IRLF

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OF THE

UNIVERSITY OF CALIFORNIA.

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Received. ...TOT .28 1892 , /^ 81OLOGY

, LIBR

?. LfC(QO h Shelf No. .

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fc

THE

MAMMOTH CAYE

AND ITS

INHABITANTS,

OR DESCRIPTIONS OF THE

FISHES, INSECTS AND CRUSTACEANS

FOUND IN THE CAVE ;

WITH FIGURES OF THE VARIOUS SPECIES, AND AN ACCOUNT OF
ALLIED FORMS, COMPRISING NOTES UPON THEIR STRUC-
TURE, DEVELOPMENT AND HABITS, WITH
REMARKS UPON SUBTERRANEAN
LIFE IN GENERAL.

A. S. PACKARD, JR., AND F. W. PUTNAM,

EDITORS OF THE AMERICAN NATURALIST.

SALEM :
NATURALISTS' AGENCY.

1872.

OP THB

tf HIV EH SIT 7

00

PRINTED AT THE

SALEM PRESS.

Corner of Liberty and Derby Streets,

SALEM, MASS.
F. W. PUTNAM & CO.

PREFACE.

THE following pages were first published in the " Ameri-
can Naturalist" for December, 1871 and January, 1872, with
the exception of the Synopsis of the family including the
Blind fishes of the cave, which was first published in the
"Annual Keport of the Peabody Academy of Science for
1871."

In bringing the several articles together in the present
form but slight changes have been made, principally in the
form of a few additional notes.

It will undoubtedly be the good fortune of some visitors
to the cave to discover other kinds of animals than those
mentioned in the following pages, and to observe new facts
relating to the habits of the various species. For it must
be remembered that all the observations thus far recorded
have been made by but a very few of the thousands who
annually visit the Mammoth Cave, and that no thorough zo-
ological exploration of the cave has yet been undertaken.
Should any new facts be observed, or unknown species dis-
covered, the authors of this little work would be pleased to
be informed of them, and communications on all such
matters are solicited for publication in the pages of the

AMERICAN NATURALIST.

THE AUTHORS.

PEABODY ACADEMY OF SCIENCE, Salem, Mass., Feb. 1872.

TJHI7WSITY

THE MAMMOTH CAVE

AND ITS

INHABITANTS.

CHAPTER I.

THE FORMATION OF THE CAVE.*

BY F. W. PUTNAM.

AFTER the adjournment of the meeting of the American Associ-
ation for the Advancement of Science, held at Indianapolis, in
August last, a large number of the members availed themselves of
the generous invitation of the Louisville and Nashville Railroad
Company, to visit this world renowned cave, and examine its pe-
culiar formation and singular fauna.

The cave is in a hill of the subcarboniferous limestone forma-
tion in Edmondson County, a little to the west and south of the
centre of Kentucky. Green river, which rises to the eastward in
about the centre of the state, flows westward passing in close
proximity to the cave, and receiving its waters thence flows north-
westerly to the Ohio.

The limestone formation in which the cave exists, is a most in-
teresting and important geological formation, corresponding to
the mountain limestone of the European geologists, and of con-
siderable geological importance in the determination of the west-
ern coal fields.

We quote the following account of this formation from Major S.
S. Lyon's report in the fourth volume of the Kentucky Geological
Survey, pages 509-10.

"The sinks and basins at the head of Sinking creek exhibit
in a striking manner, the eroding effect of rains and frost — some
of the sinks, which are from forty to one hundred and ninety feet

*From the AMERICAN NATURALIST for December, 1871.

(5)

FORMATION OF THE CAVE.

deep, covering an area of from five acres to two thousand. The
rim of sandstone surrounding these depressions is, generally,
nearly level ; the outcropping rocks within are also nearly horizon-
tal. Near the centre there is an opening of from three to fifteen
feet in diameter ; into this opening the water which has fallen within
the margin of the basin has been drained since the day when the
rocks exposed within were raised above the drainage of the coun-
try, and thus, by the slow process of washing and weathering, the
rocks, which once filled these cavities, have been worn and carried
down into the subterranean drainage of the country. All this has
evidently come to pass in the most quiet and regular manner.
The size of the central opening is too small to admit extraordinary
floods ; nor is it possible, with the level margin around, to suppose
that these cavities were worn by eddies in a current that swept the
whole cavernous member of the subcarboniferous limestone of
western Kentucky ; but the opinion is probable that the upheaving
force which raised these beds to their present level, at the same
time ruptured and cracked the beds in certain lines ; that after-
wards the rains were swallowed into openings on these fractures,
producing, by denudation, the basins of the sinkhole country, and
further enlarging the original fractures by flowing through them,
and thus forming a vast system of caverns, which surrounds the
western coal field. The Mammoth Cave is, at present, the best
known, and, therefore, the most remarkable."

So much has been written on the cave and its wonders, that to
give a description of its interior would be superfluous in this
connection, even could we do so without unintentionally giving
too exaggerated statements which seems to be the natural result
of a day underground, at least so far as this cave is concerned,
for after reading any account of the cave, one is disappointed at
finding the reality so unlike the picture. As the Association party
was accompanied by one* who, while a most enthusiastic collector
and explorer, was also a calm recorder of statements made by
the geologists of the party, we cannot do better in conveying to
our readers the general geological character and structure of the
cave than to copy his account.

" As we expected to remain within the cave a long time, our
trusty guide, Frank, had provided himself with a well-filled can
of oil, to replenish our lamps, and with this strapped upon his
back he led the way into the thick darkness. We shall attempt
no description of the cave. Its darkness must be felt to be ap-
preciated, and no form of expression, understood by mortals who
have never descended to its cavernous depths, nor trod its gloomy

* W. P. FJSHBA.CK, Esq., of the Indianapolis Daily Journal.

FORMATION OF THE CAVE. /

corridors, can convey anything like an adequate idea of the place.
After spending fifteen hours within its chambers, it is absolutely
nauseating to read the descriptions which have been current in the
letters of newspaper correspondents for a quarter of a century,
and even the vigorous and picturesque language of Bayard Taylor
becomes tame and commonplace when it attempts to describe this
subterranean wonder of the world.

How and when the cave was made, were the leading questions
in the minds of the geologists. They do not believe that the cave
was the immediate result of some violent upheaval of the strata,
which left these vast crevices and chambers of which the cave is
composed ; neither do they share the popular belief that the rapid
and violent action of some subterranean stream of water has
worn these deep channels through the limestone ; on the contrary,
they find conclusive evidence that the same agencies are at work
and the same changes in progress to-day that have been slowly,
steadily and quietly, through vast periods of time, accomplishing
the marvellous wonders that now astonish the beholder. The cave
is wrought in the stratum known as the St. Louis limestone, which
in some places reaches a thickness or depth of four hundred feet.
This stone is dissolved whenever it is subjected to the influence of
running or dripping water impregnated with carbonic acid gas.
Water exposed to the air readily absorbs this gas, and surface water
percolating through small fissures of the limestone, dissolves it.
Another fact should be stated. When, during this process of so-
lution, the water becomes thoroughly impregnated with lime, it
loses its power to dissolve the stone. The following conditions,
then, were essential to the productions of the cave, assuming what
is not disputed by geologists, that the place where the cave now is,
was once nearly solid limestone. First, that there should be fis-
sures in the strata, allowing the ingress of the surface water. Sec-
ondly, there should be a place or places of exit for the water charged
with limestone in solution. Without the latter, the water would
become charged with lime, fill up the crevices, and the dissolving
process would cease. These conditions are all present to-day, and
have remained the same during the countless ages that have passed
away while the work has been in progress. There have doubtless
been times in the history of the cave, when, owing to a greater
flow of water, the work has progressed more rapidly than at pres-
ent, but that the results have been accomplished in the manner
stated, rather than by the process of attrition by rapid currents of
large volumes of water, seems to be the general opinion of scien-
tific men. This theory is strengthened by the fact that where the
cave attains its greatest heights, and reaches its lowest depths,
the dripping waters have never ceased their labors, and are busily
at work to-day. In the Mammoth Dome, for instance — rarely
seen by visitors, on account of the dangers and fatigue incident to
the journey — where the chasm attains a height and depth of more

8 FORMATION OF THE CAVE.

than two hundred and fifty feet, a cascade falls from a great height,
and keeps the entire surface of the rocks covered with dripping
water. This, falling into a deep pit below, finds an exit through
which it bears away a portion of the lime composing the rock.
After a walk of thirteen hours, our guide informed us that he would
conduct us to the- Mammoth Dome if we felt able to bear the
fatigue of the journey. Foot-sore and weary, we were not in a
favorable condition for so arduous an undertaking, but Mr. Thomas
Kite of Cincinnati, who had visited the locality thirty years ago,
urged us to go, and told us the sight of this Dome was worth all
the rest. Provided with magnesium and calcium lights, we crawled
and climbed our way to the brink of the pit, the bottom of which
was reached by a rickety ladder, slippery and dripping with water.
A portion of the party descended, and when all were ready the
lights were ignited, and the immense dome was revealed to us in
all its majestic beauty. Upon our return, three hearty cheers were
given to the good friend at whose earnest solicitation we under-
took this part of our journey.

We are indebted to Professor Alexander Winchell. of the Unir
versity of Michigan, for the following abstract of his views con-
cerning the formation of the cave.

' The country of the Mammoth Cave was probably dry land at
the close of the coal period, and has remained such, with certain
exceptions, through the Mesozoic and Csenozoic ages, and to the
present. In Mesozoic times, fissures existed in the formation,
and surface waters found their way through them, dissolving the
limestone and continually enlarging the spaces. A cave of con-
siderable dimensions probably existed during the prevalence of the
continental glaciers over the northern hemisphere. On the dis-
solution of the glaciers, the flood of water which swept over the
entire country, transporting the materials which constituted the
modified drift, swept through the passages of the cave, enlarging
them, and leaving deposited in the cave, some of the same quart-
zose pebbles which characterize the surface deposits from Lake Su-
perior to the Gulf of Mexico. Since the subsidence of the waters
of the Champlain epoch, the cave has probably undergone compar-
atively few changes. The well one hundred and ninety-eight feet
deep, at the further end of the cave, shows where a considerable
volume of the excavatory waters found exit. The Mammoth Dome
indicates probably, both a place of exit and a place of entrance
from above. So of the vertical passages in various other portions
of the cave.'

We believe that the views of Professor Winchell are in harmony
with those of the other eminent geologists of the party, and when
it is considered that the geologists of this excursion stand in the
front rank of the most eminent scientific men of the world, their
views upon this interesting subject are well Avorthy of attention.
Before dismissing this branch of the subject, we will take occasion

FORMATION OF THE CAVE.

to correct a popular error concerning the formation of the beauti-
ful structures that adorn the ceilings of some portions of the cave.
In the dryer localities, where the floors are dusty and everything
indicates the prolonged absence of moisture, the ceiling is covered
with a white efflorescence that displays itself in all manner of
beautiful shapes. It requires no stretch of the imagination to dis-
cover among these, the perfect forms of many flowers. The lily
form prevails, and the ceilings of many of the chambers are cov-
ered with this beautiful stucco work, surpassing in delicacy and
purity the most beautiful workmanship of man. These are not
produced, as many suppose, by the dripping of water, and the
gradual deposit of sulphate of lime upon the outer portions. The
stalactite is formed in this manner, but these are neither stalacti-
form, nor are they produced in a similar way. Dripping water
is the agency that forms the stalactite, while the efflorescence
in the dryer portions of the cave cannot take place where there is
much moisture. The growth of these beautiful forms is from
within, and the outer extremities are produced first. They are
the result of a sweating process in the limestone that forces the
delicate filaments of which they are composed through the pores
upon the surface of the rock, their beautiful curved forms result-
ing from unequal pressure at the base, or friction in the apertures
through which they are forced. Mr. L. S. Burbank, of Lowell,
Mass., has kindly furnished us with the following abstract of his
opinions upon this interesting subject.

' The rosettes, wreaths, and other curved fibrous forms of gyp-
sum, in the Mammoth Cave, occur only in particular strata of the
limestone which do not appear in the first part of the long route.

Their formation may be explained in this way : that portion of
the rock where they are found consists of carbonate of lime, with
some impurities, and contained originally the sulphide of iron, or
iron pyrites, disseminated in small grains or crystals, and also in
rounded nodules or concretions, sometimes of considerable size.

By exposure to air and moisture, oxygen unites with both the
sulphur and the iron, producing sulphuric acid and oxide of iron,
which combined, form a sulphate of iron. Then a double de-
composition takes place ; the sulphuric acid unites with the lime
to form the gypsum ; the carbonic acid of the limestone combines
with the oxide of iron, forming a carbonate of iron, and this, on
further exposure, parts with the carbonic acid, and leaves the
brown coating of oxide, which is seen in many places on the sur-
face of the rock.

The gypsum is thus constantly forming in the rock, and, being
soluble, is carried by the water to the exposed surface where it
crystallizes.

The crystals appear to grow out from the rock by additions from
beneath, which continue to push the ends first formed, and if these
do not become attached to other parts of the rock, straight needle-

10 ANIMAL LIFE IN THE CAVE.

like fibres are often produced. Very commonly, however, the
crystals begin to form when a small nodule of the iron ore is ex-
posed at the surface ; the parts first formed become attached to
the surface around the edges, and as the chemical action proceeds
towards the centre of the nodules successive leaf-like layers are
thrown out, and the rosette form is the result. Along lines of
fracture in the surface of the rock, the crystals are curved in op-
posite directions.

The wreaths and other figures formed by the chains of the ro-
settes, may be caused by the chemical action described taking place
around the edges of large masses or concretions of the iron ore.

These crystalline forms occur only in the dryer parts of the
cave. Where there is more moisture, as in the ' Snow-ball room,'
the gypsum merely forms white, rounded concretions, originating
from nodules of the iron ore on the roof and sides of the cave.'"

With these general remarks on the cave we give a brief account
of its interesting fauna,* comprising representatives of the Fishes,
Insects, and Crustaceans. No Mollusks nor Radiates have as yet
been discovered, but the lower forms of life have been detected
by Tellkampf, who collected several species belonging to the gen-
era Monas, Chilomonas, and (?) Chilodon.

*In the following pages it will be noticed that the authors have expressed widely
different views as to the origin of the peculiar forms of subterranean animals.

CHAPTER II.

THE CRUSTACEANS AND INSECTS.*

BY A. 8. PACKARD, JR.

Fig. 122.

REPRESENTATIVES of all the grand divisions of the Insects and
Crustaceans have been found in this cave, and if no worms have
yet been detected, one or more species would undoubtedly reward
a thorough search.

We will enumerate what have been found, beginning with the
higher forms. No Hymenoptera (bees, wasps, and ants) or Lepi-
doptera (moths) are yet recorded as being peculiar to caves. The
Diptera (flies) are represented by two species, one of Anthomyia
(Fig. 122) or a closely
allied genus, and the sec-
ond belonging to the sin-
gular and interesting ge-
nus Phora (Fig. 123).
The species of Antho-
myia usually frequent
flowers ; the larvae live in
decaying vegetable mat-
ter, or, like the onion
fly, attack healthy roots ;
while the maggots of
Phora live in decaying
substances. It would be

Anthomyia.

presumptuous in the writ-
er to attempt to describe these forms without collections of spe-
cies from the neighborhood of the cave, for though like all the
rest of the insects they were found three or four miles from the
mouth, yet they may be found to occur outside of its limits, as
the eyes and the colors of the body are as bright as in other
species.

Among the beetles, two species were found by Mr. Cooke. The

*From the AMERICAN NATURALIST for December, 1871.

(11)

12

INSECTS OF THE CAVE.

Fig. 124.

Anoptlialmus Tellkampjli of Erichson, a Carabid (Fig. 124), and
Adelops hirtus Tellkampf (Fig. 125) allied to Catops, one of the
Silphidse or burying beetle family. The Anoptlialmus is of a pale
reddish horn color, and is totally blind ; * in the Adelops, which
is grayish brown, there are two pale spots, which may be rudi-
mentary eyes, as Tellkampf and Erichson suggest. No Hemip-
tera (bugs) have yet been found either in the caves of this coun-
try or Europe. Two wingless grasshoppers (generally called
crickets) like the common species found under stones (CeutJio-
philus maculatus Harris) , have been found in our caves ; one is
the Hadenoecus subterraneus (Fig. 126 nat. size) described by Mr.
Scudder, and very abundant in Mammoth Cave. The other spe-
cies is C. stygia Scudder, from Hickman's
cave, near Hickman's land- Fig. 123.
ing, upon the Kentucky river.
It is closely allied to the
Mammoth Cave species. Ac-
cording to Mr. Scudder, the
specimens of C. stygia were
found by Mr. A. Hyatt " in
the remotest corner of Hickman's Cave,
in a sort of a hollow in the rock, not par-
ticularly moist, but having only
a sort of cave dampness. They
were found a few hundred feet
from the sunlight, living exclu-
sively upon the walls." Even
the remotest part of that cave is
not so gloomy but that some
sunlight penetrates it.
The other species is found both in Mammoth Cave, and in the
adjoining White's Cave. It is found throughout the cave, and
most commonly (to quote Mr. Scudder) "about 'Martha's Vine-
yard' and in the neighborhood of 'Richardson's Spring' where
they were discovered jumping about with the greatest alacrity
upon the walls, where only they are found, and even when dis-

*In Erhardt's cave, Montgomery Co., Virginia, Prof. Cope found "four or five spec-
imens of a new Anoptlialmus, the A. pusio of Horn, at a distance of not more than three
hundred feet from its mouth. The species is small, and all were found together under
a stone. Their movements were slow, in considerable contrast to the activity of ordi-
nary Carabidze." Proc. Amer. Phil. Soc. 1869. p. 178.

Phora.

Fig. 125.

Anophthalmus Tellkampfii.

INSECTS OF THE CAVE. 13

turbed, clinging to the ceiling, upon which they walked easily;
they would leap away from approaching footsteps, but stop at a
cessation of the noise, turning about and swaying their long an-
tennae in a most ludicrous manner, in the direction whence the
disturbance had proceeded ; the least noise would increase their
tremulousness, while they were unconcerned at distant motions,
unaccompanied by sound, even though producing a sensible cur-
rent of air ; neither did the light of the lamp appear to disturb
them ; their eyes, and those of the succeeding species (J2. stygia)
are perfectly formed throughout, and they could apparently see
with ease, for they jump away from the slowly approaching hand,
so as to necessitate rapidity of motion in seizing them."

Mr. Henry Edwards has discovered a wingless grasshopper in a
limestone cave at Collingwood, Massacre Bay, Middle Island, New

Fig. 126.

Jfadenoecus subterraneus.

Zealand. Says Mr. Scudder, who has described the species in the
"Proceedings of the Boston Society of Natural History^' (Vol.
xii, 1869, p. 408) under the name Hadencecus Edwardsii^ "the
cave is close to the sea shore, and near a very large coal deposit,
which occasionally crops out in the interior. The Hadenceci were
rather numerous, but very difficult to catch, disappearing in the
crevices of the rocks on the approach of lights. They appeared
to be most abundant near the streams of water which percolated
through the rocks." The wingless grasshopper of the European
caves is the Hadenoecus palpatus Scudder, first described by Sulzer
under the name Locusta palpata.

The Thysanurous Neuroptera are represented by a species of
Machilis, allied to our common Macliilis variabilis Say, common in
Kentucky and the middle and southern states. So far as Tell-

INSECTS OF THE CAVE.

Fig. 127.

kampfs figure indicates, it is the same species apparently, as I
have received numerous specimens of this widely distributed form
from Knoxville, Tennessee, collected by Dr. Josiah Curtis.

It was regarded as a crustacean by Tellkampf, and described
under the name of Triura cavernicola.* He mistook the labial
and maxillary palpi for feet and regarded
the nine pairs of abdominal spines as feet.
The allied species, M. variabilis Say, is
figured in vol. v. pi. 1, fig. 8, 9 (see also
p. 94 of vol. v of the NATURALIST).

An interesting species of Campodeaf of
which the accompanying cut (Fig. 127) is
a tolerable likeness, though designed to il-
lustrate another species (C. stapliylinus
Westw.) was discovered by Mr. Cooke.
Both the European and our common spe-
cies live under stones in damp places, and
the occurrence of this form in the water is
quite remarkable. The other species are
blind, and I could detect no eyes in the
Mammoth Cave specimen.

A small spider was captured b}r Mr.
Cooke, but afterwards lost ; it was brown
in color, and possibly distinct from the
Anthrobia monmouthia Tellkf. (Fig. 128) which is an eyeless form,
white and very small, being but half a line in length. The family

*-Professor Agassiz in his brief notice of the Mammoth Cave animals, does not criti-
cise Tellkampf s reference of this animal to the Crustacea; and so eminent an authority
upon the articulates as Schiodte remarks that while " Dr. Tellkampf's account affords
us no means of forming any conclusion as to its proximate relations," that, hoAvever,
it " appears to belong to the order of Amphipoda, and to have a most remarkable
structure." TellkampPs figure of Machilis is entirely wrong in representing the labial
and maxillary palpi as ending in claws, thus giving the creature a crustacean aspect ;
and ndeed he describes them as true feet !

t Campod^a Cookei n. sp. Closely allied to C. Americana, but it is much larger; the an-
tennae are 24-jointed instead of 20-jointed as in C, Americana, and reach to the basal
abdominal segment, while in C. Americana they reach only to the second thoracic; the
terminal joints are much longer than in that species, the penultimate joint being one-
third longer. Last three abdominal segments unequal (equal in C. Americana) the penul-
timate very short, not half as long as the terminal, which is longer and slenderer than in
C. Americana, while the three are much narrower in proportion to the rest of the body
than in the other species. Hind femora longer than in C. Americana. Entirely white
and pilose. Length .25 inch, the largest C. Americana being .15 to .20 inch. (Anal sty-
lets broken off.) Several specimens were seen by Mr. C. Cooke, but only one was cap-
tured in a pool of water, two or three inches deep, in company with the C- cidotea.

Campodea.

INSECTS OF THE CAVE.

15

of Harvest men is represented by a small white form, described by
Tellkampf under the name of Plialangodes armata (Fig. 129) but
now called Acanthocheir armata Lucas. The body alone is but half
a line long, the legs measuring two lines. It should be borne in
mind that many of the spiders, as well as the Thysanura, live in
holes and dark places, so that we would naturally find them in
caves. So, also, with the Myriopods, of which a most remarkable

form* (Figs. 130, and 130 a
front of head) was found by
Mr. Cooke, three or four miles
from the mouth of the cave. It
is the only truly hairy species
known, an approach to it being
found in Pseudotremia Vudii
Cope. It is blind, the other spe-
cies of this group which Profes-
sor Cope found living in caves
having eyes. The long hairs ar-
AnthroUa monmouthia. ranged along the back, seem to

suggest that they are tactile organs, and of more use to the Thous-
and legs in making its way about the nooks and crannies of a per-
petually dark cave than eyes would be. It was found by Mr.
Cooke under a stone.

Prof. Cope has contributed to the " Proceedings of the American
Philosophical Society" (1869, p. 171) an interesting account of the

* Spirostrephon (Pseudotremia') Copei n. sp. Head with rather short, dense hairs; no
eyes, and no ocular depression behind the antennae, the surface of the epicranium being
well rounded to the antennal sockets; behind the insertion of the antennas the sides of
the head are much more swollen than in S. lactarius. Antennae slender, with short
thick hairs ; relative length of joints, the 6th being longest ; 6th, 4th, 5th, 3d, 8th, 7th, 1st,
the 7th joint being much thicker than the 8th. Twenty-eight segments besides the head ;
they are entirely smooth, striated neither longitudinally nor transversely; a few of the
anterior segments rapidly decrease in diameter towards the head. The segments are
but slightly convex, and on each side is a shoulder, bearing three tubercles in a trans-
verse row, each giving rise to a long stiff hair one-half to two-thirds as long as the seg-
ment is thick; these hairs stand up thickly all over the back, and may serve at once to
distinguish the species. No pores. Feet long and slender, nearly as long as the an-
tennae, being very slender towards the claws. Entirely white. Length of body .35
inch ; thickness .04 inch.

It is nearly allied to Pseudotremia Vudii of Cope. It will be noticed that Professor
Cope characterizes the genus Spirostrephon as having " no pores "; though we find it
difficult to reconcile this statement with that of Wood who describes S. lactarius as
having " lateral pores." Cope separates Pseudotremia from Spirostrephon for the rea-
son that the segments have "two pores on each side the median line." The present
species has no pores, but seems in other characters to be a true Spirostrephon, and we
are thus led to doubt whether Pseudotremia is a well founded genus.

16

INSECTS OF THE CAVE.

cave mammals, articulates and shells -of the middle states. He
says that " myriopods are the only articulates which can be
readily found in the remote regions of the caves [of West Vir-
ginia] and they are not very common in a living state." The
Pseudotremia cavemarum which he describes, " inhabits the deep-
Fig. 129.

Acanthocheir armatct.

est recesses of the numerous caves which abound in Southern Vir-
ginia, as far as human steps can penetrate. I have not seen it
near their mouths, though its eyes are not undeveloped, or smaller
than those of many living in the forest. Judging from its remains,
which one finds under stones, it is an abundant species, though

Fig. 130. Fig. 130 a.

Spirostrephon Copei.

rarely seen by the dim light of a candle even after considerable
search. Five specimens only were procured from about a dozen
caves." Thev second species, P. Vudii Cope, was found in Mont-
gomery Co. and he thinks it was not found in a cave. Professor
Hyatt informs me that he saw near the "Bottomless Pit" in Mam-

CRUSTACEANS OF THE CAVE.

17

moth Cave, a brownish centipede-like myriopod, over an inch in

length, which moved off in a rapid zigzag motion. Unfortunately

he did not capture it.

Next to the blind fish, the blind crawfish attracts the attention

of visitors to the cave. This is the Cambarus pellucidus (Fig.
F.; . Tii . 131, from Hagen's

monograph of the
North American
Astacidae) first de-
scribed by Dr. Tell-
k a m p f . He re-
marks that "the
eyes are rudiment-
ary in the adults,
but are larger in
the young." We
might add that this
is an evidence that
the embryo devel-
ops like those of
the other species ;
and that the inher-
itance of the blind
condition is proba-
bly due to causes
first acting on the a-
dults and transmit-
ted to their young,
until the production
of offspring that be-
come blind becomes
a habit. This is
a partial proof at
least that the char-
acters separating
the genera and spe-

Cambanis peUucidus. c[es Qf animals are

those inherited from adults, modified by their physical surround-
ings an 1 adaptations to changing conditions of life, inducing cer-
tain alterations in parts which have been transmitted with more or

MAMMOTH CAVE. 2

18

CRUSTACEANS OF THE CAVE.

Fig. 132.

Csccidotea styyia (side view).

Fig. 133.

less rapidity, and become finally fixed and habitual. Prof. Hagen
has seen a female of Cambarus Bartonii from Mammoth Cave,
" with the eyes well developed," and a specimen was also found
by Mr. Cooke. Prof. Hagen remarks that " C. pellucidus is the
most aberrant species of the genus. The eyes are atrophied,
smaller at the base, conical, instead of cylindrical and elongated,
as in the other species. The cornea exists, but is small, circular,
and not faceted ; the optic fibres and the dark-colored pigments
surrounding them in all other spe-
cies are not developed." It seems
difficult for one to imagine that our
blind craw fish was created sud-
denly, without the intervention of
secondary laws, for there are the
eyes more perfect in the young than
the adult, thus pointing back to an-
cestors unlike the species now ex-
isting. We can now understand,
why embryologists are anxiously
studying the embryology of animals
to see what organs or characteristics
are inherited, and what originate de
novo, thus building up genealogies,
and forming almost a new depart-
ment of science : comparative em-
bryology in its truest and widest
sense.

Of all the animals found in caves,
either in this country or Europe,
perhaps the most strange and unex-
pected is the little creature of which
we now speak. It is an Isopod crus-
tacean, of which the pill bugs or sow bugs arc examples. A true
species of pill bug ( Titanethes albus Schiodte) inhabits the caves of
Carniolia, and it is easy to believe that one of the numerous species
of this group may have become isolated in these caves and modi-
fied into its present form. So also with the blind Niphargus sty-
gius of Europe, allied to the fresh water Gammarus so abundant
in pools of fresh water. We can also imagine how a species of
Asellus, a fresh water Isopod, could represent the Idoteidse in our

CiEcidotea stygia (dorsal view).

CRUSTACEANS OF THE CAVE. 19

caves, and one may yet be found ; but how- the present form be-
came a cave dweller is difficult of explanation, as its nearest allies
are certain species of Idotea which are all marine, with the excep-
tion of two species : I. entomon, living in the sea and also in the
depths of the Swedish lakes, as discovered by Loven, the distin-
guished Swedish naturalist, while a species representing this has
been detected by Dr. Stimpson at the bottom of Lake Michigan.
Our cave dweller is nearly allied to Idotea, but differs in being
blind, and in other particulars, and may be called Ccecidotea sty-
giob.* (Fig. 132 side view, enlarged ; Fig. 133 dorsal view ; 6, in-
ner antenna ; c, 1st leg.) It was found creeping over the fine
sandy bottom, in company with the Campodea, in a shallow pool
of water four or five miles from the mouth of the cave.

This closes our list of known articulates from this and other
caves in this country, the result of slight explorations by a few in-
dividuals. The number will be doubtless increased by future re-
search. It is to be hoped that our western naturalists will thor-
oughly explore all the sinks and holes in the cave country of the
western and middle states. The subject is one of the highest in-
terest in a zoological point of view, and from the light it throws on
the doctrine of evolution. Professor Schiodte, the eminent Danish
zoologist, has given us the most extended account of the cave
fauna of Europe, which has been translated from the Danish into
the Transactions of tke Entomological Society of London (new
series vol. 1, 1851).

He examined four caves ; namely, that of Adelsberg, the Mag-
dalena and Luege caves, all in the neighborhood of Adelsberg,

* Generic characters. Head large, much thicker than the body, and as long as broad;
subcylindrical, rounded in front. No eyes. First antennae slender, 8-jointed (2d anten-
noe broken off). Abdominal segments consolidated into one piece. Differs chiefly from
Idotea, to which it is otherwise closely allied, by the 8-jointed (instead of 4-jointed) 1st
(inner) antenna?, the very large head, and by the absence of any traces of the three ba-
sal segments of the abdomen usually present in Idotea.

Specific chai-acters. Body smooth, pure white : tegument thin, the viscera appearing
through. Head as wide as succeeding segment, and a little more than twice as long.
Inner antennne minute, slender, the four basal joints of nearly equal length, though the
fourth is a little smaller than the basal three, remaining four joints much smaller than
others, being one-half as thick and two-thirds as long as either ot the four basal joints ;
ends of last four joints a little swollen, giving rise to two or three hairs; terminal joint
ending in a more distinct knob, and bearing five hairs. Segment of thorax very dis-
. tinct, sutures deeply incised; edges of segments pilose; abdomen flat above, rounded
behind, with a very slight median projection ; the entire pair of gills do not reach to
the end of the abdomen, and the 'inner edges diverge posteriorly. Legs long and slen-
der, 1st pair shorter, but no smaller than the second. Length .25 inch.

20 GENERAL REMARKS ON CAVE FAUNA.

and the Corneale cave at Trieste. The only plant found was a
sort of fungus, Byssus fulvus Linn. The only vertebrate is the
singular salamander, Hypochthon (Proteus) anguinus, found in the
Magdalina river. No shells were found. Regarding the articu-
lates he writes :

"On searching along the wralls within the entrance of the caves,
among the rubbish and the vegetable debris along the sides of the
river, we meet wTith a considerable number of Insecta, Myriopoda,
Arachnida and Crustacea, of various families which shun daylight ;
being such species only as inhabit promiscuously other places,
provided they are moist and feebly illumined. We find species of
Pterostichus, Pristonychus, Amara, Quedius, Homalota, Omalium,
Hister, Trichopteryx, Cryptophagus, Atomaria, Ptinns, Ceraphron,
Belyta, a grasshopper of the Locust family, probably the Raphido-
pJwra cavicola Fischer, as it was only seen in the larva state, Trich-
optera, Sciara, Psychoda, Phora, Heteromyza, Sapromyza, Tomoce-
rus, Linyphia, Gamasus, Cryptops, Julus, and Asellus. In pro-
portion as we recede from the entrance the number of species as
well as individuals greatly decreases, and at the distance which
entirely excludes the light, only single individuals are found. In
the deepest recesses these species are entirely wanting, except
some few which have been transported by the current ; only a few
Diptera are found ; namely, a species of Phora, very near P. (ma-
culata Meig., Heteromyza flampes Zett., and Sapromyza cJirysoph-
tlialma Zett., extending also very far into the caves, even to the
remotest accessible places in Adelsberg cave, more than half an
hour's walk from its entrance. Dead moths%are occasionally found
far in the caves, being left there by 'the bats; and likewise acci-
dental specimens of the parasites of the latter. Of the five ear-
lier known animals which inhabit these caves, I found Pnstonyclia
elegans Dej. rather frequently, and Homalota spelcea Er. in consid-
erable numbers. Besides these are Anopihalmus Sclimidtii, which
is very rare, and the wood louse, Titanethes alba. The new forms
he found were a beetle (Bathyscia byssina) allied to our Adelops ;*
Stagdbius troglodytes, an aberrant genus of Silphids ; a Podurid,
Anurophorus Stillicidii; and the two blind arachnidans, one a spi-
der allied to Dysdera, the Stalita tcenaria, and a false-spider, Blo-
thrus spelceus. Among the Crustacea he found Nipliargus stygius^

*Lud\vig Muller enumerates four other species of Adelops from theee caves, and
three species from France, and Maclicerites spelcetis, in Verhandl. Zool. Bot. Vereins,
Wien, 1855, p. 505. See also Heller's Beitrage zur Bsterreieh. Grotteu-Fauna. (Myrio-
poda and Crustacea.) Vienna, 1858. He describes a myriopod with rudimentary eyes
(Trachysphceria Schmidtii) allied to Glomeris, and another blind species (Brachydesmus
subterraneus) allied to Polydesmus ; also a new Titlianethes ( T. graniyer), and notices
Monolistra cceca Gerst. Wankel (18(51) also found a new rhalangid (Leiobunum troglo-
dytes) witli distinct eyes and four species of mites in the caves of Eastern Austria. The
mites are Scyphius spelceus, Linopodes subterraneus, Gamasus loricatus and G. niveus.
Mill an additional species of Trachysphama (T. HyrtHi). See also Ehrenberg's list 01
cave insects (Monabsberichte der Akad. Berlin. 18(51.)

t Several species of Niphargus occur in the wells and hot springs in Europe. Accord-

GENERAL REMARKS ON CAVE FAUNA. 21

allied to G-ammarus, which lives in small pools of water and is
white and blind ; and the cave pill bug, Titanethes albus (Koch.)."

. In conclusion Schiodte remarks that : —

"We may with propriety apply the collective term Subter-
ranean Fauna to those animals which exclusively inhabit caves,
and are expressly constructed for such habitations. Still there
is nothing in this name which would indicate that these animals
have any claim to be considered as a separate group, beyond the
mere peculiarity of their common place of abode. While a few
of them possess such an extraordinary structure as to stand in no
comparison with those animals which inhabit the light, there are
others, forming only more characteristic links in the groups of
animals more or less shy of light, of which many are found common
in the localities of the caves ; and some belong to genera having a
wide local, as well as geographical, extension. We are accordingly
prevented from considering the entire phenomenon in any other
light than something purely local, and the similarity which is ex-
hibited in a few forms (Anophthalmus, Adelops, Bathyscia) be-
tween the Mammoth Cave and the caves in Carniola, otherwise
than as a very plain expression of that analogy, which subsists
generally between the fauna of Europe and of North America.
Besides, it is clear to me that the fauna of the caves of Carniola
consists of two divisions, of which the essential character is refer-
able on the one hand to the dark locality, and on the other to the
additional confinement to staiactitic formations ; as yet we are not

ing to Bate and Westwood (British Sessile eyed Crustacea) •' the British examples have
been obtained from artificially excavated wells connected with houses for domestic
purposes. In some instances the wells have been old, in others but recently dug. In
their geological condition the habitats have been equally various. At Corsham the well
exists in the Oolite formation, at Ring-wood in chalk-flint gravel, at Mannamead in
tlate. At Corsham and Mannamead they are found on a hill, at Ringwood they lie low.
The appearance of some of these animals in a well soon after its being excavated,
raises a question of considerable interest. Thus they were found at upper Claft'ord, near
Andover and at Mannamead, near Plymouth, but not a trace of them was to be found in
the Hurroimding streams; in fact they perish in the light. It is impossible to regard
them as an extreme variety, or modification of our only fresh water Aniphipod, Gam-
marusfluviaiilus, since various parts not only differ in form, but some are altered in char-
acter; for example, the extraordinary elongation and slenderness of one of the branches
of each of the last pair of caudal appendages seem to be a special structure, having
for its object the antenna-like use of a delicate apparatus at the extremity of the body.
.... Although we can find no freshwater ally to this genus in the rivers and
streams of Europe, yet Bruzelius has taken in the deep sea, near Bohusia, a form which
he has described under the name Eriopis elongata, approximating so nearly to it that it
appears to be scarcely genetically distinct.

anus Koch, the embryology of which has been studied by V. St. George) N.fontanu»
Bate, N. Kochianun Bate. Another generic form is Crangonyx founded by Bate, which
also belongs to the subterranean fauna. " A single species as yet is all that has been
found in England; but we have little doubt but that Gammarus Ermamii of Mime Ed-
wards which was found by M. Ermann in the warm springs of Kamtschatka belongs
also to this genus. It, is curious that we should have to record that while the animals
of this genus, as in the p:-e:-e .ling(.Niphargtis) inhabit the deep artificial wells, without
being known to exist in our rivers and streams, its nearest allied form is to be found
in a marine genus, Gammarella."

22 GENERAL REMARKS ON CAVE FAUNA.

able vigorously to discriminate between the two. We shall ac-
cordingly look upon the subterranean fauna, or more properly
faunas, as small ramifications which have penetrated into the earth
from the geographically-limited faunas of the adjacent regions ;
and which, as they extended themselves into darkness, have been
accommodated to surrounding circumstances. Animals not far re-
mote from the ordinary forms, prepare the transition from light to
darkness. Next follow those that are constructed for twilight ;
and last of all those destined for total darkness, and whose struc-
ture is quite peculiar. Among these some are adapted for special
localities, those which inhabit dry localities or detached little
reservoirs being totally blind, while others, destined for running
streams, have eyes of imperfect construction, so as to receive the
impression of rays of light, but no proper image of illuminated
objects. We may therefore with tolerable precision arrange the
inhabitants of caverns under the following heads : —

Shade animals. — Extensive genera and species inhabiting cav-
erns near their entrance, and, generally, all cool, shady and moist
localities. Of these, those that fly occasionally enter far into the
caverns (Diptera).

Twilight animals. — They belong to widely spread genera, but
are peculiar to the caves, and distinguished by their small eyes.
They are principally found near the entrances to the caves, but
proceed deeper into the darkness than the shade-animals, and
although wingless, they penetrate often the whole extent of the
dark space. — (Pristonychus elegans, Homalota spelwa.)

Cave animals. — The^y form, at least in part, peculiar genera, are
wingless and colorless, as far as the consistency of their integu-
ments will admit, and exist exclusively in total darkness. The
terrestrial division is blind ; the aquatic has a perception of light.
To this group belong all the animals in the Mammoth Cave, and
among those of the caves of Carniola, Auoplithalmus, Bathyscia,
perhaps likewise Anurophorus and Hypochthon, which, however,
may belong to the following group.

Stalactite cave animals. — Insects, Arachnidans and Crustaceans
appertaining to peculiar genera, wingless, blind, brightly colored
according to the nature of their integuments, either light brown,
yellowish white, or snow white, perhaps according to the pre-
ponderance of the chitine ; living in total darkness, peculiar to
stalactite caves, in part occupying the columns and constructed
accordingly, either for ascent or hovering over them. Here belong
most of the animals treated of in this memoir — Stagobius, Blo-
thrus, Stalita, Niphargus, and Titanethes." *

A pertinent question arises as to the time of the formation of
these caves and when Lhey became inhabitable. As previously stat-

*In a note appended he adds to the list "a new cave crustacean, Pnlomnn anopthal-
mus Kollar, said to serve as food for Hypochthon [the Salamander], of which last geuus
he discriminates six species."

ORIGIN OF CAVE LIFE. 23

ed, the caves of the western and middle States are in lower Car-
boniferous limestone rocks, though the Port Kenned}^ cave explored
by Wheatley and Copef is in the Potsdam limestone. They could
not have been formed under water, but when the land was drained
by large rivers. This could not have occurred previous to the Tri-
assic period. Prof. Dana in his "Manual of Geology" shows that
the Triassic continent spread westward from the Atlantic coast "to
Kansas, and southward to Alabama ; for through this great area
there are no rocks more recent than the Palaeozoic." "Through the
Mesozoic period [comprising the Triassic, Jurassic, and Creta-
ceous periods] North America was in general dry land, and on the
east it stood a large part of the time above its present level."
Though at the close of these periods there was a general extinc-
tion of life, yet this was not probably a sudden (one of months
and even years), but rather a secular extinction, and there may be
plants and animals now living on dry land, which are the lineal
descendants of mesozoic and more remotely of Carboniferous forms
of life. So our cave animals may possibly be the survivors of Mes-
ozoic forms of life, just as we find 'now living at great depths in
the sea remnants of Cretaceous life. But from the recent explora-
tions in the caves of Europe and this country, especially the Port
Kennedy cave, with its remarkable assemblage of vertebrates and
.insects, we are led to believe from the array of facts presented by
Prof. Cope that our true subterranean fauna probably does not
date farther back than the beginning of the Quaternary, or Post
pliocene, period. We quote his "general observations" in his
article on the Port Kennedy fauna.

"The origin of the caves which so abound in the limestones of
the Alleghany and Mississippi valley regions, is a subject of much
interest. Their galleries measure many thousands of miles, and
their number is legion. The writer has examined twenty-five, in
more or less detail, in Virginia and Tennessee, and can add his
testimony to the belief that they have been formed by currents of
running water. They generally extend in a direction parallel to
the strike of the strata, and have their greatest diameter in the
direction of the dip. Their depth is determined in some measure
by the softness of the stratum, whose removal has given them
existence, but in thinly stratified or soft material, the roofs or large

t A notice of the animals found in this cave will be found in the Proceedings of the
American Philosophical Society, April, 1871. The insects there enumerated would
probably not come under the head of cave insects .

24 ORIGIN OF CAVE LIFE.

masses of rocks fall in, which interrupt the passage below. Caves,
however, exist when the strata are horizontal. Their course is
changed by joints or faults, into which the excavating waters have
found their way.

That these caves were formed prior to the postpliocene fauna is
evident from the fact that they contain its remains. That they
Were not in existence prior to the drift is probable, from the fact
that they contain no remains of life of any earlier period so far as
known, though in only two cases, in Virginia and Pennsylvania,
have they been examined to the bottom. No agency is at hand to
account for their excavation, comparable in potency and efficiency
to the floods supposed to have marked the close of the glacial
period, and which Prof. Dana ascribes to the Cham plain epoch.
An extraordinary number of rapidly flowing waters must have
operated over a great part of the Southern States, some of them
at an elevation of fifteen hundred feet and over (perhaps two thou-
sand) above the present level of the sea. A cave in the Gap
Mountain, on the Kanawha river, which I explored for three miles,
has at least that elevation.

That a territory experiencing such conditions was suitable for
the occupation of such a fauna as the deposits contained in these
caves reveal, is not probable.. The material in which the bones
occur in the south is an impure limestone, being mixed with and
colored by the red soil which covers the surface of the ground. It
is rather soft but hardens on exposure to the air.

The question then remains so far unanswered as to whether a
submergence occurred subsequent to the development of the post-
pliocene mammalian fauna. That some important change took
place is rendered probable by the fact, that nearly all the neotropi-
cal types of the animals have been banished from our territory,
and the greater part of the species of all types have become ex-
tinct. Two facts have come under my observation which indicate
a subsequent submergence. A series of caves or portions of a
single cave once existing on the southeast side of a range of low
hills among the Alleghany mountains in Wythe Co., Virginia, was
found to have been removed by denudation, fragments of the bot-
tom deposit only remaining in fissures and concavities,, separated
by various intervals from each other. These fragments yielded the
remains of twenty species of postpliocene mammalia.* This de-
nudation can be ascribed to local causes, following a subsidence
of uncertain extent. In a cave examined in Tennessee the Qssife-
rous deposit was in part attached to the roof of the chamber.
Identical fossils were taken from the floor. This might, however,
be accounted for on local grounds. The islands of the eastern
part of the West Indies appear to have been separated by submer-
gence of larger areas, at the close of the period during which they

*See Proceed. Amer. Phil. Soc. 1869, 171.

ORIGIN OF CAVE LIFE. 25

were inhabited by postpliocene mammalia and shells. The caves
of Anguilla include remains of twelve vertebrates,* of which seven
are mammalia of extinct species, and several of them are of large
size. These are associated with two recent species of molluscs
Turbo pica, and a Tudora near pup&formis.'f As these large ani-
mals no doubt required a more extended territory for their support
than that represented by the small island Anguilla, there is eA^ery
probability that the separation of these islands took place at a
late period of time and probably subsequent to the spread of the
postpliocene fauna over North America."

I think the reader will conclude from the facts Prof. Cope so
clearly presents, that the subterranean fauna of this country does
not date back of the Quaternary period. These species must have
been created and taken up their abode in these caves (Mammoth
Cave and those of Montgomery Count}r, Virginia) after the breccia
flooring their bottoms and containing the bones of Quaternary ani-
mals had been deposited ; or else migrated from Tertiary caves
farther south, which is not probable, as it has been previously
shown that those blind animals inhabiting wells immediately die
on being exposed to the light (British Sessile-eyed Crustacea, i,
p. 313), though the blind craw fish is not thus affected.

The case becomes much simpler when we consider the age of
the rocks in which the Adelsberg and other caves mentioned by
Schiodte are situated. The Alps were under water in the Middle
Eocene ; consequently the caves could not have been formed until
the close of the Tertiary. Hence the species of the cave fauna
were evidently created either at the close of the Tertiary, or more
probably the beginning of the Quaternary, as "even in the later
part of the Pliocene era there was an elevation of three thousand
feet in a part of the Island of Sicily" (Dana). We are therefore
led to conclude that the species of the subterranean fauna the
world over are recent creations, probably not older than the ex-
tinct mammals associated with man.

*Loc. cit. 1839, 183; 1870, 608. A fourth species of gigantic Chinchillid has been found
by Dr. Rijgersma, which may be called Loxomylus quadrans Cope. It is represented by
portions of jaws and teeth of three individuals. It is one of the largest species, equal-
ling the L. latidens, and has several marked characters. Thus the roots of the molars
are very short, and the triturating surface oblique to the shaft. The roots of the second
and fourth are longer than those of the first and third. The last molar has four dental
columns instead of three as in the other Loxomyli, and is triangular or quadrant-shaped
in section ; the third is quadrangular in section, and has three columns. The second is
the smallest, being only .(> the length of the subtriangular, first. Length of dental series
m .063 or 2.5 inches. Palate narrow and deeply concave. There is but little or no lat-
eral constriction in the outlines of the teeth; the shanks are entirely straight. In its
additional dentinal column, this species approaches the genus Amblyrhiza.

The large Chinchillas of Anguilla are as follows, Loxomylus lortgide
quadrant* , and AmUi/rhiza inundata.

fSee Bland, Proceed. Amer. Phil. Soc., 1871, 58.

26 ORIGIN OF CAVE LIFE.

Assuming on the principles of evolution that the cave animals
were derived from other species changed by migration from the
outer world to the new and strange regions of total darkness, it
seems evident that geologically speaking the species were suddenly
formed, though the changes may not have been wrought Until after
several thousand generations. According to the doctrine of natu-
ral selection, by which species pass from one into another by
a great number of minute variations, this time was not sufficient for
the production of even a species, to say nothing of a genus. But
the comparatively sudden creation of these cave animals affords, it
seems to us, a very strong argument for the theory of Cope and
Hyatt of creation by acceleration and retardation, xwhich has been
fully set forth in this journal. The strongly marked characters
which separate these animals from their allies in the sunlight, are
just those fitting them for their cave life and those which we would
imagine would be the first to be acquired by them on being re-
moved from their normal habitat.

On introducing the wingless locust, Ceuthophilus maculatus,
into a cave, where it must live not under stones, but by clinging to
the walls, its legs would tend to grow longer, its antennae and
palpi would elongate and become more delicate organs of hearing
as well as touch,* and the body would bleach partially out, as we
find to be the case in H. subterranea and C. stygia. The Carabid
beetle, Anopthalmus, extending farther into the cave, would lose
its wings (all cave insects except the Dipterahave no wings, elytra
excepted) and eyes, but as nearly all the family are retiring in
their habits, the species hiding under stones, its form would not
undergo farther striking modification. So with the blind Campo-
dea, which does not differ from its blind congeners, which live
more or less in the twilight, except in its antennae becoming
longer. The blind Adelops, but with rudiments of eyes, does not
greatly depart in habits from Catops, while on the other hand the
remarkable Stagobius of the Illyrian caves, which according to

* After writing this article, and without knowledge of his views, we turned to Darwin's
Origin of Species to learn what he had to say on the origin of cave animals. He attri
butes their loss of sight to disuse, and remarks :—" By the time an animal has reached,
after numberless generations, the deepest recesses, disuse will on this view have more
or less perfectly obliterated its eyes, and natural selection will often have effected
other changes, such as an increase in the length of the antenna? or palpi, as a compen-
sation for blindness." 5th Amer. Edit., p. 143. We are glad to find our views as to the
increase in the length of the antennae and palpi compensating for the loss of eyesight,
confirmed by Mr. Darwin.

ORIGIN OF CAVE LIFE. 27

Schiodte spends its life in crawling ten to twenty feet above the
floor -over the columns formed by the stalactites, to which unique
mode of life it is throughout perfectly adapted, is remarkably
different from other Silphids. Its legs are very long and inserted
far apart (the prothorax being remarkably long), with surprisingly
long claws, while the antennae, again, are of great length and
densely clothed with hairs, making them most delicate sense or-
gans.* So also are the limbs of the false scorpion, and the spi-
der and pill bug (Titanethes) of remarkable length.

But the modifications in the body of the Spirostrephon are such
that many might deem its aberrant characters as of generic impor-
tance. It loses its eyes, which its nearest allies in other, but
smaller, caves possess, and instead gains in the delicate hairs on
its back, which evidently form tactile organs of great delicacy ;
the feet are remarkably long, as also the antennae. These are not
new formations but simply modifications, apparently by use or dis-
use, of organs present in the other species. The aberrant myrio-
pod and Stagobius are paralleled by the blind fish, an animal so
difficult to classify, and so evidently adapted for its abode in end-
less darkness. And as an additional proof of the view here taken
that these cave animals are modified from more or less allied spe-
cies existing outside of the caves, we have the case of the craw
fish, whose eyes (like those of the mole), are larger in the young
than adult, indicating its descent from a species endowed with the
faculty of sight, while in the adult the appendages are modified as
tactile organs so as to make up for its loss of eyesight, in order
that it may still take its prey.

We thus see that these cave animals are modified in various
w^iys, some being blind, others very hairy, others with long ap-
pendages. All are not modified in the same way in homologous
organs ; another argument in proof of their descent from ancestors

* Schiodte remarks that "it is difficult to understand the mode of life of Stagobius
troglodytes; or how this slow and defenceless animal can escape being devoured by the
rapid, piratical Arachnidans, or find adequate support on columns, for inhabiting
which it is so manifestly constructed. We are led in this respect to consider the anten-
na?. Whatever signilicauce we attach to those enigmatical organs, we must admit that
they are organs of sense, in which view an animal having them so much developed as
Stagobius, must possess a great advantage over its enemies, if these be only Arachni-
dans. Its cautious and slow progress, and its timid reconnoitring demeanor, fully
indicate that it is conscious of life being in perpetual danger, and that it endeavors to
the utmost to avoid that danger. Darkness, which always favors the pursued more
than the pursuer, comes to its aid, especially on the uneven excavated surface of the
columns."

ORIGIN OF CAVE LIFE.

whose habits varied, as those of their out-of-door allies do at pres-
ent. Had they been specially created for subterranean life, we
should have expected a much greater uniformity in the organs
adapting them to a cave life than we actually find to be the case.

Another fact of interest in this connection is the circumstance
that these cave species breed slowly, being remarkably poor in in-
dividuals ; they are nearly all extremely rare.* Did they breed as
numerously as their allies in the outer world the whole race would
probably starve, as the supply of food even for those which do
live is wonderfully limited.

It is now known that animals inhabiting the abysses of the sea are
often highly colored : light must penetrate there, for we know that
were the darkness total they would be colorless like the cave insects.

In view of the many important questions which arise in relation
to cave animals, and which have been too imperfectly discussed
here, we trust naturalists the world over will be led to explore
caves with new zeal, and record their discoveries with minuteness,
and the greatest possible regard to exactness. The caves of the
West Indian Islands should first of all be carefully explored.
Also those of Brazil, those of the East Indies and of Africa,
while fresh and most extended explorations of our own Mammoth
Cave should be made, perhaps by a commission acting under gov-
ernment or State authority, in order that the most ample facili-
ties may be afforded by the parties owning the cave.

NOTE.— Since my article was printed, Prof. Cope's article entitled "Life in the Wyan-
dotte Cave" has appeared in the "Annals and Magazine of Natural History" (Lon-
don) for November. He enumerates the following articulates as inhabitants of this
cave; " Anophtlialmus Telllcampfii, and another species; two species of Staphylinidae;
Raphidophora ; two species of flies; an Aranea-like and Opiliolike spider; a species of
Pseudotremia; Cambarus pellucidus, an unknown aquatic Crustacean with external
egg pouches, and a Lernsean (crustacean) parasitic on the blind fish. Of these one
beetle (Anophtlialmus), the cricket (Raphidophora), a fly, the Opilio-like spider, the cen-
tipede, and the blind crawfish, are probably the same as those found in the Mammoth
Cave. Two beetles and two crustaceans are certainly different from those of the
latter, and the centipedes are much more numerous. The Gammaroid crustarean
found in the waters of the Mammoth Cave, and which is, no doubt in part, the food of
the blind fish, we did not find; but some such species no doubt exists, as we found an
abundance of a lively little tetradecapod crustacean near the mouth of a cave close by."

* The wingless grasshoppers are common however, and Prof. Hagen writes me that
the cave insects in Europe are probably not so rare as they are thought to be by natu-
ralists, since the guides do not show the best collecting places, wishing to keep a stock
on hand to sell to visitors.

CHAPTER III.

THE BLIND FISHES OF THE MAMMOTH CAVE AND THEIR

ALLIES.*

BY F. W. PUTNAM.

THE blind fish of the Mammoth Cave has from its discovery
been regarded with curiosity by all who have heard of its exis-
tence, while anatomists and physiologists have considered it as one
of those singular animals whose special anatomy must be studied
in order to understand correctly facts that have been demonstrated
from other sources ; and, in these da}Ts of the Darwinian and devel-
opment theories, the little blind fish is called forth to give its tes-
timony, pro or con.

Before touching upon this point, however, we must call attention
to the structure of the fish and its allies, and to others that are
either partially or totally blind.

In the lancelet (Branchiostoma) and the hag (Myxine) the eye
is described "as simple in form as that of a leach, consisting sim-
ply of a skin follicle f coated by a dark pigment, which receives the
end of a nerve from the brain." Such an eye speck as this struc-
ture gives would only answer for the simple perception, of light. In
the young J of the lampreys (Petromyzon) the eye is very small and

* From the AMERICAN XATTRALIST for January, 1872.

t Sae further on where Prof. Wyinan questions this structure.

t These young lampreys have been described under the generic name of Ammocoetes,
and it was not until 1S.36, when Prof. Miiller discovered the fact of a metamorphosis in
the lampreys, that their true position was ascertained. Prof. Miiller has traced the
history of the common European species and shown that it is three or four years in
attaining its perfect form. With this fact before us and with the early stages of the
Myxinoids still unknown, have we not some reason for suspecting that the Lancelet may
yet prove to be a larval form of the Myxinoids, notwithstanding that it is said to lay
eggs ? Why should we not suspect the existence in the very lowest vertebrates of some-

(29)

30 BLIND FISHES IN GENERAL.

placed in a fold of the skin of the head, and probably of little use,
as these young remain buried in the sand ; but as they attain ma-
turity, and, with it, the parasitic habits of the adult, their eyes are
developed to a fair size, thus reversing the general rule in the class.

In most other fishes the eyes are developed to a full and even
remarkable extent as to size and perfection of sight in water.
In Anableps, or the so called four eyed fish of the fresh waters of
Central and South America, which belongs to a closely allied fam-
ily with our blind fish, the Cyprinodontidve, the eyes are not only
fully developed, but are divided into an upper and lower portion
in such a way, by an opaque horizontal line, as to give the effect
of two pupils, by which the fish probably sees as well when follow-
ing its prey on the surface with its eyes out of water, as when
under water. But it is in the interesting family of cat fishes (Siiu-
ridce) that we find the most singulaf arrangement of eyes in per-
fect adaptation to the diversified modes of life of the numerous
species. In this family the eyes assume nearly every possible mod-
ification from partial and even total blindness to perfectly develop-
ed eyes, and these organs are placed in almost every conceivable
position in a fish's head ; from the ordinary large eyes on the side,
to small ones on top of the head, enabling the fish to see only what
is above ; to the oval eyes on the side, in some just back of the
mouth, situated in such a way that the fish can only see what is in
close proximity to its jaws or even below them. Many genera of
this family found in South America,* Africa f and Asia,! have the
eyes so small and buried under the skin or protected by folds or
cartilage, as evidently to be of no more use than simply to distin-
guish light from darkness.

Among the most interesting forms of this family, in this respect,
is the genus described by Prof. Cope under the name of Gronias
nigrilabris. This fish is very closely allied to our common bull

thing akin to "alternate generation," or of larvae capable of reproduction? Without
having any facts to support such an assumption, except that, on general principles, the
young of Myxine would probably be very much like Branchiostoma, and that its young
is not known, while Branchiostoma has only been found in waters where some species
of Myxinoid exists, I think that before the position of the laucelet is firmly established
we must know the embryology of the Myxinoids; for should the lancelet prove not to
be the young of the Myxinoids, it must necessarily form a distinct class of animals,
perhaps as near to the mollusks as to the vertebrates.

* Pimelodus cyclopium of Humboldt, Helogenes, Agoniosus and other genera.

f Eatropius congensis.

\Ailia, Shilbiclithys, Bagroides and other genera.

THE PENNSYLVANIA BLIND FISH. 31

pout or horned pout, and of about the same size (ten inches in
length). It was taken in the Conestoga river in Lancaster Co.,
Penn., where it is "occasionally caught by fishermen and is sup-
posed to issue from a subterranean stream said to traverse the
limestone in that part of Lancaster Co.. and discharge into the
Conestoga." We quote the following from Prof. Cope's remarks
on the fish: *-

"Two specimens of this fish present an interesting condition of
the rudimental eyes. On the left side of both a small perforation
exists in the corium, which is closed by the epidermis, representing
a rudimental cornea ; on the other the corium is complete. Here
the eyeball exists as a very small cartilaginous sphere with thick
walls, concealed by the muscles and fibrous tissue attached, and
filled by a minute nucleus of pigment. On the other the sphere is
larger and thinner walled, the thinnest portion adherent to the
corneal spot above mentioned ; there is a lining of pigment. It
is scarcely collapsed in one, in the other so closely as to give a
tripodal section. Here we have an interesting transitional condi-
tion in one and the same animal, with regard to a peculiarity which
has at the same time physiological and systematic significance,
and is one of the comparatively few cases where the physiological
appropriateness of a generic modification can be demonstrated. It
is therefore not subject to the difficulty under which the advocates
of natural selection labor, when necessitated to explain a structure
as being a step in the advance towards, or in the recession from,
liny unknown modification needful to the existence of the species.
In the present case observation on the species in a state of nature
may furnish interesting results. In no specimen has a trace of
anything representing the lens been found."

When we remember that the lens of the eye in Amblyopsis has
been found, even though the eye is less developed in all its parts
than in Gronias, it is probable that a careful microscopical exami-
nation would show its existence in this genus also.

It is interesting to note that this fish is black above (lighter on
the sides and white below), notwithstanding its supposed subter-
ranean habits, and that all the other members of the family having
rudimentary or covered eyes are also dark colored, while the blind
fishes of the Mammoth Cave and of the caves in Cuba are nearly
colorless. This want of color in the latter fishes has been consid-
ered as due to their subterranean life. If this be the cause, why
should the blind cat fishes retain the colors characteristic of the
other members of the family living in open waters ?

* Proceedings of the Academy of Natural Sciences of Philadelphia for 1864, p. 231.

THE BLIND FISHES OF CUBAN CAVES.

The fishes which in a general way, -so far as blindness, tactile
sense and mode of life are concerned, come the nearest to the blind
fishes of the Mammoth Cave, are those described by Prof. Poey*
under the names of Lucifuga subterraneus- and L. dentatus^
These fishes having the broad, flattened, fleshy head, with minute
cilia, without external eyes, and inhabiting caves so similar in
structure to the Mammoth Cave, make a comparison of them with
the fishes of the Mammoth Cave most interesting. This is greatly
enhanced by the fact that the Cuban fishes belong to a family of
essentially marine habit, quite far removed from Amblyopsis.
The fresh water ling (Lota), belonging to the same great group
of fishes (though to a distinct family or subfamily) containing .the
cod on the one hand and the Cuban blind fish on the other, is
probably the nearest fresh water relative of the Cuban fish, but

Fig. 1.

Blind Fish (Stygicola dentatus) from Caves iu Cuba.

the nearest representative yet known is the marine genus Brotula,
one species of which is found in the Caribbean Sea.

In the Cuban blind fish we find ciliary appendages on the
head and body quite distinctly developed, evidently of the same
character as those of Amblyopsis and answering the purpose of
tactile organs. These cilia are in the form of small, but 'plainly
visible, protuberances (reminding one of the single fleshy protu-
berance over the opercular opening just back of the head in Ambly-
opsis). There are eight of these on top of the head of a speci-
men I hastily examined, received from Prof. Poey by the Museum
of Comparative Zoology, and quite a number arranged in three

*Memorias Sobre la Historia Natural de la Isla de Cuba, por Felipe Poey. Tomo 2,
pp. 95-llt. Pis. 9, 10, 11. Habana, 1856-8.

fThis species was afterwards referred to the genus Stygicola Gill, on account of the
presence of palatine teeth which are wanting in the other species. There are also sev-
eral other good characters, to judge from the figures of the head, skull and brain given
by Poey, that would warrant the reference of the fish to a distinct genus from L. suiter-

FIRST NOTICE OF THE BLIND FISH. 33

rows on each side of the body, showing that tactile sense is well
developed in this fish ; though it is rather singular that the barbels
on the jaws, so usually developed as organs of touch in the cod
family and its allies, are entirely wanting in this fish.

The brain of Lucifuga subterraneus, as represented by the figures
of Poey, differs very much from that of L. dentatus and of Ambly-
opsis. In all, the optic lobes are as largely developed as in allied
fishes provided with well developed eyes. In Lucifuga subterra-
neus the cerebral lobes are separated by quite a space from the
round optic lobes, which are represented as a little larger than
the cerebral lobes, and also of greater diameter than the cerebel-
lum ; this latter being more developed laterally than in either L.
dentatus or in Amblyopsis. The three divisions of the brain
are represented, from a top view, as nearly complete circles
(without division into right and left lobes), of which that repre-
senting the optic lobes is slightly the largest. In L. dentatus the
px'ocencephalon and the optic lobes are represented as divided into
right and left lobes, as in Amblyopsis, and the cerebellum does
not extend laterally over the medulla oblongata as in L. subterra-
neus, but, as in Amblyopsis (PL 1, fig. Id), is not so broad as the
medulla, and, projecting forwards, covers a much larger portion of
the optic lobes than is the case in L. subterraneus.

The Cuban blind fish has the body, cheeks and opercular bones
covered with scales. As in Amblyopsis the eyes exist, but are so
imbedded in the flesh of the head as to be of no use. The out-
line cut here given (Fig. 1), copied from Poey, is very character-
istic of the form of the fish, but does not exhibit the fleshy cilia
or details of scaling.

The first notice that I can find of the Mammoth Cave blind fish
is that contained in the "Proceedings of the Academy of Natural
Sciences of Philadelphia/' Vol. 1, page 175, where is recorded the
presentation of a specimen to the Academy by "W. T. Craige, M.
D., at the Meeting held on May 24, 1842, in the following words : —

"A white, eyeless crayfish (Astacus Bartoni?) and ajmaall white
fish, also eyeless (presumed to belong to a subgenus of Silurus),
both taken from a small stream called the 'River Styx' in the
Mammoth Cave, Kentucky, about two and one-half miles from the
entrance."

Dr. DeKay in his "Natural History of New York, Fishes," page
187, published in J842, describes the fish, from a poor specimen in

MAMMOTH CAVE. 3

34 THE BLIND FISHES OF MAMMOTH CAVE.

the Cabinet of the Lyceum of Natural History of New York,
under the name of Amblyopsis * spelceus.^ DeKay's description is
on the whole so characteristic of the fish as to leave no doubt as
to the species he had before him, though the statement that it has
eight rays supporting the branchiostegal membrane (instead of
six), and that the eyes are "large" but under the skin, must have
been due to the bad condition of his specimen and to his taking
the fatty layer covering the minute eyes for the eyes themselves,
as pointed out by Prof. Wyman. Dr. DeKay places the genus with
the Siluridse (cat fishes) but at "the same time questions its con-
nection with the family and says that it will probably form the
type of a new family. In 1843 Prof. Jeffries Wyman \ gave an
account of the dissection of a specimen in which he could not find
a trace of the eye or of the optic nerve, probably owing to the
condition of the specimen, as he afterwards § found the eye spots,
and made out the structure of the eye. When describing the
brain, Prof. "Wyman calls attention to the fact of the optic lobes
being as well developed as in allied fishes with well developed eyes,
and asks if this fact does not indicate that the optic lobes are the
seat of other functions as well as that of sight. He also calls
attention to the papillae on the head as tactile organs furnished
with nerves from the fifth pair.

Dr. Theo. Tellkampf || was the first to point out the existence of
the rudimentary eyes from dissections made by himself and Prof.
J. Miiller, and to state that they can be detected in some specimens
as black spots under the skin by means of a powerful lens. Prof.
Wyman afterwards detected the eye through the skin in several
specimens. Dr. Tellkampf also was the first? to call attention to
the " folds on the head, as undoubtedly serving as organs of touch,
as numerous fine nerves lead from the trigeminal nerve to them
and to the skin of the head generally."

It is also to Dr. Tellkampf that we are indebted for the first
figure of the fish,^[ and for figures illustrating the brain, and inter-
nal organs. The descriptions of the anatomy of the fish by Drs.

* Obtuse vision, f Of a cave.

t Silliman's Journal, Vol. 45, p. 94.

§ Proceedings Boston Soc. Nat. Hist., Vol. 4, p. 395. 1853.

|| Miiller's Archiv. fur Anat., 1844. p. 392. Reprinted in the New York Journal of Med-
icine for July, 1845. p. 84, with pliito.

IT The only other figures of the species, that I am aware of, are the simple outlines
given in Poey's Mem. de Cuba, the woodcut in Wood's Illustrated Natural History and
the cut in Tenney's Zoology. None of these figures are very satisfactory.

THE BLIND FISH FAMILY. 35

Tellkampf and Wyman are all that have ever been written on the
subject of any importance, with the exception of the description
of the eye by Dr. Dalton, whose paper, in the "New York Medical
Times," vol. 2, p. 354, I have not seen. Prof. Poey gives a com-
parison of portions of the structure with that of the Cuban blind
fishes.

Dr. Tellkampf proposed the name of Heteropygii* for the family
of which, at the time, a single species from the Mammoth Cave was
the only known representative, and makes a comparison of the char-
acters with those of Apliredoderus Say anus, a fish found only in the
fresh waters of the United States, and belonging to the old family
of Percoids, but now considered as representing a family by itself,
though closely allied to the North American breams (Pomotis), and
having the anal opening under the throat as in the blind fish.

Dr. Storer,f not knowing of Dr. Tellkampf 's paper, proposed
the name of Hypsceidce, for the blind fish, and placed it between
the minnow and the pickerel families, in the order of Malacoptery-
gian, or soft rayed, fishes. According to the system adopted by
Dr. Giinther, it stands as closely allied to the minnows, Cyprino-
dontidce (many of which are viviparous and have the single ovary
and general character of the blind fish), and the shiners, Cyprini-
dm, of the order of Physostomi. Dr. Tellkampf, in discussing the
relations of the family, points out its many resemblances to the
family of Clupesoces, and its differences from the Siluroids, Cy-
prinodontes and Clupeoids, with which it has more or less affinity,
real or supposed. Prof. Cope in his paper on the Classification of
Fishes j places the Arnblyopsis in the order of Haplomi with the
shore minnows, pickerel and mud fish, and in an article on the Wy-
andotte Cave,§ he says that the Cyprinodontes (shore minnows)
are its nearest allies. This arrangement by Prof. Cope places
the Haplomi between the order containing the herrings and that
containing the electric eel of South America, all included with the
garpike, dog fish of the fresh waters (Amia), cat fishes, suckers
and eels proper, etc., etc., in the division of Physostomi as limited
by him.

*From the advanced po^iti >n of the terminus of the intestine being so different from
the position wliich it has in ordinary fishes.

t Synopsis of the Fishes of North America, published in 1846.

J American Naturalist, Vol. 5, p. 579, 1871.

§ Indianapolis Daily Journal of September 5, 1871. Reprinted in Ann. Mag. Nat.
Hist., Nov., 1871.

36 FISHES WITH EYES ALSO IN THE CAVE.

Prof. Agassiz in 1851* stated that the blind fish was an aber-
rant form of the Cyprinodontes.

Thus all those authors who have expressed an opinion as to the
position which the fish should hold in the natural system have
come to the same conclusions as to the great group, division, or
order, into which it should be placed. For all the terms used
above, when reduced to any one system, bring Amblyopsis into
the same general position in the S3^stem ; its nearest allies be-
ing the minnows, pickerels, shiners and herrings ; and unless a
careful study of its skeleton should prove to the contrary, we
must, from present data, consider the family containing Amblyop-
sis as more nearly allied to the Cyprinodontes, or our common
minnows having teeth on the jaws, than to any other family, differ-
ing from them principally by the structure of the several parts of
the alimentary canal and the forward position of its termination.

I have thus far mentioned only one species of blind fish from
the cave, the Amblyopsis spelceus. The waters of the cave not only
contain another species of blind fish, differing from Amblyopsis in
several particulars, especially by its smaller size and by being with-
out ventral fins, which I have identified as the Typliliclithys subter-
raneus of Dr. Girard ; but also a fish with well developed eyes,
as proved by the account given by Dr. Tellkampf and by the
drawing of a fish found by Prof. Wyman, in 1856, in the stomach
of an Amblyopsis he was dissecting. In order to call attention to
the fact that fishes with eyes are at times, if not always, in the
waters of the cave, I have reproduced the drawing by Prof. Wy-
man on plate 1, fig. 13. It is very much to be regretted that the
specimen is not now to be found, and that it was so much acted
on by the gastric juice as to destroy all external characters by
which it could be identified from the drawing, which is of about
natural size. Dr. Tellkampf's remarks on the fish with eyes are
as follows : —

"• Besides the colorless blind-fish, there are also others found in
the cave, which are black, commonly known by the name of 'mud-
fish.' I saw a dark-colored fish in the water, but did not succeed
in catching it. The latter are said to have eyes, and are entirely
dissimilar to the blind-fish."

The name "mud-fish," given to this fish with eyes, and the state-
ment that it is of a dark color, together with the drawing by Prof.

*SUliman's Journal, p. 128.

HABITS OF THE BLIND FISH. 37

Wyman of the fish found in the stomach of the blind fish, showing
the position of the dorsal fin to be the same as in the fish commonly
called mud fish in the fresh waters of the Middle, Western and
Southern States, perhaps, indicates the fish with eyes to be a spe-
cies of Melanura.* This fish is called mud fish from the habit it has
of burying itself in the mud, tail first, f to the depth of two to four
inches, and of remaining buried in the mud in our western ditches
during a time of drought. This habit, perhaps, in a measure fits
it for a subterranean life. The occurrence of a fish belonging
to the same family with the blind fish, but with well developed
eyes, in the subterranean streams in Alabama, as mentioned further
on and figured on PL 2, fig. 4, however, renders it probable that the
cave fish with eyes may be the same or an allied species, and the
drawing by Prof. Wyman would answer equally as well for it.

The farct that the Amblyopsis succeeded in catching a fish of,
probably, very rapid and darting movements, shows that the tactile
sense is well developed and that the blind fish must be very active
in the pursuit of its prey ; probably guided by the movement
which the latter makes in the water so sensibly influencing the del-
icate tactile organs of the blind fish that it is enabled -to follow
rapidly, while the pursued, not having the sense of touch so fully
developed, is constantly encountering obstacles in the darkness.

In describing the habits of the blind fish Dr. Tellkampf says : —

"It is found solitary, and is very difficult to be caught, since it
requires the greatest caution to bring the net beneath them with-
out driving them away. At the slightest motion of the water they
dart off a short distance and usually stop. Then is the time to
follow them rapidly with a net and lift them out of water. They
are mostly found near stones or rocks which lie upon the bottom,
but seldom near the surface of the water."

Prof. Cope, in describing the habits of the blind fish which he

* Dr. Gunther considers the genus Melanura of this country to be synonymous with

Umbla of Europe. In each
country only one species has
been as yet satisfactorily de-
scribed.

t See the interesting notes
on the habits of the mud min-
now, by Dr. Abbott in Amer-
ican Naturalist, Vol. 4, pages
Mud fish (Melanura Umi). 107 and 388, with figure of

the fish on page 385, which we here reproduce for comparison.

YOUNG OF THE BLIND FISH.

obtained in a stream that passes into the 'Wyandotte Cave, though
he entered it b}^ means of a well in the vicinity of the cave, says
that : —

"If these Amblyopses be not alarmed they come to the surface
to feed, and swim in full sight like white aquatic ghosts. They are
then easily taken by the hand or net, if perfect silence be pre-
served, for they are unconscious of the presence of an enemy
except through the sense of hearing. This sense is, however, evi-
dently very acute, for at any noise they turn suddenly downward,
and hide beneath stones, etc., on the bottom. They must take
much of their food near the surface, as the life of the depths is
apparently very sparse. This habit is rendered easy by the struc-
ture of the fish, for the mouth is directed upwards, and the head is
very flat above, thus allowing the mouth to be at the surface."

The blind fish has a single ovary, in common with several genera
of viviparous Cyprinodontes. In three female specimens of Am-
blyopsis which I have opened, the ovary was distended with large
eggs, but no signs of the embryo could be traced. In these three
specimens it was the right ovary that was developed, and this, as
in the figure (Plate 2, fig. Ic), was by the side of the stomach
and did not extend beyond it. The number of eggs contained in
the ovary was not far from one hundred in the specimen figured.
As the embryos develop, the mass probably pushes further
back in the cavity and also extends the abdominal walls. That
the fish is viviparous is proved by the statement made by Mr.
Thompson before the Belfast Natural History Society,* that one
of the blind fishes from the cave, four and a half inches long,
uwas put in water as soon as captured, where it gave birth to
nearly twenty young, which swam about for some time, but soon
died. These, with the exception of one or two, were carefully
preserved, and fifteen of them are now before us [at the meeting,
I wish they were here] , they were each four lines in length."

It is singular that no mention is made regarding these young, as
to the presence or absence of eyes, and, as if it was fated that this
important point should remain unnoticed as long as possible, it is
equally singular that Dr. Stein dachner omitted to examine some
very young specimens which he received from a friend a few
months since and sent to the Vienna Museum, where they will
remain unexamined until he returns there. I saw the Doctor only

* Annals and Mag. of Natural History, Vol. xiii, pp. 112, 1844.

GENERAL REMARKS ON THE BLIND FISHES. 39

a week after these, to me, interesting specimens had been sent
abroad, and he was as grieved as I was disappointed at my being
just too late to take advantage of them. (See note on p. 52.)

At what time the young are born has never been stated, but judg-
ing from such data as I can at present command, I think that it must
be during the months of September and October. Specimens col-
lected during those months would probably contain embryos in
various stages of development, the examination of which would un-
doubtedly lead to most interesting results. (See note on p. 52.)

Prof. Wyman has most generously placed in my hands his un-
published notes and drawings of the several dissections he has
made of Amblyopsis, as well as his specimens and dissections.
Many of these drawings are reproduced on Plate 1, and will, with
his notes which I here give, greatly enhance the value of this arti-
cle, as his dissections have been made with the utmost care, and
with a patience and delicacy that only a master hand attains. It
will therefore be understood that, in giving credit to Prof. Wyman
in the following pages, I refer to his unpublished notes, except
when the quotation is given from a special work. In quoting his
description of the eye and ear from " Silliman's Journal " I have
changed the references so as to refer to his drawings reproduced
on Plate 1, and not to the three cuts given in "Silliman's Journal,"
though the figures of the brain and of the otolite were copied from
those cuts.

The largest specimens I have seen of Amblyopsis are several
males and females, each from four to four and a half inches in
length, which seems to be about as large as the fish grows, though
Dr. Giinther mentions a specimen in the British Museum of five
inches in length. The largest specimen captured of late years is
said to have been taken, during the summer of 1871, and sold for
ten dollars to a person who was so desirous of securing the pre-
cious morsel that he had it cooked for his supper. The smallest
specimen I have seen was one and nine-tenths inches in length.
The general shape and character of the fish is best shown by the
figures on plates 1 and 2.

"The whole head, above and below, is destitute of scales, the
naked skin extending backwards on the sides to the base of the
pectoral fins ; the scaly portion of the body above ends in a semi-
circular edge covering the space between the upper ends of the
opercula. The skin covering the middle region of the head is

40 TACTILE ORGANS OF BLIND FISH.

smooth, but on either side is provided with numerous transverse
and longitudinal ridges (PI. 1, fig. 7), which are, on the whole,
regularly arranged. The first row of transverse ridges, eight or
nine in number, begins between the nostrils and extends back-
wards, diverging from the median line. The third ridge is crossed
at its outer end by a longitudinal one, as are also two others farther
back. The second and third rows, situated, in part, on the sides
and, in part, on the under surface, are less regular than the preced-
ing. A fourth, on the borders of the operculum, is still less Avell
defined. The transverse are also crossed here by longitudinal
ridges. About ten vertical ridges, also provided with papillae, and
similar to those on the head, are visible on the sides extending from
the pectoral fins to the tail, but are not so well defined as those on
the head. The skin of the head is of extreme delicacy and is cov-
ered by a very thin, loose layer of epithelium."— WYMAN.

"The larger ridges have between twenty and thirty papillae,
many of these having a cup-shaped indentation at the top, in
which a delicate filament is, in some instances, seen (PL 1, fig. 9).
These papillae are largely provided with nervous filaments, and, as
is obvious from their connection with branches of the fifth pair of
nerves, must be considered purely tactile, and the large number of
them shows that tactile sensibility is probably very acute and in
some measure compensates for the virtual absence of the sense of
sight. Plate 1, fig. 8, represents one of the ridges of the bead
magnified, showing the papillae of which it is made up, and figure
9 shows three papillae still more enlarged. Two of these show a
cup-shaped cavity at the^ top, and the short, slender filament al-
ready mentioned. The surface of the papillae is covered with
loosely connected epithelium cells. Fig. 10 shows the nervous fil-
aments distributed to the papillae : a, a branch of the fifth pair of
nerves passing beneath the papillary ridge and sending filaments
to each papilla. These papillary branches interchange filaments,
forming a nervous plexus in connection wTith each ridge. This
figure of the nerves was drawn with a camera lucida, from a speci-
men treated with acetic acid." — WYMAN.

"Plate 1, fig. 6, represents a double system of subcutaneous ca-
nals, which extend the whole length of the head, but were not
traced farther back than the edge of the naked or scaleless skin
which covers it. Forwards they bifurcate, nearly encircling the
nasal cavity, towards the middle line ending in a blind pouch.

THE EYES OF THE BLIND FISHES. 41

The lateral branch was not traced distinctly to an end, but seemed
to connect with the olfactory cavit}\ The walls of these canals
are exceedingly delicate and easily overlooked." — WYMAN.

"Plate 1, fig. 5, shows the globe of the eye with the optic nerve
(c), as seen under the microscope. The lens (b) is detached from
its proper place by the pressure of the glass. Irregularly arranged
muscular bands are attached to the exterior of the globe (a, a, a,
a), but were not recognized as the homologues of the muscles of
the normal eye of fishes ; nevertheless, they indicate that the globe
was movable." — WYMAN.

"In the three specimens recently dissected, the eyes were ex-
posed only after the removal of the skin, and the careful separation
from them of the loose areolar tissue which fills the orbit. In a
fish four inches in length the eyes measured one-sixteenth of an
inch in their long diameter, and were of an oval form and black. A,
filament of nerve (PI. 1, fig. 3 a) was distinctly traced from the
globe to the cranial walls, but the condition of the contents of the
cranium, from the effects of the alcohol, was such as to render it
impracticable to ascertain the mode of connection of the optic
nerve with the optic lobes.

9 Examined under the microscope with a power of about twenty
diameters, the following parts were satisfactorily made out (PI. 1 ,
fig. 3) : 1st, externally an exceedingly thin membrane, 6, which
invested the whole surface of the eye and appeared to be continu-
ous with a thin membrane covering the optic nerve, and was
therefore regarded as a sclerotic ; 2d, a layer of pigment cells, d,
for the most part of a hexagonal form, and which were most abun-
dant about the anterior part of the eye ; 3d, beneath the pigment
a single layer of colorless cells, c, larger than a pigment cell, and
each cell having a distinct nucleus ; 4th, just in front of the globe ;
a lenticular-shaped, transparent body, e [see also fig. 4], which
consisted of an external membrane containing numerous cells with
nuclei. This lens-shaped body seemed to be retained in its place
by a prolongation forwards of the external membrane of the globe ;
5th, the globe was invested by loose areolar tissue, which adhered
to it very generally, and in some instances contained yellow fatty
matter ; in one specimen it formed a round spot, visible through the
skin on each side of the head, which had all the appearance of a
small eye ; its true nature was determined by the microscope
only. It is not improbable that the appearance just referred to
may have misled Dr. DeKay — where he states that the eye exists
of the usual size, but covered by the skin.

If the superficial membrane above noticed is denominated cor-
rectly the sclerotic, then the pigment layer ma^y be regarded as the
representation of the choroid. The form as well as the position
of the transparent nucleated cells within the choroid correspond

EYES AND EARS OF THE BLIND FISH.

for the most part with the retina. All of the parts just enumer-
ated are such as are ordinarily developed from and in connection
with the encephalon, and are not in any way dependent upon the
skin. But if the lenticular-shaped body is the true representative
of the crystalline lens, it becomes difficult to account for its pres-
ence in Ambtyopsis according to the generally recognized mode of
its development (since it is usually formed from an involution of
the skin) unless we suppose that after the folding in of the skin
had taken place in the embryonic condition, the lens retreated from
the surface, and all connection with the intagument ceased. [*]

According to Quatrefages, however, the eye of Amphioxus [|]
is contained wholly in the cavity of the dura mater, and yet it has
all the appearance of being provided with a lens. If his descrip-
tion be correct, then the mode of development as well as the mor-
phology of the eye in this remarkable fish is different from that of
most other vertebrates, since the lens never could have been
^formed from an involution of the skin, nor could the eye with its
lens, as Prof. Owen asserts, be a modified cutaneous follicle.
Whatever views be taken with regard to the development of the
eye of the blind fish, the anatomical characters which have been
enumerated show, that though quite imperfect as we see it in the
adult, it is constructed after the type of the eyes of other ver-
tebrates. It certainly is not adapted to the formation of ima-
ges, since the common integument and the areolar tissue which are
interposed between it and the surface, would prevent the transmis-
sion of light to it except in a diffused condition. No pupil or any-
thing analogous to an iris was detected, unless we regard as repre-
senting the latter the increased number of pigment cells at the
anterior part of the globe.

It is said that the blind fishes are acutely sensitive to sounds
as well as to undulations produced by other causes in the water.
In the only instance in which I have dissected the organ of hear-
ing (which I believe has not before been noticed), all its parts
were largely developed, as will be seen by reference to PI. 1,
fig. 1 e. As regards the general structure, the parts do not differ
materially from those of other fishes except in their proportional
dimensions. The semi-circular canals are of great length, and
the two which unite to enter the vestibule by a common duct, it
will be seen, project upwards and inwards under the vault of the
cranium, so as to approach quite near to the corresponding parts
of the opposite side. The otolite contained in the utricle was not
remarkable, but that of the vestibule (PL 1, fig. 2) and seen in

*In birds and mammals there is a stage of development where the lids come to-
gether and firmly unite, to separate again when the animal <;gets its eyes open." In
the mole rat (Spalax typhlus) of Siberia, the lids never open, and the eyes remain
through life covered with hairy skin. It is not improbable that in Amblyopsis some-
thing analogous to this, a closing of the skin over the eye, may have taken place. — J. W.

1 1 have used the prior name of Branchiostoma in this paper when speaking of the
Lancelot.

GEOGRAPHICAL RANGE OF BLIND FISH. 43

dotted outline in fig. 1 e is quite large when compared with that of
a Leuciscus of about the same dimensions as the blind-fish here
described." — WYMAN, SillitnaiCs Journal, Vol. 17, p. 259, 1854.

The Amblyopsis spelceus undoubted!}7 has quite an extensive
distribution, probably existing in all the subterranean rivers that
flow through the great limestone region underlying the Carbonif-
erous rocks in the central portion of the United States. Prof.
Cope obtained specimens from the Wyandotte Cave and from wells
in its vicinity, and in the Museum of Comparative Zoology at
Cambridge there is a specimen labelled "from a well near Lost
River, Orange Co., Ind.," which, with those from the Wyandotte
Cave, is conclusive evidence of its being found on the northern
side of the Ohio* as well as on the southern, in the rivers of the
Mammoth Cave. I have been able to examine a number of speci-
mens from the Mammoth Cave, and have carefully compared with
them the one from the well in Orange Co., Ind., and find that the
specific characters are remarkably constant.

In 1859 1 Dr. Girard described a blind fish, received by the
Smithsonian Institution from J. E. Younglove, Esq., who obtained
it "from a well near Bowling Green, Ky." The general appear-
ance of this fish, which was only one and a half inches in length,
was that of Amblyopsis spdwus, but it differed from that species
in several characters, especially by the absence of ventral fins.
Dr. Girard therefore referred the fish to a distinct genus under the
name of Typhlichthys\ subterraneus. Dr. Giinther§ considers this
fish a variety of Amblyopsis spelceus and records the specimen in
the British Museum "' from the Mammoth Cave," as " half-grown." |j

By the kindness of Prof. Agassiz, I have been enabled to exam-
ine nine specimens of blind Jish without ventrals, in the Museum of
Comparative Zoology. Seven of these were collected in the Mam-
moth Cave by Mr. Alpheus Hyatt in September, 1859. One was
from Moulton, Lawrence County, Alabama, presented by Mr.
Thomas Peters ; and another from Lebanon, Wilson Co., Tennes-
see ; presented by Mr. J. M. Safford. It is not stated whether

* I have also been informed by Mr. Holmes of Lansing, Mich., that blind fishes have
been drawn out of wells in Michigan,
t Proceedings Acacl. Nat. Sci. Philad., p. 63.

J Blind fish.

§ Catalogue of Fishes in the British Museum, Vol. 7, p. 2, 1868.

II The largest specimen I have seen of Typhlichthys is one and seventeen-twentieths
inches in length, and the smallest Amblyopsis one and eighteen-twentieths inches.

44 OTHER SPECIES OF THE FAMILY.

these latter came from wells or caves, but probably from wells.
They are all of about the same size, one and one-half to two
inches in length, and are constant in their characters. Moreover,
four of the seven specimens from the Mammoth Cave were females
with eggs. These eggs were as large in proportion as those from
Amblyopsis. The ovary was single and situated on the right
side of the stomach, as in Amblyopsis. The difference in. the
number of eggs was very remarkable, each of the four specimens
examined having but about thirty eggs in the ovary, while in
three females of Amblyopsis (all, however, of nearly three times
the size of Typhlichthys) there were about one hundred eggs in
each. As in both species there were no signs of the embryos in
the eggs, it is not probable that any of the eggs had been developed
and the young excluded, nor is it at all likely that the great vari-
ation in the number of eggs would simply indicate different ages.
By a reference to the figures (PL 2), it will be seen that the pyloric
appendages, stomach and scales of the two fishes are different.
For these reasons, taken in connection with the absence of ven-
tral fins, I have no hesitation in accepting Dr. Girard's name as
valid for this genus, of which we thus far know of but one species,
with a subterranean range from the waters of the Mammoth Cave,
south to the northern portion of Alabama. In this connection it
would be most interesting to know the relations of the "blind
fishes" said to have been found in Michigan. For thus far we
have Typhlichthys limited to the central and southern portion of
the subterranean region, Amblyopsis to the central, and the spe-
cies in the northern portion undetermined.

In 1853, on his 'return from a tour through the southern and
western states, Prof. Agassiz gave a summary of some of his
ichthyological discoveries in a letter to Prof. J. D. Dana.* In this
letter are the following remarks : —

"I would mention foremost a new genus which I shall call Cho-
logaster, very similar in general appearance to the blind fish of the
Mammoth Cave, though provided with eyes ; it has, like Ambly-
opsis, the anal aperture far advanced under the throat, but is en-
tirely deprived of ventral fins ; a very, strange and unexpected
combination of characters. I know but one species, Ch. cornutus
Ag. It is a small fish scarcely three inches long, living in the
ditches of the rice fields in South Carolina. I derive its specific

* Published in American Journal of Sci. and Arts, Vol. 16 (2d series), p. 131, 1853.

OTHER FISHES OF THE FAMILY WITH EYES. 45

name from the singular form of the snout, which has two horn-like
projections above."

This is the only information ever published regarding this inter-
esting fish and the only specimens known are those on which Prof.
Agassiz based the above remarks.

By the kindness of Professor L. Agassiz, who has placed all
the specimens of the family contained in the Museum of Com-
parative Zoology in my hands for study, I am enabled to give a
figure and description of this interesting species from the three
specimens in the Museum, which were labelled as the originals of
Chologaster cornutus Ag., from Waccamaw, S. C., presented by
Mr. P. C. J. Weston, 1853. The largest of these specimens was
distended with eggs and I was enabled to compare the ovary with
that of Amblyopsis. From its being single and the eggs very
large, I have no doubt that it is a viviparous fish like the other
genera of the family. The position of the ovary behind the stom-
ach, as shown in fig. 2c, plate 2, and the presence of four pyloric
appendages (PL 2, fig. 2 a) instead of two, as in Amblyopsis (fig.
la) and Typhlichthys (fig. 3 a), are good internal characters, sep-
arating it from the other genera, independently of the presence
of eyes and the absence of ventral fins and papillary ridges.

The stability of the internal characters I have mentioned was
most unexpectedly substantiated by the discovery of a second
species (PL 2, figs. 4, 4 a) of the genus among the specimens in
the Museum of Comparative Zoology. I have the pleasure of
dedicating this species* to Professor Agassiz, not only in kindly
remembrance of the eight years I was associated with him as stu-
dent and assistant, but also because the fish so well illustrates the
decided position he has taken relative to the immutability of spe-
cies.

The only specimen known of this second species was drawn
from a well in Lebanon, Tenn., and presented to the Museum by
Mr. J. M. Safford, Jan., 1854. It is a more slender fish than C.
cornutus, but the intestine follows the same course and the four
pyloric appendages are present as in that species.

In the genus Chologaster f we have all the family characters as
well expressed as in the blind species, though it differs from Am-

* A Synopsis of this family with descriptions of the four species will appear in the
" Report of the Peabody Academy of Science for 1871." (Reprinted here. p. 55.)
t Literally " bile-stomach;" probably named from the yellow color of the fish.

46 ORIGIN OF THE BLIND FISHES.

blyopsis and Typhlichthys by the presence of eyes, the absence of
papillary ridges on the head and body, and by the longer intestine
and double the number of pyloric appendages, as well as by the
position of the ovary ; and agrees with Typhlichthys by the ab-
sence of ventral fins. Amblyopsis and Typhlichthys are nearly
colorless, while Chologaster Agassizii is of a brownish color similar
to many of the minnows, and C. cornutus is brownish j'ellow, with
dark, longitudinal bands.

Among the most interesting points in the history of this genus
is the fact of its occurring in two widely different localities, C.
Agassizii having been found in a well, in the same vicinity (proba-
bly in the same well) with a specimen of Typhlichthys, and
undoubtedly belonging to the same subterranean fauna west of the
Appalachian ridge, while C. cornutus belongs to the southern coast
fauna of the eastern side of that mountain chain, and is thus far
the only species of the family known beyond the limits of the great
subterranean region of the United States.

Having now given an outline of the structure, habits and distri-
bution of the four species belonging to the family, and recapitu-
lated the known facts, we are better able to consider the bearings
of the peculiar adaptation of the blind fishes, in the Mammoth
and other caves, to the circumstances under which they exist.

Prof. Cope in stating, in his account of the blind fish of the
Wyandotte Cave, "that the projecting under jaw and upward di-
rection of the mouth renders it easy for the fish to feed at the sur-
face of the water, where it must obtain much of its food," suggests
that : -

"This structure also probably explains the fact of its being the
sole representative of the fishes in subterranean waters. No doubt
many other forms were carried into the caverns since the waters
first found their way there, but most of them were like those of
our present rivers, deep water or bottom i'eeders. Such fishes
would starve in a cave river, where much of the food is carried to

them on the surface of the stream The shore minnows

are their nearest allies, and many of them have the upturned

mouth and flat head Fishes of this, or a similar famity,

enclosed in subterranean waters ages ago, would be more likely to
live than those of the other, and the darkness would be very apt
to be the cause of the atrophy of the organs of sight seen in the
Amblyopsis."

This suggestion was undoubtedly hastily made by Prof. Cope
when writing the letter which was printed in the "Indianapolis

ORIGIN OF THE BLIND FISHES. 47

Journal," and were it not tbat the article has been reprinted in the
" Annals and Magazine of Natural History," I should not criticise
the statement made in an off-hand letter for publication in a news-
paper ; for with Prof. Cope's knowledge of fishes it could simply be
a hasty thought which he put on paper, when he suggests that it is
because the Cyprinodontes have a mouth directed upwards and
are surface feeders that they were better adapted to a subterranean
life than other fishes, and hence maintained an existence, while
other species, which he supposes were introduced into the subter-
ranean streams at the same time, died out.

If the fishes of the subterranean streams came from adjoining
rivers, why were not many of the Percoids, Cyprinoids and other
forms, that are as essentially surface feeders as the Cyprinodon-
tes (many of the latter are purely "mud feeders"), as capable of
maintaining an existence in the subterranean waters as any species
of the latter ? Neither is it necessary for us to assume that the
structure of the fish should be adapted to feeding on the surface,
for not only have we in the blind cat fish, described by Prof.
Cope himself, from the subterranean stream in Pennsylvania,
an example of a fish belonging to an entirely different family of
bottom feeders, thriving under subterranean conditions, but the
blind fishes of the Cuban caves are of the great group of cod fishes
which are, with hardly an exception, bottom feeders. The fact
that the food of the blind fishes of the Mammoth Cave consists in
great part of the cray fish found in the waters of the cave, as
shown by the contents of several stomachs I have examined, and
also that one blind fish at least made a good meal of another fish,
as already mentioned, shows that the}T are not content with simply
waiting for what is brought to them on the surface of the water,
and that they are probably as much bottom as surface feeders.

Again, in regard to the sense of sight, why is it necessary to
assume that because fishes are living in streams where there is lit-
tle or no light, that it is the cause of the non development of
the eye and the development of other parts and organs? If this
be the cause, how is it that the Chologaster from the well in Ten-
nessee, or the "mud fish" of the Mammoth Cave are found with
eyes? Why should not the same cause make them blind if it made
the Amblyopsis and Typhlichthys blind? Is not the fact, pointed
out by Prof. Wyman, that the optic lobes are as well developed in
Amblyopsis as in allied fishes with perfect eyes, and, I may add,

48 ORIGIN OF THE BLIND FISHES.

as well developed as those of Chologaster cornutus, an argument
in favor of the theory that the fishes were always blind and that
they have not become so from the circumstances under which they
exist? If the latter were the case and the fishes have become blind
from the want of use of the eyes, why are not the optic lobes also
atrophied, as is known to be the case when other animals lose their
sight ? I know that many will answer at once that Amblyopsis
and Typhlichthys have gone on further in the development and
retardation of the characters best adapting them to their subterra-
nean life, and that Chologaster is a very interesting transitionary.
form between the open water Cyprinodontes and the subterranean
blind fishes. But is not this assumption answered by the fact that
Chologaster has every character necessary to place it in the same
family with Amblyopsis and Typhlichthys, while it is as distinctly
and widely removed from the Cyprinodontes as are the two blind
genera mentioned ?

Assuming, for the moment, that Chologaster is a transitional
form between the surface feeding Cyprinodontes, and Typhlichthys
and Amblyopsis, let us recapitulate the characters that distinguish
the different forms and see if they exhibit transitions, and if Cho-
logaster is traversing the slow developmental road to Ambtyopsis.

Allowing all characters embraced in the general structure of the
skeleton, brain, scales, fins, etc., as" ordinal, and common to both
Cyprinodontes and Heteropygii, we will recapitulate only such as
can be considered of family and generic value in the two groups.

CYPRINODONTES. CHOLOGASTER. TTPHLICHTHYS. AMBLYOPSIS.
Surface feeders. In part. Unknown. Partially. The same.

The same.

The same.

Viviparous. Many genera. Probably. Probably. Undoubted-

*y-

Ovary. Single in vivipa- Single and Single and The same,

rous genera* placed behind placed at eide
and placed by the stomach. of stomach,
the side of in-
testine in some
and posterior in
others.

* The ovary is also single in other genera of viviparous flahes belonging to distinct
orders.

Intestine.

Stomach tfpylor-
ic appendages.

In many genera
long and convo-
luted, in others
short and with
single turn.

In most, if not
all, stomach not
well defined
from intestine
and without ap-
pendages.

Moderately
long with two
turns.

Stomach well
defined, 003-
cal, with two
p y 1 o r i c a p-
penrlages on
each side.

Shorter with two
turns.

The same, with
one pyloric ap-
pendage on
each side.

ORIGIN OF THE BLIND FISHES.

49

Anal opening.

In normal posi-

Forward of pec-

The same.

The same.

tion.

torals.f

Air bladder.

Present in few

Present.

The same.

The same:

genera. J

Scales.

On body regu-
larly imbricat-

Irregularly ar-
ranged, firm-

The same.

The same.

ed and loosely

ly attached

attached.

by being cov-

ered in great

part by the

cuticle.

Head with scales

With scales.

Naked.

The same-.

The same.

or naked. §

Tactile papillce ||

Absent.

Absent.

Very prominent

The same.

on the head and

as ridges on

body.

the head and

sides of body.

Central fins.V

Present in most

Absent.

Absent.

Present.

genera, absent

in at least two.

Eyes**

Well developed in
all.

Well developed
and normal.

Rudiment a ryft
and of no use.

The same.

Habitat.

Fresh water ;

Limestone wa-

Limestone wa-

The same.

brackish water;

ter of subter-

ter of subter-

salt water.

ranean riv-

ranean rivers.

ers. Brackish

water ?

Geographical
range.

Nearly all parts of
the world.

One species in
subterranean

Central & south-
ern portion of

Central and
N. central

streams of S.

subterranean

portion of

central por-

fauna of Unit-

same.

tion of the U.

ed States.

S.; a 2d spe-
cies in the So.

Atl. coast fau-

na of U.S.

From this brief comparison of some of the prominent charac-
ters of the genera of the Heteropygii with the Cyprinodontes, their

f Aphredoderus and Gymnotus, and other genera of distinct orders have this forward
position of the anus also.

J The air bladder is in several families present in some species and absent in others.

§ The presence or absence of scales on the head, or on portions of it, is a generic
character subject to great variation in many families and quite constant in others.

II I cannot recall anything but the barbels on the head and jaws of many genera of
Cyprinoids, Siluroids, Gadoids, etc., etc., that can be said to be tactile organs among
fishes, with the exception of the fleshy papillae on the head and body of the blind fishes
of the American and Cuban caves, and the filaments of the fin rays of many fishes
and the fleshy ventral rays of the Gurnards.

IT Of all fins, the ventrals are the most likely to deviate from their normal structure
and position. Their presence or absence, as exhibited in many families, and often by
different ages of the same fish, and the great variation in their ppsition in different
genera of the same family, render any change in them of either generic, specific, or
individual character, or simply indicative of age (as they are lost in some adult fishes
while present in the young, and in others not developed until after the other fins).

** As I have alluded to the fact, in the first part of this paper, the eyes of fishes are
no more the constant and unvarying part of the fish structure than the ventral fins, and
like them are subject to almost every conceivable variation in position in the head, and
perfection in structure.

tfThe largest specimen I have seen of Typhlichthys, is less than two inches in length
and as the eye of an A^mblyopsis of twice the size is not over a 32d of an inch in width
it must be very small indeed in Typhlichthys, and I confess to not being able to find it
in an ordinary dissection, assisted only by a good lens.
MAMMOTH CAVE. 4

50 . ORIGIN OF THE BLIND FISHES.

acknowledged nearest allies, we can only trace what could be
regarded as a transition, or an acceleration, or a retardation of
development, in simply those very characters, of eyes and ventral
fins, that are in themselves of the smallest importance in the struc-
ture (permanence of character considered) of a fish, and, as if to
show that they were of no importance in this connection, we find
in the same cave, blind fishes with ventrals and without ; and in
the same subterranean stream, a blind fish and another species of
the family with well developed eyes.

If it is by acceleration and retardation of characters that the
Heteropygii have been developed from the Cyprinodontes, we have
indeed a most startling and sudden change of the nervous s}^stem.
In all fishes the fifth pair of nerves send branches to the various
parts of the head, but in the blind fishes these branches are devel-
' oped in a most wonderful manner, while their subdivisions take
new courses and are brought through the skin, and their free ends
become protected by fleshy papillae, so as to answer, by their deli-
cate sense of touch, for the absence of sight. At the same time
the principle of retardation must have been at work and checked
the development of the optic nerve and the eye, while accelera-
tion has caused other portions of the head to grow and cover over
the retarded eye.

Now, if this was the mode by which blindness was brought about
and tactile sense substituted, why is it that we still have Cholo-
gaster Agassizii in the same waters, living under the same condi-
tions, but with no signs of any such change in its senses of sight
and touch ? It may be said that the Chologaster did not change
because it probably had a chance to swim in open waters and
therefore the eyes were of use and did not become atrophied.
We can only answer, that if the Chologaster had a chance for
open water, so did the Typhlichtlrys and yet that is blind.

If the Heteropygii have been developed from Cyprinodontes,
how can we account for the whole intestinal canal becoming so
singularly modified, and what is there in the difference of food or
of life that would bring about the change in the intestine, stomach
and pyloric appendages, existing between Chologaster and Typh-
lichthys in the same waters ? 'To assume, that under the same con-
ditions, one fish will change in all these parts and another remain
intact, by the blind action of uncontrolled natural laws, is, to me,
an assumption at variation with facts as I understand them.

ORIGIN OF THE BLIND FISHES. 51

Looking at the case from the standpoint which the facts force
me to take, it seems to me far more in accordance with the laws of
nature, as I interpret them, to go back to the time when the
region now occupied by the subterranean streams, was a salt and
brackish water estuary, inhabited by marine forms, including the
brackish water forms of the Cyprinodontes and their allies (but not
descendants) the Heteropygii. The families and genera having the
characters they now exhibit, but most likely more numerously rep-
resented than now, as many probably became exterminated as the
salt waters of the basin gradually became brackish and more lim-
ited, as the bottom of this basin was gradually elevated, and
finalty, as the waters became confined to still narrower limits and
changed from salt to brackish and from brackish to fresh, only
such species would continue as could survive the change, and they
were of the minnow type represented by the Heteropygii, and per-
haps some other genera of brackish water forms that have not
yet been discovered.

In support of this hypothesis we have one species of the family,
Chologaster cornutus, now living in the ditches of the rice fields of
South Carolina, under very similar conditions to those under which
others of the family may have lived in long preceding geological
times ; and to prove that the development of the family was not
brought about by the subterranean conditions under which some
of the species now live, we have the one with eyes living with the
one without, and the South Carolina species to show that a sub-
terranean life is not essential to the development of the singular
characters which the family possess.

That a salt or brackish water fish would be most likely to be
the kind that would continue to exist in the subterranean streams,
is probable from the fact that in all limestone formations caves
are quite common, and would in most instances be occupied first
with salt water and then brackish, and finally with fresh water so
thoroughly impregnated with lime as to render it probable that
brackish water species might easily adapt themselves to the
change, while a pure fresh water species might not relish the solu-
tion of lime any more than the solution of salt, and we know how
few fishes there are that can live for even an hour on beino-

Of

changed from fresh to salt, or salt to fresh, water. We have also
the case of the Cuban blind fishes belonging to genera with their
nearest representative in the family a marine form, and with the

52 NOTE ON THE YOUNG BLIND FISH.

whole family of cods and their allies, to which group they belong,
essentially marine. Further than this the cat fish from the subter-
ranean stream in Pennsylvania belongs to a family having many
marine and brackish water representatives. As another very in-
teresting fact in favor of the theory that the Heteropygii were
formerly of brackish water, we have the important discovery by
Prof. Cope of the Lernsean parasite on a specimen of Amblyopsis
from the Wyandotte cave ; this genus of parasitic crustaceans be-
ing very common on marine and migratory fishes, and much less
abundant on fresh water species.

Thus I think that we have as good reasons for the belief in
the immutability and early origin of the species of the family of
Heteropygii, as we have for their mutability and late development,
and, to one of my, perhaps, too deeply rooted ideas, a far more
satisfactory theory ; for, with our present knowledge, it is but the-
ory on either side.

YOUNG OF THE BLIND FISH. — Dr. Hagen gives me the follow-
ing information about the young specimens I mentioned (page 38)
as belonging to Dr. Steindachner, which I just missed seeing
before they were sent to Vienna. These specimens were procured
by Dr. Hartung for Dr. Steindachner under the following circum-
stances. Just as Dr. Hartung was leaving the cave hotel on Oct.
21, a bottle was brought to him containing four specimens, one of
which was smaller than the others (probably Typhlichthys) , all
living. He immediately transferred them to a jar containing alco-
hol and took no notice of them until he reached Nashville, when
he discovered an addition of eight little ones in the jar.

The birth of these young was undoubtedly due to placing the
parent in the alcohol, and the date (Oct. 21) would correspond to
the time I have stated as probably that at which the young were
born.

Dr. Hagen states that he examined the young under a lens, with-
out taking them from the jar, and could not discover any eyes. The
specimens were about three lines in length.

So now we have two more facts to add to the history of the
blind fishes (though whether they apply to Amblyopsis or Typh-
lichthys is not yet settled). First, that the young are born in
October, and second, that they are without external eyes when
born. — From the AMERICAN NATURALIST for February, 1872.

\ A

I

G.A.WALKER- sc.

EXPLANATION OF PLATE ONE.

[All the figures on this plate are from original drawings by Prof. J. Wyman.]

FIG. 1. Brain, nerves and organ of hearing of Amblyopsis spelaus; enlarged; a, olfac
tory lobes and nerves; 6, cerebral lobes; c, optic lobes; d, cerebellum; «,
organ of hearing, showing the semicircular canals, with the otolite repre-
sented in place by the dotted lines; /, medulla oblongata; g, optic nerves
and eye specks.

FIG. 2. Otolite, enlarged.

FIG. 3. Eye, magnified (natural size one-sixteenth of an inch in length) ; a, optic nerve ;
6, sclerotic membrane ; c, layer of colorless cells ; d, layer of pigment cells
(iris?); e, lens.

FIG. 4. Lens, enlarged and showing the cells.

FIG. 5. Eye, enlarged, showing the muscular bands, a, a, a, a; b, the lens pressed out
of place; c, the optic nerve.

FIG. 6. Top of head, showing the canals under the skin, of the natural size. The two
black dots and lines indicate the eyes and optic nerves in position.

FIG. 7. Top of head, showing the arrangement of the ridges of papillae. Natural size.
FIG. 8. One of the ridges of papilla? fr*om the head, magnified.

FIG. 9. Three of the papilla? from the ridge, still more magnified, showing the cup-
shaped summit and projecting filament.

FIG. 10. A portion of the ridge magnified, and treated with acid, to show the arrange-
ment of the nervous plexus supplying the papillae with nerve filaments from
a branch (a) of the fifth pair.

FIG. 11. Epithelial cells from the head.
FIG. 12. Epithelial cells from the body.
FIG. 13. A fish with eyes, found in the stomach of an Amblyopsis.

(53)

EXPLANATION OF PLATE TWO.

FIG. 1. AMBLYOPSIS SPEL^EUS DeKay. Natural size.

la. Stomach and pyloric appendages. Twice natural size.

1 &. Scale, magnified.

Ic. Abdominal cavity, showing position of stomach and single ovary. Natural
size.

FIG. 2. CHOLOGASTER CORNUTUS Agassiz. Natural size.

2 a. Stomach and pyloric appendages. Twice natural size.

2 &. Scale, magnified.

2c. Abdominal cavity, showing stomach and single ovary behind the stomach.
Twice natural size.

FIG. 3. TYPHLICHTHYS SUBTERRANETJS Girard. Slightly more than natural size.

3a. Stomach and pyloric appendages. Twice natural size.

3 &. Scale, magnified.

FIG. 4. fnoT nc \ STFT? AGASSIZII Putnam. Natural size.

4 a. Stomach and pyloric appendages. Twice natural size.
4&. Scale, magnified.

The scales figured on the plate are all from the second or third row under the dorsal
fin. 4 b is represented with the posterior margin dotvn, all the others are represented
with the posterior margin on the left. The natural size of the scales is given by the
minute outline at the left of the figures above each scale ; 4 & is so small that the natural
size can hardly be represented by the black dot.

(54)

PI 2

TJHIVERSITT

CHAPTER IV.
#t*«

SYNOPSIS OF THE FAMILY HfiT-E»e£¥6«:

BY F. "W. PUTNAM.

HETEROPYGII TELLKAMPF, Muller's Arch. f. Anat., p. 392,
1844; and New York Journal of Medicine, v, p. 84, 1845.

Hypsczidce Storer, Synopsis N. A. Fish, p. 435, 1846.

Brain of ordinary development in all its parts, similar to that of
Cyprinodontes and of about the same proportions. Cerebral lobes
larger than the nearly round optic lobes. Cerebellum overlapping the
posterior third of the optic lobes. Medulla oblongata broad, with
well defined right and left sides. (On comparing the brains of the
three genera the only difference noticed \vas that in Chologaster the
cerebellum was not quite as large proportionally, but more elongated
and not quite as wide as in the other genera, while the optic lobes of
this genus with well developed eyes were no larger than in a Typh-
lichthys of the same size.)

Skeleton not studied. Giinther gives the vertebrae as thirteen ab-
dominal and nineteen or twenty caudal. The bones of the head are
thin and mostly flattened as in the Cyprinodontes. Occiput slightly
convex.

Body compressed posteriorly. Head and anterior portion of body
depressed, giving the form of a broad, flat head, with a compressed
tail.

Brancliiosteyal rays six in number and but slightly covered by oper-
cular bones ; opercular opening large.

Fins. Dorsal and anal nearly opposite and posterior to centre of
body. All the fins except the ventrals well developed, with central
rays longest and first rays simple. Pectorals close to the head, about
in the middle of the sides. (Ventrals present in Amblyopsis, absent
in Typhlichthys and Chologaster.)

Mouth opening upwards, with lower jaw slightly projecting. Mar-
gin of the upper jaw formed by the intermaxillaries. Maxillaries
placed behind the intermaxillaries, with lower third broad and below
the intermaxillaries. Several rows of fine teeth on the intermaxil-
laries and lower jaw. (Teeth on palatines in Amblyopsis and Typh-
lichthys, none on these bones in adults of Chologaster.)

Scales. None on the head. Body closely covered with small, par-
tially imbedded cycloid scales, irregularly arranged.

Lateral line absent.

* From the Annual Report of the Peabocly Academy of Science for 1871.

(55)

56 SYNOPSIS OF THE FAMILY HETEROPYGII.

Nostrils double. Anterior tubular and standing out from the end of
the snout.

Stomach well defined, coecal.

Pytoric appendages present.

Intestine with two turns.

Anus situated under the throat and forward of the pectorals.

Ovary single. (Placed by the side of the stomach in Amblyopsis
and Typhlichthys and behind it in Chologaster.)

Viviparous. (Amblyopsis.)

Testes paired. (Amblyopsis.)

Air bladder with pneumatic duct. (Amblyopsis.)

Liver with the left lobe very large and partially enclosing the
stomach.

Amblyopsis DEK.AY, Fishes of New York, p. 187, 1842.

Eyes rudimentary and imbedded under the skin.

Head with numerous transverse and longitudinal rows of sensitive
papilla3 provided with nerve branches, many of the nerve branches
terminating as free filaments outside the papilla3. Small granula-
tions on tlie spaces between the papillary ridges. Canals under the
skin.

Teeth minute, curved, and arranged in rows on the intermaxillary,
inferior maxillary and palatine bones.

Body with a prominent papilla just over the opercular opening, at
the base of a small papillary ridge, similar to those on the head.
Papillary ridges on sides of body of same character as those on the
head, and arranged at nearly equal distances from opercular opening
to base of caudal fin.

Pyloric appendages, one on each side.

Ovary situated on the right side of the stomach.

Fins. Ventrals small and placed near the anal fin. Dorsal, 9.
Anal, 9. Pectoral, 11. Ventral, 4. Caudal, 24.

Amblyopsis spelseus DEIYAY. LAEGE BLINDFISH.

CRAIGE, Procd. Acad. Nat. Sci. Philad., i, p. 175, 1842. DEKAY, Fishes N. Y., p.
187, 1842. WYMAN, Amer. Jour. Sci., xlv, p. 94, 1843; Ann. Mag. Nat. Hist., xii, p.
298., 1843. THOMPSON, Ann. Mag. Nat. Hist., xiii, p. Ill, 1844. TELLKAMPF, M til-
ler's Arch. f. Anat., p. 392,1844; N. Y. Jour. Medicine, v, p. 84, with plate, giving
three figs, of the fish ; position of internal organs ; brain ; stomach ; air bladder; scale
(profile view gives the fish without ventral fins, but ventral view shows them), 1845.
STOKER, Synopsis N. A. Fish, p. 435, 1846. OWEN, Lect. Comp. Anat. Fishes, pp.
175, 202 (fig. of brain), 1840. WYMAN, Procd. Boston Soc. Nat. Hist., iii, p. 349, 1850
"DALTON, N. Y. Medical Times, ii, p. 354, 18—." AGASSIZ, Amer. Jour. Sci. xi. p.128,
1851. WYMAN, Procd. Boston Soc. Nat. Hist., iv, p. 395 (1853), 1854; v, p. 18, 1854;
Amer. Jour. Sci., xvii, p. 259, 1854 (with figs, of brain, eye, and otolite). GIRAKD,
Proc. Nat. Sci. Philad., p. 63, 1859. POEY, Mem. de Cuba, ii, p. 101, Pis. 9, 11
(outlines of fish and of brain), 1858. WOOD, 111. Nat. Hist., iii, p. 314, figure.
1862. TENNEY, Nat. Hist., p. 344, figure. 1865. GttNTHER, Cat. Fish Brit. Museum,
vii, p. 2, 1868. COPE, Ann. Mag. Nat. Hist., viii. p. 368, 1871. PUTNAM, Amer. Nat.,

SYNOPSIS OF THE FAMILY HETEROPYGII. 57

vi, p. 6 et seq., \vith figs.., Jan., 1872. WYMAN, Mss. notes and drawings in Put-
nam, Amer. Nat., vi, p. 16 et seq., 1872. PUTNAM, Amer. Nat., vi. p. 116. Feb.,
1872 (additional note on the young).

PLATE 1 (American Naturalist, Vol. vi, Jan., 1872). FIG. 1. Brain, nerves and
organ of hearing of Amblyopsis spelceus; enlarged; a, olfactory lobes and nerves;
6, cerebral lobes; c, optic lobes; d, cerebellum; e, organ of hearing, showing the
semicircular canals, with the otolite represented in place by the dotted lines ; /,
medulla oblongata ; g, optic nerves and eye specks. FIG. 2. Otolite, enlarged.
FIG. 3. Eye, magnified (natural size one-sixteenth of an inch in length); a, optic
nerve; 6, sclerotic membrane; c, layer of colorless cells; d, layer of pigment cells
(iris?); e, lens. FIG. 4. Lens, enlarged and showing the cells. FIG. 5. Eye, en-
larged, showing the muscular bands, a, a, a, a; b, the lens pressed out of place; c,
the optic nerve. FIG. 6. Top of head, showing canals under the skin, natural
size. The two black dots and lines indicate the eyes and optic nerves in position.
FIG. 7. Top of head, showing the arrangement of the ridges of papillae, nat. size.
FIG. 8. One of the ridges of papilla? from the head, magnified. FIG. 9. Three of
the papillae from the ridge, still more magnified, showing the cup-shaped summit
and projecting filament. Fig. 10. A portion of the ridge magnified, and treated
with acid, to show the arrangement of the nervous plexus supplying the papillae
with nervous filaments from a branch (a) of the fifth pair. FIG. 11 . Epithelial cells
from the head. FIG. 12. Epithelial cells from the body.

PLATE 2. FIG. 1. Natural size; la, stomach and pyloric appendages, twice nat.
size; 16, scale, magnified (nat. size represented by the small outline on the left
over the figure); Ic, abdominal cavity, showing position of stomach and single
ovary, nat. size.

Head more than half as wide as it is long. Length of head, from
tip of jaw to end of operculum, contained nearly twice in length of
body from operculum to base of caudal fin.

Dorsal and anal fins of equal size, rounded, anal commences under
third ray of dorsal.

Pectorals pointed, reaching to commencement of dorsal.

Ventrals pointed, nearly reaching to commencement of anal.

Caudal broad, long and pointed, membrane, enclosing simple rays
above and below, continuing slightly on the tail.

Scales small, longer tljaii broad, with quadrangular centre and from
8 to 12 concentric lines, which are broken and reduced in number an-
teriorly and crossed by numerous radiating furrows posteriorly.*

Colorless, or nearly so, with transparent fins.

Measurements. Largest specimen, 4-5 inches total length. Smallest
specimen, 1-9 total length.

Geographical distribution. Subterranean streams in Kentucky and
Indiana.

Specimens examined : —

PROF. WYMAN'S COLLECTION.

7 specimens. Half grown and adults. Mammoth Cave.
MUSEUM OP COMPARATIVE ZOOLOGY.

7 specimens. No. 778. Half grown and tf $ adults. Mammoth Cave.
1 specimen. No. — . Two-thirds grown. Cave near Lost River, Orange Co., Ind.

*T,The scales described were in every instance taken from the 2d or 3d row un-
der the dorsal fin.

58 SYNOPSIS OF THE FAMILY HETEROPYGII.

BOSTON SOCIETY or NATURAL HISTORY.
2 specimens. No. 840. Half grown. Mammoth Cave.

PEABODY ACADEMY or SCIENCE.

1 specimen. No. 520. Adult 9 . Mammoth Cave. Presented to Essex Insti-
tute in 1851 by N. Silsbee.

Other specimens. Dr. Glintlier mentions six specimens and a skele-
ton in the British Museum. Mr. Thompson, an adult and newly born
young in the collection of the Natural History Society of Belfast.
Dr. Steindachncr has recently sent an adult and eight young to
the Vienna Museum. The first specimen of which we have any
record was presented to the Academy of Natural Sciences of Phila-
delphia ; the second is the one described by DeKay and then in the
Lyceum of Natural History of New York. Prof. Cope obtained three
specimens from the waters of Wyandotte Cave in Indiana. Dr. Tell-
kampf had several specimens from the Mammoth Cave, and it is prob-
able that specimens exist in nearly all the principal museums and in
many private collections, as about all that have been caught in the
Mammoth Cave for years have been sold by the guides to visitors.

Habits. But little is known of the habits of the large bliudfish.
Dr. Tellkampf states that they are solitary ; on the slightest motion
of the water they dart off a short distance, and that they are mostly
found near stones or rocks on the bottom, and seldom come to the
surface of the water. Prof. Cope states that if they are not alarmed
they come to the surface to feed, swim in full sight, and can then be
easily captured if perfect silence is preserved. He also thinks that
they are principally surface feeders.

In the stomachs of several that I have opened the only remains
found were those of Crayfish. In one specimen, opened by Dr. Wy-
man, a small fish with well developed eyes was found in the stomach.
(See Amer. Nat., vi, p. 13, PI. 1, fig. 13.)

The eggs are well developed in September, and the young are born
about the middle to last of October. The young when born are half
an inch or less in length, and are icithout external eyes. (See Amer.
Nat., Feb., 1872. The young there mentioned may possibly be those
of Typhlichthys.)

Typhlichthys GIRARD, Procd. Acad. Nat. Sci. Philad., p. C3, 1859.

Eyes rudimentary and imbedded under the skin.

Head. The same arrangement of rows of sensitive papillae as in
Amblyopsis, and the spaces between the papillae with granulations as
in that genus. (The subcutaneous canals probably exist, but have
not yet been made out.)

Teeth, as in Amblyopsis, on the maxillaries and palatines.

Body with papilla over opercular opening, and with the papillary
ridges on the sides as in Amblyopsis.

Pyloric appendages one on each side as in Amblyopsis, but of

SYNOPSIS OF THE FAMILY HETEROPYGII. 59

slightly different proportion and shape. (Stomach not so pointed
behind as in Amblyopsis.)

Ovary situated on right side of stomach, as in Amblyopsis. (Eggs
fewer in number and proportionately larger than in Amblyopsis.)

Fins. Ventrals absent. Dorsal, 7 or 8; Anal, 7 or 8 ; Pectoral, 12;
Caudal 24. (This formula is given after counting several specimens.
Girard gives, D. 7; A. 8; P. 11; 0.23.)

It will be noticed that the only characters separating this genus
from Amblyopsis are the absence of ventral fins, the shape of the
stomach and pyloric appendages, and larger eggs in less number.

Typhlichthys subterraneus GIKAKD. SMALL BLINDFISH.

GIRARD, Procd. Acad. Nat. Sci. Philad., p. 63, 1859. GUNTHER, Cat. Fish Brit.
Museum, vii, p. 2, 1868 (as a syn. of Amblyopsis). PUTXAM, Amer. Nat., vi, p. 20
et seq., with figs., Jan.., 1872.

PLATE 2 (Amer. Nat., Vol. vi., Jan., 1872). FIG. 3, slightly more than natural size;
3a, stomach and pyloric appendages, twice nat. size ; 36, scale, magnified (nat.
size represented by small outline over the figure).

Proportions and general appearance, want of color, arrangement of
papillary ridges, position and shape of fins as in Amblyopsis spela3us,
with the exception that, owing to the jaws being more obtusely round-
ed, the head is slightly blunter and broader forward.

Membrane of caudal quite prominent and extending forwards to pos-
terior base of dorsal and anal fins.

Scales broader than long. Large quadrangular centre with from 6
to 8 concentric lines reduced in number and broken up on anterior
margin. Posterior portion with numerous radiating furrows.

Measurements. Largest specimen, 1-85 inches in total length.
Smallest specimen, 1*45 inches in total length.

Geographical distribution. Subterranean streams in Kentucky, Ten-
nessee and Alabama.

Specimens examined : —

MUSEUM OF COMPARATIVE ZOOLOGY.

7 specimens. No. 780. c? $. Adults. Mammoth Cave. Collected and pre-
sented by Alpheus Hyatt, Sept., 1859.

1 specimen. No. 781. Moulton, Alabama. Presented by Thomas Peters.
1 specimen. No. 782. Lebanon, Tennessee. Presented by J. M. Safford.

Other specimens. Dr. Girard described the species from a specimen
in the Smithsonian Institution, taken from a well near Bowling Green,
Ky. Dr. Gunther mentions a specimen, in the British Museum, from
the Mammoth Cave.

Habits. Nothing is known concerning the habits of this fish. It is
evidently much rarer at the Mammoth Cave than the large species, to
judge from the small number in collections. The fact that Mr. Hyatt
obtained seven specimens when he was at the cave in September and
did not get any of the other species, may indicate some peculiar loca-

60 SYNOPSIS OF THE FAMILY HETEROPYGII.

tion in the waters of the cave where it is more abundant than in other
places. The eggs were fully developed in these specimens, but no
embryos could be detected. The fish is probably viviparous, and
very likely gives birth to its young in October.

Chologaster AGASSIZ, Amer. Jour. Sci., xvi, p. 135, 1853.

Eyes in normal position and well developed.

Head with small granulations on the surface of the skin. (No
papillary ridges.)

Teeth minute, curved and arranged in rows on the intermaxillary
and inferior maxillary bones. None on the palatines in the adults.
(Of the four specimens examined, the two larger (C. cornutus) are
without palatine teeth, while the single specimen of C. Agassizii,
which is evidently a young fish, has a few minute teeth on the pala-
tine bones. In the smallest specimen of C. cornutus the mouth is
abnormal, the intermaxillaries being reduced to a small central portion
and there are consequently no teeth in the upper jaw, but the minute
teeth on the palatines are present.*)

{Body without opercular papilla and papillary ridges on the sides.)

Pyloric appendages two on each side. Stomach rounded and turned
slightly on the side.

Ovary situated principally behind the stomach.

Fins. Ventrals absent. Dorsal, 8 or 9. Anal, 8 or 9. Pectoral, 12.
Caudal, 28.

This genus principally differs from Amblyopsis and TyphlichthjTs by
the presence of eyes, the absence of papillary ridges on the head and
body, by having two pyloric appendages on each side instead of
one, and by the posterior position of the ovary. It agrees with Typh-
lichthys in the absence of the ventrals, and the young further agree
by the presence of palatine teeth.

*I believe this is one of those interesting cases where one set of organs, or one
portion of the animal structure, takes the place of another which from accident is
wanting, and that in all probability these palatine teeth, that under normal con-
ditions wrould be cast off as the fish attained maturity, would have continued to
exist in this specimen and answer all the purposes of the intermaxillary teeth.
But that in this accidental continuance of these palatine teeth, from the mere
mechanical use forced upon them, we have the first stages of the development of
a distinct genus, to be characterized by permanent teeth on the palatines, and
reduced upper jaw bones, as many of the developmental school would argue, I do
not think will bear the test of facts observed.

A not uncommon malformation of fishes consists in the entire or partial absence
of the maxillary or intermaxillary bones. I have specially noticed this among our
common fresh water trout (Salmo) and marine Conner or sea perch (Ctenolabrus)
but there have never been recorded allied genera with these characters, while
the malformed specimens are hardly numerous enough to give support to the the-
ory that such malformations are hereditary, and it is probable that each case was
caused by the non-development of the parts from special cause during embryonic
life, or by accident to the individual.

SYNOPSIS OF THE FAMILY HETEROPYGII. 61

Chologaster cornutus AGASSIZ.

AGASSIZ, Amer. Jour. Sci., xvi, p. 135, 1853. GIRARD, Procd. Acacl. Nat. Sci.
Philad., p. 63, 1859. GUNTHER, Cat. Fish. Brit. Museum, vii, p. 2, 1868. PUTNAM,
Amer. Nat., vi, p. 21 et seq., with figs. Jan., 1872.

PLATE 2 (Amer. Nat., Vol. vi, Jan., 1872). FIG. 2. Natural size. 2a, stomach
and pyloric appendages, twice nat. size. 26, scale magnified (nat. size represented
by small outline over the left of the fig). 2c, abdominal cavity showing stomach
and single ovary behind the stomach, twice nat. size.

Head more than half as wide as it is long. Length, of head, from
tip of under jaw to end of operculum contained twice in length of
body from operculum, to caudal fin. Width between the eyes equal to
distance from eye to tip of under jaw.

Eyes of moderate size, situated just back and over the end of the
maxillaries.

Dorsal and anal fins of nearly equal size, slightly rounded. Anal
with slightly longer rays and commences under fourth ray of dorsal.

Pectoral fins pointed, reaching to line of commencement of dorsal.

Caudal fin pointed, about equal in length to the head. Membrane
above and below extending but slightly on the tail.

Scales very small and deeply imbedded in the skin. Circular with
small smooth space forward of the centre. From 15 to 20 con-
centric rings, cut by a few short radiating furrows on anterior, and
longer and more numerous ones on posterior margin.

Intestine is a little longer than in an Amblyopsis of the same size.
The two pyloric appendages on the left side are close together and
broader than the two on the right side, which are wider apart,
longer and more slender than the others.

Color. Yellowish brown, much darker above, lighter on sides, and
light yellow on under part and sides of head, belly and under part of
tail. Three longitudinal very dark brown lines on each side: the
upper commencing near the middle of top of head and following
along the back to base of caudal fin ; the middle one commencing at
the nostril and passing through the eye to upper portion of opercu-
lum, thence about in the centre of side to about the centre of base of
caudal fin ; the lower commences under the pectoral fin and follows
the ventral curve of the body to the base of caudal fin. All three
lines are darkest and broadest forward, and terminate as a series
of nearly confluent dots on the tail. Central rays of the caudal dark
brown, outer rays uncolored. Dorsal, anal and pectorals not colored.

Measurements. The three specimens are respectively 1-5, 2, and 2-3
inches in total length.

Geographical distribution. South Carolina.

Specimens examined: —

MUSEUM or COMPARATIVE ZOOLOGY.

3 specimens. No. 776. Rice Ditches at Waccamaw, S. C. Presented by P. C.
J. Weston, 1853. (Orig. of Agassiz.)

62 SYNOPSIS OF THE FAMILY HETEROPYGII.

Habits. Nothing is known concerning the habits of this species,
the only specimens observed being the three mentioned. From the
fact of its having a single ovary containing a small number (about GO)
of large eggs it is probable that it is viviparous.

Chologaster Agassizii PUTNAM.

PUTXAM, Amer. Nat., vi, p, 22 et seq., with figs. Jan., 1872.

PLATE 1 (Amer. Nat., Vol. vi, Jan., 1872). FIG. 4. Natural size; 4a, stomach
andpyloric appendages, twice nat. size; 46, scale magnified (nat. size shown lay
minute dot over left of the figure).

Head more than half as wide as it is long. Its length is contained
three times in the length of the body from the operculum to the base
of caudal fin.

Eyes proportionately large and placed over ends of maxillaries.

Dorsal and anal fins broken, but probably of about equal size. Anal
fin commences about under fourth ray of dorsal.

Pectoral fins pointed and reaching about half way to the dorsal.

Caudal fin pointed, not quite as long as the head.

Scales very minute, longer than wide, with 4 or 5 concentric
lines round a granulated centre. A few radiating furrows cut the
concentric lines on the posterior margin.

Pyloric appendages and stomach about the same as in C. cornutus.

Color. Uniform light brown, without markings except that the base
of the caudal fin is rather darker than rest of fish. Fins uncolored.

Measurements. Total length, 1-4 inches.

Geographical distribution. Subterranean streams in Tennessee.

Specimen examined: —

MUSEUM OF COMPARATIVE ZOOLOGY.

1 specimen. No. 777. From a well in Lebanon, Tenn. Presented by J. M.
Safford. Jan., 1854..

This species principally differs from C. cornutus by having a longer
body and smaller head, by having the eyes proportionately larger, and
by its coloration. Nothing is known of its habits except the fact
of its subterranean life. The scales of the single specimen known
indicate a young fish, and it is probably not over half grown.

The four species given in this synopsis are all of the family
as yet known, but that others will be discovered and the range of the
present known species extended is very probable. The ditches and
small streams of the lowlands of our southern coast will undoubtedly
be found to be the home of numerous individuals, and perhaps of new
species and genera, while the subterranean streams of the central
portion of our country most likely contain other species.

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