (4.24 ±
0.06 mm.) among the species of the horni group and has an aedeagus
which is short (the two measured are 0.78 and 0.85 mm.) so that its
genital index is low (0.19 and 0.20). The pronotum and elytra are
covered with a short and sparse pubescence. Most, but not all,
specimens have several long hairs anteriorly on the pronotum on
either side of the midline. As in chthonius the pronotal angles are
somewhat acute by virtue of the posterior margin being rather deeply
concave as it joins the rather prominent basal angle. The lateral

KREKELER: CAVE BEETLES 53

striae are somewhat more distinct in ohioensis than in chthonius, and
the apical groove of ohioensis regularly joins the 3rd stria rather than
either the 3rd or 5th as in chthonius and many other species in this
group. In contrast to all species of the horni group considered to this
point, the mesosternum of ohioensis is not distinctly angulate nor
does it have a tubercle. The general habitus otherwise is that of the
horni group. The characteristics of the aedeagus, and particularly
of the transfer apparatus, are clearly those of the horni group. Thus
there is no question of its placement here. The apex of the aedeagus
is slightly produced; the tip curves gently upward and comes to a
slightly rounded point.

P. ohioensis is the first anophthalmid to be taken from Ohio.
Freeland Cave is in a valley of a deeply dissected, relatively narrow
plateau which lies between the extension of the Bluegrass into Ohio
and the Illinoian glacial boundary on the west and the Cumberland
Plateau to the east. This plateau, the Highland Rim, corresponds
to the plateau west and southwest of the Bluegrass in Kentucky
which I refer to below, following Barr (1967b), as the Pennyroyal
Plateau, though many geomorphologists restrict the term Penny-
royal Plateau to that portion of the plateau south of the Mammoth
Cave Plateau and refer to the remainder as the Highland Rim. Free-
land Cave lies on Turkey Creek whose waters drain eastward via the
South Fork of Scioto Brush Creek and then southward via the Scioto
River to the Ohio River.

Pseudanophthalmus exoticus, new species

Holotype male from Townsend Cave, 4 miles west-northwest of
Zachariah, Estill Co., Ky., Aug. 15, 1957 (C. H. K. and J. Rittman).

Holotype male (FMNH). Fig. 7. TL 4.34, HL 1.13, HW 0.73,
PL 0.80, PW 1.00, EL 2.40, EW 1.53, DE 1.00, ANT 2.63, AED
0.70, c 0.59.

Four anophthalmid specimens were taken from Townsend Cave.
Two of them are assigned to P. exiguus and another to P. rittmani,
both described below. The fourth constitutes another species here-
with named exoticus. P. exoticus is tentatively included in the horni
group because of the form of the copulatory piece, though it differs
rather markedly from other species of this group in several respects.
The species is of moderate size (4.34 mm.) but its aedeagus is short
(0.70 mm.), hence the genital index is very low (0.16). The general
body form and humeral serrations are those of the horni group. The

54 FIELDIANA: ZOOLOGY, VOLUME 62

pronotum and elytra are covered with moderately long and dense
pubescence. The apical groove joins the 3rd stria. As in ohioensis
the mesosternum is not distinctly angulate. The following charac-
teristics of exoticus are in contrast to all other forms assigned to the
horni group: 1) the 1st discal seta is approximately at the level of the
3rd humeral papilla; 2) the discal striae are distinctly impressed and
punctate.

The apex of the aedeagus of exoticus is not produced as it is in
other species assigned to the horni group. Rather the tip of the
aedeagus is broad and blunt. The parameres are relatively short
and bear five short setae. The copulatory piece is very similar in its
ventral aspect (that shown in the drawing because the aedeagus was
flattened in preparation) to that of other species of the horni group
examined from this aspect.

Townsend Cave is in the uplands of the Cumberland Plateau im-
mediately to the east of the Bluegrass. It lies about 300 ft. above the
floodplain of the nearby Kentucky River to which its waters drain
by way of Billy Fork and Millers Creek.

Pseudanophthalmus krameri, new species

Type series. — Holotype, allotype, and 9 paratypes, from Cave
Hill Cave, 5 miles northwest of West Union, Adams Co., Ohio,
holotype, allotype, and 8 paratypes, Aug. 28, 1962 (C. H. K. and N.
Krekeler), 1 paratype, March 13, 1960 (F. Kramer and W. Men-
rath.

Holotype male (FMNH). Fig. 8. TL 3.47, HL 0.87, HW 0.67,
PL 0.70, PW 0.83, EL 1.90, EW 1.13, DE 0.67, ANT 1.93, AED
1.17, c 0.76.

Allotype female (FMNH). TL 3.63, HL 1.07, HW 0.70, PL
0.70, PW 0.83, EL 1.87, EW 1.13, DE 0.67, ANT 2.00, c 0.79.

P. krameri and the remaining species of this group, P. pholeter,
may be considered an aberrant subgroup of the horni group. In
contrast to all the other species of the horni group, they have prui-
nose microsculpture. Moreover, as in ohioensis and exoticus, the
mesosternum is not angulate. The disc of the pronotum is pubes-
cent. The recurved portion of the apical groove is relatively short
and well impressed, curving inward proximally to join the 3rd stria.
The fact that the transfer apparatus of these two species is, as de-
scribed below, so similar to that of the species of the horni group de-
scribed above, seems to warrant including them in this group.

KREKELER: CAVE BEETLES 55

P. krameri is relatively small in size, averaging 3.67 ± 0.06 mm.
in length. The posterior angles of the pronotum are somewhat acute
by virtue of the lateral and posterior margins being slightly concave ;
the basal margin is straight or very shallowly concave. The humeri
of the elytra slope more distinctly and the elytral margins are more
gently arcuate in krameri than pholeter. The striae on the disc of
the elytra of krameri are somewhat less distinct than in pholeter; this
is particularly noticeable apically: the 3rd stria, where it is joined
by the recurrent stria, is feebly impressed in krameri. The chaeto-
taxial index of krameri is high (range 0.67-0.88, mean 0.82).

The two aedeagi of krameri measured are 1.11 and 1.17 mm. in
length; the genital index is 0.32. The median lobe of the aedeagus
is quite thin; and the basal bulb, though only of moderate size, is
rather well demarcated from the median lobe. The apex of the ae-
deagus is produced and terminates in an extension which angles
sharply dorsally so that the tip has a boot-like appearance. The
copulatory piece is approximately one-third the length of the aedea-
gus (the two measured were 0.33 and 0.34 the aedeagal length) and
is covered apically with rather large denticles.

Cave Hill Cave is high on a hill from which water drains eventu-
ally into the Ohio River. The hill forms a divide so that drainage
is either 1) northeastward via Cherry Fork and thence east and south
by way of the West Fork of Brush Creek and Ohio Brush Creek to
the Ohio, or 2) south and west via the East Fork of Eagle Creek and
Eagle Creek to the Ohio. The cave lies just within the boundaries
of Illinoian glaciation. It is formed in the limestones which com-
prise the bedrock of the Outer Bluegrass. Because of the fact that
the area has been glaciated, however, geomorphologists include it
within the Central Lowlands province.

I am pleased to name this species after Frank Kramer of Cincin-
nati who first collected this form.

Pseudanophthalmus pholeter, new species

Type series. — Holotype, allotype, and 4 paratypes, from Adams
Cave, 5 miles south-southwest of Richmond, Madison Co., Ky., Aug.
15, 1964 (T. C. Barr and S. Peck).

Holotype male (FMNH). Fig. 10. TL 4.64, HL 1.23, HW 0.83,
PL 0.87, PW 1.03, EL 2.53, EW 1.60, DE 1.00, ANT 2.80, AED
1.55, c 0.55.

56 FIELDIANA: ZOOLOGY, VOLUME 62

Allotype female. TL 4.13, HL 1.13, HW 0.73, PL 0.77, PW 0.93,
EL 2.23, EW 1.33, DE 0.83, ANT 2.47, c 0.53.

P. pholeter is significantly larger (averaging 4.42 + 0.15 mm. in
length) than krameri; there is, however, an overlapping of size
ranges (krameri, 3.40-4.07 mm.; pholeter, 3.94-4.94 mm.). The
posterior angles of the pronotum are distinctly rectangular, rather
than acute as in krameri; the basal margin is concave, though shal-
lowly so. The humeri of the elytra are rounded and the elytral mar-
gins are gently arcuate. The striae on the disc of the elytra are quite
distinct and regular; the lateral striae are less distinct. The chaeto-
taxial index (range 0.50-0.72, mean 0.56) is much lower than that of
krameri. The ciliated reflexed margin of the elytra in the humeral
region extends about 45° above horizontal so that its distinct, but
shallow, serrations are not readily visible when the specimen is
viewed dorsally.

The length of the aedeagus of pholeter is much greater (that mea-
sured being 1.55 mm.) than that of krameri. But inasmuch as the
body length of krameri is smaller than that of pholeter the genital
index of the two forms is essentially the same (krameri, 0.32; pholeter,
0.33) . The median lobe of the aedeagus of pholeter is much thicker
than that of krameri; the basal bulb is large but pool v demarcated
from the median lobe. As in krameri, the produced r pex has a boot-
like appearance. The copulatory piece of the single aedeagus ex-
amined is 0.45 the length of the aedeagus. As in krameri, the copu-
latory piece is covered apically with denticles, but those of pholeter
are much smaller and less conspicuous than those of krameri.

Adams Cave is in rocks of the Outer Bluegrass and lies near Sil-
ver Creek, a tributary of the Kentucky River. Adams Cave is al-
most 90 miles distant from Cave Hill Cave where krameri is found.
In addition, the two caves are separated by the Ohio River. On the
basis of numerous common features of krameri and pholeter as de-
scribed above, particularly of the aedeagi, the two species are con-
sidered to be closely related.

INEXPECTATUS GROUP

Length 3.2-4.2 mm. Labrum with low, broad, weakly defined median tooth.
Posterior angles of pronotum sharply rectangular with straight or concave lateral
margins; disc of pronotum pubescent. Margin of elytra with ciliated obsolete
serrations throughout length, the serrations very feeble and widely spaced along
margins, less feeble and more closely spaced at humeri, the cilia along margins
larger than in horni group; striae evident though quite broad, shallow, and irregu-

KREKELER: CAVE BEETLES 57

lar, becoming indistinct laterally; 1st discal seta approximately at level of 4th
marginal humeral papilla; chaetotaxial index variable but generally high (averages
range from 0.72 to 0.99) ; recurved portion of apical groove weakly impressed and
so variable (either curving medially to the 3rd stria or curving laterally or extend-
ing subparallel to the elytral suture and interpretable as joining either the 3rd
or 5th stria) that it is of no diagnostic value.

Aedeagus moderately elongate; relatively flat ventrally and moderately arched
dorsally, with a moderate basal flexure; apex produced. Transfer apparatus pro-
portionately quite long, ranging from one-half to two-thirds the length of the ae-
deagus; composed of two elements each of which is semicylindrical at its base but
becomes blade-like (flattened, slender, and pointed) toward its apex. Parameres
moderately long, bearing 2 or 3 setae.

Barr (1967a), in reviewing the characteristics of P. inexpectatus
Barr of Mammoth Cave, pointed out the similarities of this species
to P. gracilis Valentine and P. hadenoecus Barr of the Appalachian
Valley. The similarities which are particularly striking are: 1) all
three species have an indentation in the last abdominal sternite of
the males (the only other species in which this is found are included
in the horni group; as discussed above, the indentation in those spe-
cies is not well developed); 2) the aedeagi of these species are un-
usually long and slender and contain two long and slender copulatory
pieces.

On the other hand, the Appalachian Valley species differ from
inexpectatus in the following respects: 1) the labrum lacks a tooth;
2) the humeri lack serrations; 3) the striae are very indistinct, even
on the disc; 4) the parameres bear four setae.

Though it may eventually be desirable to include the Kentucky
forms and the eastern forms in a single group, two distinct subgroups
would have to be recognized. For convenience, therefore, the Ken-
tucky forms are here considered as the inexpectatus group, a group
related to but separate from the gracilis group.

Key to the Species and Subspecies of the inexpectatus Group

1 One of the copulatory pieces denticulate 2

1' Neither copulatory piece denticulate umbratilis n. sp.

2 (1) Copulatory pieces subequal in length parvus n. sp.

2' Copulatory pieces distinctly unequal in length 3

3 (2') Longer copulatory piece at least two-thirds length of aedeagus. cnephosus

n. sp.

3' Longer copulatory piece 0.5 to 0.6 length of aedeagus 4

4 (3') Small (3.4-3.6 mm. long), narrow (width index 0.31-0.34) form from

the Bluegrass puteanus n. sp.

0.5mm

58

KREKELER: CAVE BEETLES 59

4' Forms from the region south and west of the Bluegrass.

inexpectatus Barr 5

5 (4') Smaller (3.3-3.8 mm. long) and proportionately wider (width index

0.34-0.35) form from the Mammoth Cave Plateau., .inexpectatus s. str.

5' Larger (3.6-4.2 mm. long) and proportionately narrower (width index

0.30-0.34) form from the Pennroyal Plateau.

inexpectatus orientalis n. ssp.

Pseudanophthalmus inexpectatus inexpectatus Barr, new
status

Pseudanophthalmus inexpectatus Barr, 1959, p. 10. figs. 3A and 7 (3). Type
locality: White Cave, Edmonson Co., Ky. Barr, 1962b, p. 281. Barr,
1967a, p. 24.

P. inexpectatus inexpectatus is small in size (the five specimens of
which I have measurements average 3.52 ± 0.08 mm. in length).
The chaetotaxial index is high (range 0.82-0.93, mean 0.86). The
two aedeagi I have measured are 0.98 and 1.01 mm. in length. The
copulatory pieces of the aedeagi of both i. inexpectatus and i. orient-
alis are 50-60 per cent the length of the aedeagus, and in both forms
the right copulatory piece is distinctly longer than the left and is
denticulate.

P. i. inexpectatus has been taken in small numbers from White
Cave and adjoining Mammoth Cave and from Great Onyx Cave of
the Flint Ridge cave system northeast of Mammoth Cave (Barr,
1962b). These are caves of the Mammoth Cave Plateau southwest
of the Bluegrass which lie in the Green River drainage.

Pseudanophthalmus inexpectatus orientalis, new subspecies
Type series. — Holotype, allotype, and 8 paratypes from Wilson

Cave, 1 mile southeast of Black Gnat, Green Co., Ky., Aug. 21, 1963

(T. C. Barr); 4 paratypes from Phillips Cave, 4 miles northwest of

Campbellsville, Taylor Co., Ky., June 23, 1957 (C. H. K. and J.

Rittman) .

Holotype male (FMNH). Fig. 12. TL 4.03, HL 1.07, HW 0.67,

PL 0.73, PW 0.87, EL 2.23, EW 1.30, DE 0.77, ANT 2.33, AED

0.93, c 0.74.

Allotype female (FMNH). TL 4.00, HL 1.13, HW 0.67, PL

0.73, PW 0.83, EL 2.13, EW 1.27, DE 0.73, ANT 2.50, c 0.84.

Figs. 12-16. Aedeagi of Pseudanophthalmus species of inexpectatus group.
12. P. inexpectatus orientalis n. ssp. 13. P. puteanus n. sp. 14. P. cnephosus n. sp.
15. P. parvus n. sp. 16. P. umbratilis n. sp.

60 FIELDIANA: ZOOLOGY, VOLUME 62

P. i. orientalis is slightly greater in length (3.87 + 0.04 mm.)
than the nominate subspecies. The average width of the two forms,
however, is identical (1.25 + 0.02 mm.); hence orientalis is propor-
tionately narrower (range of width index 0.30-0.34, mean 0.32; range
of elytral index 0.57-0.61, mean 0.59) than the nominate form (range
of width index 0.35-0.36, mean 0.36; range of elytral index 0.60-
0.64, mean 0.63). The chaetotaxial index of orientalis, like that of
the nominate form, is high (range 0.74-0.96, mean 0.84).

The aedeagus of orientalis is slightly shorter (the four measured
range from 0.90-0.98 mm., average 0.94 mm.) than that of inexpec-
tatus s. str. The genital index of orientalis is somewhat smaller
(range 0.22-0.26, mean 0.24), therefore, than that of the nominate
form (range 0.27-0.29, mean 0.28). In form and in the nature of the
copulatory apparatus, however, the aedeagi of these two forms are
indistinguishable. Because of their great similarity these two forms
are judged to be at most subspecifically distinct.

Wilson Cave and Phillips Cave lie near tributaries of Green River
about 40 miles east of the caves in which inexpectatus s. str. is found.
These caves are in the Pennyroyal Plateau southwest of the Blue-
grass in Mississippian limestones of an older age (Osagian series)
than those in which Mammoth Cave is formed (Meramecian series).

Pseudanophthalmus puteanus, new species

Type series. — Holotype and 1 paratype from Old Well Cave, 0.6
mile southeast of Nevada, Mercer Co., Ky., June 26, 1965 (J. R.
Holsinger).

Holotype male (FMNH). Fig. 13. TL 3.47, HL 0.90, HW 0.57,
PL 0.63, PW 0.77, EL 1.93, EW 1.17, DE 0.67, ANT 2.23, AED
1.06, c 0.75.

P. puteanus has a body length (3.46 + 0.04 mm.) comparable to
that of inexpectatus s. str. It is considerably narrower than that
form in width (1.12 ± 0.04 mm.), however, hence is comparable to
i. orientalis in width index (range 0.31-0.34, mean 0.32) and elytral
index (range 0.58-0.60, mean 0.59). The aedeagus of the single spe-
cimen measured is slightly larger in length (1.06 mm.) than the ae-
deagi of the inexpectatus forms. The aedeagal index (0.31) is there-
fore distinctly higher than that of i. orientalis and somewhat higher
than that of i. inexpectatus. The chaetotaxial index of this species
(range 0.70-0.75, mean 0.72) is a bit lower than that of the inexpecta-
tus forms. The overall form of the aedeagus of this species and the

KREKELER: CAVE BEETLES 61

shape and proportion of the copulatory pieces are very similar to
those of inexpectatus.

On the basis of morphology alone this form could be considered
another subspecies of inexpectatus. The following facts, however,
indicate that there is little, if any, possibility of gene exchange be-
tween inexpectatus and this form: 1) Old Well Cave is 35 miles dis-
tant from the nearest cave (Phillips) from which a form of inexpec-
tatus is known; 2) Old Well Cave is near a stream whose waters
eventually drain north and west via Beech Fork and Rolling Fork
to the Ohio River whereas the caves in which inexpectatus is found
are in the Green River drainage basin; 3) Old Well Cave is in rocks
of the Bluegrass which are separated from the limestones of the
Pennyroyal Plateau and the Mammoth Cave Plateau in which the
inexpectatus forms are found by a series of non-caverniferous rocks
which comprise Muldraugh's Hill.

Pseudanophthalmus cnephosus, new species

Type series. — Holotype, allotype, and 28 paratypes from Eli
Reed Cave, 6.5 miles east-southeast of Hodgenville, Larue Co., Ky.,
Aug. 17, 1957 (C. H. K. and J. Rittman).

Holotype male (FMNH). Fig. 14. TL 3.50, HL 0.90, HW 0.60,
PL 0.65, PW 0.75, EL 1.95, EW 1.15, DE 0.80, ANT 2.15, AED
0.91, c 0.79.

Allotype female (FMNH). TL 3.65, HL 1.00, HW 0.65, PL
0.70, PW 0.80, EL 1.95, EW 1.25, DE 0.70, ANT 2.20, c 0.86.

The body length of P. cnephosus (3.77 ± 0.03 mm.) is intermedi-
ate in the range of the inexpectatus group, as are the length of the
aedeagus (mean of four measured is 0.94 mm.) and genital index
(mean is 0.25). The chaetotaxial index (range 0.73-0.93, mean 0.85)
also is not unusual. What is most distinctive of this species is that
the longer of the two copulatory pieces (the right one) is more than
two-thirds the length of the aedeagus (the four measured range from
0.67 to 0.69, average 0.68, the length of the aedeagus). This is in
contrast to the other species of the inexpectatus group in which the
proportion for individual aedeagi ranges from 0.45 to 0.57. In addi-
tion, the apex of the elytra is distinctly truncate, or at least obliquely
truncate, in cnephosus whereas in the other species of the group it
tends to be more rounded.

Eli Reed Cave is formed in Mississippian limestones near the
eastern margin of the Pennyroyal Plateau. It is on the divide be-

62 FIELDIANA: ZOOLOGY, VOLUME 62

tween Rolling Fork immediately to the east and McDougal Creek
which drains westward via Nolin River into Green River.

Pseudanophthalmus parvus, new species

Type series. — Holotype, allotype, and 20 paratypes from Tatum
Cave, 1.8 miles west-southwest of Riley, Marion Co., Ky., holotype,
allotype, and 18 paratypes, Aug. 17, 1957 (C. H. K. and J. Rittman) ;
2 paratypes, Oct. 9, 1965 (T. C. Barr).

Holotype male (FMNH). Fig. 15. TL 3.45, HL 0.90, HW 0.60,
PL 0.65, PW 0.80, EL 1.90, EW 1.15, DE 0.70, ANT 2.10, AED
0.86, c 1.00.

Allotype female. TL 3.40, HL 1.00, HW 0.60, PL 0.60, PW 0.75,
EL 1.80, EW 1.05, DE 0.65, ANT 1.90, c 1.16.

P. parvus, with a length of 3.49 ± 0.03 mm., is one of the smaller
species of the inexpectatus group. The aedeagus of this species is of
intermediate length (the three measured range from 0.82-0.86 mm.,
average 0.84 mm.) and the resultant genital index (range 0.24-
0.25, mean 0.25) is also intermediate for the group. The copulatory
pieces are slightly less than half the length of the aedeagus (range
0.46-0.48, mean 0.47). The chaetotaxial index of parvus (range 0.85
-1.18, mean 0.99) is the highest in the inexpectatus group. The pro-
notum is somewhat more wide in comparison to its length in this
species than in the other species of this group. (The average pro-
notal index values are: parvus 1.22, cnephosus 1.15, puteanus 1.20,
inexpectus s. str. 1.19, i. orientalis 1.15, umbratilis 1.18; there is so
much overlapping in range between forms, however, that this char-
acteristic is of little value).

The most distinctive characteristics of this species are that the
two copulatory pieces are only slightly different in length and that
the right one is clearly denticulate. In the forms of the inexpectatus
group described above the right copulatory piece is toothed and
considerably longer than the left; in umbratilis, described below, the
right copulatory piece is longer than the left but is not denticulate.

Tatum Cave is near the southwest margin of the Bluegrass,
adjacent to Muldraugh's Hill which bounds this section to the south
and west. Followell Creek on which the cave lies drains into the
North Rolling Fork.

Pseudanophthalmus umbratilis, new species

Type series. — Holotype, allotype, and 27 paratypes from Robin-
son Cave, 4.5 miles west-northwest of Lancaster, Garrard Co., Ky.,

KREKELER: CAVE BEETLES 63

Aug. 16, 1957 (C. H. K. and J. Rittman); 6 paratypes from Dix
Dam Cave, 2 miles northwest of Buena Vista, Garrard Co., Ky., 4
on July 2, 1957 (C. H. K. and J. Rittman), 2 on Dec. 16, 1962 (M. D.
Hassel) ; 1 paratype from Arnold's Cave, 1.2 miles west-northwest of
Bryantsville, Garrard Co., Ky., July 2, 1957 (C. H. K. and J.
Rittman); 1 paratype from Reid's (=Picadome) Cave, 2 miles south-
west of Lexington courthouse, Fayette Co., Ky., July 1, 1957 (C. H.
K. and J. Rittman) ; 1 paratype from Swope Cave, 4.5 miles north
of Versailles, Woodford Co., Ky., July 3, 1957 (C. H. K. and J.
Rittman) ; 5 paratypes from Clifton Cave, 0.6 mile east-southeast of
Clifton, Woodford Co., Ky., 1 on Jan. 19, 1963 (T. C. Barr), 4 on
June 24, 1963 (T. C. Barr); 2 paratypes from Hayden Cave, 0.5
mile south-southeast of Gratz, Owen Co., Ky., July 9, 1966 (R.
Norton) .

Holotype male (FMNH). Fig. 16. TL 3.90, HL 1.05, HW 0.65,
PL 0.75, PW 0.85, EL 2.10, EW 1.25, DE 0.90, ANT 2.35, AED
0.72, c 0.74.

Allotype female. TL 4.05, HL 1.15, HW 0.70, PL 0.75, PW 0.90,
EL 2.15, EW 1.35, DE 0.80, ANT 2.40, c 0.80.

There is a great deal of variation in the body length of P. um-
bratilis (from 3.20-4.20 mm.) so that a small sample of specimens
from a given cave may appear quite different than that from an-
other. But the total range in size variation of the species is displayed
in the one cave, Robinson Cave, in which this form has been taken
in any number. The average length of the specimens from Robinson
Cave (3.86 ± 0.04 mm.) is very close to that of all the specimens
included in this species (3.75 + 0.04 mm.). The aedeagi of this
species range from 0.68-0.78 mm. and average 0.74 mm. in length,
a size considerably smaller than that of the other species of the
group. The aedeagal index (range 0.18-0.22, mean 0.20) is the
lowest in the group, though not sharply separated from that of
inexpectatus orientalis. The chaetotaxial index, which ranges from
0.62-1.00, averages 0.78.

The aedeagus of P. umbratilis terminates with a small knob just
beyond the elongate aperture. The right copulatory piece is dis-
tinctly longer than the left and about half (range 0.45-0.51, mean
0.48) the length of the aedeagus. In contrast to the other species
of the inexpectatus group, neither copulatory piece is toothed.

All the caves in which P. umbratilis is found are in Ordovician
limestones which form the bedrock of the Inner Bluegrass. More-

64 FIELDIANA: ZOOLOGY, VOLUME 62

over, all these caves are in the drainage basin of the Kentucky River
and two of its major tributaries, Dix River and Elkhorn Creek.
Interestingly, all the specimens taken of this species are from caves
in which forms of the horni group are present. But whereas this
single form of the inexpectatus group is found, several distinct forms
of the horni group are represented: horni s. str. in Reid's Cave;
h. abditus in Swope Cave; h. caecus in Clifton Cave; elongatus in
Dix Dam Cave, Arnold's Cave, and Robinson Cave; and desertus
desertus in Hayden Cave. In all but Reid's Cave and Swope Cave
umbratilis is distinctly smaller than the form of the horni group.
Though we noted in our field notes that we thought that beetles of
two different sizes were present in these caves, we were unable to
detect any difference in habitat preference between the larger and
smaller forms. The only cave in which we found P. umbratilis to be
clearly more abundant than the horni group form is in Robinson
Cave.

BARRI GROUP

Length 3.9-4.9 mm. Microsculpture of body pruinose, especially conspicuous
on elytra. Labrum with poorly defined median tooth. Posterior angles of pro-
notum rectangular with straight or slightly concave lateral margins; basal margin
straight or slightly concave, with a rather distinct median ridge extending an-
teriorly from it to disc; disc pubescent. Elytra convex, covered with short pubes-
cence; reflexed margin rather wide, ciliated throughout length, feebly serrate at
humeri; humeri rounded; striae fairly well impressed on disc, less distinct laterally;
1st discal seta approximately at level of 4th marginal humeral papilla; chaetotaxial
index intermediate and quite variable; apex truncate to rounded, slightly dehiscent;
recurved portion of apical stria well impressed, clearly joining the 3rd stria.
Inner margin of femora of anterior legs distinctly angulate (rather than broadly
rounded) about one-third the distance from the base of the femora.

Aedeagus moderately elongate, arcuate; basal plate well developed, terminal;
apex produced. Transfer apparatus about two-fifths the length of the aedeagus,
consisting of a single spatulate or twisted, scoop-shaped element covered with min-
ute acute tubercles. Parameres relatively long, bearing 3 or 4 setae.

Pseudanophthalmus barri, new species

Type series. — Holotype, allotype, and 34 paratypes from Indian
Cave, 0.5 mile southwest of Charlestown, Clark Co., Ind., holotype
allotype, and 30 paratypes, Aug. 2, 1959 (C. H. K. and W. Bloom),
4 paratypes, Jan. 26, 1957 (L. Hubricht) ; 6 paratypes from Peyton
Beechwood Cave, 3 miles southwest of Charlestown, Clark Co., Ind.,
Aug. 2, 1959 (C. H. K. and W. Bloom).

KREKELER: CAVE BEETLES 65

Holotype male (FMNH). Figs. 17, 17a. TL 3.94, HL 1.03,
HW 0.67, PL 0.77, PW 0.90, EL 2.13, E W 1.27, DE 0.83, ANT 2.47,
AED 0.99, c 0.79.

Allotype female (FMNH). TL 4.21, HL 1.07, HW 0.73, PL
0.83, PW 0.97, EL 2.30, EW 1.33, DE 0.97, ANT 2.57, c 0.58.

The two species of the barri group, 6am and troglodytes, are
virtually identical in length {barri 4.36 + 0.03 mm., troglodytes 4.28
± 0.05 mm.) and width {barri 1.39 ± 0.01 mm., troglodytes 1.40 ±
0.02 mm.). Moreover, the elytral index of the two forms is com-
parable {barri averaging 0.60, troglodytes averaging 0.62). P. barri
ranges from testaceous to dark ferrugino-testaceous in color. It has
a mean chaetotaxial index of 0.65 (range 0.52-0.79). The apex of
the elytra is truncate or obliquely truncate. The recurrent apical
stria extends anteriorly some distance in a straight line or a weak
curve subparallel to the suture then curves inward rather sharply to
join the 3rd stria.

The length of the aedeagus of barri (the four measured range
from 0.99-1.11 mm., average 1.04 mm.) is about one-fourth the
body length (range of genital index is 0.24-0.26). The median lobe
of the aedeagus is very distinctive in shape. There is a large bulge
on the left side of the median lobe just beyond half its length; this
is seen most readily in a dorsal view of the aedeagus (fig. 17a) . The
aperture is on the right side of the lobe opposite the bulge rather
than in the usual dorsal position. The apex, which is moderately
extended, curves gently downward to a blunt point. The transfer
apparatus consists of a single twisted, scoop-shaped element covered
with minute acute tubercles.

The caves in which P. barri is found lie near Pleasant Run which
drains via Silver Creek to the Ohio River. The caves are developed
in limestones which are characteristic of the Outer Bluegrass. But
inasmuch as these limestones are covered in this portion of Indiana
with glacial till of Illinoian age, geomorphologists include this area
within the Central Lowlands province.

It gives me great pleasure to name this species in honor of Dr.
Thomas C. Barr, Jr., by whose courtesy this form first came to my
attention and whose work with the anophthalmids the last decade
has set standards worthy of emulation.

Pseudanophthalmus troglodytes, new species

Type series. — Holotype, allotype, and 21 paratypes from High-

66 FIELDIANA: ZOOLOGY, VOLUME 62

baugh Cave, 4.5 miles northwest of Jeffersontown, Jefferson Co.,
Ky., Oct. 7, 1964 (N. Whitehead).

Holotype male (FMNH). Fig. 18. TL 4.47, HL 1.17, HW 0.77,
PL 0.87, PW 1.00, EL 2.43, EW 1.50, DE 1.10, ANT 2.77, AED
1.19, c 0.62.

Allotype female (FMNH). TL 4.24, HL 1.17, HW 0.77, PL
0.83, PW 1.00, EL 2.23, EW 1.37, DE 0.93, ANT 2.50, c 0.70.

It is practically impossible to distinguish P. troglodytes from
barri on the basis of external features. As indicated above, they are
virtually identical in length, width, and elytral index. P. troglodytes
tends to be darker in color, ranging from ferrugino-testaceous to
fusco-testaceous. The chaetotaxial index of troglodytes (mean 0.72,
range 0.59-0.88) is slightly higher than that of barri. The apex of
the elytra tends to be more rounded in troglodytes than barri. The
variability and wide overlapping of these several traits is so great,
however, that they are of little diagnostic value. Of somewhat
more value in distinguishing between these two species is the form
of the recurrent apical stria. In troglodytes it curves gently through-
out its length to join the 3rd stria whereas in barri it extends some
distance subparallel to the suture before curving inward.

P. troglodytes is readily distinguished from barri on the basis of
aedeagal characters. The aedeagus of troglodytes is longer (the five
measured range from 1.19-1.26 mm., average 1.24 mm.) than that of
barri. Inasmuch as the body length of the two species is the same,
this means that the genital index of troglodytes (range 0.27-0.29) is
higher than that of barri. The median lobe of the aedeagus of
troglodytes lacks the peculiar post-median bulge of barri and the
associated lateral aperture; in troglodytes the aperture of the aedeagus
is in the usual dorsal position. The transfer apparatus consists of a
single spatulate element covered with minute acute tubercles. The
apex of the median lobe is moderately extended and curves gently
upward and terminates in a small but distinct knob. The parameres
of troglodytes are more attenuate than those of barri.

Highbaugh Cave is less than 15 miles distant from the caves in
which P. barri is found, but it is separated from them by the Ohio
River. The Middle Fork of Beargrass Creek near which Highbaugh
Cave lies drains into the Ohio River. The cave is located in the
Outer Bluegrass, in rocks of Silurian or Devonian age, just beyond
the reach of Illinoian glaciation.

Figs. 17-22. Aedeagi of Pseudanophthalmus species of barri and rittmani
groups. 17. P. barri, n. sp.; 17a, dorsal view of terminal half of median lobe.
18. P. troglodytes, n. sp. 19. P. exiguus exiguus, n. sp. and ssp. 20. P. exiguus fur-
tivus, n. ssp., terminal half of median lobe. 21. P. catoryctos, n. sp. 22. P. condi-
tus, n. sp.

67

68 FIELDIANA: ZOOLOGY, VOLUME 62

RITTMANI GROUP

Length 3.5-6.6 mm. Microsculpture of body pruinose, especially conspicuous
on elytra. Labrum with weakly or well denned median tooth. Posterior angles of
pronotum rectangular with straight lateral margins; basal margin nearly straight;
disc convex, sparsely pubescent, with several long hairs on either side of midline.
Elytra convex, covered with short, sparse pubescence; reflexed margin of moderate
width, ciliated throughout length but most regularly at humeri; humeri rounded
or sloping, lacking serrations; striae of disc evident to weakly impressed, lateral
striae indistinct or absent; 1st discal seta approximately at level of 4th marginal
humeral papilla; chaetotaxial index low, averages ranging from 0.49 to 0.66; apex
truncate to somewhat pointed, slightly dehiscent; recurved portion of apical stria
distinctly to weakly impressed, distinctly to vaguely joining the 3rd stria.

Aedeagus relatively short; median lobe relatively broad and short, relatively
flat ventrally, moderately arched dorsally; basal plate well developed, terminal;
apex produced. Copulatory pieces from less than one-fourth to more than one-
half the length of aedeagus; composed of two elements, each the extension from a
basal cylinder; one element is semicylindrical and partially ensheaths the other.
Parameres moderately long, bearing 3 or 4 setae.

Pseudanophthalmus rittmani, new species

Type series. — Holotype, allotype, and 13 paratypes from Baker
Cave, 1.7 miles east of Cobhill, Estill Co., Ky., Aug. 15, 1957 (C. H.
K. and J. Rittman); 15 paratypes from Watson Cave, 0.8 mile
north-northeast of Cobhill, Estill Co., Ky., June 30, 1957 (C. H. K.
and J. Rittman) ; 10 paratypes from Betsy Cave, 4 miles south-south
east of Bowen, Powell Co., Ky., July 1, 1957 (C. H. K. and J. Ritt-
man); 1 paratype from Townsend Cave, 4.5 miles east- of Cobhill,
Estill Co., Ky., Aug. 15, 1957 (C. H. K. and J. Rittman).

Holotype male (FMNH). Fig. 11. TL 6.05, HL 1.75, HW 0.95,
PL 1.00, PW 1.20, EL 3.30, EW 2.05, DE 1.40, ANT 3.90, AED
1.42, c 0.42.

Allotype female (FMNH). TL 5.30, HL 1.50, HW 0.85, PL
0.95, PW 1.05, EL 2.85, EW 1.65, DE 1.15, ANT 3.30, c 0.48.

P. rittmani is a large (5.74 ± 0.06 mm.), moderately convex
anophthalmid. The pronotum, with sides gently sinuate the entire
length, is widest at one-fifth its length. The elytra are moderately
wide and have relatively sloping humeri. The striae of the disc
are evident though quite broad, shallow, and irregular, but the
lateral striae are indistinct; the striae are vaguely and irregularly
punctured. The chaetotaxial index is low (range 0.42-0.56, mean
0.49). The apex of the elytra is rounded or somewhat pointed. The
recurved portion of the apical stria is distinct; it is gently bowed

KREKELER: CAVE BEETLES 69

except anteriorly where it curves inward rather sharply to join the
3rd stria which is very feebly impressed at this level.

The aedeagus of rittmani is relatively large, the six measured
ranging from 1.26-1.42 mm., averaging 1.34 mm. in length. It is not
large in proportion to the body, however; the genital index ranges
from 0.21-0.24, averages 0.22. The median lobe is quite broad.
The produced apex forms an open trough which is slightly expanded
at its terminus. The copulatory pieces are half or more (0.49-0.56)
the length of the aedeagus. The two elements are extensions from
two concentric basal cylinders. The ventral part of the outer cylin-
der develops as a broad, trough-shaped, hyaline element. The
ventral portion of the inner cylinder develops as a semicylindrical
element, toothed apically, which lies within but extends a bit beyond
the other element. The parameres bear 3 or 4 setae.

The caves in which rittmani is found lie immediately east of the
Bluegrass in the Cumberland Plateau section of the Appalachian
Plateaus province, in rocks of Mississippian and /or Pennsylvanian
age. Baker, Watson, and Townsend caves are in uplands about
300-500 ft. above the Kentucky River, which are drained by deeply
incised tributaries of Miller's Creek which flows into the Kentucky
River. Betsy Cave, only some 5 miles distant, is on the other side of
a divide; it is on the side of a hollow cut by a branch of the deeply
cut South Fork of the Red River which drains by way of the Red
River to the Kentucky River.

Interestingly, in all four caves from which P. rittmani has been
taken is also found another, and clearly smaller, species of this group,
P. exiguus. In three of these caves a good series was taken of both
species. In these cases our field notes show that we recognized two
size classes and observed that they occupy two distinct micro-
habitats. The smaller beetles were found under rocks and in wet
sandy mud near the water's edge and scurried quickly when dis-
turbed. The larger beetles were found crawling on moist sandy mud
and under rocks some distance away from the water and moved
less rapidly than the smaller form when disturbed.

It is my great pleasure to name this species in honor of John
Rittman who so capably assisted me in collecting in the summer of
1957 and who has been a frequent and proficient collecting com-
panion since that time.

70 FIELDIANA: ZOOLOGY, VOLUME 62

Pseudanophthalmus exiguus exiguus, new species and subspecies
Type series. — Holotype, allotype, and 13 paratypes from Watson
Cave, 0.8 mile north-northeast of Cobhill, Estill Co., Ky., June 30,
1957 (C. H. K. and J. Rittman); 13 paratypes from Baker Cave,
1.7 miles east of Cobhill, Estill Co., Ky., Aug. 15, 1957 (C. H. K.
and J. Rittman); 13 paratypes from Betsy Cave, 4 miles south-
southeast of Bowen, Powell Co., Ky., July 1, 1957 (C. H. K. and J.
Rittman) ; 2 paratypes from Townsend Cave, 4.5 miles east of Cob-
hill, Estill Co., Ky., Aug. 15, 1957 (C. H. K. and J. Rittman); 4
paratypes from Natural Bridge Cave, 2 miles southeast of Slade,
Powell Co., Ky., 3 on June 28, 1957 (C. H. K. and J. Rittman),
1 on May 27, 1962 (T. C. Barr); 14 paratypes from Pinnacle Cave,
5.5 miles north-northwest of Heidelberg, Lee Co., Ky., Dec. 1, 1962
(T. C. Barr); 6 paratypes from Cave Hollow Cave, 2.5 miles south-
east of Crystal, Lee Co., Ky., Dec. 1, 1962 (T. C. Barr); 1 paratype
from Stillhouse Cave, 2.5 miles east-southeast of Crystal, Lee Co.,
Ky., June 29, 1963 (T. C. Barr and R. Kuehne); 1 paratype from
Ash Cave, 4 miles east-southeast of Crystal, Lee Co., Ky., Feb. 17,
1963 (P. Dickson).

Holotype male (FMNH). Fig. 19. TL 4.25, HL 1.20, HW 0.70,
PL 0.80, PW 0.90, EL 2.25, EW 1.40, DE 1.05, ANT 3.00, AED
0.90, c 0.60.

Allotype female (FMNH). TL 3.75, HL 1.00, HW 0.65, PL
0.70, PW 0.85, EL 2.05, EW 1.30, DE 0.90, ANT 2.45, c 0.62.

P. exiguus exiguus is significantly smaller (3.98 ± 0.03 mm.) than
rittmani; the largest exiguus specimen is 4.73 mm., the smallest
rittmani is 4.85. The pronotum, with sides gently sinuate the entire
length, is widest at one-third its length. The humeri of the elytra
are rounded. The 1st and 2nd striae are feebly impressed and
impunctate; the 3rd stria and those beyond it are very indistinct or
absent. The chaetotaxial index, though low (range 0.56-0.75, mean
0.65), is somewhat higher than that of rittmani. The apex of the
elytra is truncate or obtusely angled. The recurrent portion of the
apical stria is rather distinct; it extends anteriorly subparallel to the
internal margin toward the 4th stria then curves sharply but indis-
tinctly inward to the level of the indistinct 3rd stria.

The aedeagus of exiguus s. str. is significantly smaller (the 16
measured range from 0.82-0.96 mm., average 0.88 mm., in length)
than that of rittmani. The genital index (range 0.20-0.24, mean
0.22), however, is comparable to that of rittmani. The median lobe

KREKELER: CAVE BEETLES 71

is of moderate width. The elongate apex curves gently upward and
ends in a slightly swollen knob. The copulatory pieces are one-third
to two-fifths the length of the aedeagus. As in rittmani the two
elements are extensions from two concentric basal cylinders, but the
form of the elements and their relation to one another are quite dif-
ferent in the two species. In exiguus the ventral portion of the inner
cylinder develops as a long, spatulate, hyaline element. The dorsal
portion of the outer cylinder develops as a semicylindrical, partially
chitinized membrane which partially ensheaths and extends a short
distance beyond the other element. The parameres bear 3 or 4 setae.

In the series of specimens on the basis of which this form is de-
scribed there is considerable variation in size, amount of pubescence
on pronotum and elytra, distinctness and form of recurrent apical
stria, shape of produced apex of aedeagus, width of spatulate trans-
fer element, and relative length of sheathing and spatulate elements.
Some of the variation, particularly of the aedeagi, seems attributable
to differences in preparation and orientation. No pattern of varia-
tion has been recognized which would warrant division of this series
into several forms.

As indicated above, exiguus s. str. is associated with rittmani in
all four caves where that form has been found. In these caves
exiguus is recognizably smaller, occupies a different subhabitat, and
moves more quickly than rittmani.

The caves in which only exiguus s. str. is found are, like those in
which it is associated with rittmani, located in the Cumberland
Plateau in rocks of Mississippian and/or Pennsylvanian age. They
are in uplands from nearly 200 to almost 400 ft. above the Kentucky
and Red rivers. A small stream near Pinnacle Cave drains directly
into the Kentucky River. Stillhouse, Ash, and Cave Hollow caves
are along Big Sinking Creek, or its tributaries, which drains by way of
Millers Creek into the Kentucky River. Natural Bridge Cave lies
near the Middle Fork of Red River which drains by way of the Red
River into the Kentucky River.

Pseudanophthalmus exiguus furtivus, new subspecies

Type series. — Holotype and allotype from California Cave, 1.5

miles north-northeast of Ravenna, Estill Co., Ky., Aug. 16, 1957

(C. H. K. and J. Rittman).

Holotype male (FMNH). Fig. 20. TL 3.70, HL 0.95, HW 0.65,

PL 0.70, PW 0.80, EL 2.05, EW 1.30, DE 0.85, ANT 2.30, AED

0.85, c 0.73.

72 FIELDIANA: ZOOLOGY, VOLUME 62

Allotype female (FMNH). TL 3.95, HL 1.05, HW 0.70, PL
0.75, P\V 0.85, EL 2.15, EW 1.35, DE 1.00, ANT 2.40, c 0.60.

P. exiguus furtivus falls within the range of the variation expressed
in exiguus s. str. with one exception. The apex of the ensheathing
element is distinctly denticulate in e. furtivus whereas such denticula-
tions are not present in any of the specimens of the nominate form.
This peculiarity seems sufficiently important to warrant considering
this a form sub-specifically distinct from the nominate form.

California Cave. 7 miles distant from the nearest cave in which
the nominate form is found, is about 600 ft. above the Kentucky
River on a ridge of the Cumberland Plateau. The several small
streams draining this ridge are tributaries of creeks flowing into the
Kentucky River.

Pseudanophthalmus catoryctos, new species

Type series. — Holotype, allotype, and 7 paratypes from Adams
Cave, 5 miles south-southwest of Richmond. Madison Co., Ky.,
Aug. 15, 1964 (T. C. Barr and S. Peck).

Holotype male (FMNH). Fig. 21. TL 3.54. HL 0.90, HW 0.67,
PL 0.67, PW 0.83, EL 1.97, EW 1.23, DE 0.73, ANT 2.37, AED
0.68, c 0.70.

Allotype female (FMNH). TL 3.67. HL 0.97, HW 0.70, PL
0.70, PW 0.90, EL 2.00, EW 1.17, DE 0.80, ANT 2.40, c 0.66.

P. catoryctos, with a length of 3.53 + 0.07 mm., is the smallest
species of this group. As in exiguus, the pronotum is widest at one-
third its length and has gently sinuate sides, the humeri of the elytra
are rounded, and the 1st and 2nd striae are feebly impressed and
impunctate while the 3rd stria and those beyond it are very indis-
tinct. The chaetotaxial index (range 0.58-0.78, mean 0.66) is like
that of exiguus. The apex of the elytra is obtusely angled. The
recurrent portion of the apical stria is well impressed; it follows a
gently rounded course, eventually curving inward to the indistinct
3rd stria.

The aedeagus of catoryctos is distinctly shorter than that of
exiguus, each of the three measured being 0.68 mm. in length. More-
over the genital index (range 0.18-0.19, mean 0.19) is lower than
that of exiguus and rittmani. The median lobe is of moderate width.
The apex, which extends only a short distance beyond the aperture.
is quite thick and ends in a blunt angle. The copulatory pieces are

KREKELER: CAVE BEETLES 73

just less than one-fourth the length of the aedeagus. The two ele-
ments may well be, as is more clearly the case in the other species of
this group, extensions from two concentric basal cylinders. An elon-
gate hyaline element which lies above and to the left of and partially
encloses the second element may be interpreted as the extension of
an outer cylinder. The second element, a short triangular piece,
may be interpreted as the extension of an inner cylinder. The para-
meres bear 3 setae.

P. catoryctos occupies the same cave in the Outer Bluegrass south-
east of Lexington as pholeter of the horni group. It is readily distin-
guished from that species by its smaller size, lack of humeral ser-
rations, lack of angulate mesosternum, and its aedeagal character-
istics. Though catoryctos is thus found in a different physiographic
province than rittmani and exiguus, it is from that part of the Blue-
grass immediately adjacent to the Cumberland Plateau and the caves
lie separated only by a distance of just over 20 miles. The gross
morphology of the aedeagus of these three species is so different that
it is not immediately apparent that they may be included in a single
group. As described above, however, these forms are quite similar
in many features of their external morphology. That feature which
most strongly supports their inclusion in a single group is the ap-
parent development of the two copulatory pieces of each species as
extensions from two concentric basal cylinders. I have recognized
the possibility of this interpretation of the copulatory apparatus of
catoryctos after having been informed by Barr (pers. comm,) that he
has species from Jackson, Rockcastle, and Fayette counties linking
cartoryctos to exiguus.

INCERTAE SEDIS

Pseudanophthalmus conditus, new species

Type series. — Holotype, allotype, and 1 paratype from Lawrence
Cave, 0.5 mile south-southwest of Perryville, Boyle Co., Ky., Aug.
16, 1957 (C. H. K. and J. Rittman).

Holotype male (FMNH). Fig. 22. TL 4.60, HL 1.20, HW 0.85,
PL 0.85, PW 1.10, EL 2.55, EW 1.60, DE 0.95, ANT 2.75, AED
1.01, c 0.62.

Allotype female (FMNH). TL 4.50, HL 1.30, HW 0.80, PL 0.80,
PW 1.10, EL 2.40, EW 1.50, DE 0.95, ANT 2.65, c 0.61.

P. conditus is a species of moderate length (4.46 ±0.09 mm.). Its
microsculpture is pruinose, in contrast to all the other forms in the

74 FIELDIAXA: ZOOLOGY, VOLUME 62

southwestern part of the geographic range considered in this paper.
The inner margin of the profemora is broadly rounded. The meso-
sternum follows a broad convex curve to the anterior narrowing of
the mesothorax. The last abdominal sternite of the male is weakly
indented.

The head, pronotum. and elytra are pubescent, with the pube-
scence of the head being more sparse than elsewhere. The labrum
possesses a low. broad median tooth. The pronotum. which is widest
at about one-fourth its length, has rectangular posterior angles with
straight lateral margins. The humeri of Ithe elytra are broadly
rounded ; they possess more closely spaced setae than the remainder
of the reflexed margin, but are not serrulate. The striae of the disc
are weakly impressed and not punctate; only traces of striae beyond
the 3rd are distinguishable. The elytral apex is broadly rounded.
The distinct recurrent portion of the apical stria after paralleling the
suture a short distance curves gently inward to join the 3rd stria.

The single aedeagus of conditus measured is of moderate length,
1.01 mm. The median lobe is relatively slender; it is flattened
ventrally, weakly arched dorsally. The basal bulb is of moderate
size, but the basal plate is quite large. The distal aperture extends
over half the length of the median lobe. The apex extends a short
distance beyond the aperture; it is straight and thin and ends in a
blunt point. The parameres bear 4 setae. The transfer apparatus
includes an elongate, heavily chitinized element which has a weakly
bilobate tip. This element lies dorsally in the median lobe. Ventral
to this element, and so closely associated with it that the limits of
each are impossible to distinguish, is an extensively developed mem-
branous sac. This sac extends some distance beyond the chitinized
element to the tip of the aperture; it is so densely packed there that
it almost appears as another element.

Lawrence Cave is near the southern margin of the Bluegrass.
close to Muldraugh's Hill. A small stream nearby flows into Chaplin
River whose waters drain north and west via Beech Fork and Rolling
Fork to the Ohio River.

P. conditus does not appear to be closely related to any other of
the species described above. In its external features, notably in its
possession of pruinose microsculpture and lack of serrulate humeri, it
is most similar to species of the rittmani group. But its transfer
apparatus cannot, as in that group, be interpreted as extensions
from two basal concentric cylinders. Similarly, the transfer ap-
paratus of conditus shows no similarity to the characteristics of the

KREKELER: CAVE BEETLES 75

horni and inexpectatus groups. Its structure is sufficiently unclear
that it could not be ruled out as being similar to that of the barri
group species. But certain of the external characteristics of conditus,
specifically its lack of angulate profemora and serrulate humeri,
would seem to rule out the possibility of its inclusion in that group.

DISCUSSION
1

Twenty-six forms of anophthalmids are considered above, 24 of
them new, from the Kentucky Bluegrass and nearby areas. Nine-
teen of the new forms are described as species, five as subspecies.
In my study of Indiana anophthalmids (Krekeler, 1958) I con-
sidered all distinguishable forms as species. This was done on the
assumption that caves are typically isolated habitats and that
cavernicoles do not move between caves unless caves are known to
be interconnected. Forms in different cave systems, it was then
argued, would not be exchanging genes; these forms are thus repro-
ductively isolated; and reproductively isolated and distinguishable
forms, whether this isolation is extrinsically or intrinsically deter-
mined, are species. When I reviewed the dispersal of anophthalmids
in further detail (Krekeler, 1959), I decided for reasons outlined in
that paper that the burden of proof should be reversed and that
cavernicoles should be assumed to be able to move between caves
unless there is evidence of a probable barrier to dispersal. Under
these circumstances the subspecific rank is a useful tool in the descrip-
tion of anophthalmids, and is accordingly used in this paper. Those
forms which are judged by morphological criteria as probably ex-
changing genes are designated subspecies, while those judged by these
criteria as probably not exchanging genes or for which there is
evidence that extrinsic factors probably prevent gene exchange are
designated species.

The forms described above as subspecies have a geographic dis-
tribution such that, and are from caves found in strata related to one
another in such a way that, some gene exchange appears to be pos-
sible. The several subspecies of P. horni are from the Inner Bluegrass
where the limestone beds are nearly horizontal so that dispersal
between caves may occur by way of underground crevices from time
to time. But each subspecies is found in caves which are relatively
close to one another and/or of a limited drainage basin so that gene
exchange is presumably much more regular. Though inexpectatus

76 FIELDIANA: ZOOLOGY, VOLUME 62

s. str. and i. orientalis are from caves in limestone beds of different
ages, there are not extensive non-caverniferous beds intervening
which would serve as a barrier; but the caves are sufficiently far
apart (some 40 miles) that gene exchange would not be regular. In
the case of P. desertus, it is not clear how regular gene exchange is
possible between the populations listed as comprising the nominate
subspecies. All the caves seem to be formed in Ordovician limestones
of the Inner Bluegrass, but there are intervening patches of Eden
Shale. It is consistent with the general pattern being considered,
however, that the only population which can be distinguished as a
distinctive form, d. major, is that which is most widely separated
from the others so that gene exchange would be the least regular.
On the other hand, as described above, P. puteanus is so similar
morphologically to inexpectatus that using such criteria alone it
would be considered a subspecies of that form. But since the geo-
graphic and physiographic facts indicate little possibility of gene
exchange between these forms puteanus is considered a species.

The distribution of the anophthalmids considered in this paper
(see fig. 23) is of great interest for several other reasons. First of all,
as I have pointed out previously (Krekeler, 1959), the fact that some
of the inexpectatus group forms are from the Bluegrass northeast of
Muldraugh's Hill (and outliers of it called the Knobs) and others are
from the Pennyroyal Plateau and the Mammoth Cave Plateau
southwest of Muldraugh's Hill gives us some definite information
concerning the dispersal of anophthalmids. Inasmuch as the Mul-
draugh's Hill cuesta is composed of rocks in which caves are seldom
found and the strata dip in such a way that caves completely travers-
ing Muldraugh's Hill are virtually impossible, one can essentially
rule out the possibility of the dispersal of anophthalmids between the
Bluegrass and the Pennyroyal Plateau by way of caves. Dispersal
must rather be or have been overland, possibly in leaf litter or in
crevices of the soil. Inasmuch as the single American anophthalmid
known from an epigaean environment (Barr, 1967c) and the nearest
epigaean relatives of the anophthalmids are inhabitants of cool moist
habitats, it seems quite likely, as Jeannel (1949) and Barr (1967c)
have suggested, that conditions prevailing during and immediately
following the glacial advances of the Pleistocene would have been
favorable to overland movement in this region.

KREKELER: CAVE BEETLES 77

Secondly, three of the species described are from caves lying
within the boundaries of the Illinoian glacial advance : chthonius and
barri from southeastern Indiana and krameri from southwestern
Ohio. These are the first American anophthalmids of well over a
hundred known forms to be taken from glaciated areas. It does not
seem possible that these anophthalmids could have survived in the
caves in which they are now found when the area was covered by
glacial ice sheets. Even if the channels were not filled with icy water
it seems quite improbable that food necessary for their survival
would have been available. It thus appears that the anophthalmids
entered these caves sometime since the maximum Illinoian glacial
advance.

Interestingly, the three species found north of the Ohio River
within the Illinoian glacial boundaries are of groups (horni and barri)
in which other species come from caves south of the Ohio River in
Kentucky. In the tenuis group which I considered in a previous
paper (Krekeler, 1958 — at that time the species and group was
thought to be eremita, but it was pointed out by Barr in 1960 that
this was in error) the same situation exists: barberi is in Kentucky,
tenuis and other species are in Indiana. In that case I suggested
that a stream the size of the present day Ohio River would be a bar-
rier to dispersal of anophthalmids and that movement between
Indiana and Kentucky must have occurred when the Ohio River was
a much smaller and shorter stream. This would have been prior to
the glacial advance (probably Kansan) which diverted waters carried
by the preglacial Teays River to the Ohio River basin. If forms of
the horni and barri groups have, by the same reasoning, not crossed
the Ohio River since the Kansan advance and have entered the caves
in which they are currently found subsequent to the Illinoian ad-
vance, for the reasons mentioned in the preceding paragraph, how is
their present distribution to be accounted for? The present forms

Fig. 23. (See pp. 78-79). Distribution of Pseudanophthalmus species of the
Kentucky Bluegrass and vicinity. The species inhabiting a cave are shown by
the symbols for which the code is given at lower right. The numbers designate
the position of the caves as follows: 1. Reid's=Picadome; 2. Phelps; 3. Russell;
4. Church; 5. Clifton; 6. Swope; 7. Weber #2; 8. Meece; 9. Weber; 10.
Nonesuch; 11. Britton; 12. Keene; 13. Old Fort; 14. Dix Dam; 15. Robin-
son; 16. Arnold; 17. Stevens Creek; 18. Jones; 19. Price; 20. Slack's; 21.
Hayden; 22. Beaver; 23. Wilson; 24. Morris; 25. Lowry; 26. Indian; 27.
Peyton Beechwood; 28. Freeland; 29. Townsend; 30. Cave Hill; 31. Adams;
32. White; 33. Wilson; 34. Phillips; 35. Old Well: 36. Eli Reed; 37. Tatum;
38. Highbaugh; 39. Baker; 40. Watson; 41. Betsy; 42. Natural Bridge; 43.
Pinnacle; 44. Cave Hollow; 45. Stillhouse; 46. Ash; 47. California; 48. Law-
rence.

I nd i a n a

^utf,,,^'^"

<£ h. horni
X0 h. caecus
# h.abditus
^) sol ivagus
(J e I opgatus
(■£) tenebrosus
^ d. desertus
dTTD d- major

(3 chthonius
£ ohioensis
(•) exoticus
/\ kram er i
A pholeter
£ rittmani
(3 e- ex iguus
<|§> e.furtivus
^ catoryctos

p^ i. inexpectatus

[ITT] i.orientalis

§ puteanus

[■] cnephosus

| parvus

] umbratilis

V barri

▼ troglodytes

\/ conditus

79

80 FIELDIANA: ZOOLOGY, VOLUME 62

and/or their ancestors must have moved between Kentucky and
Indiana-Ohio prior to the Kansan glacial advance. Such movement
could have been across the small pre-Kansan Ohio River and or by
way of the divide between the headwaters of the pre-Kansan Ohio
(probably just east of Madison, Ind.) and the Teays River. During
the Illinoian glacial advance some of the forms already in Indiana
and Ohio could have found refuge in areas there beyond the glacial
limits. Conditions following the withdrawal of the Illinoian ice sheet
could well have been, according to the reasoning used above, favor-
able to their dispersal overland to the caves where they are now
found.

In a number of caves from which the anophthalmids considered
above have been taken two or three species coexist. In fact, this is
almost the rule rather than the exception. As I have previously
argued (Krekeler, 1958) and as Barr (1967d) more recently concludes,
it seems most likely that speciation was allopatric and that as species
have extended their ranges they have become sympatric. Interest-
ingly, as noted above, where horni and inexpectatus group forms co-
exist only one inexpectatus form (umbratilis) can be recognized,
whereas five horni group forms are recognizable. This may be be-
cause the horni group forms are the earlier occupants and have diver-
sified in longer isolation while umbratilis is a more recent invader.
Or this may be because the horni group forms are more labile in an
evolutionary sense. There is no evidence to support one alternative
in preference to the other.

In the case of the coexisting inexpectatus and horni group forms
we did not observe any habitat preference though we did suspect
while collecting that more than one species was present because of
size differences. In the caves where barri and chthonius coexist we
did not suspect that more than one species was present. We did
note, however, that many beetles in these caves (and all but two of
them were 6am) were found rather close to the entrance and in drier
places than Indiana anophthalmids are typically found — in fact,
there are no streams in either of the two caves. P. chthonius, on the
other hand, was found on wet mud banks at the type locality and
was regularly taken from under rocks on wet gravel banks in Morris
Cave. These two species thus seem to have distinct habitat prefer-
ences. In the case of rittmani and exiguus we recognized in the field
that a larger and a smaller form exhibited habitat and behavioral

KREKELER: CAVE BEETLES 81

differences as described above. Of particular interest here is that
these two species are so similar in many respects that they are in-
cluded within the same species group. Still there is no need to
postulate sympatric speciation. Whether coexisting forms are of the
same or different species groups, range extensions following allopatric
speciation would appear to be the most likely explanation for present
sympatry. The distinct differences in the aedeagi of the two species
would seem to assure reproductive isolation. Niche differences, prob-
ably related to differences in size and habitat preferences, undoubt-
edly exist to account for the continued coexistence of the two species
in all these cases of sympatry. It so happens that in the case where
two forms of the same group coexist such differences in size and habi-
tat preference are readily distinguished.

SUMMARY

Nineteen new species and five new subspecies of Pseudanoph-
thalmus are described from the Kentucky Bluegrass and vicinity.
They are divisible into four species groups. The majority of the
horni group forms (horni s. str., horni caecus, horni abditus,
solivagus, elongatus, tenebrosus, desertus s. str., desertus
major, pholeter) are from the Bluegrass; chthonius is from caves
in the glaciated portion of southeastern Indiana in the same lime-
stone beds as are included in the Bluegrass; krameri is from a cave
in the glaciated portion of southwestern Ohio in the same limestone
beds as are included in the Bluegrass; ohioensis is from the High-
land Rim which extends into southwestern Ohio; exoticus is from
the Cumberland Plateau immediately to the east of the Bluegrass.
The majority of the inexpectatus group species (umbratilis, parvus,
puteanus) are similarly from the Bluegrass; inexpectatus s. str. is
from the Mammoth Cave Plateau southwest of the Bluegrass; inex-
pectatus orientalis and cnephosus are from the Pennyroyal Plateau
southwest of the Bluegrass. Of the barri group, one species (barri)
is from caves in the glaciated portion of southeastern Indiana in the
same limestone beds as are included in the Bluegrass, while the other
species (troglodytes) is from a cave some 20 miles south in the Blue-
grass of Kentucky. Three rittmani group forms (rittmani, exiguus
s. str., and exiguus furtivus) are from caves in Kentucky on the
Cumberland Plateau immediately east of the Bluegrass, while cato-
ryctos is from the eastern portion of the Bluegrass some 20 miles

82 FIELDIANA: ZOOLOGY, VOLUME 62

distant. An additional species, P. conditus, not assigned to a spe-
cies group, is from the southwestern portion of the Bluegrass.

Forms are described as species which are judged by morphological
criteria as probably not exchanging genes or which are probably
separated from the form most similar to it by a barrier to dispersal.

From the fact that the range of the inexpectatus group forms is
divided by the non-caverniferous strata of Muldraugh's Hill it can
be concluded that dispersal of these forms or their ancestors was over-
land. P. chthonius, krameri, and barri are from caves covered
by the Illinoian glacial advance and they must have entered them
after its withdrawal. These three species, found north of the Ohio
River, are of groups with other species in Kentucky south of that
river. Movement between Kentucky and Indiana-Ohio must have
occurred prior to Kansan glaciation, for at that time the Ohio River
probably became the large stream it now is and likely a barrier to
dispersal. It is postulated that forms of these groups north of the
Ohio River found refuge there beyond the limits of Illinoian glacial
boundaries and after this glacier's withdrawal moved into the caves
they now occupy.

Two and three anophthalmid species of the same or different
species groups coexist in a number of caves. Range extension of
species following allopatric speciation appears to be the best expla-
nation for such sympatry. Differences in habitat preference were
not observed where inexpectatus and horni group forms were sym-
patric. But where chthonius of the horni group and barri of the
barri group, and where rittmani and exiguus of the rittmani group,
were sympatric, definite habitat and/or behavioral differences were
observed which may account for their continued coexistence.

REFERENCES

Barr, Thomas C, Jr.

1959. New cave beetles (Carabidae, Trechini) from Tennessee and Kentucky.
Jour. Tennessee Acad. Sci., 34, pp. 5-30.

1960. A synopsis of the cave beetles of the genus Pseudanophthalmus of the
Mitchell Plain in Southern Indiana (Coleoptera, Carabidae'). Amer. Midi.
Nat., 63, pp. 307-320.

1962a. The robustus group in the genus Pseudanophthalmus (Coleoptera:

Carabidae: Trechini). Coleopt. Bull., 16, pp. 109-118.
1962b. The blind beetles of Mammoth Cave, Kentucky. Amer. Midi. Nat.,

68, pp. 278-284.
1966. Cave Carabidae (Coleoptera) of Mammoth Cave. Psyche, 73, pp.

284-287.

KREKELER: CAVE BEETLES 83

1967a. Cave Carabidae (Coleoptera) of Mammoth Cave, Part II. Psyche,

74, pp. 24-26.
1967b. Ecological studies in the Mammoth Cave system of Kentucky. I: The

biota. Intern. Jour. Speleol., 3, pp. 147-203, pi. 37-64.

1967c. A new Pseudanophthalmus from an epigaean environment in West
Virginia (Coleoptera: Carabidae). Psyche, 74, pp. 166-172.

1967d. Observations on the ecology of caves. Amer. Nat., 101, pp. 475-491.

Erichson, W. F.

1844. Footnote in article by Th. G. Tellkampf, Ueber dem blinden Fisch der
Mammuthhoehle in Kentucky, mit Bemerkungen ueber einige andere in
dieser Hoehle lebende Thiere. Mullers Arch. Anat. Physiol., 4, pp. 381-394.

G arm an, H.

1892. The origin of the cave fauna of Kentucky, with a description of a new
blind beetle. Science, 20, pp. 240-241.

Horn, G. H.

1868. Catalogue of Coleoptera from South Western Virginia. Trans. Amer.
Entomol. Soc, 2, pp. 123-128.

1883. Synopsis of North American species of Anophthalmus. Trans. Amer.
Entomol. Soc, 10, pp. 270-272.

Jeannel, R.

1928. Monographie des Trechinae, III. Les Trechini cavernicoles. L'Abeille,
Jour. Entomol., 35, pp. 1-808.

1931. Revision des Trechinae de l'Amerique du Nord. Arch. Zool. Exptl.,
71, pp. 403-500.

1949. Les Coleopteres cavernicoles de la region des Appalaches. III. Etude
systematique. Notes Biospeologiques, Publ. du Mus. d'Hist. Nat., fasc.IV,
no. 12, pp. 37-104.

Krekeler, Carl H.

1958. Speciation in cave beetles of the genus Pseudanophthalmus (Coleoptera,
Carabidae). Amer. Midi. Nat., 59, pp. 167-189.

1959. Dispersal of cavernicolous beetles. Syst. Zool., 8, pp. 119-130.

Von Motschoulsky, T. Victor

1862. Etudes entomologiques, 11, pp. 40-42.

Thornbury, William D.

1965. Regional geomorphology of the United States. Wiley, New York.
609 pp.

Valentine, J. Manson

1932. A classification of the genus Pseudanophthalmus Jeannel with descrip-
tions of new species and notes on distribution. J. Elisha Mitchell Sci. Soc,
47, pp. 261-280.

1945. Speciation and raciation in Pseudanophthalmus (cavernicolous Cara-
bidae). Trans. Conn. Acad. Arts Sci., 36, pp. 631-672.

UNIVERSITY OF ILLINOIS-URBANA

590.5FI C001

FIELDIANA, Z0OLOGY$CHG0
62:1-51972

3 0112 009379873