•

'••'

.-:•

-

'

PLATE I.

MOUTH OF MAYFIELD'S CAVE.

MOUTH OF MAYFIELD'S CAVE.

Looking in the general direction in which the cave extends. At the lower right is shown
a portion of the shallow ravine which formerly was a part of the cave.

HELIOTYPE CO., BOSTON.

THE FAUNA OF MAYFIELD'S CAVE.

BY

ARTHUR M. BANTA.

WASHINGTON, D. C. :

Published by the Carnegie Institution of Washington
September, 1907

CARNEGIE INSTITUTION OF WASHINGTON
PUBLICATION No. 67

•

C'ORNMAN PRINTING CO.,
CARLISLK, PA.

CONTENTS.

Page

Introduction 5

The cave as a unit of environment 8

Location 8

General description 8

Moisture 11

Light 12

Temperature 12

Air-currents 15

Origin of food 15

Optimum conditions for cave life.... 16
Fauna:
Mammalia—

Vespertilionidae 17

Muridae 19

Procyonidae 20

Sciuridae 21

Amphibia—

Plethedontidfe 22

Ranidae 22

Pisces—

Amblyopsidae 23

Insecta—

Hymenoptera 25

Pemphredonidae 25

Alysidaa 25

Coleoptera 26

Carabidae 26

Silphidae 29

Staphylinidae 29

Tenebrionidae 33

Aphaniptera 33

Diptera 34

Tipulidag 34

Psychodidaa 34

Chironomidae 35

Culicidae 35

Mycetophilidse 36

CecidomyidsB 39

Dolichopodidae 39

Phoridse 39

Muscidae 40

Anthomyidae 41

Helomy zidae 41

Borboridse 45

Lepidoptera 47

Noctuidae .. ..... 47

Page

Fauna — Continued.
Insecta — Continued.

Orthoptera 51

Locustidae 51

Aptera 53

Lepismidas 53

Entomobryidae 54

Myriapoda —

Chilognatha 56

Blan julidae 56

Chordeumidae 56

Polydesmidae 57

Chilopoda 58

Scolopendridae 58

Geophilidae 58

Arachnida —

Araneida 59

Dictynidse 59

Agalenidae 59

Theridiidaa 60

Epeiridae 66

Tetragnathidae 67

Lycosidae 67

Acarina—

Eupodidae 68

Tyroglyphidae 68

Gamasidae 69

Crustacea :
Podopthalmata —

Astacidae 69

Isopoda 76

Asellidas 76

Oniscidae 77

Amphipoda 78

Gammaridae 78

Copepoda 81

Cyclopidas 81

Annellida 81

Mfcllusca 81

Platyhelminthes

Turbellaria 82

List of species 82

General considerations 89

Origin of cave life 95

Summary 106

Bibliography 109

THE FAUNA OF MAYFIELD'S CAVE.

INTRODUCTION.

This paper represents the results of an attempt at a detailed study
of the fauna of a small cave, observing its animal inhabitants through-
out the year and in all conditions to which they are normally subjected.

Mayfield's Cave was selected because of its accessibility, it being
but 4.5 miles from the laboratory of Indiana University. The cave was
visited on an average about once a week from January, 1903, to June,
1904, and somewhat less often from September, 1904, to August, 1905.

To facilitate study of the cave and ready labeling of the collections,
the cave was divided into parts, and these are referred to upon the map
(plate 2) and throughout the text of this paper by number. In col-
lecting, a carbide bicycle lamp was used and found to be very effective,
as it was easily handled and reflected a strong, steady light. Ordinarily
about three hours were spent in the cave on each trip, but often the
whole day^was taken and five to seven hours spent in the cave. Occa-
sionally bits of beef and cheese were distributed at various places in
the cave as bait to attract cave animals. It was necessary to place this
bait under small stones to prevent mammals carrying it away.

The temperature and air-currents were observed. Collections were
made, habits of the various species noted, life histories worked out as
far as possible— in fact, everything throwing light upon any phase of
cave life was sought. Several times during winter collections were
made under leaves, logs, and stones, and about sink-holes in the vicinity
of the cave. These were made with a view to determine what species
living in the cave also live outside. Any species which lives in the cave
and also outside the cave would likely be found under logs and stones
or in the ground during the winter, while the many species which do
not live through the winter are eliminated from the collections made
outside caves at this season.

From a biological standpoint Mayfield's Cave has been mentioned in
the literature a number of times. McNeill (1887, 330) described the
myriapod Conotyla bollmani taken from it and gave a short paragraph
in description of the cave. Packard (1888, 16) mentions the cave as
having been visited by Bollman and lists the species Bollman found-
Cambarus pellucidus (Tellkampf ) , Csecidotea stygia Packard, Crangonyx

6 FAUNA OF MAYFIELD'S CAVE.

sp., and a Machilis. Bollman (1888, 405) mentions the occurrence of
Conotyla bollmani McNeill within the cave, and Faxon (1889, 621) notes
the occurrence of Cambarus pellucidus (Tellkampf ) . W. P. Hay visited
the cave in 1891-92 and described a new subspecies, Cambarus pellucidus
testii, from it (1893, 283). Blatchley (1896, 127-128), during his explor-
ations in 1896, visited the cave, wrote a short description of it, and
mentioned the occurrence of 11 species within it, namely, Quedius spelxus
Horn, Anopthalmus tennis Horn, Leria (Blepharoptera) latens (Aldrich) ,
Leria (Blepharoptera) specus (Aldrich), Sinella',(Degeeria} cavernarum
(Packard), Conotyla bollmani (McNeill), Spirostrephonlactarium (Say),
Metamenardi (Latreille), Csecidotea stygia (Packard), Crangonyx gracilis
(Smith), and Cambarus pellucidus (Tellkampf).

The bibliography at the end of this paper will give a sufficient index
to the work done heretofore on the cave animals of North America.
A few papers, however, deserve special mention.

Packard's Cave Fauna of North America (Packard, 1888), based
upon material collected and explorations made by himself and others in
Kentucky caves and to some extent in southern Indiana and Virginia
caves, is a most notable and excellent work on American cave animals.
It discusses the life histories and habits of such cave animals as were
known to its author.

Cope paid a two days' visit to Wyandotte Cave in 1872, and his Report
on the Wyandotte Cave and its Fauna (Cope, 1872) is an interesting
account of the fauna of this cave and contains a discussion of cave life
in general.

Blatchley, State geologist of Indiana, with a party of four assistants,
made a five weeks' exploring and collecting trip to nineteen of the most
important caves of Indiana during the summer of 1896. An admirable
account of the work done was published (Blatchley, 1896). His paper
includes a discussion of the formation of caves, short descriptions of
the caves explored, and a list of the animals collected, together with
descriptions of new species and notes on the occurrence, habits, and
interrelations of the various cave animals. These publications* are the
nearest approach to an ecological study of a cave fauna heretofore
attempted. Eigenmann (1897-1905) and others have contributed much
to our knowledge of the structure, habits, and origin of cave animals.

I am indebted to C. H. Eigenmann and W. J. Moenkhaus, of Indiana
University, for encouragement and valuable suggestions on working up
this paper. C. H. Eigenmann personally furnished me with much of

*The following papers ought also to be mentioned in this connection: Call, 1897;
Garman, 1889; and Ulrich, 1901.

INTRODUCTION. 7

the literature which I was able to consult.* He originally proposed the
problem and repeatedly gave excellent and timely suggestions during
the progress of the paper. I was enabled to visit Mammoth and other
caves through grants from the American Association for the Advance-
ment of Science. The following persons also kindly assisted me by
specifically determining many of the animals found and in certain cases
indicating the known habits and distribution of the species: H. F.
Wickham, Iowa City, Iowa, the Coleoptera; D. W. Coquillett, United
States Department of Agriculture, C. F. Adams, Chicago University,
and C. T. Brues, Milwaukee (Aphiochssta) , the Diptera; Nathan Banks,
East End, Virginia, the Arachnida; Miss Mary Murtfeldt, Kirkwood,
Missouri, and H. G. Dyar, U. S. National Museum, the Lepidoptera; J. W.
Folsom, Champaign, Illinois, the Thysanura; R. V. Chamberlain, Salt
Lake City, Utah, the Myriapoda; A. S. Pearse, Chicago, the Copepoda;
and W. P. Hay, Washington, District of Columbia, the Crangonyx gracilis
Smith. Charles Zeleny, of Indiana University, is also credited with en-
couragement and many kindnesses during the last year spent upon this
problem. Mrs. Rosa Smith Eigenmann and later Mrs. Mary Slack Banta
kindly went over much of the text of the paper with me and made
valued suggestions on the form. C. F. Adams, of Chicago University,
has described the two species of Diptera. J. W. Beede, of Indiana
University, kindly supplied the contours for'the cave map, and E. R.
Cumings the photographs of the mouth of the cave. W. L. McAtee
and W. L. Hahn, of Washington, District of Columbia, assisted in
naming the mammals.

* After this paper was in manuscript I was enabled to consult much additional
literature on cave life at the library of the Museum of Comparative Zoology at Cam-
bridge, Massachusetts, but the pressure of other work prevented as full use of it as
was desirable.

8 FAUNA OF MAYFIELD'S CAVE.

THE CAVE AS A UNIT OF ENVIRONMENT.
LOCATION.

Mayfield's Cave is 4.5 miles northwest of Bloomington, in the south
half of section 25, township 9 north, range 2 west, Richland Township,
Monroe County, Indiana. It is in the Mitchell limestone of the upper
half of the Subcarboniferous. It is in latitude about 39.5° north. The
mean annual temperature of the region is about 53.5° F. (11.9° C.),*
and the mean annual precipitation 40. 05 inches.

The mouth of the cave is squarely in the face of a low bluff of lime-
stone, 12 feet high, which forms the head of a winding valley, so
narrow as almost to be called a ravine. This valley, once a part of
the cave, is the course of the ancient stream which ran through the cave.
The cave, which was formerly much longer than now, has been shortened
by the tumbling in of the roof near the mouth, the debris having been
carried away by the agency of water. The valley, during times of
heavy rains, still serves as an outlet for the water which floods through
the cave. The cave is a little more than one-fourth mile long
and varies from 6 to 20 feet in width and 5 to 12 feet in height to
passages so small one can scarcely crawl through them. The cave
stream still exists, but it no longer flows through the whole cave, having
found a new outlet into the head of a little valley to the east of the
mouth, t where it gives rise to a spring. From this spring flows an
above-ground stream. This outlet for the cave stream is relatively
recent, but it has existed long enough for the stream to have formed
quite a spur from the valley into which it originally made its way. A
ridge extends between the valley at the mouth of the cave and the
valley through which the present stream finds its outlet. To the south
rises a hill perhaps 200 feet in height, under the edge of which the inner
portion of the cave extends. The accompanying map (plate 2) was
made after carefully running a line through the cave, and is about
correct as to the course of the cave, except possibly the "cork-screw"
passage.

GENERAL DESCRIPTION.

The mouth of the cave is boarded up, and entrance is gained through
a small door. The cave was formerly used for storing fruit and vege-
tables during winter. The cavern leads into the hill on a level with
the ravine as a straight passage 14 feet wide by 12 feet high. The

*U.S. Weather Bureau statistics for six years give an average of 54.4° F. for eleven
months, April not included.

tSee map of cave and topographic map of surrounding region (plate 2).

PLATE 2.

Scale of feet

100 200 300

Sink hole
Cave stream

MAP OF MAYFIELD'S CAVE.

a, mouth; b, mammal hole; c, spring where the cave stream appears above ground; d, cave-
spring disappears in the cave; e, first large pool;/, "man hole;" g, "cork-screw room,"
h, "cork-screw." The contours represent 10-foot elevations, and the figures indicate the
elevations above that of mouth of cave. The numbers in quotation marks designate
different parts of the cave. The scale of feet applies to distances in the cave. The width
of the cave is everywhere exaggerated to 3 or 4 times the scale given with the map.

DESCRIPTION OF CAVE. 9

walls present considerable irregularities, with here a prominence, there
a depression, and everywhere small cracks and projections formed by
solution. The roof is a flat surface of a hard stratum of limestone.
This same stratum forms the roof of the cave to the end of the main
passage, with comparatively few breaks anywhere. This first portion
has the general character of much of the cave. At other places there
are sometimes fewer and sometimes more irregularities on the walls,
while there is now and then a small dome or widened crack in the roof.
Much of the floor is strewn with bits and slabs of limestone, with occa-
sionally a huge block fallen from the roof or side of the cave. The
references in the following pages are to corresponding references in
plate 2.

At 50 feet from the mouth, ' ' 3, " the cave bends to the south at nearly
a right angle, and 70 feet farther in, " 6, " it bends toward the east at
almost as great an angle. At "5" there is a narrow, low side passage
which runs back 30 feet and intersects another similar passage from
"6." In the earth at the intersection of these low passages is a large
mammal hole. To the left, at "11" is another side passage that more
or less directly communicates with a sink-hole, through which water
enters the cave after every rain and spreads about, so that the part of
the cave from "17" to the mouth is always pretty wet. At "10" sev-
eral large blocks of stone fallen from the roof have formed a natural
dam and collected a drift of bits of wood and plant stems, while from
this drift to "16" the bottom of the cave is covered with a considerable
deposit of soil. At "20" there is a slight elevation of the floor, which
is water- worn and strewn with slabs of stone. From "19" to "22" the
cave is usually quite dry, this being its driest portion. At "25" is an
area, probably 30 feet in length, which is kept moist by water dripping
from the roof. Following this is another dry portion, and then, at " 26, "
an area of about 80 feet kept quite moist by dripping water. Beyond
this is another dry portion which extends to the mound, "29."

The whole cave from the mouth to the mound is a fairly uniform
passage varying from 9 to 12 feet in width and 5 to 10 feet in height.
There are deposits of soil on the floor from the mouth to the second
turn ("1" to "7") and from "10" to "16." There is a bank of earth at
one side at "22," and a deposit of gravelly soil from "23" to the mound
("29"). The remainder of the floor of this part of the cave is naked
limestone, strewn with slabs and broken bits of stone.

At "27" is a side passage which winds to the east for about 50 feet
and then leads down 8 to 10 feet to a small stream of water which dis-
appears to the left. The stream has its course in a general direction
parallel to the main cave and flows toward the mouth. Here, for the

10 FAUNA OF MAYFIELD'S CAVE.

first time, we have encountered the cave stream. Less than 20 yards
from where it disappears in the cave the stream appears above ground
as a spring. The roof of this passage is a continuation of the same
stratum that forms the roof of the main cave. This branch of the
cave has been formed by the cave stream and is 3 or 4 feet wide in
parts. At places it is 15 feet high, but at other portions the stream
has worked under, leaving large masses of soil and stone above its course
over which one must crawl. After 200 feet the passage becomes too
small to follow farther. The bottom of this branch is on a lower level
than that of any other part of the cave.

From the mound to "36" the main cave is quite irregular. At "34"
the stream is found for the first time in the main cave. It here becomes
lost under the wall at the side of the passage. The continuation of this
stream is encountered in the side passage at " 27. " From ' '34' ' to ' '37' '
the stream winds along through a narrow channel 2 to 4 feet deep and
a foot wide, dissolved out of the limestone of the floor. From "37"
inward to the end of the main cave the passage is generally wide and
low, being 12 to 20 feet by 3.5 to 5 feet, with here and there small pools
connected by the stream which winds back and forth across the passage
and in places disappears at the side of the cave. At "42" the main
passage terminates; the stream comes out from under the wall. How-
ever, a slide through a small hole brings one to another part of the cave,
a long, narrow room ("43" and "44"), through which the stream runs
and which is much like a portion of the main cave. Another crawl
brings one to a room 40 feet by 25 feet by 9 feet high, from the
southeast corner of which the cave continues as a "cork-screw" passage,
a'narrow and very crooked water-worn cavern in a sort of shaly lime-
stone. It is on a higher level than the rest of the cave and appears to
be above the stratum of hard limestone which forms the roof of the
main cave. From this passage comes the stream of water so often
mentioned. The beginning of this passage is about 2. 5 feet wide by 4 feet
high. It bends back and forth many times in_ a most bewildering
manner, leading in a general southeasterly direction. It becomes grad-
ually smaller and after about 300 feet becomes too small for further
exploration.

In many places along the sides of the cave there are deposits of soil
which extend almost to the roof. The so-called "mound" is a place in
the cave where this earth fills the whole passage to within 18 inches of
the roof. The mound and these banks of earth indicate that the cave
was obstructed after having become near its present size. This obstruc-
tion was probably a large quantity of limestone which had tumbled in
from the roof near " 22. " At any rate, there was sufficient obstruction

MOISTURE. 11

to cause the cave from "22" to the cork-screw room to become filled
with mud to near the roof. The obstruction having been removed
later, the water cleared out the main passage, except at the mound,
where the stream made its way underneath the deposit. The stream,
in cutting through the earth in the cave, formed a winding course and
left the banks of soil still remaining at the sides of the main passage.

MOISTURE.

The cave stream runs only during winter, spring, and part of the
summer, or after heavy rains at any season. It is much influenced by
weather conditions, becoming swollen and clouded after heavy rains or
thaws. In late spring the stream becomes gradually smaller, until by
June it ceases to flow. The pools of the stream then become separated
from one another, but there is always sufficient water to keep them
brimming full. There are several of these pools, the largest of which
is about 15 inches in depth and covers an area 20 by 8 feet. Most of
them are much smaller, however. Usually one or both edges of the
pools extend under projecting shelves of rock at the sides of the wall.
The cave stream enters the cave from the "cork-screw" passage and
follows the main cave to ' ' 33, " where it becomes lost under the wall. It
has a winding course, wandering from side to side in the cave and often
disappearing under the wall and reappearing within a few feet.

The amount of moisture in the different parts of the cave is variable.
The part from the mouth to "17" is rather moist. From "17" to "24"
it is drier and from "21" to "24" very dry for a cave. From "24" to
"26" is a damp portion again, within which are two places where water
drops from the roof at all seasons. Beyond "26" to the mound, "29,"
it is less moist. From the mound to ' '32" it is very moist, water coming
through the roof in places. From "32" to "34" there is less moisture
again, and from ' '34' ' on the cave is everywhere quite damp. There is
considerable seasonal variation in moisture in the part of the cave from
the mouth to "34." From the mouth to "12" the cave becomes wet
after every rain by water entering through sink-holes, and if the rain
is heavy the water spreads over the floor to "17." The cave from the
mound out is very rarely flooded. Only once have I known water to
run through this part, and this was during the unusually high waters in
April, 1903. Here the cave becomes very dry, except in places where
the water seeps through the roof. After each heavy thaw or heavy
rain the amount of water coming through the roof is considerably
increased, and wet spots appear at places where during dry weather
there is little moisture.

12

FAUNA OF MAYFIELD'S CAVE.

LIGHT.

The partition at the entrance, although rather open, shuts out much
of the light from the first part of the cave and greatly increases the twi-
light, causing the stretch of the cave from the entrance to the first turn
to have a more uniform and diffused light than it would otherwise have.

The two sharp turns in the course of the cave near the entrance,
one 50 feet in and the other 70 feet farther in, regulate the rays of light
from the outside in such a way as to produce a definite area where
there are direct rays from the outside and also a definite area where
there are only reflected rays directed inward from the wall at the first
bend. Beyond the second turn one's eye, even when directed toward
the dimly lighted portion, can not discern the faintest light. This
angular course of the cave produces fairly well defined areas of strong
twilight, dim twilight, and absolute darkness, the areas of course merging
into one another to some extent.

TEMPERATURE.

The average temperature of the cave in its remote parts is the mean
annual temperature of the region. The average of temperatures taken
at "42" is 11.52° C., but during the months of July and August no
temperatures were taken here. The average for the whole year at
"42" would doubtless run very near 11.9° C., the mean annual temper-
ature of Bloomington.

Temperatures observed at Mayfield's Cave.

[Temperatures are given in degrees centigrade. The figures given at the heads of the columns
refer to points indicated by corresponding figures in plate 2.]

When tem-
peratures
were taken.

Temperature at —

Average at Bloom-
ington for preced-
ing week.

Average at Bloom-
ington for days on
which cave temper-
atures were taken.

Out
side*

iij»

"3"

"6"

"9"

"ii"

"17"

"16"

"22"

"25"

"29"

"37"

"40"

"42"

Sept.24, 1904
Oct. 15, 1004
Oct. 26, 1904
Nov.ll, 1904
Nov.21, 1903
Dec. 9, l'J04
Dec. 21, 1904
Jan. 19, 1U05
Feb. 9, 1904
Feb. 17. 1905
Feb. 19, 1904
Mar. 4, 1905
Apr. n, I'.mr,
Apr. 29, 1905
May 16. 1905
June 9, 1905
July 19, 1905

16
15
9.3
8
5
4.7
-2.5
4.4
—2.7

i2-2

-1.8
16.5
11.8
15
23
32

12.4
"8.7

11.9
11.6
9.9
87
5.9
7.4
4 3

11.8.

11.7

16 '.4'

11.6
11.5
11

9 2

11.5

11.5
11.6
11.4

11.1
11.5
11.6
11.4

11.1
11.4
11.4

11 4

11.3
11.5
11.25

11.6
11.6
11.6
11.6
11.6
11.45
11.25
11.5
10.8?
11.2
10.4
11
11.7
11.7
12
12.1

18.8
15.5
8.3
8.9
(?)
0.2
— 3.7
— 5.4
— 4.4
—11.6
— 6.55
4.5
12.9
14.5
20.1
21.5
24.3

20.6
12
11.5
3.9

(?)
1.4
— 2.8
— 0.2
- 4.17
— 6.4
- 6. 66
4.4
5.3
16.4
16.4
19.4
28.3

4.4

6.95

7.2.3
8.1

8.4
8.4
7 3

9.15

10.1

8.85

11
8.8
9

8.7'
'8.4

11. 2»
10.5
9.95
9.8
9.4
8.55
9.3
8 9
9.3

11.9
11
10 6
10.5
10.25
9.6

i6'.35
11.2
11.4

11.58
11.15

'9.6

4.9
0.6
1.8
-2.5

— i'.s

— L4

3.9
1.0
1.4
2.2

4 8

;;:;;

'21.5"
'S.Y

6.3
4.3
3.5
4.9

'5.3

"e"

7.4
8.3

9
9.8
11.05

9
9.65
10.75

... .

io. i'

10.5
10.6
10.6

11.7
11.6
11.17

10.6
12.2

9.8

»At mouth of cave. ^Passage off " 27," 11.4°.

TEMPERATURE.

13

The seasonal variation in temperature in the remote parts is less
than 2° C., the lowest temperature found at "42" being 10.4° C., the
highest 12.1° C. The variation is greatest near the mouth of the cave.
Ice is rarely found as far from the entrance as "10" in winter, but
there is little variation in the way of increase of temperature in summer.
Even at ' ', 1, " quite near the entrance, the highest recorded summer tem-
perature is 12.2° C. The

Winter temperature from Sept. Oct Nov. Dec. Jan Feb. March Apr. May June

"10" to the mound becomes
considerably lowered, a mark
of 3.5° C. being reached at
"17," and 8.55° C. at the
mound during February.
Beyond the mound there is
much less variation, a tem-
perature below 10° C. being
rarely attained. These ex-
tremes of low temperature
in the inner part of the cave
are very temporary. They
are produced only by con-
tinued severe cold and are
not maintained after the
severe weather is over.

Reference to the accom-
panying table of tempera-
tures shows that the fluctua-
tion in the cave tempera-
ture, particularly during the
winter, is not only seasonal,
but that it occurs also with
varying meteorological con-
ditions, persistently cold or
continued mild weather pro-
ducing noticeable changes
some distance within the
cave. The temperatures are
given in degrees centigrade.

The average monthly temperatures for Bloomington for the past
six years, as given by the United States Weather Bureau, are: January,
0.8° C.; February, 1.3°; March, 5.3°; April (no record); May, 18°;

-4

-12

FIG. 1. — Diagram showing average temperatures in
Mayfleld's Cave and at Bloomingtou.

14

FAUNA OF MAYFIELD'S CAVE.

42 40

a v\ fi v\
37 29

ft Yl

19

17

12'

-4'

-12°

June, 22.9°; July, 25.2°; August, 24.4°; September, 20.4°; October,
15.5°; November, 7. 8°; December, 0.5°.

In fig. 1 are graphs showing the following temperatures: (I) at
" 42," taken once for each month from September to June; (II) the
average temperature at Bloomington for the week preceding the tak-
ing of each cave temperature; (III) the average temperature for each

month from September
to June. This figure
shows at a glance the
relative constancy of
the cave temperatures.
Perhaps this is still
better shown by fig. 2,
which represents maxi-
mum and minimum
temperatures through-
out the cave and out
side. Graph I repre-
sents maximum sum-
mer and Graph II
minimum winter tem-
peratures.

Consideration of fig.
2 discloses another in-
teresting thing. Where-
as during winter the
minimum temperature
from "25" to the mouth
is much below 11.6° C.,
which may be consid-
ered the normal tem-
perature for the cave,
during summer the
maximum for the same
part of the cave is very
little above this aver-
age. This peculiarity

PIG. 2.— Diagram showing maximum and minimum tempera-
tures throughout the cave and at the mouth outside (*).

is explained in connection with the discussion of the air-currents within
the cave.

There is little fluctuation in temperature in the side passage at ' ' 26, "
where the air-currents are not effective.

ORIGIN OF FOOD. 15

AIR-CURRENTS.

There is always a current of air in the cave. The current is variable
in strength, but always discernible. The air flows in at the mouth of
the cave and out through the cork-screw passage from October to March
and in the reverse direction from April to September. In the cork-
screw passage the current is very noticeable, as it is in all the narrower
parts of the cave. During February, 1904, I twice thought I detected
the air moving in part of the time and out part of the time during the
same day. But I noticed the apparent outward movement only when
in a larger portion of the cave, where the current is difficult to determine,
and was unable to be sure this fluctuation existed.*

The cause of the currents is evident. During winter the cooler
outdoor air flows into the cave and displaces the warmer and lighter air,
which flows upward and outward through the cork-screw passage.
In summer the cooler air of the cave flows out at the mouth and the
warmer air enters the cave from above through the cork-screw.

The air-currents tend to equalize the temperature within the different
parts of the cave and to bring it on a par with the temperature outside,
but the temperature of the surrounding earth exerts a sufficient influ-
ence to counteract largely the effects of air-currents, as well as of
the temperature outside the cave. The cool air flowing into the cave
in winter produces the decided change in temperature noted from
the entrance to the mound, radiation from the rock of the cave rapidly
warming the current all along, so that the influence is much reduced
toward the mound and is little felt at "42." When the current is in
the other direction, from April to September, the cave warms up in the
reverse order, so that the remotest portion is most quickly affected by
the current, while the portion from "17" to "6" does not reach its
normal temperature until midsummer. The constant outflowing of the
current during the warm seasons keeps the temperature near the mouth
very near the mean cave temperature, radiation from the outside having
little influence.

ORIGIN OF FOOD.

For food, cave animals are directly or indirectly dependent upon
organic matter carried into the cave from the outside, and they there-
fore lead a precarious existence. The supply of food is due to chance
and accident and is irregular in the extreme.

*In Mammoth Cave, which apparently has but one opening to the exterior, the
air-currents, which are very strong, are variable, flowing sometimes in one direction
and sometimes in another, depending, it would seem, upon the temperature somewhat
and certainly upon barometric conditions. Should the outside opening into the cork-
screw passage of Mayfield's Cave become stopped with snow, fluctuating currents
might be expected in Mayfield's Cave.

16 FAUNA OF MAYFIELD'S CAVE.

There are various sources of organic matter in the cave. Mammals
leave excrement and hair and sometimes carry seeds and pieces of trash
into the cave; water sweeping into the cave through sink-holes brings
protozoans and algse, seeds, stray insects, fish-worms, pieces of wood,
and bits of plant stems; and human visitors leave drops of tallow, bits
of string, paper, etc. The fungus which grows abundantly upon
decaying organic matter in moist portions of the cave apparently serves
as food, as well as the decaying organic matter itself. I am reason-
ably sure the fungus serves as food to the cave thysanuran at least.

OPTIMUM CONDITIONS FOR CAVE LIFE.

Cave animals are most abundant under the following three conditions:
(1) A considerable degree of moisture; (2) the presence of decaying
organic matter; and (3) the occurrence of loose stones and other debris
which serve as places of refuge. The drier parts of the cave, from
"21" to about "24," from "26" to "29" and from "33" to "34" are
very poor collecting-ground. Leria defessa and L. latens may be seen
occasionally, but they are about all. A wet portion of the cave or a
moist place caused by water coming through the roof is always favor-
able collecting-ground, and particularly so provided the other favorable
conditions exist.

Most species of the cave are scavengers and are attracted to organic
matter for food. Bait placed in the cave attracts the beetles (except
Anopthalmus) and Aphiochseta, while it brings together Limosina, the
myriapods, and thysanurans in great abundance. Any decaying organic
matter, whether it is a piece of moldy paper or a drop from a candle,
becomes a Mecca for these species. Some of the predaceous species are
attracted to the baits also, Phanetta subterranea and Erigone infernalis,
for example, but possibly they simply follow up the species upon which
they prey.

As will be mentioned again later, many cave species habitually conceal
themselves under something. During the first two trips to Mayfield's
Cave I did not learn to turn over debris to look for cave inhabitants and
the collections made were very small indeed. Learning to search out
the places of concealment, on the third trip I obtained much more
material than on the first two trips combined. In parts of the cave
where there is considerable moisture, where there is an abundance of
decaying organic matter, and where there is debris to serve as places
of concealment, conditions of cave life approach the optimum and the
abundance of life is really surprising.

MAMMALIA: PIPISTRELLUS SUBFLAVUS. 17

FAUNA.

SPECIES OF MAYFIELD'S CAVE.

MAMMALIA.

Family VESPERTILIONIDAE.
Vespertilio fuscus (Beauvois). Brown Bat.

One specimen was found March 17 in a narrow horizontal crack in the
wall. It is hard to see how it could have gotten into such a narrow place.
It would not be pushed out from behind, but when a pencil was held toward
it, it fastened hold of the pencil and was pulled out by its teeth. At
another time one was found on the roof out of reach and knocked down,
when it flew and escaped. December 9, 1904, I found one of this species
in a small, dome-like corner of the wall just past the first turn of the
cave. It appeared somewhat alert when the light was thrown upon it,
but I did not disturb it further. December 21 this bat was at the same
spot, but January 19 it had disappeared. January 28 one of these bats,
perhaps the individual just mentioned, was found a few feet farther in,
flattened out in a narrow crack in the roof.

This species is probably more abundant than would appear, since
when found it has usually been in inconspicuous places. A number of
times, more often during spring and early summer, it was seen flying
about in the cave or back and forth past my light. It has also been
seen flying about in the cave in the spring after the other species have
disappeared.

Myotis lucifugus (F. Cuvier). Little Brown Bat.

Not common. One was taken March 17, 1904, and two in April,
1905, and two or three others were seen at different times.

Pipistrellus subflavus (F. Cuvier). Georgian Bat.

This bat is fairly abundant in the cave during early spring, late
fall, and winter. When fully established for the winter individuals
are to be seen in all parts of the cave. They return early, having been
found as early as September 24 (one) and September 29 (four or
five) , and remain until spring is well advanced. At first they return
in small numbers. By October 30 the number increases considerably.
Few come after this time, and probably none after November 20. They
seemed to have reached the maximum in numbers and to have been
pretty well distributed through the whole cave by November 20.

18 FAUNA OF MAYFIELD'S CAVE.

The individuals become inactive and apparently settled for the winter
almost as soon as they arrive. October 3, 1903, three seemed to have
become established, as they were dull and stupid. I never saw bats
more inactive than these. Two noted October 26 (one at "2" and
another at "33") apparently did not move again all winter. I am sure
they did not change their positions after November 11. However,
after the bats return to the cave there is some fluctuation in numbers
seen and positions occupied until about the first of December, after
which they move very little or not at all. Many, at least in 1903-04,
remained in approximately the same situations throughout the winter.
An individual was observed to remain in an identical spot from November
20 to April 5. Several others were seen so often in the same place that
they became landmarks. During the winter I have never seen one of
this species fly or change its location in the least, except when disturbed
and thoroughly aroused. When disturbed by contact with some object
this bat sometimes moves its wings and opens its mouth in a threatening
manner. If the disturbance is kept up it may be induced to squeal.
If the disturbance is the heat from a lighted candle, the bat pushes
forward its wings and rubs its head with them as if to shield it from
the heat. If left alone then it very promptly dozes off again into its
winter's sleep. On one occasion (February 17, 1905) one at "22" was
pretty thoroughly but gradually warmed. It went through the usual
motions, as if to shield its head from the heat, and opened its mouth in
the usual threatening manner and squeaked its disapproval. An hour
later it had disappeared and could be found neither on the floor nor on
the roof in that part of the cave.

During the latter part of the winter and the early spring the bats
become more 'active and are more easily disturbed. They sometimes
take wing when stimulated by the light alone and some of them can
readily be induced to fly when tormented. During April it was noticed
that at times they appeared more abundant than at others. It may be
that at this season they leave the cave at night and go to other localities,
perhaps again entering the cave in increased numbers to spend the day.
At any rate, during this season they are more abundant in the cave
at some times than at others. Later in the spring and during the
summer they are very seldom seen in the cave.

Bats do not locate in the cave with any reference to the amount of
moisture. They are found in the driest and in the dampest portions.
In the moist parts of the cave globules of water sometimes form on the
fur until the little animals seem dripping wet and look white in the
lamp-light.

MAMMALIA: PEROMYSCUS LEUCOPUS. 19

Two bats were found which had large mites attached to the tender
parts of the head, one having two clinging to its face. The mites were
so full of blood that their distended abdomens appeared red and shining.
A few Aphaniptera have been seen upon bats in the cave. *

Family MURIDAE.
Peromyscus leucopus (Rafinesque). White-footed Mouse.

Fairly abundant in all the main passages. Its distribution seems
influenced by the occurrence of suitable banks of earth or small crevices
in the rocks serving as retreats, rather than proximity to the mouth of
the cave. There were abundant signs of mice about cracks and crevices
in the wall and at all banks of earth where it was not too wet. Mice
are most abundant from near the mouth to the mound, where most
favorable conditions exist, although signs of them have been noticed
along the stream well towards "42." Five specimens were caught one
night in ten cyclone traps. They may easily be trapped in any part of
the main cave.

They feed upon nuts, seeds, and other decaying vegetable matter
washed into the cave, upon the myriapods and insects inhabiting the
cave, and upon any animal matter or refuse left by visitors to the cave.
They live in burrows in the deposits of soil and in cracks in the rocks.
Mice proved quite a nuisance by stealing the beef and cheese left as
bait for insects and other cave animals, often undermining the stone
under which the bait was placed in order to steal it. Once a mouse
trapped in the cave was partly eaten by other mice. On another occa-
sion, a bottle lost in the cave on a former trip was found with the label
off and half eaten. Feces of mice were about the place. Mice dig and
loosen up the soil in places to get at decaying seeds and other organic
matter deposited with the soil. This activity is very noticeable at times.

One of these mice was taken in a cage trap and brought out alive.
Its eyes became sore in two or three days and it soon died. Soreness of
the eyes was noted in a Neatomapennsylvanica, the cave rat of Mammoth
Cave, which seemed about to go blind in a few days after being brought
into the light and kept there. Its eyes, however, got better a few days
later, although they hardly regained their original luster and seemed
not to protrude so much as before.

*The red bat, Lasiurus borealis (Miiller) , is said to occur in the caves in this
region, but none have been recorded, although it is seen quite commonly and often taken
outside of caves. I took a single specimen of the big-eared bat, Corynorhinus
macrotis (Le Conte) , in the Upper Twin Cave at Mitchell in the fall of 1902. This is
somewhat north of the range stated and of any locality given for this species by Miller
in his Revision of the North American Bats of the Family Vespertilionida? (North
American Fauna No. 3, Biological Survey, U. S. Department of Agriculture, 1897).

20 FAUNA OF MAYFIELD'S CAVE,

Blatchley (1896, 179) found that the white-footed mice from Marengo
Cave differ much in appearence from above-ground specimens, having
larger external ears (13 by 11 mm. ) , longer whiskers (38 mm. ) , and
more protruding eyes. The same diif erences are noticed in those from
Mayfield's Cave, as compared with those from above ground in the
same region. The ears are even larger than in those from Marengo
Cave, averaging 15 by 12 mm. The whiskers show greater development
as compared with above-ground forms, and the eyes are very large and
protruding.

There is no mistaking the fact that in the case of this mouse, life in
darkness has accompanied or produced some definite modification.

Family PROCYONIDAE.
Procyon lotor (Linnaeus). Raccoon.

Tracks of the raccoon are quite abundant at all times, reappearing
immediately after a freshet, when all the old tracks have disappeared.
These signs are not confined to any special part of the cave. They are
particularly abundant along the edges of.the stream. There are small,
shallow, bowl-shaped "wallows" at several places, which possibly are
used by the raccoons. These wallows are 3 to 5 feet in width from edge
to edge and are kept worn fairly smooth. I am not sure the raccoon
has anything to do with these ' 'wallows. ' ' Probably the holes were
originally made by bears. No tracks or other signs of raccoons were
noticed in dry sand which was thickly sprinkled and left in one of these
wallows.

In the cave, raccoons probably feed upon the blind crayfish, blind
fish, and other organic matter which happens to fall in their way. Steel
traps were kept set during two months, but no raccoons or other large
mammals were caught. A raccoon or fox or ground-hog was heard
three different times while I was in the cave. On one occasion the
animal was ahead of me in the cave and fled, following me out again to
the mouth. A few days later it was heard at the first small passage to
the right at ' ' 6, " and whined from its burrow at the end of the passage
while I was near. When I went farther in it followed me to the mound,
and was heard ahead of me as I came back, remaining in the hole while
I passed and whining after I reached the mouth.

At places where the soil is banked up against the roof at one side
there are holes which follow natural breaks or arches of the rock above
and extend into the wall, often at a considerable angle. These burrows,
which were probably made by some other mammal, may be inhabited by
raccoons. They may have been made by foxes, but I do not believe
foxes inhabit the cave at present.

MAMMALIA: MARMOTA MONAX. 21

Family SCIURIDAE.
Tamias striatus (Linnaeus). Ground Squirrel.

One was seen inside the cave at "4. " It was picking up seeds which
had entered through sink-holes and drifted toward the mouth of the
cave. Signs of these squirrels were noticed several times, but not
farther in than "4." The occurrence of this mammal in the cave is
incidental, but is in no way surprising. No doubt other species, as well
as this one, occasionally wander in for short distances.

Marmota monax (Linnaeus). Ground-hog.

The occurrence of this mammal in the cave is doubtful. A hole at
"24" at the side of the cave runs upward and possibly connects with a
sink-hole. At one time, when woodcutters were at work in the edge of
the valley near which the cave runs, I heard through this hole a sound like
that of an ax. The sound probably came through the hole, which appar-
ently communicates with the surface. This hole was probably made by a
ground-hog or fox. The hole within the passage to the right at "6"
may have been made by a ground-hog, and possibly, too, the animal
whose calls I heard in the cave was a ground-hog.

Vulpes fulvus (Desmarest) (red fox), Putorius vison (Schreber)
(mink), and Putorius noveboracensis Emrnons (weasel) are doubtless
occasional visitants to the cave. Holes of considerable size in open cracks
in the rock at several places in the cave may have been made by foxes, and
the mammal mentioned above, which followed me in the cave and was
heard several times, may have been a fox. Mink tracks were recognized
in the mud at "41," and other tracks, probably made by a weasel, were
seen at the same place.

The large "wallows" at "21," at "34," and at the beginning of the
passage at "27" are too large to have been made by anything smaller
than a bear, so Ursus americanus Pallas must have formerly frequented
the cave. At "21" and "34" there are claw marks, probably of the
same animal. It is a well-known fact that bears formerly frequented
caves in this region. Blatchley (1896, p. 177) mentions finding bear
wallows and claw-marks in Eller's and Saltpeter caves, Monroe County,
and in Conneley's Cave, Lawrence County. He quotes an interesting
letter from an old resident of Greencastle, who tells of the experiences
of his father and three companions with bears in a cave near Bloom-
ington, probably Eller's Cave.

22 FAUNA OF MAYFIELD'S CAVE.

AMPHIBIA.

Family PLETHEDONTIDAE.
Plethedon cinereus (Green).

A specimen of variety cinereus was taken April 15, 1904, at ' ' 38, "
under a stone, with some trash which had drifted in. The same day
another, of variety dorsalis, was taken crawling on the mud at "29."
The occurrence of these was probably altogether fortuitous. They were
found two weeks after a violent flood which flushed the cave and washed
in drift and mud far in excess of what had been seen in the cave before.
Another specimen of variety cinereus was taken May 17, 1905. It was
well up the side of the wall to the right, just past the first turn. This
individual had without doubt entered the cave voluntarily. The occur-
rence of 5 specimens of variety cinereus and 1 of variety dorsalis on
the wall in the dark part of Donaldson's Cave, where they must volun-
tarily have gone, is evidence that this species is not averse to entering
a cave.

Plethedon glutinosus (Green).

A single specimen was found at the base of the wall at "6" in mid-
summer. It did not seem very active when picked up, but when brought
into the laboratory became very lively. Eigenmann (1901, 189) found
this species in Rock House Cave, Missouri.

Spelerpes maculicaudus (Cope). Cave Salamander.

An account of this species is published in the Proceedings of the
United States National Museum for 1906.*

Family RANIDAE.
Rana clamata Daudin. Green Frog.

The green frog is not uncommon in caves. It has been noted in the
caves at Mitchell a number of times and in Mammoth Cave (Hubbard,
1880, 83). I found 2 in Mayfield's Cave, September 30, 1903. One
was 40 feet from the mouth of the cave and the other at "40," at the
edge of a pool. About the middle of October one was seen under a
shelf of rock opposite "6." It was quite dull and inactive. However,
it seemed quite sensitive to light and attempted to get away from it.
Two weeks later it was at almost the same spot, but November 7 it had
disappeared. The occurrence of frogs in caves is accidental.

*Arthur M. Banta and Waldo L. McAtee. The life history of the cave salamander,
Spelerpes maculicaudus (Cope), Proc. U. S. Nat. Mus., No. 1443, vol. xxx, pp. 67-83,
with plates vm-x.

PISCES: AMBLYOPSIS SPEL^US. 23

PISCES.
Family AMBLYOPSIDAE.

Amblyopsis spelaeus DeKay. Blind Fish.

DEKAY, Nat. Hist. N. Y., pt. I, 1842, 187-188 (Wyandotte Cave, Indiana, and Mam-
month and Emerson's Spring caves in Kentucky). CRAIGIE, Proc. Am. Ac. Nat.
Sci. Phil., I, 1842, 175 (Mammoth Cave). WYMAN, Am. Jour. Sci. and Arts,
xiv, 1843, 94-96 (Kentucky); Ann. and Mag. Nat. Hist., 1843, xn, 298-299.
TELLKAMPF, Miiller's Archiv, IV, 1844, 381-894, taf. 9; N. Y. Jour. Med., v, 1845,
84. THOMPSON, Ann. and Mag. Nat. Hist., xin, 1844, 112. OWEN, Lectures on
Comp. Anat. and Physiol. of Vert., I, 1846, 191, 193, 202, fig. 50. STOKER, Mem.
Amer. Acad., n. s., n, 1846, 436. DALTON, N. Y. Med. Times, II, 354. WYMAN,
Proc. Bost. Soc. Nat. Hist., m, 1850 (1851), 349, 357. AGASSIZ, Am. Jour. Sci.
and Arts, ser. n, xi, 1851, 127 (Mammoth Cave). WYMAN, Proc. Bost. Soc. Nat.
Hist., IV, 1853, 395-397; Miiller's Archiv, 1853, 474-476, 3 figs.; Am. Jour. Sci.
and Arts, xvn, 1854, 258-261, 3 figs.; Proc. Bost. Soc. Nat. Hist., v, 1854, 17-18.
POEY, Mem. de Cuba, II, 1858, 104, pis. ix, XL GIRARD, Proc. Acad. Nat. Sci.
Phila., 1859, 63. WOOD, 111. Nat. Hist., in, 1862, 314, fig. TENNEY, Nat. Hist.,
1865, 344, fig. GUNTHER, Cat., vil, 1868, 2 (localities as above). COPE, Ann. and
Mag. Nat. Hist., vm, 1871, 368 (Wyandotte Cave). PUTNAM, An. Rep. Peab.
Acad. Sci., Salem, for 1871 (1872), 17-19; Am. Nat., VI, 1872, 6-30 (in part),
116, pis. I and II (Mammoth Cave; well in Orange County, Indiana); The Mam-
moth Cave and its inhabitants, Salem, 1872, 29-52 (in part), 56-58. COPE, Am.
Nat., VI, 1872, 406 (Wyandotte Cave); Rep. Ind. Geol. Surv., IV, 1872, 157-182,
fig. in. COLLETT, Rep. Ind. Geol. Surv., v, 1873, 305 (Mitchell Caves) . PUTNAM,
Proc. Bost. Soc. Nat. Hist., xvn, 1874, 222 (Mammoth Cave). JORDAN, Rep.
Ind. Geol. Surv., vi, 1874, 218 (Mammoth Cave). COPE, Rep. Ind. Geol. Surv.,
x, 1878, 483 (Little Wyandotte Cave, Indiana). HUBBARD, Amer. Entom., in,
1880, 38 (Mammoth Cave) . FORBES, Am. Nat. , xvi, 1882, 1-5 (in part) . JORDAN &
GILBERT, Syn. N. A. Fishes, 1883, 324 (subterranean streams of Kentucky
and Indiana). WRIGHT, Am. Nat., xvm, 1884, 272. PACKARD, Mem. Nat. Acad.
Sci., IV, 1888, 10, 14, 15, 90, 106 (Hamer's and Donaldson's caves, Indiana;
"Mammoth, Wyandotte, and Emerson's Spring Cave in a pool communicating with
[Green?] River; caves and wells in Indiana and Kentucky"). HAY, Rep. Ind.
Geol. Surv., xix, 1894, 234. JORDAN & EVERMANN, Fishes of N. Am., 1896, 706
(same localities as Jordan & Gilbert). BLATCHLEY, Rep. Ind. Geol. Surv., xxi,
1896, 183 (Sibert's Well Cave; caves near Mitchell, Indiana; Clifty Caves and
cave near Orleans, Indiana [Dr. John Sloan]). EIGENMANN, Rep. Brit. Assoc.
Adv. Sci., 1897, 685-686; Proc. Ind. Acad. Sci., 1897, 230; the same, 1898, 239-241;
the same, 247-251, 3 figs. EIGENMANN & YODER, Proc. Ind. Acad. Sci., 1898,
239-246, 11 figs.; Roux's Archiv ftir Entwick. der Organ., VIII Band, 4 Heft,
1899, 545-618, Taf eln x-xv, 9 figs. ; Biol. Lectures Marine Biol. Lab. Woods Hole,
8th lect., 1899, 113-126; Pop. Sci. Mo., LVI, 1899, 473-496, figs.; Proc. Ind. Acad.
Sci., 1899, 31-46, 10 figs.; the same, 239; Science, n. s., ix, 1899, 280-282, 3 figs.,
the same, 370; the same, x, 883; Proc. Am. Assoc. Adv. of Sci., 1899, 255-256;
Pop. Sei. Mo., LVII, 1900, 48-58, figs, i-vm; the same, 397-405; the same, 485;
Science, n. s. xn, 1900, 302. RAMSEY, Jour. Comp. Neur., xi, 1901, 40-47, pis.
in and iv. EIGENMANN, Science, n. s., xiv, 1901, 631; the same, xv, 1901, 523-524;
Proc. Ind. Acad. Sci., 1901, 101-105; Mark anniversary volume, 1903, 167-204,
pis. xii and xv. Cox, Rep. U. S. Bureau of Fisheries 1904 (1905), 392-394, cuts
vi-vin, pis. I, in, vi.

24 FAUNA OF MAYFIELD'S CAVE.

This species has been more widely studied than any other cave animal,
perhaps, and certainly more than any other American cave animal.
Its habits, anatomy, food, origin, etc. , are discussed in the articles listed
above. Its natural distribution includes the subterranean streams in
the Mississippi Valley east of the Mississippi River and south of the
East Fork of White River. It was introduced into Mayneld's Cave in
the summer of 1901 by Dr. Eigenmann and is now abundant, breeding
freely.

Fishes of all sizes are abundant in favorable pools where the water
reaches 6 inches or more in depth, but are most abundant in those pools
where there are shelving rocks beneath the surface or where the pools
extend under the edge of the wall. They swim slowly, often near the
surface, and pay little attention to light or sound. But if the water is
considerably disturbed they swim quickly downward and soon find their
way under a shelf of rock or into some other retreat, from which they
do not emerge for some time and then very slowly and cautiously. It
is very significant that during the unusually high water noted in April,
1904, when the cave was almost full of water which washed great
trenches at places, the fishes were not washed out of the cave. After-
wards they were apparently as abundant as before. For a fish as
sluggish as the blind fish it would seem to have involved quite a struggle
to resist the force of the current.

The young are so nearly transparent that they can scarcely be
detected, except by the shadows their almost colorless but opaque bodies
make on the bottom. Young, 8 mm. long, were seen in the shallow
water of one of the larger pools in November, 1905. There seemed to
have been young from at least two females, for a few days later young
of two sizes were seen.

In the adult Amblyopsis the eye can scarcely be located and the
alimentary canal bends forward so that the anus is below and between
the gill slits; while in young 8 mm. long there are conspicuous black
eyes, and the anus is between the ventral and the anal fins, as is usual
among fishes. By March 1 the young were 18 mm. long and had much
of the white color characteristic of the adult. The eyes were less con-
spicuous, but the anus was still behind the ventral fins. I was not able
to keep this series longer. Eigenmann (1900c, 119) finds that "in an
individual 35 mm. long the anus is situated between the origin of the
pectorals; in one 25 mm. long it lies between the pectorals and ventrals;
in the young it lies behind the ventrals."

An Amblyopsis 56 mm. long was examined to determine the food of
this species in Mayfield's Cave. The fish had been cut open and placed
in strong alcohol when caught. The stomach contents consisted of a

HYMENOPTERA: ALYSID^:. 25

quantity of food, more or less digested, in which 3 Cyclops, 2 Csecidotea
stygia, one about 4 mm., the other about 2 mm. long, and parts of other
Csecidotea were recognizable. Material taken from the stomach of an
Amblyopsis from Mitchell contained 12 Crangonyx gracilis and 7 Cseci-
dotea stygia. The amphipods ranged from 8 mm. to less than 4 mm. in
length. The Csecidotea were 6 mm. long or shorter. Eigenmann has
found larger amphipods in the stomach of Amblyopsis from Mitchell.
Putnam (1872, 24) found crayfish in the stomachs of blind fish from
Mammoth Cave, presumably A. speldzus, as he speaks of examining a
number of that species. Sloan and also Eigenmann (1899d, 481) have
known large blind fishes to swallow smaller ones, and Wyman (cf.
Putnam, 1872, 13, pi. i) found a small eyed fish in the stomach of an
Amblyopsis.

Cladocera or Copepoda, if placed in an aquarium with blind fishes,
soon disappear. I have fed Amblyopsis bits of beef, which are some-
times readily taken if moved about in front of or just above the head.
It was necessary for the bait to be kept in motion in order to be seized,
no notice being taken of the meat when not moving. Amblyopsis takes
food with a very quick movement, which suggests that it would have no
difficulty in capturing Crangonyx and Cxcidotea.

The blind fish apparently restricts the numbers of Crangonyx gracilis
and Csscidotea stygia in Mayfield's Cave. In the first and largest pool,
where there are always a number of Amblyopsis, the Crangonyx are
very scarce, and I have seen only a few smaller ones. The Csscidotea
also are much less abundant in the large pool than in the upper pools,
although very abundant in the shallow parts of the stream near by.

INSECTA.

Order HYMENOPTERA.
Family PEMPHREDONIDAE.

Three specimens of two species of wasps which I refer to this family
were taken within the cave. February 27, 1904, just within the cave,
one was found crawling slowly along on the wall. A second was taken
April 15 in about the same locality, and a third April 28, just past the
first turn, where it was rather dark. It seemed to be crawling out of a
crack and was headed toward the light. During March another was
seen on the ledge of rock just outside the mouth of the cave, from
which it apparently had come. Some of these wasps may spend the
winter in the cracks and recesses just within the cave.

Family ALYSIDAE.

A small hymenopterous insect of this family was taken on the wall
at "12" in November, 1903. It was without question a stray.

26 FAUNA OF MAYFIELD'S CAVE.

Order COLEOPTERA.
Family CARABIDAE.

Anopthalmus tenuis Horn.

HORN, Trans. Am. Ent. Soc., 1871, 327 (Wyandotte Cave). COPE, Rep. Ind. Geol.
Surv., iv, 1872, 177. PACKARD, 5th Ann. Rep. Peab. Acad. Sci., 1872, 93 (cave
at Orleans, Indiana) ; Am. Nat., x, 1876, 283-287 (Bradford and Wyandotte caves).
HUBBARD, Proc. Ent. Soc. Wash., I, 1886 (Luray Cave). PACKARD, Mem. Nat.
Acad. Sci., iv, 1888, 15, 18, 80, 84 (Wyandotte, Bradford, and Luray caves).
WICKMAN & BLATCHLEY, Rep. Ind. Geol. Surv., xxi, 1896, 193 (Wyandotte,
Sibert's Well Cave, Mayfield's, Truett's, andShiloh caves, in Indiana).

Fairly common. This beetle is the only blind one found in the cave.
It is suited to life in absolute darkness and to a region of nearly constant
temperature, for it has not been observed nearer than 400 feet from the
mouth. It is most abundant in the remote part of the cave, where it is
seen in damp places, usually crawling over mud or stones. It has been
found but twice under stones or drift, situations where the other cave
Coleoptera are most abundant. Bait of beef and cheese left under
stones at various places to attract insects did not seem to attract this
beetle. Only one was found near any bait. The fact that this species
is predaceous, while the others are not, accounts for this difference in
habit. This beetle is negatively phototropic, and if kept in a jar in a
partly darkened room seeks the darkest part of the jar.

It is a carnivorous species and presumably feeds (cf. Packard, 1876,
283) upon the Sinella cavernarum, an occasional spider or mite, and
possibly a few small rnyriapods or dipterous larva?. I have never seen
it feed, nor found its eggs or larvae to recognize them. The species is
not very abundant, however.

There is some variation in color. A very light colored individual was
taken at "42" in November, 1904. Instead of being a uniform rufous-
brown, the rufous was confined principally to the head and mandibles,
to the antennaa and prothorax, which were lighter than the head, and
to slight traces on the legs and elytra. Its light color seemed not to be
due to its being newly emerged, for it was kept alive for ten days and
took on no more of the dark color. Another of a series of 10 from
Mayfield's was considerably lighter than the average, while of the other
8, 3 were slightly darker than the average.

A series of 17 specimens from the Mitchell caves showed consider-
able variation in this respect. Two were quite dark rufous-brown;
13 were near the average in color, there being so little difference that
their positions in a color series could hardly be determined; 2 others
were very light, one having only traces of the brown, the other

COLEOPTERA— CARABID^E. 27

being much lighter than the average and about the color of the light
individual taken in Mayfield's.

The individuals from Mayfield's Cave range from 4.4 mm. to 5.7 mm.
in length, averaging 5.09 mm. Those from Truett's Cave average
5.44 mm. in length, ranging from 4.75 mm. to 6.05 mm. Packard
(1888, 80) found 18 specimens from Bradford Cave to range from 4. 1
to 5. 1 mm. in length, with the tendency toward smaller ones in smaller
"more changeable" caves.

This is the common Anopthalmiis of Indiana caves and the only blind
beetle found in Mayfield's Cave. Taken in Mayfield's, Truett's, and
Twin caves.

This genus is widely distributed in caves. Hamann (1896, 46) con-
siders it but a subgenus of Trechus. He names 51 species of the sub-
genus, principally cavernicolous, and 8 species of another subgenus of
Trechus, all of which are exclusively cavernicolous. The genus Trechus
itself has 5 European species occasionally found in caves, and others of
this genus habitually live under great stones often deeply embedded in
the earth. These and other lapidicolous Carabida?, many of them blind,
are of extremely minute size and of most sluggish habits. The genus
Trechus is found in North America, where it is represented by 4 species
living in retirement as the European species do.

There are 8 American species of Anopthalmus, all but one confined to
caves. Six are recorded from Kentucky caves. One of these A. pusio,
is also found in Virginia. Another, A. eremita, has been described
from a single specimen from Wyandotte Cave and has not since been
taken. Anopthalmus tennis has been taken only in the caves of Indiana
and in Luray Cave in Virginia.

Patrobus longicornis Say.

Trans. Am. Phil. Soc., n. s., vol. n, No. 1, 1823; Ent. Works, Le Conte ed., vol. n, 467
and 534.

A specimen of this species was found in the dirt under a stone at
"4" in May. Its occurrence in the cave was accidental.

It is an apterous species which lives in moist places under stones.

Platynus cincticollis Say.

SAY, Trans. Am. Phil. Soc., n. s., n, 52; Ent. Works, Le Conte ed., vol. n, 476.
LE CONTE, Proc. Acad. Nat. Sci. Phila., 1854, 43; Bull. Brooklyn Ent. Soc., n, 46
and 55. WICKHAM & BLATCHLEY, Rep. Ind. Geol. Sur., xxi, 1896, 194 (Porter's
Cave).

Only 2 specimens of this species were found, one at "37" on a bank
of earth, and another on the wall past the first turn.

28 FAUNA OF MAYFIELD'S CAVE.

Blatchley (1896, 194) found 5 specimens in Porter's Cave. The
species inhabits Canada and the northeastern United States, extending
south through the Middle States.*

Platynus tenuicollis Le Conte.

One specimen was obtained from Twin Cave, 1 from Donaldson's
Cave, 2 from Truett's Cave, and 3 from Mayfield's Cave. Two of those
from Mayfield's Cave and one from Donaldson's Cave were taken near
the mouth, while those in Truett's Cave and one in Mayfield's were far
back in absolute darkness. They were found under stones. Outside
of caves this species lives under stones along streams in deep ravines.

Platynus punctiformis Say.
Ent. Works, Le Conte ed., vol. II, 481.

A single specimen was found in Mayfield's at "41" two weeks after
a freshet. Doubtless the occurrence of this beetle was altogether
fortuitous.

Cope took a species of Platynus ' ' in the outer part of a branch of
Wyandotte Cave," and is quoted as authority for Platynus marginatus
having been taken in either Hamer's, Connelley's, or Donaldson's Cave
(Collett, 1873, 305). Packard (1873, 93) records P. marginatus from a
cave near Orleans, Indiana.

The occurrence of 4 species of a single genus of Carabidse in caves,
3 of which can scarcely be considered accidental, is very interesting as
illustrating the tendency of certain fitted groups to frequent caves and
take up residence therein. Six specimens of a closely related genus
(Pristonychus) are found in European caves, t

Of the family Carabida9, there are 12,000 or 13,000 species known.
Nearly all are terrestrial and there are many with rudimentary wings.
The following is taken from Sharp (Cambridge Natural History, vol.
VI, pp. 205 and 206) :

There are a considerable number of blind members of this family. Some of them
live in caverns. These belong chiefly to the genus Anopthalmus, species of which have
been detected in the caves of the Pyrenees of Austria and of North America. It has
been shown that the optic nerves and lobes, as well as the external organs of vision,
are entirely wanting in some of these cave Carabidse; the tactile seise have, however,

*H. F. Wickham, Iowa City, Iowa.

tit might be in place here to mention the finding:of Atranus pubescens Dejean in
Truett's Cave. Specimens were taken in a room 60 feet from the mouth. Of course
they may have been washed in, but 3 of them were found, while a single specimen of
Platynus tenuicollis Le Conte was the only other outside form seen, so their occurrence
seems scarcely accidental. If these had been washed in other species should have met
the same fate. This species is widely distributed in the eastern United States and
Canada.

COLEOPTERA— STAPHYLINIDvE. 29

a larger development than usual, and the insects are as skillful in running as if they
possessed eyes. Anopthalmus is closely related to our British genus Trechus, the
species of which are very much given to living in deep crevices in the earth, or under
large stones, and have some of them very small eyes. In addition to the cavernicolous
Anopthalmus, other blind Carabidaj have been discovered during recent years in various
parts of the world, where they live under great stones deeply embedded in the earth;
these blind lapidicolous Carabidss are of extremely minute size and most sluggish
habits; the situations in which they are found suggest that many successive genera-
tions are probably passed under the same stone. Not a single specimen has ever been
found above ground. The minute Carabids of the genus Aepus, that pass a large
part of their lives under stones below high-water mark (emerging only when the tide
uncovers them) , on the borders of the English Channel and elsewhere, are very closely
allied to these blind insects, and have themselves only very small eyes, which however,
according to Hammond and Miall, are covered in larger part by a peculiar shield.

Family SILPHIDAE.
Choleva sp.

A single female, probably accidental, was found under a stone at
" 37," at a piece of beef used to attract insects. Blatchley (1896, 131)
found a beetle apparently belonging to this family in Coon's Cave.
Garman (1894, 81) records Choleva alsiosa Horn from a cave near Lex-
ington, Kentucky, and Packard (1894, 732) mentions Choleva spelsea
Bilimek (1867, 902) from Cacahuamilpa Grotto in Mexico.

The genus Choleva is closely related to the genus Adelops. Adelops
hirtus is an abundant species in Mammoth and other Kentucky caves,
while there are, according to Packard (1888, 89), 59 species of Adelops
in European Caves. The genus has been discarded in Europe, however,
and the species of Adelops placed principally in the genus Bathyscia, a
genus of typically cave forms, eyeless and wingless. Catops is another
genus closely related to this genus. The species of Catops live in fungi,
carrion, or ants' nests. Cope (1872, 166) found a species of Catops
near the mouth of Wyandotte Cave. C. speluncarum (Hamann, 1896)
is found in caves in Sardinia.

Family STAPHYLINIDAE.
Rheochara lucifuga Casey.

Rheochara lucifuga CASEY, Ann. N. Y. Acad. Sci., vn, 1893, 228 (caves near Lex-
ington, Kentucky).

Calodera cavicola GARMAN, Psyche, vn, 1894, 81, fig. 1 (caves near Lexington,
Kentucky) .

Rheochara lucifuga, WICKHAM & BLATCHLEY, Rep. Ind. Geol. Surv., xxi, 1896,
195 (Truett's Cave).

This is by far the most abundant beetle found in the cave. It con-
gregates in abundance at the bait left to attract insects, a piece of meat
or cheese seldom failing to attract at least one or two, and on one occa-

30 FAUNA OF MAYFIELD'S CAVE.

sion 16 were taken from a single piece of beef which had been left
under a stone for a few days, and several others escaped. This species
has been seen crawling about over the walls and floor of the cave, but is
usually found under stones or rubbish. It is essentially a scavenger.
It has been seen feeding at the carcass of a dead mouse, upon a myriapod,
and abundantly upon the bait mentioned above. It probably feeds also
on decaying vegetable matter. It was more attracted by beef than by
other bait. None were taken in daylight or strong twilight, although
baits were placed in the light with equal frequency as in the inner
parts of the cave. It seems well able to take care of itself during a
flood; for a few days after the water receded during the flood of April,
1904, it was to be found at the baits lately submerged. A pair was
taken copulating in November, 1903, under a stone at "31," where a
piece of cheese had been left for some time and about which Sinetta
cavernarum, Rheochara lucifuga, Conotyla bollmani, and Limosina
tenebrarum had congregated in considerable numbers. The larvae and
pupae were not observed.

There is a noticeable variation in size and in the amount of dark
color on the head and near the end of the abdomen. The length ranges
from 4 mm. to 7 mm. The dark areas on the head and abdomen range
from quite dark fuscous to a faint trace of the color.

Garman (1894, 81) says of this and another species:

Both have pretty well developed eyes, and may, therefore, live at times in ordinary
situations, but they are perfectly at home in the deepest parts of caves, and are
at times very abundant there. In all my collecting in ordinary situations I have not
seen either species out of doors, and am disposed to consider them true cave dwellers.

Blatchley (1896, 195) says:

Mr. Garman is doubtless right, for no beetle is going to crawl into the deepest
recesses of caves each day and emerge again at night. So far the species has only
been found in caves and, like Quedius spelseus, has probably inhabited them too short
a time to entirely lose the eyes.

My observations entirely agree with Blatchley's views. I am of the
opinion that it is essentially a cave dweller, and if it should be found
on the outside, its occurrence there will be altogether accidental.

Quedius spelaeus Horn.

HORN, Trans. Am. Ent. Soc., 1871, 332; the same, vn, 158. COPE, Rep. Incl.
Geol. Surv., IV, 1871, 179 (Wyandotte Cave). PACKARD, Mem. Nat. Acad. Sci.,
iv, 1888, 15. WICKHAM & BLATCHLEY, Rep. Ind. Geol. Surv., xxi, 1896, 196
(Wyandotte, Mayfield's, Clifty, Donnehue's, Truett's, and Coon's caves).

Of the Coleoptera this species is next to Rheochara lucifuga Casey
in abundance. It occurs in those parts of the cave least often flooded,
and where there are considerable banks of dirt at the side or bottom of

COLEOPTERA— STAPHYLINIDvE. 31

the cave, but has not been found where the cave does not have at least
a fair degree of moisture. It has been observed a number of times in
twilight, but never quite near the mouth of the cave. It is seen quite
commonly at "6" and seems about equally abundant throughout the cave
from "6" back, where conditions are favorable. These beetles are
attracted by bait of whatsoever sort and feed upon all sorts of decaying
animal and vegetable matter. They burrow into and dig up the ground
to some extent under the stones where bait is left. Larva3 often appear
at the bait after it has been left two weeks or longer. The larva is
described by Wickham (1896, 196) and is easily distinguished from
other larvre found in the cave by its general staphylinidous form and
its large size. I have found the larvae in abundance, and have attempted
to rear them, but for some reason have been unable to keep them alive.
The eggs and pupse were not found.

Some variation is noted in the amount of dark color on the head and
elytra. More noticeable is the variation in the shade of the rufous-
brown color of the body. The body color ranges from quite dark
rufous-brown to leathery rufous. There is no correlation between the
variation in general body color and in the color of elytra and head, some
of the lighter individuals having dark elytra and some of the darker
ones having relatively light colored elytra. The variation in size is
considerable, the length ranging from about 9.5 mm. to 14.5 mm., with
an average of near 12 mm. This species seems entitled to be classed
as a true cave form. It has been found in twilight as well as in remote
parts of the cave, but has not been found in strong light. Besides the
localities mentioned above a single specimen has been recorded from
Colorado.

Quedius iulgfidus Fabricius.

Quedius fulgidus FABRICIUS, Mantissa Insectorum, I, 220.

Staphylinus iracundus SAY, Trans. Am. Phil. Soc., IV, 449; Ent. Works, ed. Le
Conte, n, 564.

Quedius fulgidus HORN, Trans. Am. Ent. Soc., vn, 158. HAMILTON, Trans. Am.
Ent. Soc., xxi, 366. PACKARD, Am. Nat., x, 1876, 286; Mem. Nat. Acad. Sci.,
iv, 1888, 74 (Weyer's Cave, Virginia, and Dixon's Cave, Kentucky). WICK-
HAM & BLATCHLEY, Rep. Ind. Geol. Surv., xxi, 1896, 195 (Donnehue's and
Marengo caves).

Not common. Only 4 specimens were found. This beetle was not
found beyond twilight in Mayfield's Cave. One of the specimens taken
was found at bait under a stone at the first bend in direct rays of light
from the outside. Another was found crawling on the roof at '"6,"
where the reflected rays of daylight reach. The two others were found
as pupae in loose earth under a stone at the first turn. They were taken

32 FAUNA OF MAYFIELD'S CAVE.

October 28, 1903. November 4 one of the pupae was drawn (fig. 3).
The next day one had emerged. The second came out November 6.
The pupa drawn measured 2.8 mm. by 7.1 mm. Blatchley (1896, 196)
found this species in total darkness in Marengo and Donnehue's cave.
Wickharn (1896, 196) says of this species:

Widely distributed (above ground) , being found according to Dr.
Hamilton's recent catalogue, over all of North America, as well as
the other continents, except South America.

Packard (1888, 74) records it from Weyer's Cave in
Virginia and Dixon's Cave in Kentucky, and considers it a
common species in the Middle and Western States. This
is evidently not a true cave species, but its breeding at the
mouth of caves and wandering into caves indicates an
inclination to become cave-inhabiting.

This genus lives ' 'under stones and bark in damp for-
ests" (Le Conte & Horn, 1883, 95). There are 19 North American
species.

Philonthus lomatus Erichson.

A single specimen taken at bait at "40" two weeks after a heavy
flood in the cave. Its occurrence was no doubt altogether accidental.

Tachinus repandus Horn.

Two specimens taken during November, 1903, under a stone just
inside the cave. It was not even in dim twilight and scarcely deserves
mention in this connection.

Lesteva pallipes var. picescens Le Conte.

Some of this species were seen on October 22, 1902, near the mouth.
June 4, 1903, several were seen under stones on loose earth at "3,"
together with their empty pupa cases. At the same time time others
were found under stones as far in as " 11," but not abundantly farther
than the rays of twilight extended.

On October 28, 1903, a bright, warm day, this beetle was extremely
abundant around the mouth of the cave. It was so abundant that 10 or
15 would be found on a single dry maple leaf, yet it was not seen more
than 25 feet from the mouth of the cave. The surface of a small log-
just at the mouth was literally alive with them. The individuals were
taking wing and all of them seemed to be headed from the cave for the
outside.

Investigation on the same day showed that where they had been so
abundant within the cave they had suddenly become scarce. A few

COLEOPTERA— APHANIPTERA. 33

were seen at the first bend, where they had been so extremely abundant
before and the next day some were still there. Two days later a few
were seen at the mouth outside. On November 7 a few were found
just inside the mouth and one at the carcass of a mouse at "11." On
November 10 and 14, none, except one at "6," could be found, although
diligent search was made for them. They were not seen again until
the following autumn.

In October, 1904, several were seen on the walls and a few under
stones from "1" to "4" while a single one was taken on the wall at
' 12. " All disappeared later. Nothing like the abundance of the year
previous was noticed, but they were fairly abundant, and those noted
may have been a few stragglers from among a great number similar to
that of the year before. During October for three successive years this
beetle was seen near the mouth in some numbers. Each time it was
very lively and soon afterwards disappeared. Once it occurred in immense
numbers and was apparently leaving the cave. This has the appearance
of migration.

The species breeds in the cave near the mouth, but where such
numbers went or what became of them, I do not know. I took specimens
of this species in Twin Cave at Mitchell in October, 1902.

Mr. H. F. Wickham says of its distribution and habits:

It lives on the banks of streams habitually. Sometimes it occurs in numbers at
Iowa City. It has a wide distribution, being known from Lake Superior and Mar-
quette, Michigan, Pennsylvania, Alabama, Maryland, Massachusetts, New Hampshire
and Iowa.

A species of Lesteva was recorded from Wyandotte Cave by Cope
(1872, 166). According to Leunis (1886, Bd. n, 95), European species
of this genus live in dark places.

There are 2 species of the genus Lathrobium of staphylinidous beetles
which live in European caves. One of them is eyeless and the eyes of
the other are very small.

Family TENEBRIONIDAE.

A larva of a species of this family was found in April, 1904, under
a piece of rotten wood at "10." It had probably been carried in by
the heavy flood of two weeks before.

Order APHANIPTERA.

A species of Aphaniptera (Siphonoptera) was seen in the cave at few
times. It was once taken off the loose, moist earth at "6," near the
carcass of a mouse, and was found upon bats. In no case was it
abundant.

34 FAUNA OF MAYFIELD'S CAVE.

Order DIPTERA.
Family TIPULIDAE. Crane Flics.

These shade-loving flies are often found at the mouths of caves.
They ;have been taken occasionally in Mayfield's near the mouth in
twilight, while a few were taken in absolute darkness — one at "11"
in November, one at "12" in October, one at "25" in July, and one on
the sand beside the stream at "38" just after the freshet in April, 1904.
I have seen flies of this family near the mouths of small caves at Bowling
Green, Glasgow, and Horse Cave, Kentucky, and also in White's Cave.
Blatchley (1896, 188) took a specimen of Ulomorpha pilosella in Porter's
Cave. Others have reported Tipulidse from caves.

These flies are at home in the twilight at the mouths of caves, but
those found farther back are probably strays.

Of those taken in Mayfield's only the following could be determined:

Trichocera sp.

Coquillett identified one specimen as belonging to this genus.

Cladura indivisa Osten Sacken.

OSTEN SACKEN, Proc. Ac-ad. Nat. Sci. Phil. 1861, 291 (New York, Massachusetts).
ALDKICH, Catalogue N. A. Dip., 1905, 85.

A specimen of this species was taken on the roof at "12" in October.
It was doubtless a stray.

Rhypholophus meigenii Osten Sacken.

OSTEN SACKEN, Proc. Acad. Nat. Sci. Phil. 1859, 226 (United States and Canada).
ALDRICH, Catalogue N. A. Dip., 1905, 84.

A single stray individual was found at "38" in April, 1904. It may
have been carried into the cave with debris during the freshet of a few
days before.

Family PSYCHODIDAE. Moth-like Flies.
Psychoda cinerea Banks.

BANKS, Canad. Ent., xxvi, 330 (New York). ALDRICH, Catalogue N. A. Dip., 1905,
106 (Eastern and Western United States, British Columbia, and Alaska).

Taken at bait or other decaying matter in the cave a number of times
and often observed upon the roof. It occurs in different parts of the
cave but has not been seen nearer the mouth than "9." It breeds in
the cave. Bait containing larvas of this species and of Limosina tene-
brarum was brought in during December, 1903, and the adults emerged
three and four weeks later. Forty or fifty came out from a piece of
decaying meat little larger than the end of a man's thumb.

Blatchley (1896, 188) records Psychoda minuta Banks from Saltpeter
Cave.

DIPTERA— CULICID^:. 35

Family CHIRONOMIDAE.
Chironomus sp.

These small flies are often seen in the cave, usually when flying after
having been disturbed or attracted by the light. They seem fairly
abundant in the remote parts of the cave and are occasionally near the
mouth. Only a few were taken.

They have been found in Twin Cave, near Mitchell, where they were
quite numerous in April, 1904. I have seen them in numbers flying
about lighted lamps in Mammoth Cave, along Echo River, and likewise
in a cave near Bowling Green, Kentucky. Osten Sacken (Packard,
1888, 80) records this genus from Mammoth Cave.

Family CULICIDAE.
Anopheles punctipennis Say.

SAY, Jour. Ac. Phil., in, 9 (all United States east of Rocky Mountains). ALDRICH,
Cat. N. A. Dip., 1905, 122 (Eastern United States and Canada).

Quite common within the mouth of the cave during winter, but not
seen in summer. It congregates in cracks and recesses and in the angle
formed by the wall and roof. It is quite abundant from the mouth to
'•' 4," where direct rays from the outside reach, but beyond the reach
of these rays its occurrence abruptly ceases. In the fall the insect
enters the cave early. It was noticed during the first half of October,
1902, and in 1903 appeared first between September 30 and October 3,
for it was not noticed on the former date and was quite abundant on the
latter date. This species has not been seen in the cave in spring as
late as April. On March 17, 1903, Anopheles was still not very active
and would hardly move when touched. I think the Anopheles had
moved very little all winter. However, before the next trip, April 4,
the Anopheles had completely disappeared. There had been several
warm and bright days during this interval.

When the Anopheles first arrives it is very active and flies the moment
the light is thrown upon it or an attempt made to approach it with the
hand. When the temperature becomes lower Anopheles becomes less
active and after the first cold snap is quite dormant, and remains more
or less so all winter, depending somewhat, however, upon the tempera-
ture. During the cold months it is so stiff that it can not fly and will
scarcely attempt to move. It seems reasonable that in regions of caves
malarial conditions might be aggravated by these mosquitoes hibernating
so freely in them.

36 FAUNA OF MAYFIELD'S CAVE.

Anopheles quadriraaculatus Say.

SAY, Etom. Works, ed. Le Conte, I, 241; n, 39 (Atlantic States, Canada, Southern
Europe). ALDRICH, Cat. N. A. Dip., 1905, 122.

A single specimen of this species was taken just inside the door,
September 24, 1903.

Culex pipiens Linnaeus.

LINN^US, Syst. Nat., 12th ed. ALDRICH, Cat. N. A. Dip., 1905, 130.

A few of this species were taken near the mouth during the fall.
Two were found at "8" during March and one near the mouth in April.
It is found in recesses of the wall or ceiling and probably hibernates in
the cave to a limited extent.

Family MYCETOPHILIDAE.
Polylepta leptogaster Winnertz.

WINNERTZ, Mon., 746 (Europe). ALDRICH, Cat. N. A. Dip., 1905, 141 (Europe and
United States).

Two specimens taken far back in the cave, along the stream.

Rymosia filipes Loew.
LOEW, Cent., ix, 36 (Connecticut). ALDRICH, Cat. N. A. Dip., 1905, 145.

Fairly common near the mouth and back to beyond the second turn
(to "8")> where there is almost absolute darkness.

Mycetophila discoidea Say.
SAY, Jour. Acad. Phil., vi, 153 (Indiana). ALDRICH, Cat. N. A. Dip., 1905, 146.

Not common in the cave. Only a few have been taken. They have
been found only in fall or spring but that is possibly a coincidence.
They occur usually on the wall or roof but one was found on the under
side of a stone at " 6. "

Mycetophila ichneumonea Say.

SAY, Jour. Acad. Sci. Phil, in, 16 (Pennsylvania). ALDRICH, Cat. N. A. Dip.,
1905, 146.

A single specimen was taken near the mouth of the cave during
October.

Mycetophila obscura Walker.

WALKER, List. Brit. Mus., I, 101 (Canada). ALDRICH, Cat. N. A. Dip., 1905, 146
(Canada and White Mountains).

Fairly common from the mouth to " 11, " and to be found occasionally
as far as to the mound. This is one of the abundant Mycetophilidse.
Taken also in Twin Cave at Mitchell. Its known distribution includes
New Jersey £also.

DIPTERA— MYCETOPHILIDjE. 37

Mycetophila umbraticus Aldrich.
Rep. Ind. Geol. Sur., XXI, 1896, 187 (Shiloh Cave); Cat. N. A. Dip., 1905, 146.

Very common. One of the most abundant Mycetophilidge in the cave.
It has been taken from near the mouth to the mound, and once beyond.
It is often quite abundant, especially from near the mouth to " 17, " and
is fairly common to the mound at all seasons. Blatchley took a single
specimen of this species from Shiloh Cave and from it this species was
described.

The following is self-explanatory:

NEW SPECIES OF MYCETOPHILA.
By C. F. ADAMS, Chicago.

In looking over some Diptera from Mayfield's Cave, Indiana, I have come across
two forms which I believe to be new. The material was collected by Mr. A. M.
Banta, of the University of Indiana, and my thanks are due him for the privilege of
studying the collection.

MYCETOPHILA ANALIS n. sp.

Male. Head brownish-black, mouth parts and basal joints of antennae yellow,
remaining joints of antennae light brown. Thorax brownish-black, lateral margins of
mesonotum and the pleurae brownish -yellow; mesonotum with short yellow pile and
black bristly hairs, the latter distributed along the sides; scutellum with an apical pair
of strong bristles. Abdomen brownish-black, apex yellow, with short yellow pile.
Coxae yellow, femora light yellow, tibiae and tarsi becoming darker distally; the front
tibias are without bristles, except the apical ones, the second are provided with a
row of indistinct setulae, and the hind tibiae have rather strong bristles; front tarsi a
little over twice as long as front tibiae, middle tarsi about twice as long as middle
tibiae, hind tarsi 1.5 times as long as their tibiae. Wings nearly hyaline; third vein
and anterior branch of the fourth vein divergent distally, furcation of fourth vein in
front of base of third vein, furcation of fifth vein considerably posterior to it. Halteres
light yellow. Length 4 mm.

One specimen. Taken just within entrance of cave, March 18, 1904.

MYCETOPHILA INCERTA n. sp.

Male. Head dark brown, mouth parts and basal joints of antennae yellow.
Thorax obscure yellow; mesonotum with two V-shaped figures, one set within the
other, resulting from the fusion of the two lateral stripes posteriorly, and with the
median one between them being divided anteriorly; scutellum and metanotum brown,
former with an apical pair of bristles; mesonotum with short yellow pile laterally
with black bristles. Abdomen obscure yellow, base of each segment dark brown
dorsally, on the posterior segments this color comes to encircle the segment; the short
pile is black. Legs light yellow, tarsi becoming tinged with brown; besides the long
apical bristle the middle and hind tibiae have each a row of shorter bristles; anterior
tarsi twice as long as their tibiae, middle tarsi 1 . 5 times as long, and the hind tarsi
as long as their respective tibiae. Wings hyaline; third vein and anterior branch of
fourth vein divergent, furcation of fourth vein beyond base of third vein. Halteres
light yellow. Length 4 mm.

One specimen. April 9, 1903.

38 FAUNA OF MAYFIELD'S CAVE.

Bolitophila hybrida Meigfen.

MEIGEN, Klassification, I, 47 (macrocera) ; Syst. Besch. Eur. zweifliigeligen Insckt.,
I, 221, pi. vin (fusca) (Europe). ALDRICH, Cat. N. A. Dip., 1905, 147 (Europe
and White Mountains).

This species is very abundant in twilight near the mouth of the cave
at times and is always common from near the mouth to " 19, " where
the drier part of the cave begins. It has twice been taken in the damp
portion of the cave between "23" and "28."

Sciara sp.

Probably 2 or 3 species. Not rare in the cave. Often seen but not
easily caught. Species of this genus are said to be very hard to deter-
mine and I got no specific determination from Mr. Coquillett, to whom
they were sent. Sciarse have been found outside the cave in the ravine,
under chunks of wood, etc. Within the cave they occur near the
mouth and through all parts of the cave. They are sometimes seen
flying about in the cave or resting on the wall. They also occur under
rocks and other debris, where they also breed. Their larvae live in
decaying organic matter. Two adults were reared in a bit of decayed
meat brought from the cave.

Sciarae were taken in Truett's Cave also. This genus was recorded
from Mammoth, Wyandotte, and Bradford caves by Packard (1888, 80),
and from Wyandotte, Saltpeter, and Donnehue's caves by Aldrich and
Blatchley (1896, 186). Call (1897, 385) records Sciara inconstans Fitch
from Mammoth Cave, where it was found breeding in a rotten apple.

Mycetophilidse have been recorded from Mammoth Cave, and Wyan-
dotte and Bradford caves in Indiana, a cave at Manitou, Colorado,
and Fountain Cave in Virginia, by Packard (1888, 21, 80), and from
Shiloh, Marengo, Saltpeter, and Donnehue's caves in Indiana (Blatchley,
1896, 186-191). They vary greatly in abundance in Mayfield's Cave.
On June 4, 1903, they were congregated by thousands just inside the
partition at the mouth, but by the last of September, when the next
visit was made, the number had reduced to normal, which is an average
of 1 to every 6 or 8 square feet on the walls and roof near the mouth.
They decrease in numbers rapidly farther in, and not many are to be
found in absolute darkness. They are found on the wall or ceiling of
the cave, where they are apt to congregate in any wide crack or depres-
sion. None except M. discoidea and Sciara were observed in situations
which could possibly suggest that they might feed in the cave. A single
specimen of the former was found under a rock where bait had been,
but it was not at the bait. Sciara breeds within the cave. These flies
may simply be sufficiently at home in the cave to hibernate there; to
wander in at various times and remain for a while; or they may feed

DIPTERA— PHORID^E. 39

there, which seems most unlikely, except the Sciara and possibly
M. discoidea.

Those near the mouth in winter seem more or less dormant, depending
upon the degree of cold, but they do not seem as dormant as the Anopheles
and do not remain in the dormant state throughout the winter to the
extent that Anopheles does, for a rise in temperature causes them to
become very active again, while Anopheles does not respond readily to
a change in temperature during the winter. These Diptera are probably
hibernating and many of them at least leave the cave in spring, but
others are seen well within the cave at all times. It seems to me very
likely that Bolitophila hybrida, Mycetophila umbraticus, and Mycetophila
obscura breed within the cave, but their larval stages were not identified,
although many unknown larva? were found.

Family CECIDOMYIDAE.

Diplosis sp.

A single specimen taken in the cave beyond the mound.

Family DOLICHOPODIDAE.
Liancalus genualis Loew.

LOEW, Neue. Beitr., vm, 70 (Middle States). ALDRICH, Cat. N. A. Dip., 1905, 298.
A specimen of this species was taken on the wall at ' '2' ' in September.

Neurigona sp.

A specimen was found near the mouth during October and another
individual of this family was found on the wall at "3."

A species of this family was recorded from one-half mile within
Diamond Cave, Kentucky, by Packard (1888, 80). Flies of this family
live in damp places, often with rank vegetation. They breed in decaying
organic matter.

Family PHORIDAE.

Aphiochaeta (Phora) nigriceps Loew.

LOEW, Cent., vn, 99 (District of Columbia). ALDRICH & BLATCHLEY, Rep. Ind. Geol.
Surv., xxi, 1886, 190 (Wyandotte Cave). ALDRICH, Cat. N. A. Dip., 1905, 337
(all eastern United States).

Fairly common. It is found in all parts of the cave. It is not more
common, at least, in twilight than in absolute darkness, and has been
more often noticed in the part of the cave through which the stream
runs. It is often seen on the wall or flying about the light, but is usually
on the floor or under a stone. It is quite generally attracted to the bait
left under stones, and has been seen at a dead mouse on two occasions.

40

FAUNA OF MAYFIELD'S CAVE.

This is one of the most active of the flies found in the cave. It crawls
rapidly and flies readily. When strong light is thrown upon it it crawls
away or takes wing quickly and when found must be caught instantly
if at all. The larvae and adults live upon decaying vegetable and animal
matter. The larvae of this species and other larvae were brought into
the laboratory in pieces of bait which had been in the cave. The larvae
were not distinguished from others, but the pupae were identified and
two drawings made (figs. 4 and 5) .

This species has been recorded from Wyandotte Cave, and I have
taken it in Twin Cave and Mayfield's.

Aphiochaeta (Phora) rufipes Meigen.

MEIGEN, System. Beschre., vi, 216 (Europe and North America). ALDRICH, Cat.
N. A. Dip., 1905, 337.

Two specimens, probably belonging to this species, were taken at the
first bend in bright twilight. It is possibly not uncommon in Mayfield's
Cave, and others of this species may have been mistaken for nigriceps.
It has been recorded from Mammoth Cave (Call, 1897, 386) , and I have
it from Twin Cave.

FIG. 4. — Pupa otAphiochceta nigriceps.
Side view. X 15.

FIG. 5.— Pupa of Aphiochteta nigriceps.
Dorsal view. X 16.

The occurrence of Aphiochseta in caves has been mentioned by many
collectors. Phora has been recorded from Mammoth, Carter, and Wyan-
dotte caves by Osten Sacken (Packard, 1871, 745, and 1888, 80), Cope
(1872a, 161), Hubbard (1880, 39), and others.

Family MUSCIDAE.
Calliphora vomitoria (Linnaeus). Bluebottle Fly.

LINNAEUS, Fauna Suec., 2ded., 452 (Sweden). ALDRICH, Cat. N. A. Dip., 1905, 520
(Europe, Canada, Alaska, United States).

One seen in the cave in February, 1905, on the side of a small stone
at " 21. " It was dormant when found, but when brought in and warmed
up became active.

Musca domestica Linnaeus. Common House Fly.

LINNAEUS, Syst. Nat., 10th ed., 1758 (Europe). ALDRICH, Cat. N. A. Dip., 1905, 528
(almost cosmopolitan) .

A single house-fly was found within the cave April 4, 1903.

DIPTERA— HELOMYZID^E. 41

Family ANTHOMYIDAE.
Hylemyia lipsia Walker.

WALKER, List of Dipt. Ins. in Brit. Mus., iv, 1849, 928. ALDRICH, Cat. N. A. Dip.,
1905, 552 (Illinois, eastern United States, and Canada).

Taken twice in the cave— an adult at "6" on the wall and a pupa
October 3, 1903, at " 17. " From the pupa an adult emerged December 3.

Pegomyia affinis Stein.

STEIN, Berl. Ent. Zeitsch., XLII, 239. ALDRICH, Cat. N. A. Dip., 1905, 558 (Pennsyl-
vania, Virginia, and Illinois).

Taken within the cave twice. A single specimen at ' '3" and another
under a stone just within the cave April 15, 1904. April 28 six speci-
mens were taken at ' ' 17, " and nearby there were several others appar-
ently of the same species.

This family is widely distributed in caves. Anthomyia is recorded
f rom Wyandotte (Cope, 1872, 161; Collett, 1873, 80), Mammoth (Tell-
kampf, 18446, 382; Packard, 1871, 745; Cope, 1872, 161; Hubbard, 1880,
37), Fountain Cave, Virginia (Packard, 1888, 80), Donaldson's Cave
(Collett, 1873, 80), and a cave near Orleans, Indiana (Packard, 1873,
93) . Some of these references probably should refer to Leria, however.

Anthomyia mitis Meigin is recorded from "Adelsberger Hb'hlen" in
Europe (Hamann, 1896, 143).

Family HELOMYZIDAE.

Oecothea fenestralis Fallen.

FALLEN, Heteromyzides, 5 (Helomyza) (Europe). MEIGEN, Syst. Beschr., vi, 56.
ALDRICH, Cat. N. A. Dip., 1905, 572 (Europe, Canada, and New York).

Fairly abundant in the cave. At times it is found in some numbers
in different places. It has been most often seen from near the mouth
to "17," but has also been taken in remote parts of the cave. This
species stays in the more retired places,
such as small, low side-passages or
cracks in the wall where there is con-
siderable moisture. For this reason it

_ • i . FIG. 6.— Larva of Oecothea fenestralis. X ».

is not so conspicuous as Lena latens

and Leria defessa, the other two abundant species of this family, but
with the exception of these two it is the most abundant Helomyzid in
the cave. This species is even more sluggish in its movements than
others of this family.

Its feeding habits were not observed. It increases in numbers in
summer and probably breeds abundantly then, although, as the following
observations show, it also breeds in winter.

42

FAUNA OF MAYFIELD'S CAVE.

FIG. 7. — Pupa of Oecothea fenestralis,
Dorsal view. X 15.

December 16, 1903, from a piece of cheese which had been left as
bait at "41," several larvae and part of the bait were taken from the cave
and stopped up in a bottle. January 8, three larvae were separated
from the others. These must have been eggs or very small larvae when
taken, as others from the same material had emerged before. January 8
one was drawn (fig. 6). The next day one of the larvae pupated.
January 11 a second pupated, and on the 15th the third. The pupae were

kept with dirt slightly moist.
February 2 two flies were in
the bottle. The third pupa had
died. One pupa was drawn in
several positions (figs. 7, 8,
and 9).

The larva is cylindrical, rap-
idly tapering toward the front
and truncate behind. It is
white and has a dark, some-
what Y-shaped area at the head.
The segments are indistinct,
but the trachea and spiracles
are easily visible. Its length
is 7. 3 mm. ; thickness, 1. 35 mm.
The pupa is plump, smooth,
and shining. It is light reddish-
brown, with the spiracles and
posterior margin darker red-
dish, becoming darker on an-
terior five segments and espe-
cially on anterior three. The
first segment is very dark;
the second and third are somewhat lighter; the fourth and fifth are
much lighter, and the sixth to last segments about a uniform tint.
Length 4. 1 mm. , width 1.36 mm., diameter 1.4 mm.

Leria latens Aldrich.

Rep. Ind. Geol. Surv., xxi, 1896, 188 (Donnehue's, Shiloh, Porter, and Mayfield's
caves in Indiana) ; Cat. N. A. Dip. , 1905, 573.

Very abundant; the most abundant species of Helomyzidae in the
cave. Found from the mouth to the mound, where it becomes scarce.
It is most abundant from "5" to " 17," and is common from the mouth
to "20." It has been rarely noted beyond the mound. During the
summer this species becomes extremely abundant near the mouth, so

FIG. 8. — Pupa of Oecothea fenestralis.
Ventral view. X 15.

FIG. 9.— Pupa of Oecothea fenestralis.

Side view. X 15.

DIPTERA— HELOMYZID^E. 43

that they average easily 3 or 4 and at times even 12 or 15 to the square
foot for considerable areas of the walls. L. latens does not commonly
occur in any abundance far above the floor of the cave, but congregates
on overhanging surfaces under the arches of low side-holes in the wall,
on the under side of projecting angles, and in similar places where there
is considerable moisture. L. latens is not often seen under a stone and
is not attracted to bait. It is probably less sluggish in its habits than
O. fenestralis, but does not crawl rapidly and seldom takes flight. The
presence of light for a short time or of the hand near is not sufficient to
cause it to stir at all. Nothing short of actual contact with a foreign
object or heat from a lamp stimulates it to motion. When it is caused to
move it crawls slowly away and if continually disturbed may occasionally
be induced to fly. Usually, however, it will not fly unless forcibly dis-
lodged and falling through the air, and sometimes it will not take wing
even then. Bright light, if thrown upon it for a time, usually causes it to
move, in many cases individuals turning squarely about and crawling
away from the scource of light. Some individuals, however, paid no
attention to the"; light, even when exposed to it for an hour or more.
I have been unable to determine the food of this species, nor have I
identified its larva or pupa, although among hundreds of dipterous larvae
found in decaying meat in the cave some, I am confident, were larvae
of this species.

The excessive abundance of this species in early summer is probably
due to its breeding more abundantly then, although it may breed in the
cave at all seasons. The apparent increase in numbers may be partly
due to individuals entering the cave from outside with the coming of
summer. This species is particularly infested with mites. Sometimes
nearly every fly has some of the parasites and some individuals as many
as 30 or 40. The parasites collect upon the back of the abdomen and
thorax and fasten themselves to all vulnerable parts of the body.

Taken in Twin Cave, where it is abundant. It was found outside
Mayfield's Cave, under chunks of wood in the ravine near the mouth.

Leria defessa (Osten Sacken).

OSTEN SACKEN, Bull. U. S. Geol. and Geog. Surv., vn, vol. in, No. 1, 168. PACKARD,
Mem. Nat. Acad. Sci., iv, 1888, 15, 17, 21, 80, 132 (Mammoth, Wyandotte, Carter,
and Hundred Dome caves, and several smaller caves near Glasgow Junction, Ken-
tukcy, New Market Cave, Virginia, and a cave at Manitou, Colorado) . BLATCHLEY,
Rep. Ind. Geol. Surv., xxi, 1896, 188 (Wyandotte Cave). ALDRICH, Cat. N. A.
Dip., 1905, 573.

Common throughout the main passage and larger side-passages of
the cave. It is most abundant from "25" to the mound, but is found
scattered throughout the cave, except in the driest portions. This

44 FAUNA OF MAYFIELD'S CAVE.

species is found over the wall and roof more than the preceding species,
and is sometimes seen on the floor or under debris. It is occasionally
attracted to bait, near which it has been seen pairing. This is the only
species of Helomyzidae which I have ever seen to fly of its own accord.
It is often attracted to one's light and occasionally flies squarely against
the glass over the front of the lamp, while I have often heard it buzzing
about or felt it fly against my body. Its reaction to a light held steadily
upon it is uncertain, as it crawls toward the light as frequently as away,
and often fails to respond at all. When disturbed by bringing the
hand near or touching it, it is more likely to take wing than any of the
others of this family. I have not identified its larva nor determined its
food, more than to observe that it is attracted to bait sometimes. There
is some increase in its numbers in summer, but it never approaches
L. latens in abundance.

A single specimen of this species was taken on the ledge outside the
cave in March.

Leria serrata (Linnaeus).

A few specimens of this species were found within the cave. It may
be more abundant than it appears, as it very closely resembles L. latens,
and among the abundance of the latter would not readily be observed.
Known from Europe.

Leria fraterna Loew.
Monograph in, 1872, 51 (Alaska) .

A single specimen was taken in January. Probably accidental in
occurrence. Known from Alaska, British Columbia, and New Hampshire.

Baron Osten Sacken (Packard, 1888, 81) remarks: ' Blepharopterse
(Helomyzidae) are often found in caves, where they are said to breed
in the excrement of bats. ' ' Their presence and abundance in Indiana
caves bear out the first part of this statement. They congregate in
abundance in recesses and cracks and on the damper portions of the
wall, usually not very far above the floor, and are particularly abundant
under the edges of projecting ledges and at the top of low arches at
the side of the cave.

Curiously enough, flies of this family are always more abundant on
the overhanging slopes than elsewhere. This is especially true of
0. fenestralis; to a smaller degree it is true of L. latens, and much less
true of L. defessa.

The larvae of 0. fenestralis have been taken from bait, and L. defessa
is occasionally attracted to bait; otherwise no indication as to their food
has been found. Fly larvae and empty pupal cases have been found

DIPTERA— BORBORID^E.

45

about bait and in soft moist earth where there is organic matter. Some
of them at least must have belonged to Helomyzidas, but with the excep-
tion of 0. fenestralis no positive identifications of larvae or pupae have
been made.

The remarkable increase in numbers of Leria latens in summer was
mentioned before. The other species likewise increase to a certain
extent at the same time. L. defessa fluctuates in numbers less than the
other species.

The relative abundance of Leria latens, L. defessa, and 0. fenestralis
in different parts of the cave is about as follows:

Leria latens.

O. fenestralis.

L. defessa.

At "6"

5

3

1

"10" to "12"

30

3

1

"15" to "19',

30

1

2

"20" to "23"

2

Rare

3

"29" to "42"

Rare

Rare

Fairly common

The activity of these flies is affected by change in temperature. In
cold weather those near the mouth, where they are subject to a temper-
ature as low as 5° C. , become quite dormant and are scarcely able to
move. In summer those nearest the mouth are as active as those
farther in.

Family BORBORIDAE.
Limosiiia tenebrarum Aldrich.

Rep. Ind. Geol. Surv., xxi, 1896, 190 (Truett's, Donnehue's, Clifty, Marengo, and
Wyandotte caves); Cat. N. A. Dip., 1905, 575.

By far the most abundant species in the cave. It is abundant in all
parts. During the winter it is not found near the mouth, where the
temperature becomes near 0° C., and is not common from "6" to "17,"
where the winter temperature is often quite low. But during the
summer it is extremely abundant near the mouth, moist places where
the ground is loose, the under side of rocks, and the ground under rocks
being sprinkled with it. On turning over a stone it hops and crawls
away in immense numbers.

This fly may be expected at almost any bait or decaying organic
matter in any part of the cave, and where the amount of decaying
matter is considerable it will be found in numbers. It is not usually
seen in any abundance upon the wall, but is always abundant under
stones or other debris and occasionally becomes numerous on the wall
near the floor. In summer, when the part of the cave subject to consid-
erable seasonal change in temperature becomes warm, these flies breed

46

FAUNA OF MAYFIELD'S CAVE.

FIG, 10. — Pupa of Limosina tenebrarum,
Ventral view, x 20.

very rapidly. This brings about a great increase in their number near
the mouth of the cave and to some extent as far back as "11," but
beyond that or at least beyond the mound, where the passage is subject
to very little change in temperature, they breed in about equal abundance
at all seasons and their number remains about constant throughout the
year. In the spring the earth from "4" to "6" where not too hard
becomes worked up and the pupal cases of this species are abundant in
the soil, on the wall above, on rocks, and everywhere about. The flies
themselves are extremely abundant, crawling through the loose soil and
over the walls. During June, 1905, an estimate of the numbers of these

_____ _ flies on the wall near the floor at

"6," where they were about
equally abundant for considerable
areas, placed their number at 75
to 100 to the square foot. This
is the extreme in abundance. But
at any season and in any part of
the cave, except near the mouth
in winter when the temperature
is low, these flies congregate in
numbers and are observed pairing
near any organic matter as soon
as putrefaction begins.

The larva? appear very soon
and in immense numbers upon any
organic matter left in the cave.
The larvse pupate in about two
weeks. The pupae are embedded
in the decaying matter usually,
but sometimes are seen on the sur-
face or fastened to a rock or other
object if quite near where the larvae feed. The adults emerge from the
pupae in about a week. Figs. 10, 11, and 12 represent pupae of this species.
As Blatchley (1896, 190) observed, these insects leap rather than fly when
disturbed. In escaping they crawl and leap a few inches, using their
wings to aid them, but never take an extended flight, even when tor-
mented and continually disturbed for some time. They do not move
very rapidly, but are much more active than the species of Helomyziclae
and respond more quickly to a stimulus. When a stone under which
are numbers of these flies is turned over they hop and crawl rapidly
away, apparently more rapidly when a bright light is thrown upon
them; however, when a light, as a candle, is left among them when they

FIG. 11.— Pupa of Limosina tenebrarum.
Side view.x 19.

FIG. 12.— Pupa of Limosina tenebra runt.
Dorsal view. X 19.

LEPIDOPTERA— NOCTUID^. 47

are at rest they are soon attracted toward it. A candle was placed on
a low shelf at " 6, " where they were very numerous. Some were observed
to approach it after a time and a few even to crawl up the candle until
they were caught in the drip. The candle was allowed to burn out,
leaving behind literally a pile of these insects in the drip. On the roof
above a lighted candle these flies move only sufficiently to escape the heat.
When kept in small bottles in a dark room to which some light was
admitted individuals of this species occupied the end of the bottles away
from the source of the light.

These flies were often reared in the laboratory from eggs or larvae
brought in from the cave. Those reared from very young larvae in
small closely-corked bottles were only two-thirds the normal size.

Another species of this genus (Limosina stygia Coquillett) occurs in
similar situations in Mammoth Cave (Call, 1897, 384).

Order LEPIDOPTERA.

Family NOCTUIDAE.
Scoliopteryx libatrix Linnaeus.

LINN.EUS, Syst. Nat., 1758, 507. SMITH, Bull. 44, U. S. Nat. Mus., 1893, 224.

This species hibernates in the cave in considerable numbers. It is
not found beyond twilight, and most individuals occur from the first
bend at "4" to about 50 feet farther in. Some remain quite near the
door and one or two were seen as far as " 6. " They usually rest upon
the roof, but a few are found on the walls and some under projecting
angles from the wall. They enter the cave early in the fall. Several
had already come September 24, 1904, and after October 15 there were
apparently no more arrivals. They leave the cave early in April. None
seemed to have moved April 4, 1904, and April 15 most of them still
had not moved, but by April 28 all had disappeared. March 4, 1905,
all yet remained where they had been during the winter, but April 8
only 6 were seen, and one of those was not where it had been during
the winter. April 29 all were gone.

These moths are rather inactive from the time they enter the cave.
Those noted on September 24 crawled a little and moved the wings some
when touched or if dislodged fluttered weakly to the ground, but showed
no inclination to take flight. Later, when it becomes cooler, they are
scarcely capable of moving and if partly dislodged often fail to regain
their footing. Toward their time of leaving the cave in the latter part
of the winter they become somewhat less dormant.

Some of this species that were brought into the laboratory became
rather active, due to the constant jarring in carrying them several miles

48 FAUNA OF MAYFIELD'S CAVE.

or to their becoming partially warmed after reaching the laboratory.
When placed in a dark room where the temperature ranged from 6° to
14° C. they became less active again, but seemed to become hardly as
dormant as before. It seems most likely that the activity of this moth
is directly influenced by temperature. However, the following peculiarity
in its behavior was noted:

September 24, 1904, when many of these moths had already entered
the cave, the temperature outside was 20.6° C., the average for the
preceding week had been 18.8° C., while the minimum had been 10.6° C.
The temperature at ''6" was 11.9° C. October 15, when all the moths
had entered the cave, the temperature outside was 12° C. ; for the pre-
ceding week it had been 15.5° C., while the minimum had been 5° C.
The temperature at "6" was 11.6° C. April 8, 1905, when nearly all
the moths had left the cave, the temperature outside was 5.3° C., the
week's average had been 12.9° C., while a minimum of 0° C. had been
reached three times. The temperature at "6" was 7.4° C. April 29,
when all had disappeared, the temperature outside was 16.4° C., for
the preceding week had been 14.5° C., and a minimum of 5° C. was
reached. The temperature at "6" was 8.3° C. From these figures it
will be seen that Scoliopteryx libatrix leaves the cave while the temper-
ature where it spent the winter is 4° C. lower than when it entered the
cave, while the average outdoor temperature was 8° C. lower than when
it went into winter quarters. The temperature just within the cave is
rapidly rising at the time the moths leave, but does not get as high as
that when they entered the cave until midsummer.

These moths seldom change their positions in the cave after becoming
settled in the fall until about time for their departure in the spring.
There are indications that some of them shift their positions somewhat
soon after arriving and just before leaving, but I doubt if at other times
they move at all unless disturbed.

In the cave this species shows an almost perfect orientation with
regard to the light. Nearly every individual takes a position with the
axis of its body in the direction of the rays of light and with its head
from the source of light. There are some exceptions to this rule, but
it is true in 80 or 90 per cent of the cases. Those individuals which are
on the wall do not seem to head from the direction of light so much as
those on the roof, but this seems accounted for in the cases of some of
them which are subjected to reflected light from different directions.
Those under overhanging shelves are sometimes subjected to light which
has become so diffused that there is no one direction from which it
seems especially to come, and very naturally these individuals are headed
without regard to the direction of the mouth of the cave. Just past

LEPIDOPTERA — NOCTUID^.

49

the bend at ' '4' ' an interesting case was noted. Three moths quite near
each other did not head from the direction of the entrance, but at a
considerable angle from it. The light from the entrance struck the
east side of the wall at ' '4' ' at the proper angle and so strongly that
the reflected rays served as the principal source of light for a region on
the roof, and it was in this area that these moths were located and to
this source of light that they had oriented themselves. It must be said,
however, that some of the moths upon the wall, and occasionally one
upon the roof, are not oriented with their heads from the source of
light. Not all the individuals orient themselves immediately upon
arrival, but in a few days nearly all are oriented. Otherwise they do
not orient themselves at all. I turned a number of individuals so that
their heads were toward the source of light or at right angles to it.
The position of each was indicated by arrows on the roof or wall and
noted so that the same individual could be observed on another trip and
its change of position, if any, noted. In the following table are given
the results of this experiment.

Table showing results of experiments as to orientation of Scoliopteryx libatrix to light.

No. of
indi-
vidual.

Position of head
with regard to light

Position of moth
changed or marked
as found.

Oct. 28, Nov. 11,
11104. 1904.

Dec. 9,
1904.

Dec. 21,

1904.

a, b, c,
d, e.

2

3
4

5

6

7

8
9
10
11
12

[• Toward

Disturbed

Marked... All gone..
Marked

Unmoved..
Moved 4
inches and
from light.
Unmoved..
! do
Marked

do

Unmoved.
Gone.

Unmoved.
Do.
Do.

Do.
Do.

Gone.
Unmoved.
Gone.
Marked.
Marked.

From

Undisturbi
Disturbed.

;d

At right angle

....do
Toward
From light spo
east wall.
Slightly to war
From light spo
opposite wa
At right angle
From

5

ton
rl

i Marked .

Undisturb*
do
do

;d.

Marked.... Unmoved.
do do

..do..

ton

11...

3

do

Marked

Unmoved..
Marked

.(In

Disturbed

Undisturbi
Disturbed.

;d..

Toward

do

do

Undisturb<
Disturbed.

3d

Found at right
angles, turned
180° and left at
right angles.

No. of

indi-
vidual.

Jan. 19,

1905.

Jan. 28,
1905.

Feb. 17,
1905,

Mar. 4,

1905.

Apr. 8.,

1905.

Apr. 29,
1905.

1
3
4
B

6

7

9
11
12

Unmoved
do
do
do

do
....do

do

Un

moved

Unmoi
do .
do.
do.

do.
do.

do

red .

Unmoved

Gone

Gone.
Gone.

do
do
do

do.,
do

do

do
do
do

do..
do

..do..

Unmoved

Gone.

do

..do..
do

..do..

do
do

..do..
do

do
do

do
do

do
Unmoved

50 FAUNA OF MAYFIELD'S CAVE.

In all the cases except one (No. 12) in which the moths were disturbed
they changed their positions so that the individuals could not be recog-
nized later. In one case (No. 2) the moth which had been moved so
that it headed toward the light turned from the light, but later moved
again and was lost sight of. In no case where the moth was left
undisturbed did it move during the winter. Two (Nos. 4 and 11) headed
toward the light and one (No. 3) at right angles to it were noted after
they had settled in the cave, and they remained unmoved until they left
the cave in the spring. It may be said by way of summary: Scoliop-
teryx libatrix when wintering in Mayfield's Cave usually takes up a
position with its head directly from the source of light. Some which
do not assume this position immediately upon entering the cave do so
within a few days; if, however, this position is not assumed within a
short time it is not taken at all. If undisturbed they do not change
their positions after becoming settled in the cave. If disturbed they
usually move and the movement in the only case actually observed was
directly from the source of light.

A number of the moths were brought into the laboratory and kept
in a dark room into which some light was admitted by leaving the door
slightly ajar. November 11 three were confined in the dark room in a
broad glass dish partially covered, so that they could not escape. The
light which reached the moths consisted of rays directly from the door,
as the black walls reflected very little. A little water was kept in the
dish, so that the air was moist. They were active when left.

Four weeks later they were found with their heads exactly from the
door. One of them fluttered a little at the light of a match held near it
outside the dish. The dish was turned so that they were headed toward
the light, only one of them having been disturbed. The next day there
was no change in their positions, except that one had crawled up the
side of the dish and was resting at about a right angle to the light.
On the second day, however, two were headed exactly from the light,
and the third, on the side of the dish, was headed at about an angle of
135° from the light. Three days later one was headed toward, one
from, and one at a right angle to the light. These three died, and
January 28 a number of others were brought in and kept in the same
manner. At first most of them seemed to react to the light and "head"
from its direction, but after the dish was shifted a few times by turning
it around so that the light struck the moths from a different direction
they reacted less definitely and very little could be determined regarding
their light reactions. However, an average of all the observations made
showed that 57 per cent headed from the light, 28 per cent toward the
light, and about 15 per cent approximately at right angles to it.

LEPIDOPTERA— NOCTUIDyE. 51

These moths were considerably shaken up in carrying them in from
the cave and had become partly warm before they were placed in the
dark room. The temperature of the dark-room remained moderately
low usually, but may have been raised occasionally when the room was
used for photographic work.

The results of this experiment were confusing and somewhat unsat-
isfactory, but seemed to indicate that the moth tended to orientate itself
with its head from the source of light wherever it was. Under more
favorable conditions the results ought to be definite and convincing.

I have seen this moth in Donaldson's Cave in autumn. According
to Smith (1893, 224) its distribution includes Nova Scotia, Hudson Bay
Territory, and south to Texas, New Mexico, and west to California, and
it is found from May to November.

Plathypena scabra Fabricius.

FABRICIUS, Ent. Syst., Suppl., 1794, 4448. SMITH, Bull. 44, U. S. Nat. Mus., 1893,
395; Bull. 48, U. S. Nat. Mus., 1895, 111.

Less abundant than the former species but also fairly common,
hibernating in the cave. On the average this species does not remain
quite so near the mouth as S. libatrix, but it never goes beyond twilight.
It, too, commonly takes up a position for the winter with its head from
the light, but this orientation is not nearly so constant as in the other
species.

It is common in Donaldson's Cave at Mitchell. According to Smith
(1893, 224) this is one of the most abundant species from June to
November, from Nova Scotia to Texas east of the Rocky Mountains.
I find no mention of its wintering over.

Order ORTHOPTERA.
Family LOCUSTIDAE.

Ceuthophilus styglus (Scudder).

Rhaphidophora stygia SCUDDER, Proc. Bost. Soc. Nat. Hist., vin, 9 (Hickman's
Cave, Kentucky [A. Hyatt]). PACKARD, Am. Nat., v, 745 (Mammoth Cave).

Rhaphidophora COPE, Rep. Ind. Geol. Surv., Ill, 1872, 167.

Ceuthophilus stygius SCUDDER, Bost. Jour. Nat. Hist., vn, 438. PACKARD, Mem.
Nat. Acad. Sci., IV, 1888, 70, 83 (White's, Fountain, Dixon, Diamond, and John
and Fred Field caves in Kentucky, and Wyandotte, Little Wyandotte, Bradford,
and a cave in Washington County, Indiana; One Hundred Dome and Laurel caves,
Kentucky [Sanborn]). BRUNNER, Monog. Stenop., 65 (Texas). BLATCHLEY,
Proc. Ind. Acad. Sci. , 1892, 148 (cave in Crawford County [W. P. Hay] ) . SCUDDER,
Proc. Am. Acad. Sci., xxx, 1894, 33 (localities as above). BLATCHLEY, Rep. Ind.
Geol. Surv., 1896, 198-200 (Wyandotte, Little Wyandotte, Sibert's Well Cave,
and Saltpeter Cave, Crawford County, Porter's, Truett's, and Strong's caves).

52 FAUNA OF MAYFIELD'S CAVE.

Ceuthophilus sloanii PACKARD, Ann. Rep. Acad. Sci., v, 1873, 93; Mem. Nat. Acad.
Sci., IV, 1888, 71, 83 (Bradford and Little Wyandotte caves); COLLETT, Rep. Ind.
Geol. Surv., 1873, 305 (Connelley's, Earner's, and Donaldson's caves at Mitchell
(Elrod & Sloan).

This Ceuthophilus is known from nearly every Indiana cave which
has been zoologically explored and from many of the Kentucky caverns.
I have taken it in Mayfield's, Truett's, and Donaldson's caves in Indiana
and in White's and Mammoth Caves in Kentucky. *

It is much more abundant in the smaller caves than in the larger
ones and near the mouths of caves than farther in. For some time
after this species was described it was not found in any of the large
caves. It is especially abundant not far from the mouth of Mayfield's
Cave, but is rarely found more than 250 feet from the mouth and usually
does not occur beyond the reach of twilight. Mr. Rothrock (cf.
Blatchley, 1896, 199) , who owns the hotel at Wyandotte Cave, says this
"cave cricket" is found as far back as Monument Mountain, far within
the cave, but Blatchley failed to find it there.

This shade-loving animal is not confined to caves. Blatchley tells
me of their occurrence in cellars and about wells and in similar situations,
and mentions their abundance in a cellar at the Wyandotte Cave Hotel.
I have specimens from under a log on the campus of Indiana University,
from a cellar in Bloomington, from a cellar in Switzerland County,
Indiana, and from beneath logs and chunks near the mouth of May-
field's Cave.

C. stygius is quite common in Mayfield's Cave, especially from the
mouth to "19," and is occasionally seen nearly as far in as the mound.
It is often seen in the well-lighted portion of the cave from the door to
" 4, " but is more common from?' '4" to "7" and at times is most abundant
from "7" to "12," where the cave is absolutely dark. Beyond "19,"
where the first dry portion of the cave begins, it rarely occurs and only
in the damper places. Small individuals are often found under debris
on the floor, but this locustid is most common on overhanging slopes of
the wall and on the roof of the cave. It is perhaps most common and
abundant on the walls quite near the roof. It occurs abundantly also
in crevices of the rock, often where they are so narrow that they barely
admit the body. It congregates to some extent where there is an over-
hanging slope near the floor.

It is not readily disturbed by light or by an object moving near, but
when touched springs from the surface to which it is clinging and
when it lands on the floor rapidly leaps away faster than one can follow

*It has not, I think, heretofore been recorded from Mammoth Cave. I found it
at the end of Bat Avenue near the top of the Mammoth Dome.

ORTHOPTERA — LOCUSTID^E. 53

it with a light. It will move but a dozen feet or so, however, and if
followed that far will be nearly as easy to approach as before. This
Ceuthophilus feeds upon organic matter. It was seen feeding upon the
decayed carcass of a mouse, and on several occasions was found feeding
upon cheese left as bait. It is sometimes found near decaying organic
matter of any sort.

According to evidence obtained this species seems to breed during
the fall. Adults were found in but small numbers and only during
September and October. No adults were seen farther back than "12."
Adults of this species were seen in White's Cave, Kentucky, about
November 25, 1903. Young appeared near the mouth of Mayfield's
Cave early in October and were seen from then until December and in
one case in February. No adults or very small individuals were seen at
other times than those mentioned. Hence it seems that the adults
occur in the cave during the summer and fall, and that they breed and
soon disappear, the young appearing in October and later. From the
size of individuals which I take to be a year old this species must require
at least two years and probably three years to mature. The eye was
moderately well developed, as Lawrence Durborow, who has given some
study to the structure of the eye of this animal, has found.

As is well known, ceuthophili commonly live under rotten logs in
shady woods, about wells and cellars, and in damp, dark places generally.
Ceuthophilus stygius has several cave-inhabiting relatives. C. ensifer
(Packard, 1888, 71) is found in Nickajack Cave, Tennessee; C. palmeri
(Scudder, 1894, 41) lives in Bat Cave near Georgetown, Texas, in Beaver
Cave, and in other localities of Texas; in Florida one species of this
genus lives in holes of the gopher turtle and in hollow trees; C. mexicanus
(Scudder, 1894, 82) is found in caves at San Pedro and San Lorenzo,
where it lives among mummies; Hadensecus subterraneus (Packard,
1888, 69 ff . ) , another relative, is extremely abundant in Kentucky and
Tennessee Caves; Phalangopsis annulata Bilmek (1867, 904) occurs in
Cacahuamilpa Grotto in Mexico. There are 2 genera and at least 5
species of ceuthophili which inhabit European caves.

Order APTERA.
Suborder THYSANURA.

Family LEPISMIDAE.
Machilis variabilis Say.

SAY, Phila. Acad. Nat. Sci., n, 1821, 12. PACKARD, Am. Nat., 1871, v, 746 (Mammoth
Cave). LUBBOCK, Collembola and Thysanura, 240, London, 1873.

Often seen from the door to the first turn. It occurs sometimes on
the wall, but more often on the roof. This is a common species in humid
places in the United States. This genus was recorded from Wyandotte

54 FAUNA OF MAYFIELD'S CAVE.

Cave by Cope (1872, 171) and Packard (1888, 67) and by Packard from
Mayfield's Cave (1888, 16). Packard (1888, 67) also mentions a white
species (Machilis cavernicola) from Mammoth Cave, where Hubbard
(1880, 37) found a Machilis, probably the same species. Tellkampf
(18446, 383) described and figured a Machilis from Mammoth Cave as
a crustacean ( Triura cavernicola) . An eyeless, brownish-yellow species
(Machilis bruneoflavea Joseph) is known from European caves, where
other species of this family also occur in caves (Hamann, 1896) .

Suborder COLLEMBOLA.

Family ENTOMOBRYIDAE.

Sinella cavernarum (Packard).

Degeeria cavernarum PACKARD, Mem. Nat. Acad. Sci., iv, 1888, 66, pi. xvi (Little
Wyandotte and Bradford caves in Indiana, Carter and Diamond caves in Ken-
tucky). BLATCHLEY, Rep. Ind. Geol. Surv., xxi, 1896, 200 (Mayfield's, Truett's,
Eller's, Shiloh, Clifty, Marengo, Little Wyandotte, and Wyandotte caves).
EIGENMANN, Science, n. s., xn, 1900, 302. ULRICH, Trans. Am. Mic. Soc., xxm,
1902, 97 (Ezell's Cave, Texas).

One of the most abundant forms of life in the cave. Found in all
moist situations and especially under pieces of stone or other rubbish,
or where there is organic matter. It is sure to be found wherever there
is sufficient moisture. Often seen in numbers about the edge of a small
puddle or at a wet place where the water drips from above. At these
places it is often seen upon the surface of the water, upon which it
is able to hop about with the same ease as when upon a more solid
footing. It has been seen in the loose earth at "4," but I have not
found it nearer the mouth than this, and beyond ' '4' ' it is about equally
abundant in favorable situations throughout the cave. Sinella caverna-
rum is attracted to any organic matter, having been found at decaying
wood, moldy paper, molding candle-drip, feces of mice, the remains
of a myriapod, and abundantly at bait, whether fresh or in any stage
of putrefaction. Probably it feeds upon fungus as well as upon decay-
ing organic matter, as it is nearly always present where mold exists.

This thysanuran is perfectly white in color and without traces of
eyes, as far as I can determine. It crawls very slowly, but is able to
spring for considerable distances v/hen disturbed. Young and adult
are seen at all seasons, but the species becomes scarce in winter in the
part of the cave subject to lower winter temperature. It is uneasy in
the presence of a light, but seems as likely to move toward the light as
from it.

Sinella cavernarum serves as food for Phanetta subterranea and
other spiders and mites and for the predaceous beetles of the cave.

COLLEMBOLA— ENTOMOBRYID^E. 55

This species is probably confined to subterranean life. It is so
transparent that the rays of sunlight would certainly penetrate to its
internal organs. Garman (1839, 229) quotes Miss Hoppin regarding
the occurrence of this insect, doubtless, in a well in southwestern Mis-
souri, as follows: 'The owner reports that at various times, a year or
more ago, the surface of the water would be covered with little white
lice or something of the kind. ' '

I found this species in Mayfield's, Truett's, Donaldson's, and Twin
caves in Indiana and Mammoth Cave, Kentucky.

Carpenter (1895, 30) found another species of Sinella (Sinella cav-
erniola) in Michelstown Cave in Ireland, which species he says is very
hard to seperate from Sinella cavernarum Packard. In fact, he seems
to incline to the view that the two are really the same species.

Tomocerus niger Bourlet.

BoURLET, Mem. Soc. Roy., Lille, 1839. LUBBOCK, Collembola and Thysanura,
London, 1873, 139-146.

Fairly common under stones and sometimes on the wall or roof from
the mouth to "11" and once or twice was taken as far in as " 25. " It is
steel-gray in color. It seems perfectly at home in the cave. This
species is sensitive to light, which it avoids. Mr. J. W. Folsom says it
differs but slightly from European specimens. According to the same
authority specimens collected out of doors in the immediate neighbor-
hood of the cave are almost typical as compared with European specimens.

Packard (1888, 66) found Tomocerus plumbeus var. pallidus in the
Carter Caves in Kentucky and in Weyer's, New Market, and Fountain
Caves in Virginia, and reports another variety (albida) from a small
cave in Utah (see also 1877, 159). This variety is bleached and many
of the individuals are almost white. Its eyes are small and dark colored.
Normal pallidus was found in Hundred Dome Cave and in other ' 'shal-
low" caves in Kentucky. Packard (1888, 67) has also described another
member of this family from caves, Lepidocyrtus atropurpureus from
Diamond Cave, Kentucky, and Smynthurus ferrugineus from New
Market and Weyer's caves in Virginia. Neither of these is a true cave
form.

Call (1897, 38) more recently found in Mammoth Cave Entomobra
cavicola Banks, a white eyeless species, and Smynthurus mammouthia
Banks, a white species with small eyes.

There are many European species of this family found in caves;
among them being two eyeless species of Tritomurus, closely related to
Tomocerus, and Tomocerus niveus Joseph (Hamann, 1896, 146).

Several other families of Aptera have cave representatives.

56 FAUNA OF MAYFIELD'S CAVE.

Class MYRIAPODA.

Order CHILOGNATHA.

Family BLANJULIDAE.

Cambala annulata (Say).

SAY, Phil. Acad. Nat. Sci., vol. n, 1811, 103; Entom. Works, ed. Le Conte, vol. II, 25
(Southern States). COPE, Proc. Am. Phil. Soc., 1869, 178 (Spruce Run Cave,
Virginia). PACKARD, Mem. Nat. Acad. Sci., iv, 1888, 18 and 65, pi. ix, figs. 1, la
(Wyandotte [Hubbard] and Little Wyandotte caves in Indiana, the Carter's caves
in Kentucky [Sanborn]; and Spruce Run Cave [Cope], Cave of the Fountains, and
Luray Cave in Virginia).

A single specimen of this abundant above-ground form was found
at ' 18' ' during May. This species is common through the middle and
southeastern United States, and judging by the number of times it has
been taken in caves is not averse to life in darkness.

Family CHORDEUMIDAE.
Conotyla bollmani (McNeill).

Triehopetalum bollmani McNElLL, Proc. U. S. Nat. Mus., x, 1887, 330 (Mayfield's

Cave) .
Scotherpes bollmani BOLLMAN, Proc. U. S. Nat. Mus., xi, 1888, 405 (Mayfield's,

Neeld's, Truett's and Coon's caves in Monroe County; Pitt's and Donnehue's caves

in Lawrence County, Indiana).
Conotyla bollmani COOK & COLLINS, Ann. N. Y. Acad. Sci., ix, 1896, 76 (same localities

as above). BLATCHLEY, Ind. Geol. Rep. xxi, 1896, 202 (localities as above and

Porter's Cave).

This is the abundant myriapod of Mayfield's Cave, and is the only
one which habitually reaches and stays in the remote parts of the cave.
It is very abundant at all seasons, particularly in the moist regions of
the cave. It is often found crawling over the floor, wall, or roof of the
cave, but is more abundant under stones and debris and is especially
abundant in the neighborhood of decaying organic matter. It has been
taken in twilight, but is not often seen there and seldom occurs nearer
the mouth than " 6. " It seems equally abundant in all the dark portions
of the cave where there is sufficient moisture.

This myriapod is attracted to all sorts of decaying organic matter.
It has been seen feeding upon rotten wood, moldy candle drip, decaying
banana peel, feces of mice, the decayed carcass of a mouse, and the
various bait used to attract cave life. Of the bait used the cheese and
the beef attracted the greatest numbers. The more decayed and ill-
smelling the bait became, the greater the abundance of myriapods.
This species when attracted to bait sometimes burrows into the soil and
loosens it up underneath and about the bait until it has a very loose and

CHILOGNATHA— CHORDEUMID^E. 57

granular appearance. This-loosening up of the soil has been observed
where the earth had previously been quite smooth and hard on the
surface.

This myriapod seems to breed at all seasons. It has been found
copulating a number of times. This has been observed only in the
neighborhood of decaying organic matter, and raises the question whether
abundance of food influences its breeding. Very young ones appear at
all seasons.

This is the only cave myriapod known from the region north of the
East Fork of White River, except Scytonotus cavernarum Bollman from
Mayfield's Cave, of which only the type is known. C. bollmani shows
no modifications due to cave life. Its eyes are well developed.

I have specimens of C. bollmani from Mayfield's, Truett's, Twin, and
other Mitchell caves, in all of which it is common.

Another species of this family, Pseudotremia cavernarum Cope, is
abundant in Wyandotte and other caves near; Scoterpes copei (Packard)
inhabits Mammoth and neighboring caves; while a third, Zygonopus
ivhitei Ryder, lives in caves of Virginia.

In European caves live 4 species of Attractosoma, 2 species of Cras-
pedosoma, and 1 other species. The Chordeumidas is the most important
family of true cave myriapods (cf. Packard, 1888, 60-64).

Family POLYDESMIDAE.
Euryurus erythropygus (Brandt).
BOLLMAN, Proc. U. S. Nat. Mus., xi, 1888, 407.

Found three times within the cave between "10" and "17." It was
also found in a sink-hole which is situated directly above that portion
of the cave. This species is common throughout the Middle West and
Eastern States, and is abundant about Bloomington.

Scytonotus granulatus (Say).

SAY, Phil. Acad. Nat. Sci., II, 1821, 105; Entom. Works, ed. LeConte, II, 27 (Pennsyl-
vania). PACKARD, Mem. Nat. Acad. Sci., iv, 1888, 65 (Indian Cave, Kentucky).
BOLLMAN, Proc. U. S. Nat. Mus., xi, 1888, 407. BLATCHLEY, Rep. Ind. Geol.
Surv., xxi, 1896, 202 (Little Wyandotte Cave).

Fairly common from the mouth to near "6," but has not been seen
farther in. It is usually found upon the wall or roof. Taken in Twin
Cave also. Blatchley (1896, 202) found a single specimen in Shiloh
Cave, 200 feet back. Packard (1888, 65) mentions a bleached specimen
of this species from Indian Cave, Kentucky, and another bleached one
from Lyon Cave, Kentucky. This is not a cave form, but very commonly
visits the mouths of caves. This species also was found at a sink-hole

58 FAUNA OF MAYFIELD'S CAVE.

over the cave. It is a common and abundant form about Bloomington,
and is distributed through the Middle West and Eastern States, where
it is also common.

Polydesmus serratus Say.

SAY, Phil. Acad. Nat. Sci., n, 1821, 106; Entom. Works, ed. Le Conte, 27. BOLLMAN,
Proc. U. S. Nat. Mus., XI, 1888, 407.

Often seen upon the wall or roof from near the mouth to " 18, " beyond
which it has not been seen. This species also was found at the sink-
hole above the cave. It is abundant about Bloomington and common in
the Middle West and Eastern United States.

Of the 3 species of the family Polydesmidse found in Mayfield's Cave
2 are fairly common. Scytonotus granulatus was not found far within
the cave, but it was abundant about the mouth. It has been found in
several other Kentucky and Indiana caves, in at least one of which it
was far along in the cave. Polydesmus serratus was fairly common
to "18."

Polydesmus cavicola from Clinton's Cave in Utah is described as
entirely white (Packard, 1877, 161; 1888, 20). A species of Polydesmus
has been found in Carter Caves in Kentucky, while 3 species of this
genus live in caves in southern Europe. Two European species of
Brachydesmus of this family are much modified and deserve to be con-
sidered true cave species. It is interesting to note that myriapods of
this family are eyeless. They are sensitive to light, however, and those
found in Mayfield's Cave keep as well within the limit of the influence
of outdoor conditions as eyed animals which inhabit the same part of
the cave.

Order CHILOPODA.
Family SCOLOPENDRIDAE.

Scolopendra woodi (Meinert).
BOLLMAN, Proc. U. S. Nat. Mus., xi, 1888, 409.

Taken once near the mouth, once at ' ' 23, " once at " 22, " and once
at "40." Distributed in the central and southeastern United States.
Not common about Bloomington.

Family GEOPHILIDAE.
Linotsenia chinophia (Wood).

Not seen by writer in Mayfield's Cave, but common across the entire
northern United States. One specimen was found at the mouth of
Mammoth Cave, Kentucky, February 3, 1903.

ARANEIDA— AGALENID^E. 59

ARACHNIDA.

Order ARANEIDA.

Family DICTYNIDAE.

Amaurobius beiinetti Blackwell.

BLACKWELL, Ann. and Mag. of Nat. Hist., xvn, 41 (Canada). MARX, Proc. U. S.
Nat. Mus., xii, 1889, 510.

Taken once in twilight just within the cave and once at "11." Its
distribution includes "Northeastern United States and Canada. It is
found under and among fallen leaves, under stumps, logs, etc. " (Nathan
Banks, East End, Virginia) .

Family AGALENIDAE.
Tegenaria derhami Scopoli.

SCOPOLI, Entom. Carnioli, 400. EMERTON, Trans. Conn. Ac., vm, 1890, 29, pi. 7,

fig. 6 (New Hampshire, Massachusetts, Connecticut). MARX, Proc. U. S. Nat.

Mus., xii, 1889, 516.

Araneus domesticus CLERK, Sv. Spindl. , 76, pi. 2, tab. 9, fig. 2.
Aranea domestica LINNAEUS, Syst. Nat., ed. x, I, 620.
Tegenaria ci vilis WALCKENAER, Tabl. d'Aran.; C. Koch, Die Arachn., vm, 37, figs.

618, 619. BLACKWELL, Spid. of Gr. Brit., i, 166, fig. 107; Spid. from Canada,

Ann. Mag. Nat. Hist., xvn, 76.
Tegenaria medicinalis HENTZ, Journ. Bost. Soc. Nat. Hist., v, 462; Spid. U. S., ed.

Burgess, 99, pi. 11, fig. 21; pi. 20, fig. 19.
Tegenaria derhamii EMERTON, Common Spiders, 1902, 97.

This spider is very common about the mouth of Mayfield's Cave. It
is most abundant just within the door and for about 20 feet in, but is
common at "4," is often seen past the turn of the cave at "4," and
occasionally occurs as far back as " 18. " It builds a sheet or platform-
like web with a funnel and tube in one corner. The web is usually
located in a convenient angle in the wall and the tube always, so far as
observed in the cave, leads to a hole or crack in the wall or back of a
projecting angle of the rock. The spider rests at the opening of the
tube or out upon the platform and at the slighest unusual disturbance,
as the turning of a bright light upon it or the molesting of its web, dis-
appears into the funnel. December 9, 1904, a male was seen upon the
web of a large female. Webs of this species contained remains of
numerous flies. This spider is very senitive to bright light, and if
prevented from entering its retreat may be driven from its web by light
alone.

"A common species in barns and cellars and has probably been
imported from Europe, where it is even more common." (Emerton
1902, 96.)

60 FAUNA OF MAYFIELD'S CAVE.

Found living outdoors in the immediate vicinity of the cave. T.
derhami is almost cosmopolitan in houses and cellars. This species
shows no signs of modification due to cave life, but is perfectly at home
about the mouth of the cave.

Blatchley (1896, 202) found 2 specimens of another species of this
genus (Tegenaria cavicola Banks) in Saltpeter Cave. Relatives of
T. derhami which live in European caves are Chorizomma subterranea
Simon, which has 6 transparent eyes, and Hadites tegenarioides Keyser-
ling, which is yellow in color and eyeless.

Cicurina paliida Keyserling.

KEYSERLING, Neue Spinnen aus Amerika, vil; Verb, der zool-botan. Gesellsch., Wien,
1887, 462 (42), pi. 6, fig. 26 (District of Columbia). MARX, Proc. U. S. Nat.
Mus., XII, 1889, 516.

Four specimens of this species were found in Mayfield's, two near
the mouth on the wall, one at "11" on the wall, and the third under
a stone at " 25. " This species is somewhat dull in color and its front
middle eyes are slightly smaller than the others. It was also found
under a log outside Mayfield's Cave and in Truett's Cave. Its distri-
bution includes all Northeastern United States and Canada.

Coelotes sp.

An immature one taken in November at "12." Its color was dull,
but this may have been because it was immature. Its middle eyes in
both rows were somewhat smaller than the others. Keyserling (Packard,
1888, 58) has described Ccelotes juvenilis from Kentucky caves.

Family THERIDIIDAE.
Theridium kentuckyense Keyserling.

KEYSERLING, Die Spinn. Am., Therid., I, 78, fig. 47 (Kentucky, Pennsylvania).
MARX, Proc. U. S. Nat. Mus., xn, 1889, 519.

This species is very abundant from the mouth back to "4" and is
occasionally found as far as "11." It has a small irregular web in a
depression or corner of the wall and feeds upon small flies caught in its
web. In color and in size of the eyes it appears about as is usual with
this family. Those taken into the laboratory sought the darkest part
of their place of confinement. An adult female kept alive in the labor-
atory spun a cocoon and laid eggs in it. The spider was misplaced and
forgotten for two months and when examined again the young had
hatched out, about 25 in number, and were dead in the bottle, while
the female had made another cocoon. The cocoon is a dull white, com-
pactly woven, somewhat flattened sphere, quite silky, 4.5 by 2.5 mm.

ARANEIDA— THERIBIID^E. 61

Taken in abundance from Mayfield's and Truett's Caves and also
from a cave at Glasgow, Kentucky.

Theridium porter! Banks.
Rep. Ind. Geol. Surv., xxi, 1896, 203 (Porter's and Truett's Caves).

Only 4 individuals of this species were found. Tkree females were
taken at " 11, " two at the edge of the roof and another within an angle
formed between a shelf and the wall. The last-mentioned individual
had a fairly well constructed irregular web. The others had only small
threads of web. The fourth individual was under a stone near the
same place and was without a web.

The front middle eyes in this species are black and somewhat reduced.
Of four specimens two had the front middle eyes scarcely more than
one-half the diameter of the other eyes, and in the fourth they were
about three-fourths as large.

Taken also in Truett's Cave. Described from specimens in Blatchley's
collection from Porter's and Truett's Caves.

T. eigenmanni Banks (cf. Ulrich, 1901, 97) is another species of
this genus from caves. It comes from San Marcos, Texas, and is some-
what dull in color.

Nesticus carter! Emerton.

EMERTON, Am. Nat., ix, 279, pi. I, fig. 28 (Carter Cave, Kentucky). PACKARD, Mem.
Nat. Acad. Sci., IV, 1888, 16, 57, 87, pi. 15, fig. 28 (Carter Cave, Kentucky). MARX,
Proc. U. S. Nat. Mus. xn, 1889, 521. BLATCHLEY, Rep. Ind. Geol. Surv., xxi,
1896, 204 (Porter's, Coon, and Marengo Caves).

A single specimen was taken at "25" under a stone. Habits like
P. subterranea and E. infernalis. Taken in Truett's Cave also. Its
color is very light gray. The front middle eyes are quite small and
black; the other eyes are possibly a little less than the usual size.

Packard (1888, 57) obtained another species of this genus (Nesticus
pallidus Emerton) in Fountain Cave, Virginia. It was light in color
and like N. carteri had the front middle eyes black and reduced in size.
Still another (Nesticus cordatus Bilimek) is described from Cacahuamilpa
Cave in Mexico (cf. Packard, 1894, 732).

In European caves occur three other species of this genus, two of
them confined to caves. Those confined to caves are dull in color and
show some decrease in size of the eyes, in one of them the front middle
eyes being absent and those still present barely discernible.

Argyrodes trigonum Emerton.

EMERTON, Trans. Conn. Ac., vi, 23, pi. 5, fig. 1. MARX, Proc. U. S. Nat. Mus., xn,
1889, 524. EMERTON, Common Spiders, 1902, 124, figs. 292-296 (New England).

Taken twice within the caveat "2." Common in New England
and all Eastern United States.

62 FAUNA OF MAYFIELD'S CAVE.

Linyphia marginata Koch.

HERR-SCHAEFF, Deutschl. Ins., 127, 21, 22. EMERTON, Trans. Conn. Ac., vi, 61, pi.

18, fig. 1 (New England). KEYSERLING, Die Spinn. Am., Therid., n,58, fig. 164.

KOCH, Die Arachn., xn, 118, figs. 1041, 1042. MARX, Proc. U. S. Nat. Mus., xn,

1889, 528.
Linyphia marmorata HENTZ, Jour. Bost. Soc. Nat. Hist., VI, 29; Spid. U. S., ed.

Burgess, 133, pi. 15, fig. 5.
Linyphia scripta HENTZ, Jour. Bost. Soc. Nat. Hist., vi, 24, 134, pi. 15, fig. 6.

Often seen inside the mouth of the cave, where its webs and young
occurred, but not found beyond " 2. " A very common spider, living in
shady woods and widely distributed in Europe and the United States.

Linyphia nigfrina Westring.

WESTRING Forteckning ofver till narrvarande tidkanda, 1851, 38 (New Hampshire,
Massachusetts, Rhode Island, Maryland, District of Columbia). MARX, Proc.
U. S. Nat. Mus., xil, 1889, 528.

Linyphia pulla BLACKWELL, Spid. of Gr. Brit., n, 234, fig. 156.

Diplostyla nigrina EMERTON, Trans. Conn. Acad. , vi, 65, pi. 20, fig. 2 (New England) .

Found in twilight near the door. Three taken at different times
suspended by single webs. This species commonly occurs under leaves,
etc. , outside of caves.

Willibaldi cavernicola Keyserling.

KEYSERLING, Die Spinn. Am., Therid., II, 123, fig. 204 (Reynolds Cave, Kentucky) .
PACKARD, Mem. Nat. Acad. Sci. IV, 1888, 58, pi. 15, fig. 32 (Reynolds Cave, Ken-
tucky). MARX, Proc. U. S. Nat. Mus., xn, 1889, 531.

Not found in Mayfield's Cave, but 2 specimens were taken in Don-
aldson's and 3 at Twin caves at Mitchell. It is a true cave form. The
colors are dull and the eyes very small. Of the three specimens in my
collections, one has eight eyes, all of which are quite small and colorless;
the middle pair in the back row are largest and the middle pair in the
front row next in point of size, while the side eyes of the front row are
difficult to see even with considerable magnification and the lateral
posterior ones are scarcely discernible. The second specimen has but 6
eyes visible, the sizes ranging as before and the ones missing being the
lateral posterior eyes. In the third specimen the eyes range in size as
in others, while one and possibly both of the lateral posterior eyes are
missing, and the lateral anterior eyes are barely visible. I have not
seen Keyserling's description, but from his figure, which Packard (1888,
plate xv, fig. 32) has copied, the eyes are small, and the front middle
ones extremely minute. Willibaldi (Linyphia) incerta Emerton (1875,
280) from Kentucky caves has small, colorless eyes, while the front
middle pair is usually lacking.

ARENEIDA— THERIDIID^E. 63

Porrhomma myops Simon, known from a cave in southern France
and another cave in Ireland, Carpenter (1895, 59) thinks, is identical
with Packard's Willibaldi incerta. Certainly, then, a close similarity
exists.

Phanetta subterranea (Emerton).

Linyphia subterranea EMERTON, Am. Nat., ix, 1875, 276, 279, pi. 1, figs. 29-31 (Carter

Caves, Kentucky, and Wyandotte Cave); Mem. Nat. Acad. Sci., iv, 1888, 16,

56, 57, pi. xv, figs. 29, 30, 31.
Phanetta subterranea KEYSERLING, DieSpinn. Am., Therid., n, 125, fig. 205. BLATCH-

LEY, Rep. Tnd. Geol. Surv., xxi, 1896, 204 (Wyandotte Cave). MARX, Proc.

U. S. Nat. Mus., xil, 1889, 531.

Very abundant in all parts of the cave. The most abundant arachnid.
It is invariably found under stones or debris where there is considerable
moisture. It sometimes has a few short strands of web. It is often
attracted to decaying organic matter and is nearly always associated
with the thysanurans. It is quite possible that the latter are attracted
to the organic matter and that the spiders are attracted by the thysan-
urans. It is quite sluggish in its movement, except when springing
upon its prey. In February, 1904, I turned over a damp rock at "23,"
on the under side of which were dozens of thysanurans and two of these
spiders. The larger spider jumped towards a thysanuran, which sprang
away and escaped, but with a second quick movement the spider caught
another, clutching it firmly, and did not release its hold even when
pursued, until it was picked up. This species probably feeds almost
entirely upon the thysanurans, but I found one which appeared to be
eating at a bit of decayed cheese. A myriapod found entangled by a
single strand of web of this spider may have served as food. This spider
often forms a small web of a few short, irregularly arranged strands,
which it attaches to the debris under which it lives and to the rock or
soil beneath.

The negative phototropism of this species is very pronounced. Indi-
viduals placed in short bottles in a dark room into which a little light
was admitted remained in the end of the bottle away from the light.
One experimented with in the cave behaved as follows: It moved
actively but irregularly for a moment after the stone was removed from
over it. Then when bright light was thrown upon it, went directly
away from the light. It was made to turn back and forth three times
by changing the direction from which the rays of light struck it, turning
squarely from the light in each case. After having been molested so
much that it seemed bewildered, it moved rather irregularly and with-
out definite direction until it accidentally ( ?) went away from the light,
when it kept moving in that direction. During all this time it was

64 FAUNA OF MAYFIELD'S CAVE.

stimulated by light alone, for when it got in the shadow of a small clod
it ceased to move. It is very susceptible to the influence of light, mov-
ing rapidly when the bright light is upon it and slowly or not at all
when it gets out of reach of the bright rays. Mechanical stimulus does
not cause it to move rapidly very long; it is just as effective in starting
the spider for a moment, but must be repeated in order to continue
effective. Light stimulus is effective in keeping the spider in active
motion as long as the stimulus is kept up, but as soon as discontinued
the effect decreases and disappears.

The cocoon of this species is white, disk-shaped, and thickened in
the middle, and 3 or 4 mm. broad. It is attached to the under side of
a stone, to a piece of wood, or to the ground under debris. It is usually in
a slight depression which serves to shape one side of the disk. It is a
well-made cocoon, and while the covering of web is not very thick it is
compactly and smoothly made. The eggs are rather large and are few
in number. One cocoon examined contained but 2, another 3, another
5, and another six eggs. I have kept the cocoons and hatched out the
young spiders. The young are very light in color; abdomen white;
cephalothorax and legs very light gray; sternum darker with light mar-
gin; length when first seen 0.7 mm. The young which were hatched
out and kept in a small bottle spun a few threads across the bottle and
rested upon them.

In Emerton's (1875, 279) specimens the colors were quite dull and
the front middle eyes were small. In those from Mayfield's and Truett's
caves some variation was noticeable in the ordinarily dull color, some in-
dividuals having legs and mandibles dark reddish-brown, others, includ-
ing a few adults and the immature ones, having little or no yellow or
reddish-brown anywhere. In individuals with the legs darkly colored
the body was also darker, but in most cases the body was very light or
even nearly white. There is considerable variation in the size of the
eyes and especially of the front middle ones which are often entirely
lacking. Twenty-seven individuals were studied in detail regarding
this point. In 2 of these the front middle eyes were about half the size
of the others; in 4 they were somewhat less than half the diameter of
the others; in 9 they were very small but fairly distinguishable; in 6
there was a dark blotch from which these eyes could be distinguished
with difficulty or not at all; and in 6 these eyes were missing altogether,
their location either being marked by a dark blotch or without any
indication of eyes or pigment.

This species is not known to occur outside of caves and probably is
confined to caverns. I have taken it in Truett's Cave and have little
doubt that it is common in Indiana caves, although so small that it has
usually escaped notice.

ARANEIDA— THERIDIID^. 65

Erigon« infernalls Keyserling.

KEYSERLING, Die Spinn. Am., Therid., 11, 180, fig. 239 (Reynolds Cave, Kentucky).
PACKARD, Mem. Nat. Acad. Sci., iv, 1888, 58, pi. 15, fig. 33. MARX, Proc. U. S.
Nat. Mus., XII, 1889, 534.

Fairly common within the cave. Twice found in pairs, a male and
female being under the same stone. Three times this species was found
with a fairly good but irregular web in an angle of the wall. It has
been taken in twilight as near the mouth as " 3, " but when in twilight
has always been found under stones and without a web. It seems more
at home in absolute darkness, where in every case but one it has been
found to have a web in an angle of the wall, under the edge of a rock
or in some such place. In one case two were found at an old bait which
had been left under a stone. It has a small web and may feed upon Dip-
tera, but probably also feeds upon the thysanurans and decaying organic
matter. It seems quite possible that this species is attracted to the bait
by the abundance of small flies and thysanurans which congregate there
rather than by the bait itself.

This species has the dull color and retiring habits of a typical cave
creature. Three individuals examined show various degrees of devel-
opment of the eyes. In one (adult female) , the eyes were about normal
and the front middle eyes were about three-fourths the diameter of the
others; in the second (adult male) the eyes were all slightly reduced in
size and the front middle eyes were about two- thirds the others; while
in the third (adult female) all were much reduced in size and the front
middle eyes were very small. Keyserling's drawing, reproduced by
Packard (1888, plate XV, fig. 33) shows all the eyes reduced in size and
the front middle ones about two-thirds the diameter of the others.

This species is not known to occur outside of caves. Taken in Twin
Cave at Mitchell.

A new Theridid, probably a new genus, was taken near the mouth.
Mr. Nathan Banks, to whom it was sent, has not yet described it.

The family Theridiidae includes the great majority of cave species
of true spiders (Araneida). This is an immense group and includes
many species which live in shady woods, in dense vegetation, under
leaves and logs, and about wells and cellars. It is not surprising that
such a group frequents caves and that some of its species become cave
inhabitants. Nine out of 16 of the true spiders of Mayfield's Cave
belong to this group, while of the 8 species which are permanent
cave residents 5 belong to this group. From other Indiana caves are 2
more species of this group and from other American caves are 5 more.
Of the total of 15 cave-inhabiting American species of this group, 7 are

66 FAUNA OF MAYFIELD'S CAVE.

not especially dull in color and have eyes about normal in size, while
the other 8 are more or less dull and have eyes with a greater or less
degree of degeneration.

Hamann (1896, 195-203) lists 19 species of this group from European
caves. Of these all but one are more or less dull in color and 16 show
decided reduction in the size of the eyes, while 5 of them have the eyes
greatly reduced.

In the Mayfield's Cave species of this group there is a direct relation
between the species inhabiting perpetually total darkness or visiting
twilight, and the degree of the degeneration of the eyes and the loss of
color. Theridium kentuckyense lives only near the mouth and usually in
bright twilight; its eyes and color show no modification from that of the
ordinary Theridium. Theridium porteri lives in dim twilight or just
beyond in darkness; its eyes are slightly reduced and its color is some-
what dull. Nesticus carteri is found well within the cave and is much
duller; its eyes are more reduced. Phanetta subterranea lives in the
depths of the cave; its color is very dull and its eyes are reduced and
part of them often lacking.

Family EPEIRIDAE.
Meta menardi (Latreille).

Aranea menardi LATREILLE, Hist. Nat. d. Crust, et de Ins., VII, 266 (Massachusetts,
Kentucky, Virginia, District of Columbia).

Metafusca C. KOCH, Die Arachn., vill, 118, figs. 685-687.

Epeirafusca BLACKWELL, Spid. of Gr. Brit., n, 349, fig. 252 (Great Britain).

Meta menardi THORELL, Rein, on Synon. of Eur. Spiders, 38. SIMON, Arachn. de
France, I, 151. EMERTON. Trans. Conn. Ac., vi, 328, pi. 34, fig. 18 (New Eng-
land). PACKARD, Mem. Nat. Acad. Sci., iv, 1888, 11, 57 (Dixon'sCave, Kentucky).
MARX, Proc. U. S. Nat. Mus., xn, 1889, 551. McCooK, Am. Spiders, III, 11.
BLATCHLEY, Rep. Ind. Geol. Surv., xxi, 1896, 203 (Mayfield's, Strong's, Donne-
hue's, Donaldson's, Clifty, Wyandotte, and Saltpeter Caves) . EMERTON, Common
Spiders, 1902, 190, figs. 443-445.

Not at all common, my only record in Mayfield's Cave being made
from 2 immature specimens. Found to be extremely abundant in the
caves at Mitchell. In Donaldson's and Lower Twin Caves it is excess-
ively abundant near the mouth, where the light is rather dim. It is
also found in darkness occasionally, but I have never taken it farther
than 200 feet from the mouth of a cave. This spider always spins a
characteristic somewhat loose and irregular orb-shaped web. I have
several times seen the large female resting in the middle of the orb-
shaped web and the much smaller male on the wall just at the edge of
the web. Flies are sometimes found entangled in the web of this species
and in a mass of debris taken from one of these webs I recognized the
remains of several Leria and one or two beetles, together with parts of
a spider cocoon, probably a discarded one of its own.

ARANEIDA— EPEIRID^E. 67

Blatchley (1896, 203) found this species to be the most abundant
spider of Indiana caves. He took 2 or 3 immature specimens from
Mayfield's Cave. According to Emerton (1902, 190) this arachnid "lives
in caves and similar cool and shady places in various parts of this country
and also in Europe."

Theridiosoma gemmosum Koch.

KOCH, Verzeichn. d. bei Niirnberg beob. Artcn., 69 (Pennsylvania, Illinois, Massa-
chusetts, Connecticut). KEYSERLING, Die Spinn. Am., ThericL, I, 218, fig. 131.
MARX, Proc. U. S. Nat. Mus., xn, 1889, 551.

Theridiosoma argenteoium, CAMBRIDGE, Ann. Mag. Nat. Hist., 1879, 194.

Microepeira radiosa EMERTON, Trans. Conn. Ac., vi, 302, pi. 24, fig. 7 (Pennsylvania,
Ohio).

Four individuals of this species were taken within the cave. All
were found during the winter and all were near the door except one
which was found suspended by a single thread of web at " 10."

Family TETRAGNATHIDAE.
Tetragnatha laboriosa Hentz.

HENTZ, Journ. Bost. Soc. Nat. Hist., vi, 27. Spid. U. S., ed. Burgess, 131, pi. 15,
fig. 3 (Massachusetts, Connecticut, District of Columbia, Maryland, Virginia,
Ohio, Utah, Nebraska, Alaska). EMERTON, Trans. Conn. Ac., vi, 334, pi. 39, figs.
7, 8, 11. MARX, Proc. U. S. Nat. Mus., xn, 1889, 552. EMERTON, Common Spiders,
1902, 202, fig. 463.

Not seen by writer in Mayfield's Cave, but found occasionally near
the mouth of Donaldson's and Twin caves at Mitchell.

Family LYCOSIDAE.
Dolomedes scriptus Hentz.

HENTZ, Jour. Bost. Soc. Nat. Hist., v, 189; Spid. U. S., ed. Burgess, 38, pi. 6, fig. 1
(Alabama). MARX, Proc. U. S. Nat. Mus., xn, 1888, 566.

Fairly abundant in fall and winter from near the mouth to "5."
These spiders appear in the cave by early fall and stay until early spring.
I have not found them in spring or summer and think they merely
hibernate in the cave. They are not very active in the fall and during
the winter are quite sluggish. One very large female was observed to
have moved less than a foot during two months of winter. Another
remained in nearly an identical spot throughout the winter. They are
found on the roof or wall and have no web. I found this species hiber-
nating under logs in the ravine below the mouth of the cave. This
spider lives habitually along the margins of streams under stones, and is
known from Alabama and the Eastern United States.

68 FAUNA OF MAYFIELD'S CAVE.

Order ACARINA.

Family EUPODIDAE.

Rhagidia cavicola Banks.

Bryobia weyerensis PACKARD, Mem. Nat. Acad. Sci., iv, 1888, 42, pi. xi (Weyer's

Cave, Virginia).
Rhagidia cavicola BANKS, Am.;Nat., xxx, 1897, 1382, pi. x, fig. 3 (Mammoth Cave).

Seen quite often and probably is fairly abundant although incon-
spicuous. It is about 1 mm. long. It is perfectly white and shows no
trace of eyes. In color, size and general appearance it very much
resembles the young of the thysanuran Sinella cavernarum. This mite
occurs in all parts of the cave in loose earth under debris or at decaying
organic matter where there is considerable moisture, and usually in
company with Sinella, for which I at first mistook it. It probably preys
upon this thysanuran and upon eggs and larvae of other cave animals.
Packard, according to Mr. Nathan Banks, took this species (probably) in
Weyer's Cave, Virginia. I have little doubt that it is generally distrib-
uted in Indiana caves, but has been overlooked because of its small size
and resemblance to Sinella.

Other species of this genus live under dead leaves and in loose moist
soil. Another species of this family (Linopodes mammouthia Banks) is
described from Mammoth Cave, where it occurs in moist dirt (Call, 1897,
p. 382). Banks (1904, p. 13) says of this family of mites:

Most inhabit the ground, but some are found on the leaves of trees. All are
predaceous and feed on various small insects or insect eggs. They seem to delight in
cold, damp places, and can be found in winter still active [among and under fallen
leaves. They are among the most common acarians in high altitudes and are also
frequent in caves, both of this country and of Europe, where their simple and primi-
tive structure is well suited to the conditions.

Family TYROGLYPHIDAE.
Tyroglyphus sp.

The Hypopus or migratorial stage of some Tyroglyphus occurred in
extreme abundance upon one of the flies in the cave (Leria latens) .
Specimens were sent to Nathan Banks, who replied as follows:

The mites are the Hypopus or migratorial stage of some Tyroglyphus— it is
impossible to tell what species. Probably it is new. Only a few species of Tyrogly-
phus are described from United States, and only one or two of these have been
connected with their Hypopus. The mites do not feed on fly.

At times the mites are to be found upon nearly every Leria latens
taken and occasionally 30 or 40 are seen clinging to a single fly. The
mites are especially abundant upon the abdomen, but attach themselves
to all parts of the body of the fly. While so very abundant on Leria

ACARINA— TYROGLYPHID^. 69

latens, I did not observe this mite upon any other species, although several
other Diptera mingled with L. latens and to such an extent that several
species are often taken near the same spot. Dozens of Leria defessa,
Limosina tenebrarum, and other species were closely examined without
finding a single one of these parasites.

Family GAMASIDAE.

Two or three times bats were seen with large mites, possibly of the
family Gamasidae, clinging to them. The mites were fastened to the
naked portions about the head of the bat, and in one case the mite's
abdomen was greatly distended and red from the blood it contained.

Carpenter (1895, 30) found Gamasus attenatus, a common British
form, among dead leaves, etc., in Michelstown Cave. Absolon (1900, 3)
mentions 9 blind Gamasida?. By experiment he found that Gamasus
niveus dies in a few minutes on exposure to daylight (1900, 4).

Hamann (1896, 215-223) lists 19 Acarina from European caves,
some of which are blind and without pigment.

CRUSTACEA.
Order PODOPTHALMATA.

Family ASTACIDAE.
Cambarus pellucidus testii Hay.

Cambarus pellucidus, PACKARD, Mem. Nat. Acad. Sci., IV, 1888, 16 (May field's Cave).

FAXON, Proc. U. S. Nat. Mus., xn, 1890, 621 (Mayfield's Cave).
Cambarus pellucidus testii, HAY, Proc. U. S. Nat. Mus., xvi, 1893, 283, fig. (Mayfield's

and Truett's Caves) ; Rep. Ind. Geol. Survey, XX, 1895, 484; the same, xxi, 1896,

209 (Mayfield's Cave). HARRIS, Kan. Univ. Sci. Bull., n, 1903, 112 (localities as

above) .

This subspecies of Cambarus pellucidus (Tellkampf ) * was described
by W. P. Hay, who bases the distinction principally upon the smoothness
(lack of spines) of the individuals from Mayfield's Cave as compared
with specimens from the more southern Indiana and from the Kentucky
caves. The subspecies is known from this cave and Truett's Cave only,
while the parent form is known from most of the Indiana and Kentucky
caves in which there are pools or streams. It is quite abundant in the
pools of Mayfield's Cave.

The subspecies is not clearly defined. All the specimens of Cambarus
pellucidus from Kentucky which I have seen have the spines on the
rostrum well developed and the tendency to spininess on the side of the

*Cambarus pellucidus was described as Astacus pellucidus Tellkampf, Arch.
Anat., Physiol. u. Wissensch. Med., 1844, 383 (Mammoth Cave).

70 FAUNA OF MAYFIELD'S CAVE.

carapace very marked; but of a number of specimens from King's Cave
at Corydon, Indiana, two are decidedly smoother than the usual type,
even approaching C. pellucidus testii, while the others are nearly as
spiny as Kentucky specimens. An occasional individual from the caves
at Mitchell is as smooth as the average from Mayfield's Cave and some
individuals from Mayfield's show some tendency toward rostral spines,
while a single specimen taken in Mayfield's was fully as spiny as the
average individual from Mitchell. In the main the distinction holds,
but the subspecies C. pellucidus testii from Mayfield's Cave intergrades
with C. pellucidus from Mitchell. Farther south one finds the increase
in number and length of spines, together with the appearance of less
conspicuous characters, very marked, and one would not hesitate to say
specimens from Kentucky belong to a species different from those living
in Mayfield's Cave if intermediate forms were not known.

Eigenman (1903, 169, footnote) says of Cambarus pellucidus from
the Mitchell caves: "We have found that the specimens of Cambarus
pellucidus from these caves have an eye structure much more degenerate
than specimens of the same species from Mammoth Cave." I have
made no examination of the structure of the eye in C. pellucidus testii,
but it seems probable that the greater degeneration noted in the eyes
of the specimens from Mitchell is also present in the individuals from
Mayfield's Cave and that the increase in amount of degeneration accom-
panies the decrease in spines so that individuals from Corydon, Indiana,
would have eyes intermediate between the eye of C. pellucidus from
Mammoth Cave and C. pellucidus from Mitchell, Indiana.

This crayfish is usually seen quietly resting on the bottom of a pool.
Rarely one is observed walking slowly. When roughly disturbed it acts
and swims much as other crayfish do when excited; that is, it swims
without regard to the edge of the pool or even the direction of the
danger and is as likely as not to swim into one's hand or out upon the
bank. However, if there is a disturbance of the water and the crayfish
becomes aware of the pursuer while at a distance or before being
touched, it, in nearly every case, swims or crawls toward protecting
rocks shelving over the edge of the pool, or to some other such means
of concealment. If there is no such protection it moves toward the
opposite side of the pool. It occasionally retreats to a hole under a rock.
C. bartoni is quite often found in such holes. Possibly C. bartoni alone
forms these holes and C. pellucidus makes use of them when deserted
by C. bartoni. Often when slightly disturbed by an object close at
hand C. pellucidus backs off, then turns around, and crawls forward.
However, when disturbed it usually starts to swim immediately. If
crowded when crawling forward toward a place of concealment, it begins

PODOPTHALMATA— ASTACID^E. 71

to swim caudal end foremost without first turning about and really
swims toward its pursuer. But the moment it begins to swim it either
turns squarely over ventral side up or turns to one side so as to move
in the direction in which it was crawling. After changing the direction
of its course it rights itself and soon disappears if there is ready means
of concealment. In its swimming and crawling motions this crayfish
is not less active than other crayfish.

This species is sensitive to a jar in the water at a distance of several
feet if the disturbance is quite pronounced, like that produced by drop-
ping a pebble into the pool. But considerable rippling or slow swishing
about in the water often fails to produce any effect upon individuals at
a little distance. It seems insensible to sound, although a heavy jar on
the bank of the pool may cause it to move. Light often fails to have
any apparent effect, but on two occasions when a bright light was sud-
denly flashed upon perfectly quiet individuals they moved immediately,
swimming rapidly from the lighted area. In these two cases there
could have been no jar or other disturbance, for I had quietly crept to
near the individuals from a distance and then suddenly thrown the light
full upon them. Sometimes when the light was held upon individuals
for several minutes they failed to respond at all; usually, however, they
moved after two or three minutes.

From observations upon C. pellucidus testii which Dr. Charles Zeleny
kept in the laboratory, I believe they molt fronf two to four or five times
a year, depending upon the size, the smaller or younger ones molting
oftener. Putnam (18756, 18) kept a specimen of C. pellucidus from
Mammoth Cave for six months and it molted twice during that time.
Just after molting C. pellucidus is very transparent, so that it is a
veritable chart of crayfish anatomy, the nerve-cord and ganglia and
alimentary canal showing through the body wall in detail. In one which
had molted two or three days previously the "stomach mill" was seen in
full operation while it was eating a bit of beef. This crayfish is ordi-
narily white, its body wall becoming more or less opaque soon after
molting. Large individuals which evidently have not molted for some
time become quite dark or dirty rusty in appearance from the collection
of dirt upon the shell.

As to the food of C. pellucidus testii within the cave, very little has
been found out. In captivity it will eat flesh of almost any animal. It
does not thrive as well on beef, however, as does C. bartoni, nor does it
eat as much. In the cave its food must be very scanty. It certainly
could not catch a blind fish, and it seems scarcely likely that it would
ever be able to catch the relatively small and active amphipods. The
isopods are less active, but are very small to serve as food for so large

72 FAUNA OF MAYFIELD'S CAVE.

an animal, which at best could probably catch very few of them. While
some decaying organic matter is being continually brought into the
cave through sink-holes, it is carried in in quantity only at time of high
water.

Putnam (1875, 16) records the following observations as to the taking
of food by Cambarus pellucidus.

The iblind species * * * darts back as soon as the food is dropped into the
water and then extends its antennas and stands as if on the alert for danger. After
a long while, sometimes from fifteen to thirty minutes, it will cautiously crawl about
the jar with its antennae extended as if using them for the purpose of detecting
danger ahead. On approaching the piece of meat, and before touching it, the animal
gives a powerful backward jump and remains quiet for a while. It then cautiously
approaches again, and sometimes will go through this performance three or four
times before it concludes to touch the article, and when it does touch it the result is
another backward jump. After another quiet time it again approaches, perhaps only
to jump back once more, but when it finally concludes that it is safe to continue in the
vicinity of the meat, it feels with its antennas for a while and then takes the morsel
in its claws and conveys it to its mouth.

I have made practically the same observations and have noted that
individuals kept in capitivity for some time may become less wary in
taking food.

Very young individuals were seen during February and March, the
earliest date being February 17. I saw quite young ones at no other
time of the year. I did not observe copulation nor see the females
with eggs.

Packard (1888, 110-112) found that the eye of C. pellucidus had lost
' 'the facets and entire cornea, the cones and rods, and the three black
pigment layers/' that the "integument of the eye, with its stalk, is
present but abnormally lessened in size," and that "the optic nerves
and ganglia are present, but the latter are small and degenerate."
Parker (1890) , on the other hand, describes the eye as degenerate, but
as not having degenerated far enough to lose the cone cells. Besides
its degenerate eye this species possesses other well-known characters
adapting it to cave life. It is without black pigment, its body is slender,
its legs are long and slender, and it has quite long and delicate antennae.
Further, in common with other cave crayfish, its antennal scale has
become very broad.

Another blind crayfish, Cambarus setosus Faxon (1889, 625), comes
from the caves of southwestern Missouri. This species has degenerate
eyes— more degenerate than those of C. pellucidus according to Parker
(1890), less according to Faxon (1889, 628) —is white in color, has slender
body and appendages, and a wide antennal scale. Cambarus hamidatus

PODOPTHALMATA— ASTACID.E. 73

Packard (1888, 40) from Nickajack Cave, Tennessee, has the same
general characters as the other cave species mentioned, though this one
is less slender in body than C. pellucidus.

Cambarus acherontis Lonnberg (Faxon, 1898, 645) is a subterranean
species found in caves and underground streams in Florida.

Cambarus barton! (Fabricius).

(?) Astacus bartonii, FABRICIUS, Suppl. Entom. Systemat., 1798, 404.

Cambarus bartoni, HAGEN, 111. Cat. Mus. Comp. Zool., ill, 1870, 75 (Mammoth Cave).
PUTNAM, Proc. Bost. Soc. Nat. Hist., xvn, 1874, 222 (Mammoth Cave). SMITH,
Am. Journ. Sci. and Arts, ix, 1875, 497 (Mammoth Cave). HUBBARD, Amer.
Ent., ill, 1880, 38 (Mammoth Cave). FAXON, Mem. Mus. Comp. Zool., x, No. 4,
1885, 41, 59 (Mammoth Cave). PACKARD, Pop. Sci. Mon., xxxvi, 1890, 393 (Mam-
moth Cave). HAY, Rep. Ind. Geol. Survey, XX, 1895, 487 (May's [?] Cave,
Down's Cave, Connelley's Cave, a cave near Paoli, and other localities) ; the same,
xxi, 1896, 210 (Strong's, Clifty, Donnehue's, and Spring caves). FAXON, Proc.
U. S. Nat. Mus., XX, 1898 (caves in Lawrence and Orange Counties). HARRIS,
Kan. Univ. Sci. Bull. , n, No. 3, 1903, 72 (cave localities as above) .

This species is common throughout the central and eastern United
States and is one of the most common of crayfishes in this part of Indiana.
It is most abundant in running water, especially in the smaller streams,
in the neighborhood of springs, and about shallow, rocky pools. It finds
its way to the very headwaters of the smallest streams, where it is
found in short burrows under rocks or under the bank.

Its distribution in caves seems to be quite general. I have found it
common in the Twin and Donaldson's Spring caves at Mitchell, Indiana,
where it is most abundant in twilight near the entrances, although
often found well within the caves. It is extremely abundant between
the Twin Caves, where a stream comes out of one cave, flows as an open
stream for about 50 yards, and then becomes a subterranean stream
within the other cave. The stream is fairly swift and cold, and its bed
is strewn with pieces of limestone, so that the locality is quite favorable
for this species. On several occasions as many as 200 large individuals
were collected at this place for class use at Indiana University. These
included the largest and finest individuals of C. bartoni I have ever
seen, some of them measuring over 100 mm. in length. C. bartoni
tenebrosus Hay (1902a, 232) from Mammoth Cave becomes larger. I
have a specimen of it 138 mm. long. This variety is also found in Horse
Cave and a cave near Glasgow, Kentucky. C. bartoni is reported from
a vast majority of the Indiana and Kentucky caves, where C. pellucidus
occurs, and is reported from Virginia and West Virginia caves, where
C. pellucidus is not found.

74 FAUNA OF MAYFIELD'S CAVE.

C. bartoni is occasionally seen in Mayfield's Cave, but is not very
abundant. It is found resting or crawling on the bottom of the pools
or, as is often the case, under a stone in the water, where it usually
constructs a burrow. Young C. bartoni in abundance appeared within
the cave about April 1 in 1905.

As W. P. Hay says of this species:

* * * Seems to take very kindly to a subterranean abode when the opportunity
is afforded (1896, 210).

Its habits throughout the range are, so far as I know, practically uniform unless
conditions are such as to preclude the customary mode of living. It is a frequenter
of cool streams, where it lives under flat rocks or in holes which it excavates among
the pebbles. * * * In its effort to secure its favorite conditions of water,
temperature, etc., it is led to ascend streams, and although this ascent is doubtless
made slowly and the attempts at ascent are of ten stopped or seriously checked by exten-
sive rapids or heavy floods, it is nevertheless almost a certainty that through this habit
the animal has gone to the very headwaters of many a mountain stream and in favor-
able seasons has crossed the divide and reached the source of some other streams.
As in Virginia, West Virginia, Kentucky, Tennessee, and Indiana many of the small
streams have their source in cave streams, it is easy to understand that C. bartoni
is of common occurrence in the caverns of the region (1895, 487).

I think this species does not take less favorably to cave life than to
out-door life. It could readily escape from Mayfield's Cave by following
the downward course of the stream during winter or spring or after any
heavy rain. In none of the caves where I have seen it does there seem
any probability that it is cut off from escape. There is good reason to
believe that it not only does not attempt to escape from caves, but that
it has remained in Mayfield's Cave for long periods of time.

A series of C. bartoni of all sizes was collected about Bloomington
and many measurements (length of body, of each antenna, of anten-
nules, of carapace, of the various appendages, width of carapace, of
areola, length and width of rostrum, etc. ) of the individuals were taken
and compared with similar measurements of individuals of a small
series from Mayfield's Cave. The results were plotted on coordinate
paper. It was ascertained that, when length of body and length of
antennas were compared, the curves showed a much greater length of
antennae for the cave forms; 58 specimens of terranean C. bartoni were
in Series I and 6 were in Series II from the cave. It was desired to
make both series larger, but more individuals of Series I could not be
obtained at the time and the Mayfield's Cave specimens were not often
found, so that after the collection of them was attempted but 6 with
complete antennas were secured. In Series I the antennas averaged
92.66 per cent of the length of the body and in Series II, 104.55 per

PODOPTHALMATA— ASTACID^E. 75

cent of the length of the body. Hence the Mayfield's Cave individuals
have antennae 11.89 per cent longer than individuals of the same species
from outside of caves in the immediate vicinity. This means that
Cambarus bartoni in Mayfield's Cave has acquired a constant character
different from C. bartoni above ground in the same locality.

There is also a noticeable difference in pigmentation in the two series,
the cave series being lighter. This difference in pigmentation is not
great, but it is fairly constant. There was no noticeable difference in the
size of the eyes nor in any other characters, although the rostrum length
seemed possibly a little greater in the cave specimens. These changes in
Cambarus bartoni have occured while the individuals might have followed
the course of the stream and escaped. Evidently individuals of this
species have voluntarily and by preference remained in the cave, where
they have become modified so that a local race has been formed.

A third series of 35 specimens from the stream which runs between
the Twin Caves, and from about the mouths and within the Twin and
Donaldson's Spring caves, was measured. This series had antennae
100.95 per cent the length of the body or 8.29 per cent longer than the
average of above-ground forms about Bloomington. It seems quite
possible that in this case the increase in length of the antennae is due
to the influence of a partial cave life.

In C. bartoni tenebrosus Hay the antennae are longer than in typical
C. bartoni. The other distinguishing characters of C. bartoni tenebrosus
do not exist in specimens from Mayfield's Cave.

The possibility of a correlation in certain cases between life in caves
and increased size suggests itself. Unfortunately the small series of
C. bartoni from Mayfield's Cave contained only individuals of aver-
age size even for typical C. bartoni, but the individuals from Mitchell
often reach more than 100 mm. in length, while individuals of C. bartoni
tenebrosus more than 100 mm. in length are common, and I have one
which measures 135 mm. I have never seen C. bartoni, living away
from caves, over 84 mm. in length; 70 mm. in length is the usual maxi-
mum attained.

This same increase in size is noted in Crangonyx gracilis from May-
field's Cave as compared with the same species outside of caves. The
greater size attained by these two species when living in caves is
remarkable in view of the apparent scarcity and difficulty of obtaining
food in caves. Possibly the scarcity of food is compensated for by the
scarcity of enemies and consequent longevity of cave inhabitants.

76 FAUNA OP MAYFIELD'S CAVE.

Order ISOPODA.

Family ASELLIDAE.

Caecidotea stygia Packard.

PACKARD, Am. Nat., v, 1871, 751, figs. 132, 133; Rep. Peabody Acad. Sci., Salem, v,
1873, 95 (Mammoth Cave, Wyandotte Cave, well near Orleans, Indiana). SMITH,
Rep. U. S. Fish Commission 1872-73, 661 (localities as above) ; Am. Nat., vil,
1873, 244; Am. Journ. Sci. and Arts, IX, 1875, 477 (Caves and wells, Indiana,
Mammoth Cave). HUBBARD, Am. Entom., n. s., i, 1880, 37-79, fig. 10. PACKARD,
Mem. Nat. Acad. Sci., IV, 1888, 16, 29, pis. ill, IV (Mammoth, White's, Diamond,
Long, Walnut Hill Spring, and Carter caves in Kentucky; Wyandotte, May-
field's and Bradford caves in Indiana); Pop. Sci. Mon., xxxvi, 1890, 391, 392.
GARMAN, Science, XX, 1892, 240 (Upper Mississippi Valley and ^Kentucky, east
to Pennsylvania). STEBBING, History of Crustacea, 1893, 377. CHILTON, Trans.
Lin. Soc. London, vi, 1874, 252. PACKARD, Am. Nat., xxvm, 1894, 742. HAY,
Rep. Ind. Geol. Survey, 1896, 207 (Strong's, Eller's, Mayfield's, Marengo, Wyan-
dotte, Saltpeter, and Evanston's caves, and from wells and springs, all in central
Indiana); Proc. U. S. Nat. Mus., XXV, 1902, 225, 421, 424, 428 (Mammoth Cave).
RICHARDSON, Bull. U. S. Nat. Mus., 54, 1905, 434, figs. 490-492 (Graham's Spring,
Lexington, Virginia; Mammoth Cave; wells in Indiana and Illinois).

Csecidotea microcephala, COPE, Am. Nat., vi, 1872, 411, 419; Rep. Ind. Geol. Survey,
iv, 1872, 163, 174. SMITH, Am. Nat., vil, 1872; 244 (Wyandotte Cave) .

Asellus stygius, FORBES, Bull. 111. State Lab. Nat. Hist., I, 1876, 11 (wells, springs,
and drains in central Illinois). UNDERWOOD, Bull. 111. State Lab. Nat. Hist., n,
1886, 359.

This eyeless and wholly bleached species is found abundantly at the
origin of the stream in the cave and in considerable numbers in all the
water of the cave. It is often found along the edge of the pools and
occasionally crawls upon the bank. More usually, however, it is found
under a stone in the water. When disturbed this isopod often retreats
to a flat stone, under which it conceals itself, or, this lacking, it goes to
shallow water or crawls out on the bank. Rarely Csecidotea stygia is
found outside of caves under stones or dead leaves in the water, but
even when quite abundant in these situations it is seldom seen away
from places of concealment.

Csecidotea stygia is a weak species. It can not swim and crawls very
slowly. It is nearly helpless out of water, its weak legs being scarcely
able to push it along. While entirely blind, C. stygia is sensitive to
light. Its habit of living under stones and dead leaves when outside of
caves has already been mentioned. In caves it usually retreats from
bright light. It is very sensitive to a jar of the water, all individuals
in a pool appearing much agitated when the water is disturbed or the
adjoining bank sharply jarred. This species serves to some extent as
food for Crangonyx gracilis; attempts to keep the two in the same dish
in the laboratory proved disastrous to the former. The isopod is not so
abundant in the upper pools of the cave, where the amphipod is most

ISOPODA— ASELLID^E. 77

abundant. C. stygia suffers also from the blind fish. The stomachs of
blind fishes examined contained C. stygia and in one case remains of
this species formed almost the entire stomach contents. This isopod is
rendered scarce in the larger pools, where the blind fishes are most
numerous, but is abundant in the more shallow parts of the stream
near by. I have never seen it feed, but judging from its habits, its food
is probably decaying organic matter. Individuals kept in the laboratory
feed upon decaying leaves in the water. Females with young in the
brood pouch have been taken at different seasons, and quite small indi-
viduals are seen at all times, so that this species must breed throughout
the year. I now have this species breeding in the light.

As already indicated, this species seems generally distributed in sub-
terranean waters in this region and often appears at the outlet of under-
ground waters. I have found it in Mayfield's, Twin, and Mammoth
caves. Outside of caves I have found it near Bloomington in a spring
and its stream, and also in a sheltered ravine.

Packard (1888, 108-110) studied the structure of the eyes of C. stygia
and found that in every case the optic ganglia and optic nerves are
missing. In nearly every case the "retinal cells" are broken and in
many cases the lenses and pigment are entirely lacking. Specimens from
a well in Illinois have the pigments and some of the lenses still present,
but in Mammoth Cave specimens this last trace of the eye is gone.

Cascidotea nickajackensis Packard (1881, 879) is known from Nicka-
jack Cave in Tennessee and from southern Georgia. C. richardsonas
Hay (19026, 424) also occurs in Nickajack Cave. C. smithii Ulrich
(1901, 93) comes from an artesian well at San Marcos, Texas. These
species of Csecidotea are white and eyeless. The genus resembles
Asellus very closely and it seems probable that the species of Csecidotea
have been derived from species of Asellus. Asellus cavaticus Schiodte
is found in caves, springs, and wells in different parts of Europe, and
Asellus forelii Blanc occurs in the depths of Lake Geneva. Both species
are white. The former is eyeless and the latter is eyeless or with
rudimentary eyes. Asellus hoppinse Faxon (cf. Garman, 1889, 232) is
described from Day's Cave, Missouri.

Family ONISCIDAE.
Porcellio scaber Latrielle.

LATRIELLE, Hist. Crust. Ins., VH, 1804, 45. RICHARDSON, Bull. U. S. Nat. Mus. 54,
1905, 621, fig. 671.

A single specimen was found upon the wall at "6" during April.
It was not bleached in the least.

Blatchley (1896, 234) found a specimen of this genus near the mouth
of Little Wyandotte Cave. Garman (1889, 237) reports one of this

78 FAUNA OF MAYFIELD'S CAVE.

family from a cave in Missouri. Acanthoniscus cacahuamilpensis(Ei\mek)
occurs in Cacahuamilpa Grotto in Mexico (Packard, 1894, 732). A closely
related family contains Haplopthalmus puteus Hay (1899, 871) from a
well at Irvington, Indiana, and Brackenridgia cavernarum Ulrich
(1902, 91), both of which are white, the former having small eyes and
the latter none. Another relative of Porcellio is Euphiloscia elrodii
from a cave at Orleans, Indiana (Packard, 1873, 97) . Several eyeless
and light-colored relatives of Porcellio are found in European caves.
One of these, Titanethes albus Schiodte, is abundant and generally
distributed in caves in Carniola and Istria, in Southern Austria.

Among other subterranean isopods are Ciralonides texensis Benedict
(1895) from an artesian well at San Marcos, Texas, Conilera stygia
Packard (1900) from a well at Monterey, Mexico, Phreatoicus assimilis
Chilton, Cruregens fontanus Chilton, and Phreatoicus typicus Chilton
from wells in New Zealand (Chilton, 1894, 185-218). All these species
are described as eyeless and white or "transparent."

Order AMPHIPODA.

Family GAMMARIDAE.

Crangonyx gracilis Smith.

SMITH, Am. Journ. Sci., ser. 3, II, 1871, 452 (Lake Superior) ; Rep. U. S. Fish Com-
mission 1872-73, 654. FORBES, Bull. 111. St. Lab. Nat. Hist. No. 1, 1876, 6 (Illinois)-
HAY, Am. Nat., xvi, 1882,241 (ponds and streams, Irvington, Indiana). PACKARD,
Mem. Nat. Acad. Sci., IV, 1888, 16, 36 (Lake Superior, Michigan, Mayfield's Cave).
HAY, Proc. Ind. Acad. Sci,, 1891, 150. BLATCHLEY & HAY, Rep. Ind. Geol. Survey,
1896, 206 (Eller's, Donnehue's, and Mayfield's caves).

This species is quite abundant in the water of the cave. The blind
fishes feed upon it, and hence its distribution is limited. It is rarely
found about the lower pools of the cave, where the fishes are most
abundant. In the upper pools, where there are no fishes, it is very
common. It does not occur in shallow water to any extent; hence is
not found in the shallow parts of the stream, where Csecidotea stygia is
so numerous.

Ordinarily this animal is to be observed quietly resting or crawling
about over the bottom of the pools. When disturbed it swims by rapid,
jerky movements and often rises toward the surface, especially when
persistently pursued. It may be driven to the very edge of the pool,
when it often crawls partly out of the water. It is quite sensitive to a
disturbance of the water, or to light, which it avoids.

This species sometimes feeds upon the cave isopod Csecidotea stygia.
I have twice seen it in an aquarium catch a living Csecidotea. In con-
finement it keeps under cover or along the dark edges of the aquarium
much of the time. It feeds upon any organic matter or even upon the
carcass of a fellow. I was sometimes suspicious that the larger ones

AMPHIPODA— GAMMARID.E. 79

attacked and fed upon the smaller. It appears to breed at all seasons,
old with young in the brood pouch being found throughout the year.

C. gracilis is found in some abundance in the Donaldson Caves at
Mitchell. Bollman (Packard, 1888, 36) and later Blatchley (1896, 206)
reported it from Mayfield's, and Blatchley (1896, 206) found it it Eller's
and Donnehue's caves. It is common in southern Illinois and in central
Indiana. I have found it to be common and very abundant about
Bloomington and Mitchell, Indiana.

The C. gracilis found in Mayfield's Cave has well-developed and
distinctly pigmented eyes, but differs from the outside form of this
region, (1) in the total lack of pigment, except in the eyes, the out-
side form being rather dark, and (2) in the larger size, the largest
from outside measuring 13 mm. and the largest from the cave more
than 18 mm. in length. Three of the largest ones from the cave meas-
ure 18.4, 18.2, and 18.2 mm. respectively; three of the largest from
outside, 12.5, 13.1 and 11.8 mm. respectively. I have kept individuals
of this species in the laboratory in the light as long as 8 months, to
test the possible effect of light upon the development of dark pigment.
Not one of the many young liberated in the light by females with young
in the brood pouch when collected (I have not had them to breed during
captivity) nor one of the old ones, showed any indication of taking on
color. A careful detailed study of a series of this variety compared
with a series of C. gracilis from outside of caves has not been made.
Such a study will probably reveal other differences between the two
forms.

W. P. Hay (1896, 206) says of the occurrence of this species in caves:

The animal is not one given to life in such situations, and I am of the opinion
they are to be regarded as accidental visitors only, having been washed by heavy
rains into the cave streams, from which they have been unable to make their escape.

Blatchley (1896, 206) says:

From the nature of the stream in the cave (Mayfield's) I do not think it is possi-
ble for it to escape, and facilities are, therefore, excellent for a future study of the
length of time necessary to bring about organic changes due to cave environment.

After having carefully laid off the course of the cave stream in
Mayfield's Cave under ground and plotted it above ground, I am con-
vinced that the cave stream has no origin above ground or that at most
it may receive water from only a sink-hole ordinarily dry. Since there
is no above-ground pool or counterpart of the under-ground stream
from which this species could be swept into the cave, it is impossible
for it to have been carried into the cave by heavy floods. After heavy
rains the cave is more or less flushed, due to sink-holes and percolating
water, and during a time of phenomenally high waters in April, 1904,
the whole cave was nearly completely filled with water. Such a flood,

80 FAUNA OF MAYFIELD'S CAVE,

however, could not carry this species in from above, but could only
make possible its entrance from below. On the other hand, the cave
stream comes out as a spring not 20 yards from where it disappears in
the cave, and to escape, this amphipod would need only to follow the
stream, which it could easily do during the winter and spring, or after
a heavy rain in any season. Similar conditions seem to exist in Donne-
hue's Cave, as nearly as I can determine from Blatchley's account.
I believe that this species has entered the cave from below and by its
own efforts, that it has been able to maintain itself in the cave, and
that it remains there in response to the proper environment. Its occur-
rence in several Indiana caves, in all of which it is fairly abundant, is,
to say the least, not inimical to this view.

Subterranean American relatives of Crangonyx gracilis are: C. vitreus
(Cope), an eyeless form, recorded from Mammoth Cave, Kentucky, and
Saltpeter Cave and wells at Orleans, Indiana (Packard, 1873, 95);
C. packardii Smith, having small eyes, recorded from Wyandotte Cave
and wells at Orleans and New Albany, Indiana (Packard, 1888, 36) ;
C. tennis Smith, an apparently eyeless form, from wells at Middletown,
Connecticut (Packard, 1888, 35) ; C. lucifugus 0. P. Hay, a pale, eyeless
form, from a well at Abingdon, Illinois (loc. cit. , 38) ; C. mucronatus
Forbes, a colorless, blind species, from wells and springs in central
Illinois (loc. cit., 37); C. flagellatus Benedict (1895, 617) and C. bowersii
Ulrich (1901, 85), blind colorless species from an artesian well at San
Marcos, Texas; Niphargus antennatus (Packard, 1888, 36), a pale or
colorless form with eyes rudimentary or lacking altogether, from Nick-
ajack Cave, Tennessee.

Several subterranean species belonging to the genera Gammarus,
Niphargus, and Crangonyx are found in Europe and some of them are
known from Algeria as well (Chevreux, 1901). These are all regarded
by Hamann (1896), perhaps wrongly, as most authorities are against
this view, as varieties of a single species, Niphargus puteanus Koch, a
colorless, eyeless form which he says is widely distributed in caves, in
the depths of Alpine lakes, and in wells in England and in nearly every
country of the continent.

Still other subterrean forms, Crangonyx compactus, Calliopius sub-
terraneus and Gammarus fragilis (Chilton, 1894, 218-244), "transpar-
ent" and without trace of eyes, are known from wells in New Zealand,
while two blind Niphargi are known from surface waters in Victoria,
Australia (Sayce, 1899 and 1901).

According to W. P. Hay, C. vitreus and C. packardii are probably
derived from C. gracilis. The form of C. gracilis found in central
Indiana caves is probably another species in process of formation.
Already it is entitled to the rank of a variety.

MOLLUSCA. 81

Order COPEPODA.
Family CYCLOPIDAE.

Specimens of Cyclops seemed fairly common in the cave stream.
Only two attempts to collect them were made, and the following species
were taken in small numbers:

Cyclops virldis (Jurine) var.?
Histoire des Monocles, 1820, 46, pi. 3, fig. 1.

Cyclops viridis brevlspinosus (Herrick).
Geol. and Nat. Hist. Surv. of Minn., 12th Ann. Rep., pt. v, Crustacea, 1884, 192, pi. 30.

Cyclops bicuspidatus Claus.
Archiv f. Naturgeschichte, July, 1857, Bd. 1, 209, Taf. XI, figs. 6 and 7.

Canthocamptus illinoisensis Forbes.
Bull. 111. Mus. Nat. Hist. No. 1, 1876, 3, 25.

According to A. S. Pearse, who identified these species, all are very
common species and appear to have normal eyes. They could easily
reach the cave straem by following up the outdoor stream as Copepods
very readily do. A blind fish's stomach contained one of these animals.

Copepods were found in the waters of Mammoth and Twin caves.
Cyclops cavernarum and C. learii, species without eyes, are reported
from an artesian well at San Marcos, Texas (Ulrich, 1902, 95) . Several
species of Cyclops are found in European caves. In some of them the
pigment is considerably reduced and two of them are described as eye-
less, but Hamann (1896, 248) thinks without sufficient grounds. C. novse-
zealandias, a normal terranean form, has been obtained from wells in
New Zealand (Chilton, 1894, 247).

ANNELIDA.

Earthworms are rather common in Mayfield's Cave, occurring in the
soil from the mouth to "10" and very commonly from "10" to "15,"
where there is considerable soil. They are occasionally seen along the
cave stream far back in the cave. Their occurrence is accidental.

An unknown chastopod was found crawling along on the bottom of
the stream at "41."

MOLLUSCA.

Slugs were occasionally seen in the cave, especially during the fall.
Many individuals of a small black one were seen on the walls and roof
near the mouth, while occasionally one was seen back as far as "17,"
where it was noted on the mud or under debris. Two individuals were
seen at "42" on the floor, near bait. A larger mottled gray slug, probabl y

82 FAUNA OF MAYFIELD'S CAVE.

a Limax, was seen a few times near the mouth. Small, usually empty,
shells of a single species of gastropod were seen from the mouth to
'17," about debris, and a similar shell was found in Truett's Cave.
Two or three times this mollusk was found alive. The occurrence of
this shell is probably accidental. It is found in locations which suggest
that it has been washed in through the sink-holes near "12."

Ulrich (1902, 85) mentions 4 species of gastropods from EzelPs Cave,
near San Marcos, Texas. Call (1897, 387) found a small species in
Mammoth Cave, which is probably a true cave form, and Chilton (1894,
246) discusses a species without pigment from wells in New Zealand.
Zonites subrupicola, found in Clinton's Cave, Utah (Packard, 1877, 163)
and later under stones in California (Packard, 1888, 21), is a bleached,
almost white form.

Class PLATYHELMINTHES.
Order TURBELLARIA.

Three flatworms were found in the pools in the cave. They were
almost or quite without black pigment, but had distinct eyes.

Packard (1888, 27, 28) found a white eyeless planarian (Vortex?
cavicolens) in Carter Cave, Kentucky, and another (Dendrocselum
percascum) , also white and eyeless, in Mammoth Cave. I found the
latter species in Mammoth Cave and also in White's Cave, near the
Mammoth Cave.

LIST OF SPECIES.

The total number of species taken from Mayfield's Cave and con-
sidered of sufficient interest to mention in this paper is 110; 75
species have heretofore been recorded from those caves of Indiana,
about 25 in number, in which collections have been made. The table on
the following pages compares my list of species from Mayfield's Cave
with Blatchley's list collected in 1896 and with other lists from all the
Indiana caves.

The species are classified as (a) strays, or accidental visitors; (6)
visitors, going into the cave and leaving at will; (c) temporary residents,
hibernating in the cave or spending a part of their existence there;

(d) permanent residents, spending their whole lives in the cave; and

(e) species which live in the cave exclusively, never being found else-
where. Other columns arrange species with regard to their relation to
light, the grouping being as follows: Those living in (a) bright twilight
only; (b) twilight only; (c) twilight, and rarely in darkness; (d) rarely
in twilight and habitually in darkness; and (e) darkness only.

LIST OF SPECIES.

Comparative table of species known from Indiana caves.

83

Species.

Previous
lists from
Indiana
caves.

01

2

'•?

Cw
k-H
<

a

0 <U

•in ^

"^

'T.

O

3
^

Grouping with re-
gard to purniii
ninic-y of habita-
tion within caves.

Grouping with
regard to light.

Remarks.

Blatchley's list.

02
+S

w
0)

fH
-*->

o

a

tc

>>

*

tt

+3
•Jl

6

g

3

'02

>

Temporary resi-
dents.

Permanent resi- ^
dents.

Cave exclusively. «

a

3

_U)

"££?

^o
o

+3

j-1

~So
M

b

jj

o

+2

w>

'%

H

c

i

S*

H

*=> a
ja—1
°f>,

%

H

d

5.M

si

cs a

5i-
'""^
•^s

Is

e

>>

"a
o

iA
rt
Q

Mfiiiuiialia:
Vrspert il'( > fuscus

X
X

x

X

X
X

• •

Not common.
Very abundant
in winter.
Not common.
Reputed to visit
caves.
Taken in Twin
Caves.
Abundant.
Common.
8een once.
Occurrence
doubtful.
Occasional.

Tracks seen.
Tracks seen. (?)

Two specimens.
One specimen.
Do.
Common.
Two specimens.

Introduced.

Mouth of Twin
Cave.

Two specimens.
One specimen.
Do.

Common.

Known from a
single specimen
One specimen.

Do.

Pipistivllus subtiavus. .
Mvotis lucifugus

X

-

X
X

_

. .

Ijiisiurus borcalis

(")

Corynorhinus macrotis.

Peromyseus leucopus..
Procyon lotor

X
X

X

x

X

X
X

X

C>}

x

X

Tamias striatus

••

X

Marmota monax

Vulpes fulvus

X

V9

X

••

Urocyon cinereoargen-
tatus.

X

X

X

x

X

(•> 1

Total Mammalia

7

1

8

3

4

1

1

2

5

••

Amphibia:

Plethedon cinereus

X

X
X

X

x

• •

P. cinereus dorsalis —
P glutinosus

x\

X

x

X

X

X

X

• •

Spelerpes maculicaudus
KM n;t clamata

X
X

X

x

Total \mphibia

2

5

4

• •

1

1

3

1

Pisces:
Amblyopsis spelwus —
Typhllchthys wyan-
dotte.

X

X

X
X

V

••

X

••

X

Total Pisces

1

3

i

1

1

Insccta:
Hvmenoptera—
Pemphredonida? sp. . .
Do

x

X

x

X

• •

X
X

X
X
X

••

••

••

Total Hymenoptera.

Coleoptera—
Carabidre- —
Anopthalmus ten-
nis.

3

1

••

2

3

••

X

X
X

X

••

••

X

••

X

Patrobus longi-
cornis.
Platynus cincti-
collis.

X

X

X
X

••

• •

••

X

••

X

••

••

84 FAUNA OF MAYFIELD'S CAVE.

Comparative table of species known from Indiana caves — Continued.

SPecies.

Previous
lists from
Indiana
caves.

Author's list from Mayfleld's
Cave.

Grouping with re-
gard to perma-
nency of habita-
tion within caves.

Grouping with
regard to light.

Remarks.

Blatchley's list.

IK

+3

01

t-i

o>

a

+a

o

a

tc

>>

8
+j

OQ

6

o5

•-*

O
-u

cc

>

X
X

Temporary resi-
dents.

Permanent resi-
dents.

Cave exclusively. ^

Bright twilight
only.

b

>>

"a
o

+3

a
.Sf

$

H

Twilight and rare- n
ly in dark.

Twilight and ha- _
bituallyindark. a

e

>>

1
X

&

Insecta — Continued.
Coleoptera— Con t'd.
Carabidfe— Cout'd.
Platynus tenuicollis
P. punctiformis

X

x

..

..

X
X

Rare.
One specimen.
Authority of
Cope.
Found in Tru-
ett's Cave.
Porter's Cave.

filler's Cave.
Clifty Cave.

Porter's Cave.
Do.

Do-
One specimen.

Very abundant.
Abundant.
Not common.
One specimen.
Two specimens.
Porter's Cave.
Very abundant.
One specimen.

Fairly common.

Rare.
One specimen.
Do.

One from Por-
ter's Cave.
Fairly common.
Saltpeter Cave.

Fairly common.
Abundant.

One specimen.
Rare.

Two specimens.

Several.
One specimen.

P. rnarginatus

X

Atranus pubescens.

Scarites subterra-
neus.
Pterostichussayi
Harpalus penusyl-
vanicus.
Stenolophus ochro-
pezus.
Bradycellus rupes-
tris.
Hydrophilidfe—
Philhydrus sp

X

X
X

X
X

X
X?

X
X
X

X

Silphidse—
Choleva sp

X

X

x

X

X

X
X

Staphylinidie—
Rheochara lucifuga.
Quedius spelseus
Q. fulgidus

V9

X
X
X
X

X?

••

X
X

X

X

••

X

Philonthus lomatus
Tachinus repandus.
CryptobiULn bicolor
Lesteva pallipes
Tenebrionidte sp. ..

••

X

• •

X

x

X
X

\

x

Total Coleoptera.. .
Aphaniptera sp

1

13

5

13

4

4

1

3

2

••

8

2

X

••

X

••

••

••

X

• •

Diptera —
Tipulidee—
Trichocera sp

X
X
X

X

X

x

X
X
X

••

••

Clad uraindi visa
Rhypholophus mei-
genii.
Ulomorpha pilosella
Psychodidse—
Psychoda cinerea...
P. minuta . . .

X

x

• •

X

x

X

• •

Chironomidse —
Chironomus sp . .

y

X

••

X
X

• •

X

X
X

••

CulicidfE—
Anopheles puncti-
pennis.
A. quadrimaculatus
Culex pipiens

X

X
X

X

X
X

x

X
X

X

Mycetophilidre—
Polylepta leptogas-
ter.
Rymosia filipes
Mycetophila discoi-
dea.

• •

••

X
X

• •

• •

••

X

X

LIST OF SPECIES. 85

Comparative table of species known from Indiana caves. — Continued.

Species.

Previous
lists from
Indiana
caves.

Author's list trom May-field's
Cave.

Grouping with re-
gard to perma-
nency of habita-
tion within caves.

Grouping with
regard to light.

Remarks.

Blatchley's list

no
+a

m

(4
(D

a
•*?

Q

«

EG

!•>>
£-1
02

b

•j'i

o

a?
>

Temporary resi-
dents.

Permanent resi-
dents.

Cave exclusively. <*

a

»3

a
.5?

-Jb

a

w 0

d

5P

''-*

S

b

>>

o
*->

s

'i

H

Twilight and rare-
ly in dark.

Twilight and ha-
bitually in dark. a

e

^

o
44

1

Imsecta— Con ti u ued.
Diptera— Cont'd.
Mycetopbllidse— Con.
Mycetophila ich-
neumotiea.
M obscura

\(

X
X

X

X
X

X
X

X
X

• •

X

Rare.

Very common.
Do.
One specimen.
Do.
Very common.
Common.
One from Tru-
ett's Cave.
One from Ma-
rengo Cave.
One specimen.

One specimen.
Few.

Very common.
Rare.
One specimen.
Do.

Two specimens.
Do.

Common.
Few .
Common.
Few.
One.
Mayrteld'sCave.
From several
caves.

Very abundant.

Wyandotte Cave

Abundant.
Do.

Abundant.

M umbraticus

X

•

•

X
X

x

M. analis

M. incerta

Bolitophilahybrida
Sciara sp

X
X

X

••

Macrocera hirsuta. .

Odontopoda say i —
Ceci d o m y i d re —
? Diplosis sp

•

X
X

\/

X,

X

X

X
X

X

Dolicopbidfe—
Liancalus genualis .
INeurigona sp
Phoridfv —
Aphioctueta uigri-
ceps.
A rufipes

X

X

•

X

-

X
X

• •

X

Muscida? —
Calliphora vomito-
ria.
Muscadomestica ...
An thorny idffi —
Hylemyia lipsia...

X

x

X

X

Pegomyia afflnis . . .
Helomyzidte —
Oecotliea fenestralis
Leria latens

X

v

X
X
X

X
X

y

X
X

X

. .

L defessa

X

X

L serrata

L. fratcrna

X

X

L specns

X
X

X

Li pubescpns

Borboridre —
Limosina tenebra-
rum.

Total Diptera

Lepidoptera —
Tineidse—
Blabophanes ferru-
ginella.
Noctuidse —
Scoliopteryx liba-
trix.
Plathypena scabra.

Total Lepidoptera . . .

Orthoptera —
Locustidte —
Ceuthophilus sty-
gius.

Total Orthoptera

X

8

X
8

X

12

••

33

10

7

9

4

9

11

••

X

X

2

X

X

X
X

1

2

••

2

X-

X

X

••

X

X

1

1

1

1

1

86 FAUNA OF MAYFIELD'S CAVE.

Comparative table of species known from Indiana caves — Continued.

Species.

Previous
lists from
Indiana
Caves.

Author's list from Maytield's
Cave.

Grouping with re-
gard to perma-
nency of habita-
tion within caves.

Grouping with
regard to light.

Remarks.

Blatch ley's list,

w

+3

GO

£H

0)

A

+3

O

a

cc
>a

rt
•~

OQ

b

i

o

'<a

!>

Temporary resi- 0
dents.

Permanent resi.
dents. a

Cave exclusively. w

a

+a

a

.£?

££

^ a

0
+a

a

5?

f*

CQ

Twilight only. cr

Twilight and rare-
ly in dark.

Twilight and ha-
bitually in dark.

e

>i
"a

0

^j
L*

c«

O

Insecta— Continued.
Aptera—
Thysauura—
Machilis varia-
bilis.
Campodea cookei.

X

X

Fairly common.

Wyandotte Cave
(Packard).

Very abundant.
Abundant.

One specimen.
Abundant.
Three specimens

Several.

Maytield's Cave
(Boll man).
Common.
Evanston's Cave.
Maytteld's Cave.

Several caves.

Sibert's Well
Cave (Boll man)

Three specimens

Two specimens.

Abundant.
Saltpeter Cave.
Few.
One specimen.

Abundant.

Three specimens
Few.
Two specimens.
Few.
Three specimens
Found in Mitch-
ell Caves.
Abundant.

Collembola —
S i n e 1 1 a caverna-
rum.
Toniocerus niger. . . .

X

X

X

1

11

X

<

Total Aptera

1

2

3

1

••

1

1

1

1

Mj/riapoda:
Chiloguatha —
Cambala annulata. . . .
Couotyla bollmani. . .
Buryurus erythropy-
gus.
Scytonotus granula-
tus.
S cavernarum

X
X

X
X

X

X
X

X

x

x

X

X

X
X

X?

X

Polvdesmus serratus.
Polyzonium rosalbum
Spirostrephon lacta-
rium.
Pseudotreniia caver-
narum.
P carterensis

V

^^

X

X

X

X

X

Cnilopoda —
Bcolopendra wood!..

Total Myriapoda .

Arachnida:
Araneida—
Dk'tynida-—
Amaurobius ben-
netti.
Agaleuidre —
Tegenaria derhami
T cavicola .

X

'

X

1

5

3

6

5

1

1

4

X

X

X
X

•

X

x

x

X

X

X
X

X
X

X

Cicurina pallida. ..
Coeletes sp

X

Thcridiida?—
Theridium kentuc-
kyense.
T porter!

X
X

v

x\

X

Nesticus carteri —
Argyrodes trigonun
Linyphia marginata
TJ. nigrina

X

X

X

X
X

X

X

Wlllibaldi caverni-
cola.
Phanetta subtcrra-
nea.

X

X

X

X

*

X

LIST OF SPECIES. 87

Comparative table of species known from Indiana caves — Continued.

Species.

Previous
lists from
Indiana
caves.

Author's list from Mayfleld's
Cave.

Grouping with re-
gard to perma-
nency of habita-
tion within caves.

Grouping with
regard to light.

Remarks.

Blatchley's list.

tc

t»

FH

<U

fH
-«-)

O

«

DQ

f~,

53

•4-3

CO

b

g"

o

«

Temporary resi- a
dents.

Permanent resi. ^
dents. ~-

Cave exclusively. ^

«

u
a
be

!i

^ a
o

4J

a

60

'C
«

b

jj,

i— i

o

a

be

'£

H

.c

I
1*

H C*

oS ^3

sa

|>?

T»

H

rf

1

««

ss

S a

sB

.SfS

•3 s

Is

e

£?

~a
o

x

1

Arachnida— Cont'd.
Araneida — Cont'd.
Theridiida? — Cont'd.
Erigone infernalis

X

V

v

Few.
One specimen.
Porter's Cave.

Wyandotte Cave
(Cope).
Few.
Two specimens.

Taken in Mitch-
ell Caves.

Common.
Donal ds on's
Cave.
Common.
Very abundant.
Few.
Wyandotte and
Clifty caves.
Wyandotte Cave
(Cope).

Abundant.
Common.

Very abundant.
One specimen.

Very abundant.
"Wyandotte Cave
Saltpeter Cave.
Wvandotte Cave
(Cope).

Not rare.
Do.

Do.
Do.

Common.
Onu specimen.

Theridid sp. nov. . . .
Tmeticus tridenta-

tUB.

Anthrobia

X

X

X

X

X

x

X

Meta menardi

X

X
X

X

X

x

X

X

X

Theridiosoma gem-
inosum.
Tetragnatha labori-
osa.
Lycosidse —
Dolomedes scriptus.

X

X

( 4

Acarina —
Rhagidia cavicola . .
Tyroglyphus sp
? Gamasus sp

X

x

X

X

X

x
x

Scotolemoii flaves-
cens.
Ctlionius packardii .

Total Arachnida. . .

Crustacea:
Podopthalmata —

Astacidse —
Cambarus pelluci-
dus testii.
C. bar ton i

10

3

19

2

5

4

5

3

c

7

6

X

X

X
X

/S
\/

X

X

X
X

X
X

X

X

X

Isopoda—
Asellidte —
Ca?cidotea sty^ia

Oniscidae —
Porcellio scaber

x

X

7

X
X

X,

Amphipoda —
Gammaridse —

Crangony x gracilis . .

Cauloxenus stygius.
Copepoda —
Cyclopida},—

X
3

X
X

••

X
X

X

x

x

C. viridis brevispi-
nosus.
C. bicuspidatus
Canthocamptus illi-
noisensis.

Total Crustacea . . .

Annelida:
? Lurubricus sp

X

X
X

9

1

4

3

1

1

8

X

X

X
1

X

• •

X

X

—

Total Annelida —

2

1

••

2

FAUNA OF MAYFIELD'S CAVE.

Comparative table of species knoivnfrom Indiana caves — Continued.

Previous
lists from
Indiana

2

Grouping with re-
gard to perma-
nencv of habita-

Grouping with
regard to light.

caves.

c

tion within caves.

1

a

b

c

d

e

a

b

c

d

e

q »

^

.

i

Species.

~l

|i

01

CD

a>

aa

1.

jsj

"a

||

||

Remarks.

~

01

~^

*~* a

~~ *-"'

>~J!

^ ^

o

rt "

rt .2

^

a

<D

DO

X

g OJ

g§

o

o

*a

*- C

*^ >>

a

JS

1—1

O

a

^H

O

c ^

ft

CS'1-1

<u

Wi

Mix.

bi'rt

o

o

*j

0>

•3

cS

a

a

0)

to

3

^— i ' '

— 5

d

^

12

05

n>

cS

^

t^ -»^

cS

M

O

"^

O3

^

O

M

EH

•H

EH-"

M

Mollusca:
Gastropoda —

X
X

x

X
X
v

Two specimens.
Common.

Several.

V

X

2

-—

—

Shell sp

3

3

Total Mollusca —

Platyhelminthes:

Turbellaria—

x

Threespecimens.

Total Turbellaria. .

'•

1

1

1

Grand total

60

21

110

17

36

24

23

10

21

12

37

38

2

The following, besides summarizing the other table, includes a column
showing the number of animals of each order now known from the
Indiana cave region.

Species.

Previous
lists from
Indiana
caves.

Author's list from Mayneld's
Cave.

Now known from all Indiana
caves.

Grouping with regard
to permanency of
habitation within
caves.

Grouping with regard
to light.

Blatchley's list,

%

a*
•„

<D

a
o

a

vi

>>

1
5

6

£
3

'in
>

Temporary resi- ft
dents.

Permanent resi- a,
dents.

Cave exclusively. »

a

+3

a

bJD

•- ^

^c

*= 0

^3

a

bo

t3

03

b

>f

"3

o

«a

a
_bc

1
H

i

4

2

i
i

'i

2

1 Twilight and rarely
• ^tc- ~i*.h-. . CD- oe< • wtol in dark.

Twilight and habit- _
ually in dark.

e

>*

a

o

X

1

Mammalia

'i

i

'3

8
5
1
3
13
1
33
2
1
3
6
19
9
'i
3
1

11
5
3
3

22
1
39
3
1
4
11
27
12
2
3
1

'i

4
'8

2
1
1

3

4

'i

io
'i

5

5

i

3

4

'2
1

7

2

4
4

1
1

'i

i

8

i
i

5
3

'i

'i

3

i

1
3
1

1

'3

2

9

'e

5
1

'2
1
11

i
i
i

6

8

i

i

i

Amphibia

Pisces

Hymenoptera

Coleoptera

13

5

Aphaniptera

Diptera

12
1
1
1
5
10
7

i

2
3
3
3

Lepidoptera

Orthoptera

Aptera

My riapoda

Arachnida

Crustacea
Annelida

Mollusca ...

Turbellaria.

Total

60

21

110

138

17

36

24

28

10

21

12

37

38

2

GENERAL CONSIDERATIONS. 89

GENERAL CONSIDERATIONS.

From the foregoing lists it will be seen that Mayfield's Cave contains
practically all the cave inhabitants and lacks very few of the true cave
forms found in any Indiana cave. Hence it appears that a small cave
like Mayfield's affords practically the entire cave fauna of a cave region.

The list of species from Mayfield's Cave is much larger than all other
lists from Indiana caves. This increase in number of species in itself
is not significant, however, except as indicating that very many species
may be found in caves occasionally. Those added to former lists are
principally species which are only incidental and temporary cave
residents.

The permanent and true cave species form but a small part of the
entire cave fauna, while visitors, strays, and temporary residents form
about 77 per cent of the life of the cave. Visitors and temporary residents
do not ordinarily breed in the cave, and do not often go into the remote
portions of the cave, where there is absolute darkness and where there
are constant conditions of temperature and moisture. Hence they are
not a part of the real cave fauna and are given little consideration in
this discussion. Nevertheless they are interesting and significant in
that they show that there is no hard and fast line between cavernicolous
and non-cavernicolous forms. In Mayfield's Cave and about its mouth
there are represented all gradations between animals whose habitat
is in the open in the woods near the mouth of the cave and other forms
(e. g., Anopthalmus tennis, Quedius spel&us, and Phanetta subterranea)
which are well adapted for cave existence and probably do not occur
outside of caves.

True cave forms are not necessarily confined to absolute darkness.
They always live in dark places, but may wander away from the absolutely
dark portions of caves into twilight or even outside of caves. The blind
isopod Cascidotea stygia is common in caves throughout Indiana and
Kentucky and is apparently nearly subterranean in habit, yet it
is not confined to this mode of existence. It has been found under
stones in streams, and in wells, springs, and drains in Virginia, Illinois,
Kentucky, and Indiana. The Ceuthophilus stygius seems fairly well
adapted to cave life, but besides being found in caves, occurs in cellars and
about wells. Sinella cavernarum, so common and abundant in Indiana
and Kentucky caves, occasionally wanders from darkness into dim twi-
light at the mouth of Mayfield's Cave, but has not been taken in full
daylight, and probably could not stand conditions less constant than those
of a cave. Banks (1893) found Scotolemon flavescens, an arachnid pre-
viously known only from Indiana caves, under stones on the Virginia
shore of the Potomac near Washington, D. C. , which does not differ from

90 FAUNA OF MAYFIELD'S CAVE,

cave specimens. Absolon (1900, 4) found that certain cave thysanurans
and mites were killed in a few minutes by exposure to daylight, while
other thysanurans and mites from outside of caves were not affected by
the same exposure. Hence while some typical cave forms wander out-
side of caves into other dark places, others are incapable of existence
except under typical cave conditions.

Many of the visitors and a few of the temporary residents occurring
just within the mouth of the cave are mere local inhabitants of caves;
but the true cave species, those structurally adapted to cave life and
living habitually in absolute darkness, are old cave forms which are
widely distributed. For example, Platynus, a genus of Carabidae, is
represented by 4 species taken occasionally in Indiana caves; Platynus
cincticollis Say is taken in Mayfield's and Truett's caves; Platynus
tenuicollis Le Conte from Mayfield's, Truett's, Twin, and Donnehue's
caves; Platynus punctiformis Say from Mayfield's; and Platynus margin-
atus from the Mitchell caves. These species have been taken in different
caves and only two of them in more than one. Their occurrence in
caves is more or less incidental. They are local cave inhabitants, prob-
ably do not breed in caves, and are not at all commonly found there.

On the other hand, there are 8 American species of the genus Anop-
thalmus of blind Carabida?, all but one of which are confined to caves
and some of them known from several localities. A. tennis is known
from almost every Indiana cave which has been zoologically explored.
This genus is also found in Europe, where there are about 50 species
living principally in caves, and where a very closely related species of
this genus lives in caves exclusively. Anopthalmus is an old genus
widely distributed and highly adapted to cave life. *

Seasonal changes in temperature are not very marked within the
cave and in the remote parts there is as little change in life. The hiber-
nating bats and the salamander larvae appear during the winter and
the young of Cambarus bartoni and Cambarus pellucidus testii are
observed toward spring, but all other forms of life seem to occur in
about uniform numbers the year round and those which breed in the
innermost parts of the cave breed when conditions of food are favorable,
regardless of the season. However, where there is any marked seasonal
change in the temperature of the cave the activity of all cave animals
is profoundly affected. From the mouth to "21," and to some extent
to the mound, the temperature varies with the season and with the lower
temperature of winter the decrease in abundance and activity of cave

*The genus Anopthalmus has been discarded in Europe, since every intermediate
stage between the eyed Trechus and eyeless Anopthalmus has been found. Anop-
thalmus is, then, considered a subgenus of Trechus, from which it differs principally
in the lack of eyes.

GENERAL CONSIDERATIONS. 91

animals is very marked. This influence is very noticeable with the
flies and spiders, and to some extent with the rnyriapods, thysanurans,
and beetles.

There seems in most cases a direct relation between the temperature
and the activity of those insects which hibernate near the mouth of the
cave. These hibernating insects become more active when subjected
to a rise in temperature even in the midst of winter, while if subject to
a temperature of 5° C. or lower they are barely able to stir. These
remarks apply to these insects when disturbed. They apparently move
very little or not at all otherwise.

Many species in the portion of the cave near the mouth retire into
cracks and small holes or short side-passages, so that the abundance of
life is apparently greatly diminished in the winter. Phanetta subterranea
and the other spiders which live under rocks and debris become quite
inactive; Theridium kentuckyense, Tegenaria derhami, and other spiders
which have webs on the wall crawl into cracks; the myriapods apparently
do likewise or conceal themselves under debris; and the helomyzid flies
and Limosina tenebrarum withdraw into cracks and crannies of the wall.
The latter also conceals itself under debris. This marked change in
the cave life is not so noticeable beyond "17." Animals which are not
near enough the mouth to be subjected to great changes in temperature
show a variation in their activity in proportion to the change to which
they are exposed. Beyond the mound the fluctuation in temperature is
so small, however, that it produces no noticeable influence upon the
activity of animals in the cave. Here the true cave forms of whatever
order are equally active at all seasons.

As far as observed cave animals do not breed during the winter in
those parts of the cave where the temperature becomes lower than about
9° C. With the coming of the higher temperature again, the spring
freshets having meanwhile left an abundance of organic matter in parts
of the cave, many cave species begin to breed abundantly. This activity
is very marked in the flies, especially the species ofLeria, Oecothea fenes-
tralis, and Limosina tenebrarum, the spiders, the myriapod Conotyla
bollmani, and the beetles Rheochara lucifuga and Quedius spelsRus. The
Diptera mentioned become extremely abundant and by about June 10
have reached their maximum in number. All species in those parts of
the cave appear to become more abundant in spring. This increase is
probably a real one due to the animals breeding freely, and an apparent
increase due to the greater activity of the individuals and their having
come out from the obscure places in which many of them spend the
winter. It seems possible that many Helomyzidse and perhaps other
Diptera are reared outside the cave and enter it early in the summer.

92 FAUNA OF MAYFIELD'S CAVE.

Concerning the food of cave animals in general comparatively little
has been published, although the food of Amblyopsishas been well under-
stood for some time. Packard (1876, 285) mentions the abundance of
Pseudotremia cavernarum about bits of candle-drip in caves and in the
same paper and later in his monograph (1888, 24, 25) mentions the
probable food of certain cave animals. Cope (1871, 13) remarks upon
the scantiness of food of cave animals and offers a suggestion as to the
source of the supply through vegetable matter swept into caves by
water. Chilton (1894, 264) confesses inability to add to the knowledge
of the food of subterranean crustaceans. Carpenter (1895, 33) notes
that Lipura gets its food by engulfing quantities of the fine earth, from
which it obtains vegetable mold, and mentions the fact that ' 'collectors
in the Carniolan caves secure insects by leaving pieces of wood as
traps." The food problem is a difficult one, and I have gotten far less
light on the subject than I had hoped.

Throughout the paper mention has been made of the food and feeding
habits of the different species so far as observed. It may be well to
recapitulate somewhat here. Sciara, Aphiochseta, Limosina, and the
staphylinidous beetles are found in greater abundance at decaying animal
matter; Ceuthophilus, the cave myriapod, and the cave thysanuran are
about equally abundant at decaying organic matter, whether animal or
vegetable. As stated before, any bit of organic matter of whatever
sort serves to attract cave Arthropods in considerable numbers. Light
disturbs these creatures, so that it is well-nigh impossible directly to
observe their feeding in the cave, but all have been so nearly caught in
the act of feeding that the cases are practically proven. The spiders are
probably predaceous in the main, but Phanetta subterranea and Erigone
infernalis may feed upon decaying organic matter also. The former
was observed to catch a thysanuran, and at another time one was found
which seemed to be feeding upon cheese. The crustaceans are preda-
ceous as well as being scavengers. The Anopthalmus is predaceous.

In fig. 13 an attempt is made to represent in a graphic way the
dependence of cave animals upon organic matter carried into the cave
and to show the food relation of cave species to one another.

The fact that all of these animals are absolutely dependent directly
or indirectly upon decaying organic matter brought into the cave from
outside by the merest chance and accident must not be lost sight of.
It will readily be seen, too, that there is a close relationship as regards
food between the different cave forms, an interdependence so complete
that were certain groups removed others must perish because of their
dependence upon them.

The habits of cave species so far as learned are exactly similar to
the habits of their near relatives outside of caves. This statement

GENERAL CONSIDERATIONS.

93

applies equally well to the food, each species taking the same food as
its epigean relatives. Attention has been called to this fact throughout
the paper. Anopthalmus is closely related to the genus Trechus, whose
almost innumerable species in Europe are much given to living in deep
crevices in the earth or under large stones, not to mention about 60
species of this genus which frequent caves. Species of Platynus live
in damp places in ravines and under stones along streams. Species of
Quedius live in damp, shady places, as do also the members of the
dipterous family Helomyzida?. Psychoda, outside of caves as well as
within caves, lives under debris and its larvae are upon decaying matter.
The spiders Phanetta subterranea, Willibaldi cavernicola, and Erigone
infernalis live under stones in the cave, spin little or no web, and feed
upon decaying organic matter, or catch their prey by springing upon it.
They have outdoor relatives which live in the same retired situations

Decaying
organic
matter

Earth worms, etc.

Fungi — Thysanura

Diptera iv

Mynapoda \\A Arachmda 1
^Anopthalmus j

Coleoptera

S(rLp— r——.

Ceuthophilus j

-^ Pe ro my sc u s

Larger Mammals
(on all the others)

Cyclops

Crangonyx
Caecidotea

Amblyopsis

Cambarus

FIG. 13. — Table of food relations of cave species. Reading from right to left the lines
leading from Peromyscus indicate that it may feed upon decaying matter as well
as all other animals in the cave, except those indicated in the lower part of the
figure. Earthworms feed upon decaying organic matter, and in turn are fed upon
by Spelerpes and Peromyscus.

and feed in the same manner. On the other hand, Theridium kentucky-
ense, T. porteri, and others which live near the mouth and spin snares
to catch their prey either themselves live outside of caves occasionally
or have near relatives which live outside and likewise spin snares to
catch prey. The cave species spin the web characteristic of the family.
The Ceuthophilus, in its habit of occasionally going under stones and
generally staying in cracks and crevices in the cave, shows a close
approach to the habits of the same species when living outside of caves
and to the habits of its relatives which live in caves, about wells, under
logs, and in the dark parts of cellars. The same thing is true of the
myriapod Conotyla bollmani and the thysanuran Sinellacavernarum, their
relatives outside generally living in damp places under logs, leaves, etc.
This list of illustrations might be considerably increased. The similarity
of habits of concealment, of obtaining food, and of rearing Iarva3 which

94 FAUNA OF MAYFIELD'S CAVE.

exists between cave species and their near relatives outside could hardly
be more striking.

This brings us to another point — the persistence of certain habits in
cave species— habits which in their outdoor relatives are absolutely
essential to self-preservation, but which in the cave species are entirely
useless. Packard (1890, 394) calls attention to this fact:

Although scarcely necessary in its changed environment, where there are no
hydrographic changes, no winter and summer, and few enemies to contend with, the
most aberrant form, the completely eyeless form Anthrobia of Mammoth Cave, still
spins a silk cocoon around its eggs; while in Weyer's Cave Nesticus pellucidus Emerton
spins a cocoon for its eggs; and either this species or its fellow troglodyte Linyphia
incerta Emerton, or both species, spin a weak, irregular web, consisting of a few
threads. Is not this a useless habit, the simple survival of ancestral traits?

W. P. Hay (1902, 436), regarding this point, says:

I would regard the habit of living under stones of Cambarus hamulaius and
Csecidotea richardsonse as a primitive instinct to which the animals cling in spite of
the fact that it is useless.

Eigenmann (1899, 477) says:

Typhlichthys, living in total darkness, has retained the habit of staying under
floating boards, sticks, and stones. Miss Hoppin noticed that Troglichthys swims with
its back to the side of the aquarium, and I have repeatedly noted the same in the young
of Amblyopsis up to 50 mm. and the still younger Amblyopsis frequently hides under
rocks.

Many cave animals habitually live under stones or debris. This is
the habit of the true cave spiders, of the cave Staphylinidae, and to
some extent of the thysanuran Sinella and of Ceuthophilus. The prev-
alence of this habit of concealment could scarcely be over-emphasized.
Small indeed would be one's collections, except for a few of the Diptera
and the Ceuthophilus, did he not hunt out these places of concealment.
One could as successfully collect in the woods in winter without looking
under the bark of trees, under logs, and other such places as collect in
caves without searching in cracks of the rock and under pieces of stone
and other debris. It is unnecessary for these animals to take to such a
habit to escape enemies, cave air is generally moist, and there is no
sunlight to avoid, hence the habit appears to serve no useful purpose.
The cave spiders spin small webs, which, certainly in the case of Phanetta
subterranea and Erigone infernalis, are entirely useless to catch prey or,
as near as can be determined, to serve any useful purpose whatsoever.
Phanetta subterranea spins a compact disk-shaped cocoon when the need
for a cocoon to protect the eggs or young is not apparent. These are
ancestral habits which, though once useful, are no longer so; yet they
persist in these troglodytes with a remarkable tenacity.

ORIGIN OF CAVE LIFE. 95

ORIGIN OF CAVE LIFE.

There are three fundamental questions which deserve attention in
considering the origin of the present cave fauna: (1) How did the
animals get into the caves? (2) What was their condition when they
entered the cave? and (3) How have they reached their present con-
dition?

HOW DID THE ANIMALS GET INTO THE CAVES ?

Most writers upon the origin of cave life make no mention of how
animals have reached caves, but confine their discussion to the modifica-
tions and the causes of the modifications which animals undergo after
having become cave inhabitants.

There is practical agreement that cave animals have descended from
non-cavernicolous forms. There is a difference of opinion as to the precise
manner of their original entrance into caves and much discussion as to
the causes of their modifications.

Lankester (1893, 389) explains the colonizing of caves and the origin
of blind cave forms as follows:

This instance [that of the blind cave animals] can be fully explained by natural
selection acting on congenital fortuitous variations; many animals are thus born with
distorted or defective eyes whose parents have not had their eyes submitted to any
peculiar conditions. Supposing a number of some species of arthropod or fish to be
swept into a cavern or to be carried from less to greater depths in the sea; those
individuals with perfect eyes would follow the glimmer of light and eventually escape
to the outer air or the shallower depths, leaving behind those with imperfect eyes to
breed in the dark place. A natural selection would thus be effected. In every suc-
ceeding generation (bred in the dark place) this would be the case, and even those
with weak but still seeing eyes would in the course of time escape, until only a pure
race of eyeless or blind animals would be left in the cavern or deep sea.

No one who has been within a limestone cavern and has thus gained
a conception of what a sudden transmission from epigean to subter-
ranean conditions must mean to an animal accustomed to secure food
by means of its eyes can seriously consider such an origin of cave
inhabitants. Further, as Eigenmann (1900, 57) suggests, cave animals,
with or without eyes (and their near relatives outside too, for that
matter), do not "follow the glimmer of light," but shun it.

Animals accidentally driven into caves are often found. The list
from Mayfield's Cave includes 17 strays belonging to 6 different orders.
Eigenmann (1900, 55, 57) cites another case, that of sunfishes carried
into caves. But there is no evidence whatever that any epigean species
has thus become established in a subterranean abode.

96 FAUNA OF MAYFIELD'S CAVE.

Packard (1894, 744-745) also introduces the element of accident, but
in a much more plausible way:

Where, however, individuals with more or less defective eyes should breed with
normal mates, any tendency to the transmission of such defects would be wiped out
by the swamping effects of crossing, owing to the immense preponderance of normal,
vigorous forms with perfect vision. The whole tendency in nature in the upper
world of light is to weed out such sporadic, defective forms. But in limestone regions
honeycombed with caves and permeated with subterranean streams, like those in the
Mediterranean regions, France, Spain, and Australia, or in those of Southern Indiana,
Virginia, Kentucky, and Missouri * * * in such regions as these, there exist the
conditions favorable to the origination and perpetuity of blind forms. To give an ex-
ample, eyed geodephagous beetles, such as the species of Trechus, of which there are
so many in Southern Europe, accustomed to burrowing in the soil under stones, when
carried down by various accidents into dark crevices or into caves from which they
are unable to extricate themselves, and too hardy and vigorous to succumb to the
deadly effects of a life in perpetual darkness, and with, perhaps, already partially
lucifugous habits, such forms under these changed conditions survive, breed, and mul-
tiply, finding just enough vigor to propagate their kind. We can easily imagine that
in time, and indeed no very long period, the newcomers would soon become adapted to
their new surroundings, an environment abnormal both from the absence of light,
and from the lack of predaceous forms to devour them; and they would live on, weak,
half -fed, half -blind, forced to make their asylum in such forbidding quarters.

But even this is not the most plausible method of explaining the
origin of cave species. Animals such as the geodephagous beetles men-
tioned above, if accidentally carried into caves, would probably establish
themselves there and I see'no objection to the possibility of the occasional
introduction into a cave by a flood or other means of a species fitted for
cave life.

The beetles referred to are light-shunning creatures and constantly
crawl about under stones and in crevices in the earth. Certainly in a
limestone region ' 'honeycombed with caves and permeated with subter-
ranean streams," as Packard suggests (and there are perhaps more
passages, many of them opening to the surface, no longer followed by
streams except in times of considerable rain, than there are passages that
always contain water), these beetles would have every opportunity to
enter underground channels in their ordinary search for food, and there
can be no reason for supposing that they would be more likely to reach
underground passages by accidental bodily transportation rather than
by migration in following their usual habits. Without denying the
possibility of the former method, it must be said that it can, at best,
be but an unusual method of the origin of cave forms, while the latter
method of entry into caves is not only the more plausible, but is sup-
ported by good evidence.

ORIGIN OP CAVE LIFE. 97

Cambarus bartoni and Crangonyx gracilis as shown before, have
selectively entered Mayfield's Cave and without constraint have remained
there. They can escape at any time. They remain within the cave
because conditions are suitable to their existence. Such is also true of
Theridium kentuckyense, Theridium porteri, and Meta menardi, which
live in twilight never far from the mouth, while Nesticus carteri, Phanetta-
subterranea, andErigone infernalis, while occasionally found in twilight,
are less abundant there than in absolute darkness far within the cave.
These spiders enter caves and remain there because they find conditions
favorable. No physical barrier prevents their escape from caves.
Crangonyx gracilis and Cambarus bartoni could not have been introduced
into Mayfield's Cave by accident, since to get into the cave from outside
they must migrate against the current. It might be said that these
forms became cave inhabitants through chance introduction into another
cave and extended their distribution to Mayfield's Cave. I consider
this very improbable and even impossible. It is certainly impossible
that Theridium, porteri and Theridium kentuckyense should have arisen
as cave inhabitants elsewhere by chance and migrated to Mayfield's.
In their present condition they could not exist far within a cave, and if
they remain near the mouth escape is always easy.

On the contrary, the fact that widely distributed species found in
epigean streams enter caves wherever such exist is strong evidence
that caves become populated by voluntary immigrants.

That species which live in caves belong to groups which are given
to life in dark and damp places is further evidence that caves become
inhabited by voluntary immigration. The Carabida? and Staphylinidae
compose nearly the whole of the coleopterous cave fauna. Many species
of both of these families live under stones and in damp ravines and in
crevices in the earth, under logs, and in similar places. The Carabid
genus Platynus, with 4 species taken occasionally in Indiana caves, is a
good illustration of the tendency of certain adapted groups to enter
caves and become cave inhabitants. Species of this genus live habitually
in retired places along ravines and in similar situations out of doors.
A closely related genus, Pristonychus, has 6 representatives in Euro-
pean caves.

A similar case is seen in the family Theridiida?, 12 species of which
have been taken in Mayfield's Cave alone. Three other species have
been taken in other Indiana caves, 5 others from caverns of Kentucky,
while 20 species of this same family are recorded from European caves.
Of this family the genus Nesticus has 2 representatives in Kentucky, 1
in Indiana, and 3 in European caves. The old genus Linyphia has 1
Indiana, 3 Kentucky, and 10 European cave species. This family "con-

98 FAUNA OF MAYFIELD'S CAVE.

sists of a great number of species of spiders living for the most part in
shady woods, among the lower branches of plants, under leaves, and in
caves and cellars. " (Emerton, 1902, 134.)

Throughout this paper constant mention has been made of the rela-
tives of cave species and in nearly every case cave species have been
shown to have relatives either in caves or in similar dark, moist places.

Species of animals found in caves belong to families and genera
which live in more or less dark and shady places outside of caves.
These families and genera are predetermined, so to speak, toward cave
life, and it is from these groups and other such groups that cave faunas
are derived.

Further, in many cases, other species of the same groups to which
cave species belong are adapted for cave life but have not found a cave
to occupy, and hence are not cave species. The Amblyopsidse (cf . Cox,
1905, 379) agree in having a relatively high degree of development of
the tactile ridges and a low grade of development of the eye. Of the
8 species known all but 2 are inhabitants of caves and those two are ap-
parently capable of cave existence. One of them lives in the rice
ditches in South Carolina and in the Dismal Swamp and in other similar
places on the Atlantic Coast. The other has been found only in Illinois
in a spring which is the outlet of an underground stream. It approaches
the Chologaster agassizii of Mammoth Cave in its adaptations for
underground existence, having been shown capable of securing its food
without the use of its eyes (Eigenmann, 1900-6, 402). The California
goby (Eigenmann, 1890) , of another family of fishes, lives in holes in
the beach between tide-marks. It is lighter in color and has more
degenerate eyes than the C. agassizii of Mammoth Cave. H. Garman
(1892, 241) found a species of Anopthalmus (A. horni) in and about
crevices of the rocks in an open quarry at Lexington, Kentucky. Sayce
(1901) has described 3 species of blind crustaceans— an amphipod and
2 isopods— from a surface stream in Australia. There are every year
being described new species of blind and pigmentless crustaceans from
wells, springs, and underground streams in various parts of the world.
They are known from the United States, England, several countries of
continental Europe, New Zealand, Tasmania, Australia, and Algeria, in
some of which countries there are no caves whatever. Specialists
doubtless are familiar with animals of many groups equally as well
suited to cave life but which live in caveless regions.

Animals become cave inhabitants because they are structurally and
physiologically suited to life in caves. They are predetermined cave
inhabitants and enter caves because there they find conditions suit-
able for their existence.

ORIGIN OF CAVE LIFE. 99

Eigenmann (1900, 57) first applied this principle to the occurrence
of blind fishes in caves:

On account of the structure of their eyes and their loss of protective pigment
they are incapable of existence in open waters. Their structure is not so much due
to their habitat as their habitat is to their structure and habit.

Davenport (1903, 19), discussing the same group, remarked:

Moreover, this family includes species that are structurally especially fitted for cave
life, even when they occur in regions where there are no caves and never have been
any. They shun the light, and live in crevices and under stones. Their bodily conditions
fit them for cave life, and when, in their constant search for dark holes, some of them
succeeded in getting into caves, they naturally thrived there.

Davenport (1903, 19), in explaining the origin of certain beach
fauna, stated this principle of adaptation as follows:

* * * that the structure existed first and a fitting environment was sought or
fallen into by the species having the peculiar bodily condition. Thus the adaptive re-
sult is, on this theory, not due to a selection of structure fitting a given environment,
but, on the contrary, a selection of an environment fitting a given structure.

The principle applies to the origin of cave animals in general as
logically as I believe it does in the case of Amblyopsidae and of the
beach fauna.

Blind and highly modified cave species do not suddenly arise from
typical out-doors forms which some accident or even the struggle for
existence has caused to enter a cave. It is, as expressed by Davenport
(1903, 21), a segregation into the fittest environment. Animals do not
possess degenerate eyes and lack pigment because they are cave ani-
mals. The eyes have in many cases degenerated and the color disap-
peared before they entered caves. They are cave animals because their
eyes are degenerate and because they lack pigment. The greater the
degree of depigmentation and degeneration of the eyes the more the
species has become confined to the dark and the greater its tendency
to subterranean existence.

Cave animals have not arisen by accidental isolation. They are
isolated in caves and other subterranean abodes because they are unfit
for a terranean life and caves are among the possible habitats.

To sum up, this voluntary migration into caves falls into two classes.
Although the accidental origin of cave species from strays which are
not structurally adapted to life in such situations is preposterous, cave
animals do sometimes arise from the collecting within the mouths of
caves of forms more or less adapted to life in partial darkness. These
animals making their homes in the mouths of caves may become further
modified and, gradually entering farther and farther into the recesses
of the cave, may finally become highly specialized and typical cave

100 FAUNA OF MAYFIELD'S CAVE.

creatures. This view was set forth by Spencer (1893, 758; 477-478)
as follows:

The existence of these blind cave animals can be accounted for only by supposing
that their remote ancestors began making excursions into the cave, and, finding it
profitable, extended them, generation after generation, further in, undergoing the
required adaptations little by little.

This class of immigrants into caves is nicely exemplified by the
spiders living about the mouth of Mayfield's Cave.

On the other hand, cave faunas may be formed directly through
colonization by animals already highly modified and entirely suited to the
rigid conditions of remote portions of caves. This is in accord with H.
Carman's view (1892, 240) :

It is to species such as these, already fitted for life in caves, that we should look,
it seems to me, as representing the ancestors of cave species; certainly not to ordinary
species with well-developed eyes. The originals of the cave species of Kentucky were
probably already adjusted to a life in the earth before the caves were formed, and it
seems probable from some facts mentioned below that they were not very different
in character from the animals now living in the caves. I can not believe that there
has been anything more than a gradual assembling in the caves of animals adapted
to a life in such channels.

WHAT WERE THE CONDITIONS OF CAVE ANIMALS WHEN THEY ENTERED CAVES?

From the foregoing discussion it will be seen that some animals
take up their abode within caves when (e. g. , Theridium kentuckyense,
Cambarus bartoni, and Crangonyx gracilis) they are little or apparently
not at all different from outside forms. Others, such as the geodepha-
gous beetles and the Amblyopsidae, before becoming cave inhabitants
at all were probably suited even to the rigid conditions of the remote
portions of caves. Animals which take up their abode in caves are not
necessarily all in the same grade of adaptation. On the contrary, it has
been seen that all grades of adaptation between extremes of little or
no adaptation and perfect adaptation may lead to cave life.

This opens the way for the conception that there is no sharp dis-
tinction between cave faunas and other dark and shade inhabiting
faunas. This conception is abundantly justified by the blind and white
goby (Eigenmann, 1890) of the California coast, the Chologaster papilli-
ferus (Forbes, 1882) living in a spring in Illinois, the blind beetle
(Carman, 1892, 24) living about crevices in open quarries in Kentucky,
the blind amphipods and isopods (Sayce, 1901) found among driftwood
in a shaded stream in Australia, and even the very highly modified
Cascidotea stygia, which occurs sometimes under stones or leaves in
shaded ravines and about the mouths of springs in the Ohio Valley,
not to mention the occurrence throughout the world of blind and color-
less animals in wells and springs.

ORIGIN OF CAVE LIFE. 101

Nevertheless, there is a real cave fauna in the sense that in caves
these dark-seeking species are accumulated in relatively greater abun-
dance than elsewhere. Furthermore, in caves species have become more
highly adapted to cave conditions than elsewhere. In this limited
sense there is a real cave fauna hedged about and shut off from the
upper world of light.

Whether cave faunas originate by the accumulation of species
highly adapted or by the gradual colonization of species which become
gradually adapted to cave life does not seem an essential point. It is
certain that the adjustment of the cave fauna occurred in dark and
sheltered places.

The question of where the adaptation to caves has arisen has an
important bearing, however, upon the determination of the age of the
present cave fauna. Could one be sure that Amblyopsis spelasus, for
example, entered Indiana caves as they were being formed as a very
slightly modified or perhaps nearly typical shade-loving fish of open
streams, one would have a possible index of the rapidity of modifica-
tions in the development of specially numerous and large sense-papilla?
and of the rate of production of retrograde modifications in the color
and the eye. Such a means of getting at the age of the cave fauna
would be of doubtful validity unless it could be shown that conditions
before the formation of the caves were such that there would be little
or no suitable opportunity for the production of shade-inhabiting forms,
and to establish this point seems to me well-nigh impossible.

HOW HAVE CAVE ANIMALS REACHED THEIR PRESENT CONDITION?

This question necessitates a statement of the peculiarities of cave
species before the causes leading to the peculiarities can be satisfac-
torily disposed of.

THE MODIFICATIONS OF CAVE ANIMALS.

The modifications of cave animals in general need only be referred
to. They consist in the possession of more slender bodies, longer
appendages, more abundant or more highly developed sense-organs,
together with a retrograde development in pigment and organs of
vision.

It is immaterial to the discussion whether these adaptations were
acquired in the cave or before the species entered the cave.

Amblyopsis is white, has eyes entirely functionless, and has the
sense-papilla? on the head more highly developed than in its near rela-
tives within or outside of caves. Czecidotea stygia has a more slender
body, more slender appendages, a greater number of sensory setse, is
white in color, and has mere rudiments of eyes as compared with its

102 FAUNA OF MAYFIELD'S CAVE.

nearest out-door relative, its parent species, the normal terranean
Asellus communis. Cambarus pellucidus has very degenerate eyes, is
entirely white, quite slender in body, and has long, delicate appendages,
and its sensory setae are longer than in ordinary crayfish. These are
familiar and typical cases.

Some less extensive but very interesting modifications are those in
species found living within Mayfield's Cave and differing very slightly
from the same species when found living outside of caves in the im-
mediate neighborhood. A comparison of individuals of Cambarus bar-
toni living out-doors with those in Mayfield's Cave shows that those in
the cave have less pigment and that there are other structural differ-
ences. In the cave forms the antennae average 104.55 per cent the
length of the body, while in the outside forms the antennae are only
92.66 per cent of the length of the body. The series obtained from the
cave was not large, but the characters of having less pigment and
longer antennae were quite constant. A similar case is seen in Cran-
gonyx gracilis. Cave specimens of this species are without dark pig-
ment and reach a larger size than the same species living outside in
streams and pools near Bloomington.

The species of Theridiidae present modifications both in the size of
the eyes and in the amount of pigment in direct proportion to the ex-
tent to which the species is subject to cave conditions. These modifi-
cations are discussed in detail and other modifications are noted in the
discussions of the different species. Packard (1888, 112, 120) noted
the same relation between the extent to which an animal lives within a
cave and its modification. Packard (1888, 120), Hamann (1896, 6),
Carpenter (1895, 29), Chilton (1894, 257, 258), and others have cited
other cases of animals living in caves possessing certain modifications
as compared with the same species outside of caves.

There is very marked and striking uniformity in the modifications
of certain cave animals. Convergence is noted in a number of cases
among cave species. Eigenmann (1899) calls attention to a remarkable
case of convergence between Typhlichthys subterraneus and Troglichthys
rosas. He says, in part (1899, 281) :

On the surface the specimens of Typhlichthys rosae very closely resemble Typh-
lichthys subterraneus from Mammoth Cave, differing slightly in the proportion and
in the pectoral and caudal fins. These fins are longer in rosas. It is, however, quite
evident from a study of their eyes that we have to deal here with a case of converg-
ence of two very distinct forms. The blind fish, Amblyopsis, may be left out of con-
sideration, since it is the only member of the family that possesses ventral fins.
Otherwise, it would be difficult to distinguish specimens of similar size of this species
from either subterraneus^pr rosse.

ORIGIN OF CAVE LIFE. 103

Two other blind fish (Typhlickthys osborni from Horse Cave, Ken-
tucky, and Typhlichthys wyandotte from Wyandotte Cave, Indiana)
very closely resemble T. subterraneus and the others. The different
species of cave crayfishes (Cambarus) resemble each other very closely,
so that, according to W. P. Hay (1902, 436), while C. hamulatus and C.
pellucidus differ considerably in their affinities, their appearances are so
strikingly similar ' 'that without a careful examination it would be ex-
ceedingly difficult to distinguish the two species." Another instance
of convergence is seen in the so-called "cave crickets." Speaking of
these cave-inhabiting Ceuthophili, Sharp (1899, 321) says:

The species with this habit, though found in the most widely separated parts of
the world, have a great general resemblance, so that one would almost suppose the
specimens found in the caves of Austria, in the Mammoth Cave of Kentucky, and in
the rock-cavities of New Zealand to be one species, although they are now referred
by entomologists to different genera.

There is not, however, a general uniformity in the peculiar modifi-
cations of cave animals. Some cave species have one structure highly
modified, others another, still others appear to have no highly modified
organs to compensate for the loss of visual organs. Chilton (1894, 269),
reviewing this point with reference to the New Zealand subterranean
Crustacea, shows that while some have additional sensory seta3, others
have not, that while some are slender-bodied, others are not, and while
the appendages in certain cases are lengthened, in others they are not,
' 'while there is no sign of a similar modification in Crangonyx compac-
tus, which has the body normally stout, the antenna and legs of only
moderate length, and the uropoda even somewhat short and stumpy"
(1894, 269). This same Crangonyx compactus is white, has very de-
generate eyes, and is an exclusively subterranean form. Further,
Chilton compares various forms and shows that when elongation of an
appendage takes place, it is in one joint in one species and in another in
another species.

The question arises whether the modified cave forms are entitled to
rank as distinct species and genera. They are not distinct species in
the de Vriesian sense. According to de Vries (1905) a new species
must have a new character. Certain cave spiders lack pigment and their
eyes are poorly developed, but they have acquired no new character.
So with other cave animals. The genus Anopthalmus has been discarded
by European authorities because species have been found showing every
gradation between a species without trace Jof eye or optic lobes, atypical
Anopthalmus, and species of Trechus with eyes. Cope's genus Orconectes
(Cope, 1872, 173), proposed for the species of blind crayfish, has failed
to meet with acceptance. Csecidotea differs from Asellus in about the

104 FAUNA OF MAYFIELD'S CAVE.

same manner that Anopthalmus differs from Trechvs and that Orconectes
differs from Cambarus. Csecidotea is more slender, lacks eyes and pig-
ment, and has longer and more slender appendages, but is otherwise
much like Asellus. The genus Csecidotea is probably as good as Orco-
nectes or Anopthalmus, and no better. It is generally accepted, however.
On the other hand, these genera are just as good as hundreds of others
which have met with general acceptance and which are not questioned.

CAUSES LEADING TO THE MODIFICATIONS OF CAVE ANIMALS.

At present I do not care to enter into any extended discussion of
the causes of the modifications of cave animals. The more slender
appendages, the more slender bodies, and the increase in number
or development of the sense-papillae and sensory setae in cave ani-
mals may be due to individual adaptations or they may be due to nat-
ural selection tending to eliminate all individuals which did not possess
variations of assistance to the animal under its unusual conditions.

It seems to me no satisfactory explanation for the loss of pigment
and degeneration of the eyes in cave animals has been offered, unless
we grant the influence of the environment upon the individual, together
with the hereditary transmission of these acquired characters. The
validity of the transmission of the effects of disuse is the contention of
Spencer (1893), Packard (1888, 1894), and Eigenmann (1899, 1903);
but as Eigenmann (1903, 197) admits:

There has always been and is yet a serious objection to the latter conclusion [i. e.,
the hereditary transmission of the effects of disuse upon the eye], because the method
of the transmission of functional adaptations to the organization of the egg so as to limit
or extend its powers is not known.

Until experiments have shown that an outside form kept under cave
conditions may be modified by its life there and this modification trans-
mitted to its offspring, the explanation of the origin of blind and color-
less cave forms through the heredity of the effects of disuse and of the
influence of lack of light can not be very satisfactory.

Adaptive structures are explained by Osborn as follows (1897, 584) :

All the individuals of a race are similarly modified over such long periods of time
that very gradually congenital or phylogenetic variations which happen to coincide
with the ontogenetic adaptive variations are selected.

Further, the same authority says (1897, 585) :

The law of determinate variation is observed to operate with equal force in cer-
tain structures, such as the teeth, which are not improved by individual use or exercise
as in structures which are so improved.

He is thus led to a belief in "fundamental predispositions to vary
in certain directions."

ORIGIN OF CAVE LIFE. 105

The occurrence of animals in caves everywhere and of similarly mod-
ified animals in every conceivable situation in which such animals are
capable of living suggests that these modifications, like those of the teeth
of mammals, may be congenital, and due to the inherent predisposition of
protoplasm to produce variations in a given direction, these variations
in the case of the animals under consideration forcing them into secluded
places, caves, and similar situations, where further modifications may
have occurred through similar causes, natural selection serving to elimi-
nate individuals modified in an unfavorable way.

There is a serious difficulty, however, in the way of offering such an
explanation for the degeneration of the eyes of the cave fishes, in which
it has been conclusively shown (Eigenmann, 1899) that the more active
elements, the muscles, retinal elements, etc. , degenerate most, and the
passive scleral cartilages little or not at all. But, on the other hand, all
cave animals do not suffer a like degeneration of the eye, some of those
living under a typical cave environment possessing eyes fairly well de-
veloped. These modifications of cave animals are hard to explain on
the hypothesis that they are the direct result of the environment when
a cave environment is so absolutely uniform.

To take an illustration, Chologaster agassizii, possessing eyes, lives
in Mammoth Cave with Typhlichthys and Amblyopsis, both practically
eyeless. It is hard to explain, if degeneration in the two cases is pro-
duced by the effects of disuse becoming hereditary, why the same
result has not occurred in the other case.*

The case of Chologaster, a species really highly suited to cave con-
ditions, is readily understood if looked at from the standpoint of deter-
minate adaptations which suited the animal to cave life but did not
involve an extensive degeneration of the eyes.

There is possibly equally good reason to suppose that the Theridiidss
about the mouth of Mayfield's Cave are becoming cave inhabitants
through determinate congenital variations which cause them to seek a
favorable habitat farther arid farther within the cave as there is to sup-
pose that they entered the cave because they were somewhat adapted
for cave life and that they are being gradually further modified by the
influence of the environment.

In conclusion, it seems clear that cave animals have originated from
forms living outside of caves and that their occurrence in caves is not
due to accident but to their finding, in their ordinary movements, an

*I am aware that to this very point the answer has been made that Chologaster
agassizii has not been in caves as long as Typhlichthys and hence has been less mod-
ified. To establish this point is, however, as difficult as to establish the heredity of
acquired characters.

106 FAUNA OF MAYFIELD'S CAVE.

environment suited to their morphological and physiological peculiari-
ties. There seems to be good evidence that caves have become colo-
nized by animals slightly adjusted to cave conditions, and by others
more or less highly modified and correspondingly well suited to cave
habitation. It is difficult to explain the adaptations of cave animals by
the action of natural selection acting upon ordinary fluctuating vari-
ations. If we assume the heredity of the effects of disuse and lack of
light there is afforded an hypothetical solution of the degeneration of
eyes and pigment. In view of the manifest difficulties in connection
with the theory of the heredity of acquired characters, it seems that
the equally well supported theory of determinate, cumulative variations
may account for the origin of cave forms and their further modifica-
tion after becoming cave forms.

SUMMARY.

(1) Conditions of temperature and moisture are quite uniform in the

innermost parts of the cave.

(2) A small cave presents the same diversity and all the essential con-

ditions of a large cave.

(3) There is little seasonal change in temperature in remote parts of

the cave.

(4) Air-currents through the cave are directly instrumental in effect-

ing whatever seasonal changes in temperature the cave under-
goes.

(5) The activity of cave animals is profoundly affected by changes in

temperature, but the changes are too small to be effective in re-
mote parts of the cave. Near the mouth, where the changes are
considerable, many animals retire into cracks and crevices with
the lower winter temperature, while with all cave species there
seems a direct relation between the temperature in the cave and
their activity.

(6) A small cave affords practically the whole cave fauna of a cave

region.

(7) Species do not often get into caves by accident.

(8) Temporary residents are perfectly at home and come into and

leave the cave freely. Except some mammals, they usually do
not go beyond twilight.

(9) Insects hibernate in the cave in considerable numbers, but seldom

in twilight.

(10) Many cave inhabitants sustain a certain definite relation to the

light, some living only in absolute darkness, others in darkness
and dim twilight, still others only in a certain degree of twilight.

SUMMARY. 107

(11) The distribution of animals within a cave, aside from consider-

ations of light and temperature, is influenced by moisture, the
presence of organic matter, and the presence of places of conceal-
ment.

(12) Many of the species taken within a cave do not form a part of

the true cave fauna; indeed the permanent and true cave species
form but a small part of the entire cave fauna.

(13) There is no hard and fast line of division between a cave fauna

and the outdoor fauna in the same region.

(14) Highly modified and truly cavernicolous animals are not neces-

sarily confined to caves, some of them occasionally being found
in the dark or in twilight in other situations. There are species
of animals adapted for cave life, but living in regions where no
caves exist.

(15) Temporary residents and some of the less highly specialized of

the permanent residents are apparently recent additions to the
cave fauna and local in their cave distribution. Highly special-
ized cave inhabitants are widely distributed and usually appar-
ently cave forms of long standing.

(16) For food cave animals are directly or indirectly dependent upon

organic matter being carried into the cave from outside.

(17) There is a close interdependence between cave animals, so that

if some forms were eliminated others would probably perish.

(18) The food and habits of cave species are similar to the food and

habits of their near relatives living in other situations.

(19) Certain ancestral, altogether useless habits persist in cave species.

(20) Certain dark or shade-loving groups of animals exhibit a tendency

to become cave inhabitants.

(21) The nearest relatives of cave forms are nocturnal, or are dark or

shade loving species.

(22) Accident plays no part, or at most a very small part, in the origin

of cave inhabitants.

(23) Animals have reached caves by active migration into places where

they find conditions suitable for their existence.

(24) Cave species were fitted for cave life before entering caves.

(25) Cave species may arise from highly modified animals living out-

side of caves going directly into the deeper parts of caves, or
they may arise gradually by the collecting about the mouths of
caves of forms slightly modified.

(26) Cave animals possess certain characteristic modifications — more

slender bodies, longer appendages, more abundant or more highly
developed sense-organs, and a retrograde development in pig-
ment and organs of vision.

108 FAUNA OF MAYFIELD'S CAVE.

(27) Animals which, through more or less adaptation, have sought life

in caves, usually become more modified after taking up such life
and are thus nicely adjusted to cave conditions.

(28) Cave animals are modified in direct proportion to the extent to

which they live under true cave conditions, or, to state it in the
reverse sense, which is just as true, cave animals are confined to
conditions similar to those of a cave, in so far as they have under-
gone modifications unfitting them for life in the open.

(29) The rate of modification of cave animals seems hard to determine,

in view of the fact that caves have been populated by animals in
all stages of adaptation to cave existence, and the age of a cave
furnishes no safe criterion as to the length of time the cave
species may have undergone modifications adapting it for cave
life.

(30) It is doubtful if cave genera and species are new in the de Vries-

ian sense, for they possess only modified or retrograde characters,
not new characters.

(31) There is a certain amount of striking uniformity in the modifica-

tions of cave animals, and in several cases convergence occurs;
on the other hand, there is too much diversity in the modifications
of cave forms for the direct effect of the uniform environment
readily to account for them all.

(32) Cave genera and species have probably not arisen as mutations,

for cave specimens of epigeal species differ from the latter
slightly in characters that better adapt them to cave conditions.

(33) Most explanations of the origin of the retrograde modifications of

cave animals involve the heredity of acquired characters.

(34) In view of the difficulties in the way of accepting the theory of

the hereditary transmission of acquired characters, it seems that
the theory of the cumulative effect of determinate variations may
explain the adaptations for cave life which make possible the
origin of cave inhabitants, and may account for their further
modifications after becoming cave forms.

BIBLIOGRAPHY. 109

BIBLIOGRAPHY.

[This list is not intended for a list of papers on cave faunas. It is meant to
include the more important titles consulted in connection with this paper. For further
bibliography of cave life see Packard (1888 and 1894), Chilton (1894 and 1900), and
Eigenmann (1899 and 1904) . Under the consideration of each species is given the
references to the literature referring to the species so far as known to me, except
in cases of many species whose importance as cave inhabitants is not great, in which
cases only references to catalogues and lists and to literature mentioning the species
as a cave inhabitant are given.

ABSOLON, P. C. K.

1900. Einige Bemerkungen iiber mahrische Hohlenf auna. <Zool. Anz., Bd.

XXIH, 1-6.
ALDRICH, J. M.

1896. On a collection of Diptera from Indiana caves. <Rep. Ind. Geol. Surv.,

xxi, 186-190.

1905. Catalogue of North American Diptera. <Smithsonian Miscellaneous Collec-
tions, XLVI, No. 1444, 1-680.
ANDERSON, A.

Blind animals in caves. < Nature, XLVII, 439.
BAILEY, L. H.

1894. Neo-Lamarckism and neo-Darwinism.<Am. Nat., xxvn, 661-678.
BANKS, NATHAN.

1893. The Phalangida Mecostethi of the United States. <Trans. Am. Ent. Soc.,

xx, 149-152.
1896. Indiana caves and their fauna: Spiders. <Rep. Ind. Geol. Surv., xxi,

202-205.
1904. A treatise on the Acarina or mites. <Proc. U. S. Nat. Mus., xxviil,

1-114, figs. 1-201.
BENEDICT, J. E.

1895. Preliminary description of a new genus and three new species of crustaceans

from an artesian well at San Marcos, Texas. <Proc. U. S. Nat. Mus.

xvill, 615-617.
BILIMEK, D.

1867. Fauna der Grotto Cacahuamilpa in Mexico. <Verh. Zool.-bot. Ges., Wien,

xvii, 901-908.
BLATCHLEY, W. S.

1896. Indiana caves and their fauna.<Rep. Ind. Geol. Surv., xxi, 121-212.

BOLLMAN, C. H.

1888. Catalogue of the Myriapods of Indiana. cProc. U. S. Nat. Mus., XI, 403-
410.

BOULENGER, G. A.

1893. Blind animals in caves. < Nature, XLVII, 608.
CALL, R. E.

1897. Flora and fauna of Mammoth Cave, Kentucky. < Am. Nat., xxxi, 377-392,

pis. x, xi.
CALL, R. E. and HOVEY, H. C.

The Mammoth Cave of Kentucky. Manual.
CARPENTER, G. H.

1895. Animals found in the Michelstown Cave. <Irish Nat., IV, 25-35, pi. i.
CHEVREUX, ED.

1901. Amphipodes des eaux souterranes de France et d'Algerie.<Bull. Soc.

Zool. France, Tom. 26, 168-179, 197-205, 211-222, 15 figs.
CHILTON, C.

1882. Notes on and a new species of subterranean Crustacea. <Trans. New Zeal.
Inst., XV, 87-92, 1 pi.; abstract in Nature, September, 28, 1882.

1894. The subteranean Crustacea of New Zealand. <Trans. Linn. Soc., 2d ser.,

Zool., VI, pt. 2, 163-284, pis. xvi-xxiii.

1900. The subterranean Amphipods of the British Isles. <Journ. Linn. Soc.
London, Zool., xxvm, 140-161, 3 pis.

110 FAUNA OF MAYFIELD'S CAVE.

COLLETT, J.

1873. Geological Report of Lawrence County (list of species from Connelley's,
Hamer's, and Donaldson's caves). <Rep. Ind. Geol. Surv., v, 1873, 305;
Am. Nat., VI, 1892, 406.
COPE, E. D.

1869. Observations on some myriapods found in and near caves of the United

States. CProc. Am. Phil. Soc. for 1869, 178, 179-182.
1871. Life in the Wyandotte Cave. < Ann. Mag. Nat. Hist. , ser. 4, vill, 368-

370; Indianapolis Journal, Sept. 5, 1871.
1872a. On the Wyandotte Cave and its fauna. "Am. Nat., vi, 406-422, figs. 109-

116; Rep. Ind. Geol. Surv. for 1872, m and IV, 157-182, 7 figs.
18726. Descriptions of species from the Wyandotte Cave; also from Mammoth

Cave.<Rep. Ind. Geol. Surv., iv, 173-182, 1 fig.

1881. The Fauna of the Nickajack Cave. < Am. Nat., vi, 1-17, pi. vii, 13 figs.
Cox, U. O.

1904. A revision of the cave fishes of North America. <TRep. Bureau Fisheries

for 1904, 379-393, pis. i-vi (Jan., 1906).
CUNNINGHAM, J. I.

1893. Blind animals in caves. <Nature, XLVII, 439.
DAVENPORT, C. B.

1903. The animal ecology of the Cold Spring sand spit, with remarks on the

theory of adaptation. <Decennial Pub. Univ. of Chicago, vol. x, 1-22,

7 figs.
DENNY, A.

See Eigenmann, C. H., and Denny, A.
DYAR, HARRISON G.

1902. List of North American Lepidoptera and key to the literature of the order.

Bull. U. S. Nat. Mus., LII, xix, 723.
EIGENMANN, C. H.

1890. The Point Loma blind fish and its relatives. <Zoe, I, 65-72, pis. ii-iii.
1897a. The Amblyopsidae, the blind fish of America. < Rep. Brit. Assoc. Adv.

Sci. for 1897, Toi-onto meeting, 685-686.
18976. The origin of cave faunas. Abstract. <Proc. Ind. Acad. Sci. for 1897,

229-230 (1898).
1897c. Amblyopsidae and the eyes of blind fishes.<Proc. Ind. Acad. Sci. for 1897,

230-231 (1898).
1898. Degeneration in the eyes of Amblyopsidas, its plan, process, and causes.

Proc. Ind. Acad. Sci. for 1898, 239-241 (1899).
1899a. A case of convergence. 'Science, n. s., ix, 280-282; Proc. Ind. Acad. Sci.

for 1898, 247-251, 3 figs. (1899).

18996. The blind fishes of North America.<Pop. Sci. Mon., LVI, 473-486.
1899c. The eyes of the blind vertebrates of North America. I. The eyes of the

Amblyopsida3.<Archiv. f. Entwickelungsmechanik, vm, 545-618, 5 pis.
1899d. Notes on the blind fishes. Science, n. s., ix, 370.
1899e. Cave animals: their character, origin, and their evidence for or against

the transmission of acquired characters. In brief.<Proc. Am, Assoc.

Adv. Sci., Columbus meeting, 255; Science, n. s., x, 883.
1899/ The blind fishes. Biol. Lectures, Marine Biol. Lab. at Woods Hole, Lecture

VIII, 111-126 (1900).
18990. Degeneration in the eyes of the cold-blooded vertebrates of the North

American caves.<Troc. Ind. Acad. Sci. for 1899, 31-46, 10 figs. (1900) ;

Science, n. s., XI, 492-503, 10 figs.

1900a. The structure of blind fishes. Pop. Sci. Mon., LVII, 48-58, figs. 1-8.
19006. Causes of degeneration in blind fishes. < Pop. Sci. Mon., LVII, 397-405.
1900c. A contribution to the fauna of the caves of Texas. < Science, n. s., XII,

301-302.
1901. Description of a new cave salamader, Spelerpes stejnegeri, from the caves

of southwestern Missouri. Trans. Am. Mic. Soc., XXH, 189-192, pis. xxvii,

xxviii.

1904. The eyes of the blind vertebrates of North America. V. The history of the

eye of Amblyopsis from the beginning of its development to its disinte-
gration in old age.<^E. L. Mark anniversary volume, 167-204, pis. xii-xv.
Abstract in Proc. Ind. Acad. Sci. for 1901, 101-105.

BIBLIOGRAPHY. Ill

EIGENMANN, C. H.— Continued.

1905. Divergence and convergence in fishes. <Biological Bulletin, vm, 1905, 59-

66, 4 figs.
EIGENMANN, C. H. and DENNY, A.

1900. The eyes of the blind vertebrates of North America. III. The structure and

ontogenic degeneration of the eyes of the Missouri cave salamander.
Biol. Bull., II, 33-40, 1 pi.
EIGENMANN, C. H. and YODER, A. C.

1898. The ear and hearing of the blind fishes. <Proc. Ind. Acad. Sci. for 1898,

242-246, 11 figs. (1899).
EMERTON, J. H.

1875. Notes on spiders from Kentucky, Virginia, and Indiana. < Am. Nat., ix,
1875, 278; Packard's cave fauna of North America. <Mem. Nat. Acad.
Sci., iv, 1888, 57-58.

1902. The common spiders of the United States. Boston, 1902.
FAXON, W.

1885. A revision of the Astacidae.<Mem. Mus. Comp. Zool., x, No. 4, 1-186,

pis. i-x.
1889. Notes on North American cray fishes. <Proc. U. S. Nat. Mus. XH, 619-634.

1898. Observations on the Astacidae.<Proc. U. S. Nat. Mus., xx, 643-694.
FORBES, S. A.

1882. The blind cave fishes and their allies.<Am. Nat., xvi, 1-5.
CARMAN, H.

1892. The origin of the cave fauna of Kentucky, with a description of a new

blind beetle. <Science, xx, 240-241.

1894. Two cave beetles not before recorded.<Psyche, VII, 81-82, figs. 1-2.
CARMAN, S.

1889. Cave animals from southwestern Missouri.<Bull. Mus. Comp. Zool., xvil,

No. 6, 225-240.
HAMANN, O.

1896. Europaische Hohlenfauna. Jena, 1896.
HARRIS, J. A.

1903. An ecological catalogue of the crayfishes belonging to the genus Cambarus.

Kansas Univ. Sci. Bull., II, No. 3, 51-187.
HAY, O. P.

1882. Notes on some fresh- water Crustacea, together with descriptions of two

new species.<Am. Nat., xvi, 143-146, 241-243.

1891. The bactrachians and reptiles of Indiana. <Rep. Ind. Geol. Surv., xvil,
407-610.

1894. The lampreys and fishes of Indiana.<Rep. Ind. Geol. Surv., xix, 147-246.
HAY, W. P.

1891. The Crustacea of Indiana. <Proc. Ind. Acad. Sci., 147-150.

1893. Observations on the blind crayfishes of Indiana, with a description of a

new subspecies, Cambarus pellucidus testii. <Proc. U. S. Nat. Mus., xvi,
283-286, pis. xliv, xlv.

1895. The crawfishes of Indiana. <Rep. Ind. Geol. Surv., xx, 475-507, 15 figs.

1896. On a collection of crustaceans from Indiana caves, annotations by W. S.

Blatchley.<Rep. Ind. Geol, Surv., xxi, 206-210.

1899. Description of ia new species of subterranean isopod. <Proc. U. S. Nat.

Mus., xxi, 871, 872-1 pi.

1901. Two new subterranean Crustacea from the United States. <Proc. Biol. Soc.

Wash., xiv, 179-180.
1902a. Observations on the crustacean fauna of the region about Mammoth

Cave, Kentucky. <Proc. U. S. Nat. Mus., xxv, 1902, 223-236, 1 fig.
19026. Observations on the crustacean fauna of Nickajack Cave, Tennessee, and

vicinity. <Proc. U. S. Nat. Mus., xxv, 1902, 417-439, figs. 1-8.
PENTZ, N. M.

1875. The spiders of the United States. Ed. Burgess, Boston Soc. Nat. Hist., 1875.
HORN, G. H.

1883. Synopsis of North American species of Anopthalmus. <Trans. Amer. Ent.

Soc., x, 270-272; Mem. Nat. Acad. Sci., IV, 1888, 75, 76.
HORN, G. H. and LE CONTE, J. L.
See Le Conte, J. L.

112 FAUNA OF MAYFIELD'S CAVE.

HUBBARD, H, G.

1880. Two days' collecting in the Mammoth Cave, with contributions to a study

of its fauna. <Amer. Ent, in, 34-40, 79-86, figs. 8-10, 19-24.
JORDAN, D. S.

1899. Manual of the vertebrates of the Northern United States. Chicago,

McClurg & Co.
LANKESTER, E. R.

1893. Blind animals in caves. <Nature, XLVII, 389, 486.
LEUNIS, J. T.

1883. Synopsis der Thierkunde, i, 1883; n, 1886.
LE CONTE, J. L., and HORN, G. H.

1883. Classification of the Coleoptera of North America. Smithsonian Miscel-
laneous Collections, xxvi, No. 507.
MARX, G.

1889. Catalogue of the described Araneae of Temperate North America.<Proc.

U. S. Nat. Mus., xii, 497-595.
Me COOK, H. C.

1893. American spiders and their spinning work. Acad. Nat. Sci. Phil., 3 vols.
MCNEILL, JEROME.

1887. Description of twelve new species of Myriapoda, chiefly from Indiana.

c'Proc. U. S. Nat. Mus., x, 328-334.
MILLER, GERRIT S.

1897. Revision of the North American bats of the family Vespertilionidas. North
American Fauna No. 13, Biological Survey, U. S. Department of Agri-
culture.

MURTFELDT, M. E.

1896. A cave-inhabiting moth. Rep. Ind. Geol. Surv., xxi, 191-192.
OSBORN, H. F., and POULTON, E. B.

1897. Organic Selection. <Science, n. s., vi, 583-585.
PACKARD, A. S.

1871. On the Crustaceans and insects of Mammoth Cave.<Am. Nat., v, 744-761,

figs. 124-133.

1873. On the cave fauna of Indiana.<Fifth Ann. Rep. Trus. Peab. Acad. Sci.,
Salem (for 1872), 93-97.

1875. The invertebrate cave fauna of Kentucky and adjoining States. <Am. Nat.,

ix, 274-278.

1876. The cave beetles of Kentucky.<Am. Nat., x, 283-287.

1877. On a new cave fauna in Utah.<U. S. Engineer Reports of Explorations and

Surveys West of 100th Meridian, in, 157-168.

1881. Fauna of Luray and Newmarket caves, Virginia.<Am. Nat., xv, 231, 232.
1883. A revision of the Lysiopetalidae, a family of Chilognath Myriapoda. <Proc.

Am. Phil. Soc., xxi, 177-197; Mem. Nat. Acad. Sci., IV, 1888, 58-65.
1885. On the structure of the brain of Asellus and the eyeless form Csecidotea.
'Am. Nat., xix, 85-86.

1888. The cave fauna of North America, with Remarks on the anatomy of the

brain and origin of the blind species. < Mem. Nat. Acad. Sci., IV, pp. 156,
28 pis., 21 figs.

1890. The effect of cave life on animals. <Pop. Sci. Mon., XXXVI, 389-397.

1894. On the origin of the subterranean fauna of North Amei'ica.<Am. Nat.,

xxvill, 727-751.

1900. A new eyeless isopod crustacean from Mexico.' Proc. Am. Assoc. Adv. Sci.,

49th meeting, 228; Science, n. s., XII, 300, 301.
PACKARD, A. S., and PUTNAM, F. W.

1872. The Mammoth Cave and its inhabitants ; or descriptions of the fishes, insects,

and crustaceans found in the cave. Salem, Naturalists' Agency, pp. 62,
figs. 8, pis. 2. Contains reprints of Putnam, Am. Nat., 1872; Putnam,
Ann. Rep. Peab. Acad., 1871; Packard, Am. Nat, 1871.
PARKER, G. H.

1890. The eyes of blind crayfishes. <Bull. Mus. Comp. Zool., xx, No. 5, 153-162,

plate.
POULTON, E. B.

See Osborn, H. F., and Poulton, E. B.

BIBLIOGRAPHY. 113

PUTNAM, F. W.

1871. Synopsis of the falmiy Heteropygii.<Ann. Rep. Trus. Peab. Acad. Sci.,

Salem, 15-23 (1872).

1872. The blind fishes of the Mammoth Cave and their allies.<Am. Nat., vi,

6-30, pis. i, ii, 2 text-figs.

1874. Fishes and crayfishes from Mammoth Cave.<Proc. Bost. Soc. Nat. Hist.,

xvn, 222-225.
1875cr. Remarks on living fishes and crayfishes of Mammoth Cave.<Proc. Bost.

Soc. Nat. Hist., xyn, 222-225.
18756. On some of the habits of the blind crawfish Cambarus pellucidus and the

reproduction of lost parts.<Proc. Bost. Soc. Nat. Hist., xvm, 1875, 16-19.
See Packard, A. S-, and Putnam, F. W.
RICHARDSON, H.

1905. Monograph of the isopods of North America.^ Bull. U. S. Nat. Mus. 54,

pp. 727, figs. 740.
ROMANES, G. J.

1874a. Natural selection and dysteleology . <Nature, ix, 361,362.
18746. Disuse as a reducing cause in species. < Nature, x, 164.

1893. Mr. Herbert Spencer on " Natural selections. "<Con temporary Review

xxm, 499-518.
SACKEN, C. R. OSTEN.

1880. Catalogue of Diptera of North America. Smith. Misc. Coll., xvt, No. 270.
SAY, T.

A Description of the Insects of North America. Edited by J. L. Le Conte.
SAYCE, O. A.

1899. On a new species of Niphargus from Australia. <Proc. Roy. Soc. Victoria,

xn, pt. 2, 152-159.
1901. On three blind Victorian fresh-water Crustacea found in surface-water.

<Ann. Mag. Nat. Hist., ser. 7, vol. 8, 558-564.
SCUDDER, S. H.

1894. The North American ceuthophili.<Proc. Am. Acad. Sci., xxx, 17-113.
SEMPER, C.

1881. Animal life as affected by the natural conditions of existence. New

York, 1881.
SHALER, N. S.

1875. On the antiquity of the caverns and cavern life of the Ohio Valley.<Mem.

Bost. Soc. Nat. Hist., II, 355-363.
SHARP, P.

1899. Insecta, in Cambridge Natural History, v.
1901. Insecta, in Cambridge Natural History, vi.
SMITH, J. B.

1893. Catalogue of the Noctuidae found in Boreal America.<Bull. U. S. Nat. Mus.

44, 1893.
SMITH, S. I.

1873. The Crustacea of the fresh waters of the United States.<Rep. U. S. Fish

Commission 1872-73, 637-665.
1875, The crustaceans of the caves of Kentucky and Indiana. <Am. Journ. Sci.

and Arts, ix, 476-477.
SPENCER, H.

1893a. The inadequacy of natural selection. ^'Contemporary Review, LXIII, 153-

167, 439-457; Pop. Sci. Mon., XLIII, 21-29, 162-173.
18936. Professor Weismann's theories. <Contemporary Review, XLIII, 743-761,

Pop. Sci. Mon., XLIII, 473-490.
TELLKAMPF, TH. G.

1844a. Beschreibung einiger neuer in der Mammuth-Hohle in Kentucky aufgef-
undener Gattungen von Gliederthieren.< Archiv. fur Naturgeschichte,
x, Bd. 1, 318-322, Taf. vm.

18446. UeberdenblindenFischder Mammothhohle in Kentucky, mit Bemerkungen
iiber einige andere in dieser Hohle lebende Thiere.<Muller's Archiv fur
Anat. u. Physiol., Heft IV, 384-394, Taf. ix.
ULRICH, C. J.

1901. A contribution to the subterranean fauna of Texas. <Trans. Am. Mic. Soc.,
xxin, 83-100 (May, 1902).

114 FAUNA OF MAYFIELD'S CAVE.

VIRE, M. A.

1899. Essai sur la faune obscuricole de France. Paris.
DE VRIES, H.

1905. Species and varieties: their origin by mutation. Chicago, Open Court Co.
WEISMANN, AUGUST.

1904. The evolution theory. Translated by J. Arthur Thomson and Margaret R.

Thomson. London. Edward Arnold, vol. n, lecture xxv.
WICKHAM, H. F.

1896. On a collection of Indiana cave beetles. <Rep. Ind. Geol. Surv., xxi, 193-197.
WYMAN, J.

1853. On the eye and the organ of hearing in the blind fish (Amblyopsis spelseus
De Kay) of Mammoth Cave.^Am. Journ. Sci. and Arts, xvn, 258-261,
3 figs.; Muller's Archiv, 1853, 474-476.
YODER, A. C.

See Eigenmann, C. H., and Yoder, A. C.

MBL/WHOl I.IBKAKY

blH IfiHX