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ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
VOLUME 42 • 1957 • NUMBERS 1 TO 14
PUBLISHED BY THE SOCIETY
The ZOOLOGICAL PARK, York
NEW YORK ZOOLOGICAL SOCIETY
GENERAL OFFICE
30 East Fortieth Street, New York 16, N. Y.
PUBLICATION OFFICE
The Zoological Park, New York 60, N. Y.
OFFICERS
PRESIDENT VICE-PRESIDENTS SECRETARY TREASURER
Fairfield Osborn Alfred Ely Harold J. O’Conaell David H. McAlpin
Laurance S. Rockefeller
SCIENTIFIC STAFF: Zoological Park and Aquarium
John Tee- Van General Director
ZOOLOGICAL PARK
Leonard J. Goss Assistant Director
and Veterinarian
John L. George Associate Curator
of Mammals
William G. Conway. .Associate Curator
of Birds
Grace Davall Assistant Curator,
Mammals and Birds
James A. Oliver Curator of Reptiles
Charles P. Gandal. . .Associate Veterinarian
Lee S. Crandall General Curator
Emeritus
William Beebe Honorary Curator,
Birds
AQUARIUM
Christopher W. Coates . Director
James W. Atz Associate Curator
Carleton Ray Assistant to the
Director
Ross F. Nigrelli Pathologist
Myron Gordon Geneticist
C. M. Breder, Jr Research Associate
in Ichthyology
Harry A. Charipper. . .Research Associate
in Histology
Homer W. Smith Research Associate
in Physiology
GENERAL
William Bridges Editor & Curator,
Publications
Sam Dunton Photographer
Henry M. Lester . . . Photographic Consultant
DEPARTMENT OF TROPICAL
RESEARCH
William Beebe Director Emeritus
Jocelyn Crane Assistant Director
David W. Snow Resident Naturalist
Henry Fleming Entomologist
John Tee-Van Associate
William K. Gregory Associate
AFFILIATES
C. R. Carpenter Co-ordinator, Animal
Behavior Research Programs
L. Floyd Clarke Director,
Jackson Hole Research Station
SCIENTIFIC ADVISORY COUNCIL
A. Raymond Dochez Carjd P. Haskins
Alfred E. Emerson K. S. Lashley
W. A, Hagan John S. Nicholas
EDITORIAL COMMITTEE
Fairfield Osborn, Chairman
James W. Atz Lee S. Crandall
William Beebe Leonard J. Goss
William Bridges James A. Oliver
Christopher W. Coates John Tee-Van
William G. Conway
Contents
3993r
SI
Part 1. May 20, 1957
PAGE
1. Miniature Circulating Systems for Small Laboratory Aquaria. By C. M.
Breder, Jr. Plates I & II; Text-figures 1-5 1
2. The Behavior of the Bottle-nosed Dolphin (Tur stops truncatus): Mating,
pregnancy, parturition and mother-infant behavior. By Margaret C.
Tavolga & Frank S. Essapian. Plates I-III; Text-figure 1 11
3. A Study of the Relationship Between Certain Internal and External Mor-
phological Changes Occurring During Induced and Natural Metamorphosis
in Rana pipiens and Rana catesbeiana. By Beulah Howatt McGovern &
Harry A. Charipper. Text-figures 1-12 33
Part 2. August 23, 1957
4. On the Bramid Fishes of the Gulf of Mexico. By Giles W. Mead. Plates
I-III 51
5. The Relation of Oxygen Consumption to Temperature in Some Tropical,
Temperate and Boreal Anuran Amphibians. By Richard E. Tashian &
Carleton Ray. Text-figures 1 & 2 63
6. Basic Patterns of Display in Fiddler Crabs (Ocypodidae, Genus Uca).
By Jocelyn Crane. Plate I; Text-figures 1-4 69
JAN 1 5 1958
Part 3. November 25, 1957
PAGE
7. Studies on the Lizard Family Xantusiidae. III. A New Genus for Xantusia
riversiana Cope, 1883. By Jay M. Savage. Text-figures 1-3 83
8. Nesting Behavior of the Crested Oropendola {Psarocolius decumanus) in
Northern Trinidad, B.W.I. By Richard E. Tashian. Plates I & II; Text-
figures 1-3 87
9. Changes in the Cytological Structure of the Adenohypophysis and Gonads
in Juvenile Bathygobius soporator after Pituitary Implantation. By Louise
M. Stoll. Plates I & II 99
10. The Ctenuchidae (Moths) of Trinidad, B.W.I. Part I. Euchromiinae. By
Henry Fleming. Plates I-III 105
Part 4. December 31, 1957
11. The Abifity of the Saprolegniaceae to Parasitize Platyfish. By Helen S.
Vishniac & R. F. Nigrelli. Plate 1 131
12. Imaginal Behavior in Butterflies of the Family Heliconiidae: Changing
Social Patterns and Irrelevant Actions. By Jocelyn Crane. Plate I.. . . 135
1 3 . Habits, Palatability and Mimicry in Thirteen Ctenuchid Moth Species from
Trinidad, B.W.I. By William Beebe & Rosemary Kenedy. Plates I & II. 147
14. Serological Relationships among Members of the Order Carnivora. By
Ludwig K. Pauly & Harold R. Wolfe 159
Index to Volume 42
167
ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
VOLUME 42 • PART 1 • MAY 20, 1957 • NUMBERS 1 TO 3
PUBLISHED BY THE SOCIETY
The ZOOLOGICAL PARK, New York
Contents
PAGE
1. Miniature Circulating Systems for Small Laboratory Aquaria. By C. M.
Breder, Jr. Plates I & II; Text-figures 1-5 1
2. The Behavior of the Bottle-nosed Dolphin (Tursiops truncatus): Mating,
pregnancy, parturition and mother-infant behavior. By Margaret C.
Tavolga & Frank S. Essapian. Plates I-III; Text-figure 1 11
3. A Study of the Relationship Between Certain Internal and External Mor-
phological Changes Occurring During Induced and Natural Metamorphosis
in Rana pipiens and Rana catesbeiana. By Beulah Howatt McGovern &
Harry A. Charipper. Text-figures 1-12 33
1
Miniature Circulating Systems for Small Laboratory Aquaria
C. M. Breder, Jr.
The American Museum of Natural History, New York 24, N. Y.
(Plates I & II; Text-figures 1-5)
Introduction
Because of the requirements of certain
experiments it became necessary to es-
tablish various small, but fully control-
lable circulating systems in small aquaria. These
have included both open and closed freshwater
systems and closed saltwater systems. As the
designs eventually worked out have proved to
be entirely satisfactory, and as many colleagues
have inquired about these systems, with a view
to building similar ones for their own purposes,
the details of construction and operation are
explained here.
Primarily these systems are the outgrowth of
work of earlier years at the old New York
Aquarium where much larger, but similar,
equipment formed the basis of operations. This
equipment itself had been developed from
schemes used by older institutions of similar
kind. Naturally, many persons had a hand in
developing the arrangements and devices em-
ployed at the New York Aquarium. For these
reasons the origins of the devices were not al-
ways clear, but those chiefly interested and re-
sponsible for them at the Aquarium were C. W.
Coates and the late C. H. Townsend, and
H. Knowles. Townsend (1928) and Breder &
Howley (1931) reported on some of these fea-
tures. It has been found that by suitable modifi-
cation of the principles of the larger devices it
is possible to develop very useful miniature
equipment. Such need, of course, applies only to
laboratories not connected with large public
aquariums and which consequently lack the
utilities usually only to be found in such places.
These devices have been worked out in connec-
tion with experimental work carried on in the
laboratories of the Department of Fishes and
Aquatic Biology of the American Museum of
Natural History, which has been supported in
part by the National Science Foundation.
Open Systems
An “open system” as here used refers to one
in which the water is used but once and not re-
circulated. That is, there is only a supply line
and a drain line. This calls for little comment in
present connections except where a very small,
well-regulated flow is required. Such apparatus
may be arranged to provide as little as a speci-
fied number of drops a minute, and wUl main-
tain a surprising accuracy if properly designed.
The overflow provided for this system is a
constant-level syphon. If such a syphon is made
by a glass blower it is expensive, subject to
breakage and is not readily cleaned. Syphons
can be quickly and cheaply made of some
straight glass tubing, a tee, some flexible rubber
or plastic tubing, and two small pieces of wood
or plastic strip. No dimensions are given, as
these will vary with the individual needs, al-
though Text-fig. 1 is drawn to scale. The two
pieces of plastic are identical and should have
two holes drilled in them that will snugly fit the
glass tubing. These parts are then assembled to
make a constant-level syphon attached to the
lip of an aquarium as shown in Text-fig. 1 and
PI. I, Fig. 1. The lower piece of plastic may be
fastened to the aquarium by small clamps or be
cemented to it. The open, upper end of the tee
vents the syphon. If a cap or plug is placed on
this it immediately becomes a simple syphon
and will drain the aquarium to the level of its
inlet tube. This is sometimes found to be an
added handy feature. The level of the water in
the aquarium will be that at which the overflow
water spills out through the horizontal leg of
the tee. Adjustments of this to a fine point may
be made by raising or lowering the tee through
the hole in the plastic support, or this whole
external assembly may be moved up or down by
altering the position of the straight length of
tubing which passes through the hole in the other
piece of plastic connecting it with the tube in-
1
2
Zoologica: New York Zoological Society
[42:1
Text-fig. 1. Side and end view of constant-level syphon made up of standard parts.
side the aquarium. By arranging the outside
part of the syphon to lie along the aquarium
wall, as shown, the danger of its being in the
way of operations is reduced. It is obvious that
cleaning presents no problem with this type of
syphon. In most cases a plastic or glass strainer
is placed over the intake end of the syphon. If
something does nevertheless block the syphon
tube from the aquarium, it almost always can be
cleared by blowing into the open end of the tee
and restarting the syphon by drawing on this
same open end of the tee while the outlet tube
is held shut. It is possible and sometimes more
convenient to attach the outer portion of the
constant-level syphon to a small board which is
affixed to a pivot at its upper end so that it is
free to rotate on the stationary part attached to
the aquarium frame. A small handle pointing
upward from there makes its adjustment simple
and marks on the latter in reference to some
stationary part make return to a former rate of
flow exactly possible. The action is simply that
by rotating the part of the syphon so that the
horizontal part of the tee raises or lowers, the
level in the aquarium follows accordingly. This
in turn affects the float valve, which is described
below. The rate of flow will increase if the
syphon outlet is lowered and decrease if it is
raised. This is useful where the exact level of
water is not of any importance but where it is
desired to vary the amount of water flowing
through the aquarium by specific amounts and
where it is necessary to repeat such changes in
flow at will.
If the water supply has considerable pressure,
such as is ordinarily encountered in city water
systems, or approaches it, a pressure-reducing
valve which may be regulated should be em-
ployed. This can bring the pressure down to a
value which will not burst or otherwise destroy
the light equipment to be employed. This valve
placed someplace in the supply line should be
set so as to deliver little more than the maximum
amount of water which will be required of it.
Another way to accomplish the same purpose is
to permit the supply water to run into a small
reservoir of no more than sufficient height to
provide enough head of pressure. Into this reser-
voir the supply water is allowed to run continu-
ally, of a little more volume than the aquaria will
ever need. This is necessary to maintain a con-
stant head in the reservoir. A small excess will
overflow and go to waste by this method. It is
economic only where water saving has no sig-
nificance.
The water flowing into this aquarium is con-
trolled by a float-valve constructed of a glass
stop-cock, a suitable-sized chemical flask and
some small parts of either wood or plastic. These
are assembled as shown in Text-fig. 2 and held
together by iron screws and two pieces of strip
steel. This metal is mentioned because of the
danger of toxic salts forming if brass were used,
since such corrosion might fall into the aqua-
rium. A one-hole cork is bolted to a piece of
Incite and then inserted into the mouth of the
flask as shown. A dowel or plastic rod is inserted
in the other hole in the plastic piece and secured.
1957]
Breder: Miniature Circulating Systems for Small Laboratory Aquaria
3
Text-fig. 2. Top and side view of float-valve for control of inflow of water and additives.
A similar piece of plastic is movably secured on
this rod and on a similar one at right angles to
it, extending from the valve. This is so arranged
that the center of the flask comes to rest directly
below the horizontal rod extending from the
valve. This is best seen in the plan view of the
device. By loosening the two set screws in the
upper plastic piece the flask may be moved
vertically on the one and horizontally on the
other. PI. I, Fig. 2, shows one arrangement of
this device.
The extending glass tubes which are an in-
tegral part of the stop-cock are inserted through
two snug holes in the wood or plastic end-pieces
of the valve and these are held in position by
the two steel strips which are held in place by
four wood screws. In the center of one of these
steel pieces a hole is tapped into which is
screwed a set screw with a pointed end (about
60°) and a lock nut as shown. The stem handle
of the stop-cock is imbedded in a piece of wood
cut about as shown. For this purpose a suitable
space is hollowed out in the block into which the
stem is inserted, the space around being filled
with plastic wood or similar product. The face
of the block and the stop-cock stem must be at
right angles. On the outer face of the block a
small steel strip is affixed with a small drill-tip
impression at its center on the axis of the stem.
Into this the pointed set screw fits as shown. This
is adjusted so that the valve works freely with-
out being too tight or leaking. This prevents the
glass stop-cock from working loose and leak-
ing after a time of long-continued operation.
The diagonal dotted line on the block indicates
the position of the hole through the plug as well
as that of the handle on the stem. It is shown in
a position just fully closed. It is obvious that
with a fall in the water level the valve will open
proportionally to the change in water level and
shut itself off as the water level rises.
The interaction between the constant-level
syphon and this valve is indicated in Text-fig. 3.
It is clear that danger from flooding could come
only from some damage to the equipment. If,
for instance, something clogs the overflow in any
way, the float valve shuts itself off when it has
reached whatever predetermined point for which
it has been set.
As an extreme point of precaution a safety
alarm or shut-off could be built as an entirely
separate system. Such a device, which has never
failed so far as the writer’s experience goes, con-
sisted of an old pair of contacts such as are to
be found on relays to one member of which was
fastened a shell vial. This hung over the water
in such a manner that when the water rose over
a specified place it lifted the vial and pushed the
two contacts together. It operated on two dry
cells to ring a doorbell but could be used with a
relay to switch on house current to operate any
suitable device. This could be a normally open
solenoid valve placed in the supply line. Such
extreme caution would only be warranted where
a little flooding would be disastrous.
It is obvious that this float-valve could be used
under certain experimental procedures to add
chemicals to an aquarium at a prescribed rate by
inactivating the float and fixing the rate of flow
by hand. It also could be used to bring the con-
centration of some chemical to a fixed limit and
then hold it at that point in flowing water
4
Zoologica: New York Zoological Society
[42:1
Text-fig. 3. Diagram of interaction between float-valve and constant-level syphon. Dashed line
represents level of water in aquarium and syphon arm.
aquaria. The water supply would operate as
above described and a second float-valve regu-
lated to add much less chemical than the water
flow would move with it and act as a follower
to the other if there was any fluctuation in the
flow of water, thus holding the additive in pro-
portion to the change of water. Also a single
float could be arranged to operate the two valves
in proportion to the setting of each.
While the designs of these float-valves have
varied from time to time, all have been built
embodying the same principles as herein de-
scribed. The first and somewhat primitive one
has, at this writing, been in service continually
for more than four years and is still entirely
satisfactory and dependable.
Closed Systems
The term “closed systems” refers to circu-
lating systems in which the water is returned
to the aquaria after filtration or other treatment
and none is allowed to run to waste during
normal operations. Some such system is manda-
tory for the maintenance of marine forms re-
mote from a ready supply of sea water, and often
convenient or necessary for various experimental
procedures involving freshwater aquaria. This is
especially true of cases where it is necessary to
maintain close control of some feature such as
temperature, chemical quantities and the like.
By use of such means it is possible to maintain
a series of aquaria with absolutely identical
water conditions, as the water in all is part of a
common body. Consequently no matter what
transpires in one aquarium there is no oppor-
tunity for the water of that one to depart from
the characteristics of the rest since it is moving
freely from one aquarium to the other and is
being continually and effectively mixed.
An especially useful arrangement for some
purposes is one in which the flow between
aquaria may be continuously varied from maxi-
mum in one direction through zero flow to
maximum flow in the opposite direction. This
may be readily accomplished by the adjustment
of four valves while the pump runs continuously
in one direction at constant speed. The details
of the arrangement of these valves are shown
diagrammatically in Text-fig. 4. PI. II, Fig. 3,
is a photograph of such a device. In operation
the action is as follows. With valves A2 and B1
closed and the others open, the flow is out
through pipe A and returns through pipe B, as
indicated by the arrows, at maximum flow. If
these valves are reversed so that A1 and B2 are
closed and the others open, the flow through
pipes A and B is reversed, although the flow
through the pump remains as indicated by the
arrow on it. To pass uniformly from the first
position, as shown in Text-fig. 4, through a state
of no flow to the reverse, either valve A2 or B1
can be gradually opened. This reduces the speed
of water movement because of “back leakage.”
After one of them has been opened fully the
opening of the other can further retard flow.
When it, too, has been fully opened, that is with
1957]
Breder: Miniature Circulating Systems for Small Laboratory Aquaria
5
Text-fig. 4. Diagram of piping and valves for continuously variable flow from maximum in
one direction through zero flow to maximum in the opposite direction.
all valves fully open, there should be no flow
through pipes A and B, as there is as much pump
pressure in one branch of both A and B pipes
as in the other. Then by beginning to close either
valve A1 or B2 the flow begins to move in the
opposite direction. When these two are fully
closed, the maximum flow in the opposite direc-
tion has been reached.
If three valves are arranged on either line A
or B or such a set on both, various water treat-
ment devices may be placed in the series, such as
a heating or cooling device, in which case the
water may be best passed through a glass coil
for heat exchange purposes. This arrangement
could equally well be used for any kind of de-
contamination which might be required or for
the introduction of specified materials being
mixed with the passing water. Unlike most
aquaria plumbing, in this case the water must
pass from one aquarium to another, so that as
many as may be needed can be placed in series.
A diagram of a more usual arrangement for
a closed circulating system is shown in Text-fig.
5. This is the form which is perhaps most useful
for general laboratory purposes. Here each
aquarium is respectively supplied and drained
directly from a common supply and return. It
is advisable to use a pump of somewhat greater
capacity than needed for the purpose. With this
means the excess water can be returned to the
reservoir without passing through the aquaria.
which greatly assists in the eflBcient application
of whatever water treatment is being given and
insures adequate pressure for the system. Simple
filters may be made by wedging a piece of glass in
a small aquarium and filling the intake side with
suitable filter material, while the reservoir may
be used for whatever chemical or other treat-
ment is to be applied. Two may be provided, as
shown in the figure, and used alternately or
together. A constant-level syphon takes the
water to the first reservoir aquarium. Only one
syphon is shown, which may be switched to the
other aquarium for cleaning purposes.
The pipes and fittings used in this system are
standard hard rubber. Connections between
aquaria at the same level may be made by
“jumpers” which are preferably of hard rubber.
Their use is indicated in the lower level aquaria
shown in Text-fig. 5. These have been found to
be fully satisfactory and in several years’ opera-
tion have not clogged nor have they become
airbound. They are, however, not suitable for
the overflow lines of the upper series of aquaria.
Here constant-level syphons may be used as
shown in PI. I, Fig. 1, or, preferably, a hole may
be drilled in the slate bottom of each aquarium
and a one-hole rubber stopper holding a glass
tube inserted. Still better is the installation of a
hard rubber standpipe locked in place with fit-
tings. The drilling of slate is not easily accom-
plished and there is considerable danger of
6
Zoologica: New York Zoological Society
[42:1
cracking or otherwise damaging the aquarium.
The manufacturers will supply aquaria with such
holes drilled on order.
Since the supply to the aquaria of this system
is preferably from the top, as indicated, the drain
line as above described may be made to draw
water from the bottom of an aquarium by the
following simple means. A tube of glass, or other
material, of larger diameter than the drain tube
and as long as the depth of water in the aqua-
rium, is placed over it, reaching nearly to the
bottom of the aquarium. Since the larger tube
extends above the surface, water leaving the
aquarium must enter the annular space between
the two tubes and pass up between them to spill
into the open upper end of the inner tube. In
addition to giving the aquarium a better circu-
lation, much detritus is drawn up through this
arrangement and delivered automatically to the
filters. The outer tube, if of glass, may be posi-
tioned by slipping a short piece of plastic tubing
on its lower end and cutting various openings or
notches in the plastic. The area of these passages
should of course be at least equal to the cross-
section area of the inner tube. The annular space
between the inner and outer tubes should also
have this much area, at least. On the other hand,
it is best not to make the annular space much
larger than needed because this will cause the
water flowing through it to move with less speed.
The value of this arrangement as a detritus re-
mover is thus lessened, for the slower-flowing
water will not lift as heavy particles as will the
faster.
As such a system is usually intended to be
operated continuously for long periods without
attention, a safety feature may be built in which
would shut down the pump if the water in the
reservoh' rose too high or fell too low. The one
in current use, shown in PI. II, Fig. 4, was im-
provised from the tube of a mercury switch. This
was mounted on a rotatable glass shaft running
through a support of plastic. It was actuated by
a chemical flask float by means of a thread over
a small drum so that the motion of the float was
transmitted to the pump switch. Any unusual
change in the water level, either positive or nega-
tive, would indicate some radical failure at some
point in the system. Since the aquaria which held
the fish were drained by an overflow they would
continue to hold their water level so that stop-
ping the pump would insure the retention of
water there. Even if one of the aquaria leaked
and lost its contents the others would not suffer
by draining through the system to it because of
this protective device.
To prevent normal evaporation from stop-
ping the pump, a supply was provided which
operated in conjunction with the protective
switch. This supply was administered by a float
valve identical with that shown in Text-fig. 2.
A very nice adjustment was found possible with
these two float-actuated mechanisms, so that the
dripping from the float-valve supply became
1957]
Breder: Miniature Circulating Systems for Small Laboratory Aquaria
1
directly proportional to the evaporation, without
at any time tripping the protective cut-off ffoat-
valve. On very humid days it could be seen that
the number of drops per minute was notably
less than on a clear dry day when evaporation
was high. In the case of saltwater aquaria this
device had an added important application
which is discussed under the treatment of salt
water.
Because of the nature of the controlling de-
vices above described, it is necessary to observe
certain details in starting the system. The levels
of water in the lower series of aquaria will be
different when the pump is not running than
when it is in operation. This is mostly because
the drain lines of the upper series of aquaria
empty themselves into the lower aquaria when
the pump is stopped. Therefore the water is
carried at a lower level in these aquaria so that
there will be no overffowing when the circula-
tion has been stopped. For this reason a switch
should be shunted around the cut-off float to be
used in starting the system before the operating
level is reached. It will not suffice to wedge the
float into a position where its switeh will be
closed, because its free action is necessary to
establish its proper level of operation. After a
dynamic equilibrium has been achieved by ad-
justing both the cut-off device and the density
control device, the shunt switch should be
opened, after which the system should control
itself. If it does not at first, very obvious adjust-
ments of either or both will bring them into the
proper relationships.
The diagram of the closed marine circulating
system shown in Text-fig. 5 employed seven
aquaria for holding experimental fishes, only
three of which are shown in the illustration.
Three “reservoir” aquaria were used, of which
only two are shown. These were standard com-
mercial aquaria measuring 2' X 1' X 1'. The
two smaller, used as filters measured 10" X
8" X 6". The pump was driven by a 1/10 hp
motor and was rated at 10.8 gals, per min. at
zero head. The pipe sizes are not indicated, as
they would naturally vary with the needs of each
system. In this one the flow was slow but suf-
ficient at about three gals, per hour through
each of the seven top row aquaria. At the right
of Text-fig. 5 the supply pipe is extended up-
wards for some distance and with the upper end
open. This permits building up whatever head
of water is desired without subjecting the pipe
to pressure greater than that produced by
gravity.
Although the upper series of aquaria were
intended for holding fishes and the lower series
were regarded as treating reservoirs, the latter
too may be, and have been, used to hold fishes,
that is, all but the one from which the pump
draws water, since the suction and turbulence
here would be destructive to most small fishes.
Aerating stones and a standard aquarium glass
heater and thermostat comprised the rest of the
water-treating equipment. The heater which
turned off when the water reached 74°F. was
sufficient to keep the water throughout the sys-
tem close to that temperature as it was only
slightly higher than the normal room tempera-
ture. It was found that the aerating stones made
it possible to permit the flow of water in the
lower aquaria to run through submerged pipe
outlets and thereby reduce the amount of splash-
ing and consequent salt deposits. This was not
found necessary in the upper series, for each
supply pipe carried only one-seventh of the flow
in the lower pipes.
Materuls
It is strongly recommended for all the pur-
poses for which these devices were developed
that only hard rubber or some biologically inert
plastic be used. In fresh water, iron plumbing is
adequate for many purposes but for sea water
no metals whatever should be used if any de-
gree of satisfaction is to be obtained. Hard rub-
ber and acrylic resin or vinyl chloride-acetate
copolymer plastics have been used throughout
for those parts which come in contact with the
water, including the pumps.i Also it is important
to see that no brass or other such metals are
used in positions over the aquaria in order to
prevent possible corrosion falling into the water.
In all cases involving the use of pumps for
aquarium purposes it is best to have a spare
stand-by duplicate pump and motor unit as a
precaution against the failure of either motor
or pump. It is then possible to change such a unit
in a few minutes, in the case of accident, with
no serious interruption to the operation of the
system. It is most convenient to use flexible
connectors between the pump and the rigid
plumbing leading to the aquaria. It is then only
necessary to unfasten two screw clamps and in-
sert the new unit in place. This type of arrange-
ment is shown in PI. II, Fig. 3.
An additional advantage of this kind of con-
nection is that it dampens any vibrations, from
the pump or motor, which tend to travel
throughout the system along rigid connectors.
The pump in PI. II, Fig. 3, was suspended by
iln the systems described, the plastics used in the
construction of the apparatus go under the trade names
of Lucite and Plexiglas and the tubing under the name
Tygon. The pumps have been supplied, complete with
motors, by Broadbent-Johnston, Inc., Compton, Cali-
fornia.
8
Zoologica: New York Zoological Society
[42:1
four light cords, a means which is also very
effective in quieting such small machines.
Treatment of Water
The treatment of fresh water for aquarium
purposes is too well known to warrant comment
m present connections and is usually necessary
only under special situations. The maintenance
of sea water in a satisfactory condition is quite
another matter, however. It is not the purpose
here to discuss the theoretical aspects of the
chemical and physical conditions of sea water.
Such matters may be found extensively treated
by Sverdrup, Johnson & Fleming (1942) and
Harvey (1955). The following is intended
purely as a guide for the practical application
of principles which have been found adequate
to maintain a variety of marine fishes. Under
this treatment regular reproductive behavior
was quickly estabhshed in both Histrio and
Bathygobius, which had been reared from juve-
niles. It also permitted a variety of volunteer
algae and microorganisms to establish them-
selves. Incidentally these aquaria were kept
under conditions of no daylight, the illumina-
tion being supplied by fluorescent tubes of the
“warm white” type necessary for satisfactory
plant growth. The periods of light and darkness
were controlled by a time switch.
The equipment found necessary for the con-
trol of sea water consisted of a small hydrometer,
a colorimetric pH device and some simple titrat-
ing equipment.
The filters were provided with bone charcoal
and the bottoms of the aquaria and reservoirs
were floored with so-called coral sand, and
aerating stones were placed in various conven-
ient places, but not in the aquaria containing
fishes.
The specific gravity and pH were taken every
day until the rate of change was established and
from then on were taken at less frequent inter-
vals. This rate of change will vary with the
quantity of water, the bulk of the organisms con-
tained and the temperature of the water. At less
frequent intervals titrations were made to de-
termine the variously-called excess base, titra-
tion alkalinity, or alkaline reserve. This method,
which measures the bound CO2, is not especially
accurate but is sufficient for the present pur-
poses. It consists of titrating a sample with
N/lOO hydrochloric acid to which brom-cresol
purple has been added as an indicator. After the
purple color has vanished the sample is repeat-
edly boiled and further titrated until the purple
color no longer reappears on heating. If the
sample consists of 100 cc. to which five drops
of indicator have been added, the final burette
reading in cc. multiplied by 0.1 gives the bound
CO2 or bicarbonate in mfilimols/liter. This
method is not to be generally recommended for
accurate work but is sufficient as a comparative
measure of how far and how fast the aging water
is departing from its original value.
With this information, corrective measures
may be taken. The specific gravity is nearly
taken care of by automatic means involving the
use of the float-valve already discussed. Under
normal operations distilled water is used to make
up for the evaporation of sea water, which of
course tends to increase its density thereby.
This has been satisfactorily supplied from a five-
gallon carboy on a shelf higher than the float-
valve. The operation of the float-valve holds the
amount of water in the system at a constant
volume, which means also that the dissolved
salts will remain at a constant amount. If it is
desired to increase the density of the water, in-
stead of using distilled water as an additive, sea
water may be used until the specific gravity has
reached the desired level. If it is desirable to re-
duce the salinity, water may be withdrawn from
the system while distilled water is used in the
float-valve supply. This may be conveniently ac-
complished by means of a syphon with a small
hose clamp so that the flow is restricted to a
drip slow enough to permit the float-valve to
follow. Although distilled water was customarily
used, in its absence tap water was used with no
detectable effect on the fishes or the system.
If the pH falls to lower values it may mean
that there is an increase in the amount of free
CO2 present. This could indicate too many or-
ganisms for the volume and temperature of the
water or too much decomposition for the anti-
acid components of the system to dispose of
rapidly. The calcium carbonate in the sand
should react with the acids formed and unless
there is overcrowding this type of decreasing
alkalinity usually does not present a problem.
If the placing of fresh activated bone charcoal
in the filter results in an abrupt increase in the
pH, it is almost certain that there is too much
free CO2 present. The use of charcoal renewed
at short intervals will bring the CO2 content
down but the charcoal rapidly becomes satu-
rated and cannot be thought of as a regular part
of the regulatory process. An increase in the
number of aeration stones or amount of air they
pass, while much slower in its effects, is a much
more satisfactory way to insure against the ac-
cumulation of CO2.
If on a falling pH, none of the procedures
above mentioned increase the pH significantly,
the titration reading should be carefully checked
and it too should show a decrease. This would
indicate a lowering of the bound CO2 which does
1957]
Breder: Miniature Circulating Systems for Small Laboratory Aquaria
9
not normally occur in an imoverloaded system
in the presence of calcareous sand. If it does,
however, more sand may be added, or sodium
bicarbonate may be dissolved and administered
with the distilled water through the float-valve.
Since the sand alone tends in a long-term sense
to disproportionately increase the Ca in solution
as compared with the Na, the occasional use of
sodium bicarbonate, which tends to do the re-
verse, aids in keeping these two quantities in
more nearly normal proportions. See Breder &
Smith (1932).
The described procedures may seem to be
somewhat complicated, but they are, in fact, not
much more complex than those involved in
maintaining a similar number of standing fresh-
water aquaria. After the equipment is built and
regulated, so that valves and controls are in
balance, there is nothing to be done with them
at any time and in fact there should be no tam-
pering with them at all. It is probably wise to
post warnings to this effect. There is little aquar-
ium cleaning to be done, as most of the accumu-
lating detritus is automatically deposited in the
filters. Aside from feeding the fishes and some-
times cleaning algae off the glass sides, the latter
being controlled by adjusting the lighting ar-
rangements, there are the following routine
matters to be done. These will vary with each
installation but may be approximated by the
regimen under which the described installation
of seven aquaria were controlled, as follows.
Read pH and Sp. G Twice a week 5 min.
Titrate sample Once a week or
less 15 min.
Fill distilled water Twice a week 5 min.
carboy
Clean filters Once in 2 weeks 15 min.
Make adjustments based Once a month or
on above information longer Various
The need for changing the pump and motor is
such a rare occurrence as not to figure in the
above schedule and should not take more than
5 minutes. Every attempt has been made to re-
duce the maintenance of the system to its mini-
mum. It is not uncommon for the system to be
left alone for as long as three days, as over a
long week end. The only thing to normally expect
at the end of this period is some extra-hungry
fishes. It should be borne in mind that the
smaller the system— that is, its total amount of
water— the more rapidly decomposition or con-
tamination can spread through it. It is prudent
not to reduce the size of the equipment more
than necessary.
References
Breder, C. M., Jr., & T. H. Howley
1931. The chemical control of closed circulating
systems of sea water in aquaria for tropical
marine fishes. Zoologica, 9 (11): 403-442.
Breder, C. M., Jr., & H. W. Smith
1932. On the use of sodium bicarbonate and
calcium in the rectification of sea-water in
aquaria. Jour. Marine Biol. Assoc. N. S.,
18 (1): 199-200.
Harvey, H. W.
1955. The chemistry and fertility of sea waters.
Cambridge University Press, Cambridge,
i-viii: 1-224 pp.
Sverdrup, H. U., M. W. Johnson & R. H. Fleming
1942. The oceans. Prentice-Hall, Inc., New York,
i-x: 1-1087 pp.
Townsend, C. H.
1928. The public aquarium, its construction,
equipment and management. Report U. S.
Commissioner of Fisheries. Appendix 7:
249-337.
10
Zoologica; New York Zoological Society
[42:1:1957]
EXPLANATION OF THE PLATES
Plate I
Fig. 1. A constant-level syphon in operation.
Fig. 2. Float- valve for the control of inflow. Note:
This is the arrangement as used in the
closed saltwater system. The placement is
for convenience, but usually valves are
placed close to one end.
Plate II
Fig. 3. Pump provided with reversible flow de-
vice. The plastic pump here shown is
powered by a 1/80 hp motor and is rated
to deliver 3.7 gals, per min. at zero head.
Fig. 4. Safety control for closed circulation. Its
relation to the system is indicated in Text-
fig. 5. The pipes at the extreme right are
the pump intake and excess return.
BREDER
PLATE I
FIG, 1
FIG. 2
MINIATURE CIRCULATING SYSTEMS FOR SMALL LABORATORY AQUARIA
BREDER
PLATE II
FIG. 3
FIG. 4
MINIATURE CIRCULATING SYSTEMS FOR SMALL LABORATORY AQUARIA
2
The Behavior of the Bottle-nosed Dolphin {Tur slops truncatus):
Mating, pregnancy, parturition and mother-infant behavior^
Margaret C. Tavolga
Department of Animal Behavior,
American Museum of Natural History, New York
Frank S. Essapian^
Marineland Research Laboratory, Marineland, Florida
(Plates I-III; Text-figure 1)
Introduction
Because Cetacea bear their young in
water, data on the normal progress of
pregnancy and parturition, together with
information on mating behavior and post-natal
development, have been either lacking or only
incompletely available to biologists interested in
these animals. The greatest part of the informa-
tion as yet recorded has been received from
whalers and other sea-faring men who have
observed these animals in the open sea, where
conditions are far from ideal for such observa-
tions (Liitken, 1888; Pedersen, 1931; Slijper,
1936). James (1914) first observed the birth of
a Common Porpoise (Phocaena phocaena (Lin-
naeus) in captivity in the Brighton Aquarium,
but this was a stillbirth. Khvatov (1938) and
Sleptzov (1940) both observed what they be-
lieved to be normal births in the Common Dol-
phin {Delphinus delphis Linnaeus) in shallow
portions of the Black Sea, or in large seines in
which a number of animals were confined prior
to being drawn toward shore. McBride & Hebb
(1948) and McBride & Kritzler (1951) were
the first, however, to record their observations
under conditions almost ideal for this purpose.
In their 1951 paper, McBride & Kritzler pub-
^The authors wish to extend their thanks to the staff
of the Marineland Research Laboratory, Marineland,
Florida, for their cooperation and for the use of their
facilities and equipment. They also wish to express their
appreciation to Dr. Lester Aronson of the American
Museum of Natural History, Mr. F. G. Wood, Jr., of
the Marineland Research Laboratory, and Dr. Frank A.
Beach of the Department of Psychology, Yale Uni-
versity, for their special help in reading the manuscript.
^Present address: Seaquarium, Miami, Florida.
lished on this subject the latest data taken at
Marine Studios, Marineland, Florida, where the
activities of the Bottle-nosed Dolphin {Tursiops
truncatus Montagu) in a 75-foot circular tank
may be viewed conveniently through portholes
set in the walls. At that time five pregnancies
resulting in normal births had been under ob-
servation. Three animals had conceived in the
tank. Since then five additional pregnancies, all
resulting in normal births, have been observed,
providing many additional data. The present re-
port is based upon a series of observations in-
stituted by one of us which was conducted over
a period of slightly more than three years, and
which for the last year and a half was on a
daily basis. During parts of the latter portions
of these observations, both authors were able to
be at Marineland at the same time, and obser-
vations could be made almost constantly, so that
a steady daily record was kept which provided
the observers with continuous data, not only on
the group as a whole but also on the activities
of any individual animal.
Up to the time of the 1951 paper by McBride
& Kritzler, new animals were introduced at ir-
regular intervals and some deletions were made
in order to keep the population at a desirable
level and at the same time to minimize the pos-
sibilities of combat which resulted on a previous
occasion in injuries to a newborn dolphin (Mc-
Bride & Kritzler, 1951). Since then, the colony
has become a stabilized and self-perpetuating
group, in which the only additions are those by
birth. This fact has enabled the observers to ob-
tain data on the growth and maturation of sev-
eral individuals. One of the females born in the
tank has reached the beginning of her eighth
11
12
Zoologica: New York Zoological Society
[42:2
year and terminated her first pregnancy in
March, 1954.3
Table 1 shows the names, sexes, dates of birth
and relationships of the members of the stable
population of the tank. In addition. Happy, a
fully-grown adult male, and presumably the
father of all the infants conceived in the tank,
was present throughout the three years covered
by the present data. Frank and Floyd, born in
the tank, were removed in December, 1953.
actively sought the company of the selected
female by such stratagems as chasing her when
she left him, hovering over her when she hunted
for fish or was otherwise occupied in one spot,
and by returning to her immediately after both
had been engaged in other activities, such as
feeding. Such exclusive companionship lasted
from three or four days to several weeks.
This period of prolonged companionship of-
ten, but not always, led to further activities on
Table 1. Stable Population of Tursiops truncatus in Tank at Marine Studios.
Date of Sex of Name of Name of Duration
Birth
Infant
Infant
Mother
of Birth*
Remarks
Feb. 26, 1947
Female
Sprayt
Mona
28
April 23, 1948
Male
Mrs. Jones
21
Died of injuries
May 8, 1949
Male
Algiet
Susie
117
May 12, 1950
Male
Frank
Pudgy
May 15, 1950
Male
Floyd
Mrs. Jones
Feb. 7, 1953
Female
Mamie
Mona
Died June 13, 1953
Feb. 23, 1953
Female
Maggie
Mrs. Jones
Feb. 27, 1953
Female
Nellie
Susie
May 7, 1953
Male
Mitch
Pudgy
68
Died Feb. 22, 1954
March 4, 1954
Female
Peggy
Spray
48
Died March 19, 1954
*Duration of birth of infant in minutes, from first appearance to complete emergence. Where no figure appears,
the birth took place at night and was unobserved.
■[These animals were conceived in the wild before the capture of the mother. AU other infants in the table
were conceived in the tank.
Precopulatory Behavior
In the spring of 1952, 1953 and 1954 (Feb-
ruary to May inclusive), there occurred a num-
ber of special activities which appeared to be
the preliminaries to copulation, since the copu-
latory pattern was frequently seen to follow
them. The aggregate of these activities over a
period of time between two animals of opposite
sex we term courtship. It should be understood
that not all of these activities need necessarily
take place between any two particular animals,
and further, that they need not follow any par-
ticular order.
During the major portion of the year the
adult male changed his companions often, swim-
ming with one animal for a few minutes or a
few hours, and rarely remaining with it for more
than a few days. His companions during this
time were other males, females or young ani-
mals of either sex. He also remained alone for
long periods. During the early spring, however,
he showed preference for the company of one
particular female and remained with her for
prolonged periods. He either accepted the pres-
ence of other animals or avoided them, but he
®This paper was submitted for publication on Feb-
ruary 13, 1956.
the part of the male, depending on the at-
tendant circumstances. At times, without any
further activity, the male suddenly broke off the
partnership and shifted his attention to other
interests, and there was no further association
between the male and female for weeks or
months. On the other hand, he often suddenly
increased his attention to her and engaged in
activities which were more aggressive and more
directly associated with precopulatory behavior.
These activities were quite varied and included
posturing, stroking, rubbing, nuzzling, mouth-
ing, jaw clapping and yelping. Any or all of
these were included in any sequence and varied
in intensity, becoming most intense just before
exhibition of the copulatory pattern.
Posturing. — Although this type of act ap-
peared at any time during precopulatory be-
havior, it was usually one of the first indications,
aside from prolonged companionship, that pre-
copulatory activities were about to become more
intense. The male swam toward the female and
took up a position in front of her, or slightly to
one side, but well within her line of vision. He
then bent his body in such a way that from the
side it took on the shape of an S-curve, head
pointing up, flukes down, as in Text-fig. 1. He
maintained this position for about two to five
seconds, and then resumed normal swimming.
1957]
Tavolga & Essapian: Behavior of the Bottle-nosed Dolphin
13
The action was repeated immediately, after
several minutes, or not at all, depending on the
female’s general responsiveness and the other
activities taking place in the tank. All males
postured sometimes, but males were seen to
begin courtship without posturing. Therefore it
appears to be a subordinate portion of the pre-
copulatory pattern of behavior. Although pos-
turing was usually shown by the male, it was
observed in the female under certain circum-
stances which will be described later.
Text-fig. 1. Male dolphin in posturing position.
Stroking. — During normal swimming the
male took up a position just behind and below
the female. When this happened, the female
often altered the normal motion of her flukes
in such a way as to stroke or lightly pat the
head of the male as she swam. At times the posi-
tions were reversed, and the male stroked the
female’s head. The stroking motion was easily
distinguished from the normal swimming mo-
tion, and also from the lashing of the flukes
given by another animal in an attempt to avoid
contact or to get away. Such action on the part
of the two animals was usually assumed when
they were resting, and was the least vigorous
and energetic of the sexual activities. At other
times either male or female swam in such a
way as to bring one or both flippers in contact
with some part of the other’s body (PI. I, Fig. 1 ) ,
and by swimming along or across the other’s
body produced a stroking effect. Another form
of the same activity took place when one of the
animals turned on its back and, swimming be-
neath the other in the same direction and at the
same speed, moved its flippers rhythmically back
and forth across the other’s body. The other
member of the couple responded by repeating
the action at the some time so that the two sets
of flippers stroked each other. Such actions
could, of course, be continued only until it was
necessary for one or the other to rise to breathe,
but it was resumed immediately following this,
and with interruptions for respiration was some-
times continued for half an hour or more.
Stroking was also carried out by gentle move-
ments of the head against the body of the
partner. Another variation took place when one
animal placed itself beside or beneath the other.
and stroked the genital area of the partner with
the tip of one fluke.
Rubbing. — This was a much more strenuous
action in which one member, usually the male,
swimming at a fast pace, advanced upon the
female and rubbed part of his body vigorously
against hers (PL I, Fig. 2). This advance took
place from any direction relative to the two
animals, but was most dramatic when the male
approached the female head on, as if to engage
in a head-on collision, and then turned aside at
the right moment and just enough to produce
vigorous contact of the two bodies along their
lengths. This type of inter-animal contact rarely
lasted for more than a few minutes, during
which the male made repeated rushes at the
female. Trumpeting and whistling sounds were
sometimes heard during this time. The female
sometimes turned after each rush and swam
toward the male as he repeated the maneuver,
enhancing the vigor of each contact, or she
rushed away, leaping out of the water in a real
or apparent attempt to escape. When the latter
took place, the male usually rushed to the spot
where she was to reenter the water and rubbed
energetically against her as she did so. (PI. I,
Fig. 3).
Nuzzling. — Nuzzling took place when one
animal applied its closed snout to some part of
the body of the partner and moved it around
in that area. Although this action took place in
a number of other situations, such as the nuzzl-
ing which goes on between mother and infant,
the male often nuzzled at the genital area of the
female, possibly as an investigative measure,
and also possibly stimulating her thereby. The
female, at times, nuzzled the male in the same
way.
Mouthing. — (PI. I, Fig. 4) Mouthing was an
activity similar to nuzzling, but took place with
the mouth open. Although used during precopu-
latory activity in the same manner as nuzzling,
it also took place in other ways. When in an
appropriate position, one animal took a flipper,
fluke or sometimes even the head or snout of
the partner between the teeth (PI. I, Fig. 4) , and
gently closed the mouth upon it, thereby holding
the part. This grasp was a gentle one, and was
released as the partner pulled away, without
leaving tooth marks or gashes. This is in con-
trast to the grips taken on each other by two
fighting males, which result in deep cuts and
gashes inflicted by the sharp teeth. Either nuz-
zling or mouthing of the male’s genital region
by the female sometimes resulted in an erection.
Jaw Clapping. — At times during the com-
panionship of the male and female, a second
14
Zoologica: New York Zoological Society
[42:2
male interfered in the relationship between
them, either by swimming between the two,
separating them, or by nuzzling the female
while she swam with the first male. In such a
case, the male’s response was rapid and often
violent. He is able, by clapping his jaws together
forcefully, to produce a sharp loud sound, which
can be heard at a distance of fifteen feet or more
from the outer wall of the tank. As early as
1940, McBride recognized this sound as “the
principal form of intimidation” when produced
by a dominant animal. Often jaw clapping was
sufficient to discourage the intruder, but if not,
the first male resorted to violent lashes with his
flukes, sometimes throwing the offender against
the side of the tank with tremendous force; or
he actively attacked the intruder with his teeth,
inflicting serious gashes upon the other’s flukes,
flippers, back and head. Jaw clapping was also
used if the female swam away for a short time,
or if another female interfered. In these cases,
however, the final results were rarely violent,
since the partner rejoined the male immediate-
ly, and an interfering female hastily left. In
other situations, jaw clapping was used by non-
dominant animals as a sign of displeasure and
not necessarily as intimidation.
Yelping. — During the more intense stages
of precopulatory behavior, another sound, char-
acteristic of this situation, was often heard in
the tank. This sound has been interpreted to be
a type of mating call (Essapian, 1953; Wood,
1953). It was a high-pitched yelp, delivered
as a series of short sharp cries. The number of
yelps in a series varied from two or three to
seven or eight, and the series might be repeated
after an interval of about fifteen seconds, or
might not be repeated for a much longer inter-
val, depending upon the circumstances. There
can be no doubt that the male produced these
sounds, since when he was in the proper posi-
tion relative to the observer, the flap of the
blowhole and the area behind the blowhole
could be seen to move as if they were being
depressed at the time that the sound was heard.
No bubbles escaped from the blowhole with the
production of this sound. The male yelped very
rarely when he and the female were swimming
quietly together. On the other hand, if the fe-
male strayed away from his vicinity, or stopped
to investigate another animal or an object in the
tank, in the majority of cases the male promptly
produced a series of the yelping cries and con-
tinued to do so at short intervals until the fe-
male returned to him. If she did not respond
at once, he continued to yelp until she did. Im-
mediately upon her return to him the yelping
ceased. If another female responded, she was
ignored. Since, as far as the investigators could
determine, yelping was heard only during the
mating season, and the selected female re-
sponded, it is felt that the interpretation of this
sound as a mating call is valid.
Although the male consistently swam with
a single female during the earliest days of the
spring season, the most intense of the precopu-
latory activities did not take place at this time.
Such activities seemed to begin shortly after the
birth of the first infant of the year, and evidence
is accumulating that true precopulatory activity
may be triggered by this birth. In 1953, the first
infant was born on February 7, and the first dis-
play of intense sexual activity on the part of
the male, accompanied by yelping, was seen
three days later. Since no observations were
taken on the two previous days, it may have
taken place even earlier. In 1954 the first infant
was born on March 4, and the beginning of
yelping and intense sexual aggressiveness on the
part of the male began the same afternoon.
Since the March date is quite late in the year
for the appearance of the first infant, and con-
tinuous observations were carried out through
all of February, it can be seen that there was a
possible connection between the two events.
However, more evidence is necessary to estab-
lish a clear relationship between them.
The pattern of the male’s sexual activities
previous to copulation was not a consistent one.
Any or all of the actions described above were
involved, and any or all of them could take
place within an hour or two. On one day the
male engaged in each of them and periods of
intense activity alternated with periods when
the two animals swam quietly together for an
hour or more. On another day he persistently
engaged in one of them, rubbing for instance,
though the entire day, and displayed none of the
others, resting only a few minutes at a time
before resuming his activity.
Initiation of precopulatory behavior by the
female.— The part of the female in a paired sit-
uation was less energetic than that of the male.
When she was the chosen companion, it was
only rarely that she engaged in any but the
mildest of precopulatory activities toward him.
She swam close to him, perhaps maintaining
contact with him by an extended flipper touch-
ing his body, and she was occasionally seen to
initiate mild stroking activity. She actively co-
operated with him, however, when he initiated
the more intense actions, and there was one
occasion when she actively took the lead. As
mentioned above, during the height of the rub-
bing activity on the part of the male, she often
1957]
Tavolga & Essapian: Behavior of the Bottle-nosed Dolphin
15
rushed away from him at great speed, and
leaped far out of the water, perhaps to escape
his attentions. On such occasions, he sometimes
joined her in her leaps, but more often he swam
rapidly to the spot where she was expected to
enter the water, where he rubbed against her
even more energetically as she came down. This
sequence of events was often repeated a number
of times before both animals reverted to swim-
ming together quietly. Since after such activity
the female immediately rejoined the male and
continued to swim close to him, and because
copulatory activity often followed such a dis-
play, it is possible that the display aroused more
intense activity on the part of the male.
In the case of an unpaired female, however,
the situation was quite different. At various
times an unpaired female was seen engaging in
somewhat unusual activities, such as scraping
her ventral side along the bottom of the tank
on the coquina gravel, and rubbing her genital
area vigorously and repeatedly over the brushes,
which are set up in the tank as back-scratchers
for the animals. These actions have led the ob-
servers to suspect that such a female was in a
state of high sexual receptivity, and in some
cases this opinion has been confirmed by her
subsequent actions. An unpaired female in this
state often postured before the male, and some-
times initiated stroking and rubbing activities.
She approached him from all angles and at-
tempted to maintain physical contact with him.
She was often very persistent in these activities
even though the male paid no attention to her.
She nuzzled and mouthed his genital region, and
this latter behavior was sometimes followed by
an erection on his part. If this happened, she
often stationed herself within his line of vision
and remained motionless in the water, her ven-
tral side toward him as if inviting his attention.
If he did not respond, she sometimes repeated
some or all of these actions. In addition, she
sometimes sought out the younger of the two
males, who just reaching maturity, and initiated
these activities with him. Since this younger
male was often rejected as a sexual partner by
the mature females, he usually responded to
such attentions, and if he did so, the female al-
lowed him to begin copulatory behavior. Such
a female was seen to hang motionless and almost
vertical at the surface of the water with only
her blowhole exposed, while the younger male
persistently engaged in copulatory behavior.
Uusually he was not completely successful. At
times, the mature and dominant male was at-
tracted to the scene of this activiy, and par-
icipated in it himself, in which case the two
males alternately approached the female.
Copulatory Behavior
The males exhibited erections during the
more vigorous phases of sexual activity. Erec-
tion and the copulatory pattern often followed
the leaping display of the female or a period of
strong rubbing.
The copulatory activities which have been
observed may be divided as follows: erection,
approach, intromission, ejaculation, and with-
drawal.
Erection.— The penis of the Bottle-nosed Dol-
phin erects suddenly ( 1 to 2 seconds) , the body
of the organ curving outward and ventrally
from the genital slit in which it is ordinarily
enclosed, followed by the tip which springs rap-
idly away from the body. The erect penis is
directed craniad, and deviates from the longi-
tudinal axis of the animal an estimated 20 de-
grees to the left.
Approach.— Because of the sinistral deviation
of the penis, the male usually approached the
female on her right side, and partially rolling
on his right side, brought the penis close to her
genital opening. He approached from the rear
and parallel to her, at an angle perpendicular
to her path, or at any angle between these two
extremes. He swam toward her, dorsal side up,
rolling over as he approached her, or he turned
on his back at a distance and swam toward her
in that fashion. Sometimes he approached from
her left side but this was unusual.
At his approach, the female did any of three
things. Sometimes she swam away from the
male, or continued swimming at his side, with-
out changing her position or speed. In both of
these cases, entry was almost impossible, and
the male usually desisted after a short time.
At other times, she rolled over on her left
side and presented her ventral side to the male,
at the same time slowing her speed. This posi-
tion allowed the male to effect penetration more
easily.
Intromission.— (Bl.l, Fig. 5) When the male
was close to the female, with his ventral sur-
face approximating hers, he brought the erect
penis into contact with her ventral side in the
region of the vagina. This contact was often
repeated many times before intromission was
achieved, since both animals were swimming
slowly throughout, and other animals and
water currents sometimes caused the copulat-
ing pair to move apart. If there was no outside
interference, and the female matched her
speed to that of the male, intromission was
effected. In most of the cases seen by the
present observers, the penis was inserted no
more than to half its length into the vagina.
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Pelvic thrusts were often observed, both be-
fore and after intromission was effected. They
usually began as soon as the ventral side of
the erected penis came into contact with the
ventral side of the female. They continued
only as long as the contact was maintained,
and at times did not occur until penetration
took place. A series of these thrusts usually
lasted for two or three seconds, but the series
was sometimes repeated a number of times.
The intromission lasted for not more than ten
seconds, after which withdrawal took place.
Ejaculation— observers do not know
whether ejaculation took place during intro-
mission, since the animals could not be cap-
tured and sperm counts made. On one occa-
sion, after a partial intromission of short
duration such as those mentioned above, the
animals broke apart suddenly, and a stream of
milky fluid was ejected into the water from
the penis of the male. It is thought that this
was semen.
Withdrawal— Atier an intromission lasting
from two to ten seconds, the animals broke
apart suddenly. Each righted itself and
swam normally. The penis at this time was
still erect. Often the male made a new ap-
proach immediately, and the above activities
were repeated. Sometimes the penis withdrew
suddenly into the genital slit and the two an-
imals swam off in different directions. At
other times the copulatory pattern was re-
peated at intervals of one to seven or eight
minutes for a half hour or more.
The greatest part of copulatory activity by
the bull took place at night or in the early hours
of the morning. It is not certain whether the
reason for this lay in the diversions created dur-
ing the day by feeding periods or whether it was
the result of nocturnal preferences by the ani-
mals. In the past when there were fewer feed-
ing periods the animals also tended to copulate
at night. Their behavior in the wild in this re-
spect is, of course, unknown.
Just after dawn on March 1, 1953, a member
of the Marine Studios staff reported seeing a
copulation between an adult male and a mature
female. Although he did not observe the two
coming together, he reported that the male
swam on his back beneath the female. The ven-
tral surfaces were closely pressed together so
that the penis of the male was not visible. This
position was retained for about thirty seconds.
When the pair broke apart, the penis of the
male was erect. No traces of the ejaculate were
seen in the water. Since the female involved
produced an infant the following spring, it is
possible that it was at this time that conception
took place. This, of course, is far from certain
and such data can be used only to narrow the
time of conception to within a week or two,
since the female was paired with the male dur-
ing that period of time.
The single instance described above was of
longer duration and there was complete pene-
tration of the penis. Those seen by the present
observers were of shorter duration and involved
partial penetration. The observers were not able
to determine whether the single instance was a
typical copulation and the others incomplete,
or whether a short copulation with partial pene-
tration is typical and the single instance atypical.
There were certain complicating factors in-
volved in such observations, since other animals
were attracted to the site of copulatory activities.
Young males, particularly, exhibited erections
and competed for the female by crowding and
pushing one another aside. In addition to inter-
fering with the copulating animals, they also
effectively obscured the view of the observers.
Behavior During Gestation
Since the report of McBride & Kritzler
(1951) five conceptions have taken place and
the resulting pregnancies have been carried to
completion, each terminating in the birth of a
normal healthy infant. Four of these were rec-
ognized during the latter months of the gesta-
tion period. These resulted in the four births
during the spring of 1953. The flfth (1954
birth) was recognized as a possibility from the
time of conception, and its progress recorded
throughout.
“Spray,” a young female, was born at Marine
Studios on February 26, 1947, and was the first
normal infant porpoise born in captivity. She
was, from the first, a healthy animal and despite
bruises and lacerations inflicted on the day of
her birth by sexually aroused males in the tank,
she grew and progressed normally. She was a
gregarious animal, and learned early to partici-
pate in all of the activities of the other animals,
as well as to respond to the attentions of human
beings when members of the staff played with
her. Although McBride & Kritzler believed that
she might initiate her first pregnancy in the
spring of her fourth year, it was not until the
spring of 1953 (the beginning of her seventh
year) that this event took place.
On February 10, 1953, the only adult male
dolphin in the tank began to show a persistent
interest in Spray, and continued to seek her com-
pany for the following 13 days without eliciting
any interest on her part. He was heard yelping
on several occasions, also without any response
1957]
Tavolga & Essapian: Behavior of the Bottle-nosed Dolphin
17
from her. He continued his pursuit of her, and
on February 23 they were seen swimming to-
gether for prolonged periods for the first time.
At this time the “courtship” took on what might
be considered a typical pattern, including all
the types of precopulatory behavior described
in the previous section. Yelping by the male was
often heard and Spray responded to it during
this period. In addition. Spray was often seen
leaping out of the water, followed by the male
who persisted in keeping in close physical con-
tact with her. No erections were seen during
this activity. Early on the morning of March 1 ,
it was reported that the bull and Spray were seen
copulating. No further such incidents were re-
ported, although it is probable that others took
place. The two kept up their close association
until about March 9, when it was observed that
they were no longer constantly together. Dur-
ing the several days following this, the associ-
ation was abandoned except for occasional short
contacts, and for the remainder of the spring
season the two animals were rarely seen together,
each seeking the company of other dolphins. The
male was seen either alone or in the company of
other females. Spray reverted to her most con-
sistent occupation, swimming with one or an-
other of the new infants.
In the spring of 1949, a male infant was born
to another of the adult females. From the time
of his birth. Spray was almost constantly in his
company, and took on the role of playfellow,
not only to him, but to all of the later infants
born in the tank. She often remained with them
when their mothers were at the feeding platform,
and was often seen swimming about the tank
with them when their mothers were otherwise
occupied. It was natural, therefore, that she
should continue this occupation with the infants
born in 1953, and she continued in these acti-
vities well into the summer.
It has been reported in McBride & Kritzler
(1951 ) that during the latter half of pregnancy,
the female tends to withdraw from association
with others. In the case reported in that paper,
two females, both pregnant, spent most of their
time together, apart from the rest of the tank
population. In the present case there were no
other pregnant females present. Although Spray
was often sought out as a companion by other
members of the tank community, she showed an
obvious preference from her earliest days for
the company of her mother, “Mona,” and this
preference has continued to the present. There-
fore, when she began to withdraw from the com-
pany of the other animals, she sought her mother
as a companion. The first sign of this association,
which was to continue throughout her pregnan-
cy, was seen on June 27. On that day it lasted
for only a short time, but was repeated on July 1,
July 3 and again on July 7. Thereafter it became
a more regular companionship, and the time
Spray spent with Mona gradually increased from
a few minutes a day till, toward the end of her
pregnancy, almost all the time she spent in com-
pany was with her mother. The remainder of the
time in company was spent with “Pudgy,” an-
other mature female, and her recent infant,
“Mitch” with both of whom Spray had spent
much time since the infant’s birth in May, 1953.
Although closely observed throughout, it was
seldom after July 8 that she was seen in the
company of any other animal, and when such
instances occurred, they lasted for brief periods
only. It was also noted that in most of these in-
stances, the other animal involved approached
Spray, while she displayed little interest in seek-
ing out other company. At this time and there-
after throughout her gestation, it was also noted
that her activities decreased in vigor, and she
adopted habits that were slower and more lei-
surely.
On June 30, the increasing roundness of her
abdomen was noted for the first time, and al-
though the animals have been known to gain
and lose weight for various unknown reasons, at
this time the possibility of her pregnancy be-
came, in the minds of the observers, a prob-
ability. PI. II, Figs. 6 & 7, show the difference in
contour between a non-pregnant dolphin and one
in a late stage of pregnancy.
On October 2, it was noted for the first time
that Spray’s mammary glands showed percep-
tible growth, and the area just forward and on
each side of the genital opening, which had
heretofore been rather flat and undemarcated
from the surrounding region, now became round-
er and fuller, as the glands increased in size.
This definition of the glands became greater as
term approached, although the glands never
reached the size of those of some of the other
mature, lactating females.
On December 9, straining or body flexion was
observed for the first time in this pregnancy.
Straining always occurred when the animal was
in the horizontal position, often when she was
swimming. If the animal was resting at the sur-
face she remained there, with only the blowhole
out of water. If she was swimming below the
surface, she usually came to a complete halt.
The flukes and caudal peduncle moved ventral-
ly (PI. II, Fig. 8), and as the flexion reached
its maximum, the tail approached an angle of
45 degrees from the horizontal, forming an angle
with the body of about 135 degrees. Often the
head also bent ventrally, and as the strain
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[42:2
reached its climax, the entire body formed an
arch, and the muscles appeared to be taut. This
position was held for a few seconds. When the
animal relaxed, the tail passed the horizontal
and was raised dorsally, where it was held up at
an angle almost as great as in the other direction
(PI. II, Fig. 9). This recurved position was also
held for a few seconds, and then the animal re-
laxed again and swam off in normal fashion.
The arch might occur without the recurve but
the recurve never took place alone. Although
body flexion has been seen in animals other than
pregnant females, it is most noticeable in such
females, and is considered a characteristic ac-
tion in the latter months of pregnancy because
of its repeated appearance. It was not until
January 12 that Spray was seen straining again,
but after this date it was seen often. From the
beginning of February, flexions were seen to oc-
cur at the rate of about one a day, gradually in-
creasing in frequency until the rate was two or
three a day by the middle of the month. Toward
the end of February, the total number of flex-
ions per day increased, and the pattern changed
substantially. During this period Spray might
go through an entire day during which no flexi-
ons were seen by the observers, while on another
day she could be observed to strain almost con-
tinuously for an hour or more, about two or three
minutes elapsing between flexions. A period
without flexions then followed, succeeded in
turn by another period in which straining was
frequent. This pattern continued until just be-
fore the birth took place, and as term approach-
ed, the periods of flexion succeeded each other
with decreasing periods between until just be-
fore term, when Spray was recorded as strain-
ing almost continuously throughout several hours
every day. On the morning of the day before
she gave birth, she was seen to flex seventeen
times in thirty-seven minutes.
As term approached, there were also other
activities in which Spray was seen to participate,
and which were not usually seen in non-pregnant
animals. In a number of instances, she came
down over the brushes which were anchored to
the floor of the tank and applied her ventral
side to them, scratching vigorously. At times she
was seen to actually insert some of the bristles
into the genital opening and wriggle back and
forth over them. She also glided down to the
piles of rocks on the floor, and as she approached
them, she lowered the posterior end of her body,
so that she bumped her ventral side against them
with considerable force. Twice she did this with
such force that large rocks were displaced. The
activities could, perhaps, be correlated with in-
creased vaginal irritation or tension.
During the latter months. Spray’s size in-
creased considerably, especially in the abdomi-
nal area. Although porpoises, which are exceed-
ingly streamlined animals, do not show the dis-
tortions in body shape exhibited by some other
animals during late pregnancy, even casual visi-
tors to the exhibit were able to observe her
greater girth and often commented upon it. At
this time her navel, which is normally a slightly
depressed scar, became slightly elevated from the
body surface. Formerly a fast and graceful
swimmer, she became at times almost clumsy in
her body movements.
Several weeks before the birth Spray was ob-
served to change her habits of respiration. She
frequently rose to the surface, and hovering
there, respired a number of times in succession,
the total time elapsed being about two or three
minutes. Thereafter, she resumed normal swim-
ming. A minute or two later the behavior was
repeated. This might be followed by an interval
of normal breathing. As term approached the
unusual breathing pattern became more frequent,
and consequently she came to spend most of her
time swimming at the upper level of the tank
quite close to the surface. Only occasionally did
she descend to the floor, usually to rub herself
over the brushes or rocks. During this time she
was also observed to open her mouth widely at
intervals, as if yawning, and she sometimes pro-
jected her tongue beyond the tip of her upper
jaw, as if licking it. She did this while beneath
the surface of the water, and its significance is
unknown.
At this time there was also an increase in the
number of defecations per day. This number in-
creased until during the last few days before
birth defecation was almost continuous. Nor-
mally the animals defecate after feeding, and
occasionally between feedings. Since the feces
dissipate rapidly into the water, it was difficult
to estimate the amount, but it is believed that the
total amount of feces was also increased. It is
probable that the increased numbers of both
respirations and defecations were caused by
pressure of the growing uterus upon the other
internal organs. Feeding continued normally
throughout this time and through the day pre-
ceding the birth.
During the last few weeks, any exertion on
Spray’s part produced another visible sign that
parturition was not far off. During flexions, or
during the spurts of vigorous swimming in which
she occasionally indulged, the vaginal opening,
ordinarily a closed slit, was seen to dilate to a
width of about a half inch to an inch (PI. II, Fig.
10). As term drew near, these dilations became
more frequent and also more pronounced. The
1957]
Tavolga & Essapian: Behavior of the Bottle-nosed Dolphin
19
opening widened to approximately two inches
and the dilation sometimes lasted as long as five
to ten seconds before relaxation set in.
During this last month the movements of the
infant inside Spray’s abdomen were easily seen.
At times, distinct bulges could be seen on various
parts of her abdomen, bulges which shifted from
place to place, sometimes in a matter of seconds.
When this happened distinct movements could
be observed which were not due to the action
of the mother’s muscles. When a particularly
violent movement of the foetus took place, she
often halted abruptly and waited in mid-stroke
until the movement had stopped and the foetus
was quiet again. On February 4, a bulge ap-
peared on her abdomen, posteriorly and ven-
trolaterally on the right side. This was persist-
ent, lasting for several days. On February 13,
Spray’s contours changed again, and evidently
the foetus was now situated more anteriorly than
had previously been the case, since bulges were
seen on both sides in the anterior part of the
abdomen. Later the foetus shifted more to the
left, and on February 23 an area on her left side
just behind her flipper was noticeably protrud-
ing. This condition persisted until the time of
parturition.
Parturition
On the morning of March 4, both observers
were called to the tank at 5 a.m. The birth was
observed to begin at 4:55 a.m. by a member of
the Marine Studios night crew, who had been
previously alerted as to the imminence of the
event. Spray was seen swimming slowly and nor-
mally around the tank with her mother, Mona.
The flukes of the infant were protruding from
the vaginal opening and were lying in a horizon-
tal plane. Most of the other animals in the tank
were acting normally, and only when one of
them came close enough to touch Spray or the
baby did she speed up or swim evasively. At 5 : 1 5
a.m. the flukes turned at right angles to their
former position, and the infant was seen to be
lying on its left side. At 5:20 a.m. about three-
quarters of the caudal peduncle had emerged. As
she swam. Spray flexed mildly and the foetus
emerged still further, but as she relaxed, it re-
ceded again. The situation continued in this way
until 5:40 a.m., when the entire length (about
14 inches) of the caudal peduncle had appeared.
PI. II, Figs. 11 & 12, show two stages of birth.
At this time Spray was still swimming slowly, but
in a normal fashion, with Mona. At 5:43 a.m.
there was suddenly a great deal of excitement in
the tank. The animals gathered in the center
amid whistling and a variety of other noises. Al-
I though banks of lights had been set up in the
I event of a birth at night, the excited racing of the
animals and the flurry of bubbles in the water
which this produced, plus the darkness of the
hour, kept the observers from seeing what took
place during the exact instant when the infant
fell free from the mother. The infant was first
seen at the surface a few seconds later, against
the light at the far side of the tank. The total
time elapsed was 48 minutes. A minute or two
later the animals had again broken up into
groups, and the infant, a female, was seen swim-
ming uncertainly, but energetically, between the
dorsal fins of Spray and Mona. The baby’s dor-
sal was inclined to the left and her flippers and
flukes were soft and flabby. A number of ver-
tical creases without pigment were seen along
her sides. This is characteristic of newborn dol-
phins. A few minutes later, at 5:47 a.m., the
adult male, possibly aroused by the birth, gave
chase to Mona, and her place beside the baby
was taken by Pudgy, the other female which
had spent much time with Spray during her
pregnancy. Shortly thereafter, however, the bull
turned his attention to Pudgy, and Mona re-
turned to swim with Spray and the infant. These
three continued together, and the two adult
animals guarded and guided the new infant
throughout the first days of its life.
As soon as it became light, the skin of Spray’s
abdomen was seen to be patterned in a series of
fine wrinkles, as in crepe paper. Probably this
was the result of the strong contraction of the
muscles beneath, which resumed their normal
tonus before the skin could do so. The crepe
paper effect lasted for several days before it
disappeared.
At 9:15 am. Spray, who had been swimming
slowly with the infant and Mona, suddenly
speeded up and raced across the tank. This was
the first rapid swimming she had done for several
days, and as she did so the placenta emerged
from the vaginal opening as though forcibly ex-
pelled. There was no blood visible as it was
ejected, and Spray paid no further attention to
it. Immediately after the placenta was expelled,
Spray slowed down again, and swam off with
the baby. She did not approach the placenta
again.
At 10:55 a.m. the new infant began to nuzzle
her mother’s side. Spray neither slowed down
nor rolled over at this time. At 11:18 the baby
nuzzled again, this time more persistently, but
without success. Again Spray did not cooperate.
At 1:00 p.m., however. Spray rolled over on her
side and adjusted her speed so that her mammary
region was close to the baby’s snout. The in-
fant displayed no reaction to this, and did not
nurse. At 1:22 p.m. the baby nuzzled again.
This time Spray rolled over, and after several
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[42:2
thrusts of her snout, the infant found the nip-
ples, and suckled successfully for the first time
(PI. Ill, Fig. 13). Another nuzzle a minute or
so later was unsuccessful, but others shortly
thereafter and later in the afternoon showed that
the young one was locating her objective more
rapidly. Within 24 hours she was seen to suckle
regularly without any preliminary searching.
In 1953, the year previous to the parturition
just described, four other births occurred. Each
of these produced a normal, living infant. All
the conceptions took place in the tank at Marine
Studios. There were certain features in which
each differed from the one just described.
During the last few days of January, 1952,
Mona was observed to be spending most of her
time with the adult bull porpoise, and was re-
ported in copulation with him at that time.
Shortly thereafter, she ate very little for a period
of almost a month, although still in the com-
pany of the bull. She remained with the bull
through most of February, although observa-
tions indicate that during the latter half of Feb-
ruary it was Mona who sought the bull’s com-
pany, rather than the reverse.
It was only in November, 1952, that her in-
creased size attracted attention to her condition.
On February 6, 1953, Mona’s abdomen showed
a shallow, concave area ventrally between the
region of the navel and that of the mammary
glands, which could be seen only when viewed
from the side. Anterior to this region the dis-
tension of her abdomen was conspicuous. This
concavity did not appear on Spray, and its signi-
ficance is not known, though it may be con-
cerned with the position of the foetus in the
uterus. Some time during the night of February
6, a female infant was born to Mona, and when
first seen the next morning, she was swimming
between the dorsal fins of Mona and Spray. By
that time the dorsal fin of the infant was already
stiffened and upright, so that it could not be de-
termined to which side it had inclined. Also, the
placenta had been passed, and the baby was
suckling regularly. The infant showed five deep
transverse creases on the left side, six on the
right. All of these were much lighter than the
rest of the animal. The skin of Mona’s abdomen
showed the fine wrinkles which are character-
istic of the new mother.
Although “Mrs. Jones,” a mature female, was
not observed to spend more than a day or two
with the bull in the spring of 1952, abdominal
distension in November of that year made it
evident that she, too, was pregnant. On Feb-
ruary 14, 1953, she was reported to be strain-
ing and dilating her vaginal opening. During
this month she spent most of her time either
alone or with Pudgy. On three occasions before
the birth, Mrs. Jones showed flexion and on two
of these dilations were also observed. She showed
no concavity in the posterior abdominal area.
On February 23, she gave birth to a female
infant during the afternoon. When the infant
was first seen, shortly after 4 p.m., its dorsal
was inclined to the right. The mother showed
the characteristic wrinkling of the abdominal
skin.
On February 11 and 18, 1952, the bull’s
courtship activities were observed to be directed
toward “Susie,” another female, and she re-
sponded to his yelps. In November it was noted
that Susie was pregnant. On February 6, 1953,
the shallow concavity was seen on her abdomen.
This persisted until she gave birth. During the
latter months of her pregnancy, Susie was not
accompanied by another female, but swam alone
and avoided the company or contact of any other
animal. She flexed intermittently during the
month before parturition and her genital slit
dilated when she strained or when she broke
into a rapid swim. She was also seen to defecate
often and copiously during the last few days be-
fore birth.
Early on the morning of February 27, she
gave birth to a female infant, whose dorsal fin
was inclined to the right. Susie was not accom-
panied by any other female on the morning of
the birth in spite of attempts made by Pudgy to
remain near her, and she cared for the infant
without the aid of any other animal. This was
quite unusual, considering the conduct of all the
other mothers on which data are available.
Late in April, 1952, Pudgy was seen in the
company of the bull for several days. It was not
until April, 1953, that her pregnant condition
was suspected. Although she was the stoutest
animal in the group, her weight was well distrib-
uted over her entire body, and the roundness
of her abdominal region was not outstanding.
Her previous pregnancy had gone unsuspected
until May, 1950, when she gave birth to a male
infant. Therefore, when her size increased in
April, 1953, and especially when she avoided
the company of other animals, she was again
suspected of being pregnant, despite the absence
of other symptoms.
On May 5 and 6, Pudgy was seen in the com-
pany of Spray, but with no other animal. On
the 6th, she flexed a number of times. On the
morning of the 7th, she strained almost continu-
ously, and between flexions rose to the surface
where she hung, respiring frequently in a pattern
similar to that shown by Spray. At this time the
shallow concave area was seen on her abdomen.
During the afternoon of the 7th, Pudgy gave
1957]
Tavolga & Essapian: Behavior of the Bottle-nosed Dolphin
21
birth to a male infant. The birth began at 2:30
p.m. when two or three inches of the baby’s
flukes were seen to emerge immediately after a
strong flexion. At this time Mrs. Jones was swim-
ming beside Pudgy. The following outline, ex-
tracted from the notes taken on the day of the
birth, give a clear picture of the events during
this parturition.
2:30 p.m. Pudgy arches and recurves. Suddenly
there appear approximately 2-3 inches of
the baby’s flukes. Pudgy contracts abdomi-
nal muscles.
2:35 p.m. Pudgy swimming slowly in center of
tank. 3-4 inches of flukes showing. Flexion.
2:37 p.m. Flukes completely visible. Horizon-
tal, but tips curled ventrally.
2:38 p.m. Pudgy swimming close to floor of
tank with Jones. She flexes and recurves.
Other animals nosing around at distance
of 2-3 feet.
2:40 p.m. Pudgy almost scrapes the bottom with
her ventral side. Slows down till barely
moving.
2:43 p.m. 2 inches of caudal peduncle showing.
Pudgy maintains steady unhurried rate of
speed.
2:45 p.m. 3 inches of caudal peduncle emerged.
Jones with Pudgy.
2:47 p.m. Pudgy flexes. Spray swims just behind
and below Pudgy, her head pointing up-
ward.
2:50 p.m. 4-5 inches of caudal peduncle have
emerged. All other animals quiet. Pudgy
strains.
2:51 p.m. Foetus slips back an inch or two.
2:55 p.m. Foetus has turned, now lies on its
right side.
2:57 p.m. Pudgy swimming close to surface and
wall.
3:02 p.m. 5-6 inches of caudal peduncle have
emerged.
3:07 p.m. Feeding show in progress. All animals
rushing around.
3:08 p.m. Diver feeding underwater. All animals
feeding except Pudgy, who swims in cir-
cles away from points of disturbance.
Pudgy alone. Infant’s tail hangs limp.
3:15 p.m. Still 5-6 inches of caudal peduncle
showing.
3:22 p.m. 7-8 inches of caudal peduncle show-
ing. Pudgy swims slowly, flexes.
3:27 p.m. Foetus slides partially out and then
recedes as Pudgy flexes and relaxes. Pudgy
is alone, swimming short circles in N. E.
corner of the tank.
3 : 36 p.m. Pudgy now with Jones. Still 7-8 inches
of caudal peduncle are all that can be seen
of infant.
3:38 p.m. Pudgy flexes again, and recurves
strongly, swimming partially on her right
side. At this point, less than an inch of the
baby’s dorsal becomes visible, and in the
next 4-5 seconds, the foetus is expelled,
followed by a stream of blood.
Contrary to previous reports (McBride &
Kritzler, 1951) Pudgy did not whirl as the infant
was expelled, but continued swimming on and
up toward the surface. The umbilical cord broke
as it pulled taut. As in the case of Spray, all the
dolphins gathered at the point where the birth
took place, whistling and producing a clamor
of other noises. Mrs. Jones pushed the baby out
of the way of the other animals. Then she placed
her snout beneath it and gave it one thrust
toward the surface, where it took its first breath,
5 or 6 seconds after its birth. It is well known
that infant dolphins are able to reach the sur-
face without assistance immediately after birth
(McBride & Kritzler, 1951), and there is no
reason to suppose that this infant was an ex-
ception, especially as it was seen swimming
immediately after the nudge. As a normal infant,
it would undoubtedly have reached the surface
for its first breath unaided, and the push given by
Mrs. Jones was of secondary importance. Spray
positioned herself on the other side of Mrs.
Jones and the infant swam between their dorsal
fins. Pudgy trailing behind. This situation con-
tinued for the next two or three minutes, when
Pudgy moved into position beside the baby, and
Mrs. Jones dropped out. The infant was ener-
getic as it labored to keep pace with the adults.
The baby’s dorsal was inclined to the left. The
total time elapsed in this birth was 68 minutes.
Behavior of Mother and Infant
As stated by McBride & Kritzler (1951), and
confirmed in later cases by the present observers,
the newborn dolphin remains very close to its
mother during the first month. During the first
few days while its swimming is still uncertain,
particularly in regard to direction, the infant will
often stray slightly, but the mother invariably
swims toward it and pushes it gently, guiding its
direction until it is close to her side again, usually
next to her dorsal fin. This position close to her
dorsal fin is maintained by the infant during its
first month or two, and gradually the baby then
becomes accustomed to what the observers con-
sider the typical rest position, under the mother’s
tail, with the top of its head lightly touching her
abdomen. During the first few weeks, at least,
the infant sleeps in the position next to the
mother’s dorsal fin.
22
Zoologica: New York Zoological Society
[42:2
The mother does not allow the young infant
to come into close contact with other animals,
and when one approaches, will swiftly sweep the
young one away with her, usually placing it on
the side away from the intruder.
After about two weeks the infant begins to
swim about its mother and vary its position in
relation to her. It may dart around her head,
slip under her tail, or swim beside her at a dis-
tance of a foot or two. However, if the young
one increases the distance between them to as
much as ten feet, the mother will immediately
swim toward it and reorient it toward her. At
feeding time, the new mother will guide her in-
fant to a position a few feet from the feeding
platform, where she leaves it swimming in a
small circle. She will approach the feeding plat-
form and secure one fish, then rush back to the
baby. Often she will not approach the platform
again, but spend her time with the infant on the
side of the tank opposite to the feeding platform,
maneuvering herself and her infant away from
dolphins who race for food thrown on the sur-
face of the water. Because of this situation, spe-
cial feedings are often provided for new mothers,
who will accept fish thrown to them while swim-
ming with the offspring after the regular feedings
are over.
The four young ones born in 1953 prospered
and grew quickly. Nursing began promptly and
all the calves were able to find the nipples rapidly
within 24 hours after birth. They suckled in the
general pattern noted by McBride & Kritzler
(1951), three to nine times within as many
minutes, followed by an interval of 10 to about
25 or 30 minutes.
At about two weeks of age all the babies began
to move away from their mothers, and were re-
peatedly retrieved. As mentioned under Spray’s
behavior during pregnancy, she spent a good deal
of her time in the company of the young dol-
phins. From the time they were less than a month
old, she was seen swimming with them and their
mothers, and often swept away each of them with
her in a rapid trip around the tank. If any other
animal came close to one of the infants, the
mother promptly showed her displeasure by slap-
ping the intruder with her flukes, or by swiftly
removing the calf from his vicinity. However,
each mother soon allowed Spray to escort her
infant without evidence of concern, and Spray
usually returned the infant to its mother within
a minute or two.
When Spray had been accepted as a trusted
escort for the infants, all the mothers occasion-
ally left their calves in her company while they
were occupied at the feeding platform, and she
was often seen surrounded by two, three or four
youngsters at this time. In addition, one mother
could sometimes be seen guarding a group of
infants while the others fed (PI. Ill, Pig. 14) . The
mother who had received her food then returned
and relieved the escort so that she could get her
share. At the end of feeding time, each mother
retrieved her own calf, and there was never the
slightest hesitation about picking her own in-
fant from the group. No mother was ever seen
to make a mistake and choose the wrong infant.
By the time they were six weeks old, the young
dolphins were swimming away from their
mothers regularly, and associating with each
other or some of the adult animals. Their
mothers, by this time, had relaxed their vigilance
somewhat, and the infants were no longer kept
within a few feet at all times, although they were
still kept under close watch. A short excursion
or play period was usually followed by a longer
rest period spent under or close to the mother.
The infants began to show an interest in the
objects and fishes in the tank and were seen to
investigate, and if small enough toss about, any
new object which came to their attention. If the
object was a familiar one in the tank, a bit of
rock, a turtle, or a small fish, the mothers
watched nearby while the calf investigated, and
sometimes joined it. However, if the object was
an unfamiliar one, such as a new animal, or an
object dropped by a visitor, the mother displayed
agitation and set up a loud and continuous
whistling while steering the young one away,
and only when she had removed the baby from
the disturbing object did she resume her normal
behavior. This extreme vigilance and care were
typical of the dolphin mother during the first
months of her infant’s life, and continued in
somewhat lesser degree throughout at least the
first year. .
In direct contrast to the above situation. Spray 1
showed a distinct lack of care toward her infant
born in 1954. At first the situation seemed nor- *
mal. The baby began to nuzzle her mother’s ‘J
side, as stated above, five hours after birth, and f
it was 2 hours and 27 minutes after this initial ' |
effort that she began to suckle. After this, nurs- J
ing proceeded successfully, and the baby suckled
in the same general pattern as the above infants.
The baby seemed to be in good health and swam j
strongly from the first.
After a few days, however, it was seen that .(
she was not growing as other infants had done.
Newborn dolphins lose their thin appearance i
quickly, and after a few days their necks become j
less distinct and their abdomens become round-
ed. This was not true in the case of Spray’s in-
fant.
In addition. Spray did not care for the baby
1957]
Tavolga & Essapian: Behavior of the Bottle-nosed Dolphin
23
as vigilantly as other mothers had done. She
remained at the feeding platform throughout
almost the entire feeding, returning to the calf
only once or twice during this five- or six-min-
ute period. The infant was alone during this
time, except for the momentary visits of her
mother.
On March 13, Spray’s infant attempted for
the first time to swim away from her mother.
Spray and Mona swam after her, but did not
hurry to her side. This, too, was in contrast to
the conduct of other dolphin mothers. On
March 14, the baby persistently left Spray and
swam among the other animals. If she strayed
more than about twenty feet away. Spray
I brought her back, but if she stayed within this
distance, no attempt was made to restrict her
movements. Twice, however, when she strayed
to the other side of the tank. Spray went to her.
Then Spray turned on her back, positioned the
baby on her chest between her flippers, and rose
to the surface, pushing the young one out of
water. Other dolphin mothers have been seen
; to do this when their young infants swim eva-
sively away. The effect is to temporarily immo-
bilize the young one.
On March 16, Spray left the baby entirely
alone during the feeding. On this date, the in-
fant nuzzled its mother’s side immediately after
|i the feeding, but several minutes elapsed before
Spray slowed down to allow her to suckle. On
i' March 17, Spray allowed the infant to swim
around alone for several ten-minute periods. On
i that day, the baby appeared to be nursing almost
[i continuously, only very short intervals elapsing
! between periods of suckling.
On March 18, the baby left Spray several
? times, and sometimes entered another group of
' dolphins. Spray did not follow her, but allowed
her to work her way out of the group alone.
During the morning of March 19, the infant
i alternately remained close to Spray and per-
I sistently suckled, or wandered far away from
her, while Spray was occupied with other ani-
mals. At these times. Spray did not bring her
back or swim after her. The baby was seen to
suckle several times at 9:30 a.m.
At 12:15 p.m. it was reported to the observers
that the infant was in distress, and both hastened
to the tank. Unfortunately, by the time they ar-
rived, a minute or two later, the infant was dead,
and it was necessary to gather reports from eye-
witnesses, and to attempt to fit together what-
' ever facts could be supplied by them into a co-
herent account.
Early on the morning of the 19th, it was re-
ported by an employee of Marine Studios that
Spray and another unidentified dolphin, prob-
ably Mona, pushed the baby down to the floor
of the tank. By moving their bodies over her,
they temporarily prevented her from rising to
the surface. There was no apparent reason for
this action, and its significance is not clear. This
report was substantiated by two other employees
wno happened to be looking into the tank at the
time. Smce both observers had seen the infant
following this incident, and she appeared to be in
good condition, no particular importance was
attached to it at the time.
At about 12:06 p.m., the infant swam to the
surface to breathe, apparently under her own
power. Spray met her as she came down, and
the two rose to the surface again. Spray pushing
the young one up ahead of herself. As the baby
sank, the second time hubbies of air streamed
from her blowhole, and she did not swim again.
Spray pushed her about the tank, nudging her
up toward the surface every minute or so, but
the infant did not breathe again. The baby’s
mouth opened and her tongue hung out to the
side. The above account came from another
employee who was watching through the port-
hole at the time, and it was at this moment that
the authors arrived on the scene and it was ob-
served that the baby was dead. A diver entered
the tank immediately to bring her out. Spray
actively avoided the diver and pushed the dead
calf ahead of her, but the body was finally taken
from her and removed from the tank. From the
time the infant was first seen to be in distress to
the time when she was removed from the tank,
only 12 minues had elapsed. The body was taken
to the laboratory and an autopsy was performed.
This revealed that the calf was extremely thin,
her stomach was completely empty, and she had
suffered a fractured left mandible. Her lungs
contained water, and it was therefore assumed
that the immediate cause of death was drowning.
Judging from the extremely fresh condition of
the fracture, it is believed that the accident that
caused it must have taken place within 24 hours
of her death. At the time of her death at the
age of 15 days, the infant measured 42.5 inches
in total length, and weighed 30 pounds. Her
weight may have been slightly less than normal
for her age, since she was very thin. She pos-
sessed eight hair follicles on the left side of her
snout, and six on the right side, but only one
vibrissa was visible, in the last hair follicle on
the right side. No structural ahnormalities were
found at the autopsy.
Whatever the reason may have been. Spray’s
infant was not receiving the usual amount of
care that is given to newborn dolphins. Her
mother left her for unusually long periods of
time, and failed to remove her from situations
in which she might have been injured inadvert-
I
24
Zoologica: New York Zoological Society
ently by the other animals. On March 13, the
baby swam directly under the platform while
the other animals were feeding, and was struck
by at least one dolphin falling back into the
water. On March 18, while Pudgy and the bull
were engaged in some of the more vigorous
courtship activities, the baby swam between
them, and may have been lashed by a tail be-
fore she found her way out. In neither of these
instances did Spray make any attempt to rescue
her young one, and it is possible that the latter
incident was the cause of the infant’s fractured
mandible.
Two of the 1953 infants died before the end
of their first year, one by accidental drowning,
and the second because of an unknown infec-
tion, on which an investigation is being carried
out at the present time. The remaining two young
dolphins born in February, 1953, were weighed
and measured in September of that year, and
their lengths at that time were 68 and 69 inches,
their weights 146 and 141 pounds. At the pres-
ent writing they are 16 months old. They still
nurse regularly, usually after feeding time, al-
though it is thought that they are about to end
the nursing period, since they occasionally skip
a nursing. They began to take small pieces of
squid in August, 1953, and early in December
they were observed to swallow fish for the first
time. Since then both young dolphins have de-
veloped excellent appetites for fish and eat them
regularly at feeding times. The nursing period
usually ends gradually over a period of months
as the young animals show an increasing prefer-
ence for fish. They race the adults for the fish
even after their appetites have been satisfied.
They have often been seen to rush for a fish
thrown to them, then drop it to race after an-
other on the other side of the tank. They seem
to make a special effort to take fish from just
in front of another animal, and often do so.
They may play with the fish so taken in a num-
ber of ways. They will drop a dead fish in front
of the rocks where the small tank inhabitants
hide, and just as they come out to eat, the
young dolphin will snatch the fish away again,
only to repeat the performance a minute later.
The young animals indulge in this type of sport
only after they have eaten and are satisfied, but
often the fish are caught by a still hungry adult
and the youngsters are deprived of their play-
thing. Every activity of the young animals now
in the tank shows them to be normal healthy
dolphins.
Discussion
The dolphins are a specialized group of mam-
mals adapted in both structure and function to
a life lived completely in the water. Superficially,
[42:2
they resemble fish. Their forelimbs are flippers,
without use as grasping organs, and probably
serve mostly as balancers, or in steering. They
possess no hind limbs. Their tails, furnished with
broad horizontal flukes, are excellent propulsive
organs. Their streamlined form enables them to
move gracefully and rapidly through the water.
Since they were derived from land mammals,
they bear their young alive in typical mammalian
fashion, and suckle them for an extended period
of time. They also display mammalian care for
the young. Since they possess no limbs which
can be used as manipulative organs, their mouths,
which are provided with a set of efficient teeth
for holding their prey, serve also to grasp ob-
jects and to inflict wounds in their own defense.
The powerful muscles of the tail also aid in
defense.
Together with these features, the dolphin pos-
sesses a highly developed and exceptionally large
brain, of which the largest part is cerebrum. The
brain of one nine-months-old Tursiops weighed
1230.4 grams, and its volume was 1225 cc. The
length of this male specimen at the time of its
death was 64 inches, and its weight was 137
pounds.
The large brain and the probable resulting
high level of adaptiveness might be expected to
lead, in animals with manipulative limbs, to
complex varieties of behavior. The lack of these
limbs results in the ingenious use of flukes and
flippers. These can be delicately adjusted to pro-
duce minute changes in movement and position.
Such changes enable the animal to make the
best possible use of the only grasping organ it
possesses, the mouth. The remarkable use of
these organs under the control of a well-devel-
oped central nervous system leads to behavior
patterns which are peculiar to these animals and
their close relatives.
No accurate observations on the precopulatory
and copulatory behavior of dolphins were pos-
sible before the tanks at Marine Studios were
established, and therefore such observations as
were available were recorded by whalers and
transmitted to interested scientists, or such sci-
entists reported their own observations from the
decks of ships. Lillie (1910) reported that whal-
ers saw two adult sperm whales (Physeter) rush
together near the surface, then turn up ver-
tically at the end of the rush. They believed that
they had seen copulation, and said that the males
were exhausted and easy to capture after these
activities. Morch (1911) observed coition in blue
whales during the summer, but gave no details
of their behavior. Tomilin (1935) noted what
he considered to be sexual attachment in various
species, and differences between the species and
1957J
Tavolga & Essapian: Behavior of the Bottle-nosed Dolphin
25
between individuals in the same species. He
stated that this attachment sometimes reached
great intensity and was strongly shown especially
It one of a pair was wounded. He gave no details
of copulation. Hamilton (1945) reported that
while sailing off the coast of Brazil, he saw two
Steno swimming on their sides with their genital
areas pressed together. They were slightly sep-
arated anteriorly, and one’s mouth was slightly
open. This posture was maintained while they
remained in sight, which, according to the au-
thor, was “not for very long.” The ship was
moving at 10 knots, and the animals maintained
the speed of the ship while in the above position.
Such fragmentary observations do not give
any indication of the complex pattern of pre-
copulatory and copulatory behavior in this group
of animals and it was not until they could be
observed under water for extended periods of
time that details could be reported. Although
many of the events leading to copulation are
reported here, the sequence is variable, and it is
by no means certain that copulation can be pre-
dicted following any particular stage. The ob-
servers have seen much copulatory behavior,
some of it at least partially successful. In at least
one instance an ejaculation of semen has been
seen after withdrawal of the penis from the
vagina of the female. Sudden erection and re-
traction of the penis occur most often during
the spring months both with and without true
copulatory activity, so that retraction is no indi-
cation that ejaculation has taken place during a
partial intromission. The one instance in which
full penetration took place occurred before seven
in the morning and was not seen by the authors.
The employee who witnessed it is a reliable ob-
server who has made a hobby of watching the
animals over a long period of time. It is not
known whether the copulations involving partial
penetration over a short time are as effective as
those with full penetration extending over 30
seconds. Although the authors have spent a
number of hours watching during the evening
hours and also in the early morning, they have
not seen the longer copulations. Since there is no
provision for adequate lighting at night, ob-
servations during this time are impossible. Per-
haps long-continued early morning observations
over a long period of time will provide more
data.
Although McBride, in 1940, recognized the
jaw clapping of the bull as “the principal form
of intimidation,” the entire use of this sound was
not recognized until much later. As late as 1948,
McBride & Hebb again stated that this sound
was used by a dominant animal toward sub-
ordinates. The observations of the present au-
thors indicate that this is not always the case.
and the jaw clap has, under direct observation,
been made or attempted by almost every animal
in the tank. It has occurred under a number
of circumstances but these were always such
that the jaw clapping animal appeared to be
displeased, dissatisfied or annoyed. Pudgy clap-
ped her jaws when the bull, who is dominant
to her, made sexual advances to her and she
was not responsive. All the mothers clapped
their jaws when another animal interfered with
or approached their young. Spray has been seen
to clap her jaws when the young male swam
between her and the infants in her company.
The younger animals, such as Spray and the
young male, produce a much less resounding
clap than Pudgy and the bull, but their efforts
were unmistakeable, and as they gained more
experience and practice, the sound produced be-
came louder. Apparently this evidence of dis-
pleasure is limited to the adults, since the 1953
youngsters have not as yet attempted it.
Townsend (1914) gives some evidence to
show that Tursiops breeds yearly in the wild. Of
51 specimens taken in November, 34 were fe-
males. All the females taken were in milk at that
time. Although he does not state how many of
the females were killed, all those that were killed
were not only in milk but contained young. Un-
fortunately, the age of the suckling infants could
not be determined. Therefore, it is not known
whether all infants in the wild suckle for 18 to
21 months or more, or whether there is a yearly
reproductive cycle. Since young dolphins at Ma-
rineland have been known to suckle for periods
up to 20 months, it is possible that Townsend’s
data would support the idea of a two-year cycle
in the wild.
At Marine Studios, no female has as yet pro-
duced young two years in succession. Several
animals have produced young at two-year in-
tervals. There is in the tank only one fully adult
male, and therefore no significant competition
for females. There is the possibility that females
unaccompanied by newborn young are prefer-
red partners. It is to be hoped that some future
study may show whether the two-year cycle seen
at Marine Studios is unchanged from the wild
condition.
McBride & Kritzler (1951) believed that the
female Bottle-nosed Dolphin becomes sexually
mature at four years of age and may bear her
first infant in the spring of her fifth year. Al-
though this may be theoretically true, the only
evidence now at hand indicates that the first in-
fant may not be conceived until several years
later. Spray, the only animal that was born and
reached sexual maturity in captivity, did not
conceive until she was six years of age, and
26
Zoologica: New York Zoological Society
[42:2
delivered her first infant just after her seventh
birthday, although she was known to be in ex-
cellent condition throughout her life. It is, of
course, not known whether she would have con-
ceived earlier in the wild, and further evidence
on the age at which sexual maturity is reached
will depend on the time at which the young
females now in the tanks first conceive.
There has been much speculation on the length
of the gestation period in these animals, and
estimates which have run from 10 to 14 months
have in the last few years narrowed to 11 to 13
months. It is now possible to state with a reason-
able degree of accuracy that the gestation period
is very close to 12 months. Spray’s association
with the bull lasted for a period of two weeks,
from February 23 to March 9, 1953, and her
infant was born on March 4, 1954. Similarly,
the associations of Mona and Susie with the bull
were almost exactly a year before the birth of
their young ones. Although Pudgy was observed
to be in the company of the bull in late April,
1952, and her infant was born in early May of
the following year, the authors believe that this
discrepancy comes within the limit of error, and
that the twelve-month period is the most valid
one.
McBride & Kritzler (1951) noted the with-
drawal from association with other animals
shown by pregnant females, especially during the
latter part of pregnancy. They also mentioned
the tendency of two pregnant females to associ-
ate with each other. These tendencies have been
confirmed in the present paper. In 1952, Pudgy
and Mrs. Jones, although they conceived several
months apart, spent most of their time together
during their pregnancies, though they had not
been close companions before this time. When
Pudgy’s infant was born, Mrs. Jones was the
animal which escorted the infant during the first
few minutes of its life, and she and Spray kept
the infant out of the way of other excited ani-
mals, and accompanied Pudgy and the new baby
for the first few weeks. When Spray’s infant was
born it was Mona, her mother, and Pudgy who
performed these functions alternately, so that
there was almost always another mature female
swimming beside the baby. Such protection
proved to be important, since the records show
that the birth of a young one seems to stimulate
the adult male to begin or renew his courting
activities. Often these are directed toward the
new mother, whose evasive movements become
frantic in her efforts to protect her newborn
infant. In 1947, when Spray was born, there
were several adult males in the tank, and in their
attacks on the new mother and other females
and each other. Spray’s skin was badly lacer-
ated by their teeth. Since then, these competing
animals have been removed, but the danger to
the new infants still exists, though to a lesser
degree.
In one case, however, the new mother was not
accompanied by another female. Susie is an ex-
tremely retiring animal, and remained alone
from the time she was first captured and placed
in the tank. She conceived in February, 1952,
and remained alone throughout her pregnancy,
a situation which was unusual enough to war-
rant the attention of the observers. When her
infant was born in February, 1953, she was not
accompanied by any other animal. Immediately
after the birth Pudgy made persistent efforts to
swim with her and remain near her, but Susie
avoided Pudgy as diligently as she did all the
other animals. Her behavior, though unusual,
was consistent, since both before and after her
pregnancy she showed no inclination to asso-
ciate with any other dolphin. She was captured
in 1949 and gave birth three days later to a male
infant, an event which took place in the receiv-
ing tank. This tank contains very shallow water,
and the environment was quite unnatural to her.
Her timidity may have stemmed from that time.
The young male has now almost reached ma-
turity, and the two ceased their association prior
to the arrival of the new infant. It took more
than a year of training before Susie took her
food at the feeding platform. Either her inherent
timidity or the shock of her arrival and the sub-
sequent birth may account for her unusual
conduct.
The young of members of the Delphinidae are
usually born tail first. In every birth at Marine
Studios except one (Spring, 1955) the infant
emerged in this manner.
In the earliest papers describing the birth of
small cetaceans, (James, 1914), the author as-
sumed that the normal presentation was head
first, and that it was mishandling or other im-
proper treatment which caused the tail presen-
tation. The birth described by James took place
in Phocaena phocaena and was a stillbirth.
James mentioned mishandling as the reason for
the tail presentation, and the tail presentation as
the reason for the stillbirth. It is now known
that although mishandling my lead to stillbirth,
the reasons are other than those described above.
Caudal presentation has been described in de-
tail by a number of authors (Wislocki & Enders,
1941; Slijper, 1949). McBride & Kritzler (1951)
discussed it in connection with data available to
them from Marine Studios, and all agree that
the foetus is, or becomes, oriented in such a way
that the head is pointed toward the tubal end of
the uterine cornu, while the tail extends toward
1957]
Tavolga & Essapian: Behavior of the Bottle-nosed Dolphin
27
the vaginal opening. Sleptzov (1940) observed
635 embryos of Delphinus delphis in situ during
various stages of development, and came to the
conclusion that during early embryology the
foetus may lie in either direction. Later the head
tends to become oriented toward the tubal end of
the cornu, and by the time reversal is no longer
possible, because of size, the vast majority of
embryos are situated in this way, so that tail
presentation is inevitable.
The problem remains, however as to why the
orientation takes place in this manner. Slijper
(1949) , citing Williams & DeSnoo, discusses the
dangers attendant upon breech presentation in
mammals which produce foetuses with com-
paratively large heads, and believes that the com-
pression of the umbilical cord between the pel-
vis of the mother and the head of the infant
which occurs in breech presentation may cause
asphyxia if the infant is not quickly extracted.
Although Odontocetes possess no pelvic girdle
other than two small pelvic bones, the pelvic
region contains large masses of tough fibrous
connective tissue which could exert considerable
pressure, and it is possible that such compression
may be exerted in the case of these animals.
Whether or not this is true, it does not explain
the orientation of the foetus in the uterus, but
only illustrates an effect of such orientation.
Slijper’s discussion of the role of uterine con-
tractions in the orientation of the foetus is more
pertinent to the question. Mammals other than
primates show, during pregnancy, uterine con-
tractions of a peristaltic nature which begin at
the tubal end of the uterus, and Slijper believes
that the head should be impelled in the direction
of the cervix under the influence of these con-
tractions, in mammals with a small head and
long neck. Since this formation does not apply
to cetaceans, he considers that the body form of
the foetal cetacean might favor caudd presen-
tation.
Schumann (1914) discusses the possible
causes of almost universal cephalic presentation.
He states that in multiparous animals the foetuses
lie indifferently, in either breech or cephalic
presentation, whereas in unipara, the head pre-
sents in 90 to 98 percent of the cases. In man the
foetal head is heaviest and gravitates to the most
dependent position, nearest the cervix. In uni-
parous quadrupeds, however, the hindquarters
and trunk of the foetus are its heaviest parts,
and consequently sink to the lowest portion of
the uterus, which with its cornua lies low in the
abdominal cavity. The cervix rises to meet the
vagina.
In dolphins, which are uniparous, the uterus
lies in a situation similar to that seen in the
quadrupeds, so that the heaviest portion of the
foetus would tend to gravitate to the lowest por-
tion, the tubal end of the cornu. These animals,
however, possess no hind limbs, only remnants
of the pelvic girdle, and the tail is long and
slender. The head, which is relatively very large,
and the pectoral region are heavier than the
hindquarters. Under these circumstances, the
gravitation of the heaviest portion toward the
tubal end of the uterus would result in tail
presentation. The present authors believe that
this situation may be of considerable influence
in the determination of tail presentation in the
dolphins.^
It is necessary, however, to consider the fold-
ing of the uterine cornu in evaluating this ex-
planation. The non-pregnant dolphin uterus is
not folded. Presumably the fold occurs when the
foetus is too long to extend to its full length in
the abdomen of the mother. At this time it
would also be too large to turn easily on its own
axis. Examination of the pregnant uterus of
many delphinids has shown that the foetus ap-
pears, at first glance, to have its head oriented
toward the vaginal end of the uterus. Closer ob-
servation, however, has usually shown that the
head is enclosed in a fold, which is oriented
toward the caudal end of the mother, but that on
following this fold in the direction in which the
head is pointing, it is found that the uterus con-
tinues from this point toward the Fallopian
tubes. Following the body of the foetus in the
opposite direction reveals that it folds close to
the origin of the tail, and the tail occupies the
lower end of the uterus, and extends toward the
vaginal opening.
These facts may seem to contradict the weight
theory outlined above, but if we consider that the
head of the foetus, as in most mammals, devel-
ops more rapidly than the other parts, and be-
comes the heaviest part of the foetus in an early
stage of development, before the fold occurs,
it seems possible that both events might occur
in sequence, with the usual observed result of tail
presentation. It is unfortunate that Sleptzov
(1940), who had at his disposal more than six
hundred foetuses in all stages of development,
did not record more fully the sizes, weights and
developmental characteristics of these animals,
together with the condition and configuration of
the uteri in which they were found.
Although McBride & Kritzler (1951) stated
that in all cases observed by them the dorsal fin
of the newborn dolphin was folded to the right,
^Recently E. J. Slijper (1956. Some remarks on
gestation and birth in cetacea and other aquatic mam-
mals. Hvalradets Skrifter. Scientific Results of Marine
Biological Research, No. 41, pp. 1-62) reported addi-
tional data and conclusions concerning tail presenta-
tion in cetacea which confirm those given above.
28
Zoologica: New York Zoological Society
[42:2
at least two cases were seen by the present ob-
servers in which the dorsal was folded to the
left. In one of these cases the infant was seen
to be lying on its left side when only partly ex-
truded from the mother’s body. We do not know
whether this correlation holds true in all cases.
The symmetrically placed vertical creases on
each side of the trunk at birth are not neces-
sarily six in number, as stated by McBride &
Kritzler. In the group of births recorded here,
the creases have numbered five, six or seven, and
there were sometimes more on one side than on
the other. Also, one or more may be continuous
over the dorsal part of the animal. They lacked
pigment and were easily visible for several weeks
after birth, when they gradually acquired pig-
ment and disappeared. The young dolphin to
which Spray gave birth, and which died at 15
days of age, showed two creases which were not
continuous over the dorsal side but which ap-
peared on both sides, three which were con-
tinuous over the dorsal, and two ventral creases,
which were not seen previous to her death be-
cause of the lack of pigment on the ventral side
of the body.
It may be significant to note that the dorsal
fin of the infant which died at 15 days was
slightly inclined to the right when the body was
removed from the tank. At birth it was distinctly
folded to the left. It has been noted by many ob-
servers that the dorsal fins of all the adults in-
cline to the right, and it is thought that the
swimming tendencies of the animals may be the
cause of this phenomenon. Since the animals in
the tank usually swim in a clockwise direction
through a counterclockwise current, the force
of the water against their dorsal fins may be the
cause. It should be noted, however, that this
inclination in no way hampers their swimming
in any other direction. It seems significant that
the tilt to the right should have taken place so
quickly in an infant whose dorsal was inclined
sharply in the opposite direction only 15 days
before.
The mother of a newborn dolphin has never
been seen to eat the placenta, which is expelled
several hours after birth. Since eating this organ
appears to play a significant role in the behavior
of so many other mammals, this point seemed
to be of particular interest. In all the cases on
record at Marine Studios no mother dolphin has
even approached the placenta after expelling it,
and unless removed by divers and preserved, the
organ is torn to shreds by the turtles and fish in
the tank.
Although McBride & Kritzler (1951) reported
that a mother dolphin, immediately after the
infant’s body falls free, will whirl about and
thus snap the umbilical cord, this does not ap-
pear to be necessary in every case. At least one
female in the present group spurted forward in
a straight fast swim as the baby was released and
the cord snapped quickly as it pulled taut. The
survival value of the whirl, in case the infant
fails to reach the surface, is obvious, but the
pattern does not seem to be an invariable one.
The problem of suckling in the dolphin has
long been of interest to observers, and the mech-
anism has been fairly well worked out. Milk is
poured from the mammary glands into large
sinusoids, which in turn open to the nipples,
situated on each side of the genital slit. When
the infant grasps the nipple, abdominal contrac-
tions of the mother pour the secretions into the
baby’s mouth. The question remained as to
whether the stimulation of the nipple by the in-
fant was necessary in order to produce a flow of
milk, or whether the contraction of the ab-
dominal muscles was under the voluntary con-
trol of the mother. An instance that took place
in the spring of 1954 shows that, at least in ex-
traordinary circumstances, the flow of milk is
entirely under the mother’s control. On February
22, 1954, a nine-months-old young dolphin,
born the previous May, died at Marine Studios
of an unknown skin infection. At the time of his
death, he was still getting the main part of his
food from his mother, and was only beginning
to show an interest in other food. On the day
before his death, he suckled very few times, and
his mother’s nipples protruded from their slits,
apparently pushed outward by the engorged
sinusoids. On the day of his death and for sev-
eral days afterwards this condition continued.
Twice in the first several days after the young
one’s death, the mother was seen to turn on her
side in the water and visibly contract her ab-
dominal muscles, raising her tail as she did so.
As she did this, a stream of milk spurted from
her nipples. This stream was ejected with a
force strong enough to send it a distance of
more than two feet before it dissipated into the
water. It is not known whether such a feat can
be accomplished in circumstances where the
sinusoids are not filled to excess, but it shows
that a female dolphin can eject milk without
stimulation of the nipple under certain con-
ditions.
Several Russian authors, particularly Khvatov
(1938) and Sleptzov (1940) have reported on
births of Delphinidae (mainly Delphinus del-
phis) which they observed in the Black Sea. In
most of these births, the animals concerned
were captured dolphins, and were seen while
in seines being drawn toward shore. The rest
were in shallow areas near shore. It is possible
1957]
Tavolga & Essapian: Behavior of the Bottle-nosed Dolphin
29
that the births may have been abnormal in some
cases, particularly in the case of the captured
animals, which could have been shocked and
frightened by the procedure of capture. Also the
conditions for observing the births were un-
favorable, since many animals were milling
about in the seine, and in all cases the births
were observed from above the surface of the
water. In spite of these difficulties, there are
descriptions of reverse flexions, leaps, dives, and
in some instances, the moment of birth when the
infant fell free. In all cases the infant was born
tail first, but no description of the moments
immediately following birth is recorded. In the
most complete description, the birth took one
hour and fifteen minutes from the time the
author (Sleptzov, 1940) first saw the tail of the
infant protruding from the mother, and the in-
fant’s dorsal fin was folded to the right. The
general description of the birth is similar to
those seen at Marine Studios. Khvatov, after see-
ing an apparently abnormal birth, observed an
infant attached to its mother by the umbilical
cord, and postulated that this was a normal con-
dition. He thought that the infant suckled while
still attached to the mother. In the specimens
examined at Marine Studios, no umbilical cord
was found to be long enough to permit such a
procedure. Whether the umbilical cord of the
species he studied was sufficient in length is not
known, but Sleptzov, after seeing what he be-
lieved to be the same species, thought Khvatov’s
theory erroneous. He observed a larger number
of births than did Khvatov, and in each instance
the infant separated from the mother promptly
with no apparent ill effects to either. All evi-
dence from births in the wild indicates that they
parallel closely those seen at Marine Studios.
The reaction of Spray toward her first infant
presents an interesting problem. She was seven
years of age when she gave birth to this calf. It
has been thought that dolphins mature sexually
at an earlier age, and therefore it is doubtful that
she was too young to care for it. She had been
active for several years in escorting and protect-
ing other young dolphins in the tank and it
seems reasonable to assume that therefore she
had sufficient general background experience.
However, never before had she had the constant
care of an infant, and she had not suckled an
infant before. Although the immediate cause of
the infant’s death was drowning, it is not certain
whether the baby drowned because of a shock
reaction to the fractured mandible, whether it
starved until it was too weak to swim, or whether
there was another reason, undiscernible to the
investigators. The observers could find no ab-
normalities in the digestive tract to account for
its thinness or its inability to suckle. Young
dolphins use their tongues, rather than their
lower jaws, in suckling. The youngster was quite
obviously hungry, judging from the number of
times it approached its mother’s mammaries,
and its persistent nuzzling there. It was im-
possible to determine whether Spray produced
sufficient milk as she could not be removed from
the tank for examination. Her mammary glands
remained distended for several days and gradu-
ally receded over a period of several weeks. It
is possible that more evidence may be found
when Spray gives birth to a second infant and
her reactions to it can be observed.
Although this report contains many data that
have not been previously presented, it is ex-
tremely difficult to quantify, since records of the
earlier births, both by McBride and his co-
workers and by the present investigators, did not
contain all the details which are now con-
sidered essential to an analysis of the behavior
of these animals. In addition it is probable that
further details, up to now unrecorded, will later
present themselves. Further observation by ex-
perienced investigators and the recording of
seemingly non-essential details will produce a
more complete picture, and one that will allow
a more accurate comparison with other
mammals.
Summary
The group of captive Bottle-nosed Dolphins
(Tursiops truncatus) at Marine Studios is an
actively reproducing colony. Mating and par-
turition take place from February to May,
usually in a two-year cycle. The report com-
prises the data on five pregnancies and births,
one of which was observed from conception.
Precopulatory behavior consisted of a pro-
longed companionship between a male and fe-
male. There followed such activities as postur-
ing, stroking, rubbing, nuzzling, mouthing, jaw
clapping and yelping on the part of the male.
These occurred in any sequence and if the fe-
male responded, copulatory behavior followed.
This included erection, approach, intromission,
ejaculation and withdrawal. Most of the copula-
tory activity took place at night or in the early
morning, although some of the activities were
seen at all hours during the day.
The gestation period was about twelve
months. After about four or five months, the
pregnant female tended to withdraw from the
society of other dolphins, and usually estab-
lished at this time the one or two associations
which she maintained throughout the remainder
of her pregnancy. The rest of her time was spent
alone. Her activities gradually diminished in
vigor, and as term approached she became slow
and sometimes clumsy. Toward the end of preg-
30
I
[42:2
Zoologica: New York Zoological Society
nancy a labored respiratory pattern was seen,
yawning and body flexion appeared, and defeca-
tion increased in frequency.
The infants were born tail first. The mother
either whirled or swam rapidly ahead, and the
umbilical cord broke as it pulled taut. The un-
pigmented creases on each side of the body of
the infant were not symmetrical in every case,
and sometimes there were more on one side
than on the other. The infants swam from birth.
The dorsal fin was folded sharply either to the
right or left, but stiffened in an upright position
within a few hours. The placenta was expelled
several hours after birth. The mother did not
eat it, nor did she approach it again.
Dolphin mothers kept close to their infants
at all times for the first months, and removed
them promptly from dangerous situations. Dur-
ing the first weeks, the infants were not allowed
to stray more than ten feet from the mother.
Nursing was established during the first 24
hours. The infants were weaned at approxi-
mately 18 months.
One female dolphin, born and raised at Ma-
rine Studios, did not care for her infant as care-
fully as the other mothers. She left the baby
alone at feeding time and for other short periods.
She did not remove it from contact with other
animals or from dangerous situations. The infant
died at 15 days of age.
Literature Cited
Essapian, F. S.
1953. The birth and growth of a porpoise.
Natural History, 62 (9): 392-399.
Hamilton, J. E.
1945. Two short notes on Cetacea. 1— Coitus—
Steno rostratus. 2— Function of the flippers
and tail— Balaenoptera and Orcinus. Proc.
Zool. Soc. London, 114: 549-550.
James, L. H.
1914. Birth of a porpoise at the Brighton
Aquarium. Proc. Zool. Soc. London,
1914: 1061-1062.
Khvatov, J. P.
1938. New information on the durability of re-
tention of the corpus luteum in the dol-
phin. Biol. Med. Exper. U.R.S.S., 5: 27.
Lillie, D. G.
1910. Observations on the anatomy and general
biology of some members of the larger
cetacea. Proc. Zool. Soc. London, 1910:
769-792.
Lutken, C.
1888. Was die Gronlander von der Geburt der
Wale wissen wollen. Zool. Jahrb., Abth.
Syst., 3: 802.
McBride, A. F.
1940. Meet Mr. Porpoise. Natural History, 45:
16-29.
McBride, A. F. & D. O. Herb
1948. Behavior of the captive bottle-nose dol-
phin, Tursiops truncatus. Journ. Comp,
and Physiol. Psychology, 41(2): 111-123.
McBride, A. F. & H. Kritzler
1951. Observations on pregnancy, parturition,
and post-natal behavior in the bottlenose
dolphin. Journ. Mammal., 32: 251-266.
Morch, j. a.
1911. On the natural history of the whalebone
whales. Proc. Zool. Soc. London, 1911:
661-670.
Pedersen, A.
1931. Fortgesetzte Beitrage zur Kenntnis der
Saugetier- und Vogelfauna der Ostkiiste
Gronlands. Ergebnisse einer zweijahrigen
zoologischen Untersuchungreise in Ost-
gronland. Medd. Gronland, 77: 343-507.
Schumann, E. A.
1914. The mechanism of labor from the stand-
point of comparative anatomy. Amer.
Journ. of Obstetrics, 69: 637-658.
Sleptzov, M. M.
1940. On some particularities of birth and nutri-
tion of the young of the Black Sea por-
poise Delphinus delphis. Zoologich Zhur-
nal, 19: 297-305.
Slijper, E. j.
1936. Die Cetaceen — vergleichend-anatomisch
und systematisch. Capita Zoologica— vol.
VII. Die Lage des Embryo bei das Geburt
— Kapitel 16, ss. 455-465. (Published by
Martinus Nijhoff, The Hague, 1936).
1949. On some phenomena concerning preg-
nancy and parturition of the Cetacea.
Beijdragen tot de Dierkunde, 28: 416-446.
Tomilin, a. G.
1935. Maternal instinct and sexual attachment
in whales. Bull. Soc. Nat. Moscau, 44:
351-361.
Townsend, C. H.
1914. The porpoise in captivity. Zoologica, 1
(16): 289-299.
WiSLocKi, G. B. & R. K. Enders
1941. The placentation of the bottle-nose por-
poise {Tursiops truncatus). Amer. Journ.
Anat., 68: 97-125.
Wood, F. G., Jr.
1953. Underwater sound production and con-
current behavior of captive porpoises
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3: 120-133.
19571
Tavolga & Essapian: Behavior of the Bottle-nosed Dolphin
31
EXPLANATION OF THE PLATES*
Plate I
Fig. 1. Young male dolphin (Algie) above, and
adult female (Mona) below. The male
strokes the back of the female with his
flippers.
Fig. 2. Young male and female during rubbing
activity. The two have approached each
other head-on and their bodies will rub
together (Algie and Spray).
Fig. 3. Young male and female during rubbing
activity (Algie and Spray).
Fig. 4. Male and female during mouthing.
Fig. 5. Adult male (Happy) and young female
(Spray) in copulation. Note partial inser-
tion of penis into vaginal opening.
Plate II
Fig. 6. Adult female (Susie) in non-pregnant
state.
Fig. 7. Adult female (Susie) in late pregnancy.
Fig. 8. Adult female (Mrs. Jones) in late preg-
nancy, in body flexion.
Fig. 9. Adult female (Mona) in late pregnancy.
Recurve of body flexion.
Fig. 10. Female in late pregnancy. Recurve of
body flexion while rubbing over rocks.
Note vaginal distension and protruding
nipples.
Fig. 11. Birth in progress. Flukes and part of
caudal peduncle are seen projecting from
vagina of mother.
Fig. 12. Birth in progress. Taken just before com-
plete emergence. Note umbilical cord, also
presence of companion female on the left.
Plate III
Fig. 13. Suckling. Young female (Spray) and in-
fant (Peggy) a few days after birth.
Fig. 14. A new mother (Pudgy) escorts her own
infant and two others during feeding pe-
riod while mothers of the other two feed.
♦Photographs by F. S. Essapian.
9
II
liTAVOLGA a ESSAPIAN PLATE I
FIG. 5
THE BEHAVIOR OF THE BOTTLE-NOSED DOLPHIN (TURSIOPS
TRUNCATUS); MATING, PREGNANCY, PARTURITION AND
MOTHER-INFANT BEHAVIOR
FIG. 12
THE BEHAVIOR OF THE BOTTLE-NOSED DOLPHIN (TURSIOPS
TRUNCATUS): MATING, PREGNANCY, PARTURITION AND
MOTHER-INFANT BEHAVIOR
TAVOLGA a ESSAPIAN
PLATE III
FIG, 13
FIG. 14
THE BEHAVIOR OF THE BOTTLE-NOSED DOLPHIN (TURSIOPS
TRUNCATUS); MATING, PREGNANCY, PARTURITION AND
MOTHER-INFANT BEHAVIOR
3
A Study of the Relationship Between Certain Internal and External
Morphological Changes Occurring During Induced and Natural
Metamorphosis in Rana pipiens and Rana catesbeiana
Beulah Howatt McGovern^ & Harry A. Charipper
Department of Biology, Graduate School of Arts and Science, New York University
(Text-figures 1-12)
The dramatic metamorphosis of the
tailless Amphibia is the subject of an
extensive literature resulting in large
part from Gudernatsch’s (1912, 1914) accelera-
tion of the process by thyroid feeding, and from
the subsequent discovery of the thyroid-pitui-
tary relationship by Adler (1914), Allen (1916)
and Smith (1916a). A chronology of the ex-
ternal metamorphic events was established by
Etkin (1932), who also showed (1935) that
thyroxine treatment did not change the order of
events but did change their spacing. Knowledge
of internal metamorphosis both normal and ac-
celerated is quite complete, with many excellent
and detailed studies of one, two, or more in-
ternal organs. Elowever, there is no single analy-
sis of internal changes comparable to Etkin’s
for the external pattern. Disharmonies in de-
velopment during accelerated metamorphosis
have frequently been noted, and several theories
of the control of the sequence and spacing of
metamorphic events have been reported (see the
review of Lynn & Wachowski, 1951 ) .
During normal metamorphosis, the rate and
extent of change in the several organs are so
correlated that a given degree of change in an
external organ, such as the developing hind limb,
is a measure of the change to be expected in-
ternally (Allen, 1929). Although the available
evidence indicates that this is not true during
accelerated metamorphosis, more information as
to the degree of deviation from the normal corre-
lation between internal and external changes in
accelerated metamorphosis seemed desirable.
1 Submitted in partial fulfillment of the requirements
for the degree of Doctor of Philosophy in the Graduate
School of Arts and Sciences, New York University,
New York, N. Y.
Therefore, a comparison of the development of
several internal organs in normal and thyroxine-
treated animals at similar external metamorphic
stages was undertaken. Rana pipiens and Rana
catesbeiana were both studied in order to obtain
differing lengths of larval period for the purpose
of comparison.
Grateful acknowledgement is made to Drs.
E. G. Reinhard and W. Gardner Lynn of The
Catholic University of America; to the former
for interest in the problem and generosity in ex-
tending facilities for research, and to the latter
for valued discussion and advice.
Materials and Methods
1 . Plan of the Study
Rana pipiens and Rana catesbeiana were the
species chosen for study because of the difference
in the lengths of their larval periods.
To compare the integration of internal and
external changes in normal metamorphosis with
that in accelerated metamorphosis, larval stages
were selected at which the external morphology
of the normal and treated animals was considered
comparable. Normal and accelerated animals
were killed at each of these stages and certain
of their internal organs compared. Because of
the occasional rather substantial variation in age
between normal and accelerated tadpoles at ap-
parently the same external metamorphic stage,
untreated tadpoles of the same age as the ac-
celerated were also killed and studied con-
currently.
To select accelerated stages most comparable
morphologically to normal stages, the effects of
the various thyroxine concentrations were ob-
served. It was concluded that only the forelimb
emergence stage was recognizably similar to the
33
34
Zoologica: New York Zoological Society
[42:3
normal in all series, and therefore a more com-
prehensive study was made of this stage. Earlier
accelerated stages in certain respects overlapped
several normal stages. Although accelerated ani-
mals were studied at a stage later than forelimb
emergence, this later stage has been named
“terminal stage” rather than “taU stub” stage
as in the normal series.
The following stages were studied:
(1) . Initial stage (normal animals, 12 and 21
days old)
(2) . Two mm. hindlimb stage {R. pipiens
only)
(3) . Forelimb emergence stage (both fore-
limbs emerged)
(4) . Tail stub (terminal stage)
The effects of rate of acceleration on meta-
morphic integration were studied by the use of
two aqueous thyroxine concentrations:
1 ; 10,000,000 and 1 : 100,000,000. Each has been
used previously (e.g., Allen, 1932; Etkin, 1935).
In addition, two age groups of tadpoles were
treated with each concentration. One group was
12 days old at the beginning of treatment; the
other 21 days old. Treatment by immersion was
continuous.
The following groups for each stage in both
species were studied and compared:
(1) . Animals metamorphosing normally in
the laboratory;
(2) . Animals accelerated by 1:10,000,000
thyroxine from 12 days of age;
(3) . Untreated animals of the same age as
were those in (2) when killed for ex-
amination;
(4) . Animals accelerated by 1:10,000,000
thyroxine from 21 days of age;
(5) . Untreated animals of the same age as
were those in (4) when killed for ex-
amination;
(6) . Animals accelerated by 1:100,000,000
thyroxine from 12 days of age;
(7) . Untreated animals of the same age as
were those taken in (6) when killed for
examination;
(8) . Animals treated with 1:100,000,000
thyroxine from 21 days of age;
(9) . Untreated animals of the same age as
were those in (8) when killed for exam-
ination.
At the selected stages, internal metamorphosis
was studied morphologically following dissec-
tion, and the forelimb stage of R. pipiens by
microscopic examination of serial section. The
internal organs studied were:
(1) . Tongue: length, development, presence
or absence or premetamorphic papillae;
(2) . Intestine: length, histology (in R. pi-
piens) at the forelimb stage;
(3) . Gall bladder: color, size;
(4) . Urinary bladder: presence, degree of
development;
(5) . Pancreas: size, histology (in R. pipiens,
forelimb stage) ;
(6) . Pituitary: (in /?. p/p/cni, forelimb stage)
size, development;
(7) . Thyroid: (in /?. pip/eni', forelimb stage)
follicle number, mitotic activity, posi-
tion.
In addition, body length, tail length and hind-
limb length were recorded throughout the course
of normal and accelerated metamorphosis in
both species.
Because of R. catesbeiana’s long tadpole per-
iod, 17 late tadpoles were collected from the
Kenilworth Lily Ponds, Washington, D. C.,
March 24, and brought to the laboratory to
serve as the normal series for the stages of fore-
limb emergence and tail resorption. The rest of
the R. catesbeiana tadpoles used for the study
of younger normal stages and for the experi-
mental series were laboratory raised from an
egg film collected at the same place, July 2.
2. Method of Rearing and Feeding the Tadpoles
The tadpoles were reared in flat, white enam-
eled pans (10" X 15" X Wz"). For R. pipiens,
20 animals were placed in each pan with 1,000
cc. of water or thyroxine solution. For R. cates-
beiana, 1 0 animals were placed in each pan with
2,000 cc. of water or thyroxine solution. The
thyroxine solutions and water were changed
daily. Food (chopped spinach and Pablum)
was provided in excess. The experiments termi-
nated after forelimb emergence when the ani-
mals’ tails were resorbed to dark stubs or when
deaths had reduced the number of animals in
each series to 10 for R. pipiens and to 5 for R.
catesbeiana.
3. The Normal and Experimental Series
The experimental animals in each species
were divided into four series. In the table below
are listed the designation, the treatment and the
number of animals in each series.
Naturally metamorphosing animals consti-
tuted the series designated as “SC” (Stage Con-
trols). Separate series of untreated tadpoles
were raised and observed, and individuals from
these series were fixed concurrently with those
from the experimental series. These constituted
the “Age Controls.” For R. pipiens there was a
total of 100 tadpoles in the Age and Stage Con-
1957]
McGovern & Charipper: Metamorphosis in Rana pipiens and Rana catesbeiana
35
trol series; for R. catesbeiana, a total of 96 tad-
poles.
Number of
ANIMALS
Desig-
R. pi-
R. cates-
nation
Treatment
piens
beiana
A-1
1:10,000,000 thyroxine
from age 12 days
60
30
A-2
1:10,000,000 thyroxine
from age 21 days
60
30
B-1
1:100,000,000 thyroxine
from age 12 days
60
30
B-2
1 : 100,000,000 thyroxine
from age 21 days
60
30
4. External Measurements
Measurements were made as often as changes
in rate of development seemed to require. In
both species, measurements were made daily
during the course of the experiments using
1:10,000,000 thyroxine. For the 1:100,000,000
thyroxine series, generally daily measurements
were made for R. pipiens and less frequently for
R. catesbeiana (once a week or ten days). At
such times, 10 animals in each series were meas-
ured. To facilitate the process of measuring, the
normal and experimental animals of both species
were anesthetized with MS 222.
5. Dissection
Two specimens of R. pipiens and three speci-
mens of R. catesbeiana for each normal stage
and each experimental stage, along with an equal
number of age controls, were dissected. Gall
bladder color changes and presence or absence
of the urinary bladder were investigated in addi-
tional dissections made so that these observa-
tions were based on at least five dissections for
each stage and series in both species. Dissection
was done using 7X and 17 X magnifications.
The entire digestive tract was removed from the
body cavity by freeing the cloaca from the
ureters, cutting the mesentery and lifting the
entire mass from the dorsal body wall anteriorly
at the glottal level of the pharynx. Then the color
of the gall bladder and its size relative to the
liver were recorded. The shape of the ventral
pancreas was sketched and its dimensions taken
with dividers recorded on the sketch. The liver
and pancreas were removed, the bile duct sev-
ered and the intestinal mesentery cut so that the
entire intestine could be uncoiled. Measurement
was accomplished by cutting the intestine into
convenient lengths and straightening these
lengths on a millimeter rule. Finally the presence
or absence of the urinary bladder and its relative
degree of development were recorded.
The shape of the tongue was sketched and
(for R. catesbeiana) the anterior-posterior di-
mension as taken with dividers was recorded.
The number and disposition of the premeta-
morphic papillae were also recorded.
6. Histological Study
Two animals in the forelimb emergence stage
in each of the R. pipiens series were serially sec-
tioned and prepared for histological study. Age
controls were also serially sectioned. Specimens
were fixed in Bouin’s solution, sectioned in par-
affin at 10 micra and stained with Masson’s
stain. Observations
1. Growth Measurements
It has been noted already that the normal
series of R. catesbeiana represents animals ob-
tained at different times; (first) mature tadpoles
collected in the field and raised in the laboratory
through the rest of the tadpole period; and (sec-
ond) tadpoles hatched in the laboratory from
an egg film collected later in the season. The tad-
poles from the egg film provided both the experi-
mental series and their normal Age Controls.
The Age Controls were observed during a period
of 173 days and were fixed at the end of that
time with the remaining experimental. By this
time, a number of the Age Controls were as
large and as well developed as the late tadpoles
(used as the Stage Controls) had been when
collected. Therefore the growth data for the
two normal groups were plotted as a single curve
(Text-fig. 1) broken by a time lapse equal to
that between December (when the Age Controls
were fixed) and March (when the Stage Con-
trols were collected).
The growth curves and the time required for
response (Text-fig. 2) were approximately the
same in both species of organisms treated with
1 : 10,000,000 t^roxine. The experiments termi-
nated in death after 16 days for R. pipiens and
10 or 11 days for R. catesbeiana. With 1:100,-
000,000 thyroxine, the growth curves for the
two species are very similar (Text-figs. 3 & 4) .
The number of days required for response in
R. catesbeiana was greater than for R. pipiens,
but, considering the normal length of the re-
spective tadpole periods, the effect of treatment
appeared earlier in R. catesbeiana (see Text-
fig. 5). It should be pointed out, however, that
tail resorption was carried further in R. pipiens
(Text-fig. 6) during this time.
The age difference in the groups of tadpoles
treated with the same thyroxine concentrations
caused no appreciable difference in the growth
curves in either species.
The curves obtained for hindlimb growth of
tadpoles treated with 1:100,000,000 thyroxine
(Text-figs. 3 & 4) are almost identical with that
published by Etkin (1935) for thyroidectomized
R. cantabrigensis tadpoles. Relative to body
length, the hindlimbs of the 1 : 1 00,000,000 thy-
roxine-treated animals were longer than normal
in both species (Text-fig. 5).
I
36
Zoological New York Zoological Society
[42:3
Text-fig. 1. Normal growth curves for (a) body
length, (b) tail length, and (c) hindlimb length in
R. pipiens and R. catesbeiana. One unit of age for R.
pipiens has been made equal to five units for R.
catesbeiana for easier comparison of the curves.
Continuous tail growth of both species of ani-
mals treated with 1:100,000,000 thyroxine
(Text-fig. 6) was unexpected in view of Blacher’s
( 1928) observation that short exposure to dilute
concentrations of thyroidin produced tail short-
ening.
2. Dissection Results
a. Intestine
Kuntz (1922) found that the average reduc-
tion in intestine length during the normal meta-
morphosis of R. pipiens was 82.2%. In the pres-
ent investigation the average reduction in the
intestine length of control animals was found to
be 86.2% for R. pipiens and 83.7% for R.
catesbeiana. These averages are in very good
agreement if it is remembered that dietary dif-
ferences may affect intestine length by as much
as 58% (Babak, 1905).
Text-fig. 7a shows the changes in the ratio of
intestine length to body length for the normal
MM
days
MM
40
30
(b)
/ Body a
L. Tail
Body
A- 1
R pipiens
10 20 30 40
A-2
R catesbeiana
Legend
control
treated
ACCELERATED GROWTH
Text-fig. 2. Growth under treatment with
1:10,000,000 thyroxine. A-1 curves are for tadpoles
treated from 12 days of age; A-2 are for tadpoles
treated from 21 days of age; (a) and (b) show
data from treated R. pipiens tadpoles; (c) and (d)
from R. catesbeiana.
and for the accelerated R. pipiens series; in Text-
fig. 7b, comparable data are presented for R.
catesbeiana. It can be seen that, during normal
metamorphosis, the ratio for the tadpole stage
is 13 to 14 and it decreases to a value of approxi-
mately 2 at the tail resorption stage. By the time
of forelimb emergence, one-half of the total
shortening had occurred in the intestine of R.
catesbeiana and three-quarters of the total short-
ening of the intestine of R. pipiens had been
effected. Treatment with 1 : 10,000,000 thyroxine
produced shortening of the intestine, giving a
ratio (intestine length to body length) of 2;
however, shortening had been completed by the
forelimb emergence stage. Under the influence
of 1:100,000,000 thyroxine, little shortening
had taken place by forelimb emergence. By the
terminal stage, the intestine length-body length
ratio in R. pipiens is almost normal as compared
with the ratio for R. catebeiana in which the
intestine remains considerably longer than twice
the body length.
1957]
McGovern & Charipper: Metamorphosis in Rana pipiens and Rana catesbeiana
37
ACCELERATED GROWTH
R. PIPIENS
Text-fig. 3. R. pipiens, growth of tadpoles under
1:100,000,000 thyroxine treatment. B-1 curves are
for tadpoles treated from 12 days of age; B-2 for
tadpoles treated from 21 days of age.
Blacher (1928) and Allen (1932) established
the order of sensitivity to thyroid treatment of
the several tadpole organs. Both Blacher and
Allen reported the intestine to be one of the
more sensitive organs by virtue of its shortening
after even brief periods of exposure to dilute
concentrations. In Text-fig. 7, the curves show
that the intestine of tadpoles treated with
1:100,000,000 thyroxine continues to grow for
some time prior too the onset of shortening. The
data can be reconciled if, instead of shortening,
one postulates a period of slower intestinal
growth in the treated tadpoles than in controls
of the same age.
b. Pancreas
In the normal animal the pancreas progres-
sively increases in size up to the time of meta-
morphosis when regression begins. The shape
and the size of the pancreas were the same in
treated and in untreated individuals of both
species of the same age. Because regression cut
short the normal growth period, the experi-
mental animals’ pancreas began regression at a
MM
B-l
ACCELERATED GROWTH
R. CATESBEIANA
Text-fig. 4. Growth of R. catesbeiana tadpoles un-
der 1:100,000,000 thyroxine treatment, (a) B-l
curves are for tadpoles treated from 12 days of age;
(b) B-2 curves for tadpoles treated from 21 days of
age.
smaller size than normal. Treatment with
1 : 10,000,000 thyroxine elicited a marked regres-
sion at the forelimb emergence stage and further
shrinkage was observed in the terminal stage.
There was no apparent regression in the pan-
creas of animals in the forelimb emergence
stage which were treated with 1:100,000,000
thyroxine. At the onset of the terminal stage,
regression was evident in both species.
c. Gall Bladder
Early in normal development the gall bladder
was large as compared with the liver (about 1 /5
its size). Later, it was relatively smaller (1/8
to 1 /lO of the size of the liver). By the time of
forelimb emergence, the gall bladder again ap-
peared larger ( 1/5 to 1/4 the size of the liver) .
No measurements were made to show whether
these size changes were changes in the gall
bladder or in the liver, though it was apparent
that the liver first increased and then decreased
in size. Hoskins & Hoskins (1919a&b) reported
I
38
Zoological New York Zoological Society
[42:3
/^ge ,n 50 100 150 200 250 300 350 400
Doys (b) R.catesbeiano
RELATIVE HINDLIMB
LENGTHS
Text-fig. 5. Relative hindlimb lengths in normal
tadpoles and tadpoles treated with 1:100,000,000
thyroxine from 21 days of age. (a) R. pipiens; (b)
R. catesbeiana. Age in days is plotted against the
ratio of body length to hindlimb up to the tail
stub stage in normal animals and to the “terminal”
stage in the treated.
that the liver decreased in size during meta-
morphosis, and Kuntz (1922) reported an 80%
reduction in liver weight which took place when
tail resorption and intestine reduction were well
advanced.
The gall bladders of animals treated with
1:10,000,000 thyroxine remained relatively
large. At the terminal stage, they appeared rela-
tively larger than in normal animals at the tail
stub stage and the livers appeared to be more
than normally reduced in size. The gall bladders
of tadpoles treated with 1:100,000,000 thy-
roxine were still 1/8-1/10 of the size of the
liver at the forelimb emergence stage.
Color change in the gall bladder during accel-
erated metamorphosis was studied in Hyla versi-
color, H. crucifer, R. clamitans, Bufo ameri-
canus and Ambystoma maculatum (A. puncta-
tum fide Speidel) by Speidel (1926). He noted
MM
40
30
(o)
20
/ ''
10
-
0
10 20 30 40 50 60 70
R. pipiens
MM
-
50
40
Legend
(b) 30
/ / Control
20
]/ Treated (8-2)
10
L 1 1 1 1 1 1 1 1
Age in 50 100 150 200 250 300 350
R.catesbeiano
TAIL GROWTH
Normal and Accelerated
Text-fig. 6. Tail growth in normal tadpoles and
tadpoles treated with 1:100,000,000 thyroxine from
21 days of age. (a) R. pipiens; (b) R. catesbeiana.
no exceptions in several hundred observations
from the series: “yellow-green,” “brighter-
green,” “emerald green” and a “very dark
green.” In normal R. pipiens, the gall bladder
was dark green at the time of forelimb emer-
gence. In R. catesbeiana the gall bladder was
emerald green at this time, and it was dark green
at the tail stub stage. In most of the 1:10,000,-
000 thyroxine-treated R. pipiens tadpoles, the
gall bladder was dark green at the forelimb
emergence stage; in R. catesbeiana, at the ter-
minal stage, the gall bladder was still emerald
green as noted above. In the 1 : 100,000,000 thy-
roxine-treated tadpoles of both species, the gall
bladder was emerald green at the forelimb emer-
gence stage and approximately half of these in-
dividuals had dark green gall bladder at the
terminal stage.
d. Urinary Bladder
The urinary bladder developed late in the
normal premetamorphic climax period and was
small but well developed in the tadpole with
fully developed hindlimbs.
At the forelimb emergence stage of both spe-
cies which had been treated with 1:10,000,000
1957]
McGovern & Charipper: Metamorphosis in Rana pipiens and Rana catesbeiana
39
Text-fig. 7. Relative intestine lengths. Age in days is plotted against the ratio of intestine length to body
length for R. pipiens in (a) and for R. catesbeiana in (b). A-1 curves are for tadpoles treated with
1:10,000,000 thyroxine from 12 days of age; A-2 for those treated with 1:10,000,000 thyroxine from
21 days of age; B-1 for those treated with 1:100,000,000 thyroxine from 12 days of age; B-2 for those
treated with 1:100,000,000 thyroxine from 21 days of age.
thyroxine, approximately half had urinary blad-
ders. At the terminal stage, all but one or two
animals had small, poorly developed urinary
bladders. In both species, the animals treated
with 1:100,000,000 thyroxine had well-devel-
oped urinary bladders by the time of the fore-
limb emergence period.
e. Tongue
No measurements of the tongue were made for
R. pipiens. In R. catesbeiana, normal develop-
ment of the tongue began (when the tadpoles
reached 21-27 mm. body length) as a white con-
densation of tissue in the anterior part of the
floor of the mouth. At a body length of 35-38
mm., the tongue was clearly outlined and the
tissue mass was about 1 mm. long. At the time
of forelimb emergence, the tongue was a well-
formed organ, 2.5-3 mm. long, with the poste-
rior edge free and bifurcated. Between the end
of forelimb emergence and the onset of the tail
stub stage, the tongue grew rapidly and almost
doubled its length to 5 mm.
All the experimental animals of both species
showed more advanced tongue development
than normal animals of the same age. In the
groups treated with 1 : 10,000,000 which were
examined at the terminal stage, the tongue was
poorly differentiated, i.e., a free but not bifurcate
posterior edge. In the groups treated with
1:100,000,000 thyroxine, the tongue appeared
to be the same as in corresponding stages of
normal animals. The amount of tongue growth
(in length) between the forelimb emergence
and the terminal stages for thyroxine-treated R.
catesbeiana is compared with that in the normal
in the table below. Also included is the time in
days between the two stages. (The A-1 series,
treated with 1:10,000,000 thyroxine from 12
days of age, is omitted because the tongue meas-
ured less than 0.5 mm in length at the forelimb
emergence stage).
The slower growth rate of the tongue in or-
ganisms treated with 1:100,000,000 thyroxine
is marked (See B-1 and B-2).
In the normal R. pipiens tadpole there are two
premetamorphic tongue papillae; in R. cates-
beiana there are four. These begin to be resorbed
in the late tadpole stages and disappear by the
onset of tail stub stage. In all the R. pipiens
series, the papillae were gone by the beginning
of the forelimb emergence, but in R. catesbeiana.
40
Zoological New York Zoological Society
[42:3
Tongue Growth (in mm.)
Stage
Control
.4-2*
B-1**
B-2**
Forelimb Emergence
2.7
0.5
1.3
1.3
Tail Stub (Terminal)
5.0
0.7
2.2
1.8
% Increase
85%
40%
69%
38%
Days between Stages
7
2
82
90
* Treated from 21 days of age with 1:10,000,000
thyroxine.
** Treated from 12 days of age with 1:100,000,000
thyroxine.
***Treated from 21 days of age with 1:100,000,000
thyroxine.
papillae were still present at the terminal stage
of animals treated with 1:100,000,000 thy-
roxine.
The histology of the premetamorphic tongue
papillae of R. sylvatica was studied by Helff &
Mellicker (1941) who found four papillae, the
characteristic number for that species. In their
search of the literature, they found that one
previous investigator had described these struc-
tures. Schulze (1870, cited in Helff & Mellicker,
1941) found two papillae in Pleobates fusca, R.
esculenta, R. temporaria and Bufo cinereus. R.
catesbeiana thus resembles R. sylvatica in pos-
sessing four papillae, whereas R. pipiens re-
sembles the species studied by Schulze.
3. Histological Studies: Forelimb Stages, R.
pipiens
a. Pituitary Gland
i. Anatomy
In Text-fig. 8 are reproduced Atwell’s (1918)
reconstructions of the development of the
anuran pituitary gland. It can be seen that two
changes take place: first, the two lobes of the
pars lateralis come to lie anterior and lateral to
and (in the adult) separate from the pars an-
terior and pars intermedia; second, the mass of
the gland grows posteriorly, the Anterior at a
more rapid rate than the Intermedia (so that in
the adult the former is the most posterior part of
the gland. Also in Text-fig. 8, the length of each
of the three components of the pituitaries of
single normal and experimental animals is
plotted in relationship to other parts. These dia-
Text-fig. 8. The pituitary in normal and treated R. pipiens. The drawings numbered 9 through 14
above the block graphs are redrawn from Atwell’s (1918) reconstructions of pituitary development in
Anura. The block graphs show the length and spatial relationships of the pituitary parts of normal and
treated R. pipiens tadpoles obtained by counting the 10 micra sections each part occupied in each
animal. A-1 is data from a forelimb stage animal 24 days old treated with 1:10,000,000 thyroxine from
12 days of age; A-l-C is from an untreated tadpole 24 days old; A-2 is from a forelimb stage animal 34
days old treated with 1:10,000,000 thyroxine from 21 days of age; A-2-C is from an untreated animal 34
days old; B-1 is from a forelimb stage animal 50 days old treated with 1:100,000,000 thyroxine from
12 days of age; B-l-C is from an untreated tadpole 50 days old; B-2 is from a forelimb stage animal
52 days old treated with 1:100,000,000 thyroxine from 21 days of age; SC is from an untreated fore-
limb stage animal 67 days old.
1957]
McGovern & Charipper: Metamorphosis in Rana pipiens and Rana catesbeiana
41
grams result from plotting the number of 10
micra sections constituting each part of the
pituitary.
In the 12-day-old, 1:10,000,000 thyroxine-
treated forelimb stage (A-1), pituitary relation-
ships correspond to those in Atwell’s youngest
animal (stage 9), while the untreated animal of
the same age (A-l-C) had progressed to Atwell’s
stage 10. In the 21 -day-old, 1:10,000,000 thy-
roxine-treated forelimb stage animal (A-2),
these appear to be the same as in the untreated
animal of the same age. Both correspond to
Atwell’s stage 10 or 11. The higher concentra-
tion of thyroxine did not cause precocious ana-
tomical development of the pituitary.
The pituitaries of the animals treated with
1:100,000,000 thyroxine from 12 days of age
(B-1) and those treated from 21 days of age
(B-2) were smaller than those of either the
normal tadpole of the same age (B-l-C) or of
the normal forelimb stage animal (SC). The
block graphs do not show the progressive
change in the connection between the two lobes
of the pars lateralis with the pars anterior illus-
trated by Atwell’s figures 12, 13 and 14. Exami-
nation of the slides showed that the normal tad-
pole (B-l-C), on the basis of lateralis position,
had reached a stage between Atwell’s 12 and 13
while the normal animal (SC) was between
stages 13 and 14. The forelimb animals treated
with 1:100,000,000 thyroxine (B-2 and B-4)
appear to resemble the normal tadpole more
than the normal forelimb emergence stage. The
lower concentration of thyroxine, moreover,
showed no accelerating effect on the anatomical
development of the pituitary.
ii. Relative size
Text-fig. 8 also shows that the pituitaries of
PITUITARY PART LENGTHS
AND BODY LENGTHS AS 0/0's
OF NORMAL FORELIMB
STAGE VALUES
B Experimental
(H Control
Text-fig. 9. Pituitary part lengths and body lengths of thyroxine-treated forelimb stages compared with
those of the normal forelimb stages in R. pipiens. The length of each pituitary part and the body
length of the experimental forelimb stages and of normal tadpoles of the same age as the treated are
shown as percents of the normal forelimb stage values (SC, last set of four 100% blocks). In A-1, data
from a forelimb stage animal treated with 1:10,000,000 thyroxine from 12 days of age are shown in
black blocks superimposed upon that of an untreated animal of the same age in white. In A-2 the data
from a forelimb stage animal treated with 1:10,000,000 thyroxine from 21 days of age are shown in
the same way against the data from an untreated tadpole of the same age; B-1 are data from a forelimb
stage animal treated from 12 days of age with 1:100,000,000 thyroxine shown against that for an un-
treated tadpole of the same age; B-2 data from a forelimb stage animal treated from 21 days of age
with 1:100,000,000 thyroxine are shown against the data for an untreated tadpole of the same age.
42
Zoologica: New York Zoological Society
[42:3
all of the experimental forelimb-emergence ani-
mals were shorter than those of untreated ani-
mals of the same age. To determine whether the
reduced size of the pituitary was proportional
to the reduced body size of the thyroxine-treated
animals and to determine whether or not all
pituitary parts were equally affected, the length
of each pituitary part and the body length of the
normal forelimb stage animal were taken as
100%. The same measurements for each ex-
perimental forelimb animal and for its age con-
trol were recalculated as percents of the lengths
of the corresponding parts in the normal fore-
limb animal. In Text-fig. 9, the profiles produced
by graphing these values for the experimental
animals were superimposed in black upon the
profiles of their respective age controls. The last
profile to the right (a block of four 100% col-
umns) represents the values for the normal fore-
limb stage animal.
It can be seen that the bodies of those treated
with 1:10,000,000 thyroxine and those treated
with 1:100,000,000 thyroxine appeared to have
bodies and pituitaries about equally inhibited
(with regard to length) at the forelimb stage,
and in these animals the pars anterior appeared
to be the pituitary component most inhibited,
iii. Cellular Differentiation
Distribution and differential counts of the
pituitary cell types were not considered because
of the thickness of the sections. The cytoplasm
in the cells of all the lobes was less abundant in
the experimental animals than in either the age
controls or the normal forelimb stage. Pigment
granules which were large and coarse in young
normal stages were more evident in all the ex-
perimentals than in the stage controls.
b. Thyroid Gland
i. Size of the Gland
Conclusions as to thyroid size were based on
the length of the glands determined by counting
the number of 1 micra sections which the two
halves of the gland occupied in each animal, and
on a relative area per section derived from count-
ing the number of follicles containing visible
colloid. To show how the thyroids of the thy-
roxine-treated forelimb stage animals differed
from those of the normal forelimb stage animal
and from the untreated tadpoles of the same age,
body length, length of thyroid and average num-
ber of colloid-filled follicles per section in the
normal forelimb stage animal were arbitrarily
rated as 100%. The same data for the treated
animals (forelimb stage) and for their respective
age controls were recalculated as percents of
these norms. In Text-fig. 10 are five profiles re-
sulting from this treatment of the data. At the
extreme right is the normal forelimb stage pro-
®/
4>
H Experimental
□ Control
THYROID LENGTHS
Text-fig. 10. Thyroid size and body size in normal
and thyroxine-treated R. pipiens. Body length, length
of thyroid determined by counting the number of
10 micra sections the gland occupied and the aver-
age number of follicles per section for the accele-
rated forelimb stages and for untreated animals
of the same ages as the treated are shown as per-
cents of the normal forelimb stage values (SC, the
last set of three 100% blocks). The values for each
treated forelimb stage animal are shown in black
blocks superimposed on the same values for a normal
tadpole of the same age shown by white blocks.
A-1 shows the values for a forelimb stage animal
treated from 12 days of age with 1:10,000,000
thyroxine and for a normal animal of the same
age; A-2, for a forelimb stage animal treated from
21 days of age with 1:10,000,000 thyroxine and
for a normal tadpole of the same age; B-1 for a
forelimb stage animal treated from 12 days of age
with 1: 100,000,000 thyroxine and for a normal tad-
pole of the same age; B-2 for a treated forelimb
stage animal treated from 21 days of age with
1:100,000,000 thyroxine and for a normal tadpole
of the same age.
file. To the left are the profiles for the untreated
tadpoles. Superimposed upon these are the pro-
files of the treated forelimb stage animals of the
same age.
Text-fig. 10 shows that the thyroids of the
treated animals at the forelimb emergence stage
were inhibited as reflected by their size when
compared with the normal forelimb stage and
1957]
McGovern & Charipper: Metamorphosis in Rana pipiens and Rana catesbeiana
43
...
untreated animals of the same age. Inhibition as
indicated by area (determined by the number of
follicles) was greater than that indicated by
length.
ii. Mitotic Activity
In the following table are given the number of
10 micra sections occupied by both the right and
left halves of the thyroid glands, and the number
of mitoses in the entire glands.
Sections
Mitoses
A-1
30
3
A-l-C
39
47
A-2
49
12
A-2-C
62
120
B-1
95
14
B-l-C
130
406
B-2
131
24
SC
211
861
The normal series-
-A-l-C, A-2-C,
B-l-C and
SC— gave evidence of higher mitotic activity than
did the treated animals.
iii. Glandular Activity
A thyroid gland made up of vacuolate col-
umnar cells, with vacuolate colloid, and of many
collapsed follicles, possesses the generally ac-
cepted histological criteria of a physiologically
active gland. The forelimb stage animals treated
from 12 days of age with 1:10,000,000 thy-
roxine (A-1) had thyroid glands made up of
cuboidal cells with scant cytoplasm, with few
follicles, and with no vacuolation. In those
treated with 1:100,000,000 thyroxine from 21
days of age (A-2), the thyroid cells were
cuboidal. The cells had more cytoplasm than
those of the A-1 animals but less than those in
animals of the same age. As in the age controls,
a few follicles showed vacuolation of the colloid.
The animals from both age groups treated with
1 : 1 00,000,000 thyroxine had thyroids in which
the cuboidal cells were approaching a columnar
architecture. The thyroids of the normal fore-
limb stage animals bore all of the criteria of high
activity.
iv. Pigment
In the younger normal animals (as was noted
by D’Angelo & Charipper, 1939) large aggre-
gates of pigment are conspicuous features of the
thyroid gland. By the time of forelimb emer-
gence, however, such aggregates were rare.
Moreover, the granules are distributed and the
pigmentation is not conspicuous. All of the ex-
perimental animals showed pigmentation either
to the same degree or to a greater degree than
their respective age controls.
V. Thyroid Position
Text-fig. 11 shows outline drawings from
Text-fig. 11. Thyroid position relative to the gill
skeleton in the normal forelimb stage and in the
forelimb stage produced by treatment with
1:100,000,000 thyroxine from 12 days of age.
Graphic reconstruction from serial sections. Normal
stage magnified 5X; treated 11.25X. The thyroid
is shaded.
graphic reconstruction of the thyroid gland and
its relationship to the ventral aspect of the
hyobranchial cartilages. Text-fig. 11a shows the
normal forelimb emergence stage (5X). Text-
fig. 11b is the forelimb emergence stage after
1:100,000,000 thyroxine treatment from 12
days of age (enlarged 11.25X). The thyroid
gland assumed a more anterior position under
the gill-bearing skeleton in the treated animals
than in the normal. D’Angelo & Charipper’s
(1939) Figure 1 , illustrating the normal changes
in thyroid position up to and including meta-
morphosis, indicates that the more anterior posi-
tion is characteristic of normal tadpoles of the
same age as the treated. The position of the
thyroid in the normal forelimb stage agrees with
their observations from the same stage. Thus
1:10,000,000 thyroxine treatment did not effect
a precocious migration of the thyroid.
c. Small Intestine
In the normal forelimb stage animal, the
histology of the small intestine is well-repre-
sented by Kuntz’s (1922) figures for the same
stage of R. pipiens and by lane’s (1934) illus-
tration for this stage in thyroid-fed tadpoles of
R. clamitans. There were many mitoses in the
basal cells of the new mucosal layer. The old
mucosa was evidenced by conspicuous necrotic
masses. The submucosa and muscularis had
44
Zoological New York Zoological Society
[42:3
thickened considerably. The forelimb stage ani-
mals treated with 1:10,000,000 thyroxine had
the intestinal histology of the normal forelimb
stage. However, the cytoplasm of the cells in all
layers was very scant and the resulting histology
was superficially quite different from the normal.
Forelimb stage animals treated with 1:100,-
000,000 thyroxine had the same intestinal his-
tology as that of the untreated tadpoles of the
same age.
d. Pancreas
In the normal forelimb stage animal regres-
sion of the pancreas was far advanced. The his-
tological picture represented a condition inter-
mediate to Kaywin’s (1936) stages 3 and 4 of
thyroxine-treated R. catesbeiana. The acini
showed three stages of metamorphic changes:
( 1 ) some were necrotic, (2) some were made up
of syncytial cells, and (3) some were made up
of cells with large nuclei and sparse, deeply-
staining cytoplasm. There were pronounced con-
nective tissue spaces.
In the forelimb stage of animals treated with
1 : 10,000,000 thyroxine, the pancreas was com-
parable to the normal forelimb stage pancreas
as regards the degree of metamorphic change.
However, pigment was very conspicuous. Fore-
limb stage animals treated with 1:100,000,000
thyroxine failed to show metamorphic change in
the pancreas. The glands were smaller than
those of the untreated tadpoles of the same age,
but there were no signs of loss of definition of
cell boundaries or necrosis in the acinar cells.
e. Summary of Observations on the Histology
of the Forelimb Stages of R. pipiens
Kahn (1916) found that the pituitaries of
tadpoles treated with horse thyroid were much
larger than normal. Schliefer (1935) found that
thyroid extract had no effect on the pituitary
although the development of the gland was ac-
celerated along with the rest of the body. Under
the conditions of the present study the anatom-
ical development, at the forelimb emergence
stage, of the pituitary of thyroxine-treated ani-
mals was not found to be more advanced than
that of untreated animals of the same age. In
fact, the glands were smaller. Treatment with
1:10,000,000 thyroxine was less inhibitory to
pituitary growth than it was to body growth.
Thus, in agreement with Kahn, the pituitaries
were larger than those of untreated tadpoles of
the same age. After treatment with 1:100,000,-
000, body size and pituitary size were in a more
normal proportion; with this concentration, the
pars anterior appeared to be more inhibited than
the other pituitary components. With both con-
centrations, the cytoplasm of the cells of all
parts of the pituitary gland was sparse and pig-
ment remained conspicuous in the pars anterior
and pars lateralis.
The thyroids of treated animals showed evi-
dence of inhibition in every respect considered
at forelimb emergence. They were much smaller,
had a lower mitotic rate and give histological
evidence of a lower degree of differentiation and
physiological activity than the glands of either
the normal forelimb stage or of the untreated
tadpoles of the same age. These findings agree
with those of Clements (1932), Etkin (1935)
and Brinks (1936).
The small intestine and pancreas in animals
treated with 1 : 10,000,000 thyroxine showed the
same type of histological change at the time of
forelimb emergence as that of normal animals at
the same stage. On the other hand, the intestines
and pancreases of those treated with 1 : 100,000,-
000 thyroxine did not show metamorphic
change.
4. General Summary of Observations
A comparison of the condition of seven in-
ternal organs, in normal metamorphic stages
and in comparable stages produced by treat-
ment with two concentrations of thyroxine, has
shown that at each stage, the accelerated ani-
mals are different not only from the normal but
also from each other. This is summarized, in the
table below, for the forelimb stage.
Difference in the length of the tadpole period
in R. pipiens and in R. catesbeiana did not
greatly affect the results of thyroxine treatment
as studied by growth measurements and dissec-
tion. R. catesbeiana reacted earlier, considering
its normal tadpole period, to both concentra-
tions of thyroxine than did R. pipiens. In days,
R. catesbeiana reacted more quickly to 1:10,-
000,000 thyroxine and less quickly to 1:100,-
000,000 thyroxine than R. pipiens. As judged
from the degree of response, R. catesbeiana was
less sensitive than was R. pipiens, as shown by
the fact that resorption of the tail and shorten-
ing of the intestine after 1:100,000,000 thyrox-
ine treatment was not as advanced. In compari-
son with their respective normal stages, never-
theless, the animals showing accelerated devel-
opment behaved similarly in both species.
Discussion
The thyroid glands themselves of animals un-
dergoing thyroxine-accelerated metamorphosis
apparently do not modify the effects of thyroxine
treatment. Etkin (1935) reached this conclu-
sion by comparing the effects of thyroxine on
thyroidectomized and partially thyroidectomized
R. cantabrigensis tadpoles. The present study
supports that conclusion by the detailed simi-
larity of the external metamorphosis of acceler-
Normal Development in Normal and Treated Forelimb Stage Animals
1957]
McGovern & Charipper: Metamorphosis in Rana pipiens and Rana catesbeiana
45
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46
Zoological New York Zoological Society
[42:3
ated intact R. pipiens and R. catesbeiana to that
of his accelerated, thyroidectomized animals.
The development of the thyroids of treated R.
pipiens at the time of forelimb emergence was
very much inhibited and resembled that of hypo-
physectomized tadpoles described by Atwell
(1935) seventeen months after operation. The
growth of the pituitaries of these animals was
also inhibited. D’Angelo (1941) correlated the
growth and differentiation of the thyroid with
the marked growth and basophilic differentiation
of the pituitary pars anterior during the period
of rapid hindlimb growth. Inhibited pituitary
development is a plausible explanation for the
inhibited condition and apparent inactivity of
the thyroids of thyroxine-treated tadpoles.
In attempting to duplicate the normal pattern
of sequence and spacing of the external meta-
morphic events, Etkin (1935) found that no one
concentration of thyroxine could effect this re-
sult. However, by starting with low concentra-
tions and gradually increasing the dosage, he
obviated both the simultaneous occurrence of
events normally spaced in time which resulted
from the use of single high concentrations and
the greater than normal intervals between events
which resulted from the use of single low con-
centrations. Schreiber (1934 a & b) and Cache
(1940) account for both the sequence and the
spacing of events in normal metamorphosis as
results of differences in the response thresholds
of the several organs at the level of thyroxine
increases. Etkin, having found the sequence of
events unchanged by different concentrations of
thyroxine, considered that this sequence was in-
herent in the tissues, all of which were sensitive
even to the lowest concentrations, and concluded
that only the spacing of events and rate of re-
sponse were thyroxine controlled.
If the sequence of events is controlled by in-
creasing concentrations of hormone reaching
successively the low thresholds of early events
and the higher thresholds of later events, then a
particular concentration capable of initiating, let
us say, a late event must also be able to initiate
other events normally occurring earlier or at
the same time. If the sequence of events is in-
herent in the tissues and is independent of the
concentration of thyroxine, then if a particular
event has already taken place in both normal
and accelerated animals, other events already
begun in normal animals should also have begun
in the accelerated animals regardless of the con-
centrations used.
In the present experiments, normally meta-
morphosing and accelerated animals were ex-
amined at a time when, in each, the same ex-
ternal metamorphic event had just occurred.
namely, forelimb emergence. In the normal ani-
mal, the histology of the intestine and pancreas
is midway in the change to the adult condition.
In animals treated with 1:100,000,000 thyrox-
ine, there was no histological change apparent
in either organ, while those treated with 1:10,-
000,000 thyroxine resembled the normal in his-
tology. The urinary bladder is absent in half the
forelimb stage animals treated with 1:10,000,-
000 thyroxine, but it is present in the normal
forelimb stage, the normal late tadpole, and in
the 1:100,000,000 thyroxine-treated animals in
the forelimb stage. The gall bladders of
1:100,000,000 thyroxine-treated animals retain
the tadpole size relation to the liver, but in the
normal forelimb stage and in the 1:10,000,000
thyroxine-treated forelimb stage, the gall blad-
ders are increased in size relative to the Uver.
If the delayed events cited were just begun
although not far advanced, then these devia-
tions in the accelerated animals from the normal
integration of events could be explained as con-
centration effects upon the reaction rates and,
thus, upon the spacing of events, rather than as
an alteration of sequence. This would corrobo-
rate Etkin’s views. Despite the difficulty imposed
by the conditions of the present experiments in
ascertaining precisely the initiation of an in-
ternal metamorphic event, the probability of its
having occurred in normal sequence regardless
of the thyroxine concentration can be further
explored if its position as one of a sequence of
three events is followed.
In normal animals, the urinary bladder ( 1 ) is
well formed and intestinal shortening (2) is
1/2 to 2/3 complete when the forelimbs emerge
(3). In 1:100,000,000 thyroxine-treated ani-
mals, the urinary bladder (1) is well formed
at forelimb emergence (3) but there has been
little or no shortening of the intestine (2). In
1 : 10,000,000 thyroxine-treated tadpoles, all have
fully shortened intestines (2) at forelimb emer-
gence ( 3 ) but half have no urinary bladders ( 1 ) .
At any given moment in normal metamorpho-
sis, the rising concentration of thyroxine is the
same for all organs and although, as Etkin sug-
gests, each event may have its own rate to be
speeded or retarded hy concentration, this rate
at normal concentrations is necessarily related
to those of other events taking place at the same
time. If all the events were initiated in the natural
sequence in the development of the animals
treated with the two concentrations, the cited
deviations in developmental pattern from the
normal in the two groups can be ascribed en-
tirely to the effects of the experimental concen-
trations on rate. It seems improbable that the
urinary bladder, normally formed before intes-
1957]
McGovern & Charipper: Metamorphosis in Rana pipiens and Rana catesbeiana
A1
tinal shortening begins, should not be visible
under 17 X magnification in half the 1:10,000,-
000 thyroxine-treated animals at the forelimb
stage, all with fully shortened intestines, unless
the beginning of its development had been de-
layed relative to the natural or normal devel-
opmental sequence.
If the normal sequence of events is not a
matter of successive threshold responses (since
a concentration that produced one event did not
produce all the other naturally concurrent
events) and is not independent of thyroxine con-
tration (since both concentrations apparently
produced altered sequence), another explana-
tion for the control of metamorphic sequence
must be sought.
The lack of significant differences in response
to the same thyroxine concentrations in the two
species studied indicates that differences in tissue
sensitivity are not the cause of the species dif-
ferences in the length of time before the onset
of metamorphosis. This, taken with the preco-
cious thyroid activity and metamorphosis pro-
duced by Ingram (1928) in R. catesbeiana with
R. clamata pituitary implants; with Etkin’s
(1950) demonstration of tissue sensitivity to
thyroxine as early as the opercular closure stage;
and with the well-known effects of hypophysec-
tomy on metamorphosis, leads to the conclusion
that the onset of metamorphic change is pitui-
tary-controlled. It has been accepted that this
control is exerted through trophic action on the
thyroid gland. If the sequence of metamorphic
events is neither solely an effect of the thyroid
in producing rising thyroxine concentrations nor
independent of concentration and inherent in
the tissues, perhaps the control of metamorphic
sequence may be found in a thyroid-pituitary
interrelationship at the tissue level.
It is evident that a dynamic equilibrium must
be maintained between growth and differentia-
tion during the late tadpole and early metamor-
phic period. Tadpole organs continue to grow
up to the time the changes take place which
cause them either to disappear or differentiate,
producing the adult condition. Potential adult
organs both grow and differentiate during the
tadpole period. That both the thyroid and pitui-
tary are involved has been shown in the results
of hypophysectomy and thyroidectomy. Smith
(1916 a & b, 1918) showed that although the
initial growth rates of normal and hypophysec-
tomized tadpoles were very similar, a definite
retardation of growth became evident in the
second half of the tadpole period during which
growth is normally rapid. Thus, the effect of
the pituitary upon growth begins to be exerted
at the period of rapid thyroid, hindlimb and
general body growth. Hoskins & Hoskins (1919
a & b) found that thyroidectomized tadpoles
grew more rapidly and were almost twice the
size of normal animals at the time they should
have metamorphosed. Gordon, Goldsmith, &
Charipper (1943, 1945) also reported excessive
growth in chemically thyroidectomized tadpoles.
These results suggest that, after an initial inde-
pendent period, growth during the second part
of the tadpole period is affected by two factors:
( 1 ) a factor from the pituitary-promoting
growth and (2) a factor from the thyroid-in-
hibiting growth. That the latter normally over-
takes the former may be inferred from the ces-
sation of growth at metamorphosis when thy-
roid activity is at its peak. It is of interest to note
here that in accelerated metamorphosis, in an
organ like the pancreas which normally develops
a typical structure that is replaced by a different
adult structure at metamorphosis, the accelera-
tion does not consist of a telescoping in time of
the normal anatomical history of the organ. In-
stead, the tadpole part is dropped out. The ini-
tiation of metamorphic sequence may depend
upon a succession of tissue-determined critical
ratios between the concentrations of the pitui-
tary growth factor and thyroxine.
In an early period, the initiation of develop-
ment in adult organs might take place in an
order determined by the concentration of thy-
roxine and the growth stimulus, while tadpole
organs unaffected by the low thyroxine inhibi-
tion continue to grow; in an intermediate period,
growth of all organs continues followed by a later
period in which growth slows in all organs and
as the growth/thyroxine ratio changes, succes-
sive tadpole organs begin metamorphosis.
This explanation, based on the growth curves
shown in Text-fig. 2 and applied to the conditions
under 1:10,000,000 thyroxine treatment, might
be present as follows.
Because the experiments began in the period
which Smith found independent of hypophysec-
tomy, the cessation of growth observed would
indicate that pituitary independent growth is also
inhibited by thyroxine in this concentration.
Tadpole growth rapidly ceased, regressive
changes were marked, and adult organs ap-
peared but were small and underdeveloped. The
pituitary and thyroid were retarded compared
with the glands of untreated tadpoles of the
same age.
In the 1:100,000,000 thyroxine experiments,
tadpole growth continued. At first, this was a
virtually normal rate, but as time went on the
growth curves flattened. The conditions in
1:100,000,000 thyroxine acceleration might be
represented as follows (Text-fig. 12). Here “a”
is the period of pituitary independent growth.
48
Zoological New York Zoological Society
[42:3
TIME
Text-fig. 12. For explanation see text below.
“b” is growth affected by 1:100,000,000 thy-
roxine (“c”) applied from the time indicated
by the arrow. Three different periods in the rela-
tionship of “a”— “b” to “c” can be recognized,
but each differs from the corresponding period
in the “normal” diagram. In period ti, 1:100,-
000,000 thyroxine is applied; the growth rate in-
creases whereas the level of th}Toxine does not.
The thyroxine concentration is too low to stop
tadpole growth, but is sufficient to initiate the
growth of the adult organs.
In period t2, the relation of “b” and “c” is
fairly constant, as can be seen (Text-fig. 12).
In period ts, “b” decreases slowly and “c” re-
mains constant. As a result “b” approaches “c”
more slowly and the period ts is longer. This
lateration in the amount of change and the tim-
ing of the change in the growth/thyroxine rela-
tionship could account for the relatively longer
limbs of the 1:100,000,000 thyroxine-treated
animals, the slower-than-normal tail resorption
and the delay in intestinal shortening. That tail
resorption eventually takes place in some speci-
mens, as does intestinal shortening, can be ex-
plained by the growth factor falling slowly in
value to a point where, although the thyroxine
concentration has not been changed, the normal
“critical” ratio between the two factors is ap-
proached. This would also bring Allen’s (1932)
statement that “an apparently subminimal con-
centration of thyroxine may be effective if it
acts long enough” into conformity with subse-
quent evidence that thyroxine is not stored in
the tissues (Etkin, 1935).
Experiments on starved tadpoles have pro-
duced results of great interest in this connection.
D’Angelo, Gordon & Charipper (1938) showed
that starvation imposed before the 5-8 mm.
hindlimb stage in R. sylvatica retarded develop-
ment and resulted in failure of metamorphosis,
but starvation imposed after this resulted in pre-
cocious metamorphosis. Later it was found
(1941) that early starvation resulted in extreme
retardation of the thyroid gland and failure of
cell differentiation in the pituitary. Tadpoles
starved at later stages had thyroids which showed
secretory activity for some time. It was suggested
that early metamorphosis may be the result of
precocious thyroid activity possible from the in-
creased sensitivity to thyrotrophic hormone.
There is also a possibility that once the thyroid
secretes hormone, a reduction in the growth po-
tential by starvation could precipitate metamor-
phosis at a level of thyroxine production which
would be insufladent normally. It may be neces-
sary to point out that the smaller body size of
the 1:100,000,000 thyroxine-treated animals in
the present experiments is not the result of star-
vation from precocious cessation of feeding. The
intestines were distended with food when exam-
ined histologically at the forelimb stage and
feces were abundantly present in the thyroxine
solutions at the daily changes until weU after
this period.
The explanation above has been offered to
show how the normal sequence of metamorphic
events might be controlled by a series of critical
ratios between the growth-promotmg influence
of the pituitary and the effects of the thyroid.
The experimental data offer little evidence con-
cerning the function of the pituitary-accelerated
metamorphosis. The pituitary glands were
smaller than normal, on an absolute basis, in
the treated animals but not relatively, as com-
pared with body size. The pars anterior seemed
to be more reduced than the other parts. The
anatomy was tadpole rather than normal at
the forelimb stage. The appearance of the thy-
roid indicated deficiency of thyrotrophic hor-
mone. On such evidence elaboration of theory
is premature and must await further data such
as might be afforded, for example, by a compari-
son of the effects of thyroxine treatment on
metamorphic sequence in intact and in hypophy-
sectomized tadpoles.
Summary
1. Comparison of the development of the in-
testine, pancreas, urinary bladder, tongue, gall
bladder, pituitary and thyroid glands in normal
and thyroxine-accelerated R. pipiens and R.
catesbeiana tadpoles at similar metamorphic
stages has shown that the accelerated animals
differ from the normal and that those accelerated
by 1:10,000,000 thyroxine were different from
those accelerated by 1:100,000,000.
2. In the same animals, different organs
showed either the same or more advanced devel-
opment than did the corresponding organs of
normal animals.
3. Treatment of R. pipiens tadpoles with
either concentration resulted in less developed
thyroids and pituitaries than were found in
1957]
McGovern & Charipper: Metamorphosis in Rana pipiens and Rana catesbeiana
49
naturally metamorphosed animals of the same
stage.
4. The responses of the tadpoles of the two
species to the same experimental treatment were
much alike in every point compared despite the
difference in the length of their tadpole periods.
5. The data fail to support the theory that
the sequence of metamorphic events is depen-
dent upon a series of thyroxine thresholds in the
several organs, but indicate some probability that
the sequence is dependent in part on thyroxine
concentration.
6. It is suggested that normal metamorphic
sequence may result from a thyroid-pituitary
relationship which changes with time and oper-
ates at the tissue-cellular level rather than at the
organ level.
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D’Angelo, S. A. & H. A. Charipper
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Gasche, P.
1940. Beeinflussung der Umwandlungsgeschwin-
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(Thyroxineinwirkung und Injektion von
thyreotropen Hypophysenvordenlappen-
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1943. The effect of thiourea on the development
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fluence of specific organs on growth and
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knowledge of organs with internal se-
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further contribution to the knowledge of
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Helff, O. M., & M. C. Mellicker.
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sylvatica, including the histogenesis of
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and the mucous glands. Amer. J. Anat.,
68: 339-369.
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1919b. Observation on thyroidless R. sylvatica
larvae kept through the second season of
normal metamorphosis. Anat. Rec., 16:
152.
Ingram, W. R.
1928. Interrelation of pituitary and thyroid in
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Jane, R. G.
1934. Studies on the amphibian digestive system,
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1916. Zur Frage der Wirkung von Schildriise
und Thymus auf Froschlarven. Pfliig.
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1936. A cytological study of the digestive sys-
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Kuntz, a.
1922. Metamorphic changes in the digestive
system in Rana pipiens and Ambystoma
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Lynn, W. G., & H. E. Wachowski
1951. The thyroid gland and its functions in
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SCHLIEFER, W. *
1935. Die Entwicklung der Hypophyse bei
Larven von Bufo vulgaris bis zur Meta-
morphose. Zool. Jb. Anat. u. Ontog., 59:
383-454.
SCHREIBER, G.
1934a. L’applicazione delle leggi d’azioni degli
ormoni alia metamorfosi degli Anura.
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1934b. Le disarmonie de la metamorfosi degli
anfibi e loro significato nella fisiologia
dello sviluppo. Boll. Soc. Ital. Biol.
Sperim., 9: 1211-1212.
Smith, P. E.
1916a. Experimental ablation of the hypophysis
in the frog embryo. Science, 44: 280-282.
1916b. The effect of hypophysectomy in early
embryo upon the growth and later de-
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64.
1918. The growth of normal and hypophysec-
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Speidel, C. C.
1926. Studies of hyperthyroidism. III. Bile pig-
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J. Exptl. Med., 43: 703-712.
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ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
VOLUME 42 • PART 2 • AUGUST 23, 1957 • NUMBERS 4 TO 6
PUBLISHED BY THE SOCIETY
The ZOOLOGICAL PARK, New York
Contents
PAGE
4. On the Bramid Fishes of the Gulf of Mexico. By Giles W. Mead. Plates
I-III 51
5. The Relation of Oxygen Consumption to Temperature in Some Tropical,
Temperate and Boreal Anuran Amphibians. By Richard E. Tashian &
Carleton Ray. Text-figures 1 & 2 63
6. Basic Patterns of Display in Fiddler Crabs (Ocypodidae, Genus Uca).
By Jocelyn Crane. Plate I; Text-figures 1-4 69
4
On the Bramid Fishes of the Gulf of Mexico^
Giles W. Mead^
(Plates I-III)
Contents
I. Introduction 51
II. Material and Acknowledgements 51
III. Taractes longipinnis 52
IV. Collybus drachme 58
V. Summary 59
VI. Literature Cited 59
I. Introduction
Berg (1947: 474) includes the Pteraclidae
and Steinegeriidae within the Bramidae.
Although this arrangement may be a
natural one, it is convenient here to follow the
older classifications {e. g. Jordan, 1923: 181)
and restrict the family to the genera Brama,
Taractes, Collybus and Eumegistus and their
synonyms. The Pteraclidae can be separated
from the Bramidae by the longer bases and
more anterior origins of their vertical fins. The
Steinegeriidae includes only Steinegeria rube-
scens Jordan & Evermann (1887) of which only
the type is known. This specimen was collected
from the stomach of a red “grouper” which was
caught in the Gulf of Mexico off Pensacola,
Florida. It is in poor condition. Although the
systematic position of this fish has not been es-
tablished, its nearly vertical mouth, non-falcate
dorsal and anal fins, the enlarged spines of the
caudal peduncle and its nearly flat forehead sepa-
rate it from Taractes, Collybus and Brama of
similar size. Steinegeria rubescens will be the
subject of a later paper, but is excluded from
further comment here. A revision of the major
classification of the bramid-like fishes is needed
as badly as an analysis of the species and of the
allometric growth associated with each.
The family, as so restricted, is represented in
^Contribution No. 973, Department of Tropical Re-
search, New York Zoological Society.
^Ichthyological Laboratory U. S. Fish and Wildlife
Service, U. S. National Museum, Washington 25, D. C.
the Gulf of Mexico by two species: Collybus
drachme, reported here for the first time from
five off-shore localities; and a species of Taractes
which I have called T. longipinnis and which is
also first recorded here from the Gulf of Mex-
ico. Although my material is inadequate for an
analysis of the non-Gulf bramids, I have re-
viewed some of these in order to more clearly
establish the identity of these Gulf of Mexico
specimens.
“Length” as used here refers to standard
length unless otherwise qualified. All measure-
ments over 100 mm. were made with dividers
and recorded in millimeters. Measurements be-
tween 5 and 100 mm. were made with dividers
or dial calipers and recorded in half millimeters.
Measurements of less than five mm. were taken
with a calibrated ocular grid micrometer and
were recorded to the nearest tenth of a milli-
meter.
II. Material and Acknowledgements
The first Gulf bramid which came to my
attention was a Taractes 74.0 mm. in standard
length (85 mm. fork length) taken by the U.S.
Fish and Wildlife Service vessel Oregon at sta-
tion no. 1043, an experimental tuna long-line
station occupied on May 11, 1954, at Lat. 24°
16' N., Long. 92° 00' W. The specimen was
found in the stomach of a yellowfin tuna (Neo-
thunnus albacares) which was caught at a depth
of less than 35 fathoms. The superficial fin mem-
branes had been largely digested and some of
the scales, head bones and fin rays were dam-
aged, but the fish was otherwise in good con-
dition.
Four adult Taractes were caught on later
Oregon cruises. All were taken by the long-line
at depths less than 35 fathoms. These specimens,
376, 371, 357 and 349 mm. in standard length
(418, 413, 394 and 390 mm. in fork length
respectively), came from the following two
Oregon stations:
51
52
Zoologica: New York Zoological Society
[42: 4
1173 28° 54' N. Lat., 88° 02' W. Long.,
August 25, 1954
1317 28° 23' N., Lat., 88° 43' W. Long.,
June 11, 1955
Three of these fish are now in the collections of
the U.S. National Museum, where they bear the
catalogue numbers US 157793 and US 164328.
These specimens, and the types of Collybus
drachme and Steinegeria rubescens, were made
available by Dr. Leonard P. Schultz, Curator
of Fishes.
Ten juvenile specimens of Collybus drachme
were collected at the following Oregon stations :
1043 24° 16' N. Lat., 92° 00' W. Long.,
May 11, 1954
1065 20° 34' N. Lat., 95° 37' W. Long.,
May 21, 1954
1377 27° 43' N. Lat., 88° 43' W. Long.,
August, 1955
1484 20° 50' N. Lat., 95° 53' W. Long.,
April 3, 1956
1486 22° 25' N. Lat., 97° 00' W. Long.,
April 4, 1956
I am indebted to Stewart Springer and Harvey
R. Bullis, Jr., of the U.S. Fish and Wildlife
Service, Pascagoula, Mississippi, and Edgar L.
Arnold, Jr., of the U.S. Fish and Wildlife
Service, Galveston, Texas, for the original
preservation of this Oregon material.
G. E. Maul, Curator of Fishes in the Museu
Municipal do Funchal, Funchal, Madeira, has
kindly sent information regarding Taractes from
Madeiran waters, and Dr. E. H. Bryan, Jr.,
Curator of Collections in the Bishop Museum,
Honolulu, has sent a photograph of a cast of
Collybus drachme.
Comparative bramid material from the Ber-
muda collections of Dr. William Beebe is now
deposited in the Natural History Museum of
Stanford University and was made available
by Dr. George S. Myers.
The large Taractes from Nova Scotia which
was reported by Bigelow & Schroeder (1929;
MCZ no. 31598) and a photograph of a 570
mm. (fork length) Portuguese Taractes sent to
these authors by Dr. Alfredo Ramalho were
made available by Drs. Henry B. Bigelow and
William C. Schroeder of Harvard University
and the Woods Hole Oceanographic Institution.
III. Taractes longipinnis
Since my study of the six western North At-
lantic Taractes listed above indicates that they
are specifically identical regardless of certain
differences among them, it is necessary to de-
scribe the more striking similarities and differ-
ences among the individuals of this series.
A 357 mm. Gulf specimen (PI. I, Fig. 1) was
first compared with the 618 mm. fish reported
by Bigelow & Schroeder (1929) from Cape
Sable, Nova Scotia. These fish differ in several
respects. In the smaller fish the ventral fins
originate under the middle of the bases of the
pectoral fins, while in the Nova Scotia specimen
their origins are beneath the anterior edges of
the pectoral bases. The 357 mm. specimen also
has relatively longer dorsal and anal lobes than
the 618 mm.i specimen, but, as Barnard (1948:
357) has suggested and as will be more fully
discussed below, a progressive relative shorten-
ing of these fin lobes accompanies an increase
in length of the adult fish. The 357 mm. indi-
vidual also differs in having spines on the pos-
terior body scales, and its eye is a little smaller
and more nearly circular than that of the larger
fish. Differences such as these accompany the
development of Brama rail and are therefore of
doubtful phylogenetic significance here. The fin
membrane connecting the posterior dorsal and
anal fin rays extends beyond the scaly sheath to
the tips of the fin rays in the 357 mm. fish, while
in the 618 mm. one this membrane is limited
to the proximal two-thirds of the fin; it does not
extend beyond the scaly sheath.
In all other details, such as the distinctive
shape of the caudal fin, the presence and nature
of the transverse caudal grooves which lie im-
mediately in advance of the dorsal and ventral
procurrent caudal rays, the dentition, the gill
rakers and their minute accessory spines and
tubercles and the general body form (exclusive
of fins), the 357 mm. Gulf of Mexico individual
is the same as that from Nova Scotia.
The differences between the 74 mm. juvenile
(PI. I, Fig. 2) and the 357 mm. adult are more
striking. In pattern and number of scales, if not
in the shape of the individual scale, the speci-
mens are similar. The larger fish has 44 rows of
scales between the upper margin of the gill
opening and the base of the mid-caudal rays
(where a sharp change in scale size occurs);
the smaller one has 45. The number of longi-
tudinal rows, from the mid-dorsal line in front
of the origin of the dorsal fin to the mid-ventral
line before the anal origin, is 27 in the larger
fish and 26 in the smaller. The adult has 17
scales along the ventral mid-line between the
origin of the ventrals and anus while the small
fish has 15. Although the fin membranes are
partially missing on the juvenile, traces of scale
pockets similar to those of the adult can be
found on the dorsal, anal and caudal fins. The
distinctive axillary scales of both pectoral and
ventral fins are alike in the two fish. The snout,
branchiostegal membrane and rami of the lower
jaw are equally scaleless in the two individuals.
The character of the individual scales, on the
1957]
Mead: On the Bramid Fishes of the Gulf of Mexico
53
other hand, differs greatly between these two
fish. Each body scale of the 74 mm. individual
bears a strong spine, a characteristic of all known
juvenile bramids and pteraclids. Considering
only the spines along the median row of scales,
those anterior to the 34th scale are directed
backward; the last ten are directed forward. In
the 357 mm. fish only the antrose spines on the
posterior half of the body remain. All scales of
Bigelow & Schroeder’s 618 mm. fish are smooth.
Each scale spine of the 74 and 357 mm. speci-
mens originates near the center of the scale
and protrudes through a notch in the posterior
edge of the preceding scale. As the spines de-
crease in size anteriorly in the 357 mm. indi-
vidual, these notches become less prominent.
The same trend is apparent when a scale of the
74 mm. specimen is compared with one from the
same point on the body of the 357 and 618 mm.
specimens. Accompanying this change in scale
form is a decrease in height of each scale. In
the juvenile, the greatest height of a scale taken
from the side of the body behind the pectoral
fin is 3.5 to 4 times its greatest width, a ratio
approaching the extremes encountered in such
fishes as the grammicolepid zeoids and the
bramid genus Collybus. The height of a scale
from the same body location on the larger speci-
mens is only one and one-half times its great-
est width.
The premaxillary and mandibular teeth of the
357 and 74 mm. fish are very similar. The larg-
est teeth of both jaws are the inner, anterior
series of recurved canines. Some of the outer
teeth of both jaws are directed slightly outward,
a condition more apparent in the juvenile, in
which the lips are somewhat macerated. The
palatine teeth of the larger fish are better de-
veloped than those of the smaller. The vomer
is toothless in both.
In number and shape of gill rakers and in
position and degree of development of the ac-
cessory spines and hooks on each raker there
are no significant differences between the 74
and 357 mm. fish. Exclusive of the rudiments,
each has a raker count of 2-f-7. In addition, the
epibranchial has three rudiments, which increase
in length ventrally, and the hypobranchial has
four. Each epibranchial raker has one or two
medially-directed spines, while each raker of
the ceratobranchial has three or four along its
distal half. Each rudiment has a cluster of spines
at its tip. In each fish there are also 4-f9-|-4
spine-bearing tubercles on the inner side of the
first gill arch. The 357 mm. specimen differs
from the juvenile only in having 4 rather than 3
rudiments on the epibranchial, 6 instead of 4
on the hypobranchial, and in better developed
spinules on the gill rakers and tubercles.
It is in the position and nature of the fins that
the greatest differences between the juvenile
and the adult are seen. Paramount among these
is the nature of the dorsal and anal fins— falcate,
with the rays filamentous and scarcely branched
in the juvenile, less attenuated and with the rays
branched in the larger fish (Table 1). The ap-
parent difference in the position of the ventral
fin could easily accompany growth from 74 to
357 mm., as could an increase in the amount
of dermal material which supports the dorsal,
anal and caudal lobes. A reduction in the length
of filamentous fin rays is also undoubtedly a
consequence of growth; such losses are known
to occur during the development of many diverse
species. Many fishes, conservative in form as
adults, are adorned with various filamentous
appendages as young. The change from lack of
true branching in the fin rays of the juvenile
cannot as readily be attributed to growth as
these other fin differences, although juveniles
of Brama have undivided or very weakly divided
rays which later become fully branched (Liitken,
1880, pi. 4; Schmidt, 1918: 5).
No lateral line is visible on the 618 mm.
specimen reported by Bigelow & Schroeder. The
357 mm. specimen, on the other hand, displays
a series of about 1 3 modified scales which appear
to represent a poorly-developed lateral line.
This series begins above the upper end of the
gill opening and arches posteriorly to a point
below the middle of the dorsal lobe. A series
of more typical lateral-line pores can be found
on the corresponding scale row in the 74 mm.
specimen. This series, also composed of 13
scales and terminating beneath the dorsal lobe,
is slightly more arched than in the larger fish.
In both adult fish the premaxillary terminates
anteriorly on a horizontal with the center of
the eye and extends posteriorly to beneath the
center of the eye. The free edge of the opercular
bone is thin, entire and irregular in outline. The
margins of the sub-, inter- and preopercle are
membraneous. There are slight differences in
degree of ossification and in the shape of the
orbit. In both fish the orbit is nearly circular,
while in Bigelow & Schroeder’s 618 mm. fish
the orbit is vertically elongate (the horizontal
axis is contained 1.3 times in the vertical).
Both specimens have prominent transverse
caudal grooves which lie above and below the
caudal peduncle anterior to the procurrent
caudal fin rays.
Although a direct comparison of the three
additional Gulf of Mexico fish with the 618 mm.
Taractes from Nova Scotia was not possible,
no significant differences were found between
these and the 357 mm. fish discussed above. The
counts and measurements of the sLx western
54 Zoologica: New York Zoological Society [42:4
Table L— Counts and Measurements of Six Specimens of Taractes longipinnis
FROM THE Western North Atlantic
Bigelow & Schroeder,
1929. Nova Scotia
Oregon Station 1173
Oregon Station 1173
Oregon Station 1112
Oregon Station 1317
Oregon Station 1043
Standard length (mm.)
618
376
371
357
349
74
Fork length (mm.)
670
418
413
394
390
85
Counts:
Dorsal fini
35
381/2
371/2
371/2
371/2
37
Anal fini
28
291/2
301/2
291/2
281/2
28
Pectoral fin
20
22/22
21/21
21/20
21/21
20/21
Scales^
43
46
45
44
47
43
Gill rakers^
2-1-7
2-f7
2-f8
2+7
2+7
2-1-7
Measurements (% of standard length)
Length of head
32.4
29.8
31.3
29.4
29.8
34.5
Length of snout
10.2
7.2
9.2
8.4
7.2
8.1
Length of premaxillary
14.7
14.4
15.1
14.6
13.8
16.9
Horizontal diameter of eye
6.6
5.9
6.5
5.9
6.3
10.8
Vertical diameter of eye
8.4
6.4
7.3
6.7
6.9
10.1
Width of interorbital
12.9
12.5
12.7
12.3
12.3
12.1
Greatest depth of body
52.1
54.5
55.0
55.5
55.3
58.8
Least depth of caudal peduncle
7.4
6.6
6.2
5.9
6.3
8.1
Snout to origin of anal fin
56.74
53.7
55.2
58.8
56.1
54.0
Snout to origin of ventral fin
36.54
32.5
34.2
37.5
33.8
30.4
Snout to origin of dorsal fin
44.84
41.8
44.2
41.2
44.4
43.2
Height of dorsal lobe
28.3
49.5
51.7
47.9
50.7
63.5
Height of anal lobe
28.2
53.5
54.7
53.8
53.0
70.3
Length of mid-caudal rays
8.5
11.2
11.3
10.4
11.7
14.9
Length of pectoral fin
38.2
39.9
37.7
35.0
36.4
21.6
Length of ventral fin
7.0
8.5
6.7
8.4
8.3
12.1
lA combined count is used here since it is impossible to distinguish accurately between spines and soft rays
without removing the overlying skin and scales.
“The row of scales which extends from the upper end of the gill cleft to the base of the caudal fin. exclusive
of the small scales which cover the caudal base.
^Exclusive of rudiments.
^From Bigelow & Schroeder’s figure (1929, pi. 1).
North Atlantic specimens are recorded in
Table 1.
I mention the shapes of the caudal fins of
these four Gulf of Mexico adults since they may
be of later taxonomic interest. Two nominal
species of Taractes, T. brevoorti and T. saussuri,
are characterized by their biconcave caudal fins,
i. e. the central rays of that fin are longer than
those fianking them. In one of the four adult
Taractes from the Gulf the central rays equal
in length those on either side; in two others they
are very slightly shorter, and in the fourth they
are slightly longer. In none, however, is the
margin of the caudal as strongly biconcave as
in T. brevoorti.
In summary, the six western North Atlantic
specimens of Taractes differ in the following
respects: the shape of the caudal and vertical
fins, form of the individual scales, presence or
absence of poorly-developed lateral-line scales,
shape of the eye, slight differences in body pro-
portion, presence or absence of branching in
dorsal and anal rays, and extent of the dorsal
and anal fin membranes. Growth from juvenile
to adult occurs in a less conservative manner
in the bramids than in most acanthopterygian
1957]
Mead: On the Bramid Fishes of the Gulf of Mexico
55
fishes, in which an array of distinguishing fea-
tures such as these would be of considerable
taxonomic and nomenclatorial interest. In the
Bramidae, however, differences between indi-
viduals of different size must be evaluated with
reference to the great ontogenetic changes which
are known to take place.
I do not hesitate to ascribe the decrease in
relative height of the dorsal and anal lobes with
increasing size to normal aUometric adult
growth (PI. II, Fig. 3; Barnard, 1948: 374).
Similarly, all known bramid juveniles have a
spine on each scale which is gradually lost with
growth; it is therefore appropriate to believe that
the reduction in scale armature with increase in
size in our series is a function of growth. I like-
wise ascribe the progressive shortening of the
relative height of the individual scale to normal
ontogenetic change, but feel hesitant about the
significance of those scales in the 74 and 349-376
mm. fish which appear to represent a lateral
line. Because of the intricate pattern and bizarre
form of all scales it is difficult to trace the reduc-
tion in these possible lateral-line rudiments, if
such a reduction occurs. Fitch (1953: 539) re-
ported a lateral line in his 590 mm. California
Taractes, and Iordan’s Eumegistus illustris, ob-
viously closely allied to, if not congeneric with,
Taractes, has a lateral line. The lateral line may
be variable in Taractes and will deserve close
attention when a larger series becomes available.
If this evaluation is correct, the five western
North Atlantic adult Taractes can be referred
to a single species, for the only difference which
distinguishes the four Gulf of Mexico fish from
the larger one from Nova Scotia is the extent
of the posterior dorsal and anal fin membranes.
In the fish of 618 mm. the tips of the posterior
dorsal and anal rays are free from the mem-
brane; in the Gulf of Mexico fish the tips lie
within this membrane. This characteristic is
among those listed by Bigelow & Schroeder
(1929: 45) as diagnostic of Taractes princeps.
I hesitate to afford it such distinction, since the
only Taractes which possess it are the largest
known individuals: Bigelow & Schroeder’s of
618 mm., Johnson’s types of T. princeps (27 to
33 inches), Fitch’s California fish (590 mm.)
and, if it be admitted to Taractes, Jordan’s
Eumegistus illustris (about 608 mm.). In fishes
such as some scombrids and carangids, the
dorsal and anal finlets, separate and free from
a connecting membrane in the large adult, are
completely or partially enclosed in a fin mem-
brane as juveniles and young adults. Parallel fin
development mav occur among the bramids.
An important difference between the 74 mm.
Gulf juvenile and the five adults makes its iden-
tity with them less certain. In the juvenile Tar-
actes the posterior dorsal and anal rays are
unbranched, although there is an indistinct line
distally which separates the anterior half from
the posterior. In the adults these rays are
branched nearly from their bases. Since the rays
of the juvenile bramids figured by Liitken
(1880), Sanzo (1928) and others are similar to
those of our juvenile although the adults have
completely divided rays, the separation of
species by this character can be questioned.
However, because of this difference, the mor-
phometric differences shown in Table 1, the
great difference in size between this 74 mm. fish
and the next-smallest known Taractes (300 mm.
fork length— Barnard, 1948: 375, pi. 10), and
the general confusion attending past work on
the juveniles of most pelagic spiny-rayed fishes,
this specimen was compared with other species
closely related to it in appearance if not in
phylogeny.
The scale pattern, fin structure, general phy-
siognomy and especially the presence of distinct
transverse caudal grooves distinguish our juven-
ile from all of the bramids other than Taractes
to which I have compared it. Brama raii and
Collybus drachme of about the same length as
our fish (85 and 81 mm. in fork length respec-
tively) are represented in the collections of Stan-
ford University’s Natural History Museum.
These are totally different from the juvenile
Taractes. Although the young of Eumegistus
illustris, T. brevoorti, T. raschi and T. saussuri
are unknown, none of the adults has the promi-
nent transverse groove on the caudal peduncle,
a groove which is well developed in our juvenile.
The prolonged dorsal and anal fin rays, which
are filamentous, with only the suggestion of
branching, are reminiscent of those of the
heterogeneous assemblage of species reported by
Jordan (1919), but since the dorsal fin of our
Taractes originates well behind the eye and the
anal a considerable distance behind the ventral
fins, I did not study in detail Pteraclis and the
various nominal species of Pterycombus and
Centropholis (Barnard, 1927: 598; Kuronuma,
1941: 56; Belloc, 1927: 239; and others). The
juvenile Taractes is also quite distinct from the
berycoid fishes which probably resemble it when
young: Trachyberyx barretoi and Trachichtodes
spinosus. There is little reason to believe that
the 74 mm. Gulf juvenile belongs to a genus
other than Taractes.
Taxonomic status of the western North At-
lantic Taractes.— Although most of the nomen-
clatorial and taxonomic difficulties which con-
fronted Bigelow & Schroeder when they reviewed
this genus in 1929 are still extant, more recent
work and additional material justify a reap-
praisal of Taractes taxonomy here.
56
Zoologica: New York Zoological Society
[42: 4
With the exclusion of Collybus Snyder and
Eumegistus Jordan & Jordan, the Bramidae can
be divided into the two natural groups proposed
by Smitt (1892-95) and recognized by Bigelow
& Schroeder (1929): those forms with more
than 70 scales in a median series, Brama, and
those with fewer than 60, Taractes. I am con-
cerned here only with the few-scaled species, of
which the following nominal representatives are
known:
Taractes asper (Yowe, 1843) Madeira
T. longipinnis (Lowe, 1843) Madeira
T. brevoorti (Poey, 1861) Cuba
T. raschi (Esmark, 1862) Norway
T. pnnce/75 Johnson, 1863 Madeira
T. saussuri (Lunel, 1866) Cuba
T. steindachneri (Doderlein, 1883) Japan
T. platycephalus Matsubara, 1936 Japan
T. miltonis, Whitley, 1938 Australia
I have tentatively omitted Eumegistus illustris
Jordan & Jordan (1922: 36) from Taractes, al-
though its close resemblance cannot be ignored.
Because of its lateral line and smooth-edged
scales, Eumegistus has been considered generi-
cally distinct from Taractes, but I have noted
above the variation in the lateral line found in
specimens of Taractes, and both the scale spine
and the notch in the rear edge of each scale in
young Brama and Taractes disappear with age.
Jordan & Jordan’s type specimen of E. illustris
was a large individual, about 608 mm. in length.
There are, however, meristic differences between
E. illustris and the nominal species of Taractes.
Fowler (1938: 44) based his description of
Brama leucotaenia on a juvenile specimen 22.5
mm. in standard length from the Philippine
Islands. Fowler compares his fish with Brama
raii and distinguishes it from that species by the
pigmentation of the dorsal fin. However, his
comparison of B. leucotaenia with B. raii is of
little moment since the large scales (53 in a
median longitudinal series) show it to be more
closely related to Taractes. Although I have ex-
amined Fowler’s type (U.S. National Museum
no. US 98817) , I have not undertaken a detailed
study of its relationship. Since in scale count,
number of gill rakers (about 5-1-10) and posi-
tion and extent of the lateral line it more nearly
resembles Eumegistus illustris than any known
Taractes, I consider this juvenile fish generically
and probably specifically identical with Eume-
gistus illustris.
Collybus bears little resemblance to Taractes
or Brama. More will be said of Collybus later.
The identity of Taractes as per is uncertain.
Lowe’s description is inadequate and the type
(from Madeira) cannot be located. This fish has
traditionally been considered a young stage of
T. longipinnis, which Lowe described on the
preceding page of the same paper (1843: 82).
However, T. asper may be the young of an en-
tirely different bramid-like fish. Mr. G. E. Maul
(in correspondence) has pointed out to me the
similarity between Lowe’s description of T.
asper and the berycoid species Trachyberyx
barretoi Roule. I can contribute no original in-
formation in clarification of this question and
will omit Taractes asper from further considera-
tion here.
Omitting Eumegistus, Collybus and T. asper,
a natural subdivision of the remaining Taractes
species suggests itself. Gross differences in body
proportions and fin structure distinguish Tar-
actes longipinnis of Lowe and its allies from the
type of T. raschi, a second specimen referred to
T. longipinnis by Smitt et al. (1892-95 : 80) , and
Matsubara’s T. platycephalus. In body form, fin
structure and scale pattern, if not in more de-
tailed features, there is little in common between
these three fishes and representatives of the T.
longipinnis group of similar size. The type fig-
ures of T. raschi and T. platycephalus are repro-
duced here (PI. II, Fig. 4; PI. Ill, Fig. 5). It is
unfortunate that Matsubara did not consider in
more detail the resemblance between his T. pla-
tycephalus and Esmark’s type of T. raschi rather
than accepting Smitt’s synonymy, which places
T. raschi in the synonymy of T. longipinnis, and
basing his comparison on Bigelow & Schroeder’s
paper. I have seen neither species, but if the fin
structure and shape are subject to growth
changes as pronounced as those in Brama and
T. longipinnis, and if the vomerine dentition
is as variable in Taractes as Lunel found it to be
in Brama, it is difficult to characterize T. platy-
cephalus.
The second subdivision of Taractes includes
the deeper-bodied species in which both young
and adults have strongly falcate dorsal and anal
lobes, more steeply inclined foreheads, and dis-
tinctive squamation: T. longipinnis (Lowe, not
of Smitt), T. princeps, T. brevoorti, T. stein-
dachneri, T. saussuri and T. miltonis.
Doderlein’s T. steindachneri {in: Steindach-
ner & Doderlein, 1883, pi. 7) has been referred
to the synonymy of T. longipinnis by Steindach-
ner & Doderlein (1884: 174), a disposition ac-
cepted by Bigelow & Schroeder. It was resur-
rected by Jordan, Tanaka & Snyder (1913: 134),
Matsubara (1936) and others on the basis of
slight differences in scale and fin-ray counts. The
species might better have remained in synonymy.
Taractes brevoorti and T. saussuri (PI. Ill,
Fig. 6) are alike in having a double-concave
caudal fin. Although one of the Gulf of Mexico
adults which I have examined has central caudal
rays slightly longer than those flanking them.
1957]
Mead: On the Bramicl Fishes of the Gulf of Mexico
57
no recent specimens of Taractes approach the
extreme condition shown by T. saussuri. These
two species must be retained. T. brevoorti ap-
pears to be closely related to T. longipinnis. T.
saussuri, on the other hand, is intermediate be-
tween T. longipinnis and T. raschi in body form
and fin structure. Nothing further can be said
about these species until specimens become
available.
Whitley (1938: 193) distinguishes his Aus-
tralian Taractes miltonis from the fish described
by Bigelow & Schroeder (1929) as follows:
“Head, body, scale, and fin characters agreeing
excellently with the detailed description of the allied
Taractes princeps (Johnson) recently given by Bige-
low and Schroeder (Bull. Mus. Comp. Zool. Har-
vard, Ixix., 2, February, 1929, p. 45 and plate — )
but is of slightly larger size and is distinguished by
having the eye-diameter about one-fourth, instead
of about one-fifth the length of the head; anal lobe
considerably shorter than head; comparatively
longer pectoral and ventral fins; distance from ven-
tral origin to anal origin notably less than length of
head; different gill-rakers, etc.”
Whitley does not describe the differences in
gill rakers. The horizontal diameter of the eye
of his specimen is 7.2 percent, of standard
length cf. 5. 9-6.6 in the western North Atlantic
adult specimens (Table 1) ; the vertical diameter
is 8.3 cf. 6. 4-8.4. The head length he reported,
200 mm. or 27.8 percent, of standard length, is
2 percent, shorter than any known western
North Atlantic adult. I have noted elsewhere
the relative reduction in the height of the anal
fin lobe with increasing length of fish in Taractes
longipinnis and Whitley’s measurement, 140
mm. or 19.5 percent, of standard length, is not
out of accord with this growth change. There is
no notable difference in the distance between
the origins of the ventral and anal fins between
Whitley’s fish and Bigelow & Schroeder’s (about
1 percent, of standard length), and his measure-
ments of the lengths of the pectoral and ventral
fins (35.9 and 7.65 percent, respectively) are
within the range of the adult western North
Atlantic material (35.0-39.9 percent, and 6.7-
8.5 percent, respectively). Taractes miltonis, as
deseribed by Whitley, is devoid of distinguishing
eharacteristics. This species, along with T. stein-
dachneri and T. princeps, should go into the
synonymy of T. longipinnis.
Taractes longipinnis and T. princeps were sub-
jected to a detailed comparison by Bigelow &
Schroeder (1929). These authors tentatively
retained both species, listing the following com-
bination of characters as diagnostie of T. prin-
ceps (p. 45) :
“1. Very deeply lunate tail.
2. The fact that the low rays of the anal and
dorsal fins are distaUy free from the mem-
brane for about a third of their length.
3. Great length of the anterior dorsal and anal
rays.
4. Scales smooth, without hooks or spines (at
least in adult) .
5. Caudal pits present.”
Bigelow & Schroeder’s recognition of both
species reflects a careful study of the problem
and is the more conservative course. Since
neither type is still extant, an understanding of
the relationship between Taractes longipinnis
and T. princeps awaits the comparison of series
of specimens from the type locahty, Madeira.
Maul, however, who has seen many Taractes
during his long association with the Madeiran
fishery, can distinguish but one species, which
he refers to T. longipinnis (personal communi-
cation) . If it seems probable that a large series
of specimens would show that the two nominal
species are identical, it would be reasonable to
combine the two at this time— the course rec-
ommended by those ichthyologists who have
been able to examine more than one specimen.
I prefer this alternative, for my material influ-
ences the interpretation of three of the distin-
guishing characteristics afforded T. princeps by
Bigelow & Schroeder and the remaining two are
of dubious significance in the absence of con-
firming material. If my 74 mm. juvenile and
Barnard’s 268 mm. adult are correctly referred
to Taractes, the shape of the caudal fin changes
with growth from a shallow fork to the deeply
lunate form seen in the largest specimen. Simi-
larly, there can be little doubt that the lobes of
the dorsal and anal become relatively shorter
during adult growth, and that there occurs a loss
of scale spines and a change in the shape of the
scale itself during development. Two peculiari-
ties remain to characterize T. princeps: the pres-
enee of caudal pits and the absence of an inter-
radial membrane between the posterior dorsal
and anal rays. Lowe did not mention caudal
grooves in his brief description of T. longipinnis.
They may or may not have been present. As
noted elsewhere, the dorsal and anal rays are
free from the interradial membrane only in the
largest Taractes, and this difference alone seems
inadequate for the separation of T. longipinnis
and T. princeps.
I have not attempted to review the generic
nomenclature. I cannot concur in deBuen’s
(1935: 102) union of Brama and Taractes or
with his suppression of the generic name Brama
in favor of Lepidotus. Lepidotus Asso (1801)
was used by deBuen (1935: 102), Whitley
(1938: 191) and Fowler (1949: 74) in place of
Brama (Bloch & Schneider, 1801: 98) — the
needless suppression of a generic name which
58
Zoologica: New York Zoological Society
[42: 4
had been universally accepted for more than a
century. The respective dates of publication,
within the year 1801, have not been determined
or at least were not discussed by the recent pro-
ponents of the generic name Lepidotus. Tar-
actes is used here because it has been applied
customarily to the species discussed in this paper
and because there is no nomenclatorial reason
for its suppression.
Since I have concluded that all of the indi-
viduals which are known from the western North
Atlantic are alike and that this series possesses
features which weaken the argument provided
by Bigelow & Schroeder for the separation of
T. princeps and T. longipinnis, I suggest that
the population represented by these individuals
should bear the name Taractes longipinnis Lowe.
I propose the following tentative synopsis of
the species of Taractes:
I. Fewer than 50 scales in a median longitudinal
series exclusive of the small scales overlying the
base of the central caudal fin rays.
A. Caudal fin “rounded.” (A species of un-
certain identity) T. asper
AA. Caudal fin emarginate, forked or bicon-
cave.
B. Standard length more than twice the
greatest body depth. Pectoral fin with 17-18
rays.
C. Caudal fin emarginate. Forehead con-
cave, less than an eye’s diameter between
upper edge of eye and dorsal profile of
head.
D. Vomer toothed. (North Atlantic;
syn.: T. longipinnis of Smitt, not of
Lowe) T. raschi
DD. Vomer toothless. (Japan).
T. platycephalus
CC. Caudal fin biconcave; tips of cen-
tral rays on a line with tips of the outer
caudal rays. More than an eye’s diameter
between eye and dorsal profile. Forehead
convex. (Cuba) T. saussuri
BB. Standard length less than twice the
greatest body depth. Pectoral fin with 19-
21 rays.
E. Caudal fin emarginate. (Atlantic and
Pacific; syn.: T. steindachneri, T. prin-
ceps, T. miltonis, not T. longipinnis of
Smitt) T. longipinnis
EE. Caudal fin biconcave. (Cuba).
T. brevoorti
IV. Collybus drachme
As well as from the type locality (Hawaii),
Collybus drachme Snyder (1904: 525; fig. 7)
has been caught off Bermuda (Kanazawa, 1952:
80). Twelve specimens have been taken in the
Gulf of Mexico, all from the stomachs of yel-
lowfin tuna and lancet fish (Alepisaurus) caught
at the five Oregon stations listed earlier in this
paper. They range in standard length from 33.0
to 52.5 mm. and are in various stages of diges-
tion. A 42.5 mm. fish was cleared and stained
and examined for skeletal characteristics. I have
also compared the twelve specimens with
Snyder’s type and cotypes, which are now in
the U.S. National Museum and in the Natural
History Museum, Stanford University.
I use Snyder’s trivial name, drachme, for these
Gulf of Mexico specimens since I have found no
significant differences between representatives
from the two oceans.
The largest known Collybus which has been
described or figured is Snyder’s 81 mm. Ha-
waiian type. Fowler (1928: 138) reported three
larger (167-186 mm.) specimens obtained in
Honolulu and now in the Bishop Museum and
Jordan & Jordan (1922: 35) referred to a cast
of a large individual, also in the Bishop Museum.
A photograph of this cast portrays a fish about
155 mm. in length, but few details of the original
fish can be discerned. The physiognomy and
shape and position of the fins are similar to
those of Brama and to Snyder’s larger specimens
of Collybus. This cast was certainly not made
from a Taractes. Some authors have suggested
that Collybus represents the young of a species
of Brama or Taractes, a view that I am reluctant
to accept. The scale count separates Collybus
from Brama (45-55 cf. 70-80 in Brama) and
our juvenile Taractes, 74 mm. long, is much
thicker-bodied and has more falcate fins than
any Collybus that I have seen. Collybus may
represent a young T. raschi or Eumegistus, but
pending evidence to the contrary, Collybus
drachme should be recognized.
The following diagnosis is a composite taken
from my twelve Gulf individuals, no one of
which is undamaged:
Body compressed, eliptical in outline with a
ventral profile more strongly convex than the
dorsal. Eye large and circular, not entering into
the dorsal profile, eye diameter 2.5-2.1 in head.
Mouth oblique, premaxillary extending to be-
neath middle of pupil. Lower jaw coterminal
with upper or slightly protruding. Head 3. 0-3. 4
in standard length; opercular bones smooth, al-
though the elongate scales overlying the free
edges are serrated. Greatest depth of body, at
origin of dorsal fin, 1.5-1. 8 in standard length.
Least depth of caudal peduncle 3. 5-4.5 in head.
Lateral line usually absent, occasionally present,
complete or incomplete. Head and body lightly
pigmented, a sprinkling of melanophores below
1957]
Mead: On the Bramid Fishes of the Gulf of Mexico
59
orbit, along bases of all fins, on peritoneum and
along dorsal edge of body.
D.: 32-34. A.: 28-29. P.: 20-21. Gill rakers:
2-3 + 8-9. Scales in a median series: 46-54;
about 18 horizontal rows on body, counted
obliquely upward and backward from the origin
of anal fin. Vertebrae (one specimen only) : 38.
Snout and forehead in front of center of eye
scaleless but covered with small pores. Re-
mainder of head and body scaled. Scales along
bases of dorsal and anal fins and in axil of pec-
toral and ventral fins. Scales varied, those on
head more strongly ctenoid than those on body.
All scales vertically elongated, those on middle
of body extremely attenuated, the width con-
tained about nine times in the height. All scales
with a vertical ridge and a central protruding
spine or knob. About 14 keeled scales along the
ventral midline between origin of pelvic fins and
that of anal.
Teeth on jaws; none on vomer or palatines.
Mandibular teeth in a single row posteriorly, a
band anteriorly. Outermost anterior teeth re-
curved and enlarged. Two fangs at inner edge of
anterior band at tip of mandible. Premaxillary
teeth similar to those on mandible, but without
anterior fangs.
Measurements, expressed as percent, of stand-
ard length: length of head, 29.9-33.9; length of
snout, 5.3-7.V; length of premaxillary, 14.7-16.9;
diameter of eye, 11.4-12.7. Greatest depth of
body, 56.8-66.2 (decreasing with increasing
length of fish) , least depth of caudal peduncle,
8.3-12.7. Height of dorsal lobe, 21.1-23.9; height
of anal fiji, 8.4-10.4; length of pectoral fin, 27.4-
32.2; length of ventral fin, 10.6-12.3.
V. Summary
Except for the controversial Steinegeria rube-
scens, no species of non-pteraclid bramid fish
has hitherto been reported from the Gulf of
Mexico. The off-shore collections of the U.S.
Fish and Wildlife Service vessel Oregon contain
representatives of two such species, which are
identified here with Taractes longipinnis Lowe
and Collybus drachme Snyder. Both are de-
scribed, growth changes in Taractes longipinnis
are discussed, and the nominal species of Tar-
actes are reviewed.
VI. Literature Cited
Asso Y DEL Rio, Ignacio Jordan de
1801. Introduccion a la ichthyologia oriental de
Espana. Anales Ciencias Nat., vol. 4, pp.
28-52. [Also published separately, Madrid,
1801, 28 pp.; not seen].
Barnard, K. H.
1925-27. A monograph of the marine fishes of
South Africa. Ann. South African Mus.,
vol. 21, 1065 pp., 37 pis.
1948. Further notes on South African marine
fishes. Ann. South African Mus., vol. 36,
pp. 341-406, 5 pis.
Belloc, Gerard
1927. Note preliminaire sur un poisson nouveau
du genre Centropholis. Ann. Soc. sci. nat.
Charente-Inferieure (La Rochelle), 1927,
pp. 239-243, 1 pi.
Berg, Leo S.
1947. Classification of fishes both lecent and
fossil. Edwards, Ann Arbor, 517 pp.
Bigelow, Henry B., & William C. Schroeder
1929. A rare bramid fish {Taractes princeps
Johnson) in the northwestern Atlantic.
Bull. Mus. Comp. Zoology, Harvard Coll.,
vol. 69, pp. 41-50, 1 pi.
Bloch, Marc Elieser, & Johann Gottlob
Schneider
1801. Systema Ichthyologiae iconibus ex illu-
stratum . . . , Berlin, 584 pp., 1 10 pis.
DE Buen, Fernando
1935. Fauna ictiologica. Catalogo de los peces
Ibericos: de la planicie continental, aquas
dulces, pelagicos y de los abismos proxi-
mos. Part 2. Notas y Res., Inst. Espan. de
Oceanogr., ser. 2, no. 89, pp. 91-149.
Esmark, Lauritz
1862. Beskrivelse over en ny fiskeart, Brama
raschii Esm. Forh. Vidensk. Selsk. Christi-
ania, (1861), pp. 238-247.
Fitch, John E.
1953. Extensions to known geographical distri-
butions of some marine fishes on the
Pacific coast. California Fish and Game,
vol. 39, pp. 539-552.
Fowler, Henry W.
1928. The fishes of Oceania. Mem. Bernice P.
Bishop Museum, vol. 10, 540 pp., 49 pis.
1938. Descriptions of new fishes obtained by the
United States Bureau of Fisheries Steamer
“Albatross,” chiefly in Philippine seas and
adjacent waters. Proc. U.S. Nat. Mus.,
vol. 85, pp. 31-135.
1949. The fishes of Oceania, Supplement 3.
Mem. Bernice P. Bishop Mus., vol. 12, no.
2, pp. 37-186.
Hilgendorf, Franz Martin
1878. Ueber das Vorkommen einer Brama-Art
und einer neuen Fischgattung Centropho-
lis aus der Nachborschaft des Genus
Brama in den japanischen Meeren. Sitzber.
Ges. Naturf. Freunde Berlin, pp. 1-2.
Johnson, James Yate
1863. Descriptions of five new species of fishes
obtained at Madeira. Proc. Zool. Soc. Lon-
don, 1863, pp. 36-46.
60
Zoologica: New York Zoological Society
[42; 4
Jordan, David Starr
1919. On Elephenor, a new genus of fishes from
Japan. Ann. Carnegie Mus., vol. 12, nos.
2-4, pp. 329-342, pis. 54-58.
1923. A classification of fishes, including families
and genera as far as known. Stanford Uni-
versity Pubis., Univ. Ser.-Biol. Sci., vol. 3,
no. 2, pp. 79-243 -|- i-x.
Jordan, David Starr, & Barton W. Evermann
1887. Description of six new species of fishes
from the Gulf of Mexico, with notes on
other species. Proc. U.S. Nat. Mus., vol. 9,
pp. 466-476.
Jordan, David Starr, & Eric Knight Jordan
1922. A list of the fishes of Hawaii, with notes
and descriptions of new species. Mem.
Carnegie Mus., vol. 10, no. 1, pp. 1-92,
pis. 1-4.
Jordan, David Starr, Shigeho Tanaka &
John Otterbein Snyder
1913. A catalogue of the fishes of Japan. Journ.
Coll. Sci., Tokyo Imp. Univ., vol. 33, art.
1, 496 pp.
Kanazawa, Robert H.
1952. More new species and new records of
fishes from Bermuda. Fieldiana— Zoology,
vol. 34, no. 7, pp. 71-100.
Kuronuma, Katsuzo
1941. Notes on rare fishes taken off the Pacific
coast of Japan. Bull. Biogeographical Soc.
Japan, vol. 11, no. 8, pp. 37-67, pis. 1-2.
Lowe, R. T.
1843. Notices of fishes newly observed or dis-
covered in Madeira during the years 1840,
1841 and 1842. Proc. Zool. Soc. London,
pt. 11, pp. 81-95.
Lunel, Godefroy
1866. Revision du genre castagnole (Brama) et
description d’une espece nouvelle Brama
saussurii. Mem. Soc. Phys. et Hist. Nat.,
Geneve, vol. 18, pp. 165-196, pis. 1-2.
Lutken, C. F.
1880. Spolia Atlantica— Bidrag til Kundskab om
Formforandringer hos Fiske under deres
Vaext og Udvikling, saerlight hos nogle af
Atlanterhavets H0js0fiske. Vidensk. Selsk.
Skr., 5. Raekke, naturvidenskabelig og
mathemetisk Afd., vol. 12, no. 6, pp. 413-
613.
Matsubara, Kiyomatsu
1936. A new bramid fish found in Japan. Bull.
Japanese Soc. Sci. Fish., vol. 4, no. 5,
pp. 297-300.
PoEY Y Aloy, Felipe
1851-61. Memorias sobre la historia natural de
la isla de Cuba, acompanadas de sumarios
latinos y extractos en frances . . ., Havana,
2 vols., 427 pp.
Sanzo, Luigi
1928. Contributo alia conoscenza di uova e
larve di Brama raji Bl. Mem. Comitato
Talassografico Italiano, no. 147, 9 pp., 1 pi.
Schmidt, Johs.
1918. Bramidae, in Johs. Schmidt and A. Strub-
berg, Mediterranean Bramidae and Trichi-
uridae. Rep. Danish Oceanogr. Exped.
1908-10 to the Mediterranean and Adj.
Seas, vol. 2 (Biology), (no. 4). A. 6.,
15 pp.
Smitt, F. W. (ed), B. Fries, C. U. Ekstom
& C. SUNDERVALL
1892-95. A history of Scandinavian fishes, ed.
2, 1240 pp., 54 pis., Stockholm.
Snyder, John Otterbein
1904. A catalogue of the shore fishes collected
by the steamer Albatross about the Ha-
waiian Islands in 1902. Bull. U.S. Fish
Comm., vol. 22, pp. 513-538, 13 pis.
Steindachner, Franz, & L. Doderlein
1883. Beitrage zur kenntniss der fische Japans
(I). Denkschr. Akad. Wiss. Wien., vol. 47,
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1884. Beitrage zur kenntniss der fische Japans
(III) . Denkschr. Akad. Wiss. Wien., vol.
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Whitley, Gilbert
1938. Ray’s bream and its allies in Australia.
Australian Zoologist, vol. 9, pp. 191-194.
pi. 19.
1957]
Mead: On the Bramid Fishes of the Gulf of Mexico
61
EXPLANATION OF THE PLATES
Plate I
Fig. 1. Taractes longipinnis from Oregon station
1112 (Gulf of Mexico). Standard length:
357 mm. (Photograph courtesy of the
Woods Hole Oceanographic Institution).
Fig. 2. Taractes longipinnis from Oregon station
1043 (Gulf of Mexico). Drawn, with some
reconstruction, from a damaged juvenile
74 mm. in standard length. (Drawn by
Janet Roemhild).
Plate II
Fig. 3. Profiles of five specimens of Atlantic Tar-
actes longipinnis. From the smallest to the
largest, the data from which these profiles
were drawn were taken from (a) the 74
mm. Gulf of Mexico juvenile; (b) a South
African adult of about 268 mm., from Bar-
nard, 1948; (c) a 357 mm. Gulf of Mexico
adult; (d) a 618 mm. adult from Nova
Scotia, from Bigelow & Schroeder, 1929;
and (e) a South African adult of about 706
mm., from Barnard, 1948.
Fig. 4. Taractes raschi. After Smitt et. al., 1892-
95, p. 80, fig. 24; from Esmark, 1862, pi. 1.
Plate III
Fig. 5. Taractes platycephalus. From Matsubara,
1936, p. 297, fig. 1.
Fig. 6. Taractes saussuri. From Lunel, 1866, pi. 2.
Fig. 7 Collybus drachme from Oregon station
1065 (Gulf of Mexico). Standard length:
33.0 mm. (Drawn by Janet Roemhild).
MEAD
PLATE I
FIG. 1
FIG. 2
ON THE BRAMID FISHES OF THE GULF OF MEXICO
MEAD
PLATE II
FIG. 4
ON THE BRAMID FISHES OF THE GULF OF MEXICO
MEAD
PLATE III
FIG. 7
ON THE BRAMID FISHES OF THE GULF OF MEXICO
5
The Relation of Oxygen Consumption to Temperature in
Some Tropical, Temperate and Boreal Anuran Amphibians^
Richard E. Tashian^ & Carleton Ray^
Department of Tropical Research and New York Aquarium, New York Zoological Society
(Text-figures 1 & 2)
PREVIOUS studies on the geographic vari-
ation in climatic adaptation for poikilo-
therms have revealed much of physiolog-
ical, ecological and evolutionary importance.
The literature in this field has been thoroughly
reviewed by Prosser (1955) and Bullock (1955).
In respect to amphibians, investigations on the
geographic variation of developmental adapta-
tion in anurans (e.g. Moore, 1949; Volpe, 1954)
have been particularly significant. Relatively
little data, however, are available in which meta-
bolic rate and temperature have been compared
in amphibians from different latitudes. It is the
purpose of this study, therefore, to present an
analysis of the respiratory metabolism of some
tropical, temperate and boreal anurans when
compared at two similar temperature levels.
The tests were carried out on five species of
tropical frogs; Hyla maxima and Hyla crepitans
(Hylidae) ; Leptodactylus typhonius and Eu-
pemphix pustulosus (Leptodactylidae) and
Prostherapis trinitatis (Dentrobatidae). The
temperate and boreal forms included the toads
Bufo boreas boreas and Bufo boreas halophilus
(Bufonidae) ; and the frogs Rana clamitans and
Rana sylvatica (Ranidae) and Hyla crucifer.
The experimental work was undertaken at
Simla, field station of the Department of Trop-
ical Research, New York Zoological Society,
Arima Valley, Trinidad, B.W.I., and the Depart-
ment of Zoology, Columbia University, New
York, from December, 1954, to July, 1955.
^Contribution No. 974, Department of Tropical Re-
search, New York Zoological Society.
^Present address: University of Michigan, 1135 E.
Catherine St., Ann Arbor, Mich.
®This study was aided in part by the Arctic Institute
of North America through funds granted by the Office
of Naval Research.
We wish to thank Ernest Karlstrom of the
University of California for sending us the Bufo
boreas material from California. The assistance
of Rosemary Kenedy during the work at Simla
is also gratefully acknowledged. The respirom-
eters were kindly loaned to us by F. John Vern-
berg of Duke University.
Materials and Methods
All tropical frogs were collected in the imme-
diate vicinity of Simla in the northern range of
Trinidad at an elevation of around 800 ft. be-
tween December 26 and March 30. Hyla cru-
cifer was collected near New York City in late
April and early May, and Rana clamitans and
Rana sylvatica from Bondville, Vermont (1,500
ft.), on July 3 and 4. Breeding individuals of
Bufo boreas boreas were taken seven miles
northwest of Juneau, Alaska (sea level), be-
tween June 1 and 10, and post-breeding forms
of Bufo boreas halophilus from California at the
following localities: Richmond, Contra Costra
Co. (sea level) ; Russian River area, Sonora Co.
(sea level to 1,000 ft.) and Sequoia National
Park, Tulare Co. (7,500 ft.) between June 1 and
19. The five tropical species ranged in weight
from 0.5 to 45.5 gms., and the four temperate
and boreal species from 0.84 to 56.1 gms.
After collecting, the tropical frogs were main-
tained in aquaria at room temperature (20-25°
C.), and the temperate and boreal forms were
kept in aquaria in a constant temperature (15°
C.) room. All were fed regularly on adult or
larval insects.
Closed system volumetric respirometers of the
type developed by Flemister & Flemister ( 1951 )
were employed for the determination of the res-
piration rates. Tests were conducted in water
baths maintained at 25 ± .5° C. and 10 ± .5° C.
63
64
Zoologica: New York Zoological Society
[42: 5
Text-fig. 1. The relation of oxygen consumption to weight in five species of tropical anurans at two tem-
perature levels (h = regression coefficient; r = coefficient of correlation). Dashed lines represent regres-
sion slopes for temperate species.
for the tropical forms and 24 ± .1° C. and 14 ±
.1° C. for the temperate and boreal animals.
After allowing a half hour for thermal equilib-
rium the tests were then run for a period of three
or four hours. The volumes obtained for oxygen
consumption have been corrected to standard
temperature.
Results
Text-figures 1 and 2 represent the double-
logarithmic regression plots of weights to metab-
olism where the log of the rate of oxygen con-
sumption (cc./hr.) equals log a b log W,
or metabolic rate equals the constants a
and b being the regression coefficients of the
intercept and slope and W the wet body weights
(gms.).. Because of the slight depression for the
values at 14° C. the data for Bufo boreas, al-
though plotted, were not included in computing
the regression lines for Text-figure 2.
At both temperature levels the respiratory
rates for the tropical frogs averaged lower than
those from the northern latitudes at similar tem-
peratures. This appears to be especially evident
when we compare the smaller animals, and is
reflected in a steepening of the slope for the
tropical weight regression coefficients. Although
the average regression coefficient of .71 for the
temperate animals at the two testing tempera-
tures is but slightly higher than an exponent of
% (.67) that we might expect in keeping with
the surface law of metabolism (metabolism/
time = the average slope of .84 for the
tropical species seems significantly greater.
There appears to be little correlation between
temperature and regression slope in animals
from the same general latitude.
With the exception of Bufo boreas, the tem-
perature coefficients (Qio) did not vary signifi-
cantly between the tropical and temperate
1957]
Tashian & Ray: Oxygen Consumption and Temperature in Anurans
65
Text-fig. 2. The relation of oxygen consumption to weight in four species of temperate and boreal anurans
at two temperature levels (b = regression coefficient; r = coefficient of correlation). Dashed lines rep-
resent regression slopes for tropical species. Data for B. boreas were not used in plotting the regression lines.
species (Table 1). The oxygen consumption
rates for Bufo boreas at both temperature levels
showed little variation both within and between
the Alaskan and Californian forms. In fact, at
both 14° and 24° C. the two races of Bufo
boreas showed a metabolic rate typical of the
temperate frogs tested at 24° C. This relative
insensitivity to temperature change is reflected
in their low temperature coefficients.
Discussion
The weight regression coefficients (.703 and
.707) for the temperate animals studied here
are supported by the findings of others. Davison
( 1955) found a value of approximately % (.67)
for the exponent in seven species of temperate
anurans (weight range 1 to 350 gms.) that he
measured at 25° C., and Rubner (1924) reports
an exponent of .67 for Rana esculenta. Regres-
sion coefficients of .80 and .85 are given by
Scholander et al. (1953) in the tropical and
arctic animals (fishes, crustaceans, insects and
spiders) that they measured at 0° C. (arctic
species) and 20-30° C. (tropical species). No
shift in the weight regression slope is apparent
between the arctic and tropical forms. Because
of the limited number of species tested by us in
the present study, the steeper regression slopes
(.825 and .861) found in the tropical species
can not be considered as conclusive.
The compensatory metabolic rates exhibited
by the northern or cold-adapted anurans studied
here are in keeping with the general concept
(Bullock, 1955) that activity rates are greater at
a given temperature in cold-blooded animals
from northern latitudes when compared with
the same or closely related southern forms.
Scholander et al. (1953) state that climatic
adaptation in terrestrial poikilotherms is not
Table 1. Summary of Data on Respiratory Metabolism for Adult Tropical, Temperate and Boreal Anurans at Two Temperature Levels.
66
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Tashian & Ray: Oxygen Consumption and Temperature in Amirans
67
significantly reflected by over-all oxygen con-
sumption in the tropical and arctic insects and
spiders that they tested at 0° and 20° C. Prosser
(1955) also concludes that metabolic differences
in populations are to be found in aquatic but
not in terrestrial poikilotherms. If we consider
the amphibians studied here as semi-terrestrial,
we should then expect to find a significant, but
not pronounced, difference in climatic adapta-
tion between the tropical and northern forms.
This appears to be the case. It would seem that
a similar study of tropical and temperate rep-
tiles would be of particular interest in this
respect.
It is also generally accepted (Bullock, 1955)
that cold-adapted poikilotherms tend to show a
lower Qio when compared with the same or
closely related warm-adapted species. This con-
cept becomes apparent among the anurans of
this study only in the toad Bufo boreas. The
other frogs show no significant variation in Qm
between tropical and temperate forms (Table
1). This seemingly inherent low Qio for Bufo
boreas could have been a contributing factor
in its successful northern extension into south-
ern Alaska. It is interesting to note, however,
that in Rana sylvatica, which ranges even fur-
ther north than Bufo boreas, we find no in-
herently lower temperature coefficient. These
results can probably be partially explained by
the limited number of determinations made as
well as by the fact that we are here grossly com-
paring different species. Moreover, familial dif-
ferences such as the more terrestrial habits of
Bufo boreas could conceivably explain the dis-
similarities. Breeding condition, size and sea-
sonal fluctuations may also have had an
influence. However, with the exception of Hyla
crucifer, all northern animals were collected
between June 1 and July 4, and only adults of
similar weight within a species were used in the
determination of the temperature coefficients.
Bufo boreas boreas was the only form in breed-
ing condition at the time of collecting. It is sur-
prising that individuals of Bufo boreas exhibited
such a persistently similar metabolic picture,
coming as they did from two places so widely
separated in latitude and altitude (sea level to
7,500 ft.).
There is good evidence in support of the con-
clusion (for references see Dehnel, 1955) that
at their normal environmental temperatures, ac-
tivity rates of closely related northern and
southern animals are usually similar. The annual
range of mean monthly temperatures for Juneau
vary from -1.6° C to 13.0° C (1921-1950) with
an annual mean of about 5.8° C. compared with
a range of 25.6° to 27.3° C. and an annual mean
of 26.5° C. for Trinidad (Port-of -Spain) . If we
take the mean monthly temperature for June at
Juneau (12.0° C.) and the average January to
March monthly temperature for Simla (ca.
24° C.), we can then use the lower testing tem-
perature (14° C.) for Bufo boreas boreas and
the upper testing temperature (25° C.) for the
tropical species as being reasonably close to their
normal mean habitat temperature at the time of
collecting. In examining the two extreme lati-
tudinal forms of similar size (Table 1), we find
Bufo boreas boreas with a mean oxygen coeffi-
cient (QO2) of .152 at 14° C., which is higher
than that of all but one of the tropical frogs
tested at 25° C. At their normal habitat tempera-
tures, then, the northern species showed the
more active metabolism. Other studies (Scho-
lander et ah, 1953, and Tashian, 1956) showed
a higher QO2 for the tropical forms at their
normal environmental temperatures.
Summary
1 . The oxygen consumption of five species of
tropical frogs from Trinidad, B.W.I., was de-
termined and compared with that of four species
of temperate and boreal anurans from Vermont,
New York, California and Alaska at two tem-
perature levels.
2. The respiratory metabolism for the north-
ern forms averaged higher than that of the
tropical animals at both temperatures. This
higher metabolism for the cold-adapted animals
is more evident in the smaller species, and is
reflected in a steepening of the weight regression
slope for the tropical species at both tempera-
ture levels.
3. With the exception of a lower Qio for both
races of Bufo boreas when compared with any
of the other forms tested, no significant variation
could be detected in the Qio of northern and
southern forms.
4. At their normal habitat temperatures, the
northern species had a higher rate of respiration
than the tropical species.
Literature Cited
Bullock, T. H.
1955. Compensation for temperature in the
metabolism and activity of poikilotherms.
Biol. Revs., 30: 311-342.
Davison, J.
1955. Body weight, cell surface, and metabolic
rate in anuran amphibia. Biol. Bull.,
109: 407-419.
Dehnel, P. A.
1955. Rates of growth of gastropods as a func-
tion of latitude. Physiol. Zool., 28: 115-
144.
68
Zoologica: New York Zoological Society
[42: 5: 1957]
Flemister, L. J. & S. C. Flemister
1951. Chloride ion regulation and oxygen con-
sumption in the crab Ocypode albicans
(Bose). Biol. Bull., 101: 259-273.
Moore, J. A.
1949. Geographic variation of adaptive char-
acters in Rana pipiens Schreber. Evolu-
tion, 3 : 1-24.
Prosser, C. L.
1955. Physiological variation in animals. Biol.
Revs., 30: 229-262.
Rubner, M.
1924. Aus den Leben des Kaltbluters, 11 Teil,
Amphlbien und Reptilien. Bioch. Zeitschr.,
148: 268-307.
SCHOLANDER, P. F., W. FLAGG, V. WALTERS & L.
Irving
1953. Climatic adaptation in arctic and tropical
poikllotherms. Physiol. Zool., 26: 67-92.
Tashian, R. E.
1956. Geographic variation in the respiratory
metabolism and temperature coefficient
in tropical and temperate forms of the
fiddler crab, Uca pugnax. Zoologica,
41: 39-47.
VOLPE, E. P.
1954. Hybrid inviability between Rana pipiens
from Wisconsin and Mexico. Tulane stud.
Zool., 1: 111.
6
Basic Patterns of Display in Fiddler Crabs
(Ocypodidae, Genus Uca)
Jocelyn Crane
Department of Tropical Research, New York Zoological Society,
New York 60, N. Y.
(Plate I; Text-figures 1-4)
Contents
I. Introduction 69
II. General Form of Display in
Fiddler Crabs 70
III. Basic Wave Patterns 70
IV. Advanced Stages of Display 73
V. Ecological Differences in Distribution
of Display Types 74
VI. Geographical Distribution in Relation
to Display 76
VII. Evolutionary Trends 76
VIII. Summary 81
IX. Referenees 81
I. Introduction
This contribution is a preliminary result of
a general study of ocypodid crabs. The
elaborate and varied displays characteris-
tic of the fiddler crabs (genus Uca) appear to
contribute substantially to an understanding of
the phytogeny of the group. An interim report is
presented because the increasing use of fiddler
crabs in biological research makes their relation-
ships of timely interest.
Specific differences in display behavior were
described in an earlier study (Crane, 1941)
where it was found that in a group of sympatric
species of Uca on the west coast of Central
America every species could be distinguished
from every other species by the form of the dis-
play. Even species which could not be otherwise
distinguished without a lens were easily identi-
fied at a distance by the characteristic form and
tempo of the “waving” of the large cheliped. It
1 This study has been supported by a grant from the
National Science Foundation.
2 Contribution No. 975, Department of Tropical Re-
search, New York Zoological Society.
was found further that closely related species
had similar types of display which were divis-
ible into a number of groups.
During subsequent years other American spe-
cies of Uca have been studied in the field on
both sides of the continent, from California to
Guayaquil, Ecuador, and from Massachusetts
to Rio de Janeiro, Brazil (Crane, 1943, 1944 and
unpublished).
Now, through a grant from the National Sci-
ence Foundation, the work has been expanded
into a world-wide program. In 1955 and 1956
the crabs were studied in Pakistan, Ceylon, Ma-
laya, Singapore, Sarawak, the Philippines, north
and northeast Australia, Fiji, Samoa, Tahiti and
adjacent islands. Early in 1957 Panama was
revisited, and previously known species reviewed
in the light of the recent studies of Indo-Pacific
forms. The behavior of the crabs, particularly
of displaying individuals, was recorded in color
on more than 9,000 feet of 16 mm. motion pic-
ture film. This material, along with the correlated
and supplementary field notes, forms a perma-
nent record for comparative laboratory analysis.
The displays of about 19 good Indo-Pacific spe-
cies of Uca have been recorded in this manner
and 29 American forms. Although extensive field
work remains to be done, particularly in Africa,
the principal outlines of the development of dis-
play in the group appear now to be clear.
All detailed supporting data, discussion on the
functions and releasing mechanisms of display,
specific correlation of display with morphology
and discussion of display in relation to infra-
specific categories and speciation are reserved
for a monographic report on the genus, now in
preparation. The following subjects more or less
closely associated with display will also be treated
only in the later publication : color change, sound
69
70
Zoologica: New York Zoological Society
[42; 6
production, territoriality, physical combat be-
tween males and the erection of structures near
the burrows.
Among the many people who have facilitated
the field work I wish at this time to express my
appreciation particularly to Dr. William Beebe,
Director Emeritus of the Department of Tropical
Research, New York Zoological Society, and to
Dr. Waldo L. Schmitt, Chief Curator of Zoology
at the United States National Museum. Both
were instrumental in inaugurating the study and,
through the years, have given most helpful ad-
vice and encouragement.
II. General Form of Display in
Fiddler Crabs
The outstanding characteristic of displaying
fiddler crabs is a rhythmic elevation and lower-
ing of the hypertrophied major cheliped of the
male. In the few species usually familiar to
western observers in the field this takes the form
of “waving” or “beckoning,” and is character-
istic of the breeding season. The significance of
the gesture has been described by various work-
ers as non-sexually territorial, sexually territorial,
a sex attractant, a challenge to other males and
as various combinations of all of these possi-
bilities. Although it is not proposed at this time
to contribute further to the discussion, it may
be said that field observations and motion pic-
ture records prove conclusively two points., First,
all the suggested functions occur, sometimes all
in a single species. Furthermore, in some ad-
vanced species there are distinct differences
separating territorial, male-to-male and male-
to-female displays, the latter being the most
highly developed.
The motion of the cheliped is by no means
always wavelike; in some species it is the merest
slight raising of the flexed manus and chelae in
front of the buccal region. In others it is a vio-
lent, very rapid shaking of the same parts at the
level of the eye-stalks; in others the base of the
manus is rapped against the ground, the waving
is in jerks, or the cheliped is held high and re-
volved in circles. Among these extremes there
are all degrees. Tempo is exceedingly various
and specific, although unrelated crabs often wave
at about the same rate of speed. Single waves
range from one wave lasting 13 seconds to five
waves being crowded into a single second. The
first extreme of timing is found in certain South
American populations of U. pugnax rapax under
certain conditions and the second in an appar-
ently undescribed species from Port Darwin,
Australia. The waving rate of most species falls
between one-half and two seconds per wave.
Motions associated with waving include ele-
vation of the carapace on the ambulatories, re-
volving, moving from side to side, and various
“bounces” and “curtsies” accomplished by rapid-
ly lowering and raising the body on one or both
sides.
In the following pages the display motion of
the large cheliped will for convenience be termed
“wave,” whether or not it bears an anthropo-
morphic resemblance to such a gesture.
III. Basic Wave Patterns
Most of the species of Uca may be clearly
divided into two groups depending on a basic
difference in the direction of the first part of
the motion of the major cheliped. These two
behavioral complexes are characteristic of the
majority of species in, respectively, the “narrow-
fronted” and “broad-fronted” groups, as they
have been termed in most systematic treatments.^
3 Bolt, 1954, basing his decision on a collection of
specialized Central American species of Uca, gave the
narrow-fronted and broad-fronted groups each generic
standing, reserving Uca for the narrow-fronts and pro-
posing Mimica for the broad-fronts. Peters (1955),
working on ecological and behavioral aspects of the
same material, reduced Minuca to subgeneric rank. The
present writer currently feels that Minuca should be
given at most subgeneric status. This view is dependent
on the extensive series of intermediate species distributed
in other parts of the world, and on the distinctness and
homogeneity of Uca, sensu lata, in comparison with all
other genera in the family. Until further comparative
morphological work is completed, it seems preferable
to use simply the non-technical terms, “narrow-fronts”
and “broad-fronts,” as a practical division which for
the majority of species appears to be phylogenetically
justified.
Vertical Waves. (Text-figs. 1; 4, A-B). In the
great majority of narrow-fronts, the cheliped at
the beginning of the wave remains flexed in front
of the buccal region and is raised up and slightly
forward from there, without unflexing, until it
reaches the level of the eyeballs. In the simplest
displays it reaches no farther and therefore is
never unflexed. In more advanced displays its
elevation continues obliquely above the eyes,
through the unflexing of the manus and chelae.
Regardless of the amount of elevation or degree
of unflexing, the cheliped is lowered back into
rest position in the same plane in which it was
elevated. This type of display will be called a
“vertical wave.”
Lateral Waves. (Text-figs. 3; 4, D-F). By con-
trast, the characteristic wave form of almost all
species of broad-fronted Uca commences with
a sweep to the side, rather than with a vertical
elevation of the cheliped; this kind of gesture
will be called a “lateral wave.” In moderately
intensive display, in which differences among
species are most apparent, the flexed cheliped
is pushed away from the body at the beginning
1957]
Crane: Basic Patterns of Display in Fiddler Crabs
71
of the wave and more or less unflexed toward
the side; it then sweeps up to a completely un-
flexed, high-reaching position. Finally it is flexed
once more in front of the buccal region. Since
it is often lowered in a direct path, close to the
body, a more or less circular motion is com-
pleted. The amount of deviation from the path
of the first part of the wave differs among species,
and within species depending on the degree of
display intensity; therefore the circularity varies
widely; sometimes, in fact, lateral displays are
altogether single plane. In displays of the lowest
intensity of all, even laterality is often absent,
the cheliped motion closely resembling the slight,
flexed, single-plane display of the vertically wav-
ing narrow-fronts, as described in the preceding
paragraph (Text-fig. 4 C) .
Intermediate Waves. The waving pattern of a
few species is intermediate between vertical and
lateral types. Morphologically these forms also
show some characters intermediate between
those of narrow-fronts and broad-fronts.
Position of Body during Waving. In addition
to the form of the wave there are several funda-
mental differences in the position of the body
during waving. In some species it is scarcely
raised at all; in many it is raised on the ambula-
tories and lowered with every wave; in still others
it is raised and held erect during a series of
waves. The amount of elevation varies some-
what within species in accordance with the in-
tensity of waving.
Systematic Distribution of Basic Wave Pat-
terns. An arrangement of species characterized
by various types of wave is presented in Table 2.
Authorities for the species, geographical distri-
butions and localities where display has been
observed are listed in Table I.
As will be seen from Table 2, the vertical type
of wave is typical of Indo-Pacific narrow-fronts.
These species in turn include those forms in which
Uca display is simplest and, it seems unquestion-
ably, most primitive. These uncomplicated dis-
plays are found in manii, rhizophorae (Text-fig.
1, A-B), rosea, ischnodactyla and rathbunae; in
these species the body is scarcely if at all raised
above the ground during display. U. dussumieri
is somewhat further advanced. Although specific
differences in the waving pattern of all of these
are clear-cut, the group similarity is obvious to
the observer. In all of these species the cheliped
is not raised high overhead, and the amount of
time devoted to waving is negligible in compari-
son with that so spent in the more advanced
species.
The waving of still other narrow-fronts from
the Indo-Pacific is considerably more advanced,
as in marionis, signata (Text-fig. 1, C-D) and
zamboangana (Text-fig. 1, E-F). Although the
pattern is still vertical and single plane rather
than lateral in character, the reach is higher,
the tempo tends to be swifter, the proportion of
time devoted to waving greater and the eleva-
tion of the body conspicuous.
Neotropical members of the narrow-fronted
group are aberrant as well as intermediate, both
morphologically and behavioristically. The dis-
play is characterized by a lateral, not vertical,
type of wave. Except in low intensity display
the gesture is spiral, rather than merely circular,
since the cheliped, after its initial lateral eleva-
tion, is revolved throughout a series of waves
without being lowered to the ground. This late-
ral, spirally circular display has been observed
in heteropleura, heterochelos, stylifera, princeps,
maracoani and insignis (Text-fig. 2). It reaches
its maximum development in the two latter
species, which are allopatric counterparts
in the Atlantic and Pacific. In both of them,
which attain large size and a correlated remark-
able development of the heterogonic major cheli-
ped, the chelae are directed upward, spread wide
apart and then rotated without pause, often for
several minutes at a time. Even in their low in-
tensity displays these two species are of the
lateral type; in the others, however, low intensity
waving shows single-plane, vertical-wave affini-
ties. Most of the group hold the carapace high
during a series of waves, although in stylifera
and princeps the hind legs bend during the high-
est reach of the cheliped, tilting the posterior
part of the carapace downward. U. heteropleura,
heterochelos and young princeps raise and lower
the carapace with each wave.
Two narrow-fronted crabs, the Australian
longidigitum and the Indo-Pacific tetragonon,
as well as the Indo-Pacific broad-fronted gai-
mardi, all show wave characteristics intermediate
between the vertical and lateral types. The dis-
plays are very distinct, however, and the three
species are not morphologically closely related
to one another.
Alone among the known displays of broad-
fronted crabs the neotropical Atlantic thayeri
has a vertical, single-plane type of wave similar
to those of the Indo-Pacific narrow-fronts. Mor-
phologically, also, thayeri shows affinities with
those species.
The remaining broad-fronts, in which display
has been observed, are all characterized by a
strongly lateral type of wave, usually with a
variable degree of circularity (Text-fig. 3).
In a number of the very rapidly waving neo-
tropical broad-fronts, such as beebei and salti-
tanta, the last half of the wave is so swift that
the eye does not record it. Hence although the
motion was often described in field notes as a
single-plane wave, subsequent examination of
72
Zoologica: New York Zoological Society
[42: 6
Text-fig. 1. Examples of vertical types of wave in three species of narrow-fronted Uca from the Indo-
Pacific region. Illustrations on left show rest positions between waves, those on right the maximum eleva-
tion of the cheliped, which is raised and lowered in a single plane. Note in the series, reading from the
top down, the progressively higher reach of the cheliped and greater elevation of the carapace. See text,
p. 70 ff.. Table 2 and Text-fig. 4, A-B.
A, B, U. rhizophorae (photographed in Singapore); C, D, signata (Philippine Is.); E, F, zamboan-
gana (Philippine Is.). Drawings by Dorothy F. Warren, after motion picture frames and mounted specimens.
motion picture frames proved that circularity
was clearly evident.
Examples of a strongly circular type of lateral
wave include the Indo-Pacific annulipes and lac-
tea (Text-fig. 3, A-D) and the eastern Pacific
latimanus.
In a few species, such as U. terpsichores, the
rest position of the cheliped during display is
with the chelae directed forward.
An aberrant group of broad-fronts, of which
pugnax rapax (Text-fig. 3, E-F) is typical, is an
important element in neotropical Uca; two At-
lantic representatives extend even into the north
temperate region {pugnax, minax). They are all
characterized by exceedingly broad fronts and
by a jerking, obliquely-lateral wave which dur-
ing moderate intensity is circular. They have no
Indo-Pacific representatives.
The lateral wave of the broad-fronts, in the
progressively specialized species, shows increas-
ing speed, maximum unflexing of the cheliped
and more prolonged periods of time devoted
to waving.
It has already been noted that in species in
which waving is poorly developed {rhizophorae,
manii, etc.) the body is raised scarcely or not
at all during waving. In the majority of species,
among both narrow-fronts and less advanced
broad-fronts, the carapace is raised on the am-
bulatories and lowered with every wave. In some
1957]
Crane: Basic Patterns of Display in Fiddler Crabs
73
lateral wavers, the body is only slightly elevated,
but is held in this position throughout the series.
(In a few of this group, for example in beebei,
an impression of raising-and-lowering is given
by the tendency of the crab to bend the hind
legs during the peak of the cheliped reach, thus
tilting the carapace down posteriorly; this is es-
pecially true when the crab is displaying on
soft mud; c.f. Peters, 1955, who reported that
this form raises and lowers the carapace with
every wave). In advanced species among both
narrow-fronts and broad-fronts, the body is held
high on the ambulatories throughout a series of
waves.
IV. Advanced Stages of Display
One behavioral distinction between the nar-
row-fronts and the broad-fronts is found in all
the species so far observed except for a few
instances. This concerns the final stages of pre-
mating behavior.
Advanced Display in Narrow-fronts. In the
narrow-fronts the male pursues a female or ap-
proaches her at the mouth of her burrow; this
behavior does not necessarily immediately follow
display, although the male always has been dis-
playing during the same low-tide period. The
approach, however, often does follow display
directed more or less toward a female, although
this individual is not necessarily the one which
is eventually approached. There is no marked
increase in tempo of waving during the approach,
and additional elements of display are lacking,
except for a few special steps in the aberrant
neotropical stylifera, insignis and maracoani.
When within reach, the male seizes the female
and, after brief tapping or stroking of her cara-
pace with his ambulatories, attempts copulation
at or near the mouth of her burrow. This has
been observed by Altevogt (1955) in marionis
in India; and by the present author as follows:
marionis in Fiji, the Philippines, Australia and
Singapore; dussumieri in the Philippines, Sara-
wak and Singapore; manii in Singapore and
Penang; rathbunae in the Philippines; two unde-
scribed new species in Australia; heteropleura,
stylifera and insignis in Ecuador and Pacific
Panama; and maracoani in Trinidad, northern
South America and Brazil. Actual copulation
was seen in marionis (four times), rhizophorae
(twice), manii (twice), dussumieri (once),
Australian new species (once) and stylifera
(once). The last stage of courtship, involving
stroking, has been observed probably 75 times
at a conservative estimate, although these latter
episodes did not, after display, end in copulation.
A few instances of apparently atypical be-
havior is known in the narrow-fronts. Females
of stylifera (Crane, 1941, p. 172) insignis and
Text-fig. 2. Lateral circular type of wave in the
neotropical crab, Uca insignis (photographed in
Panama). Carapace is held high on the ambulatories
throughout a series of high-reaching circular waves,
during which the cheliped is never brought down
into the flexed position of rest. A, maximum reach
of cheliped; B, “low” position, between waves. See
text, p. 71, and Table 2. Drawings by Dorothy F.
Warren after motion picture frames and mounted
specimens.
signata were seen on one occasion each to follow
a displaying male into his burrow. Two small
princeps {ibid, p. 170) seized females. Each
tried unsuccessfully to drag the female down
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[42: 6
his own burrow, grasping her with the ambula-
tories of the minor side.
During all of the above instances where I have
observed final stages of display and copulation
above ground I have never once seen the male
seize and hold the female with the major cheli-
ped. Therefore the definite holding observed by
Altevogt near Bombay in copulating marionis
(1955.2, p. 518) apparently is of rather excep-
tional occurrence.
Advanced Display in Broad-fronts. In the
broad-fronted group, in addition to an increased
tempo of display at the approach of a female,
there are specialized steps, curtsies or rappings
which are only elicited at this time.
Another group distinction is apparent at the
end of courtship. In the broad-fronts the usual
procedure is for the male, after high intensity
display, to precede the female down his own
burrow; the female may or may not follow, and
may or may not stay below, presumably copu-
lating, for any length of time. I have seen the
female actually follow the male into the burrow
in the Indo-Pacific annulipes in Karachi, Singa-
pore, Sarawak and Davao; in lactea in Pakistan,
Singapore, the Philippines, and Fiji; in the Paci-
fic neotropics in stenodactyla, beebei, batuenta,
saltitanta and latimanus; and, finally, in the
western Atlantic in pugnax, pugnax rapax, pugi-
lator, cumulanta, leptodactyla and in two appar-
ently undescribed new species.
In only two broad-fronts, stenodactyla and
beebei in Panama, was copulation seen; this was
at the surface by the female’s burrow as in nar-
row-fronts (Crane, 1941, pp. 193, 197). A num-
ber of females in each of these two species,
however, have been seen to follow the male
in the usual broad-front fashion. Throughout
the broad-fronted group, therefore, underground
mating in the male’s burrow is presumably the
rule and the female, instead of being pursued
toward her own burrow by the male and often
seized more or less forcibly, is attracted by rapid
and characteristic display into the burrow of the
male which she enters after he has disappeared.
In the broad-fronted gaimardi (= pulchella
Stimpson) in Tahiti both the first and the last
stages of display appear almost perfectly inter-
mediate betwen those typical of the primitive
narrow-fronts and of the broad-fronts. The in-
termediate character of the wave has already
been mentioned; advanced stages of display were
lacking and the male pursued the female as in
narrow-fronted crabs. Even during high intensity
display the carapace was scarcely elevated. Mor-
phologically, as usual with crabs having inter-
mediate displays, the species shows intermediate
characters.
Another broad-fronted species of particular
interest is saltitanta, from the Pacific neotropics.
As noted previously (1941, p. 166) this form
represents the extreme in development of the
rapping type of display, which occurs in a num-
ber of related species on both coasts of tropical
America. In these species a crab, at the end of
a wave, may bounce the lower edge of the major
manus and pollex on the ground (Text-fig. 4 F).
Usually this bouncing or rapping occurs only
when the crab is excited, often in the final stages
of courtship. It is typical immediately before
the male’s descent into his own burrow, after
display has been directed toward a particular
female. In saltitanta the rapping has been pushed
forward to become an integral part of routine
waving, whether or not the crab is particularly
excited and whether or not display has been
directed toward a female.
The basic characteristics of the lateral wave
itself may be superceded during the specialized
motions of advanced display. Depending on the
species, the cheliped may be held motionless
overhead, or in front, whUe the crab bobs and
races to and fro. Again, the cheliped, having
passed through stages of low-intensity single-
plane and high-intensity circular display, reverts
once more to a single-plane return; now, how-
ever, the cheliped is opened laterally and raised
to maximum height while the waving tempo is
greatly increased. In a number of species the
cheliped is not returned to its usual rest position
between waves, but performs an aerial circle,
somewhat similar to that found in the aberrant
neotropical narrow-fronts.
In Table 2 only a slight indication is given
of the types of advanced display occurring
among the broad-fronts. In spite of group simi-
larities, the display of each species is so distinct
that, if seen in moderate intensity and advanced
stages, none could ever be confused in the field
with that of any other species, even from other
parts of the world.
As in studies of the complex displays of cer-
tain vertebrates, it is obviously essential, both
for accuracy of description and for an approach
to understanding, that the displays be observed
for prolonged periods and under a variety of
conditions.
V. Ecological Differences in the
Distribution of Display Types
In 1941 (p. 160) the present investigator noted
that the most highly developed displays in west-
ern Central American crabs were found in spe-
cies living on the shores of bays and estuaries
which, while protected from waves, were freely
confluent with the open sea and consequently of
1957]
Crane: Basic Patterns of Display in Fiddler Crabs
75
Text-fig. 3. Examples of lateral types of wave in broad-fronted Uca. A-D, U. lactea (photographed
in Fiji Is.), showing maximum development of the lateral circular wave, in which the cheliped starting
from the flexed position (A) is unflexed outward (B), then raised (C), and Anally returned (D) to the start-
ing point. This wave is best developed in displays of moderate intensity; at low intensity or at high inten-
sity during advanced display, the wave may be of a vertical or lateral single plane type. See text, p. 71,
Table 2 and Text-fig. 4, C-E. E, F, U. piignax rapax (Venezuela). Rest position and maximum cheliped
reach of lateral circular wave, characteristic of moderate intensity display. Cheliped is unflexed outward,
raised and lowered in a series of jerks. See text, p. 72, and Table 2. Drawings by Dorothy F. Warren,
after motion picture frames and mounted specimens.
relatively high salinity. Recent observations in
the Indo-Pacific have shown agreement. In both
the narrow-fronted and broad-fronted groups
the most actively displaying species, which ap-
parently are also most morphologically ad-
vanced, are those in similar localities. Examples
of Indo-Pacific crabs with well developed dis-
plays are the narrow-fronts tnarionis and tetra-
gonon and the broad-fronts annulipes and lactea;
all live typically on rather open shores or inlets
of protected bays, as do their neotropical coun-
terparts including insignis and maracoani among
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[42: 6
the narrow-fronts and stenodactyla, deichmanni,
terpsichores, latimanus and leptodactyla among
the broad-fronts. Those with the least developed
displays, the Indo-Pacific rhizophorae, manii,
etc. (p. 71), are also those morphologically
the least specialized for an amphibious life; all
inhabit less saline situations, sometimes living in
almost fresh water, on more sheltered mudflats
and streambanks.
An interesting point is that the extremely
broad-fronted American group, including pug-
nax and mordax, has spread into sheltered locali-
ties extending even, in mordax, far up tropical
rivers. The tempo of their displays is slow, as
in sheltered, unrelated species in the Indo-Paciflc,
but it is clearly of the lateral, highly developed
broad-fronted type, and the females, as usual
in the broad-fronts, are attracted down the bur-
rows of the males.
VI. Geographical Distribution in Relation
TO Display
Those narrow-fronted species with the sim-
plest and apparently most primitive display
{manii, etc.) are all confined to the Indo-Malay-
an region, which undoubtedly is the center of
distribution for the family. The intermediate
narrow-front, tetragonon, which currently
reaches Tahiti and was formerly found also in
Hawaii, shows in its display as in its morphology
certain intermediate elements which seem to
form a link with the American narrow-fronts.
The displays of the latter are more highly evolved
than any yet known in the group in the Indo-
Pacific, and show affinities to those of the broad-
fronts; the species are also specialized morpho-
logically, being characterized for example by
extreme narrowness of the front, by unique
peculiarities of the spoon-tipped hairs of the
second maxilliped, and by male abdominal ap-
pendages more similar to those of broad-fronts
than of Indo-Pacific narrow-fronts.
The broad-fronted displays are represented
by both simple and advanced examples in both
hemispheres. There are, however, many more
American than Indo-Pacific species, and most
of the American species are more specialized in
both behavior and morphology than any of the
observed Indo-Pacific broad-fronts with the ex-
ception of lactea.
U. gaimardi, the only Indo-Pacific broad-front
reaching as far east as Tahiti, appears to be,
as is tetragonon among the narrow-fronts, an
intermediary between Indo-Pacific narrow-fronts
and American broad-fronts.
Three Indo-Pacific species have been observed
in display over a horizontal distribution of 5,000
to 8,000 miles. These are marionis, annulipes
Text-fig. 4. Ground projection of path of various
types of wave in Uca. In each path the beginning is
at upper right boundary of the black line. In A, B
and C, the length of the line represents the extent
of the obliquely forward thrust of the cheliped dur-
ing its elevation. A, U. rhizophorae, a low vertical
wave, single plane; B, z^mboangana, a high vertical
wave, single plane; C, lactea, a vertical, single plane
wave occurring in low intensity display; D, same, a
lateral circular wave, typical of display of moderate
intensity; E, same, a lateral, single-plane wave found
during advanced stages of display; F, saltitanta, a
lateral circular wave, with straight line marking route
as cheliped retraces beginning of path, rapping
ground in front of crab as the claw bounces back
into rest position. Cf. text, p. 70, and Text-figs. 1
and 3.
These projections were obtained by placing a pre-
served, fiexible specimen of each species on a thick
sheet of modelling clay and fastening a needle to
the tips of the major chelae with scotch tape in such
a way that, regardless of the elevation of the cheli-
ped, the needle cut partially through the modelling
clay. Starting from the flexed, rest position the
cheliped was then moved as though in natural dis-
play, in accordance with motion picture records, the
tip of the needle marking the projected path.
and lactea. No gross variations in display be-
havior have been observed among the various
populations within each species, although de-
tailed analyses of motion picture film are ex-
pected to show minor distinctions. The one major
difference, noted among the populations of lac-
tea, was that no “shelters” (Crane, 1941, p. 157)
were built by the species even at the height of
display in Singapore, the Philippines or Fiji.
They were, however, built by about one-fourth
of the adult male lactea in Karachi, Pakistan, in
mid- June.
VII. Evolutionary Trends
The varying patterns of Uca display, from the
simplest to the most complex, show a trend prev-
alent in many branches of the animal kingdom
in groups where vision is well developed. In Uca,
as in numbers of birds, reptiles, fishes, spiders
and mantids, for example, it is axiomatic that
the display increases the conspicuousness of the
animal— regardless of the function of the par-
ticular display under consideration.
1957]
Crane: Basic Patterns of Display in Fiddler Crabs
11
A primary component of the conspicuousness
is usually a temporary or apparent increase in
size. In fiddler crabs this increase is attained in
three ways; first by a progressively higher reach
with the cheliped; second by the broadening of
the wave into an expansive lateral gesture re-
placing a close-to-the-body motion; and, third,
by the higher and more prolonged elevation of
the body on the increasingly straightened walk-
ing legs. In some advanced laterally-waving
species, however, the body is not held very high,
the walking legs being braced far out at the
sides; this position seems to be an aid in balanc-
ing. It is probably a necessary stance in these
species in which both heterogony and the lateral
extension of the cheliped are extreme.
There are other display characteristics which
contribute to the conspicuousness of the crab
and which are attained most fully in those species
in which waving is highly developed. These char-
acteristics include faster tempo of the wave, a
rhythm accented by jerks or other specializations
and an increase in time devoted daily to waving.
The trend toward the attraction of females
to the male burrow, typical of the broad-fronts,
may well have adaptive value, since pairs copu-
lating at the surface are fully exposed to preda-
tion by birds. A probable factor in the evolution
of this pattern is the fact that among broad-
fronts the male is often considerably larger than
the female. Because of this size discrepancy males
cannot easily descend into the narrow burrows
of the females.
Many species with highly developed displays
resemble primitive forms when waving at low
intensity. This is especially well shown when a
crab is feeding and waving at the same time,
when tidal or meteorological conditions are un-
favorable, or when the crab is not ph3'siologically
in full display condition. Under any of these
circumstances even species as far apart phylo-
genetically as maracoani,stenodactyla and lactea
may raise the cheliped only slightly and lower
it in the same, nearly vertical plane, scarcely
elevating the body. This low intensity display
therefore is very similar to the full display of
rhizophorae, manii and their relations.
There appear to be no structural reasons pre-
venting vertical wavers from displaying in the
lateral fashion. The form of the condyles, the
proximal concavities of articulating segments
and the elasticity of the connections are similar
throughout, although Peters (1955, pp. 489 ff.)
illustrates minor differences in the degrees of
possible motion in two closely related species.
Throughout the genus the cheliped is often wide-
ly extended, regardless of the type of wave per-
formed during display, in combat between males
when the chelae of the combatants are actually
engaged.
Altevogt (1955.2, p. 514) recorded differences
in the display of very large and moderate-sized
individuals of marionis, the giants having a less
out-reaching form. He thinks this is probably
associated with the altered center of gravity. It
seems to the present investigator that the evolu-
tion of the form of display in the gigantic species
of neotropical narrow-fronts, maracoani and
insignis, may have been guided by similar prob-
lems of weight and balance.
Evidence has recently been found that dis-
placement behavior sometimes occurs during
Uca display, and it seems likely that such
behavior varies among the species roughly in
accordance with their phylogenetic position.
Gordon (1955) first reported displacement feed-
ing (in the sense in which the term has been used
by Armstrong, 1950, and Tinbergen, 1952) in a
number of African species. I fully agree with
her interpretation, having often observed similar
types of activity not only in Uca but in other
genera of displaying ocypodids, notably Ilyoplax.
Gordon’s observations were of feeding, often
in a token, non-functional fashion, usually after
fighting or after high intensity display and a
thwarting of mating behavior.
The current studies present evidence of two
more aspects of the subject. First, displacement
behavior is most prevalent in species character-
ized by highly developed display. Second, in
these species there occurs not only displacement
feeding, as described by Gordon, but also dis-
placement claw-cleaning. The large manus and
chelae are buffed and polished by the small, or
at least the requisite motion is made, although
the terminal brush may not actually touch the
large chelae. The gesture is repeated frequently
even though the crab has been above ground
for many minutes and the cheliped is altogether
free of mud. This activity seems to occur espe-
cially when there is conflict between the urge
to display and that to escape, such as when the
camera or observer is very close to the crab.
Current observations indicate furthermore
that there will prove to be specific differences in
the tendency toward one type or another of
displacement behavior. For example, festae in
Ecuador, deichmanni in Panama and cumulanta
in Trinidad and Venezuela all appear to be
especially prone to displacement claw-cleaning.
U. lactea in the Indo-Pacific and leptodactyla in
Brazil, on the other hand, seem to have a stronger
tendency to displacement feeding. In contrast,
in the primitive Indo-Pacific narrow-fronts {rhi-
zophorae et al.) no displacement behavior has
been noticed; it appears likely that in these
78
Zoologica: New York Zoological Society
[42: 6
Table 1. Species of Uca in which Display Has Been Studied During the Present Investigation
Note: The majority of these forms are undoubtedly good species. It is expected that some will be
reduced to subspecific rank, especially certain narrow-fronted crabs with narrow ranges in the Indo-Pacific.
This however does not alter the fact that the displays of all the forms listed below are distinct.
The 43 species are arranged alphabetically since a phylogenetic presentation must await the publica-
tion of correlated morphological evidence. Display has also been observed in at least ten additional species;
since these are apparently new and undescribed, they have been omitted from the list. Motion picture
records have been obtained of more than three-quarters of all the species observed.
In the second column are given references to descriptions of waving published after 1940. An account
of earlier work on Uca display is given in Crane, 1941, p. 152. Since these early reports lacked a compara-
tive ethological viewpoint they remain, for present purposes, chiefly of historical interest and will not be
repeated here. The only references included below, therefore, are recent wave descriptions which are to
some degree specifically diagnostic. Published records which are extremely incomplete (e.g. of minax,
Crane, 1944) are omitted, as are displays not described but mentioned in connection with another subject
(e.g. Gordon, 1955, on displacement behavior).
In addition to the displays listed below which Peters (1955) has also recorded, he has described (loc.
cit.) those of macrodactyla glabromana Bott and leptochela Bott, neither of which species has been
observed by the present author. Hediger (1934) gives a helpful account of tangeri in a reference inadvert-
ently omitted by Crane, 1941.
Species
Description of
waving
since 1940
General Range
Localities where display
was observed by author
annulipes (Latreille)
—
Indo-Pacific
Pakistan (Karachi),
Ceylon, Penang, Singapore,
Sarawak, Philippines
batuenta Crane
Crane, 1941
Peters, 1955
E. Pacific
Panama, Ecuador
beebei Crane
Crane, 1941
Peters, 1955
E. Pacific
Panama, Ecuador
cumidanta Crane
Crane, 1943
W. Atlantic
Venezuela, Trinidad
deichmanni Rathbun
Crane, 1941
E. Pacific
Panama
dussumieri (Milne Edwards)
—
Indo-Pacific
Singapore, Sarawak,
Philippines
festae Nobili
—
Ecuador
Ecuador
gaimardi (Milne Edwards)
—
Western &
Central Pacific
French Oceania (Tahiti,
Raiateia, Bora Bora)
galapagensis Rathbun
—
E. Pacific
Ecuador
heterochelos (Lamarck)
—
W. Atlantic
Venezuela
heteropleura (Smith)
Crane, 1941
E. Pacific
Panama, Ecuador
inaequalis Rathbun
Crane, 1941
Peters, 1955
E. Pacific
Panama, Ecuador
insignis (Milne Edwards)
—
E. Pacific
Panama, Ecuador
inversa (Hoffmann)
—
Indo-Pacific
Pakistan (Karachi)
ischnodactyla Nemec
—
Fiji
Fiji
lactea (de Haan)
Indo-Pacific
Pakistan (Karachi),
Singapore, Philippines,
Fiji, Samoa
latimana (Rathbun)
Crane, 1941
Peters, 1955
E. Pacific
Panama, Ecuador
leptodactyla (Guerin)
—
W. Atlantic
Venezuela, Brazil
limicola Crane
—
E. Pacific
Panama
longidigitum (Kingsley)
—
E. Australia
Nr. Brisbane
1957] Crane: Basic Patterns of Display in Fiddler Crabs 79
Table 1. Species of Uca in
WHICH Display
Has
Been Studied During
THE Present Investigation
( Continued)
Species
Description of
waving
since 1940
General Range
Localities where display
was observed by author
manii Rathbun
—
Indo-Pacific
Singapore
maracoani (Latreille)
Crane, 1943
W. Atlantic
Venezuela, Trinidad,
British Guiana, Surinam,
Brazil
marionis (Desmarest)
(incl. nitida (Dana))
Altevogt,
1955.1, 1955.2
Indo-Pacific
Singapore, Sarawak, East
& North Australia,
Philippines, Fiji
minax (Le Conte)
—
W. Atlantic
New Jersey
mordax (Smith)
Beebe, 1928
Crane, 1943
W. Atlantic
Guatemala, Venezuela,
Trinidad, Brazil
oerstedi Rathbun
Crane, 1941
E. Pacific
Panama
olympioi Oliveira
—
S. Brazil
Rio de Janeiro
panamensis (Stimpson)
—
E. Pacific
Panama
princeps (Smith)
Crane, 1941
Peters, 1955
E. Pacific
Panama, Ecuador
pugilator (Bose)
Crane, 1944
Burkenroad, 1947
W. Atlantic
Connecticut, New York,
Florida
pugnax (Smith)
Crane, 1944
W. Atlantic
Massachusetts, Connecticut,
New York, Florida
piignax rapax (Smith)
Crane, 1943
W. Atlantic
S. Florida, Guatemala,
W. Indies, Colombia,
Venezuela, British Guiana,
Surinam, Brazil
rathbunae Pearse
—
Philippines
Manila, G. of Davao
rhizophorae (Tweedie)
—
Malaya & Borneo
Singapore, Sarawak
rosea (Tweedie)
—
Malaya
Penang
saltitanta Crane
Crane, 1941
Peters, 1955
E. Pacific
Panama
si gnat a (Hess)
—
Indo-Pacific
Australia (Gladstone),
Philippines
speciosa (Ives)
—
W. Atlantic
S. Florida
stenodactyla (Milne Edwards
& Lucas)
Crane, 1941
Peters, 1955
E. Pacific
Costa Rica, Panama,
Ecuador
stylifera (Milne Edwards)
Crane, 1941
E. Pacific
Panama, Ecuador
terpsichores Crane
Crane, 1941
E. Pacific
Panama, Ecuador
tetragonon (Herbst)
—
Indo-Pacific
French Oceania (Bora
Bora)
thayeri Rathbun
—
W. Atlantic
Trinidad, Brazil
zamboangana Rathbun
—
Philippines
Gulf of Davao
species display is so feebly developed that, when
any conflict or frustrating situation arises, the
crab simply stops waving.
No comment can yet be made on the role
displacement behavior, through ritualization.
may have played in the evolution of display
motions.
Studies on heliconiid butterflies in Trinidad
(Crane, 1955 and in ms.) apparently show con-
clusively the frequent occurrence of displace-
80
Zoologica: New York Zoological Society
[42: 6
Table 2. Arrangement of Uca spp. According to General Type of Wave.
Key: Species names in italics', narrow-fronts. Species names in bold-face: broad-fronts.
IP : Indo-Pacific. curtsy ^ general type of
EP: Eastern Pacific. rapping V outstanding characteristic
WA: Western Atlantic. special steps \ in advanced display.
No AD: No special advanced display characteristics.
(Where distribution symbol is not followed by one of these key designations, advanced display is incom-
pletely known. See text).
Wave Vertical
Wave Lateral
Body position
at moderate
display intensity
(Cheliped raised up-
ward at beginning of
wave; always re-
turned to position in
same plane)
(Cheliped extended to side
at beginning of wave; often
returned to position via a
circular route)
Body raised
throughout a
series of waves
Body conspicuous-
ly raised and
lowered with
each wave.
Body not raised,
at all, or mini-
mally raised and
lowered with
each wave.
zamboangana (IP)
dussumieri (IP) (No AD)
marionis (IP) (No AD)
signata (IP) (No AD)
thayeri (WA) (No AD)
annulipes (IP) (curtsy)
beebei (EP) (special steps)
festae (EP) (special steps)
galapagensis (EP)
inversa (IP)
lacfea (IP) (curtsy)
Eatimanus (EP) (curtsy)
leptodactyla (WA) (curtsy)
lEinicola (EP)
minax (WA)
mordax (WA) (curtsy)
olympioi (WA) (curtsy)
pugnax (WA) (curtsy)
stenodactyla (EP) (special steps)
terpsichores (EP) (special steps)
insignis (EP) (special steps)
maracoani (WA) (special steps)
princeps (EP)
stylifera (EP) (special steps)
batuenta (EP) (rapping)
cumulanta (WA) (rapping)
deichmanni (EP) (rapping)
inaequalis (EP) (rapping)
cerstedi (EP)
panamensis (EP)
pugilafor (WA) (rapping)
saltitanta (EP) (rapping)
speciosa (WA) (rapping)
heterochelos (WA)
heteropleura (EP) (No AD)
gaimardi (IP) (No AD)
longidigitum (Australia)
tetragonon (IP) (No AD)
ischnodactyla (IP) (No AD)
manii (IP) (No AD)
tathbunae (IP) (No AD)
rhizophorae (IP) (No AD)
rosea (IP) (No AD)
1957]
Crane: Basic Patterns of Display in Fiddler Crabs
81
ment behavior in these insects. Its probable
occurrence has been mentioned in salticid spid-
ers {idem, 1948, p. 202), mantids {idem, 1952,
p. 288) and Drosophila (Bastock & Manning,
1955, p. 104) . It now seems likely that displace-
ment behavior will prove to be a noteworthy
factor in the ethological study of many higher
invertebrates.
VIII. Summary
Two basic patterns of display have been dis-
tinguished during field studies of more than
fifty species of fiddler crabs {Uca). The first
pattern is characteristic of a group of species
with narrow fronts. It is distinguished by a
simple, more or less vertical gesture (“wave”)
made with the major cheliped of the male, and
by the male’s pursuit of the female toward her
burrow; copulation in the known instances takes
place on the surface of the ground. The second
pattern is typical of broad-fronted species in
the genus. It is characterized as follows; the
cheliped is unflexed laterally, rather than verti-
cally elevated, and sometimes completes a cir-
cular motion in returning to rest position; there
is in addition a distinct second stage of display
which is usually elicited by the approach of a
female and which depends both on special move-
ments of the various appendages and on an
increased tempo of waving; finally, in the last
stage of display the male attracts the female
down his own burrow, which he enters first.
A few species with intermediate types of be-
havior have been observed, especially in the
Indo-Pacific.
The simplest and most primitive of the nar-
row-fronted displays are found in a group of
closely related Indo-Pacific species of narrow
distribution which inhabit protected estuaries
and tidal streams; manii and rathbunae are ex-
amples. Displays of higher development are
found among species inhabiting more saline,
exposed locations; typical of these are tetra-
gonon and zamboangana, both from the Indo-
Pacific area. The narrow-fronted crabs reach
their highest display development, along with
their greatest morphological specialization, in
the neotropical representatives, culminating in
insignis and maracoani. All favor relatively ex-
posed habitats.
The broad-fronted, laterally waving fiddler
crabs are distributed in both the Indo-Pacific and
American regions but, unlike the narrow-fronted
species, they are poorly represented in the Indo-
Pacific. As in the other groups, the highest dis-
play development occurs among species living
in more seaward niches. Representatives are
found of both simple and highly evolved dis-
plays. As examples, gaimardi, from the western
and central Pacific, illustrates the simplest known
type of lateral display, while complex specializa-
tions are shown variously by lactea from the
Indo-Pacific, saltitanta and terpsichores from the
eastern Pacific and pugnax from the western
Atlantic.
Three wide-ranging Indo-Pacific species have
been observed in display over distances ranging
from 5,000 to 8,000 miles. No gross intraspecific
differences were noted on the peripheries or
elsewhere in their ranges, except that some dis-
playing lactea built small structures of sandy
mud in Karachi, but not in either the Philippines
or the Fiji Islands, at least during the period
of observation.
As is usual in a number of animals, the gen-
eral trend in display evolution in Uca is toward
increased conspicuousness. This increase is
attained principally by higher speed in waving,
by greater complexity in the rhythms and forms
of display motions and by increased apparent
size, through extension of the appendages either
vertically or horizontally. The species in which
display is highly evolved also spend a greater
portion of their time in display than do other
members of the genus.
Displacement behavior during Uca display is
briefly discussed, including its possible role in
the systematics of the group.
IX. References
Altevogt, a.
1955.1 Some studies on two species of Indian fid-
dler crabs, Uca marionis nitidus (Dana)
and U. annulipes (Latr.). Jour. Bombay
Natural History Soc., 52:700-716.
1955.2 Beobachtungen und untersuchungen an
indischen winkerkrabben. Z. Morph, u.
Okol. Tiere, 43:501-522.
Armstrong, E. A.
1950. The nature and function of displacement
activities. Symposia of the Society for
Experimental Biology, No. 4. Physiologi-
cal mechanisms in animal behavior. Aca-
demic Press, N. Y. 361-384.
Bastock, M. & A. Manning
1955. The courtship of Drosophila melanogaster.
Behaviour, 8:85-111.
Bott, R.
1954. Dekapoden (Crustacea) aus El Salvador.
1. Winkerkrabben {Uca). Senck. Biol.,
35:155-180.
Burkenroad, M. D.
1947. Production of sound by the fiddler crab,
Uca pugilator Bose, with remarks on its
nocturnal and mating behavior. Ecology,
28:458-461.
82
Zoologica: New York Zoological Society
[42: 6: 1957]
Crane, J.
1941. Eastern Pacific Expeditions of the New
York Zoological Society. XXVI. Crabs of
the genus Uca from the west coast of
Central America. Zoologica, 26:145-208.
1943.1 Crabs of the genus Uca from Venezuela.
Zoologica, 28:33-44.
1943.2 Display, breeding and relationships of fid-
dler crabs (Brachyura, genus Uca) in the
northeastern United States. Zoologica,
28:217-223.
1952. A comparative study of innate defensive
behavior in Trinidad mantids (Orthoptera,
Mantoidea) . Zoologica, 37:259-293.
1955. Imaginal behavior of a Trinidad butterfly,
Heliconius erato hydara Hewitson, with
special reference to the social use of color.
Zoologica, 40:167-196.
Gordon, H. R. S.
1955. Displacement activities in fiddler crabs.
Nature, 176 (4477) :356-357.
EXPLANATION
Plate I
Oblique views of different types of wave in fiddler
crabs. For comparison with frontal views. Text-figs.
1 and 3.
Fig. 1. Vertical wave: Uca dussumieri displaying
near Sasa, Gulf of Davao, Philippine Islands.
Maximum elevation of cheliped.
Fig. 2. Vertical wave: Uca marionis in Singapore.
Hediger, H.
1934. Zur biologie und psychologic der flucht
bei tieren. Biol. Zentralbl. Leipzig, 54:21-
40.
Peters, H. M.
1955. Die winkgebarde von Uca und Minuca
(Brachyura) in vergleichend-ethologi-
scher, okologischer und morphologisch-
anatomischer hetrachtung. Z. Morph, u.
Okol. Tiere, 43:425-500.
Tinbergen, N.
1951. The study of instinct. Oxford, at the Clar-
endon Press. 228 pp.
1952. “Derived” activities; their causation, bio-
logical significance, origin, and emancipa-
tion during evolution. Quart. Rev. Biol.,
27: 1-32.
Vervey, J.
1930. Einiges fiber die biologie ost-indischer
mangrovekrabben. Treubia, 12:169-261.
OF THE PLATE
Major cheliped partially elevated. Note for-
ward direction of merus and carpus, and
compare their position in Fig. 4, below.
Fig. 3. Lateral wave: Uca latimanus near Panama
City, Panama. Maximum elevation of cheli-
ped.
Fig. 4. Lateral wave: Uca annulipes near Santo-
bong, Sarawak. Cheliped partially elevated.
Note lateral direction of merus and carpus.
CRANE
PLATE I
FIG. 1
FIG. 3
FIG. 2
FIG. 4
BASIC PATTERNS OF DISPLAY IN FIDDLER CRABS (OCYPODIDAE, GENUS UCA)
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ZOOLOGICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
VOLUME 42 • PARTS • NOVEMBER 25, 1957 • NUMBERS 7 TO 10
PUBLISHED BY THE SOCIETY
The ZOOLOGICAL PARK, New York
Contents
7. Studies on the Lizard Family Xantusiidae. III. A New Genus for Xantusia
river siana Cope, 1883. By Jay M. Savage. Text-figures 1-3
8. Nesting Behavior of the Crested Oropendola (Psarocolius decumanus) in
Northern Trinidad, B.W.I. By Richard E. Tashian. Plates I & II; Text-
figures 1-3
9. Changes in the Cytological Structure of the Adenohypophysis and Gonads
in Juvenile Bathygobius soporator after Pituitary Implantation. By Louise
M. Stoll. Plates I & II
10. The Ctenuchidae (Moths) of Trinidad, B.W.I. Part I. Euchromiinae. By
Henry Fleming. Plates I-III
PAGE
83
87
99
105
7
Studies on the Lizard Family Xantusiidae. III.
A New Genus for Xantusia riversiana Cope, 1883
Jay M. Savage
Department of Biology, University of Southern
California, Los Angeles 7, California
(Text-figures 1-3)
IN the course of investigations leading toward
a complete revision of the night lizard fam-
ily Xantusiidae, it has become apparent
that Xantusia riversiana Cope, 1883, is so dif-
ferent from other members of the group that a
new genus is required for its reception. The
decision to place riversiana in a separate genus
is based upon an analysis of the scutellation,
body proportions, coloration, life history and
habits of all known xantusiids, combined with
an extensive study of the osteology of the family.
Detailed consideration of these features, char-
acterization of the genera and species of xantu-
siids and discussion of the evolutionary patterns
within the family are reserved for the revisional
report. However, it seems advisable to propose
the new genus in advance of the appearance of
the larger work in order to make the name avail-
able to others investigating various aspects of
the biology of the Xantusiidae. To increase the
usefulness of the present paper an artificial key
to the genera of night lizards is appended to the
description of the new genus.
There has been considerable confusion in the
past concerning the status and relationships of
several genera and species of Xantusiidae be-
cause of the lack of a standardized terminology
for the squamation of these lizards. In order to
analyze the scutellational differences and simi-
larities among members of the family it has been
necessary to revise completely the existing system
employed for the scales on these lizards. A de-
tailed discussion and definition of scutellational
terms will be included in my revision of the fam-
ily, but for the sake of consistency the new sys-
tem of scale terminology is used throughout the
present report. The head shields of riversiana are
illustrated in Text-figures 1-3. The terms radials
and femorals refer to the scales along the anterior
surface of the forearm and thigh, respectively.
Walker (1955) has utilized my system of scale
nomenclature in his descriptions of new Lepi-
dophyma from Mexico.
It is a great pleasure to have the opportunity
of naming what may well be the last new genus
of recent reptiles from western North America
in honor of Dr. Laurence M. Klauber of San
Diego, who has contributed so magnificently to
our understanding of the herpetofauna of this
region.
Klauberina, new genus
Type of Genus— Xantusia riversiana Cope,
1883.
Definition.— External characteristics: (1) one
frontonasal; (2) a large median; (3) two front-
als; (4) two parietals; (5) no pretemporals; (6)
temporals a series of enlarged plates; (7) two
rows of supraoculars; (8) nostrils pierced at
juncture between nasal, postnasal, rostral and
first supralabial; (9) pretympanics granular;
(10) postmentals and infralabials distinct; (11)
anterior postmentals paired; (12) anterior pre-
gulars granular; (13) gulars enlarged rectangular
plates, much larger than pregulars; (14) dorsal
scales granular; (15) dorsals essentially homo-
geneous in size and shape; (16) ventrals in 16
longitudinal rows; (17) radials and femorals
forming enlarged plates; (18) caudals faintly
keeled; (19) all scales in caudal whorls of same
size. Osteological characteristics: (20) nasals
meeting one another for most of their length;
83
84
Zoologica: New York Zoological Society
[42: 7: 1957]
Text-fig. 1. Diagram of
dorsal head scutellation of
Klauberina riversiana
(Cope, 1883). R = rostral;
FN - frontonasal; M =
median; F - frontal; P =:
parietal; O = supraocular
series.
Text-fig. 2. Diagram of
lateral head scutellation of
Klauberina riversiana
(Cope, 1883). N =: nasal;
PN - postnasal; PT = pre-
tympanics; S = supralabial
series; T = temporals.
(21) premaxillary not reaching to frontals; (22)
frontals paired; (23) frontals not roofing over
orbits above, outer margins concave; (24) pari-
etals paired; (25) squamosal touching parietal;
(26) prefrontal extending onto top of skull, ex-
cluding a portion of frontal from orbit; (27)
jugal moderately broad; (28) postfronto-orbital
not expanded medially, gradually decreasing in
width from anterior to posterior; (29) anterior
bony palate of paleochoanate type; (30) fenes-
tra vomeronasalis externa bordered by prevomer
and maxillary; (31) epipterygoid process from
prootic; (32) ectopterygoid and palatine in con-
tact or narrowly separated; (33) basisphenoid
and basioccipital distinct; (34) teeth on both
jaws strongly triconodont; (35) clavicles per-
forate; (36) sternum without posterior projec-
tion; (37) phalangeal formula of hand 2-3-4-S-3.
Diagnosis.— new genus may be readily sep-
arated from Cricosaura Gundlach & Peters, 1863,
and Lepidophyma Auguste Dumeril, 1851 (in-
cluding Gaigeia H. M. Smith, 1939), by the
characters presented in the accompanying key.
In addition Klauberina is distinct from these
genera in a number of the features mentioned
in the definition above (from Cricosaura in char-
acters 1-5, 8-9, 11-12, 14-16, 18-22, 24-25, 27-
32, 34-37, and from Lepidophyma in 5, 7, 8,
10-11, 13, 15-17, 23, 26-28, 34), but a complete
summary of the differences between the various
genera will be given elsewhere.
From its nearest ally, Xantusia Baird, 1859,
Klauberina is distinguished by having (charac-
ters for Xantusia given in parentheses) : external:
no pretemporal scales (pretemporals present);
temporals formed of a series of enlarged plates
(temporals formed of small granules) ; two rows
of supraoculars (one row of supraoculars) ; ven-
tral scales in 16 longitudinal rows (ventrals in
12-14 rows) ; caudal scales slightly keeled (caud-
als smooth) ; osteological: jugal moderately
broadened (jugal reduced to a narrow sliver);
ectopterygoid and palatine in contact or nearly
meeting (ectopterygoid and palatine widely
1957]
Savage: Studies on the Lizard Family Xantusiidae
85
Text-fig. 3. Diagram of squamation on underside
of head of Klauberina riversiana (Cope, 1883).
I = infralabials; PM = postmentals. The pregular
scales are the small scales lying median and pos-
terior from the postmentals and anterior to the
gular fold; the gulars are the scales on the gular fold.
separated); teeth strongly triconodont (teeth
simple) .
Klauberina riversiana is markedly different
from the recently described Eocene fossil xantu-
siid, Palaeoxantusia fera Hecht, 1956, based
upon a portion of a lower jaw from Elk Moun-
tain, Wyoming (Bridger Formation), in having
strongly triconodont teeth. The fossil genus has
simple rounded teeth.
The nominal night lizard genus Impensodens
erected on the basis of an incomplete lower jaw
from Post-Pleistocene deposits in Yucatan, Mex-
ico (Langebartel, 1953), appears to be synony-
mous with Lepidophyma according to Hecht
(1956, p. 3). In any event this mandible is dis-
tinct from that of Klauberina in having weakly
triconodont teeth with the secondary cusps lo-
cated on the inner surface of the teeth and not
in the same longitudinal plane as the primary
median cusp.. Klauberina is distinctive within the
family in having strongly triconodont teeth with
all the cusps on a given tooth located along the
same longitudinal plane.
Included Species. — Klauberina riversiana
(Cope, 1883) of the California Channel Islands,
Santa Barbara, San Clemente and San Nicholas,
is the unique member of the genus.
Notes on the Family.— In a recent discussion
of the position of the Xantusiidae within the
suborder Sauria, McDowell & Bogert (1954, pp.
94-98) acknowledge the aid of my unpublished
study on the family as a source for certain of
their remarks. Unfortunately the information
presented by them cannot be derived from the
data provided by me. Corrections therefore seem
in order.
McDowell & Bogert state that among other
features the family Xantusiidae is characterized
by paired parietal bones, absence of a parietal
foramen and ovulation involving but two eggs
at one time (with rare exceptions). Contrary to
these statements, one xantusiid, Cricosaura
typica Gundlach & Peters, 1863, has the parietal
bones fused to form a single element. Neither
can the family be characterized as lacking a
parietal foramen. Members of the genus Xan-
tusia always have this aperture and Klauberina
also consistently has the foramen (as clearly
shown on figure 25 in McDowell & Bogert),
although it may be partially or entirely covered
by osteoderms on the outer surface of the skull
in adult examples. The published studies of
Brattstrom (1951) and Shaw (1949) conclu-
sively show that Klauberina may produce four
to ten ova at one time and my own investiga-
tions indicate that Lepidophyma usually has
about six eggs formed at one period of ovulation.
The species of Xantusia produce one to three ova
but the usual number is two. In addition to these
points, it may be noted that postanal bones and
sacs are present in males of both Xantusia ari-
zonae Klauber, 1931, and Xantusia vigilis Baird,
1 859, not in vigilis alone as stated by McDowell
& Bogert.
An Artificial Key to the Recent Genera
OF Night Lizards
la. A single frontal scale; no parietal scales; a
single anterior postmental scale; nasal bones
completely separated from one another by
nasal process of premaxillary; frontal bone
single; parietal bone single; phalangeal
formula of hand 2-3-4-4-3 Cricosaura
lb. Two frontal scales; two parietal scales; a pair
of anterior postmentals or postmentals fused
with infralabials; nasal bones in contact with
one another for most of length; two frontals;
two parietals; phalangeal formula of hand
2-3-4-5-3.
2a. Supraoculars well-developed; postmentals
and infralabials distinct; scales on back
and sides essentially homogeneous in size
and shape; radials and femorals enlarged
into plates; orbit not completely roofed
over above by frontal bone, outer margin
of frontal concave; prefrontal bone ex-
tending onto top of skull, separating an-
terior portion of frontal from orbit;
86
Zoologica: New York Zoological Society
[42: 7
postfronto-orbital not expanded medially,
gradually decreasing in width posteriorly.
3a. No pretemporal scales; temporal
scales enlarged plates; two rows of
supraoculars; ventrals in 16 longi-
tudinal rows; jugal expanded; ecto-
pterygoid and palatine in contact or
narrowly separated from one another;
teeth triconodont Klauberina
3b. Pretemporals present; temporals
small, granular; one row of supra-
oculars; ventrals in 12-14 longitudinal
rows; jugal reduced to a narrow sliv-
er; ectopterygoid and palatine widely
separated; teeth simple. . . .Xantusia
2b. Supraoculars reduced to a fleshy flap
projecting from side of frontal scale;
postmentals and infralabials fused into a
single series; scales on back and sides a
heterogeneous mixture of granules and
enlarged scales; radials and femorals not
enlarged, same size as other scales on
limbs; orbit completely roofed over
above by frontal bone, outer margin of
frontal even, not concave; prefrontal
bone not extending onto top of skull, not
separating anterior portion of frontal
from orbit; postfronto-orbital expanded
medially, not gradually decreasing in
width posteriorly Lepidophyma
Literature Cited
Brattstrom, Bayard Holmes
1951. The number of young of Xantusia. Her-
petologica, vol. 7, pt. 3, pp. 143-144.
Cope, Edward Drinker
1883. Notes on the geographical distribution of
Batrachia and Reptilia in western North
America. Proc. Acad. Nat. Sci. Philadel-
phia, 1883, pp. 10-35.
Hecht, Max Knobler
1956. A new xantusiid lizard from the Eocene
of Wyoming. American Mus. Novitates,
no. 1774, 8 pp., 2 text-figs.
Langebartel, Dave A.
1954. Faunal and archeological researches in
Yucatan caves. 4. The reptiles and am-
phibians. Bull. 33, Cranbrook Inst. Sci.,
pp. 91-108, 2 text-figs.
McDowell, Samuel Booker, &
Charles Mitchell Bogert
1954. The systematic position of Lanthanotus
and the affinities of the anguinomorphan
lizards. Bull. American Mus. Nat. Hist.,
vol. 105, art. 1, pp. 1-142, pis. 1-16, text-
figs. 1-43.
Shaw, Charles Edward
1949. Notes on broods of two xantusiids. Her-
petologica, vol. 5, pt. 2, pp. 23-26.
Walker, Charles Frederick
1955. Two new lizards of the genus Lepidophy-
ma from Tamaullpas. Occ. Paps. Mus.
Zool. Univ. Michigan, no. 564, pp. 1-10.
8
Nesting Behavior of the Crested Oropendola (Psarocolius decumanus)
in Northern Trinidad, B.W.I/
Richard E. Tashian^
Department of Tropical Research,
New York Zoological Society, New York 60, New York
(Plates I & II; Text-figures 1-3)
[This paper is one of a series emanating from the
tropical Field Station of the New York Zoological
Society, at Simla, Arima Valley, Trinidad, British
West Indies. This station was founded in 1950 by
the Zoological Society’s Department of Tropical
Research, under the direction of Dr. William Beebe.
It comprises 200 acres in the middle of the Northern
Range, which includes large stretches of undis-
turbed government forest reserves. The laboratory
of the station is intended for research in tropical
ecology and in animal behavior. The altitude of the
research area is 500 to 1,800 feet, with an annual
rainfall of more than 100 inches.]
Contents
Introduction
The Nests
The Pre-incubation and Incubation Periods
The Nestling Period
Vocalization and Display
Cowbird Activity
The Nesting at Colony B
Miscellaneous Observations
Discussion and Summary
Literature Cited
Introduction
The oropendolas or giant caciques con-
stitute an assemblage of neotropical birds,
within the family Icteridae, made up of
five genera (Ocyalus, Zarhynchus, Clypicterus,
Gymnostinops and Psarocolius) and 12 species.
They are among the largest passerine birds of
the New World tropics, reaching 21 inches in
iContribution No. 977, Department of Tropical Re-
search, New York Zoological Society.
^Present Address: University of Michigan, 1133 E.
Catherine St., Ann Arbor, Michigan.
Page
. 87
. 88
. 89
. 90
. 92
. 94
. 95
. 95
. 96
. 96
length, and are characterized by similar social
habits, colonial nesting, vocalizations and dis-
play patterns. Their dominant colors are shades
of black and brown with yellow or orange char-
acteristically present in the tail and bill. Another
closely related group of icterids are the true
caciques (i.e., Cassiculus spp.) which, although
quite similar in habits and appearance to the
oropendolas, average smaller in size. Previous
studies on the 12 species comprising the oro-
pendolas have been limited to the life history
studies of the Wagler Oropendola {Zarhynchus
wagleri) by Chapman (1928) and the Monte-
zuma Oropendola {Gymnostinops montezuma)
by Skutch (1954).
The Crested Oropendola or Giant Crested
Cacique {Psarocolius decumanus insularis) is
uniform brownish-black in general coloration,
becoming chestnut on the rump and under tail
coverts; the tail is lemon yellow with the middle
pair of rectrices brownish-black; the bill pale
greenish-yellow; the tarsi and toes black and
the iris blue. The sexes are similar in color but
differ in size, the female averaging 13 inches
in length and the male 17 inches. Five subspecies
of this oropendola are recognized, ranging from
Panama across northern South America south
to southern Brazil, Paraguay, northeastern Ar-
gentina, eastern Peru and eastern Bolivia. The
race insularis is restricted to the islands of
Trinidad and Tobago and northeastern Vene-
zuela. The genus Psarocolius also includes the
Green Oropendola (F. viridis), the D’Orbigny
Oropendola (F. atro-virens) and the Chestnut
Oropendola (F. angustifrons) .
Two nesting colonies of the Crested Oro-
87
88
Zoologica: New York Zoological Society
[42: 8
pendola were discovered in early January, 1955,
in the cocoa plantation and secondary forest
near Simla about four miles north of the town
of Arima in the Northern Range of Trinidad.
This area is situated in the Arima Valley at an
elevation of 800 feet, and is bordered by lower
montane rain forest which was, in all proba-
bility, the original forest type of the study area.
For further ecological details of meteorology
and biotic zones of this region see Beebe (1952).
Both nesting sites were established in long-boled
mountain immortelle or erythrina trees {Ery-
thrina micropteryx) with the sack-like nests
hanging some 60 to 80 feet from the ground.
These immortelles are familiar nesting sites of
the Crested Oropendola in the Arima valley,
and are common trees on the cocoa plantations
where they were originally introduced as cover
for the cocoa.
The birds were still in the process of nest-
building when their sites were found. A blind
was constructed on a slope opposite the more
accessible colony (Colony A) at a level with
most of the nests and about 100 feet away. This
colony was visited daily from Jan. 12 to March
29 for a total of 303 hours. The second colony
(Colony B), about a quarter of a mile from the
first, was visited weekly from Feb. 6 to April 3
for a total of 15 hours. Observations were made
with the aid of 7 X 50 binoculars, and moving
pictures and photographs of special activities
were taken through a six-inch or 300 mm. tele-
photo lens. The majority of the observation
periods took place between 9:00 A.M. and noon
in the morning and 2:30 P.M. to 6:00 P.M. in
the afternoon. However, a number of observa-
tions were also made between dawn and 9:00
A.M. and from 6:00 P.M. to darkness.
Since it was not possible to look into the in-
accessible nests, it was especially important to
keep them under frequent observation in order
to determine any change in rhythm of the nest-
ing visits. In this manner a good approximation
of the dates of egg laying and hatching could
be made. It was felt that the observation periods
were of such regularity that determination of
these dates is accurate to within 24 hours. Un-
less otherwise stated, the figures given for the
frequency of the attentive and inattentive pe-
riods of the parent at the nest during the various
nesting phases are included only if the period
of observation exceeded three hours. A nest visit
implies complete entry into the nest, except on
those occasions during advanced feeding when
the female fed the nestling without entering
the nest.
The average monthly maximum and mini-
mum temperatures at Simla for Jan., Feb. and
March, 1955, were 61-89, 66-88 and 66-88° F.
respectively. The total monthly rainfall for the
same months was: Jan., 2.20; Feb., 2.29; March,
2.68 inches.
I am especially grateful to Dr. William Beebe
for his many helpful suggestions during this
study as well as for placing his highly informa-
tive notes at my disposal. I also wish to thank
Miss Rosemary Kenedy for assisting me so gen-
erously in the photographic phases of this work.
Dr. Edward J. Alexander of the New York Bo-
tanical Garden kindly identified the nesting
material.
The Nests
Twenty-three nests were begun at colony A
(Plate I, Fig. 1), of which 11 were completed,
and at colony B, 14 nests were begun, all of
which were completed. The nests at colony A
were fairly evenly distributed throughout the
tree, whereas at colony B they were bunched
together on one side of the tree. Incidentally, at
another colony of Crested Oropendolas, found
on March 14 in an adjacent valley at an eleva-
tion of about 1,500 feet, the nests were sus-
pended in an immortelle tree some 50 feet
above the ground and were all arranged very
closely together for a horizontal distance of only
about 20-25 feet. Eighteen nests appeared to
have been started, of which 14 seemed complete.
During a half hour’s watch feeding of the nest-
lings was observed at four separate nests.
High winds apparently take a considerable
toll of nests. Four nests (three occupied and one
unoccupied) at colony B, and one unoccupied
nest at colony A, fell during heavy winds. Two
of these fallen nests seemed typical and measure-
ments are given in Table 1.
A great increase in the length of the nest open-
ing and the dispersal of the leaf lining is ap-
parent between incubation and the third week
of feeding. Another unoccupied nest, obviously
abnormal, measured seven feet in length and
weighed 312 grams.
Examination of these nests showed that they
were composed largely of the stripped leaf fibers
of bromeliads (Bromeliaceae) and heliconia
(Musaceae) and stems of vines (Cucurbita-
ceae). The leaves used in lining the bottom of
the nests on the inside were almost exclusively
those of the immortelle (^Erythrina microp-
teryx). According to Goeldi (1897, 1903) the
Crested Oropendola from southern Brazil em-
ploys the fibers of the hromeliad Tillandsia
usneoides almost entirely as a nesting material,
whereas near the Amazon, orchid roots and
elongated roots of the rhizomorph Marasimius
sp. are used. Young (1929) describes a nest
1957]
Tashian: Nesting Behavior of Crested Oropendola
89
Table 1. Measurements of Two Nests of Psarocolius decumanus, in Incubating
AND Feeding Nesting Stages
Length of
opening
Weight (gms.)
Nesting phase
Length
Width
Fiber
Leaf
lining
Total
Incubating
2T0"
8"
5"
93.1
87.7
180.8
Feeding
2' 11"
9"
1'5"
178.8
4.1
182.9
from British Guiana as made up of narrow
strips of cane leaves, plantain bark, cocoanut
fiber and vines. He also states that two or three
nests are sometimes built by the same female.
The color of some broken egg shells recov-
ered from a fallen nest was a pale bluish-green
or, by comparison with Ridgway (1912), Pale
Glaucous Green.
The Pre-incubation and Incubation Periods
Dr. William Beebe (unpublished notes) noted
nest building in the Crested Oropendola to be
under way in the Arima valley as early as Dec.
5 (1952) and Dec. 23 (1953) and as late as
March 4 (1950). Belcher & Smooker (1937)
reported that in Trinidad and Tobago nest build-
ing began in December and eggs were found
from mid-January onwards, with May 24 as the
latest date.
The nesting colonies near Simla were not dis-
covered until Jan. 9, 1955, at colony B and
Jan. 11 at colony A, at which time nest building
was well advanced at both sites. There had been
no activity at either tree in late December, 1954,
and nest building therefore must have begun
either in late December or early January and
lasted about two weeks.
Sixteen of the 23 nests started at colony A
were being actively worked on during the obser-
vation period on Jan. 12. On the following day,
however, with 12 nests still incomplete, only
two nests were visited by the birds during almost
eight and one-half hours of observation. Al-
though most of the nests were no longer actively
visited after Jan. 12, there was considerable ac-
tivity in the nest tree between Jan. 13 and Jan.
18. Periodically small groups made up mainly
of females, many of which carried nesting mate-
rial, would fly into the top of the tree. Here
they would remain for varying periods of time,
clucking noisily, but with only a few exceptions
they made no further attempts at nest building.
Of the 16 nests being built on Jan. 12, only
eight were observed to be revisited for the pur-
pose of nest building on subsequent days; one
was visited once, two were visited twice, one
was visited five times and one was visited six
times, and at only three were visits continued
to egg laying and incubation. These last three
nests (1, 2 and 3) showed no similarity in the
length of time between the end of nest-building
and the start of incubation. This pre-incubation
activity is summarized in Table 2.
The earliest onset of incubation took place
at nest 1 on Jan. 20. In all probability, this nest
was completed prior to Jan. 12. Nest 2, how-
ever, was not completed until Jan. 14, and
incubation did not begin until Feb. 5. Nest 3
appeared to be complete on Jan. 12, with incu-
bation commencing on Feb. 6.
Just why colony A was seemingly abandoned
is difficult to explain, for it appeared to be
thriving. The observation post was well camou-
flaged with canvas and concealed by vegetation,
in addition to being situated some 100 feet from
the nest tree. Moreover, these birds are not par-
ticularly known for their shyness, and active
colonies are to be found in well-populated areas.
In fact, colony B flourished despite its proximity
to a rock quarry where frequent blasting was
taking place. Dr. Beebe has noted that in the
previous three years, as well as the following
year (1956), this site always contained medium-
sized colonies.
As can be seen by Table 2, nests 2 and 3 were
visited frequently and fairly regularly before in-
cubation. It appeared that incubation was under
way at nest 3 on the afternoon of Jan. 20, as
the female remained in her nest for periods
ranging from 23 to 112 minutes during three
and a half hours of observation. However, from
Jan. 21 to 30 no further activity was noted at
nest 3. With this one exception, the length of
time spent in the nests at each visit was usually
less than a minute, and exceeded five minutes
on only three occasions.
Oftentimes, nests 2 and 3 were visited in
seemingly unaccountable spurts over relatively
short periods of time. The most outstanding
example of this behavior took place at nest 2
on the morning of Jan. 29 where, within a period
of 44 minutes, the nest was visited briefly 22
times.
The significance of this pre-incubation activity
90
Zoologica: New York Zoological Society
[42: 8
Table 2. Pre-incubation Activity by the Female Parents at Colony A
Date
Total observation
time (min.)
Number of visits per hour
Nest Nest Nest
1 2 3
Average time in nest (min.)
Nest Nest Nest
1 2 3
Jan.
12
173
0.6
0.3
1.3
<1.0
<1.0
<1.0
13
501
0
0.1
0
0
<1.0
0
14
540
0
0.4
0
0
<1.0
0
15
575
0
0
0.1
0
0
<1.0
16
285
0
0
0
0
0
0
17
410
0.8
0
1.1
<1.0
0
<1.0
18
444
0.9
2.0
1.8
3.7
1.3
<1.0
19
240
0.7
2.7
0.8
<1.0
2.0
9.0
20
210
*
1.1
1.1
0.8
58.6
21
180
0
0.6
0
<1.0
22
350
4.1
0
3.1
0
23
245
1.7
0
1.3
0
24
370
0
0
0
0
25
215
2.0
0
2.3
0
26
226
0.5
0
<1.0
0
27
361
4.1
0.1
4.5
<1.0
28
131
3.2
0.4
6.0
<1.0
29
131
11.9
0.9
0.8
<1.0
30
190
2.0
1.2
6.7
4.7
31
352
1.7
0.8
4.4
<1.0
Feb.
1
170
0.3
0.3
20.0
<1.0
2
350
0.6
1.3
7.5
<1.0
3
80
0
0
0
0
4
83
0.7
2.4
<1.0
<1.0
5
341
0
0
^
^Onset of incubation.
is not clear. Since nests 1, 2 and 3 were com-
plete by Jan. 14, after which no nesting mate-
rial was brought to the nests, and no nest weav-
ing detected, these facts appear to eliminate
the possibility that the hens involved were giv-
ing “finishing touches” to their nests. Activity
ranged from the apparent pseudo-incubation
behavior at nest 3 on Jan. 20 to the extremely
frequent nesting visits at nest 2 on Jan. 29.
The beginning of incubation was determined
by observing the increase in the duration of
the attentive periods by the female at her nest.
In colony A, the laying of the first egg at nest 1
occurred on Jan. 20, at nest 2 on Feb. 5 and at
nest 3 on Feb. 7. The incubation period at all
three nests lasted approximately 15 days. During
this time the average time spent on the eggs
was 74.6% at nest 1, 72.0% at nest 2 and
67.9% at nest 3. The frequency and duration of
the attentive periods are indicated in Table 3
and Text-figures 1-3. The percentage of time
spent in the nest was determined by dividing
the average length of the attentive periods by
the sum of the average attentive periods and
multiplying by 100. The number of visits per
hour at all nests ranged from 0.5 to 1.9, averag-
ing 1.0 at nests 1 and 3 and 0.8 at nest 2.
For at least seven days prior to egg laying,
the female parents at nests 2 and 3 were seen
to return to their empty nests at the onset of
darkness, and presumably spent the night there.
The times of these final entries, as well as those
noted during incubation proper, varied from
about 5:55 to 6:20 P.M.
The Nestling Period
With the sudden increase in the number of
nest visits per hour, the hatching of the first egg
was considered to have taken place. This oc-
curred at nest 1 on Feb. 4, at nest 2 on Feb. 21
and at nest 3 on Feb. 22. As with incubation,
all parental care was undertaken by the females.
The length of nestling life varied from 31 to
32 days at nest 1 and 34 or 35 days at nest 2
to 36 days at nest 3. The amount of time that
the hen spent in the nest with the newly-hatched
young decreased gradually to less than about
20%. We can refer to this as the brooding
1957]
Tashian: Nesting Behavior of Crested Oropendola
91
period. At nest 1 this brooding period lasted for
seven days, at nest 2 for 11 days and nest 3 for
nine days. After the last day of brooding the
frequency of visits ranged from 3.2 to 8.5 per
hour, averaging 5.7, 5.6 and 5.9 at nests 1, 2
and 3 respectively. Time spent at the nest was
usually less than 5% at nests 2 and 3 during
the post-brooding period, the length of the aver-
age visit being but a few seconds. At nest 1,
however, it varied from 5% to 15%, a factor
which might possibly have been due to the
presence of more than one nestling.
After the cessation of brooding, the females
stayed with their nestlings all night for a varying
number of days. At nest 1, this period extended
for 11 days after hatching, or 34% of the nes-
tling stage; at nest 2 for 26 days (74%), and at
nest 3 for 23 days (63%).
The exact nature of the food brought to the
netslings could not be determined; however,
large orthopterous insects or lepidopterous larvae
were frequently noted. Excrement packets were
periodically removed from the nests.
By the 18th or 19th day of feeding, the
whining of the nestlings, as well as their move-
ments within the nests, could be detected. The
young birds are either able to see through the
nest wall, or can hear the wing sound of the ap-
proaching parent, as often this whining began
even before the hen had reached the nest. It
usually continued on through the feeding and
for a few minutes after she had left. The male
can also bring about this whining, for on one
occasion a male flying to a branch near an occu-
pied nest caused the young to whine. This, in-
cidentally, induced the male to explore the nest
curiously, but he did not enter.
During the early days of feeding the hen en-
tered the nest entirely. But as the nest opening
became enlarged by her frequent passage and
the nestling increased in size, she fed it either
while perching at the nest entrance (Plate I,
Fig. 2) , or while clinging to the side of the nest.
In the last days of nestling life, the young bird
at nest 3 was sometimes seen to climb entirely
out of the nest. Then, while clinging to the side
and flapping its wings, it would at times com-
pletely circle the nest before re-entering.
Nest-leaving took place at nest 1 sometime
between 11:30 A.M. on March 7 and 3:30 P.M.
on March 8 (31 or 32 days after hatching), at
nest 2 between 3:49 P.M. on March 27 and
8:09 A.M. on March 28 (34 or 35 days after
hatching) and at nest 3 at 9:07 A.M. on March
29 (36 days after hatching). The following
notes were taken on the morning of nest-leaving
at nest 3:
7:49 A.M. Observer entered blind.
7:50-8:53 Nestling fed on 5 separate occa-
sions.
9:02 Nestling had head out of the nest
opening before hen reached nest;
after being fed it climbed com-
pletely out of the nest, perching at
the entrance and peering about for
a few minutes before re-entering.
9 : 07 After being fed, the nestling perched
at the nest entrance for a few sec-
onds, then exercised its wings mo-
mentarily and flew off in a fairly
strong, slightly descending flight,
alighting in a tree some 75 feet
away.
9:08 Fledgling whined and vibrated
wings while being fed.
Table 3. Per cent, of Time Spent in Their Nests
BY THE Female Parents at Colony A during the
Incubation and Brooding Periods
Number of
days after
egg laying
Nest 1
Jan. 20*
Nest 2
Feb. 5*
Nest 3
Feb. 7*
Incubation
1
81.8
81.0
2
98.8
—
71.8
3
41.6
81.3
4
81.6
79.8
73.0
5
78.1
—
58.5
6
59.4
63.9
66.1
7
60.7
75.1
8
68.1
84.5
75.4
10
60.9
—
73.9
1 1
60.9
—
71.2
12
74.0
69.5
65.1
13
—
72.9
85.0
14
42.6
61.5
_
15
—
83.2
72.0
Brooding
16
_
56.0
17
31.9
27.1
59.0
18
—
52.3
23.0
19
22.8
38.3
34.1
20
21.5
44.3
34.5
21
—
25.0
22
31.6t
—
24.7
23
42.1
28.4
24
—
23.7t
25
32.4
26
22.lt
*Probable dates of egg laying.
tLast day brooding observed. Percentages not includ-
ed if observation period was less than 3 hours.
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Text-fig. 1. Frequency of nest visits and duration of attentiveness by female parent at nest 1 in colony A.
9:09
Fledgling flew to another perch
about 25 feet away where it re-
mained perching quietly.
9:25
Fledgling fed by hen.
9:30
Fledgling flew to another tree about
50 feet distant, again remaining
quiet after landing.
9:43
Fledgling flew off again; no longer
observable.
No more than one nestling was seen at any of
the nests at colony A.
Vocalization and Display
At first hearing, any description of the various
calls of the male Crested Oropendola appears
hopeless, as there does not seem to be any par-
ticular pattern to his polyphonic and diverse
utterances. However, after prolonged and care-
ful listening, they were recognized as falling into
four groups:
PER CENT. TIME IN NEST
1957]
Tashian: Nesting Behavior of Crested Oropendola
93
7 §
O
3:
Uj
3 ^
Text-fig. 3. Frequency of nest visits and duration of attentiveness by female parent at nest 3 in colony A.
Call 1. An introductory whaaa cheeee with
the second phrase higher and fol-
lowed by a continuous, unmusical
trUI;
whaaa
Call 2. Introduction a rising unmusical trill
(rrrrrrrrrup) followed by cheow
wow, eow woo woo or cheeeeeow,
not unlike the meow of a cat:
cheow wow
Call 3. An introductory cheeeee (falling in
pitch), or a rising brrrrrree,
cheeeeeow or rrrrrrrrrup, usually
diminishing in intensity, followed by
lob lob baah or ko ko plo, repeated
three or more commonly four times;
the first two notes are given rapidly
and the last is drawn out:
ko ho plo ko ko pio ko ko plo
Call 4. An introductory peeeeeow or
cheeeeee (falling) or rrrrrrrup (ris-
ing), diminishing in intensity, fol-
lowed by a series of ko, plo or kok
notes usually repeated five times on
the same level:
ko ko ko ko ko
The most common was Call 1 which com-
prised 45% of all of those heard. Call 2 was
given 24% of the time, Call 3, 16%, and Call
4, 15%.
On numerous occasions the male was heard
to utter a confusion of stuttering, gurgling notes,
containing some of the elements of a typical
display call but wholly lacking in pattern:
oooorrrrrup whup whaaaat upka brrrrrr woo
woe whup wheel brrrrreee woo, etc. This was
accompanied by much gurgling, feather rustling
and wing flapping. It was almost as though a
young male were attempting to imitate the call
of an adult and failing badly. Every once in a
while a male would rather half-heartedly begin
a display, but never really finish. Both the abor-
tive gurgling and the unfinished calls were heard
during all nesting periods.
Calls 1, 2 and 3 usually accompanied a dis-
play, whereas Call 4 was apparently given in the
absence of any display.
The male was seen to go through his complete
display not only in the presence of females, but
also when alone or in the company of males. Dr.
Beebe observed two males simultaneously going
through a complete display while facing each
other. They were some distance from the nest
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Zoologica: New York Zoological Society
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tree, and no females were noted in the vicinity.
In the typical display, the bird begins by
bending forward in a deep bow until his head
is well below the perch. At the same time he
ruffles his body feathers and brings the extended
wings together over the back. The wings are
then vibrated rapidly with the primaries open.
Almost immediately after the wings are ex-
tended, the tail is brought forward quickly over
the back until it makes an approximate 90°
angle with the back (Plate II, Figs. 1 & 2), and
remains in this position from 1.5 to 1.9 seconds.
Then, as the tail begins to lower to its normal
position, the wing beats become less rapid and
take on a flapping or fanning quality which lasts
from 1.1 to 2.6 seconds. Sometimes, however,
the wing beat does not slacken until the tail has
been completely lowered. Finally, the wing flap-
ping stops and the bird comes out of his bow,
the entire performance lasting about four to
five seconds. These times are based on the
analysis of moving pictures taken of five com-
plete displays. The average duration of the four
main phases of the display are as follows:
1. Tail elevated forward over
back. 0.20 seconds
2. Wings brought together over
back and vibrated. 1.74 seconds
3. Tail lowered to normal
position. 0.36 seconds
4. Wing fanning or flapping 1.97 seconds
The intervals between beats in the wing-fan-
ning phase were measured at 0.03 seconds. Al-
though not actually measured, many of the dis-
play calls seemed to last around five to seven
seconds. In all probability, the figures above
would have averaged higher if more displays
had been analyzed.
Although the spreading of the tail feathers
was not noted in the present study. Young
(1929) reported it in Psarocolius decumanus
from British Guinea.. This tail spreading during
the display is probably characteristic of all
oropendolas, as Fuertes (1916) observed it for
the Montezuma Oropendola and Chapman
(1928) for the Wagler Oropendola.
Displays were frequently performed on the
nests proper. In this type of display, the male
flew directly to the nest and, clinging to its side,
began the display almost immediately upon
alighting. The head was always directed down-
ward. In such a position only the initial bowing
of the perching display was, of necessity, omit-
ted. Twenty-eight per cent, of all displays at the
nest tree were of this type. Crandall (1914) also
described this type of nest display for the Monte-
zuma Oropendola in Costa Rica.
During the 303 hours of observation at colony
A, 683 displays were counted either at the nest
tree or from the immediately surrounding trees.
The displays from the nest tree numbered 320,
and those from the nearby trees, 363. The fre-
quency of displays given on or near the nest
tree ranged from 0.1 to 10.2 per hour, with an
average of 2.2 per hour throughout all nesting
periods. At no time of the day was the display
observed to be given more frequently.
Often a male would fly to a nest and perform
no display. On the other hand, there were in-
stances where a male displayed on as many as
five different nests in succession. There seemed
to be no particular preference in selecting the
nests to display on, and oftentimes an occupied
nest was flown to while a hen was incubating,
whereupon she would usually vigorously chase
the male just as she would a cowbird. Sometimes
when a group of males were in the nest tree and
one of them flew to a nest, another male would
chase it off and, as though stimulated by this
chase, then go through a complete nest display.
On one occasion this type of behavior was re-
peated three times in the course of about three
minutes by the same pair in the same respective
roles. It was a fairly common practice for males
accompanying returning females to the nesting
colony to display almost as soon as they arrived
in the nest tree.
The most common note of the female was a
cluck which she sometimes uttered repeatedly
for considerable periods of time while perch-
ing, preening or flying about the nest tree. Often,
when she first came to the nest tree after an
inattentive period, during either incubation or
feeding, she clucked continuously sometimes up
to 15 minutes before finally entering her nest.
Less frequently she carried on this clucking after
leaving the nest. When annoyed or disturbed,
the clucking became more rapid. Although not
noted for certain, the male probably also gives a
similar cluck note. Skutch (1953) reports this in
the male Montezuma Oropendola. In chasing
cowbirds, males or other females from her nest
or its vicinity, the hen generally voiced a harsh,
nasal mewing rising at times to a high-pitched
whine.
As is characteristic of aU oropendolas and
caciques, a warning cack is given by the male
at the approach of danger. This seemed to be
most often brought on either by the sight or
scream of the Gray Hawk {Buteo nitidis), or
the sudden appearance of man. Frequently after
the sounding of an alarm cack, the oropendolas
present failed to leave the nest tree or its imme-
diate area, and incubating hens rarely left their
nests.
Cowbird Activity
The visitations of Giant Cowbirds (Psomo-
1957]
Tashian: Nesting Behavior of Crested Oropendola
95
colax oryzivorus) to colony A were carried on
with persistant regularity between Jan. 13 and
March 4. On the latter date the nestlings were
26 to 43 days old. After March 4 only one
further cowbird visit was noted, on March 23.
On 34 of the 50 days between these dates, cow-
birds were seen coming to the nest tree at one
time or other on 52 separate visits. Cowbird
activity was recorded on 77% or 29 of the 37
days in which the observation periods lasted
longer than three hours.
It was interesting to note that on 34% of their
visits, the cowbirds came to the nest tree follow-
ing returning female oropendolas. In the ma-
jority of cases only one cowbird visited the
colony at any one time. However, on five occa-
sions two cowbirds came to the nest tree to-
gether, and once a group of four came in
together.
Usually the cowbirds immediately began in-
specting a series of nests. At colony A most of
the nests were unoccupied. The process of the
inspecting consisted in momentarily peering
into the nest opening, and only on two occa-
sions were cowbirds seen to enter the nests com-
pletely. If the nest happened to be occupied by
an incubating female, she would always give
her scolding whine and usually give immediate
chase. The cowbird was chased for only a short
distance, after which the female oropendola
would usually immediately return to her nest.
Often the same cowbird would return immedi-
ately, only to be chased again from the same or
other nests. Only three instances were noted
when an incubating or brooding female did not
leave her nest to chase a cowbird, but instead
merely voiced a scolding whine from within.
On several occasions cowbirds were chased from
unoccupied nests by females who happened to
be either returning or leaving their nests. These
chases were not especially energetic, however,
and pursuit was not vigorous.
Male oropendolas were rarely seen to chase
cowbirds, and when both cowbirds and males
were in the nest tree at the same time, neither
seemed very disturbed by the other’s presence
even when perched only a few feet apart. If
cowbirds remained quietly in the nest tree they
were never observed being chased even by
females.
Cowbirds were never noted to remain for very
long in the nest tree during inspecting visits. The
average length of time was only two to three
minutes.
From all indications, cowbirds appeared to
have had little success in parasitizing any nests
at colony A. Only two nests were ever observed
to be entirely entered, and these for but a few
seconds.
The Nesting at Colony B
The various nesting phases at both colonies
were relatively comparable. On the first day of
observation at colony B (Feb. 6) nestlings
were present in three nests and incubation was
under way in five of the 1 1 nests eventually oc-
cupied. On the same date at colony A, one nest
contained young and two were incubating. As
shown in Table 4, three of the 14 original nests
were never occupied, and of the remaining 11,
three fell, leaving eight (73%) of the occupied
nests to produce fledged young.
Nest number 3, which the nestlings had left
by March 6, was seen to be revisited on March
13. During observation periods in the following
three weeks a female was seen to visit it, but the
nature of the visits could not be determined
with any certainty. This might possibly be an
instance of the rearing of a second brood. In
1956 Dr. Beebe noted nest activity by females
at colony A as late as June 4, again indicating
the possibility of a second brood
The general activity at colony B was much
the same as that at colony A, but naturally on
a larger scale. The times of the various nesting
phases were similar, with incubation lasting
about two weeks and feeding from about four
to five weeks.
Miscellaneous Observations
Most of the time the females went about their
various activities independently. However, it
was not uncommon to see two hens leave or
return to their respective nests at the same time.
This was observed not only when they left the
nests for the first time in the morning and at
the final entry at the coming of darkness, but
also during regular daytime activity. For ex-
Table 4. Summary of the Nesting Phases for
the Occupied Nests at Colony B. (I=incubat-
ing; N=nestling period; F=nestlings fledged;
X=nest fell; R=nest reoccupied).
Nest
February
March
6
13
20
27
6
13
20
27
1
I
X
2
N
N
N
X
3
N
N
N
N
F
R?
4
N
N
N
N
F
5
I
I
I
N
N
X
6
I
I
N
N
N
F
7
I
I
I
N
N
N
N
F
8
—
I
I
N
N
N
N
F
9
—
I
I
N
N
N
N
F
10
I
I
I
N
N
N
N
N
11
I
I
I
N
N
N
N
N
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Zoologica: New York Zoological Society
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ample, on March 4 the three females at colony
A, who were feeding their nestlings, either en-
tered or left their nests 218 times during six and
a half hours of observation., On 23 (10%) of
these nest entries or departures, two females
were seen either to leave or arrive at their nests
together.
Copulation was seen on but one occasion at
the nest tree; this took place in the late after-
noon (6:00 P.M.) on Feb. 2. The male ap-
proached one of a pair of females and displayed,
after which he began to peck at her cloacal area.
The female fluttered her wings but otherwise
remained fairly still. The male then engaged in a
brief copulatory act and followed this by peck-
ing at her cloacal region once more, while she
squatted and fluttered her wings again slightly.
The male then went through another complete
display.
Other than the cowbirds, all species of birds
that visited the nest tree at colony A remained
unmolested. Once a Gray Hawk {Buteo nitidis)
flew into the nest tree and perched undisturbed
for a few minutes before flying off. It was ap-
parently unseen by the oropendolas of the
colony, as the recognized presence of a hawk in
the vicinity would invariably elicit the sounding
of the alarm cack. Once at colony B, when a
Gray Hawk flew overhead, several oropendolas
took off in pursuit.
Discussion and Summary
The nesting activity of two colonies of the
Crested Oropendola (Psarocolius decumanus
insularis) in the Northern Range of Trinidad,
B. W.I., was studied from the end of nest build-
ing to nest leaving by the young. Fourteen nests
were completed at one colony all of which were
later occupied, and 11 were completed at the
other, of which three were occupied. The rela-
tive length of the various nesting phases as well
as the general activity were similar at both
colonies. Nesting success at the colony with the
three occupied nests was 100%, and at the
colony with the 11 occupied nests, 73%. Nest
building and care of the young were under-
taken entirely by the females., The following
discussion and summary are based on the obser-
vations at the colony containing the three occu-
pied nests.
The period from the completion of nest build-
ing to the beginning of incubation varied from
7 to 25 days with the incubation period proper
lasting 15 or 16 days. This is but slightly longer
than the incubation of 11 to 14 days reported
in the literature for the family Icteridae. The
average time spent on the eggs ranged from
68% to 75%.
The nestling period varied from 31 to 36 days,
and brooding from 7 to 1 1 days. These nestling
periods do not vary significantly from those of
30 to 37 days given by Chapman (1928) and
Skutch (1954) for two other oropendolas:
Gymnostinops montezuma and Zarhynchus
wagleri. After the cessation of brooding the
females remained with the nestlings at night
from 11 to 26 days, or 34% to 74% of the
nestling period.
The different phases of the male perch dis-
play were analyzed with the aid of motion pic-
tures. Comparisons with the descriptions in the
literature of the display of other oropendolas
{Psarocolius decumanus, Chapman, 1894,
Young, 1929; Gymnostinops montezuma,
Fuertes, 1916, Skutch, 1954; and Zarhynchus
wagleri, Chapman, 1928) emphasize the marked
similarity of the sequence of display phases
among oropendolas. This similarity is brought
out in Plate II, which compares the perch dis-
plays of the Crested and Green Oropendola. Dis-
plays were frequently performed on the nests
proper, and 28% of all displays at the nest
tree were of this type. The frequency of all dis-
plays given on or near the nest tree averaged
2.2 per hour throughout all nesting periods.
The various calls of the male were analyzed
and found to fall into four separable groups.
These calls, although of a similar quality to
those reported for other oropendolas (op. cit.),
appear to differ considerably, not only within
the oropendola complex, but also within the
same genus. Beebe (unpublished notes) records
Psarocolius decumanus from Trinidad as calling
wholly unlike Psarocolius angustifrons from
Venezuela; and Friedman & Smith’s (1950) de-
scription of the call of Psarocolius decumanus
from northern Venezuela does not seem to re-
semble closely any of the calls of the Trinidad
birds.
Persistant cowbird activity was noted from the
termination of nest building on to advanced
feeding, at which time the nestlings were 26 to
43 days old. During this period cowbirds were
seen to visit the colony to inspect nests on about
77% of the observation days. This type of per-
sistance by Giant Cowbirds was also noted by
Skutch (1954) in a colony of Montezuma Oro-
pendolas.
Literature Cited
Beebe, W.
1952. Introduction to the ecology of the Arima
valley, Trinidad, B.W.I. Zoologica, 37:
157-184.
Belcher, C. & G. D. Smooker
1937. Birds of the colony of Trinidad and To-
bago. Ibis, pp. 226-550.
I
.1
I
I
■ji
^ i;
1
1957]
Tashian: Nesting Behavior of Crested Oropendola
97
Chapman, F. M.
1894. On the birds of the island of Trinidad.
Bull. Am. Mus. Nat. Hist., 6: 1-86.
1928. The nesting habits of Wagler’s Oropen-
dola (Zarhynchus wagleri) on Barro
Colorado Is. Bull. Am. Mus. Nat. Hist.,
58: 123-166.
Crandall, L. S.
1914. Notes on Costa Rican birds. Zoologica,
1: 325-343.
Friedmann, H. & F. D. Smith, Jr.
1950. A contribution to the ornithology of north-
eastern Venezuela. Proc. U. S. Nat. Mus.,
100: 411-538.
Fuertes, L. a.
1916. Impressions of the voices of tropical birds.
Smithsonian Report for 1915 (Pub. 2392),
pp. 299-323.
Goeldi, E.
1897. On the nesting of Cassicus persicus, Cassi-
cus oryzivora, Gymnomystax melanicterus
and Todirostrnm maculatus. Ibis, pp. 361-
370.
1903. On the nesting of O. decumanus near
Para, and the materials of its nest. Ibis,
pp. 630-632.
Ridgway, R.
1912. Color standards and color nomenclature.
Publ. by the author. Washington, D. C.
Skutch, a. F.
1954. Life histories of Central American birds,
families Fringilidae, Thraupidae, Icteri-
dae, Parulldae and Coerebidae. Pacific
Coast Avifauna No. 31, pp. 287-304.
Young, C. B.
1929. A contribution to the ornithology of the
coastland of British Guiana. Part III.
Ibis, pp. 221-261.
98
Zoologica: New York Zoological Society
[42: 8: 1957]
EXPLANATION OF THE PLATES
Plate I
Fig. 1. View of colony A from observation blind
(February).
Fig. 2. Female parent feeding nestling in nest 3
at colony A (March).
Plate II
Figs. 1 & 2. Male Crested Oropendolas (Psarocolius
decumamts) displaying in nest tree at
colony A.
Fig. 3. Beginning of display in the male Green
Oropendola (Psarocolius virens) . New
York Zoological Park.
Fig. 4. End of display in the male Green Oro-
pendola. Note bow is deeper than in P.
decumanus.
TASHIAN
PLATE 1
NESTING BEHAVIOR OF THE CRESTED OROPENDOLA (PSAROCOLIUS DECUMANUS) IN NORTHERN TRINIDAD, B.W.I
TASHIAN
PLATE II
FIG. 1
FIG. 3
FIG. 4
NESTING BEHAVIOR OF THE CRESTED OROPENDOLA IPSAROCOLIUS DECUMANUS) IN NORTHERN TRINIDAD, B.W.I.
9
Changes in the Cytological Structure of the Adenohypophysis and
Gonads in Juvenile Bathygobius soporator after Pituitary Implantation^
Louise M. Stoll
American Museum of Natural History, Department of Fishes and Aquatic Biology
(Plates I & II)
Introduction
The relationship of the pituitary and the
gonads has been studied and acknowl-
edged in all vertebrate classes. General
seasonal changes in the pituitary and gonads of
the goldfish and carp were described by Scruggs
(1951) and correlated with the previous liter-
ature on the subject in teleosts. No report has
been found on the histology of the the specific
cell types of the pituitary correlated with the
histology of the gonads after experimentally in-
duced changes in the two tissues. The present
report is a histological study of gonads and trans-
itional lobes of pituitaries of juvenile fish after
pituitary implantation.
All fish used were collected in shallow water
areas around North Bimini Island, Bahamas,
B. W. I. Experimental work was done at the
Lerner Marine Laboratory on Bimini.
The help given during this study by Dr. C. M.
Breder, Jr., and Miss Priscilla Rasquin is grate-
fully acknowledged.
Materials and Methods
The marine teleost Bathygobius soporator
(Cuvier & Valenciennes) was used. This species
is very hardy and will survive the handling in-
volved during implantation. Since these fish
do not engage in active swimming to a large
extent, the incision healed rapidly and no
stitches were necessary.
Twenty-five young Bathygobius soporator of
an average standard length of 38 mm. were given
intraperitoneal implants of whole pituitary
^This study was supported in part by a grant from
The National Science Foundation.
glands from larger fish of the same species. The
average standard length of the donors was 66
mm. All 25 fish received three pituitaries each.
Pituitaries were obtained by the following pro-
cedure. The donor fish was heavily anesthetized
in ethyl urethane dissolved in sea water. The
lower jaw was removed and the floor of the
cranium with its covering membranes was cut
away with small bone clippers, exposing the
pituitary. The area for cutting was determined
by the position of the saccus vasculosus, which
could be seen through the bone. Pituitaries were
removed by grasping the infundibulum with
watch-maker forceps and pulling gently. The
pituitaries were held in physiological saline
(0.6%) until all three were collected.
The recipient fish was lightly anesthetized
with ethyl urethane and placed on cotton sat-
urated in sea water. A slit was made in the ab-
dominal wall lateral to the midline with watch-
maker forceps and the three pituitaries were
introduced through the slit. A dissecting micro-
scope was used white removing and implanting
the glands. Immediately after implantation, the
fish were placed in individual two-gallon tanks
with running sea water.
The gonads of the donor fish were removed
and fixed in Bouin’s picro-formol solution, em-
bedded in paraffin, sectioned at 10^, and stained
with Harris’s haematoxylin and eosin. The fish
receiving the implants were killed after varying
lengths of time. Eighteen fish were killed at
seven to eight days, and seven fish from two and
one-half to three days post-operative. Their
bodies, minus the heads, were fixed in Bouin’s,
sectioned at lO/i, through the region of the gonads
and stained with Harris’s haematoxylin and eos-
in. Heads of the 25 experimental fish were fixed.
99
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Zoologica: New York Zoological Society
[42; 9
embedded in paraffin and sectioned at 5/u.. The
modification of Halmi’s paraldehyde fuchsin
stain by Sokol (1953) provided an excellent
stain with which to observe the response of any
specific cell group of the pituitary to a given
experimental procedure. Bouin’s fixative and the
paraldehyde fuchsin stain were used for the
lieads of 18 of the experimental fish, and Baker’s
fixative and acid haematin stain were used for
the seven other experimental fish. The paralde-
hyde fuchsin was used specifically for the study
of basophils and the acid haematin for acidophils.
A group of 11 normal fish were killed and
prepared for histological study by the same
methods used for experimental fish. The normal
fish ranged from 80 mm. to 29 mm. in standard
length. All stages of normal gonad development
and maturation were represented in this series.
Tavolga (1955) reported that B. soporator
collected in the area of Marineland, Florida,
were at the height of the spawning season in
July and August., However, the largest gobies
which were collected for the present study in
Bimini waters were judged by gross dissection
and histological section to be in or near spawn-
ing condition in March and April. Testes were
swollen, nearly white in color and had high
epithelium; ovaries were bright yellow and thin
walled as described by Vivien (1941) as charac-
teristic of the spawning condition for the
European goby, Gobius paganellus.
The largest female collected by Tavolga
(1954) was 65 mm. in standard length, while
males of 90 mm. or more were found. In Bimini,
the largest female found was 87 mm. and the
largest male 108 mm. The mean, of course, was
lower, the average of 43 females being 66 mm.
and of 56 males, 74 mm.
Testes of eight normal Bathy gobius were
stained with Sudan Black B and by the Baker
technique for phospholipids. The testes of one
fish which had received three pituitary implants
and maintained for one week was also stained
with Sudan Black B.
Results
Implants of pituitaries of mature fish to juve-
nile fish caused degranulation of the basophils of
the adenohypophysis and stimulation of the
gonads. The acidophils of the transitional lobe
were not affected. In both experimental and con-
trol fish, the acidophils were large, well-rounded
and heavily stained with the Orange G of the
paraldehyde fuchsin stain. The acidophils
stained with acid haematin did not show degranu-
lation.
Degranulation of the basophils occurred in
nine fish which had received implants and were
maintained for one week before being killed.
Little or no degranulation was seen in seven fish
sacrificed three days after receiving pituitary im-
plants, nor in two fish maintained for one week
before killing. The latter had gonads in very
early stages of maturation and it is thought that
the tissue may be refractory at this early stage.
Partially degranulated basophils of a three-day
implant are shown in Plate I, Figure 1, and may
be contrasted with the advanced degranulated
condition found in a seven-day implant shown
in Figure 2. Normal acidophils are shown in
Figure 3.
The degranulated basophils are found along
the periphery of the pituitary and in groups
along the region of juncture with the inter-
mediate lobe., The centers of the cells are clear
and the cells somewhat shrunken and collapsed.
Dark-staining granular material may be seen
between the cells. The basophils of the control
and non-reactive experimental fish are large and
have either very small clear areas or none. The
basophils take a very heavy and dark stain.
Atz (1953) described basophils similarly lo-
cated in the fresh water teleost, Astyanax mex-
icanus. The correlation of changes in the peri-
pheral basophils with changes in the gonads led
her to consider these cells as gonadotrophs. In
all but one case basophils were reported to have
increased in number near the time of onset of
the spawning season. In no case was there a re-
port of degranulation occurring at this time. In
the present experiment no increase in the num-
ber of basophils was noted.,
The possibility of basophilic degranulation
occurring as a result of stress and loss of ACTH
was considered. However, two fish which sur-
vived seven days showed no degranulation and
the seven fish which were killed after three days,
an even shorter time to adjust to stress, showed
little or no degranulation.
The testes of Bathygobius soporator have an
unusual amount of tissue which is not sperma-
togenic. The functional germ cell tissue of the
testis is spirally wrapped around a core of tissue
of very different nature. The core tissue has some
resemblance to interstitial tissue but the amount
seems out of proportion to the need or to the
amount of interstitial tissue found in other
teleosts. The intimate relation of the germ cell
tissue and the core tissue would indicate that the
core is a functional part of the testis but no defi-
nite function has been described.
Gonads, especially the ovaries, were stimu-
lated in the fish showing degranulation. The
stimulation caused rapid maturation of about
half the ova and a third of the spermatogonia
but little increase in over-all size of the gonad.
1957]
Stoll: Changes in Bathygobiiis separator after Pituitary Implantation
101
However, the core tissue of the testes hypertro-
phied. Stimulation was greater when the pituitary
donor had gonads in intermediate stages of
maturation than when the gonads were very ad-
vanced and approaching or in spawning condi-
tion. An ovary from an untreated juvenile and
an untreated mature fish are shown in Plate I,
Figures 4 and 5. Figure 6 shows the response
of an ovary to the experimental procedure. The
ovary from the mature untreated fish shows
more uniform graded development than that of
the treated juvenile fish where several ova are
highly developed while the rest are in a uniform
early stage.
Testicular and core tissue of an untreated
juvenile and untreated mature fish are shown
in Plate II, Figures 1 and 2. The normal condi-
tion of the core tissue of a juvenile fish stained
with Sudan Black B to show lipids is shown in
Plate II, Figure 3, and is contrasted with Figure
4 of the same magnification which shows the
hypertrophied core tissue from a fish which had
received pituitary implants. Baker’s acid hae-
matin and pyridine extraction techniques were
used on the core tissue to indicate some of the
cellular components. Staining results of the core
tissue of normal fish showed a positive reaction
to acid haematin (Plate II, Figure 5) and a neg-
ative one to pyridine extraction (Plate II,
Figure 6), indicating the presence of phospho-
lipids.
Discussion
The histology of the pituitary and the identi-
fication of cell types as sites of specific hormone
production is still to some extent controversial.
In the teleosts the transitional lobe has been
found to be homologous to the anterior lobe in
other vertebrates (Charipper, 1937).
The pituitary basophils of the rat were de-
scribed as the site of thyrotropic and gonado-
tropic hormone elaboration and storage by
Purves & Griesbach (1951). A subsequent re-
port (1954) designated the specific basophils
which produce TSH, FSH and LH.
However, there is some disagreement as to
whether the gonadotropic hormone is a single
biochemical entity or two or even more. Evans
& Simpson (1950), in a review of the literature
concerning the gonadotropins, called attention to
the fact that it is not certain that there are two
distinct gonadotropic hormones identical with
the two substances which have been isolated
from pituitary tissue or that both hormones are
necessary in both sexes for normal reproductive
function. The gonad maturity at the time of pit-
uitary injection for experimental purposes is an
important factor.
Matthews (1939), using injections of mam-
malian pituitary extract on Fundulus hetero-
clitus, found that the injections had no stimulat-
ing effect in the males and only four of 35 fe-
males responded to any degree. Hypophysectomy
caused regression of the gonads especially in the
males. On the basis of this work the pituitary
gland of a teleost was seen to exert a controlling
influence on the seasonal cycle and this influence
was considered to be . . of greater importance
in maturation than in proliferation of the germ
cells.” The present study is in agreement, as
pituitary implantation caused an acceleration in
the rate of maturation of the germ cells.
The effect of adult Fundulus pituitary im-
plants to immature fish of the same species was
reported by Matthews in 1940. Adult pituitary
implanted intraperitoneally into immature fish
at three-day intervals caused gonad stimulation
by the end of four weeks. The males were espe-
cially responsive and large numbers of mature
spermatozoa were seen. The pigmentation of
the fish was characteristic of the adult in breed-
ing season. Included in this report was a review
of the use of mammalian pituitary extracts for
injection in fish and the conflicting results and
data collected; in many cases the mammalian
preparation had no effect. A similar review of
the use of injecting fish pituitary preparations
in fish showed a fairly uniform response. All
species tried showed enlargement of the gonads
and in some cases expulsion of eggs and sperm.
All experiments involving hypophysectomy
showed gonad regression.
Burger (1941) used pituitaries from adult
Fundulus as implants to other adult male Fundu-
his which had been hypophysectomized at the
time of maximal testicular development and
which showed inhibition of the testes after hypo-
physectomy. The implants caused recrudescence
of the testes within two weeks. The implants
were made intraperitoneally and each fish re-
ceived five at a time for four times. Burger con-
cluded that the Fundulus pituitary contained
gonadotropic material which was responsible
for spermatogonial proliferation and for the ma-
turation phenomena.
Riley & Fraps (1942) investigated the gonad-
stimulating activity of anterior pituitary in the
female domestic fowl. Glands from hens with
regressed or quiescent ovaries produced a greater
stimulation of the gonads in immature mice than
glands from hens in full reproductive condition.
Greater gonad stimulation of the gobies in the
present report resulted when pituitary donors
were in intermediate stages of gonadal develop-
ment.
The cyclic changes in the pituitary of the uro-
dele amphibian (Taricha torosa) were described
by Miller & Robbins (1955). In this form the
102
Zoologica: New York Zoological Society
[42: 9
delta basophils of the pituitary increase in num-
ber and granulation in relation to spring sperma-
togenesis and oogenesis and late fall final gonad
maturation. The beta basophils were considered
to be related to increased thyroid activity.
As part of a study of light and temperature
effect on the sexual cycle of the bitterling Rho-
dens amarus, Verhoeven & van Oordt (1955)
studied the adenohypophysis of the fishes which
had an experimentally induced sexual cycle.
They briefly note that the beta cells of the gona-
dotropic zone had become more numerous and
staining response of these cells to PAS was a
deep purple coloration which in rats indicates
gonadotropic hormone. The method by which
an increase in the number of beta cells was ascer-
tained was not given.
Rasquin & Stoll (1955) described the asso-
ciation of degranulated centrally located baso-
phils with hypertrophied adrenals in the fresh-
water teleost Astyanax mexicanus after injec-
tions of pitressin. The degranulation was thought
to be a result of loss of ACTH, as the thyroid
tissue was not stimulated. The peripheral baso-
phils (suspected of gonadotropin elaboration)
did not degranulate.
The only possible explanation for the appear-
ance of degranulated basophils after pituitary
implantation in the present study is that the
basophils are not elaborating secretion granules
at the normal rate because of the excess pituitary
substance which was added., The absence of gran-
ules in cells usually crowded with them is gener-
ally interpreted as a release of hormone, but
since the fish were given more pituitary there
would appear to be no need for the pituitary of
the implanted animal to add its secretion to a
system already over-supplied. The fact that the
degranulation was so gradual— little or none
could be detected after three days of implanta-
tion-supports the idea that degranulation in this
case is a result of an inhibition of elaboration
rather than a stimulated release of secretory
products which would presumably occur more
rapidly.
Sections of the core tissue of testes of Gobius
auratus Risso were stained and found to be faint-
ly fuchsinophilic and unblackened by iron
haematoxylin (Eggert, 1931). The granules
within the cells of the tissue apparently con-
tained lipids. Included in the report is a survey
of the literature on the core tissue and a discus-
sion of the possibility that this is the interstitial
tissue. Eggert claimed that the fine vacuolation
and granulation indicated that the cells were not
interrenal. He did not find core tissue in the
testes of G. panizzae Verga, G. buchichi Stein-
dachner, G. quagga Heck or G. jozo Linnaeus.
Coujard (1941) described the core tissue in
Gobius niger and G. minutus as being made up
of cells with large nuclei and nucleoli, abundant
in protein and containing lipids. He did not
attribute a definite function to the tissue but
considered that it undoubtedly had a function
correlated with glandular development and the
nervous mechanism of spawning.
The procedure for demonstrating phospho-
lipids has been described in detail by Baker
(1946) . He used the two-stage technique of acid
haematin staining and pyridine extraction on sev-
eral tissues, including the testes of the mouse.
The secretion droplets of the interstitial cells
reacted positively to acid haematin and nega-
tively to pyridine extraction, which identified the
secretory granules as phospholipid. j
Melampy & Cavozos (1954) made a compara- i
tive study of lipids in the vertebrate testis, using ;
Sudan dyes and Baker’s acid haematin-pyridine
extraction technique. Using the Sudan dyes, a
positive reaction was obtained from the inter-
stitial cell cytoplasm, basement membrane, cyto-
plasm of Sertoli cells, spermatogonia, spermato-
cytes and spermatids of the bull, ram, boar,
guinea pig, rooster and horned lizard. The tele-
ost, the bluegill (Lepomis macrochirus), showed
a negative reaction with the Sudan dyes. The
authors suggest that the failure to get a positive
reaction from bluegill testis might be due to the
cyclic breeding season of this form and that at
the stage of the test the reactive quantity of
cholesterol in the testis was absent. Baker’s test
was not applied to the bluegill testis.
The implantation experiments reported here
show that the core tissue of the testis is respon-
sive to pituitary stimulation. The marked hyper-
trophy of the cells, the high phospholipid content
and the close proximity of the core tissue to the
spermatogenic cells seem sufficient evidence for
designating the core cells as interstitial tissue.
Summary
1. Pituitary implants from adult fish to juvenile
fish caused degranulation of the peripheral baso-
phils of the transitional lobe of the hypophysis
and gonad stimulation in receiver fish maintained
for one week after implantation.
2. Basophils of fish maintained for three days
after implantation showed little or no degranu-
lation.
3. Acidophils of the adenohypophysis were
not affected by the experimental procedure.
4. Degranulation which occurred gradually
may be the result of inhibition of hormone
elaboration rather than stimulation of release
of hormone.
5. Basophilic degranulation, when it occurred,
1957]
StoU: Changes in Bathygobius soporator after Pituitary Implantation
103
was the same in both sexes but acceleration of
maturation of the ovary was greater than that
in the testis.
6. The core tissue of the testes hypertrophied
after pituitary implantation and had a high phos-
pholipid content. On this evidence, the core
tissue is designated as interstitial tissue.
7. The degree of gonad maturity of the pitui-
tary donors was an important factor in the re-
sults. Greater stimulation resulted when donors
were in intermediate maturation stages than
when donors were in advanced stages.
Bibliography
Atz, E. H.
1953. Experimental differentiation of basophil
cell types in the transitional lobe of the
pituitary of the teleost ft^h, A sty anax mexi-
canus. Bull. Bingham Oceanogr. Coll.,
14:94-116.
Baker, J. R.
1946. The histochemical recognition of lipine.
Quart. J. Microscop. Sci., 87:441-470.
Burger, J. W.
1941. Some experiments on the effects of hy-
pophysectomy and pituitary implantations
on the male Fundiilus heteroclitus. Biol.
Bull., 80:31-36.
Charipper, H. a.
1937. The morphology of the hypophysis in
lower vertebrates, particularly fish and
amphibia, with some notes on the cytology
of the pituitary of Incarassius (sic) auratiis
(the goldfish) and Nectiirus maculosus
(the mudpuppy). Cold Spring Harbor
Symposia on Quantitative Biology, vol. 5,
Cold Spring Harbor, New York, pp. 151-
164.
COUJARD, R.
1941. Sur I’existence d’une glande testiculaire et
d’une glande genitale annexe chez les
gobies. C. R. Soc. Biol., 135:570-574.
Eggert, B.
1931. Die Geschlechtsorgane der Gobiiformes
und Bleniiformes. Zeit. f. wissen. Zool.,
139:249-558.
Evans, H. M., & M. E. Simpson
1950. Physiology of the gonadotrophins. The
Hormones, Vol. II. Ed. Pincus and Thi-
man, pp. 351-404.
Matthews, S. A.
1939. The relationship between the pituitary
gland and the gonads in Fundulus. Biol.
Bull., 76:241-250.
1940. The effects of implanting adult hypophysis
into sexually immature Fundulus. Biol.
Bull., 19:101 -lU.
Melampy, R. M., & L. F. Cavazos
1954. Comparative study of lipids in vertebrate
testes. Proc. Soc. Exp. Biol. & Med., 87:
297-303.
Miller, M. R., & M. E. Robbins
1955. Cyclic changes in the pituitary gland of
the urodele amphibian, Taricha torosa
(Triturus torosus). Anat. Rec., 122:105-
114.
Purves, H. D., & W. E. Griesbach
1951. Pituitary basophils as the source of the
thyrotrophic and gonadotrophic hormone.
Proc. U. Otago Med. Sch., 29:3-4.
1954. The site of follicle stimulating and lutein-
ising hormone production in the rat pitui-
tary. Endocrinology, 55:785-793.
Rasquin, P., & L. M. Stoll
1955. Effects of pitressin and water injections on
the secretions of brain and hypophysis in
a teleost. Anat. Rec., 122:452-453.
Riley, G. M., & R. M. Fraps
1942. Relationship of gonad-stimulating activity
of female domestic fowl anterior pitui-
taries to reproductive condition. Endocrin-
ology, 30:537-541.
Scruggs, W. M.
1951. The epithelial components and their sea-
sonal changes in the pituitary gland of the
carp (Cyprinus carpio, L.) and goldfish
(Carassius auratus, L.). J. Morph., 88:
441-469.
Sokol, H. W.
1953. Selective staining of the pituitary gland of
teleosts demonstrating at least six tinctorial
cell types in the adenohypophysis. Anat.
Rec., 117:582 and personal communica-
tion.
Tavolga, W. N.
1954. Reproductive behavior in the gobiid fish,
Bathygobius soporator. Bull. Am. Mus.
Nat. Hist., 104:427-460.
1955. Effects of gonadectomy and hypophysec-
tomy on prespawning behavior in males
of tire gobiid fish, Bathygobius soporator.
Physiol. Zool., 28:218-233.
Verhoeven, B., & G. j. van Oordt
1955. The influence of light and temperature on
the sexual cycle of the bitterling, Rhodeus
amarus (preliminary communication). Ko-
nink. Neder. Akad. von Wetenschappen.
Proc. Series C, 58:628-634.
Vivien, J. H.
1941. Contribution a I’etude de la physiologie
hypophysaire dans ses relations avec I’ap-
pareil genital, la thyro'ide et les corps
suprarenaux chez les poissons, selaciens et
teleosteens. Scylliorhinus canicula et
Gobius paganellus. Bull. Biol. France et
Belg., 75:257-309.
104
Zoologica: New York Zoological Society
[42: 9: 1957]
EXPLANATION OF THE PLATES
Plate I
Fig. 1. Partially degranulated basophils in a 41
mm. fish three days after pituitary im-
plantation. Paraldehyde fuchsin, sagittal
section, 1500 X.
Fig. 2. Peripherally located degranulated baso-
phils from a 40 mm. fish killed one week
after pituitary implantation. Paraldehyde
fuchsin, sagittal section, 1500X.
Fig. 3. Normal acidophils and basophils from 43
mm. control fish, paraldehyde fuchsin,
sagittal section, 1500X.
Fig. 4. Ovary of 55 mm. untreated juvenile.
Haematoxylin and eosin, cross section,
lOOX.
Fig. 5. Ovary of 73 mm. untreated adult. Haema-
toxylin and eosin, cross section, 100 X.
Fig. 6. Stimulated ovary from 42 mm. fish after
one week of pituitary implantation.
Haematoxylin and eosin, cross section,
lOOX.
Plate II
Fig. 1. Core and spermatogenic tissue of 41 mm.
untreated juvenile. Haematoxylin and
eosin, cross section, lOOX.
Fig. 2. Core and spermatogenic tissue of 80 mm.
untreated adult. Haematoxylin and eosin,
cross section, lOOX.
Fig. 3. Core tissue of 41 mm. untreated juvenile
stained with Sudan Black B, cross section,
500X.
Fig. 4. Core tissue of 35 mm. juvenile after one
week of implantation. Sudan Black B,
cross section, 500 X.
Fig. 5. Core tissue of 81 mm. adult stained with
acid haematin, cross section, 1500X.
Fig. 6. Core tissue of 74 mm. adult after pyridine
extraction, cross section, 1500X.
STOLL
PLATE I
FIG. 2
FIG. 3
•
CO ,
s> ’«s»*
FIG. 4
FIG. 1
FIG. 5 FIG. 6
CHANGES IN THE CYTOLOGICAL STRUCTURE OF THE ADENOHYPOPHYSIS AND GONADS IN JUVENILE
BATHYGOBIUS SOPORATOR AFTER PITUITARY IMPLANTATION
STOLL
PLATE II
FIG. 3
FIG. 4
• #
A
'V'
FIG. 5 FIG 6
CHANGES IN THE CYTOLOGICAL STRUCTURE OF THE ADENOHYPOPHYSIS AND GONADS IN JUVENILE
BATHYGOBIUS SOPORATOR AFTER PITUITARY IMPLANTATION
10
The Ctenuchidae (Moths) of Trinidad, B. W. I.
Part I. Euchromiinae/
Henry Fleming
Department of Tropical Research,
New York Zoological Society, New York 60, N. Y.
(Plates I-III)
[This paper is one of a series emanating from the
tropical Field Station of the New York Zoological
Society, at Simla, Arima Valley, Trinidad, British
West Indies. This station was founded in 1950 by
the Zoological Society’s Department of Tropical
Research, under the direction of Dr. William Beebe.
It comprises 200 acres in the middle of the Northern
Range, which includes large stretches of undisturbed
government forest reserves. The laboratory of the
station is intended for research in tropical ecology
and in animal behavior. The altitude of the research
area is 500 to 1,800 feet, with an annual rainfall of
more than 100 inches.
[For further ecological details of meteorology
and biotic zones see “Introduction to the Ecology of
the Arima Valley, Trinidad, B.W.I.,” William Beebe.
(Zoologica, 1952, Vol. 37, No. 13, pp. 157-184).].
Contents Page
Introduction 105
Euchromiinae 106
Pseudosphex 106
Pleurosoma 108
Sphecops 108
Homoeocera 108
Isanthrene 108
Phoenicoprocta 109
Loxophlebia 113
Mesothen 114
Pheia 115
Chroslosoma 116
Leucotmemis 116
Nyridela ...116
Cosmosoma 117
Dixophlebia 118
Pseudomya 118
Rhynchopyga 119
Saurita 119
^Contribution No. 978, Department of Tropical Re-
search, New York Zoological Society.
Psoloptera 122
Dycladia 122
Syntomeida 122
Histiaea 123
Macrocneme 123
Calonotos 127
Introduction
This paper concerns the species of moths
belonging to the subfamily Euchromiinae
that have been reported in the literature or
collected by the Department of Tropical Re-
search of the New York Zoological Society at its
biological station at Simla, Arima Valley, Trini-
dad, B.W.I. The subfamily Ctenuchinae will be
treated in a subsequent paper which will also
include keys to all the genera of Ctenuchidae
found in Trinidad, as well as the bibliography.
I have followed Travassos (1935: 437-451)
in the selection of the family name for this group
of moths. Ctenuchidae has priority over other
names in use as follows;
Ctenuchidae, Kirby, 1837.
Syntomidae, Snellen, 1867.
Euchromiidae, Neumoegen & Dyar, 1893.
Amatidae, Jansen, 1917.
While the type genus of Syntomidae, Syntomis,
has been found to be a junior synonym of A mata,
the emendation of the XIVth International Con-
gress of Zoology at Copenhagen, 1953, states
that a family group taxon based on a synon-
omized genus is not to be changed (Copenhagen
Decisions on Zoological Nomenclature: 36, par.
54 (1) (a), 1953). The Syntominae are limited
to the Old World and do not concern us in this
paper.
105
106
Zoologica: New York Zoological Society
[42: 10
No attempt has been made to make the refer-
ences under the species complete. References
to the original description, pertinent or new
synonomy, colored figures, helpful descriptions
to the species or a specific reference to Trinidad
have been cited, however.
Three publications cited whenever possible
are:
Hampson, G. F.: Catalogue of the Lepidop-
tera Phalaenae, I, Syntomidae, 1898; Sup-
plement I, Amatidae, 1914.
Seitz, A.: Macrolepidoptera of the World,
VI, Euchromiidae, 1915 and 1917.
Kaye, W. J. & N. Lamont: A Catalogue of
the Trinidad Lepidoptera Heterocera. Mem.
Dept. Agric. Trinidad and Tobago, No.
3, 1927.
This paper includes keys to the species of
moths found in Trinidad and photographs of the
species collected at Simla, Arima Valley, in an
attempt to make it useful to biologists working
on ctenuchids in Trinidad. Five new species of
Euchromiinae are described.
The species of Ctenuchidae of Trinidad are
continental rather than Caribbean. Trinidad and
the adjacent island of Tobago are geologically
part of South America rather than of the West
Indies, and the faunistic and floristic character
of the two islands is decidedly related to Guiana
and Venezuela.
Our own collecting almost without exception
has been confined to the Arima Valley in the
Northern Range of Trinidad. However, Kaye
& Lamont (1924) listed the species known to
occur on the island as a whole and Lamont &
Callan (1950) added two species to the
Euchromiinae. The species reported by these
authors are included in this paper. There are
extremely few records from the southern part
of the island and additional species may be ex-
pected when this region has been investigated.
My thanks go to Miss Rosemary Kenedy,
who made notes and took photographs of many
of the holotypes of the ctenuchid species in the
British Museum (Natural History) which aided
in the determination of some of the species in
question. She also collected the majority of the
department’s ctenuchid collection. Thanks go
also to Dr. William Beebe and Miss Jocelyn
Crane for their part in assembling the collec-
tion and for advice and criticism.
Euchromiinae
The absence of vein Sc in the hindwing sepa-
rates the Euchromiinae from other families of
Trinidad moths. In this subfamily vein M2 of
the hindwing is rudimentary or absent but often
represented by a vein-like line of scales, whereas
in the subsequent subfamily, Ctenuchinae, vein
M2 of the hmdwing is present. Veins Cui and
Cu2 are stalked or united in the Euchromiinae,
but in the Ctenuchinae, with the exception of
the Horama and related genera, they are widely
separated.
PsEUDOSPHEX Hubner
This and the following two genera are wasp-
like in appearance, with constricted abdomens.
Pseudosphex kenedyae, new species
(PI. I, fig. 1; PI. II, fig. 1)
Pseudosphex melanogen Beebe, not Dyar, 1955;
32, fig. 6.
Length of forewing 1 1 mm.
Closely related to Pseudosphex laticincta
Hampson. P. kenedyae differs from P. laticincta
principally in lacking the dorsal band on the
first abdominal segment and on the lateral bullae.
Ma/e.— Antennae bipectinate. Processes on
first segment of antennae small but gradually
increasmg in length to the seventh segment
where it reaches a length of .142 mm. The pro-
cesses are but slightly dilated, pubescent anteri-
orly and with a bristle at the distal end. Near
the distal third of the antennae the pectinate
processes decrease in length and resolve into
three dentate segments followed by 14 serrate
segments. The serrate segments are blackish-
brown and the remainder of the antennae red-
dish-brown.
Palpi concolorous yellow with a fan-shaped
tuft of hairs on the basal palpal segment yellow
interspersed with black hairs. Front yellow but
in rubbed specimens may have a whitish cast.
A large black spot on vertex of head surrounded
by yelTow. Side of head behind eyes yeUow with
occasional black hairs or scales.
Patagia yellow with a broad black bar behind
middle. Tegulae yellow edged with black; broad-
ly edged with black at the costal margin of the
fore wing where it projects slightly on the sub-
costal and cubital veins. Mesothorax (scutum)
black with two diagonal yellow stripes and a
mid-dorsal line of yellow scales broadening cau-
dally. Hind edge of scutum with yellow scales.
Metathorax yellow.
Legs yellow. The inner face of the forecoxae
shining white. The distal edge with a few scat-
tered yellow scales. Inner face of forefemur
shining white but somewhat variable in width
in different lights, with the cephalad edge yellow.
Frequently on the outer edge of the distal tip
of the femur of the meso- and metathoracic
legs a very small patch of shining white scales.
In different lights the yellow color of the
thoracic structures may take on an orange-yel-
low cast.
1957]
Fleming: Ctenuchidae (Moths) of Trinidad, B.W.I.
107
Forewing hyaline. Costal area, area beyond
discal cell, part of discal cell above discal fold
and area below vein 2dA fuscus brown. Re-
mainder of wing with scattered setae.
Hindwing hyaline with scattered setae except
the costal area above the cell and vein Rg which
is fuscus, similar in color to the respective area
of the forewing.
Dorsum of first abdominal segment and bul-
lae yellow with the anterior margin finely edged
with black. Second abdominal segment (pedun-
cle) translucent yellow. The subsequent caudal
segment with the anterior half tan brown and
the posterior half translucent yellow. The mid-
dorsum of this segment may have a slight mid-
dorsal line of darker scales. Fourth, fifth and
sixth segments of the abdomen with the two-
thirds of the cephalad portion of the segment
blackish-brown and the caudal third translucent
yellow. Seventh abdominal segment dark and
yellow area approximately equal. Last segment
with only the cephalad quarter of the segment
blackish-brown. Terminal tuft of the abdomen
yellow.
Ventrum of the abdomen with the anterior
part of the valve (peduncle) relatively unsealed,
usually gray but sometimes dark brown., Often
with scattered occasional yellow scales. Approxi-
mately at the point where the caudal lateral
edge of the dorsal sclerite of the peduncle seg-
ment touches the ventral vein, the valve be-
comes translucent yellow. The valve is fringed
on its caudal edge with white scales approxi-
mately a fifth of the length of the yellow part
of the valve. Remainder of the ventrum yellow
peppered with black scales.
Female similar to the male except wing ex-
panse greater (12 mm.), and the basal and
medial part of the antennae prismatic. The last
12 distal antennal segments are slightly serrate,
dark grayish-brown in our single female and
the three preceding segments light brown and
dentate rather than prismatic. The remaining
segments of the antennae are warm reddish-
brown. The legs are concolorous yellow with
no white areas.
Closely related to P. laticincta from which it
differs by having a concolorous first abdominal
segment and bullae whereas in P. laticincta the
first abdominal segment has a dorsal stripe and
the bullae are banded.
Rosemary Kenedy compared part of our
series with Hampson’s type of P. laticincta in
the British Museum (Natural History) and
noted that in the male of P. kenedyae the an-
tennal shaft is darker than the type of P. lati-
cincta so that the basal darkening is not so
apparent. P. kenedyae has a longer and more
restricted antennal extremity preceded by a con-
trasting lighter area and the pectinations slightly
longer and not so dilated. The light brown ab-
dominal band on the second segment of P.
kenedyae is a black band in P. laticincta and
the same color as the subsequent black abdomi-
nal bands. The dark abdominal bands of P.
laticincta are narrower and extend to slight
points behind mid-dorsally, more so than in
P. kenedyae. In P. kenedyae only the fifth and
sixth abdominal segments consistently have pro-
nounced black points.
It is also important to note from Miss Ken-
edy’s memoranda that the palpi of P. laticincta
are similar to P. kenedyae. The palpi are yellow
with a fan-shaped process from the basal palpal
segment with yellow scales interspersed with
black scales. In other words, the palpi are not
banded in the usual meaning of the term in P.
laticincta, as the original description would lead
one to beheve.
P. kenedyae difi'ers from P. melanogen Dyar
in having more yellow on the thorax and much
wider yellow bands on the abdomen. The same
considerations are true for P. deceptans Zerny
which was synonymized under P. melanogen by
Hampson. Neither P. melanogen nor P. decep-
tans are described as having white forecoxae.
For purposes of Trinidad identification, the
broad fuscus costal margin of the forewing wUl
separate this species from the species in the fol-
lowing two genera.
I take pleasure in naming this species for Miss
Rosemary Kenedy, Research Assistant of the
Department of Tropical Research, whose initia-
tive in the use of Heliotropium as an attractant
for euchromids brought to light this new species.
Material— AW of the types were taken at Simla,
Arima Valley, Trinidad, on Heliotropium indi-
cum. Holotype, male. Catalog No. 5639,
20-11; allotype, female, (5640) 29-IV; 15 para-
types, males, (5641) 14-1, (5642) 15-1, (5643)
20-1, (5644) 28-1, (5645) lO-II, (5646) 20-11,
(5647) 22-11, (5648) 23-11, (5649) (5650)
(5651) 4-IV, (5652) 29-IV, (5653) 1-V,
(5654) 27-XII,(5655) 31-XII.
Disposition of type material.— The. Depart-
ment of Tropical Research, New York Zoolog-
ical Society, will retain three paratypes. Catalog
Nos. 5650, 5651 and 5655. Paratypes with Cata-
log Nos. 5648 and 5653 are in the British Mu-
seum (Natural History) collection and paratypes
with Catalog Nos. 5644 and 5652 are in the
United States National Museum collections. The
holotype, allotype and remaining paratypes are
in the American Museum of Natural History,
New York.
108
Zoologica: New York Zoological Society
[42: 10
Pleurosoma Orfila
This and the following genus may be distin-
guished from the preceding genus, Pseudosphex,
by vein Rs of the forewing arising basad of vein
R.3.
The genus Astridia Kiriakoff, 1948: 267, is
an absolute synonym of Pleurosoma Orfila,
1935: 178.
Both genera were erected with Sphecosoma
angustatum Moschler as the type.
Pleurosoma trinitatis (Rothschild)
(PI. Ill, fig. 1)
Sphecosoma /rmhatw Rothschild, 1911: 24.
Sphecosoma trinitatis, Rothschild, 1913: 471, pi.
XIV, fig. 5.
Sphecosoma trinitatis, Hampson, 1914: 90.
Sphecosomatrinitatis,Y>XdL\id.i,\9l5'. 40,fig. lOh.
Sphecosoma trinitatis, Kaye&Lamont, 1927: 1.
Pleurosoma trinitatis, Orfila, 1935: 178.
Described by Rothschild from four male
specimens collected at Caparo, Trinidad, in De-
cember, 1905, by S. M. Klages.
The figures in Rothschild and Seitz are mis-
leading, for they indicate that the color of the
abdominal segment following the peduncle is
black, similar to the subsequent segments, rather
than rufous brown. The length of the forewing
in our specimens is 12 mm. The length of the
forewing of the holotype in the British Museum
is 12 mm., rather than the 14 mm. given by
Rothschild.
Female similar to male except antennae pris-
matic rather than bipectinate.
Material— VitiQen males and eight females.
Rfl/rge.— Trinidad.
Sphecops Orfila
Differs from Pleurosoma in lacking the two
thoracic bladder-like processes beneath the base
of the first abdominal segment. First segment of
abdomen much shorter in Sphecops than Pleuro-
soma. The males of Sphecops have a ventral
valve covering the second and third abdominal
segments which is absent in Pleurosoma.
Sphecops aurantiipes (Rothschild)
(PI. Ill, fig. 2)
Sphecosoma aurantiipes Rothschild, 1911: 25.
Sphecosoma aurantiipes, Rothschild, 1913: 471,
pi. XIV, fig. 7.
Sphecosoma aurantiipes, Hampson, 1914: 91.
Sphecosoma aurantiipes, T>Tavidt, 1915: 40, fig.
lOh.
Sphecops aurantiipes, Or^Xa, 1935: 181.
Four of our Trinidad specimens have been
compared with Rothschild’s type from San Es-
teban, Venezuela, in the British Museum by Miss
Rosemary Kenedy. Rothschild gives the length
of the forewing as 14 mm. in his original descrip-
tion, whereas our measurement of the type is
12.5 mm. The Trinidad specimens have a fore-
wing length of 10 to 11 mm. In this respect they
resemble material that Rothschild had before
him from Paraguay and Bolivia which he re-
ported as having a wing length of 10 mm. The
yellow bands on the abdomen of the Trinidad
specimens are narrower than in the type from
Venezuela.
Material.— Seven males.
Range.— Venezuela, Paraguay and Bolivia. A
new record for Trinidad.
Homoeocera Felder
A genus related to Gymnelia, consisting of
large, heavy-bodied species. We have not taken
any species of this genus in the Arima valley.
Homoeocera magnolimbata Dognin
Homoeocera magnolimbata Dognin, 1911 : 11:3.
Homoeocera magnolimbata, Hampson, 1914:
1:95, pi. V, fig. 1.
Homoeocera magnolimbata, Draudt in Seitz,
1915: VI:46, pi. 27b.
Kaye & Lamont (1927) report this species
from Trinidad. One specimen appears to have
been taken in Trinidad by F. W. Urich at St.
Ann’s in October, 1899. This specimen is not in
the Urich Insect Collection at the Victoria Mu-
seum in Port-of-Spain. It is a large insect, ap-
proximately two inches in expanse, with hyaline
wings and black abdomen with conspicuous
white spots and some metallic blue.
Kenedy reports another specimen from Trini-
dad in the British Museum (Natural History) in
the Joicey Collection, collected in 1909. It was
part of a series from French Guiana, Amazons,
East Bolivia and Peru.
Isanthrene Hubner
The thorax of the species in this genus is
smoothly scaled in contrast to the species of
Homoeocera which have hairy thoraxes.
Isanthrene tryhanei Rothschild
Isanthrene try/zanei Rothschild, 1911: XVIII:
26.
Isanthrene tryhanei, Rothschild, 1913: XX:
471, pi. XIV, fig. 24.
Isanthrene tryhanei, Hampson, 1914: I: 98.
Isanthrene tryhanei, Draudt in Seitz, 1915: VI:
47, pi. 10b.
1957]
Fleming: Ctenuchidae (Moths) of Trinidad, B.W.I.
109
Rothschild described this species from St.
Aim’s, Trinidad, from one female. It does not
appear to have been collected since the original
specimen was taken. It is another large species
approximately two inches in expanse, with hya-
line wings. The wings have a yellowish cast
particularly along the costal and inner margin,
in contrast to Homoeocera magnolimbata.
Phoenicoprocta Druce (not Hampson)
Stephens (1850) used the name Hyela for a
generic name in Noctuidae. Walker (1854) used
Hyela for one of his groups in the genus Glau-
copis with the intention that the species he
placed under Glaucopis would henceforth take
the various group names as generic names.
Hampson (1898) realized that Walker had em-
ployed a name, Hyela, that was preoccupied by
Stephens, and erected a new name Phoenico-
procta. Druce (1898) was apparently cognizant
of Hampson’s monograph of the Ctenuchidae
and described a presumed new species, Phoeni-
coprocta metachrysea. This species employing
Hampson’s new name was published in May,
1898, in advance of Hampson’s monograph,
which was not published until September, 1898.
Thus Druce must be credited with the generic
name Phoenicoprocta. The type of the genus is
Phoenicoprocta metachrysea by monotypy
which Dyar (1915) synonymized under
coprocta vacillans Walker (1856) on the basis
of breeding carried out by H. W. B. Moore in
British Guiana.
The males of this genus are easily distin-
guished in Trinidad by the blue or red spotted
abdomen and carmine anal tufts. The Aethria
species with a carmine tuft have a concolorous
blackish abdominal dorsum. The females have a
Calonotus-\\ke abdomen and may be distin-
guished from Calonotus by vein R2 in Phoeni-
coprocta being forked with veins R3+1+5 rather
than from the cell as in Calonotus.
Phoenicoprocta vacillans (Walker)
PI. Ill, figs. 3, 4, 5
Eunomia vacillans Walker, 1856: VII: 1617
(male).
Phoenicoprocta vacillans, Hampson, 1898: I:
197, pi. VII, fig. 11 (male).
Phoenicoprocta vacillans, Draudt in Seitz, 1915 :
VI: 47, pi. 12a (male). 1917: Supp.: 199.
Phoenicoprocta metachrysea Druce, 1898: (7)
I: 404 (male).
Phoenicoprocta metachrysea, Hampson, 1898:
I: 196, pi. VII, fig. 12 (male).
Phoenicoprocta metachrysea, Draudt in Seitz,
1915: VI: 47, pi. 12a; (male). 1917: Supp.:
199.
Phoenicoprocta chrysorrhoea Hampson, 1898:
I: 196 (male).
Phoenicoprocta chrysorrhea, Draudt in Seitz,
1915; VI: 57, pi. 12a (male). 1917: Supp.:
199.
Phoenicoprocta trinitatis Strand, 1915: 21
(male).
Phoenicoprocta trinitatis, Draudt in Seitz, 1915 :
VI: 58, pi. 27f (male).
Phoenicoprocta nigropeltata Strand, 1915: 22
(male).
Leucotmemis albigutta Schaus, 1905; XXIX,
No. 1420: 185 (female).
Leucotmemis albigutta, Hampson, 1914: Supp.
1 : 1 5 1 , pi. VIII, fig. 2 (female) .
Leucotmemis albigutta, Draudt in Seitz, 1915:
VI: 70, pi. 27k; 1917; Supp.: 201 (female).
Leucotmemis thoracica Schaus, 1905: XXIX,
No. 1420: 186 (female).
Leucotmemis thoracica, Hampson, 1914: Supp.:
I: 150, pi. VIII, fig. 1 (female).
Leucotmemis thoracica, Draudt in Seitz, 1915:
VI: 70, pi. 27k; 1917: Supp.: 201 (female).
Antichloris trinitatis Rothschild, New Syntomi-
dae. 1912: XIX: 154 (female).
Autochloris trinitatis, Hampson, 1914: Supp. 1:
104, pi. V, fig. 17 (female).
Antichloris trinitatis, Draudt in Seitz, 1915: VI;
136 (female).
Autochloris trinitatis, Draudt in Seitz, 1917:
VI: Supp.: 197, pi. 27c (female).
Paramya chrysonota Hampson, 1898, I: 165,
pi. VII, fig. 3 (female).
Paramya chrysonota, Draudt in Seitz, 1915 : VI :
44, pi. 10k (female).
Calonotus hoffmannsi Rothschild, 1911 : XVIII;
30 (female).
Calonotus hoffmannsi, Rothschild, Some unfig-
ured Syntomidae, 1913: XX: 470, pi. XIII,
fig. 35 (female).
Leucotmemis hoffmannsi, Hampson, 1914:
Supp. I: 150 (female).
Paramya? hoffmannsi, Draudt in Seitz, 1915:
VI: 44 (female).
Calonotus hoffmannsi, Draudt in Seitz, 1915:
VI: 108, pi. 17k; 1917: Supp.: 201 (female).
Antichloris trinitatis, Kaye & Lamont, 1927 ; No.
3; 9.
Phoenicoprocta trinitatis, Kaye & Lamont, ibid:
1927; 1.
Phoenicoprocta rubiventer? Kaye & Lamont,
1927: 1.
Mydropastea chrysonota, Kaye & Lamont,
1927: 1.
110
Zoologica: New York Zoological Society
[42: 10
The extreme variability of this species is indi-
cated by the above synonymy, which undoubt-
edly is incomplete. Species designation has been
based on the color of the collar, shoulder-covers,
thorax, legs, abdomen and the ventral valve at
the base of the abdomen in the males; and in the
females, aside from the fact that they have been
placed in the wrong genera, largely on the
amount of hyaline areas in the wings.
In all of our specimens of Phoenicoprocta
from Trinidad, British Guiana and Venezuela,
and specimens loaned to us by the American
Museum of Natural History from Mexico, Brit-
ish Guiana, Brazil and Peru, there is a small
accessory cell in the forewing formed by a sec-
torial cross-vein between vein R2 and vein
R3+4+5. This sectorial cross-vein emerges from
R3+4+5 typically at the same point at which vein
Rs forks but may vary to the extent of being
one millimeter more basad. Hampson (1898, p.
196) makes no mention of this cross-vein, which
appears to be a good generic character. In addi-
tion, he states that vein 3 (vein Cui) of the
hindwing is absent whereas it is invariably pres-
ent though shortly stalked near the margin of
the wing.
Furthermore, the genus is not divisible into
two sections on Hampson’s characters. In the
species that he places in the first section, the
discocellulars in the hindwing are not oblique
throughout while in the forewing, vein 3 (vein
Cui) is variable in a series from the same local-
ity.
The commonest form at Simla, Trinidad, is
nigropeltata Strand which was described from
Trinidad. The character distinguishing this form
from the typical vacillans is a black ventral valve
with a white posterior edge, rather than a white
valve with a black base and lateral edges. We
have fifteen specimens that agree with this diag-
nosis of nigropeltata, but in four additional
specimens the white edge is replaced by pink.
Hampson (1898, p. 197), in his diagnosis of
vacillans, stated “forecoxae white and crimson”
(forecoxae white, red inside, as Kenedy noted
on holotype). Eight of our 19 specimens agree
in this respect, although more pink than red,
and the remaining 1 1 specimens have the coxae
white in front and blackish-brown inside. Four
of these latter specimens have the pink-edged
valve.
Ten additional specimens appear to be trini-
tatis Strand. This species is described as having
short palpi. In our specimens the palpi appear
to be the usual length. The origin of vein Cui
in the forewing is variable. The amount, posi-
tion and presence of blue, crimson and black
on the collar are extremely variable, encompass-
ing trinitatis, sanguinea and new forms. The
colors of the patagia and tegulae vary consider-
ably in different lights because of their iridescent
cast alone. The subdorsal macular stripes on the
abdomen also vary considerably; sometimes they
are large and distinct and in others, particularly
near the base of the abdomen, they may be quite
small. The ventrum of the abdomen in these
specimens varies between black and blackish-
brown. The ventral valve is light to dark pink
and in one specimen with a considerable mix-
ture of black scales. The posterior edge of the
valve is finely white. The forecoxae in these
ten specimens are white. The inside of the fore-
coxae and the remaining coxae usually red but
sometimes blackish-brown.
In an additional specimen, an eleventh speci-
men, the shoulder covers are dark brown, fore-
coxae white and remaining coxae blackish-
brown, the last four abdominal segments with a
macular red subdorsal line and the ventral valve
black with a white edge. This specimen is an
unnamed, more melanotic, form of the preced-
ing ten specimens.
Two specimens are very similar to sanguinea
Walker. Our specimens differ in having a black
transverse line at the base of the anal tuft and
a narrower discocellular bar in the forewing.
In the forewings of sanguinea Walker there is
a small hyaline spot above vein R5 which is
definitely absent in all of our specimens. In
Draudt’s figure in Seitz (fig. 15a; 1915) the
shoulder covers and the abdomen are reddish-
brown rather than the correct color, crimson.
Hampson (p. 198, 1898) gives the type locality
as Honduras. Walker (1854, p. 172) in his origi-
nal description did not state the origin of his
specimen. The holotype in the British Museum
(Natural History) has a hand-printed label
“Honduras” in neither Walker’s nor Hampson’s
writing.
The sexes in Phoenicoprocta vacillans are
dimorphic. Besides, the amount of scaling in
the wings of the females in Trinidad is extremely
variable.
The female specimens will run in Hampson’s
generic key (1898) to Mydropastea (Phaeo) or
Paramya (Methysia) since Hampson’s dicho-
tomous couplet (1898: 23 (B, a^), 24 (b^)) is
based on the presence of abdominal tufts which
are a male character. The abdominal aspect of
the females differs radically from that of the
males, which have a typically maculated series of
red or iridescent blue spots whereas the females
have blackish and iridescent blue longitudinal
stripes. The abdomens of both males and females
are somewhat bulbous caudad of the third or
fourth abdominal segments.
1957]
Fleming: Ctenuchidae (Moths) of Trinidad, B.W.I.
Ill
The abdomens of all the female specimens are
similar in showing a mid-dorsal stripe of irides-
cent blue or blue-green scales. This line is con-
tinued on the metathorax, which has a single
mid-dorsal spot of the same color. The mid-
dorsal stripe of the abdomen is bordered by a
blackish-brown stripe on each side commencing
on the first abdominal segment and followed
laterally by another stripe of iridescent blue or
blue-green which is replaced on the first ab-
dominal segments by iridescent spots on a black-
ish-brown ground. Laterally the last subdorsal
iridescent stripe is bordered by a fine blackish-
brown line. In old or rubbed specimens the iri-
descence of the abdomen may have a coppery
cast. The ventrum of the abdomen is dark brown
to brownish-black and iridescent in some lights.
There are iridescent blue points beneath the
wings on the thorax and a line of iridescent blue
on the caudal edge of the metathorax continuous
from each side. The amount of carmine on the
under side of the coxae is variable but is present
in all female specimens on the distal end of
the prothoracic coxae and frequently on the
whole underside of the forecoxae but usually
only at the distal apex of the meso- and meta-
thoracic coxae. The anterior surface of the
prothoracic coxae is marked with iridescent blue
on a brown background, most distinct on the
outer edge of the coxae.
Female Form /.—Two specimens. Both the
fore and hindwings of this form are immaculate
blackish-brown with no hyaline spots whatso-
ever. Some blue scales along the radius vein
of the forewing. Kenedy compared this form
with the holotype of Antichloris trinitatis Roth-
schild in the British Museum (Natural History)
and believes them to be identical.
Female Form //.—This form differs from
Form I in having a short elliptical hyaline spot
below the middle of the cell and hyaline spots
below the cell on each side of vein Ms in the
forewing. The hindwing has a hyaline spot below
the cell, a small spot at the forking of vein
Cui+2 and vein M,s and a large spot above vein
Ms to vein Mi. This last mentioned hyaline spot
decreases in size approximately one-third above
the line of scales representing vein Ms. One of
the three specimens has crimson patagia. The ab-
dominal and wing patterns of this form are simi-
lar to the figure in Draudt in Seitz (pi. 101;
1915) of Mydropastea chrysonota Hampson.
Female Form ///.—Differs from Form II in
having two or three small hyaline spots above
the larger hyaline spots on each side of vein Ms
of the forewing. The spot anterior to the large
spot in cell Ms, if present, is minute. The hyaline
area beneath the discal cell is larger and may
extend slightly below the anal vein. A slight
hyaline streak within the discal cell along the
cubital vein. In the hindwing the hyaline spot
below the cell is larger than in Form II and may
extend slightly below the anal vein.
Female Form /F.— Three specimens were
taken of this form. The hyaline areas in the
forewing are progressively more extensive by
the addition of a hyaline spot in the forewing
between veins Cui and Cus, a large hyaline
streak below the anal vein and the posterior por-
tion of the discal cell. In the hindwing the hyaline
area extends below the anal vein. None of the
above four female forms is to be interpreted as
being stable, as they grade into one another.
Paramya chrysonota Hampson is a synonym
of Phoenicoprocta vacillans Walker. One speci-
men of chrysonota was collected at Itaituba,
Amazons, Brazil, and described as a male.
Kenedy inspected the holotype of chrysonota in
the British Museum (Natural History) and dis-
covered that, in point of fact, it is a female
rather than a male. It agrees with our female
series in Form II from Simla, Trinidad.
Some discussion of the past history of Par-
amya chrysonota would not be out of place.
Section I of Paramya Hampson (p. 164; 1898)
establishes with Paramya intersecta Hampson
(p. 164; 1898) as the genus type is synonymized
by Hampson (p. 384; 1914) under Methysia
Butler with the genus type species Methysia
(Glaucopis) notabilis Walker (1854) on the
basis that Hampson’s figure (fig. 205, 1898) of
Methysia notabilis is correct. Thus Methysia
contained the following species: notabilis Walk-
er, intersecta Hampson, senetus Schaus, picta
Druce and melanota Hampson. This left brice-
noi Rothschild, flavia Schaus, picta Druce and
chrysonota Hampson with no proper generic
name as these species were in Hampson’s section
II of the genus Paramya.
However, Hampson was not the legitimate
author of Paramya nor was he privileged to
select a type species, as Druce in May of the
same year (1898) and also, for that matter,
Schaus in June had each described new species
employing the generic name Paramya. Hamp-
son in his Cat. Lep. Phal. I, published in Sep-
tember, included both Druce’s and Schaus’s new
species in the second section of his presumed
new genus. Neither Druce nor Schaus wrote
generic descriptions and what apparently hap-
pened was that both were familiar with the new
genus that Hampson was to erect in his mono-
graph and described their new species in that
genus. Unfortunately their descriptions were
published a few months previous to Hampson’s
monograph. The type species of Paramya is
Paramya picta Druce.
112
Zoologica: New York Zoological Society
[42: 10
However, Paramya was preoccupied by Con-
rad (1860) in Mollusca and, consequently was
unavailable for Druce’s species. Travassos
(1946) erected a new name, Metamya, for
Paramya Druce. The genotype is Metamya picta
Druce, and Paramya flavia Schaus and Ichoria
bricenoi Rothschild are congeneric. Paramya
chrysonota Hampson (1898), with which we
are concerned, is not congeneric with either
section of Hampson’s original conception of the
genus and is a female Phoenicoprocta.
Kaye & Lamont (1927) placed Paramya
chrysonota Hampson (1898) in the genus
Mydropastea Hampson (1898). I presume they
did this because Paramya as conceived by Hamp-
son has veins Rg and M, of the hindwing long
stalked. Apparently the specimens that Kaye &
Lamont possessed from Trinidad resembled our
specimens from Trinidad in which veins Rg and
M, are approximate. Thus, they placed the
species in Mydropastea, albeit incorrectly.
In Hampson’s Supplement (p. 209; 1914) he
synonymized his genus Mydropastea (1898)
under Phaio Neumoegen (1894). Draudt in
Seitz picked up the synonymy but changed the
spelling of the genus to Phaeo following Wagner
(1912).
Calonotus hoffmannsi Rothschild (1911) is
a synonym of Phoenicoprocta vacillans Walker.
Draudt in Seitz (p. 108, 1915; p. 201, 1917)
was correct in his suggested synonymy, namely,
Calonotus hoffmannsi (1911) is a synonym of
Paramya chrysonota (1898). Hampson in his
Supplement (p. 150; 1914) placed hoffmannsi
in Leucotmemis. Calonotus hoffmannsi was
described from Itaituba, Brazil, the same type
locality as Paramya chrysonota.
The male genitalia of the specimens from
Trinidad, two from Caripito, Venezuela, and a
specimen from British Guiana are similar. How-
ever, the genitalia of two specimens, one from
Candelopa, Peru, and another from Pitaguaya,
Bolivia, while very similar in other respects,
have a distinctly different pair of protuberances
on either side of the base of the uncus. They
are rounded and bulbous in the latter specimens
and pointed in our vacillans. The general facies
of the Peruvian and Bolivian specimens appears
to be the same as our material.
Judging from the preceding discussion it is
quite likely that a large number of the species
described in Phoenicoprocta will be synony-
mized. It is necessary, however, to study geni-
talia of the holotypes or in lieu of that, of topo-
typical material. In the subsequent paragraphs
I shall give an indication of the probable dis-
position of some of the species of Phoenico-
procta.
Phoenicoprocta mexicana (Walker) will have
to be removed from the genus. The sexes are not
dimorphic. The wing venation differs in the fore-
wing by lacking the sectorial vein in the radials.
It differs in the hindwing by veins Cui and Cu2
being completely united and veins Rg and M,
being approximate, not connate or very shortly
stalked.
Forbes (1930) correctly removed Zygaena
parthenii Fabricius from the genus Mallodeta
where Hampson (1898) placed it. Nonetheless,
it does not appear to be a Phoenicoprocta. The
venation differs in the forewing by lacking the
sectorial vein of the radials. It also differs in
the forewing in that vein R5 forks from the
radial midway between vein R2 and the forking
of veins R3 and R4, whereas in vacillans it sepa-
rates very near the forking of veins R3 and R4.
I have not seen a female nor has the female been
described, so I do not know if the sexes are
dimorphic. The hindwing of partenii is similar
in that veins Cu2 and Cus are forked near the
margin of the wing and veins M, and Rg are
slightly stalked.
Phoenicoprocta paucipuncta Dyar is not a
Phoenicoprocta but probably, as Forbes (1939)
suggests, a color form of Cosmosoma gemmata
Butler.
The following species may well be only color
forms of vacillans. Males: rubiventer Hampson
from Panama; sanguinea (Walker) , the species
type for which Hampson gives the type locality
as Honduras though Walker did not know the
type locality in his original description, the hand-
printed label on the holotype stating Honduras
being in neither Walker’s nor Hampson’s writ-
ing; astrifera (Butler) from Braga (Amazons)
Brazil; haemorrhoidalis (Fabricius) from Brazil;
ffavipicta Hampson from British Guiana; varia-
bilis Kaye from from Panama; intermedia Fors-
ter from Venezuela and nigriventer Gaede from
Venezuela; females: biformata Gibbs and the
aberration atrapennis Strand from British Hon-
duras; insperata (Walker) from Para, Brazil.
Phoenicoprocta astrifera (Butler), as Draudt
in Seitz figures it (pi. 12c; 1915), has wide
terminal wing margins. The holotype has wing
margins as in vacillans. The following were de-
scribed as forms of vacillans and should be re-
appraised: aurantipatagiata Draudt, aurifiua
Draudt, nigricoxa Zerny and punicea Strand.
While I have seen no specimens from the West
Indies, the form existing there appears to be a
distinct species. Hampson (1914: 125) synony-
mized thomae (Lucas), cubana Druce and
selecta (Herrich-Schaffer) under capistrata.
Fabricius’s type locality for capistrata is
“America”, which meant the mainland, and
1957]
Fleming: Ctenuchidae (Moths) of Trinidad, B.W.l.
113
Herrich-Schaffer’s type locality for selecta is
Brazil. Both are females and are more likely
the female forms of vacillans while exima (Her-
rich-Schaffer), described from Cuba, is the ac-
tual female form from the West Indies. If the
above suggestions prove to be correct, thomae
(Lucas) has priority and capistrata (Fabricius)
would preoccupy vacillans (Walker).
Phoenicoprocta jamaicensis Schaus is based
on a female type. Phoenicoprocta lydia (Druce) ,
with the synonyms thera (Druce) and demona
(Druce), is a distinct species from Mexico. It
has the same fore- and hindwing venation as
the vacillans we have from Trinidad and is
sexually dimorphic in the same fashion.
The forms of Phoenicoprocta from South
Brazil, Paraguay, Uruguay, Peru and Bolivia ap-
pear to be distinct from the forms found from
the Amazons north. Jorgensen’s species schrei-
teri and sieboldi appear to be the slight variants
of latimarginata Gaede (1926). The wider mar-
gins of the wings alone would seem to distinguish
these southern forms from vacillans. Two geni-
talia from Bolivian and Peruvian specimens that
were mentioned before are distinct from vacil-
lans though the facies of the insects are similar.
Phoenicoprocta teda (Walker) is a distinct
species but quite possibly not a Phoenicoprocta.
Loxophlebia Butler
In our species of Loxophlebia the discocellu-
lars are not oblique throughout in the hindwing
as Hampson (1898: 206) states in his generic
diagnosis. Furthermore, vein Cu2 of the hind-
wing is present but very shortly stalked with
vein Cui; the forking occurs at the inner edge
of the black wing border. The following genera,
Mesothen and Loxophlebia, may be separated
from other Trinidad genera by vein M2 of the
forewing originating well above vein Ms, more
than a third but less than half the distance up
between veins Mi and M3. The same vein, M2,
in Pheia is approximately a fifth in our species.
The genus Mesothen is very close to Loxophlebia
and I am unable to separate these genera except
superficially on the basis of the specimens on
hand, so the following key will serve to dis-
tinguish the Trinidad species in both genera.
Surinam and British Guiana forms of Loxo-
phlebia bisigna in the key would run to Meso-
then aurantegula, so I have included an addi-
tional couplet in the event that these forms are
found in Trinidad in the future.
1 . Abdomen with some red or orange color
2 (Loxophlebia)
Abdomen with no red or orange color
4 (Mesothen)
2. Abdomen with broad lateral orange bands
diaphana
Abdomen with no lateral bands of any
color 3
3., Abdomen blackish-brown with subdorsal
white spots on basal segment and sub-
dorsal red spots on fourth abdominal seg-
ment bisigna (Trinidad form?)
Abdomen black with the last four segments
orange postflavia
4. Disc of thorax orange -red pyrrha
Disc of thorax black or blackish-brown . . 5
5. Tegulae with orange patches 6
Tegulae concolorous brownish-black. . . .7
6. Abdomen concolorous blackish-brown
aurantegula
Abdomen with subdorsal white spots on
basal abdominal segments
bisigna (extra-limital forms)
7. Male with abdominal ventral valve. Female
with white subventral abdominal band
endoleuca
Male without abdominal ventral valve.
Female without white subventral abdomi-
nal band desperata
Loxophlebia diaphana (Sepp)
Glaucopis diaphana Sepp, 1848: II: 185, pi. 81.
Glaucopis discifera. Walker, 1854: I: 178.
Laeinocharis bura Butler, not Herrich-Schaffer,
1877: I: 33, pi. 11, fig. 12.
Chrysostola albifronsy[os,ch\&r, 1872: XXXIII:
344.
Loxophlebia diaphana, Hampson, 1898: I: 209.
Loxophlebia diaphana, Draudt in Seitz, 1915:
VI: 62, pi. 12g.
Loxophlebia diaphana, Kaye & Lamont, 1927:
No. 3 : 2.,
We have not collected this species at Simla,
but Norman Lamont collected a specimen at
Palmiste in January, 1921.
Surinam and Amazons.
Loxophlebia bisigna (Kaye)
(PI. Ill, fig. 6)
Pheia bisigna 1911: XLIV: 146.
Pheia bisigna, Hampson, 1914: Supp. I: 128, pi.
VI, fig. 17.
Loxophlebia klagesi Rothschild, 1811: XVIII :
29.
Loxophlebia klagesi, Rothschild, 1913: XX:
471, pi. 14, fig. 21.
Loxophlebia clagesi, Hampson, 1914: Supp.:
135.
114
Zoologica: New York Zoological Society
[42: 10
Pheia bisigna, Draudt in Seitz, 1915: VI: 61;
1917: Supp.: 199, pi. 27f.
Loxophlebia klagesi, Draudt in Seitz, 1915: VI:
63, pi. 12g.
Loxophlebia bisigna, Kaye & Lament, 1927 : No.
3: 2.
I have followed Draudt’s (1915, 1917) syn-
onymy of this species. The abdominal charac-
teristics of the original type material of klagesi
are variable. Rothschild’s paratype of klagesi
from Caporo {sic. Caparo), Trinidad, is similar
to our three specimens in having paired reddish
subdorsal spots on the fourth abdominal seg-
ment. Two paratypes from Maripa, Caura River,
Venezuela, have paired reddish spots on the third
and fourth abdominal segments. The four Suri-
nam types along with Kaye’s holotype of bisigna
from the Potaro River, British Guiana, lack
these reddish spots. Kaye’s bisigna was described
two months earlier than Rothschild’s klagesi.
The label on the holotype of klagesi in the Brit-
ish Museum (Natural History) has on the local-
ity label “Aroewarwa Creek, Maroewyn valley,
Surinam, May 1905. (S. M. Klages)” whereas
in Rothschild’s original description the material
from this locality is given as February.
/?fl77gc.— Trinidad, Venezuela and Guianas.
Loxophlebia postflavia Druce
(PI. Ill, fig. 7)
Loxophlebia postflavia Druce, 1898: I: 407.
Loxophlebia postflavia, Hampson, 1898: I:
210, pi. VIII, fig. 4.
Loxophlebia postflavia, Druce in Seitz, 1915:
VI: 64, pi. 12h.
Two specimens were collected. This is a new
record for Trinidad.
Described from French Guiana.
Specimens in British Museum from British
Guiana and Surinam.
Mesothen Hampson
In this genus, as in the former genus, Cui and
Cu2 of the hindwing are forked within the dark
terminal band. Thus Cu2 is present, not absent,
as stated in Hampson’s original description (p.
212; 1898). The species of this genus have been
keyed in the genus Loxophlebia.
Mesothen aurantegula (Jones)
Loxophlebia aurantegula D. Jones, 1914: 4, pi.
1, fig. 6.
Mesothen aurantegula, Hampson, 1914: Suppl.
I: 146, pi. VII, fig. 25.
Mesothen aurantegula, Draudt in Seitz, 1917:
VI: 200, pi. 27h.
Mesothen aurantegula, Kay & Lamont, 1927:
No. 3: 2.
An unusual record based on one specimen col-
lected by Norman Lamont at Palmiste in Janu-
ary. The species was described from one female
taken at Santos, southern Brazil.
Rrtngg.— Southern Brazil and Trinidad.
Mesothen endoleuca Druce
(PI. Ill, fig. 8)
Mesothen endoleuca Druce, 1905: XV: 460.
Mesothen endoleuca, Hampson, 1914: Supp. I:
146, pi. VII, fig. 27.
Mesothen endoleuca, Draudt in Seitz, 1915: VI:
66, pi. 27i.
Female with a ventrolateral band of white
scales on abdomen, broadest at the base of the
abdomen. Black border at the apex and margins
of the forewing much wider than in male. Fore-
coxae white in both sexes.
Eleven specimens were collected, of which
five were females.
Venezuela. A new record for Trini-
dad.
Mesothen desperata (Walker)
Pseudomya desperata Walker, 1856: VII: 1602.
Mesothen desperata, Hampson, 1898; 215, pi.
VIII, fig. 3.
Mesothen desperata, Hampson, 1914; 146, fig.
20.
Mesothen desperata, Draudt in Seitz, 1915; 66,
pi. 271.
Mesothen desperata, Lamont & Callan, 1950:
197.
One specimen reported by Lamont & Callan
(1950: 197) was collected by Lamont in April
at Palmiste. It is an aberrant species for the genus
as the male lacks a ventral valve according to
Hampson (1914: 146). Since our females of
endoleuca have a ventrolateral band broadest
at the base of the abdomen, not mentioned in the
description of desperata, I have used this char-
acter in the key to separate the female specimens
of these two species.
Range.— Panama, Guianas, Brazil, Paraguay
and Argentina.
Mesothen pyrrha (Schaus)
(PI. Ill, fig. 9)
Dycladia pyrrha Schaus, 1889: V: 89.
Dycladia pyrrha, Druce, 1896: II: 348, pi. 71,
fig. 27.
Mesothen pyrrha, Hampson, 1898: I: 214.
1957]
Fleming: Ctenuchidae (Moths) of Trinidad, B.W.I.
115
Mesothen pyrrha, Draudt in Seitz, 1915: VI: 66,
pi. 12i.
Mesothen pyrrha, Kaye & Lament, 1927 : No. 3 :
2.
Only one of our male specimens has the valve
white. In the remaining specimens it is brown
with the margins at the base of the valve broadly
and irregularly white. Only the proximal end of
the coxae is white, while the remainder of the
coxae is brown.
Material.— Fom males and one female.
Range.— MEXICO to Peru and the Guianas.
Pheia Walker
Very close to Cosmosoma but the lower side
of the discal cell is very short and the discocellu-
lar veins oblique. The only Trinidad species is
considerably smaller than any of the Trinidad
Cosmosoma. The wingspread of the smallest
Trinidad species of Cosmosoma, klagesi and
remotum, is 32 mm. whereas the following spe-
cies is 22 mm.
Pheia beebel, new species
(PI. I, fig. 2; PI. II, fig. 2)
Length of forewing of male 11-12 mm., of
female 13 mm.
Closely related to Pheia gaudens (Walker),
from which beebei differs in its smaller size and
narrower apical black patch on the forewings.
Pheia gaudens was described from Para, Brazil,
and has been reported from Venezuela and Peru.
Antennae bipectinate with a tuft on the apex
of each pectination. Pectinations along shaft of
antennae on the respective sides of the an-
tennae of similar length except near base and
apex. Pectinations on outside of antennal shaft
slightly longer than on the inside. Shaft of an-
tennae of male blackish-brown and of female
brown. Scape of antennae with bright yellow tuft
of scales most conspicuous on the inside anterior
edge.
Palpi, front of head and behind eyes bright
yellow. Basal segment of palpi clothed with long
hair of irregular length, thus somewhat ragged
in appearance. The remaining two segments of
palpi finely scaled and upturned to vertex of
head. Vertex of head black with some scattered
iridescent blue scales most conspicuous in the
center.
Collar bright yellow. Tegulae bright yellow,
finely edged with black scales on the outside and
broadly edged with black on the inside edge. A
patch of iridescent blue scales within the black
border of the inside edge.
Dorsum of the mesothorax yellow and of
metathorax iridescent blue. Caudal edge of meta-
thorax with a narrow line of bright yellow scales.
Pleura and ventrum of thorax bright yellow.
Legs yellow with small brown patches on the
mesothoracic and metathoracic legs. Forecoxae
immaculate bright yellow. Some brown on distal
segments of all tarsi.
Veins and margins of wings brown-black.
Margins of wings narrow but broadening at apex
of forewings (.91 mm. along vein Rs). In the
forewing the Sc vein and area between discal
cell and Sc vein to slightly beyond the end of the
cell yellow. A small patch of yellow at base of
wing between the discal cell and a line of black
scales marking the anal wing fold. Another yel-
low patch more densely scaled between the wing
fold to slightly beyond vein 2dA and terminating
distally at a small black patch which lies on the
anal fold. The distal end of the small black patch
is slightly more than 2 mm. from the base of the
wing. Some scattered black scales on the part of
vein 2dA which extends into the yellow basal
patch, but the basal part of the vein yellow.
Scales representing vein M2 extend within the
discal cell for approximately half the length of
the cell.
Hindwing with creamy yellow patch at base
of wing extending along anterior part of discal
cell for two-thirds the length of cell. The black
margin of the wing diminishing gradually and
disappearing completely near the base of the
wing. This part of the wing is normally hidden
beneath the forewing. The creamy patch extends
within the base of the cell and terminates at a
small black spot. Area below discal cell hyaline.
A small streak of yellow scales in brown-black
anal margin at base of wing.
Basal segment of abdomen including bullae
bright yellow but each of the subsequent seg-
ments except the terminal segment with the
anterior margin bright yellow and posterior mar-
gin brown-black. These latter brown-black bands
expanding into triangular marks on the dorsum
and obliterating the yellow on the mid-dorsum.
The brown-black bands have scattered patches
of iridescent blue or blue-green scales in vari-
ous lights. The terminal segment of the abdomen
is brown-black. Ventrum of abdomen yellow.
I take pleasure in naming this species after
William Beebe, Director Emeritus of the De-
partment of Tropical Research, New York Zo-
ological Society.
Material.— AU types were taken at Simla,
Arima Valley, Trinidad. Holotype, male. Cata-
log No. 57207, 16 XII; 2 paratypes (57208)
15-1 and (57209) 21-1.
Disposition of type material.— The Depart-
ment of Tropical Research, New York Zoologi-
cal Society, will retain two paratypes, Catalog
116
Zoologica: New York Zoological Society
[42: 10
Nos. 57208 and 57209. The holotype, Catalog
No. 57207, is in the American Museum of Natu-
ral History.
Chrostosoma Hubner
Distinctive in this group of genera by having
vein Ri of the forewing stalked with the other
radial veins and not free from the cell.
Chrostosoma viridipunctatum Rothschild
(PI. Ill, fig. 10)
Chrostosoma viridipunctatum, Rothschild, 1911:
30.
Chrostosoma viridipunctatum, Rothschild, 1913;
471, pi. XIV, fig. 34.
Chrostosoma viridipunctata, Hampson, 1914;
148.
Chrostosoma viridipunctatum, Draudt in Seitz,
1915: 67, pi. 12k.
Without an examination of the genitalia of
the holotype in the British Museum (Natural
History), it is impossible to be certain of the
above identification. A genitalic examination of
all the echemus-like species of this genus is
necessary to determine the valid species and
their respective ranges.
Hampson’s (1898: 215) division of the genus
into two sections on the basis of the presence
or absence of vein Cui in the hindwing is incor-
rect, as this vein is only forked nearer the margin
of the wing in the first section than in the second
section of the genus. Kenedy found that the
holotypes of decisum and viridipunctum had
both preserved vein Cui.
The series of viridipunctatum in the British
Museum (Natural History) contains three fe-
male specimens from Trinidad, one from Caparo
and two from Tabaquite. One of the latter speci-
mens has metallic coloring similar to Roth-
schild’s type, whereas in our series of males
and females any metallic coloring if present is
slight. In this same British Museum series are
specimens from Argentina, Paraguay, southern
Brazil, French Guiana and Surinam containing
both males and females.
The holotype is a female. The type locality
is Yungas de la Paz, Bolivia, 1,000 meters.
In our series of viridipunctatum the iridescent
blue, while variable, is never pronounced. Some
iridescent blue on the front and vertex of the
head, collar and shoulder covers and faint traces
on the subdorsum of the abdomen.
Materials— specimens (5 males and 4
females). A new record for Trinidad.
Guianas to Argentina and Bolivia,
Leucotmemis Butler
Leucotmemis differs from Chrostosoma in
having vein Ri of the forewing rising from the
cell rather than stalked on R2-Rs. Vein Cui of
the forewing from close to angle of the cell.
Leucotmemis nexa (Herrick-Schaflfer)
Loemocharis nexa Herrick-Schaffer, 1854: f.
254.
Leucotmemis nexa, Hampson, 1898: 224.
Leucotmemis nexa, Draudt in Seitz, 1915: 70,
pi. 13a.
Leucotmemis nexa, Kaye & Lamont, 1927: 2.
This species, reported from Trinidad by Kaye
& Lamont from St. Ann’s, Palmiste and Rock
Penal Road, has not been taken at Simla. It is
probably improperly placed in Leucotmemis as
the facies is entirely different from the re-
mainder of the genus with the exception of L.
insperata (Walker) . L. nexa resembles a female
Phoenicoprocta vacillans or a Calonotus. It may
be separated from the female vacillans by having
white lateral spots on the first abdominal seg-
ment. The sex of nexa has never been reported.
The facies of L. insperata can be encompassed
within the range of variation of the female
P. vacillans. Walker described insperata from
one female collected at Para, Brazil. There does
not appear to be any distinctive venational dif-
ference between Phoenicoprocta and Leucot-
memis, though most likely the accessory cell be-
tween vein R2 and vein R3+4+.'5 of the forewing
in Phoenicoprocta is sufficient to separate the
genera. In the only female of Leucotmemis I
have seen, L. lemoulti (Rothschild), the anten-
nal pectinations are equal on both sides of the
antennal shaft, whereas in Phoenicoprocta and
Calonotus the antennal pectinations in the fe-
male are much longer on the outside of the
antennal shaft than on the inside. In addition,
Calonotus has vein R2 of the forewing rising
from the cell as well as vein Ri. Calonotus hoff-
mannsi Rothschild, which was removed to Leu-
cotmemis by Hampson (1914), is a synonym
of Phoenicoprocta vacillans. Rothschild similar-
ly described lemoulti in Calonotus.
Range.— Mexico to Brazil.
Nyridela Lucas
A genus containing only two species, which
may be distinguished from related forms by
their large size. Vein Rr of the forewing rises
from the cell and Cu2 from near the angle of
cell.
Nyridela chalciope (Hubner)
Isanthrene chalciope Hubner, 1827: 20, figs.
469, 470.
1957]
Fleming: Ctenuchidae (Moths) of Trinidad, B.W.I.
117
Glaucopis acroxantha Perty, 1834: 156, pi. 31,
fig. 4.
Nyridela chalciope, Hampson, 1898: 218.
Nyridela chalciope, Draudt in Seitz, 1915: 69,
pi. 9k.
Nyridela chalciope, Kaye & Lament, 1927, 2.
A large hyaline-winged species with black
margins and a black band running from the
middle of the costal margin of the forewing to
the anal angle. Antennae with yellow tips and
the abdomen black with iridescent bluish-green
spots.
We have not collected this species at Simla
but Kaye & Lamont report it from St. Ann’s
Valley in the western part of the Northern
Range. This species is the South American
counterpart of the closely related species
from Central America, Nyridela xanthocera
(Walker).
Range— Colombia and Brazil.
CosMOSOMA Hubner
A large, somewhat confused, genus that needs
revision. Many of the species that Hampson
(1898) originally included he subsequently
(1914) placed in the genus Gymnelia which
has a lobed hindwing. Eight species are recorded
from Trinidad, of which we record six species.
1 . Discal bar broad 2
Discal bar narrow 4
2. Tegulae and patagia bright red
rubriscapulae
Tegulae and patagia brown or blue. ... 3
3. Abdomen with scarlet subdorsal streaks on
first segment melathoracia
Abdomen without scarlet subdorsal streaks
on first segment subflamma
4. Antennae with white tips 5
Antennae without white tips 7
5. Costal margin and terminal band of fore-
wing immaculate black anoxanthia
Patches of orange-red along costal margin
and terminal band at tornus of fore-
wing 6
6. Abdomen with a dorsal red stripe . achemon
Abdomen without a dorsal red stripe
remotum
7. Abdomen concolorous orange-yellow
klagesi
Abdomen yellow, second and third seg-
ments black with lateral blue spots . pytna
Cosmosoma rubriscapulae Kaye
(PI. Ill, fig. 11.)
Cosmosoma rubriscapulae Kaye, 1901: 116.
Cosmosoma rubriscapulae, Hampson, 1914:
156.
Cosmosoma rubriscapulae, Draudt in Seitz,
1915: 71, fig. 13b.
Cosmosoma rubriscapulae, Kaye & Lamont,
1924: 3.
The large size combined with the scarlet pa-
tagia and tegulae renders this moth distinctive
in Trinidad from other Cosmosoma species. We
have collected three males and three females
of this species at Simla. In our specimens the
discal bar of the forewings of the females is
broader and heavier than in the males.
Range.— Reported only from Trinidad.
Cosmosoma subflamma subflamma (Walker)
(PI. Ill, fig. 12)
Glaucopis subflamma Walker, 1854: 159.
Laemocharis panopes Herrich-Schaffer, 1854:
243.
Cosmosoma subflamma, Hampson, 1898: 227.
Cosmosoma siibflammum, Draudt in Seitz,
1915: 71, fig. 13b.
Cosmosoma subflamma, Kaye & Lamont, 1924:
3.
The bright red legs distinguish this species
from other Cosmosoma species in Trinidad.
Schaus described a species, lucia, from St. Lucia,
B.W.I., which Hampson (1898) records as a
subspecies of subflamma in which the legs are
largely black.
Material.— Wq have taken only one male. An-
other specimen was taken by F. W. Urich in
1918 in the Arima Valley and Lamont captured
a specimen on May 28, 1921, at Palmiste.
Range.— St. Lucia to south Brazil on the east-
ern side of South America.
Cosmosoma melathoracia Kaye
(PI. Ill, fig. 13)
Cosmosoma melathoracia Kaye, 1901: 115, pi.
V, fig. 10.
Cosmosoma melathoracia,'Hav!ypson, 1914: 157.
Cosmosoma melathoracia, Draudt in Seitz,
1915: 73, fig. 13e.
Cosmosoma melathoracia, Kaye & Lamont,
1924: 3.
Male with dorsal line of iridescent blue scales
and a brown valve at base of abdomen edged
laterally with white.
Material.— Six males.
Range.— Described and reported only from
Trinidad.
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Cosmosoma anoxanthia Druce
(PI. Ill, fig. 14)
Cosmosoma anoxanthia Druce, 1905: 460.
Cosmosoma bolivarensis Klages, 1906: 536.
Cosmosoma anoxanthia, Hampson, 1914: 165,
pi. VIII, fig. 25.
Cosmosoma anoxanthium, Draudt in Seitz,
1914: 80, fig. 27m.
Cosmosoma achemon, f. bolivarensis, Draudt
in Seitz, 1914: 79.
Cosmosoma anoxanthia, Kaye & Lament,
1924: 3.
Female similar to male except that on the
forewing the oblique quadrate patch of blackish-
brown scales from the lower extremity of the
discal cell to tornus (between veins Cui and
Cuo) is absent in the female.
Material —Six males and three females.
Trinidad, Venezuela and Bolivia.
Cosmosoma achemon (Fabricius)
(PI. Ill, fig. 15)
Zygaena achemon Fabricius, 1781: 162.
Euchromia tyrrhene Hubner, 1827: 23, figs.
483-484.
Cosmosoma voltumna Druce, 1897: 303.
Cosmosoma achemon, Hampson, 1898: 247.
Cosmosoma achemon, ab. tyrrhene, Hampson,
1914: 527.
Cosmosoma perfenestratum Dyar, 1899; 175.
Cosmosoma achemon, Draudt in Seitz, 1914:
79, fig. 14c.
Cosmosoma perfenestratum, Draudt in Seitz,
1914: 80, 1917: 201.
The very bright red dorsal abdominal stripe
will separate this species from all other Trinidad
Cosmosoma. A new record for Trinidad.
Material— One male from St. Augustine,
Trinidad.
Range.— Jamaica, Haiti, Venezuela and Brazil.
Cosmosoma remota (Walker)
Glaucopis remota Walker, 1854: 170.
Cosmosoma remotum, Hampson, 1898: 248,
pi. IX, fig. 22.
Cosmosoma remotum, Draudt in Seitz, 1914:
80, fig. 14c.
Cosmosoma remota, Kaye & Lamont, 1924: 3.
Reported by Kaye & Lamont from Tobago.
This species is very close to achemon, from
which it may be distinguished by the absence
of the red abdominal dorsal stripe. We have not
taken this species in the Arima Valley but Kaye
has specimens from Trinidad in his collection.
Range.— Trinidad, Tobago and Venezuela.
Cosmosoma klagesi Rothschild
(PI. Ill, fig. 16)
Cosmosoma klagesi Rothschild, 1910: 509.
Cosmosoma klagesi, Rothschild, 1913: 470, pi.
XIII, fig. 11.
Cosmosoma clagesi, Hampson, 1914: 170.
Cosmosoma klagesi, Draudt in Seitz, 1914: 82,
fig. 14g.
Cosmosoma klagesi, Kaye & Lamont, 1924: 4.
Mater/a/.— Fifty-one males and twenty fe-
males. Our commonest Cosmosoma in the Ari-
ma Valley.
Range.— Trinidad and Brazil.
Cosmosoma pytna Druce
Cosmosoma pytna Druce, 1906: 78.
Cosmosoma pytna, Hampson, 1914: 157, pi.
VIII, fig. 12.
Cosmosoma pytna, Draudt in Seitz, 1914: 83,
fig. 27m.
Cosmosoma pytna, Kaye & Lamont, 1924: 4.
We have not collected this species in the Arima
Valley nor, insofar as the literature reveals, has
it been collected since the holotype was taken
in Trinidad.
Dixophlebia Butler
The fringes of hair along the lower radial,
median and cubital veins of the forewing will
distinguish this genus from other genera of
Ctenuchidae.
Dixophlebia holophaea Hampson
(PI. Ill, fig. 17)
Dixophlebia holophaea Hampson, 1909: 346.
Dixophlebia holophaea, Hampson, 1914: 174,
pi. IX, fig. 9.
Dixophlebia holophaea, Draudt in Seitz, 1914:
85, fig. 14k.
A grayish-black moth with hyaline in the dis-
cal cell areas in both wings.
Material— Two males.
Raaga.— Surinam and British Guiana. A new
record for Trinidad.
PsEUDOMYA Hubner
A genus probably derived from Saurita, from
which Pseudomya may be distinguished by hav-
ing the hind tarsal joints fringed with scales in
the males.
1957]
Fleming: Ctenuchidae (Moths) of Trinidad, B.W.l.
119
Pseudomya melanthus (Stoll)
(PI. Ill, figs. 18, 19)
Sphinx melanthus Stoll, 1782: pi. 367, C.
Pseudomya melanthus, Hampson not Stoll,
1898: 264. (/ngM/fa Walker).
Pseudomya melanthus, Hampson, 1914: 175.
Pseudomya melanthus, Draudt in Seitz, not
Stoll, 1914: 87, fig. 14m. (trigutta Walker).
Pseudomya melanthus, Draudt in Seitz, 1917:
202.
Pseudomya melanthus, Kaye & Lamont, 1927:
4.
The facies of this species is very similar to
Pseudomya sanguiceps Hampson from Panama
(Hampson, 1898: 264, pi. X, fig. 4) figured in
Seitz on line 14m. The medial black band of the
forewings is darker and wider than shown in
the Seitz figure and extends from the costal to
the inner margin in the male. Our single female,
while having a very distinct quadrate patch of
dark scales at the discal veins, has only a scat-
tering of scales below the cell. Length of fore-
wing of the male 10 mm. and of the female
11 mm.
Material— One. male and one female.
Trinidad and Surinam.
Rhynchopyga Felder
Differs from Saurita and Pseudomya in hav-
ing Cui and Ms on a long stalk in the hindwing.
Rhynchopyga flavicollis (Druce)
(PI. Ill, fig. 20)
Amycles flavicollis Dmce, 1884: 46, pi. 7, fig. 11.
Rhynchopyga flavicollis, Hampson, 1898: 270.
Rhynchopyga flavicollis, Draudt in Seitz, 1914:
90, fig. 15e.
The long, very narrow brown wings of this
slender, small species is distinctive. Our single
male specimen lacks the orange streak on the
patagia present on the holotype. Our specimen
also has less orange on the tegulae.
Material.— One male. A new record for Trini-
dad.
Guatemala, British Honduras, Costa
Rica, Panama and Colombia.
Saurita Herrick-Schaffer
Probably not a natural genus but attempts
to divide it have thus far been unsuccessful.
Veins Ms and Cui of the hindwing arise from
the same point or are very shortly stalked.
1 . Abdomen of male with ventral valve cover-
ing basal segments. Abdomen of male and
female large, with iridescent blue sublateral
spots and crimson anal tufts .... cassandra
Abdomen of male with no ventral valve.
Abdomen of male and female other-
wise 2
2. Thorax with crimson patches only 3
Thorax orange red or crimson 6
3. Abdomen with whitish patches 4
Abdomen with brown or black patches . . 5
4. Discal cell of forewing brown-scaled
lacteata
Discall cell of forewing hyaline . . orimensis
5. Forewing uniformly hyaline or smoky
hyaline perspicua
Forewing with post-discal milky hyaline
band clusia
6. Vertex of head crimson red salta
Vertex of head black 7
7. Wings fully scaled with only slight thin-
ning of scales between veins concisa
Discal cell and area below discal cell of
forewing hyaline 8
8. Underside of body yellow te menus
Underside of body black afflicta
Saurita cassandra (Linnaeus)
(PL III, fig. 21)
Sphinx cassandra Linnaeus, 1758: 494.
Saurita cassandra, Hampson, 1898: 274, fig.
111.
Saurita cassandra, Draudt in Seitz, 1915: 93,
fig. 15f.
Saurita cassandra, Kaye & Lamont, 1927: 4.
A robust, brown-winged Saurita with the dis-
coidal patch darker and extending somewhat
along the cubital veins. Males and females simi-
lar except for the longer pectinations on the
male antennae.
Material.— One male.
Range.— Venezuela to Argentina.
Saurita clusia (Druce)
(PI. Ill, figs. 22, 23)
Laemocharis clusia Druce, 1897: 303.
Hypocharis clusia, Hampson, 1898: 271, fig.
125.
Hypocharis clusia, Draudt in Seitz, 1915: 91,
fig. 15e.
Hypocharis clusia, Kaye & Lamont, 1927: 4.
Saurita clusia, Forbes, 1939: 121.
Forbes (1939: 121) placed clusia in Saurita,
pointing out that the character Hampson (1898:
271) employed to erect the genus Hypocharis
is equally true of Saurita, namely, vein Mi of
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[42: 10
the forewing is more or less from below the
angle of the cell in all species of Saurita as well
as Hampson’s Hypocharis.
This species will key to Saurita nox (Druce)
in Hampson’s key (1898; 275). Hampson’s
footnote indicates that he examined the holo-
type of nox which was in the Staudinger collec-
tion. In his description of the species he men-
tions “a broad diffused whitish band from just
beyond middle to termen.” He gives the wing
expanse as 22 mm.
Druce, on the other hand, neither in his orig-
inal description (1896: 30) nor in the Biologia
(1897: 341, pi. 71, fig. 11) where he copies his
original description, makes any mention of a
whitish postmedian band but states simply,
“primaries and secondaries smoky hyaline, with
the veins all black.” His figure (/.c.) illustrates
a smoky hyaline-winged moth with blackish
veins. Druce records the wing expanse as 114
inches.
Draudt in Seitz (1915: 93, fig. 15g) shows a
figure resembling the figure of Druce. Forbes
conjectures that Draudt’s figure is of S. fumosa.
In his description, however, Draudt writes of a
“postdiscal, faded, whitish spot.” He gives the
wing expanse as 22 mm. He compares the
species to Saurita lacteata Butler which has a
different-shaped inner margin of the hindwing.
(See Forbes, l.c. for grouping of species of
Saurita) .
Hampson (1914: 186) synonymizes Chrosto-
sonia maratha Druce as the female of Saurita
nox. The holotype is in poor condition with the
wings torn and rubbed and the abdomen miss-
ing. However, the whitish postdiscal band
crosses the forewing in the photograph slightly
distad of the discoidal veins. The facies of the
forewing looks very similar to that of clusia.
Specimens from Trinidad are included within
the series of Saurita clusia in the British Museum
(Natural History) collection. These Trinidad
specimens and also those in Kaye’s private
collection under clusia are similar to the speci-
mens we collected. However, if I were to follow
my interpretation of Forbes (1939: 121) I
should have identified them as nox. Forbes states
that clusia is smaller than nox, the white post-
medial band narrower sex for sex, and the blue
spotting is distinctive, and for nox, “There are
no blue spots; in the male the white area invades
the outer third or half the cell, in the female
the extreme apex of the cell may be pale.” First
of all, our series of specimens varies from having
a distinctively blue spotted abdomen to a com-
pletely unspotted concolorously colored ab-
domen with the completely unspotted section
representing approximately half the collection.
Our specimens are much larger than the speci-
mens in the American Museum of Natural His-
tory from Panama identified as clusia by Forbes.
All of our specimens have the postmedial band
invading the discal cell to the extent of half or
more. Lastly, our specimens differ in genitalia
from those in the American Museum under
clusia.
Hampson (1898: 271) mentions the blue
spotting in regard to clusia. His figure of clusia
is similar to our photograph of the holotype of
clusia, but his wing expanse is much larger (30
mm.) (and Druce 114") than the six specimens
named clusia by Forbes (23-25 mm.) in the
American Museum of Natural History. In other
than the size difference, Hampson and Forbes
agree.
Aside from the discrepancy between Druce’s
original description of nox and the subsequent
descriptions by other authors which make the
identity of nox questionable, it is quite possible
that we are concerned with more than two
species. However, I consider that it would be
unwise to describe the Trinidad form without
having material throughout the range of the
above two species.
Saurita lacteata (Butler)
Dycladia lacteata Butler, 1877: 34, pi. 17, fig. 3.
Saurita lacteata, Hampson, 1898: 276, fig. 128.
Saurita lacteata, Draudt in Seitz, 1915: 93,
fig. 15g.
Saurita lacteata, Kaye & Lamont, 1927: 4.
This species is listed by Kaye & Lamont as
occurring in Trinidad but the specimens in
Kaye’s collection under this name are similar
to the following new species. The specimens
under lacteata in the British Museum (Natural
History) from Trinidad are likewise in agree-
ment with the following species. Butler describes
lacteata from the Rio Jutahi, Amazons.
Saurita orimensis, new species
(PI. I, fig. 3; PI. II, fig. 3)
Length of forewing of male 10.5-11.5 mm.
Related to Saurita lacteata Butler from which
arimensis differs in having the discal cell hyaline
and in not having a yeUowish-white patch below
the cell and another yellowish-white patch be-
yond the cell.
'Antennae bipectinate in the male with each
pectination dilated and bristled on the distal
end. Palpi brown, reaching vertex of head with
fan-shaped brown tuft on first segment. Whole
head uniform brown.
1957]
Fleming: Ctenuchidae (Moths) of Trinidad, B.W.I.
121
Patagia and disc of thorax uniform brown
with a small white patch on the mid-dorsum of
the metathorax. Tegulae brown with a crimson
patch on anterior margin near patagia (shoul-
ders) and a white bar at discal edge of tegulae
on the anterior margin of forewing. The inner
half of tegulae along the disc of the thorax
crimson.
Some white on the outer edge of the basal
half of the forecoxae on the episternum (2nd).
The remainder of the legs and the lateral and
ventral parts of the thorax brown.
Fore- and hindwings hyaline. Veins brown.
The forewings narrowly bordered with brown
but the apex of the wings broadly brown. The
brown patch at the apex extending within the
wing to the point where veins R4 and R5 fork.
A relatively broad brown discal bar (1.25 mm.) .
Hindwings with narrow brown margins broaden-
ing at apex. At the 1st anal fold the brown color
extends within the wing for approximately one-
third of the length of the wing from the margin,
and a brown patch extending into the wing at
the anal angle for approximately the same
distance.
Abdomen with the dorsum of the first and
second segments and to a variable extent the
third segment creamy white. The bullae creamy
white. A creamy white band extends from bullae
along the spiracular region of the abdomen for
four segments. The band is broadest at the
base and narrows toward the distal end of the
abdomen. The remainder of the abdomen brown.
The name arimensis has been taken from the
type locality, Arima Valley.
Material— All types were taken at Simla, Ari-
ma Valley, Trinidad. Holotype, male, Catalog
No. 57200, 2-V; 6 paratypes, (57201) 2-VI,
(57202) 16-III, (57203) 21-IV, (57204) 13-11,
(57205) 7-1, (57206) 3-VI.
Disposition of type material— The Depart-
ment of Tropical Research, New York Zoologi-
cal Society, will retain two paratypes. Catalog
Nos. 57205 and 57206. Paratype, Catalog No.
57204, is in the collection of the British Museum
(Natural History) and paratype. Catalog No.
57203, is in the United States National Museum
collection. The holotype, Catalog No. 57200,
and the paratypes. Catalog Nos. 57201 and
57202, are in the American Museum of Natural
History.
Saurita perspicua Schaus
Saurita perspicua Schdios, 1905: 187.
Saurita perspicua, Hampson, 1914: 185, pi. IX,
fig. 31.
Saurita perspicua, Draudt in Seitz, 1915: 93,
fig. 28e.
Saurita perspicua, Kaye & Lamont, 1927: 4.
This species has not been collected since the
original type. Subsequent mention in the litera-
ture has been based on the original description.
The type is a female in the U.S. National Mu-
seum. Schaus (1905) states that the wings are
smoky hyaline and Hampson (1914) simply
says that they are hyaline. We have not taken
this species at Simla.
Trinidad.
Saurita salta (Schaus)
Thrinacia salta Schaus, 1894: 226.
Saurita salta, Hampson, 1898: 277, pi. X, fig. 12.
Saurita salta, Draudt in Seitz, 1915 : 94, fig. 15h.
Saurita salta, Kaye & Lamont, 1927: 5.
Described from Venezuela. Kaye &
Lamont report it from Trinidad, with no data.
Saurita temenus (Stoll)
Sphinx temenus Stoll, 1781: pi. 367, D.
Saurita temenus, Hampson, 1898: 279.
Saurita temenus, Draudt in Seitz, 1915: 94, fig.
16b.
Saurita temenus, Kaye & Lamont, 1927: 5.
See following species, Saurita afflicta, for dis-
cussion.
Rnn^e.— Surinam, Amazons.
Saurita afflicta (Walker)
(PI. Ill, figs. 24, 25)
Glaucopis (Pseudomya) afflicta Walker, 1854:
144.
Glaucopis afflicta, Butler, 1877 : 29, pi. 7, fig. 12.
Saurita temenus, Hampson, 1898: 279 (in part).
Saurita venezuelensis Klages, 1906: 538.
Saurita venezuelensis, Draudt in Seitz, 1917:
94, fig. 15h.
Saurita temenus, Draudt in Seitz, 1917: 94, fig.
16b (in part).
Saurita afflicta, Forbes, 1939: 123.
The Saurita temenus record of Kaye & La-
mont (1927: 5) may be in error and their
specimens rightfully placed in this species. The
specimens we have taken at Simla are S. afflicta.
This species was listed by Hampson (1898: 279)
as a synonym of S. temenus and subsequent
authors until Forbes (1939: 123) followed
Hampson. Forbes was the first to point out that
the two names represented different species most
easily separated by the yeUow ventrum of S.
temenus and the blackish ventrum of S. afflicta.
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[42: 10
However, since S. afflicta was described from the
Amazons and S. temenus from Surinam, it is
possible that both species inhabit Trinidad.
The females may be distinguished from the
males by having the spaces between the veins
beyond the cell hyaline.
Material —11 specimens: 12 males and five
females.
Honduras to the Amazons.
Saurita concisa (Walker)
Euchromia concisa Walker, 1854: 243.
Thrinacia afflicta L)T\ice, 1884: 56 (not Walker).
Saurita concisa, Hampson, 1898: 279.
Saurita thoracica Klages, 1906: 538.
Saurita concisa, Hampson, 1914: 189.
Saurita thoracica, Draudt in Seitz, 1915: 94.
Saurita concisa, Kaye & Lamont, 1927: 5.
Kaye & Lamont placed Saurita venezuelensis
Klages as a synonym, but I have followed Forbes
and synonymized venezuelensis under Saurita
afflicta Walker.
Kaye & Lamont record this species from
Palmiste, Trinidad, but we have not collected it
as yet at Simla.
Panama to the Amazons.
PsoLOPTERA Butler
A small genus closely related to Saurita, from
which it differs by having vein Ri forked with
R2 in the forewing.
Psoloptera leucosticta (Hubner)
Glaucopis leucosticta Hubner, 1827: t. 162.
Psoloptera leucosticta, Hampson, 1898: 285.
Psoloptera leucosticta, Draudt in Seitz, 1915:
96, fig. 15 m.
Psoloptera leucosticta, Kaye & Lamont, 1927: 5.
Kaye & Lamont report this species from
Trinidad, one specimen from Guaico at the
southern foot of the Northern Range and two
specimens from Palmiste. We have not taken
it at Simla, but we collected the species at Cari-
pito, Venezuela, across the Gulf of Paria from
Trinidad. The wings and body are purplish-
black with two white points at the base of the
abdomen.
Range.— Venezuela, Trinidad, Guianas and
Amazons.
Dycladia Felder
A small genus of moths that are very beetle-
like. The following species in particular, like the
species of the genus Correbidia in the next sub-
family, resemble a Lycidae beetle.
1. Orange lateral line of the abdomen not
reaching beyond middle of the abdomen
and no black spot at the base of the fore-
wing correbioides
Orange lateral line of the abdomen absent
only on the last segment and base of fore-
wing with large black spot. . . basimacula
Dycladia correbioides Felder
Dycladia correbioides Felder, 1874: pi. 102,
fig. 20.
Dycladia correbioides, Hampson, 1898: 293,
fig. 139.
Dycladia correbioides, Draudt in Seitz, 1915:
99, fig. 16d.
Dycladia correbioides, Kaye & Lamont, 1927 : 5.
Kaye & Lamont record this species from Pal-
miste but see following species, Dycladia basi-
macula Schaus.
Dycladia basimacula Schaus
(PI. m, fig. 26)
Dycladia basimacula Schaus, 1920: 9.
It is most likely that the D. correbioides of
Kaye & Lamont is this species. D. correbioides
is the name used for the form found in Colombia
and Panama, and D. emerita for the form from
Costa Rica to Mexico. Schaus described D. basi-
macula from Venezuelan and Trinidad material.
Material— Vova males.
Range.— Trinidad and Venezuela.
Syntomeida Harris
The three posterior veins from the discal cell
of the hindwing are forked.
Syntomeida melanthus (Cramer)
Sphinx melanthus Cramer, 1779: pi. 248, C.
Sphinx nycteus Stoll, 1780: pi. 325, F.
Euchromia apricans Walker, 1854: 224.
Syntomeida albifasciata Butler, 1876: 366.
Syntomeida melanthus, Hampson, 1898: 306,
fig. 138.
The lustrous blue-black wings with a variable
number of quadrate yeUow spots and orange-
red and black bands on the abdomen separate
this insect from other Trinidad ctenuchids. The
names albifasciata, nexilis and nycteus apply to
variants in either the number of yellow wing
spots or abdominal coloration.
Kaye & Lamont report this species from San
Fernando.
1957]
Fleming: Ctenuchidae (Moths) of Trinidad, B.W.l.
123
Mexico to Uruguay and Peru.
Histiaea Walker
The largest of the Trinidad ctenuchids, with
densely scaled, brown wings variously marked
with red and yellow or grayish spots.
1. Abdomen with two basal segments com-
pletely yellow dorsally. Forewing with ex-
tensive longitudinal red rays running from
base meldolae
Dorsum of abdomen brown with two sub-
dorsal spots on basal segment, and triangu-
lar, subdorsal yellow spots on the anterior
parts of the following two segments. Fore-
wing with at most red or reddish-tinged
spots or small streaks cepheus
Histiaea meldolae Butler
(PI. Ill, fig. 27)
Histiaea meldolae Butler, 1876: 362.
Histiaea meldolae, Druce, 1884: 42, pi. 6, fig.
14.
Histiaea meldolae, Hampson, 1898: 311.
Histiaea meldolae, Draudt in Seitz, 1915: 101,
fig. 16g.
Histiaea meldolae, Kaye & Lamont, 1927: 5.
Material —ThrQC males and three females.
Range— Ths holotype is a Trinidad specimen.
British Guiana, Venezuela, and Panama.
Histiaea cepheus (Cramer)
(PI. Ill, fig. 28)
Histiaea cepheus Cramer, 1780: pi. 109, E.
Histiaea cepheus, Hampson, 1898: 313.
Histiaea monticola Klages, 1906: 538.
Histiaea monticola, Hampson, 1914: 203.
Histiaea cepheus, Draudt in Seitz, 1915: 101,
fig. 16g.
Histiaea monticola, Draudt in Seitz, 1915: 102.
Histiaea cepheus, Kaye & Lamont, 1927: 6.
Ma/ma/.— Forty-six males and 17 females.
TJange.— Venezuela to Surinam.
Macrocneme Hubner
This genus of iridescent wasp-like moths is
in a very confused condition. The iridescence
of the wings, while diagnostic in many instances,
is extremely difficult to describe. Forbes (1939)
has written the only paper that is of any real
assistance. The descriptions in Hampson (1898
and 1914) are valuable but his taxonomic
treatment is inadequate. Genitalic studies of the
types and series of specimens must be made
and associated with other characters before
identification in this genus will have any relia-
bility.
The following key is based on specimens we
have collected and some unidentified Trinidad
material loaned by the British Museum (Natu-
ral History).
1. First segment of abdomen with four white
spots; two subdorsal and two lateral. No
ventral valve in male. Large species with
a wing length of 16 mm. or more 2
First segment of abdomen with only two
iridescent blue or green subdorsal spots
on conspicuous abdominal bullae. Ven-
tral valve present in male. Smallest
species with a wing length of 14 mm.
or less 6
2. Males 3
Females 5
3. At least the basal 2/3 of the ventrum of the
abdomen with a broad uninterrupted
white band thyra
Otherwise; white of underside of the ab-
domen broken up into spots 4
4. Forecoxae iridescent blue species?
Forecoxae white spinivalva
5. Iridescence of the forewing uniform green
to end of discal ceU except slight black
along base of costal margin and small
black point on the base of the anal vein.
(True of males also) thyra
Iridescence of forewing interrupted by a
black band from inner margin to at least
the anterior part of discal cell. (True of
males also) spinivolva
6. Forewing with streaks of iridescent blue or
green from base of wing to end of discal
cell vittata
Forewing with only blue or green spots at
base of wing 7
7. Abdomen with sub ventral white spots on
two medial segments plumbea
Abdomen without subventral white spots
albitarsia
Macrocneme plumbea (Hampson),
new combination
(PI. Ill, fig. 29)
Poliopastea plumbea Hampson, 1898: 337, pi.
XII, fig. 26.
Poliopastea plumbea, Draudt in Seitz, 1915:
110, fig. 18b.
Our specimens agree with Hampson’s original
description of plumbea but the description is
inadequate for proper determination. Hampson
states that the abdominal valve is blue edged
124
Zoologica: New York Zoological Society
[42: 10
with white, with a white patch behind it. Our
specimens have these characters, but in addition
the two segments caudad of the white patch have
small subventral white spots. Draudt in Seitz
may have had similar specimens, as he states:
“—the next rings indistinct, white sublateral
spots.” Kenedy, in notes, writes that the type
has small, sublateral spots on the next segment
and sublateral metallic blue spots on next two
segments. One of our specimens shows only faint
traces of the second pair of spots. AH of our
specimens have subventral blue spots on the sub-
terminal abdommal segments but they may be
very faint. The hind legs are missing on the holo-
type, but in the series of specimens of plumbea
(not type material) in the British Museum
(Natural History) the hind tarsi are white.
Hampson’s male holotype came from the lower
Amazons, Paratins. Draudt records the species
from the Amazon and French Guiana, and
specimens in the British Museum (Natural His-
tory) collection are from Trinidad, Venezuela,
British Guiana and Sao Paulo, Brazil. My identi-
fication of the Trinidad material is provisional
on a genitalic examination of the holotype in the
British Museum.
Hampson erected a new genus, Poliopastea,
for plumbea and made plumbea the type species
of the genus. Hampson’s conception of the dif-
ferences between Macrocneme and Poliopastea
appears to be the porrect palpi in Poliopastea,
veins Cui, Ms and Ms of the forewing close to
the angle of the cell and vein Rs of the fore-
wing from the cell. The specimens that I have
seen from Trinidad have individuals with both
porrect and upturned palpi. Vein Cui of the
forewing is variable in Macrocneme so that the
character, veins Cui, Ms and Ms from close to
angle of cell, has no significance and some spe-
cies of Macrocneme have vein Rs of the fore-
wing from the cell. Consequently, I consider
Poliopastea a synonym of Macrocneme. I have
not examined any of the other species that have
been placed in Poliopastea but it seems likely
that obscura (Wallengren) and viridis (Druce)
can be included in Macrocneme. Hampson
(1914: 207) placed ochendeni Rothschild in
Macrocneme. He erected a new genus Pseudo-
phaio for rosenbergi Rothschild and provision-
ally transferred verdivittata to Calonotus. Polio-
pastea pava (Dognin) was aberrant in the genus
and required a separate section. It was originally
described by Dognin in the genus Thysanopry-
ana, which is a synonym of Baritius in the family
Arctiidae.
Kaye & Lamont listed plumbea as a synonym
of eacus along with vittata and nigritarsia. The
figure of eacus in Stoll (1781, fig. 335 C) has
evanescent, iridescent blue in the ceU and some-
what beyond, and vittata (see above under
vittata) has iridescent blue above and below the
cell as well as within the cell. Hampson’s nigri-
tarsia is a larger moth than eacus and the iri-
descent blue is more distinct and extensive.
Lamont & Callan (1950: 197) report albitarsia
from Trinidad (Palmiste) . This species may be
separated from plumbea by the absence of the
white spots on the ventrum of the abdomen.
Both eacus and nigritarsia have black hind tarsi,
whereas albitarsia, plumbea and vittata have the
terminal segments of the hind tarsi white.
Material.— Four males from Simla and two
males from the British Museum (Natural His- |
tory) from Caparo and San Fernando, Trinidad. j
Macrocneme thyra thyra Moschler
Macrocneme thyra Moschler, 1883: 334, 1. 18, ij
fig. 24. 1
Macrocneme thyra, Hampson, 1898, 321. \
Macrocneme thyra, Draudt in Seitz, 1915, 103,
fig. 17a.
Macrocneme albiv enter Dognin, 1923 : 2.
Macrocneme thyra, Kaye & Lamont, 1927 : 6.
Macrocneme thyra, Forbes, 1939: 129, 1. 1,
fig. 3 (genitalia).
This is the nomenclatural type that was de-
scribed from Surinam. One male specimen from
the collection in the British Museum (Natural
History) I have tentatively assigned to this form. |
It was collected at Tabaquite, Nariva District, ;
Trinidad. This single specimen has a very short
streak of iridescent blue at the base of the wing !
in the lower part of the cell and just below the
cell. Iridescent blue in the distal half of the cell,
just above the cell and belov/ the anal vein, the |
latter extending almost to the base of the wing. I
The area between the cell and the anal vein black |
but with a trace of iridescence below the irides- ;
cence in the cell end. This specimen has more
extensive black in the basal part of the wing
than Hampson indicates for thyra. The ventrum
of the abdomen is white. While I have not re-
moved the genitalia, it is well extruded and I
have been unable to see any significant difference
between the genitalia of this specimen and the
genitalia of thyra that Forbes (1939) figures,
and the following race.
Range.— Guianas, Brazil, Peru to Panama?
Macrocneme thyra intacta Draudt
(PI. Ill, fig. 31)
Macrocneme thyra, subspecies 1, Hampson,
1898: 321.
Macrocneme thyra intacta, Draudt in Seitz,
1915: 103.
1957]
Fleming: Ctenuchidae {Moths) of Trinidad, B.W.l.
125
I am unable to find any essential details of the
genitalia of the Trinidad specimens different
from the genitalic figures of Forbes (1939).
Both processes of the male valves are curved as
in the figure and the upper process of the left
valve has a distinct tooth on the inner edge. This
last feature is not shown in the figure but stated
in the text. The juxta is squarely cut off and
short. The uncus is the same shape as in the
figure, but with small, narrow, lateral-winged
margins which I do not see in the figure.
This is subspecies 1 of Hampson. Draudt ap-
pears to have given the subspecies a name on the
basis of Hampson’s description without having
specimens before him. Hampson (1898) char-
acterized it in part by stating “Forewing with
blue-green at base entire.” This is true of our
Trinidad specimens except as noted in the key;
namely, the Trinidad specimens have black on
the base of the costal margin and a small black
streak or spot on or near the base of the anal
vein. This last-mentioned black streak or spot is
never very conspicuous and is occasionally
absent.
Material— lAA specimens from Simla (137
males and 7 females) and 9 specimens (6 males
and 3 females) from Port of Spain, Guaico,
Ariapite Valley and St. Ann’s borrowed from
the British Museum (Natural History).
Stated by Hampson as Trinidad and
Colombia.
Macrocneme spinivalva, new species
(PI. I, fig. 4; PI. II, fig. 4)
Length of forewing of male 17 mm., of female
18 mm.
Antennae dark brown, bipectinate. Each
pectination tufted on distal end and length of
pectinations rapidly decreasing at proximal and
distal ends of antennal shaft. Pectinations
shorter in male than in female.
Palpi upturned to vertex of head. First seg-
ment of palpi clothed with ragged, dark brown
scales with an immaculate white medial tuft of
broad scales on anterior face, for two-thirds to
three-quarters the length of the second segment
of the palpi from the proximal end. The line of
white scales absent in female. Ground color of
second and third palpal segments dark brown in
both sexes.
Front of head dark brown with two white
spots on upper edge beneath the antennal scapes
in both sexes. Vertex of head brown.
Patagia (coUar) with a pair of dorsal white
spots and a pair of subdorsal white spots. In
the male, but not the female, each dorsal spot
joined to its respective subdorsal spot by a fee
line of white scales on the anterior margin of
the patagia. Immediately below the subdorsal
spots and hardly separated from them, anterior
to the bases of the tegulae and approximately in
line with the middle of the eyes, a small white
spot. A rectilinear episternal white spot in line
with the lower part of the eyes and base of palpi.
The episternal spot is present in the female
though usually smaller than in the male, but the
spot below the subdorsal spot is absent in the
female. Tegulae dark brown with iridescent
green reflections most pronounced on the an-
terior edge. Dorsum of thorax dark brown with
iridescent reflections in various lights, most dis-
tinct on the metathorax.
Forecoxae of the male white but the re-
mainder of the forelegs brown with faint irides-
cent reflections except for the caudal edge of the
femur of the forelegs which is gray. Each of the
forecoxae of the female with two small white
spots on the proximal end, one laterad and one
dorsad. The remainder of the female legs brown
with varying iridescent reflections. Mesothoracic
legs in both sexes brown with some blue reflec-
tions, with a small white spot on the distal end
of the femur. Metathoracic legs of the same
brown color with reflections but with the distal
one-half to two-thirds of the tibia in both sexes
fringed. The metatarsus with long fringe. The
fringe on the third, fourth and fifth tarsal seg-
ments white.
Forewing with two white spots on base of
wing near tegulae. Distinct blue iridescence in
the forewing is confined to the following five
areas :
(1) A streak the length of the discal cell
above the discal cell,
(2) to the distal half of the discal cell,
(3) to the same respective part of the wing
below the discal cell except that the iridescence
foUows vein Cu2 in the direction of the tornus
a little beyond the end of the discal cell,
(4) a basal patch below the base of the discal
cell, and
(5) a median streak below the anal vein.
The remainder of the wings including the veins
through the iridescent areas blue to brownish-
black depending on the age of the specimen. The
above description of the forewing pattern may
be summarized by saying that the iridescence is
confined to the plane of the end of the discal cel!
interrupted by a dark transverse band perpen-
dicular to a dark anal streak. Underside of fore-
wing with the basal half of the wing to cell end
iridescent blue.
Upperside of hindwing bluish or brownish-
black depending on the age of the specimen.
Underside of hindwing iridescent blue except
126
Zoologica: New York Zoological Society
[42: 10
for apical and anal area. In old specimens the
iridescence may be restricted to the cell and
costal area.
Abdomen with two pairs of white spots, one
pair subdorsal and the other on the hoods or
bullae. First segment of abdomen blackish-
brown and the remainder of the abdominal seg-
ments iridescent blue-green with bluish-black
subdorsal longitudinal bands. The background
color may become quite brassy in old specimens.
Ventrum of male without a basal ventral valve
but a pair of subventral white spots on the first
segment in both sexes. Subsequent segments of
the abdomen in both sexes with a midventral
white spot except the last abdominal segment
which, like the first segment, has a pair of white
subventral spots.
Male genitalia with base of uncus broadly in-
flated and the edges thin in cross-section with
the distal process normally directed ventrally
and slightly spatulate. Valves large with two
processes, the dorsal process curved with a dis-
tinct thorn-like structure on inner edge of the
distal half and a ventral process which is slender
and hirsute. Juxta very slender, so slender as to
be remarkable for the genus.
Female genitalia with signa of bursa copula-
trix spherical, with long spines, sea-urchinlike,
with small anterior portion unspined. In the
female genitalia of thyra, the other species of
this section of the genus in Trinidad, the signa
is ovoid or almost subquadrangular with short
spines.
This is most likely the species mentioned by
Forbes ( 1939: 126, pi. II, fig. 8) which he refers
to as Macrocneme species. It will run to this
species in his key to the species of Macrocneme
and his figures are extremely similar. The tegu-
men and uncus are less inflated laterally than
in spinivalva and do not show any part of the
distal end of the uncus. The thorn or tooth on
the dorsal process of the valve appears more
pronounced and acuminate than in Forbes’s
figure of the valve, but this may be a matter of
perspective.
This species seems to be most closely related
to thyridia with which we had it confused in our
collection. The tooth or thorn-like structure on
the upper process of the valve and extremely
slender juxta will separate spinivalva from
thyridia. The spined condition of the valve can
often be seen without removing the genitalia in
the male. Thus far, we have not found thyridia
in Trinidad.
The specific name spinivalva, meaning thorn-
valve, refers to the thorn-like structure on the
upper process of the male genitalic valve.
Material.— male. Catalog No. 5716,
Simla, Arima Valley, 27-III; allotype, female.
Catalog No. 5717, Simla, 26-III; paratypes,
male and female in coitu, (5718) Simla, 22-XII;
paratypes, males, (5719) Simla, 16-11— (5720)
#288, Trinidad, B.M.-(5721) June, 1902, Aria-
pite Valley, B. M.— (5722) June, 1902, Ariapite
Valley, Trinidad, B. M.; paratypes, females,
(5345) Simla, 17-III-(5724) Simla, 6-III-
(5725) Simla, 27-III-(5726) Simla, 13-III-
(5727) Trinidad, B. M.-(5728) Trinidad, B. M.
—(5729) Trinidad, Dr. Jackson, 1921-1922,
B. M.-(5730) Trinidad, A. HaU, Feb. 1930,
B. M.— (5731) Ariapite Valley, July, 1902—
(5732) Trinidad, B. M.— (5733) Trinidad,
B. M.— (5734) Trinidad, F. Shade, Nov. 1920,
B. M.-(5735) Trinidad, B. M.-(5736) Trini-
dad, B. M.
Disposition of type material.— The Depart-
ment of Tropical Research, New York Zoo-
logical Society, retains the two paratypes in
coitu (5718). Those paratypes listed above with
abbreviation “B.M.” are specimens borrowed
from the British Museum (Natural History)
and have been returned to England. Paratype
5723, female, is in the United States National
Museum collection. The holotype, allotype and
the remaining paratypes are in the American
Museum of Natural History, New York.
Macrocneme vittata Walker
(PI. Ill, fig. 30)
Macrocneme v/rtota Walker, 1854: 249.
Macrocneme vittata, Hampson, 1898: 249, PI.
XII, fig. 23.
Macrocneme nigritarsis, aberration 1, Hampson,
1898: 326.
Macrocneme caurensis Klages, 1906: 540 (after
Hampson: 1914: 207).
Macrocneme v/ttatfl,Draudt in Seitz: 1915: 105,
fig. 17d.
Macrocneme nigritarsis, aberration trinitatensis.
Strand, 1917: 84.
Macrocneme vittata, Forbes, 1939: 127, 133.
This species has not been reported by Kaye &
Lamont. They did, however, list it as a synonym
of eacus (Stoll). Macrocneme eacus does not
seem to have been found in Trinidad. It is a small
Macrocneme with a suffused bright blue fore-
wing in contrast to vittata which has definite if
variable iridescent blue marks. The latter seems
closely related to alesa Druce, which differs in
having more green than vittata. The costal green
of vittata extends but little beyond the discal cell
from the base, whereas in alesa it runs to near
the termen. The blue streak on the anal vein ex-
tends beyond the middle of the wing in alesa,
but in vittata from Trinidad this streak in the
1957]
Fleming: Ctenuchidae (Moths) of Trinidad, B.W.l.
ni
specimen with it most enlarged is still only half
the length of the discal cell. This species will key
to alesa in Hampson (1898: 322) since in the
key vittata is separated on the basis of having
green only in and above the cell, which is con-
trary to what Hampson states in his description
(1898: 325). Forbes (1939: 127) similarly has
vittata restricted to blue in costal half or less.
The amount of blue in our specimens is variable
in the anal region of the forewing; from only a
spot near the base of the wing to streaks above
and below the anal vein reaching to approxi-
mately the end of the discal cell, above the anal
vein (origin of vein Cu2) and half the length of
the discal cell below the anal vein. The holotype
of alesa is a female from Bolivia and the holo-
type of vittata from Para, Brazil.
Hampson (1898: 326) on the basis of one
female specimen identified as nigritarsia, de-
scribed an aberration from Trinidad distin-
guished by having white sublateral spots on the
four medial segments of the abdomen. Hampson
did not mention in his description that this speci-
men has some white on the tarsi, which has been
seen by Kenedy. Later Strand (1917: 84) named
this aberration of Hampson trinitatensis. Hamp-
son’s female specimen can hardly be anything
else than a normal vittata female, and thus,
since the Trinidad locality record for nigritarsis
appears to be dependent on this single specimen,
nigritarsis cannot be considered as having been
taken in Trinidad.
Material.— ffimQ specimens (5 males and 4
females). Thirty specimens (14 males and 16
females) in the British Museum (Natural His-
tory). One from Caparo, four from Ariapite
Valley and the remainder labelled only Trinidad.
Range.— Amazon^ and Venezuela (caurensis) .
Calonotos Hubner
Key to the Trinidad Species of Calonotos
1. Wings without hyaline spots, .helymus
Wings with hyaline spots 2
2. Abdomen cupreous with black
stripes tiburtus
Abdomen iridescent green with
black stripes 3
3. Basal segment of abdomen
black with a pair of subdorsal
white spots and lateral hoods
white craneae
Basal segments of abdomen
green without a pair of white
spots, but with dorsal black
stripes and lateral white
hoods tripunctatus
Calonotos helymus (Cramer)
Sphinx helymus Cramer, 1775 : 1, pi. 2, figs. D,
E.
Glaucopis aterrima Sepp, 1848: p. 17, pi. 97.
Calonotos helymus, Hampson, 1898: 335.
Calonotus helymus, Draudt in Seitz, 1915: 109,
fig. 18a.
Calonotus helymus, Kaye & Lamont, 1927: 7.
This species was collected at Palmiste by
Lamont. We have not collected it in the Arima
Valley as yet. Hampson in his key (1898: 333)
states, “Wings without hyaline spots,” which
separates this species from all other Calonotos
collected to date from Trinidad. Draudt’s figure
in Seitz (1915: 18a) has a single hyaline spot in
the discal cell area of the forewing, although his
text states that the wings are unspotted.
Range.— French and British Guiana.
Calonotos tiburtus (Cramer)
(PI. Ill, fig. 32)
Sphinx tiburtus Cramer, 1780: pi. 237C.
Calonotos tiburtus, Hampson, 1898: 333, fig.
154.
Calonotus tiburtus, Draudt in Seitz, 1915: 108,
figs. 17i 3 & $.
Calonotus tiburtus, Kaye & Lamont, 1927: 6.
The bright, metallic coppery stripes on the
abdomen serve to distinguish this species from
other Calonotos of Trinidad. We have found the
species quite common flying in the daytime in
forested parts of the Nariva Swamp south of
Brigand Hill. Male genitalia with a pair of lobe-
like processes at base of uncus. Uncus curved
ventrally and tapering abruptly to a point at
distal end. Dorsal edge of harpe sharply curved
ventrally at distal end to form a large ventral
process and from inner edge of dorsal edge of
harpe a smaller terminal process. From near the
base of the harpe on the ventral edge a long
slender process. Scoup (juxta?) curved dorsally
at end and terminating in a thorn-like spine pre-
ceded by a pair of double-pointed thorns.
Aedeagus terminating in two long but unequal
spines.
Strand (1915: 25), on the basis of a male and
two females collected in Trinidad, named a
local form trinidadensis. The description simply
states “Without white spots on the palpal base
and on the thorax. Wing expanse 41, length of
forewing 20 mm. female.”
Materials.— Two male specimens were col-
lected on May 5 and June 17 at Simla.
Range.— Costa Rica to Surinam.
128
Zoologica: New York Zoological Society
[42: 10
Calonotos cmnoae, new species
(PI. I, fig. 5; PI. II, fig. 5)
Length of forewing of males and females
average 19-20 mm. Females average but slightly
larger than males.
Antennae of male bipectinate with the distal
end of each pectination dilated and bristled. The
pectinations on each side of the antennal shaft
subequal. In the female the antennae are similar,
but the pectinations are shorter than in the male
and the pectinations on the inside of the antennal
shaft of the female are very short in comparison
with the pectinations on the outer side of the
antennal shaft. Antennae in both sexes with the
pectinations decreasing in size towards the apex
of the shaft. The dorsal side of the distal end of
the antennal shaft white (approximately the
distal fifteen segments) but with the terminal
one or two segments brown.
Palpi normal in shape and position for the
genus. The fan-shaped tuft on the base of the
first segment of the palpi concolorous brown, as
well as the three palpal segments.
Front of head blackish-brown with two white
spots on upper lateral edges below antennae.
Vertex of head, collar, tegulae, thorax and legs
concolorous dark brown. A small amount of
white on the distal end of the hindcoxae in both
sexes but smaller and less conspicuous in the
female. In some lights there is a slight bluish-
green reflection from the dark brown of the
various structures mentioned above.
Forewings blackish-brown. Typically with
three white spots, one elongate spot below the
middle of the discal cell, a subspherical spot be-
yond the cell bordering on the discoidal veins
proximally and between vein Mi and vein Ms
and a third spot between veins Cui and Cus.
However, this last-mentioned spot is always
smaller in the males than in the females and
may be absent, although usually a few white
scales persist. In addition, in one female this
same spot is so enlarged as to extend half way
between veins Cus and Ms. A short iridescent
blue sub-basal streak on upper edge of basal
third of cell but not extending to base of wing. In
two males and one female there is a short iri-
descent blue steak within the basal part of the
discal cell.
Hindwing concolorous black-brown with one
white hyaline spot beyond the cell.
Underside of forewing concolorous dark
brown except for the inner margin below the
anal vein which is light brown. The underside of
the hindwing concolorous dark brown with iri-
descent blue-green patches above and within
the discal cell. The whitish-hyaline spots on the
upper side of the wings are present on the under-
side of the wings.
Abdomen shining iridescent, silvery blue with
two longitudinal subdorsal black-in-brown
stripes. The basal segment black-brown with two
gray or whitish subdorsal spots. These spots are
seldom conspicuous and may be all but absent.
Two lateral white spots on bullae which are
larger than the subdorsal spots. A narrow black-
ish-brown lateral stripe with a very narrow stripe
below of iridescent silvery blue ground color.
This silvery blue stripe diminishes and finally
disappears on the terminal segments of the ab-
domen. The ventral surface of the abdomen
blackish-brown with a midventral white stripe.
This white stripe is always present but is stronger
in some specimens than others.
Male genitalia massive and symmetrical. A
pair of appendages arises cephalad and dorsad
of the uncus. These processes extend caudally
beyond the uncus and have a lobe-like structure
subventrally at their bases. The base of the
uncus is in the plane of the tegumen, but the
uncus narrows and bends ventrally at right
angles for a distance longer than its horizontal
length. The harpe with two processes. One
process long and slender with long hair on its
ventral edge produced from the ventral edge of
the harpe. The ventral edge of the harpe nar-
rows abruptly after this process to produce just
before the terminus of the harpe a small slender
process directed ventrally. Scoup (juxta ?) long,
broad and massive, terminating in two long
processes bent at right angles in a dorsal direc-
tion.
Most closely related to triplagus Hampson
and chalcipleurus Hampson. In both of these
species, as in craneae, the iridescent blue mid-
dorsal line commences at the anterior edge of
the second abdominal segment rather than at the
anterior edge of the first abdominal segment
which is characteristic of tripunctatus. The wing
expanse of the holotype of triplagus is 38 mm.
as against 42-45 mm. of craneae. The collar of
triplagus has paired white spots while craneae is
concolorous black-brown. The dorsum of the
thorax is spotted metallic green in triplagus but
is immaculate black-brown in craneae. The
coxae and extremities of the femora are spotted
white in triplagus and unspotted in craneae. The
hyaline white on the wings of triplagus and
craneae appear to be similar. The type locality
of triplagus is Manaos, Brazil. The legs and
thorax are also spotted white in chalcipleurus
and the dorsum of the thorax with golden green
spots, contrary to craneae in which these areas
are black-brown. The pectinations of the an-
tennae appear to be longer in chalcipleurus than
in craneae. The ventrum of the abdomen with a
series of white spots in chalcipleurus, whereas in
craneae it has a ventral white stripe. Hampson
1957]
Fleming: Ctenuchidae (Moths) of Trinidad, B.W.I.
129
gives the wing expanse of chalcipleurus as 46
mm., which is considerably larger than craneae.
I take pleasure in naming this species after
Jocelyn Crane, Assistant Director of the De-
partment of Tropical Research, New York Zoo-
logical Society.
All of the type material was collected at Simla,
Arima Valley, Trinidad. Holotype, male. Cata-
log No. 5656, I-VI; allotype, female (5657)
25-IV; paratypes, 29 males, (5658) 3-1, (5659)
7-1 (5660) ll-I, (5661) ll-I, (5662) 12-III,
(5663) 27-m, (5664) 2-IV, (5665) 2-IV,
(5666) 3-IV, (5667) 6-IV, (5668) 8-IV,
(5669) 16-IV, (5670) 18-IV, (5671) 25-IV,
(5672) 25-IV, (5673) 27-IV, (5674) 29-IV,
(5675) 1-V, (5676) 2-V, (5677) 4-V (5678)
7-V, (5679) 8-V, (5680) 10-V, (5681) 12-V,
(5682) 12-V, (5683) 13-V, (5684) 13-V,
(5685) 28-V, (5686) 1-VI; 23 females, (5687)
7-1 (5688) 5-II, (5689) 13-11, (5690) 20-11,
(5691) 27-11, (5692) 28-11, (5693) 2-III,
(5694) 6-III, (5695) 7-III, (5696) 14-III,
(5697) 20-III, (5698) 27-III, (5699) 27-III,
(56100) 29-m, (56101) 4-IV, (56102) 19-IV,
(56103) 27-IV, (56104) 3-V, (56105) 4-V,
(56106) 6-V, (56107) 16-V, (56108) 28-V,
(56109) 1-VI.
Disposition of type material.— The Depart-
ment of Tropical Research, New York Zoologi-
cal Society, will retain four paratypes. Catalog
Nos. 5662, 5669, 5699 and 56108. Paratypes
with Catalog Nos. 5683 and 56106 are in the
British Museum (Natural History) collections
and paratypes with Catalog Nos. 5682 and
56109 are in the United States National Museum
collections. The holotype, allotype and remain-
ing paratypes are in the American Museum of
Natural History, New York.
Calonotos tripunctatus Druce
(PI. Ill, fig. 33)
Calonotos tripunctatus TiTMCg, 1898: 401.
Calonotos tripunctatus, Hampson, 1898: 335
(in part), pi. XII, fig. 7.
Calonotus tripunctatus, Draudt in Seitz, 1915:
109, fig. 18a.
Calonotus tripunctatus, Kaye & Lamont, 1927:
7.
Superficially similar to craneae but may be
easily separated by the dorsal green band in
tripunctatus running to the thorax and no white
points on either side of this line on the basal
segment of the abdomen.
Kaye & Lamont synonymized chalcipleurus
Hampson under tripunctatus, and while I have
not seen chalcipleurus, it appears to be a valid
species. First of all the basal abdominal segment
of chalcipleurus is black-brown as in craneae
and tiburtus and does not have the dorsal silvery
blue that is present on tripunctatus. A pair of
subdorsal white spots above the bullae on the
basal abdominal segment of chalcipleurus which
is absent in tripunctatus. The thorax and legs of
tripunctatus are black-brown, whereas in chal-
cipleurus the legs and thorax are spotted white
with the dorsum of the thorax with metallic
green spots. The type locality of chalcipleurus
is Aroa, Venezuela.
Male genitalia with large flat lobes on base
of uncus. Uncus curved ventrally with distal end
inflated but with acuminate terminus. Harpe
with ventral process long and slender and dorsal
process with a broad ventral spine near bifurca-
tion of dorsal and ventral harpal processes and
a long finger-like process directed caudally at
approximately the mid-point. Terminus of the
dorsal process blunt. Scoup (juxta ?) with a
single small spine at caudal end and directed
laterally. Caudal end of aedeagus with long
process at one side.
One female collected on May 14 with a Cata-
log Number 5491 is provisionally placed in this
species. It has only one white spot on the fore-
wing but otherwise appears identical.
Material.— Tvitniy-one specimens (7 males
and 14 females).
Holotype from Trinidad. Reported
from St. Vincent and Venezuela.
130
Zoologica: New York Zoological Society
[42: 10: 1957]
EXPLANATION OF THE PLATES
Plate I
Fig. 1. Pseudosphex kenedyae, dorsolateral view
of male genitalia.
Fig. 2. Pheia beebei, lateral view of male genitalia.
Fig. 3. Saurita arimensis, dorsal view of male
genitalia.
Fig. 4. Macrocneme spinivalva, dorsal view of
male genitalia.
Fig. 5. Calonotos craneae, lateral view of male
genitalia.
Plate II
Fig. 1. Pseudosphex kenedyae, new species. Holo-
type.
Fig. 2. Pheia beebei, new species. Holotype.
Fig. 3. Saurita arimensis, new species. Holotype.
Fig. 4. Macrocneme spinivalva, new species. Hol-
otype.
Fig. 5. Calonotos craneae, new species. Holo-
type.
Plate III
The following figures are of species collected at
Simla, Arima Valley, Trinidad, except for Cosmo-
soma achemon collected at St. Augustine, Trinidad.
Fig. 1. Pleurosoma trinitatis 2.
Fig. 2. Sphecops aurantiipes $.
Fig. 3. Phoenicoprocta vacillans (nigropeltata) $.
Fig. 4. Phoenicoprocta vacillans 2-
Fig. 5. Pheonicoprocta vacillans 2.
Fig. 6.
Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 11.
Fig. 12.
Fig. 13.
Fig. 14.
Fig. 15.
Fig. 16.
Fig. 17.
Fig. 18.
Fig. 19.
Fig. 20.
Fig. 21.
Fig. 22.
Fig. 23.
Fig. 24.
Fig. 25.
Fig. 26.
Fig. 27.
Fig. 28.
Fig. 29.
Fig. 30.
Fig. 31.
Fig. 32.
Fig. 33.
Loxophlebia bisigna
Loxophlebia postflavia $.
Mesothen endoleuca $.
Mesothen pyrrha $.
Chrostosoma viridipunctatum $.
Cosmosoma rubriscapulae 2.
Cosmosoma subflamma subflamma fi.
Cosmosoma melathoracia $.
Cosmosoma anoxanthia
Cosmosoma achemon $.
Cosmosoma klagesi
Dixophlebia holophaea
Pseudomya melanthus $.
Pseudomya melanthus 2-
Rhynchopyga flavicollis $.
Saurita cassandra
Saurita clusia $.
Saurita clusia 2*
Saurita afflict a
Saurita afflicta 2-
Dycladia basimacula $.
Histiaea meldolae $.
Histiaea cepheus $.
Macrocneme plumbea $.
Macrocneme vittata $.
Macrocneme thyra intacta $■
Calonotos tiburtus 2.
Calonotos tripunctatus $.
FLEMING
PLATE 1
FIG. 1
FIG. 3
FIG. 4
FIG. 5
THE CTENUCHIDAE (MOTHS) OF TRINIDAD, B.W.I.
PART I. EUCHROMIINAE
FLEMING
PLATE II
FIG. 2
FIG. 3
FIG. 4
FIG. 5
THE CTENUCHIDAE (MOTHS) OF TRINIDAD, B.W.L
PART I. EUCHROMIINAE
FLEMING
PLATE III
PART I. EUCHROMIINAE
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200L0GICA
SCIENTIFIC CONTRIBUTIONS OF THE
NEW YORK ZOOLOGICAL SOCIETY
VOLUME 42 • PART 4 • DECEMBER 31, 1957 • NUMBERS 11 TO 14
PUBLISHED BY THE SOCIETY
The ZOOLOGICAL PARK, New York
Contents
PAGE
Part 4. December 31, 1957
11. The Ability of the Saprolegniaceae to Parasitize Platyfish. By Helen S.
Vishniac & R. F. Nigrelli. Plate 1 131
12. Imaginal Behavior in Butterflies of the Family HeUconiidae: Changing
Social Patterns and Irrelevant Actions. By Jocelyn Crane. Plate I.. . . 135
13. Habits, Palatabihty and Mimicry in Thirteen Ctenuchid Moth Species from
Trinidad, B.W.I. By William Beebe & Rosemary Kenedy. Plates I & II. 147
14. Serological Relationships among Members of the Order Carnivora. By
Ludwig K. Pauly & Harold R. Wolfe 159
Index to Volume 42 167
11
The Ability of the Saprolegniaceae to Parasitize Platyfish'
Helen S. Vishniac & R. F. Nigrelli
Department of Microbiology, Yale University, and
New York Aquarium, N. Y. Zoological Society
(Plate I)
A NUMBER of species of fungi belonging
to the Saprolegniaceae have been iso-
^ lated from naturally occurring infec-
tions in fish and other aquatic animals (Table 1 ) .
The list is so diverse, from a taxonomic point of
view, as to suggest that any saprolegniaceous
fungus might be capable of parasitism under ap-
propriate conditions. The experiments reported
here were undertaken to determine whether
genera and species not previously reported to be
parasitic, as well as saprophytic isolates of species
known as parasites, could infect platyfish under
conditions which allowed infection by two fungi
isolated from naturally occurring infections. The
platyfish is known to be susceptible to Sapro-
legnia (Gordon, 1936). However, there is no
evidence of host specificity for these fungi even
though there is variation in the susceptibility of
different fish in captivity (Tiffney, 1939a) as well
as in nature. Tiffney (1939a) reported that
Saprolegnia parasitica can attack at least 16
species of fishes and 2 species of amphibians.
Saprolegnia parasitica also attacks aquatic rep-
tiles both in nature and in captivity. Other species
have been reported in crustaceans (Atkins,
1954; Hohnk & Vallin, 1953; Prowse, 1954;
Smith, 1940), lamellibranchs (Atkins, 1954),
1 It is a pleasure to acknowledge our debt to Dr.
Myron Gordon and the Genetics Laboratory of the
New York Zoological Society for the fish used in these
experiments. We are indebted to Dr. J. R. Raper, Dr.
A. W. Ziegler and Dr. L. Shanor for the isolates desig-
nated by their initials. The experimental infections were
made at Haskins Laboratories, New York City, in the
laboratory of Dr. S. H. Hutner, for whose interest we
are grateful. This study was supported in part by re-
search grant E 1179 from the National Institute of
Allergy and Infectious Diseases of the National Insti-
tutes of Health, U. S. Public Health Sendee.
larval mosquitoes (Rioux & Achard, 1956) and
rotifers (Valkanov, 1931).
Mycosis caused by Saprolegniaceae is a disease
of temperate and tropical freshwater and possibly
brackish water fishes. Those affecting temperate
water fishes, e.g. carp, have been recognized in
Central Europe since the Middle Ages. The num-
ber of fish species susceptible to infections by
Saprolegniaceae has never been properly re-
corded. From more than 25 years’ experience
with the diseases of fish, it can be stated without
exaggeration that most freshwater fishes are sus-
ceptible to infection by some species of Sapro-
legniaceae, especially under tank and hatchery
conditions and following injury of the skin or
gills. In fishes, primary infections occur in in-
dividuals that have lesions caused by injury.
Lesions caused by protozoan and metazoan
parasites are the loci of secondary infections. At
the New York Aquarium, more than 45 deaths
were reported in a single year as being caused by
“Saprolegnia” (Nigrelli, 1943). A re-check of
the Aquarium’s records for 1939, 1940 and 1941
showed that 24 species of temperate and tropical
freshwater fishes, 4 species of urodeles and 4
species of turtles died from such infections.
Saprolegnia-VikQ infections were also found in
the feet of alligators which had developed ulcers
from other causes. No efforts were made at the
time to determine which species of Sapro-
legniaceae were involved in these deaths.
The results of our experiments indicate that
although the family Saprolegniaceae is primarily
saprophytic in nature, the potentiality of para-
sitic existence is a familial characteristic. More-
over, imder our conditions, at least, the infecting
fungus can be a primary invader.
131
132
Zoologica: New York Zoological Society
[42: 11
Table 1. Saprolegniaceae Occurring as
Parasites of Aquatic Animals
Fungus
Reference
Achlya flagellata
see Tiffney, 1939b
A. polyandra
see Tiffney, 1939b
A. prolifera
see Tiffney, 1939a, b
A. racemosa
see Tiffney, 1939b
Achlya sp.
Tiffney, 1939b
Aphanomyces laevis
Smith, 1940
Ap. astaci^
see Prowse, 1954
Ap. daphniae^
Prowse, 1954
Ap. hydatinae^
Valkanov, 1931
Ap. ovidestruens'-
see Prowse, 1954
Dictyuchus monosporus Tiffney, 1939b
Leptolegnia baltica
Hohnk & Vallin, 1953
L. caudata
see Coker, 1923
L. marina
Atkins, 1954
Saprolegnia diclina
Rioux & Achard, 1956
S. ferax
see Tiffney, 1939a, b
S. mixta
see Tiffney, 1939b
S. monoica
see Tiffney, 1939a, b
S. parasitica
see Tiffney, 1939a, b
S. torulosa
see Tiffney, 1939b
1 If these are indeed valid species.
Materials and Methods
The 19 isolates of saprolegniaceous fungi used
are listed in Table 2. Brevilegnia unisperma and
Dictyuchus monosporus were originally included
in this group, but were omitted when they failed
to produce zoospores under the conditions of
the experiments, since infection in nature is
normally accomplished by zoospores (Tiffney,
1939a). Two of the fungi listed were isolated
from naturally infected aquarium fish: Achlya
sp. from an electric catfish (Malapterurus elec-
tricus) and Saprolegnia parasitica from the mos-
quito fish (Gambusia affinis).
The Achlya sp. was similar to the sterile
Achlya described by Tiffney (1939b) from the
shell of Chelydra serpentina, except in the ap-
pearance of the “chlamydospores.” In our isolate,
these structures were spherical to ovoid, never
clavate. They were formed first terminally, later
on racemose branches or less frequently in basi-
petal chains. These “chlamydospores” were
shown in our isolate to be oogonia. When mats
(grown in pure culture) of Achlya sp. were
placed in a petri dish in proximity to mats of
male (E247) and female (355) thalli of A. bi-
sexualis, abundant antheridial initials formed on
E247 within three to four hours. After 24 hours,
well differentiated oospheres could be seen in the
“chlamydospores” of Achlya sp. A few eccentric
oospores maturing in the oogonia of Achlya sp.
attested to the completion of the mating reaction
with E247. There was no reaction between our
Achlya sp. and the female isolate 355. This fun-
gus cannot, however, be considered a female
isolate of A. bisexualis or of any other described
heterothallic species of Achlya. Not only were
the majority of oospores aborted in the mating
just described, but the characteristic of produc-
ing oogonial initials in the absence of hormonal
stimuli provided by the presence of antheridial
initials is present in no other heterothallic Achlya
(see Raper, 1955). Since specific distinctions in j
this family of fungi are made largely on the basis j
of characteristics of sexual reproduction, we con-
sider it inadvisable to formally describe this
species in the absence of a male isolate giving
a normal complete mating reaction.
Table 2. The Ability of Various
Saprolegniaceae to Infect Platyfish
^ _ Uninfected Fish
Infected Fish
J7UU^US JTiCSiCUL
Alive \
Died
Alive 1
Died
None
2
2
0
0
None
4
0
0
0
Achlya sp.
1
0
1
2
A. ambisexualis
E87 S (J.R.R.)
0
0
0
4
A. ambisexualis
302 $ (J.R.R.)
0
0
0
4
A. bisexualis
BIM $ (J.R.R.)
0
0
0
4
A. bisexualis
355 $ (J.R.R.)
0
0
0
4
A. glomerate
(A.W.Z.)
2
2
0
0
A. Klebsiana (L.S.)
0
0
0
4
A. Sparrowii^
0
0
0
4
A phanomyces
laevis (A.W.Z.)
0
0
0
4
Calyptralegnia
achlyoides (L.S.)
0
0
2
2
Isoachlya
monilifera
3
0
0
1
Protoachlya
paradoxa (L.S.)
0
1
0
3
Saprolegnia
delica (L.S.)
0
0
0
4
S. ferax
0
0
0
4
S. megasperma^
0
1
0
3
S. mixta
0
0
0
4
S. parasitica
1
0
0
3
Thraustotheca
clavata
0
0
0
4
T. primoachlydr
(A.W.Z.)
0
0
2
2
1 Johnson (1956) considers this species to be synony-
mous with A. racemosa Hildebrand. Since A. racemose
has centric oospores while A. Sparrowii has subcentric
oospores (a distinction which Johnson considers, in the
same study, to be of subgeneric rank), we prefer to
retain A. Sparrowii.
2 Two broods of young were born in the course of
this experiment. Normal young (9 and 5) remained
healthy; premature fry (i.e. with visible yolk sac) be-
came parasitized and died (5 and 3). It was the mother
of one of these broods that died without becoming
parasitized by Saprolegnia megasperma.
1957]
Vishniac & Nigrelli: Ability of Saprolegniaceae to Parasitize Platyfish
133
The Mexican platyfish, Xiphophorus macu-
latus, was used as host in these experiments.
The conditions chosen for these experiments
were determined, by means of preliminary ex-
periments, to provide the greatest chance of in-
fection by the two parasitic isolates (Achlya sp.
and Saprolegnia parasitica) together with the
lowest mortality from causes other than induced
fungal infection. Pyrex kitchen trays (ca. 8"X
12"X2") were washed thoroughly, steamed,
filled with 1.5 liters of tap water, and placed on
an illuminated bench at room temperature (ap-
proximately 20° during the course of these ex-
periments). An excess of fish food was then
placed in them and inoculated with a pure cul-
ture of a fungus. When the growth of the fungus
on the fish food was producing an abundance of
zoospores, four platyfish, roughly evenly dis-
tributed as to size and sex, were placed in each
tray, together with a few strands of Nitella. Just
before exposure to the fungi, these fish were in-
jured by scraping the scales from an area ap-
proximately 2X2 mm. on one side of the caudal
peduncle. Uninjured fish were not attacked; fish
injured by mere rubbing or nicking of the
caudal peduncle were not consistently attacked.
The fish were then observed until death, at which
time they were removed from the tray, examined
to verify the identity of the fungus and preserved
in formalin (Plate I) , or until the disappearance
of the fungus from the tray.
Paraffin sections of the diseased peduncle were
prepared and stained with haematoxylin-eosin
and with Masson’s trichrome stain; whole
mounts of the skin and scales showing the my-
celia were treated with a modification of Mal-
lory’s method in which the following stains were
used: Harris’ haematoxylin, Phloxin B and Stir-
ling’s anilin crystal violet. The tissues were then
then treated with Gram’s iodine and differen-
tiated with several changes of anilin oil until no
more color was removed. The stained material
was then passed through several changes of
xylene and mounted in Permount. With this
method, the mycelia stained blue and the spo-
rangia red and the extent of the infection was
followed with ease.
Results
The results of these experiments are given in
Table 2. It is evident that under the conditions
which permit infection by the two parasitic
isolates, nearly every saprolegniaceous fungus
used can attack, and usually kill, platyfish. The
two apparent exceptions, Achlya glomerata and
Isoachlya monilifera, did not in fact present
quite the same conditions as the parasitic isolates.
Achlya glomerata grew sparsely, although what
growth there was produced zoospores. Isoachlya
monilifera grew moderately well before the ad-
dition of the fish, which promptly ate up the
mycelium, thus greatly reducing their chance of
infection. Consumption of mats of living my-
celium of Saprolegnia parasitica by fish which
suffered no harm thereby has been noted by
Tiffney (1939a). The fungus appearing on in-
fected fish was in every case the species with
which the tray had been inoculated.
The first signs of infection always appeared at
the injured area on the caudal peduncle in the
form of a tuft of hyphae. Later, in fatal infec-
tions, hyphae often emerged from the gills and
mouth, and in tufts over the body, or the entire
fish became covered with the fuzzy growth of
the fungus.
The pathological lesions were more or less
similar, varying only in degree. In relatively
light infections, the mycelia penetrated the epi-
thelium of the scales and skin with some necrosis
of the involved areas. In heavy infections, the
epithelium was often sloughed and the scales and
fin rays were softened or completely destroyed.
The growth penetrated the deeper tissues, the
hyphae often passing into the muscle bundles and
resulting in hyalinization or complete destruc-
tion. Macrophages filled with melanin, cellular
debris and blood cells, together with lympho-
cytes, were invariably found massed in the areas
of the mycelial growth. Inflammatory reaction of
varying intensity occurred in regions immediately
adjacent to the infection. Surprising as it may
seem, little or no bacterial infection was present
in any of the sections studied, indicating that the
tissue destruction was due almost exclusively to
the fungi.
Discussion and Conclusions
The criteria listed by Henle in 1840 for es-
tablishing a causal relationship between an
organism and a disease in its putative host begin
with the isolation of the organism from cases of
the disease. We have shown that given a set of
conditions which allow infection by two sapro-
legniaceous fungi isolated from diseased fish, all
other saprolegniaceous fungi for which the same
conditions (including abundant sporulation)
could be provided would infect platyfish. Some
of the species used, although not isolated from
diseased animals in this instance, have pre-
viously been reported to be parasitic, viz.
Aphanomyces laevis, Saprolegnia ferax and S.
mixta. The remaining 12 species have not been
so reported. Since from these results and pre-
vious reports at least 27 species in 10 genera of
the Saprolegniaceae have been found capable of
attacking animal hosts, we may conclude that
134
Zoologica: New York Zoological Society
[42: 11:1957]
potential parasitism is a familial characteristic.
But from the ecological point of view the 12
species not known to be involved in natural in-
fections are not parasites. There is, to be sure,
a very marked difference in frequency of oc-
currence of reported parasites. Saprolegnia
parasitica is universally reported as most com-
mon; other species are reported in frequencies
ranging down to only a single established case.
It would hardly be surprising if continued search
should disclose that the forms we have desig-
nated as potential parasites are actually parasitic
in nature. The reasons for the rarity or absence
of the parasitic habit in some species of this
family are probably to be found in their specific
ecology. A successful parasite in nature must be
abundantly sporulating under the conditions of
temperature, etc., that exist when and where
susceptible animals appear. While the report of
Coker (1923) on seasonal occurrence of sapro-
legniaceous fungi and the extensive studies of
Hohnk (1934, 1956), of Hohnk & Bock (1955)
and of Bock (1956) are outstanding as contribu-
tions to our knowledge of the specific ecology of
the Saprolegniaceae, more precise quantitative
techniques would be desirable before attempts
are made to correlate the production of zoospores
and infections by various species of the Sapro-
legniaceae in nature.
Summary
Eighteen of 19 isolates of saprolegniaceous
fungi, including 16 species belonging to 7 genera,
infected platyfish having a standardized wound
on the caudal peduncle. The infection usually
resulted in the death of the fish. Histological ex-
amination indicated that tissue destruction was
due almost exclusively to the infecting fungus.
Literature Cited
Atkins, D.
1954. Further notes on a marine member of the
Saprolegniaceae, Leptolegnia marina N.
Sp., infecting certain invertebrates. J. Mar.
Biol. Ass. U. K., 33: 613-625.
Bock, K. J.
1956. Zur Okologie und Systematik saprophy-
tischer Wasserpilze aus dem Silbersee bei
Bremerhaven. Veroff. Inst. Meeresforsch.
in Bremerhaven, 4: 25-44.
Coker, W. C.
1923. The Saprolegniaceae. U. of North Caro-
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Gordon, M.
1936. Fishes, Beware the Fungus. Nature Maga-
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Hohnk, W.
1934. Saprolegniales und Monoblepharidales
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1955. Ein Beitrag zur Okologie der saprophy-
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Hohnk, W., & St. Vallin
1953. Epidemisches Absterben von Eurytemora,
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Johnson, T. W., Jr.
1956. The Genus Achlya. U. of Michigan Press,
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Nigrelli, R.
1943. Causes of Diseases and Death of Fishes
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Prowse, G. a.
1954. Aphanomyces daphniae Sp. Nov., Para-
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Rarer, J. R.
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and Growth. E. G. Butler, ed. (12th Sym-
posium Soc. for the Study of Development
and Growth). Princeton U. Press, Prince-
ton, N. J. Pp. 119-140.
Rioux, J.-A., & F. Achard
1956. Entomophytose mortelle a Saprolegnia
diclina Humphrey 1892 dans un elevage
d'Aedes berlandi Seguy 1921. Vie et
MUieu, 7: 326-335.
Smith, R.
1940. Studies on two strains of Aphanomyces
laevis found occurring as wound parasites
on crayfish. Mycologia, 32: 205-213.
Tiffney, W. N.
1939a. The host range of Saprolegnia parasitica.
Mycologia, 31: 310-321.
1939b. The identity of certain species of Sapro-
legniaceae parasitic to fish. J. Elisha
Mitchell Sci. Soc., 55: 134-151.
Valkanov, a.
1931. fiber Morphologie und Systematik der
rotatorien-befallenden Pilze. Arch. Pro-
tistenk., 74: 5-17.
EXPLANATION OF THE PLATE
Plate I
Fig. 1. Platyfish infected with Isoachyla moni-
lifera (upper) and Saprolegnia mega-
sperma (lower). Preserved in formalin.
2X.
VISHNIAC & NIGRELLI
PLATE I
THE ABILITY OF THE SAPROLEGNIACEAE TO PARASITIZE PLATYFISH
I
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V
12
Imaginal Behavior in Butterflies of the Family Heliconiidae:
Changing Social Patterns and Irrelevant Actions^
Jocelyn Crane
Department of Tropical Research, New York Zoological Society, New York 60, N. Y.
(Plate I)
[This paper is one of a series emanating from the
tropical Field Station of the New York Zoological
Society at Simla, Arima Valley, Trinidad, British
West Indies. The Station was founded in 1950 by
the Zoological Society’s Department of Tropical Re-
search, under the direction of Dr. William Beebe. It
comprises 200 acres in the middle of the Northern
Range, which includes large stretches of undisturbed
government forest reserves. The laboratory of the
Station is intended for research in tropical ecology
and in animal behavior. The altitude of the research
area is 500 to 1,800 feet, with an annual rainfall of
more than 100 inches.
[For further ecological details of meteorology and
biotic zones see “Introduction to the Ecology of the
Arima Valley, Trinidad, B.W.I.,” William Beebe.
(Zoologica, 1952, Vol. 37, No. 13, pp. 157-184.)]
Contents
I. Introduction 135
II. Historical Review 136
III. Materials and Methods 137
IV. Survey of Social Behavior in Six Species
of Trinidad Heliconiidae 137
V. Changes in Social Patterns with Age. .. . 138
VI. Irrelevant Actions 139
VII. Discussion 141
VIII. Summary 143
IX. References 143
I. Introduction
This contribution is one of a series on the
biology and ecology of butterflies of the
neotropical family Heliconiidae. It con-
cerns post-imaginal changes in social behavior
patterns, and various kinds of irrelevant be-
havior, including the displacement of actions
^Contribution No. 979, Department of Tropical Re-
search, New York Zoological Society.
characteristic of one pattern by actions from
another behavioral field.
The observations presented arose as unex-
pected incidental results of a comparative etho-
logical study of six species common on the island
of Trinidad, British West Indies. The publication
of these data appears to be a prerequisite to the
adequate exposition of comparative specific
characteristics and to an illumination of the
group’s phylogeny. A first paper in the series, on
Heliconius erato hydara Hewitson, has been
published in which the effect of age on behavior
and the occurrence of displacement behavior
were mentioned (Crane, 1955). Five additional
species, along with H. erato, are considered
below. These consist of Dry as Julia Julia (Fab-
ricius), Heliconius melpomene euryades Rif-
farth, H. sara rhea Cramer, H. ricini insulana
Stichel and H. isabella isabella Cramer. Details
of their behavior patterns, accounts of releasing
mechanisms and discussion of phylogenetic as-
pects are being reserved for more appropriate
inclusion in forthcoming contributions.
The risks of inaccuracy and incompleteness in
the description of butterfly behavior became ap-
parent very early in the work, particularly in
the study of courtships. The patterns seemed at
once surprisingly variable within a species, un-
expectedly similar among species and altogether
too unpredictable to be characteristic of arthro-
pods, with their high degree of dependence on
stereotypic forms of behavior.
Since the butterflies were being studied prin-
cipally in outdoor insectaries, it was thought at
first that the discrepancies might be the result of
unnatural conditions— although even if that were
so the interest of the variability of the be-
havior would scarcely have been lessened.
135
136
Zoologica: New York Zoological Society
[42: 12
Enough corroborative observations have now
been made in the field, however, to show that
the apparent eccentricities are equally charac-
teristic of free-flying butterflies. Further cor-
roboration of the natural prevalence of the types
of behavior under consideration comes from the
fact that, now that the causes are better under-
stood, the patterns shown in the insectaries are
highly predictable.
Some of the behavioral discrepancies have
proved to be due simply to age differences in the
individuals observed. The full courtship pattern
characteristic of the species, for example, is
elicited only in and by individuals between cer-
tain ages, although successful reproduction can
take place throughout a much longer portion of
imaginal life.
Other irregularities appear when, as Tin-
bergen (1952, p. 26) detoes the conditions for
displacement activities, “a strongly activated
drive is denied discharge through its own con-
summatory act(s).” However, in the present
paper the term “irrelevant behavior” (suggested
by Rand, 1943) will be employed as a more
general term than “displacement behavior.” The
latter, it seems, may be usefully restricted to the
definition given by Bastock, Morris & Moynihan
(1953, p. 25): “A displacement activity is an
activity belonging to the executive motor pat-
tern of an instinct other than the instincts
activated.”
This distinction between terms seems desir-
able since some of the butterfly actions under
consideration do not at all appear to belong to
the motor pattern of another instinct, and hence
will be simply referred to as “irrelevant actions.”
Others, which fulfil the conditions of the more
restricted definition of displacement activities,
will be so designated here and treated as a sub-
division of irrelevant actions.
My thanks go to the National Geographic
Society for a grant-in-aid, to Dr. William Beebe,
Mr. Henry Fleming and Dr. D. W. Snow for
helpful suggestions, and to Miss Barbara P.
Young for rearing numerous larvae.
II. Historical Review
The special aspects of social behavior under
consideration are little-known fields in the study
of invertebrates.
The gradual development of behavior pat-
terns in physiologically adult vertebrates has
been extensively studied, and it is well known
that changes occur in response to physiological
alterations due both to increasing age and to
seasonal causes.
Corresponding information has been gath-
ered on few invertebrates, although the Hymen-
optera include outstanding exceptions. The
sequence of changes in colony functions have
been studied in worker honeybees (Rosch,
1925), Polistes (Steiner, 1932) and in various
ants {e.g. Buckingham, 1910). Verlaine (1932)
reported differences between young and old
mason wasps in nest-repairing and provision be-
havior, that of old bees late in the season being
incomplete; this observation invalidated a con-
clusion drawn by Fabre (1879; ed. 1920) from
experiments which he performed, using aged
individuals. Pardi (1947) found that age was
one of the factors determining the status of in-
dividual Polistes females in a dominance hier-
archy. Nielsen & Nielsen (1952) reported that
the migratory period of a pierid butterfly
(Astia) was confined to a single day of the five-
day life-span. As an example of intrageneric
variation, the parasitic wasp genus (Opius) may
be cited: males of certain species cannot mate
for five days or more after emergence, although
in other species they do so early in the imaginal
period (Hagen, 1953). Finally, entomologists
would probably agree that female insects that
have recently molted into the final instar are in
general more attractive to males than are older
individuals. It will be noted, however, that use
of the inexact term “recently” is necessary.
Rockstein (1956) discussed the unreality of
a sharp boundary between the pupal and imag-
inal stages in insects, citing recent reasearch on
biochemical changes occurring after emergence
in the worker honeybee, house fly. Drosophila,
moths and the Japanese beetle. In Drosophila
there are concomitant increases in glycogen con-
tent and wing-beat frequency during the first
week of imaginal life. Although this frequency
change is not directly related to social behavior,
the phenomenon illustrates the kind of corre-
lation which may be brought to light in investi-
gations linking invertebrate physiology and be-
havior.
Studies of irrelevant actions, including dis-
placement behavior, in vertebrates are increas-
ing in number, following the pioneer work of
Lorenz, Tinbergen, Makkink, Koorlandt and
Armstrong. General accounts and references are
given by Armstrong (1950), Lorenz (1950),
Tinbergen (1951, 1952), and Bastock, Morris
& Moynihan (1953). Recent studies on particu-
lar species of birds and fish include those of
Hinde (1953), van lersel (1953), Moynihan
(1953) and Morris (1954).
The probable occurrence of equivalent be-
havior in invertebrates has apparently not been
suggested until recently. Armstrong (1950, pp.
379 ff.) summarized the situation as it appeared
at the time of his writing as follows: “Probably
displacement activities are commoner in some
other groups besides birds than is at present
1957]
Crane: Imaginal Behavior in Butterflies of the Family Heliconiidae
137
known, but they have reached their highest de-
velopment in birds ... So far as is at present
known they have a comparatively insignificant
role among insects. A thwarted solitary wasp,
Bembex rostrata, when forced to remove pebbles
repeatedly from the mouth of its burrow merely
buzzed loudly and ran around in a wide arc
(Nielsen, 1945). When experimenting with but-
terflies attracted to pieces of coloured paper
Dr. D. Use noticed movements which might
have been displacement activities (personal
communication). Possibly one of the factors re-
sponsible for the apparently slower speciation of
insects than birds (Mayr, 1942) is the greater
displacement-proneness of the latter.”
A few recent observations and comments,
however, suggest that displacement activities
may prove, after all, to be widespread among
higher arthropods. References to date appear
to be confined to the following: salticid spiders
(Crane, 1949), mantids {idem, 1952), the but-
terfly erato {idem, 1955) , Drosophila
(Bastock & Manning, 1955) and fiddler crabs,
genus Uca (Gordon, 1955, and Crane, 1957).
III. Material and Methods
The studies were all conducted in out-of-door
wire mesh insectaries in Trinidad (Crane & Flem-
ing, 1953; Crane, 1955) between 1954 and
1957. During the past two seasons a new in-
sectary, designed as were the earlier ones by
Henry Fleming, has been in operation. Con-
structed entirely of aluminum, it measures 24 X
36 feet, the dimensions of the larger of the two
earlier structures, but it is higher than its prede-
cessor, measuring 12 feet at the ridgepole. It
also has two doors with a small vestibule be-
tween, forming a baffle which has proved very
useful in preventing the escape of butterflies. A
small pond and bog have been added near one
end; they form an efficient aid in maintaining
the necessary high humidity.
The aluminum netting reflects far more heat
than does bronze mesh; it also diffuses the light
better, making it excellent for photography.
Finally, species suitable for keeping in a cage
of this size tend to bat against the netting less,
even when they have just been released into it,
than they did against the bronze netting of the
previous insectary.
For this reason, as well as because of the in-
sectary’s relative coolness, heavy vines giving
large areas of dense shade have been found to
be not only unnecessary but undesirable. Instead,
the planting is kept to several well-separated
major groups of shrubs, saplings and wild
bananas of varying degrees of height, density
and leaf size. The rest of the space is occupied
by flowering weeds and by the pond. Branches
with bromeliads attached stand upright at inter-
vals against the netting, as a further aid in main-
taining humidity and natural conditions. The
wild banana group {Heliconia) beside the bog
forms the coolest, most shady corner of the cage.
H. erato, melpomene and other shade-lovers in-
variably seek it out during the heat of the day.
In contrast, other species, such as H. isabella,
which is most active around noon, frequent the
open center of the cage around a group of Lan-
tana, Bidens and Asclepias. In this cage all of
the latter, favorite food blossoms of the heli-
coniids, thrive and are allowed to grow freely
over most of the cage. This scattering of a
natural food supply encourages a normal amount
of flight by the butterflies.
The success of the new design is attested by
the fact that all of the six species (p. 135) of
heliconiids discussed in the present paper feed,
court, mate and lay eggs. Many individuals re-
main alive, barring accidents, for one to three
and one-half months, although differences in
viability are shown. H. isabella is the most dif-
ficult to maintain, and observations on this
species are still somewhat deficient. Two locally
rare heliconiids, Philaetraea dido (Linnaeus)
and Heliconius wallacei Reakirt, have not been
successfully maintained. They live a few days
and feed, but do not “settle in” and spend most
of their active periods batting against the roof.
Presumably a higher cage is needed.
Broods from aU the species of heliconiids in-
cluded in this study were raised in the labora-
tory. The young imagoes were kept in small
cages out-of-doors until needed for observation
or testing in the large insectary. General meth-
ods are given in an earlier paper (Crane, 1955).
Table 1 gives an idea of the number of healthy
imagoes that were used in the preparation of
this study. Substandard specimens, as well as
those obesrved in the early seasons before tech-
niques were perfected and the present problems
formulated, are not included.
IV. Survey of Social Behavior in Six Species
OF Trinidad Heliconiidae
The social behavior of all six of the Trinidad
species discussed below consists of three general
types— courtship, “social chasing” and roosting.
These have already been described in some de-
tail for H. erato (Crane, 1955). Except for minor
differences they are characteristics of the other
five species as well. Although these slight specific
differences are of great potential interest from a
phylogenetic point of view, their detailed dis-
cussion belongs in subsequent papers on the
ethology of the genus.
138
Zoologica: New York Zoological Society
[42: 12
Table I. Numbers of Individuals upon which
Present Data Are Based
(From broods reared during the seasons of
1954-1957, incl.).
Species
Males
Females
Dryas julia
49
46
Heliconius melpomene
27
28
Heliconius erato
71
62
Heliconius ricini
38
27
Heliconius isabella
21
22
Heliconius sara (1957 only)
16
19
A. Courtship. Since the similarities of the
behavior within the genus are far greater than the
differences, a comparative chart (Table 2) of
the patterns of fully developed courtship will in-
dicate the trends to the extent needed for present
purposes.
It will be seen from the table that courtship
usually begins and always ends similarly in all
species, while differentiation is shown principal-
ly in the first and second stages of the second,
sedentary phase.
In brief summary, the sequence in its most
complete form is as follows:
1. Aerial Phase.
Stage 1. Nudging. A flying male approaches
a resting female from the rear. She then takes
wing, usually without his actually touching her
in any way.
Stage II. Flight. The male chases and over-
takes the female, rises above and in front of her
and fans her with the rapid vibration of his wings,
so spreading the products of his scent scales. She
then descends or is forced down to a perch.
Chases sometimes include mutual circling and
spiralling in all the species. At these times the
circling by the female is apparently always
merely the result of her temporarily successful
effort to duck out from under and behind the
flying male and rise above and in front of him;
he in turn repeats the manoeuvre and the result-
ing vertical circling may continue for several
minutes. Horizontal spiralling is less frequent
but seems to have a similar basis. In the pre-
viously published account of erato (Crane, 1955)
this flight stage, which is little developed in that
species and often omitted, was not separated
from Stage I of the Sedentary Phase below.
2. Sedentary Phase.
Stage I. Primary Fanning. The male fans
the alighted female from the front or rear, de-
pending on the species, facing in the same
direction as the female. The latter, meanwhile,
flutters her wings, also characteristically, elevates
the abdomen and, in this stage or the next, ex-
trudes the subterminal scent glands.
Stage II. Secondary Fanning. The male fans
the female from a different position from that in
Stage I, but still facing in the same direction as
she. The female meanwhile extrudes the sub-
terminal scent glands if she has not already done
so.
Stage III. Alighting and Engaging. The male
alights beside the female, either just in front of
or just behind her, moves back or forward ap-
propriately and, as she closes her wings, curves
his abdomen up between her posterior wings, en-
gages her genitalia with his harpes, and swings
around so that the two insects now face in op-
posite directions.
B. Social Chasing. Discussed first in 1955, by
Crane, in erato, “social chasing” was the term
given to social flights that are not apparently di-
rectly of a sexual nature. It was found to take
similar form in the other five species. In all,
it consists of the pursuit of males and old females
by males of any age, and of either sex by old
females. It will be discussed on pp. 141 and 142.
C. Roosting. Four of the six species roost
gregariously, namely H. erato, melpomene, ricini
and Sara. All return to the same bush or vine,
and often to the same twigs or tendrils, night
after night. Usually the perch selected is dry.
Although the four species tend to maintain sep-
arate roosts, erato, melpomene and ricini often
roost together, as do ricini and sara.
A trace of gregariousness is found in Dryas,
which sometimes hangs up for the night near
one or two others of its kind. H. isabella, how-
ever, always roosts alone. The two latter species
always hang from beneath green leaves. Roost-
ing will be further considered on p. 139.
V. Changes in Social Patterns with Age
None of the social behavior patterns sum-
marized in the foregoing section is fully ex-
pressed when the insect emerges from the chrys-
alid, and it now appears that the maturation
time for the various responses is similar in all
six species. Table 3 shows these periods. “Age”
indicates time after emergence from chrysalid.
It will be seen from the table that the full
pattern of courtship— including Stage II of the
Aerial Phase and the specifically characteristic
wing flutters of the Sedentary Phase— is evinced
by females only on their second and third days,
that is, between 24 and about 56 hours after
[DS
Species
y Phase
Stage III
M;
Female
Male
Female
Dryas jidia
Nudges
rear
ification of Stage I
Alights beside 9 head, moves
backward, curves abdomen
forward between 9 wings, en-
gages harpes, swings around
to face in opposite direction
from 9
Closes wings; lowers abdo-
men; withdraws scent glands
Heliconius
melpomene
Dryas
As in Dryas
As in Dryas
Heliconius
erato
Stage I, but lower in-
j
As in Dryas
As in Dryas
Heliconius
ricini
Dryas
Alights beside 9 abdomen,
moves forward, then pro-
ceeds as in Dryas. (Rarely
alights beside 9 head as in
Dryas).
As in Dryas
Heliconius
isabella
Dryas
As in Dryas
As in Dryas
Heliconius
Sara
Dryas
As in Dryas
As in Dryas
■ 1.
■»
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I
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t
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Table II. Basic Motion Sequence in Fully Developed Uncomplicated Courtships of Trinidad Heliconuds
Species
Aerial Phase
Sedentary Phase
Stage I
Stage n
Stage 1
Stage n
Stage III
Male
j Female
Male I Female
Male
Female
Male
Female
Male
Female
Dryos jiilia
Nudges from
rear
Takes flight
Pursues; aims to fly above Aims to back out from under
and in front of $. (Scent and rise above Eventually
scales probably in use) . alights on exposed perch.
Front fanning,
facing in same
direction as $.
All wings flattened and vi-
brated in narrow arc, the
hindwings more flattened
than forewings and vibrated
more rapidly; abdomen
raised; scent glands extruded
Brief dorso-posterior
ning; then return to
fanning
fan-
front
Intensification of Stage I
Alights beside $ head, moves
backward, curves abdomen
forward between $ wings, en-
gages harpes, swings around
to face in opposite direction
from 9
Closes wings; lowers abdo-
men; withdraws scent glands
Heliconius
melpomene
< Entire Aerial Phase essentially as in Dryas but often omitted >
As in Dryas
All wings flattened equally
and slightly vibrated; other-
wise as in Dryas
Rear fanning, then return to
front fanning
As in Dryas
As in Dryas
As in Dryas
Beliconius
erato
< Entire Aerial Phase essentially as in Dryas but often omitted >
Rear fanning
Forewings closely appressed;
hindwings flattened and
quivered; otherwise as in
Dryas
Front fanning
As in Stage I, but lower in-
tensity
As in Dryas
As in Dryas
Heliconius
ricini
^ Entire Aerial Phase essentially as in Dryas ^
As in Dryas
As in H. melpomene
Dorso-rear and rear fanning
As in Dryas
Alights beside 9 abdomen,
moves forward, then pro-
ceeds as in Dryas. (Rarely
alights beside 9 head as in
Dryas).
As in Dryas
Heliconius
isabella
< Entire Aerial Phase essentially as in Dryas >
As in Dryas
1st day: As in H. erato
2nd & 3rd days: As in Dryas
As in Dryas
As in Dryas
As in Dryas
As in Dryas
Heliconius
Sara
< Entire Aerial Phase essentially as in Dryas >
As in Dryas
As in H. melpomene, but
wings vibrated strongly
through wider arc
As in Dryas
As in Dryas
As in Dryas
As in Dryas
1957]
Crane; Imaginal Behavior in Butterflies of the Family Heliconiidae
139
Table III. Approximate Ages after Emergence
AT WHICH Various Types of Activity Occur in
Six Species of Heliconiids
Activity
Age
Flutters to ground when disturbed
1 hr.
First flight when undisturbed
2 hrs.
First feeding, rarely
6 hrs.
First feeding, usually
2nd day
Earliest copulation, female
10-30 minutes
Earliest copulation, male
3rd day
Complete courtship pattern
elicited, female
2nd & 3rd days
Complete courtship pattern
3rd day—
elicited, male
2% months
Latest copulation, unmated females
6th-8th day
Latest copulation, males (2 species)
2% months
First eggs laid
4th- 12th day
Maximum ages reached, males and
females (2 species)
'iVx months
emergence. In Isabella, it is not even possible to
see all the specLQc characteristics at one time.
During the female’s first day, the wing fiutters
of the sedentary phase are very similar to those
of erato and the aerial phase is, as usual, normal-
ly non-existent; yet by the second day the special-
ized flutter is already disintegrating into the more
generalized Dryas-type flutter (see Table 2). In
all the species after their third day the wing-
flutterings of the females are diminished pro-
gressively in intensity and characteristic form.
In males, copulation does not occur until the
third day (48 hours after emergence) and may
not take place until the fourth or fifth; these
later dates are apparently always characteristic
of H. Sara. No social activity by the males what-
ever is shown until the second day, 24 hours
after emergence, when males sometimes nudge
young females from the rear. After noon of the
second day slight chasing may occur, but this
behavior does not usually take place until the
third day.
Unlike females, males court and can mate
practically throughout life. In older males, al-
though complete and successful courtship is
swiftly elicited by second-day females, relatively
little attention is paid to older unmated or egg-
laying individuals.
Males chase each other freely throughout life,
especially in the absence of young females, ex-
cept during their first two days.
In both sexes and all species, roosting accord-
ing to the species habit begins on the second or
third night. Even in the most gregarious species
{erato, melpomene and sara), however, young
females often hang up for the night alone, unless
courting or mating has proceeded during the late
afternoon. At such times the female, too, often
roosts with the group.
VI. Irrelevant Actions
A. Males. Under certain conditions the court-
ship pattern characteristic of the various species
of heliconiids becomes atypical. Instead of court-
ship proceeding in the usual fashion to copula-
tion or, alternatively, to the point where one
partner stops responding and both go their sep-
arate ways, the male continues special behavior
that never ends in copulation. On the very rare
occasions when mating soon ensues between the
^ same partners following the first stages of this
irregular behavior, the male has returned to an
early stage of courtship, and then followed the
typical sequence.
Irrelevant actions never begin before the male
has reached Stage II of the Sedentary Phase.
In other words, he is in the final phase of fanning
above, in front of, or behind the alighted female,
his position depending on the pattern character-
istic of his species (Table 2). Sometimes the
male has reached Stage III, having alighted be-
side the female and tried unsuccessfully to attach
his harpes.
One of two major types of irrelevant behavior
follows, depending on whether the female flies
away and evades the male or whether she stays
in place.
In the first type, having lost track of the fe-
male, the male flies about at unusual speed for
up to five minutes at a time, without pausing,
feeding or making any apparent “searching”
motions (as he may do, on other occasions,
among the vegetation) . This type of behavior has
been artificially induced by the observer’s simply
picking up the female and keeping her tem-
porarily out of sight.
The second type of irrelevant behavior follows
when the female stays alighted and often appears
to the observer to be making full courting re-
sponses. The first manifestation is always poor
orientation in the fanning (PI. I, Figs. 7-8).
Normally in all the species the male faces in
exactly the same direction as the female, al-
though the longitudinal axes of the two insects
are usually more or less oblique to each other
since the male fans characteristically from down-
in-front to up-in-back. In disoriented fanning the
male may face in any direction, even backward
140
Zoologica: New York Zoological Society
[42: 12
with respect to the female, and frequently
changes the direction without any regular se-
quence.^
After disoriented fanning has continued for
any period up to five minutes in length, there are
a number of alternate sequels.
Most frequently the partners separate, either
the male or the female flying off, and neither one
thereafter evinces either excitement or special
behavior.
Ocaasionally the female takes wing and eludes
the male, whereupon the male flies about rapidly
as described under the first type of irrelevant
behavior.
2In the study of H. erato already published (Crane,
1955), Plate I, Figs. 5 and 6, are excellent examples of
disoriented fanning. A third example from the same
series is published as Plate I, Fig. 8, of the present con-
tribution. At the time these photographs were made,
the existence of this type of irrelevant behavior had not
been recognized. It is now clear that the sequence of
stiU photographs in the earlier plate could not be taken,
with present-day still camera apparatus, to illustrate a
complete, normal consumated courtship between the
same two individuals, photographed in sequence. This is
because these courtships always proceed too svsdftly to
permit recharging of the speedlamp between flash shots.
Only during the prolonged repetitions of irrelevant be-
havior can still photographs be made of a single pair
during a consecutive period of courting behavior. A mo-
tion picture camera, operated at high speed, is of course
suitable for the work.
In a common sequel to disoriented fanning,
the male settles near the female. He may be
beside her, back of her or, most frequently,
facing her. He then extends the proboscis, some-
times uncoiling and coiling it repeatedly and
with speed and force; when uncoiled it often
touches or even palpates the female’s head,
thorax, legs or, rarely, abdomen (Oane, 1955,
PI. Ill, Figs. 13-14).
Sometimes the male palpates similarly with
his antennae. In these cases the proboscis is not
uncoiled.
Once only a male Dryas, after prolonged dis-
oriented fanning, settled obliquely to the side of
the female, facing her, and alternately brushed
each of his own eyes with the palp of the cor-
responding side. The motion was repeated sev-
eral times and the performance was repeated an
hour later.
Occasionally a period of proboscis extension
or antennal palpation will be followed by re-
sumed disoriented fanning, and then a second
period of the more extreme form of irrelevant
behavior. Sometimes the male persists in alter-
nate disoriented fanning and its sequels until he
appears altogether exhausted and remains
perched motionless nearby. At these times he
cannot even be urged into flight by a sudden
close wave of the observer’s hand. Under ordi-
nary conditions such a gesture sends into the
air even butterflies that are fully accustomed to
moving human beings and insectary conditions.
Nine situations regularly elicit irrelevant be- ■
havior in courting males. As previously stated, i]
an advanced stage of courtship must have been
reached before irrelevant behavior begins.
1. Courting of an unmated female, four to
eight hours after her emergence; that is, on the
afternoon of her first day.
2. Courting of a young mated female. It is
now certain that second matings in females are
very rare, and are practically confined to indi-
viduals that have already reached egg-laying age
and are being persistently courted by young
males in the absence of young unmated females.
3. Courting of a mated female about to lay
eggs.
4. Courting of a male on his first or second
day.
5. Courting of a very young imago of either
sex, belonging to another species.
6. Loss of an unmated female, during court-
ship, through her flying out of sight.
7. Courting of females, unmated or mated,
more than three days old by males more than
about one month old.
8. Overcrowding of the insectary. This results
in somewhat indiscriminate courting activity,
apparently clearly attributable to the operation
of the principle of heterogeneous summation
and resulting in the courting of unsuitable in-
dividuals.
9. Unresponsiveness of unmated young fe-
males, or other failure in the final stages of
courtship, due to unknown causes.
B. Females. Equivalent irrelevant behavior of
females following broken-off courtships has not
been detected. However, later in life a female’s
resistance to courtship appears to change in char-
acter to such an extent that it may be termed
irrelevant behavior.
A recrudescence of courtship of egg-laying
females is usual in all species. Very rarely these
attempts end in copulation. The usual procedure
is for the male to chase the female when she is
beginning to hover around the foodplant, and to
try to force her to alight for the Sedentary Phase
in the usual fashion. At these times the female,
flying with increasing strength as she is freed
of the weight of eggs, usually manages to duck
out from under the flying male and fly well above
him. Then, instead of flying away as unrespon-
sive younger females do, she hovers, mounting
higher as he in turn tries to come on top. As
1957]
Crane: Imaginal Behavior in Butterflies of the Family Heliconiidae
141
she grows older her flight above him more and
more resembles the swift, vigorous flight of the
fanning male, in which the fore- and hindwings
are well separated (whether or not the friction
surfaces in the species concerned hold the scent
scales) . The male usually soon gives up and flies
away, without irrelevant behavior (see p. 139).
As the female grows older she chases passing
butterflies with increasing frequency, regardless
of their age or sex. Some chasing of other spe-
cies, either related or of similar size and color,
also occurs. This behavior continues even after
egg-laying has ceased and, at least in H. erato,
is strongest in those females that have completely
finished egg-laying; in that species no eggs are
laid after the age of about six weeks, while the
female’s imaginal life often extends to more than
three months. Only in the last two weeks of life
does this energetic chasing decrease.
The following special behavior has been ob-
served only in females of H. sara. It occurs only
on the two days preceding their first egg-laying
or, alternatively, when another female has al-
ready laid on the only available space. (This
species in Trinidad lays eggs in a cluster on the
terminal leaflets of Passiflora auriculata). On
these occasions sara uncoils the proboscis and
with it touches either the spot on which she
would normally lay, or the eggs already laid by
another female. This procedure never occurs
when she is actually ready to lay her own eggs,
or when no other eggs are already in place.
VII. Discussion
A number of points which have emerged from
the observations described in the preceding pages
now require comment.
It is clear that, in this group of butterflies,
there can be no accurate description of court-
ship or other social behavior that is based on a
few observations and individuals, any more than
this is possible in the case of a higher vertebrate.
The ages and past experience of each butterfly
observed, particularly females, must be known
in order properly to describe and evaluate their
activities.
A. Age. As in many birds, the patterns of
reproductive behavior are not closely linked with
successful reproduction. In the first place, the
full specific courtship pattern is unnecessary for
copulation. Before pairing with first-day females,
males omit all except Stage III of the Second
(Sedentary) Phase of courtship, and the young-
est females gives no overt responses at all. These
exceedingly simple courtships, in which prac-
tically all appetitive behavior is almost always
omitted, seem to depend predominantly on fe-
male scent, as do courtships in moths. All spe-
cific differentiation shows in courting females
later in the first afternoon and, particularly,
on the second or third days. Beyond that age,
unmated females are receptive for at least three
to six days, although their characteristic actions
decline in both intensity and specialization.
Courting of very young males by other males,
or of young individuals of other species, is un-
doubtedly due to the strong family odor of re-
cently emerged insects. It is interesting that this
odor apparently does not develop until the insect
is at least ten minutes out of the chrysalid. Once
a male H. sara actually copulated with a female
H. melpomene when the latter, aged about one
hour, was still unable to fly. Usually these two
species, which are strikingly different both in
color and, even to the human sense, in odor,
have very little attraction for each other.
It is unlikely that females, which spend the
first day practically inactive, are often found by
males during these early hours. Odor alone is
not an adequate releaser. A male, after losing
sight of a newly alighted young female that has
slipped underneath a leaf, sometimes searches
around with every appearance of vagueness and
inefficiency, and only rarely locates her once
again.
B. Social Chasing and Related Topics. It
now seems clear that all the activities referred
to in the paper on H. erato (Crane, 1955) as
“social chasing” are appetitive fragments of the
courtship pattern. They correspond to instances
in numerous other animals where, as in frogs, a
male in response to an incomplete stimulus situa-
tion embraces another male and, in the absence
of appropriate response, releases his grip.
It is possible that in the wild some species of
heliconiids maintain territories. All that can be
said now is that in this family no trace of terri-
torial behavior, or of a dominance hierarchy,
has been observed either in the insectaries or
during field observations. There is no patrolling
of definite routes, no special display that lends
itself to an interpretation of threat behavior,
no overt fighting, no individual that usually is
the pursuer in the frequent inter-male chases.
On the other hand all of the chases of one male
by another can be explained satisfactorily as
merely the chasing phase of normal courtship
which, because of the inadequacy of the sexual
situation, breaks down. The break usually occurs
after more or less mutual circling, which exactly
resembles the circling resulting from the evasion
attempts of unresponsive females (p. 138).
When two males are courting the same female.
142
Zoologica: New York Zoological Society
[42: 12
the latter often escapes in the general excitement
and the two males continue for a short time
chasing and fanning each other. Even when,
through the breakdown of the releaser sequence,
they “discover” the mistake there is no evidence
of agonistic behavior; the two simply separate
and go their ways. Rarely one or both of the
males may continue rapid flight, apparently of
an irrelevant character (p. 139).
It seems likely that the phylogenetic origin of
gregarious roosting was the continuation of
social chasing until time to hang up for the
night. This could have become flxed, through
the action of natural selection, into the stereo-
typed patterns of the species, as it came to have
protective value. Presumably the strong species
odors, which laboratory tests show to be un-
pleasant to predators, become intensified through
crowding and could be a strong deterrent to
nocturnal enemies that depend largely on the
sense of smell. These enemies are probably
chiefly reptiles and small mammals.
The increase of chasing by older females is
of particular interest. This masculine type of
activity would, in a vertebrate, be subject to a
hormonal interpretation, in which the decline
of female reproductive hormones leads to visible
effects of male hormone activity. However, in
insects, morphological sexual characters, at least,
are not glandularly controlled; rather “. . . the
sex of every part is controlled directly by the
chromosome constitution of the cells composing
it. Hence the sexual characters are unaffected by
the removal of the ovary or the testes or even
by their transplantation” (Ford, 1945, p. 192).
Schneirla (1953, pp. 677-678) , after citing refer-
ences concerning mating behavior in insects that
were castrated or otherwise sexually abnormal,
commented, “Such results suggest that factors
governing susceptibility in male and female in-
sects are not directly dependent upon testes or
ovaries, but may concern hormones already in
the blood ... the activity of some cephalic
endocrine secretion, or other physiological
agencies such as neural processes.” The present
instance, therefore, of pseudo-masculine be-
havior in aged female butterflies, is one more
instance of the desirability of cooperative studies
between physiologists and students of behavior.
C. Irrelevant Behavior. Irrelevant behavior
proves to play a large part in the apparently
normal social life of all butterflies under obser-
vation. It is not confined to species in captivity,
since it has also been frequently observed in the
wild. In fact, if only a few pairs of butterflies of
a given species are observed in the Sedentary
Phase in the field, it is far more likely that the
observer witnesses irrelevant behavior of some
kind than a specifically characteristic courtship.
The various phases of irrelevant behavior and
the situations which produce it have proved to
be similar in all six species. In males, it is known
certainly to take place only following an ad-
vanced stage of an incomplete courtship that
is directed toward an unsuitable individual. It
occurs in females after their most receptive
period (see preceding section). Experiments to
determine whether irrelevant behavior takes
place in hungry individuals that are prevented
from reaching visible and olfactorily detectable
food have so far been inconclusive.
The instances of irrelevant behavior described
are in completely different categories from “so-
cial chasing” since they are in no sense merely
unfinished portions of the regular reproductive
or roosting patterns. Like “social chasing,” how-
ever, they never end in copulation.
Two of them, irrelevant proboscis-uncoiling
and excessive, rapid, undirected flight, can ap-
propriately be termed displacement activities.
Occurring when the strongly activated sex drive
is denied expression through its own consum-
matory act, the motions are clearly associated
with other patterns of behavior, namely feeding
and flight.
The once-observed eye-rubbing with the palps
may be equivalent to the apparently displaced
cleaning motions described in mantids (Crane,
1952), Drosophila (Bastock & Manning, 1955)
and ocypodid crabs (Crane, 1957).
The two remaining types of irrelevant be-
havior in courting male heliconiids cannot be
further classified at present. They should not, it
seems, be termed displacement activities in the
restricted sense (p. 136) since they do not occur
in other behavior patterns found within the spe-
cies. These motions are disoriented fanning and
palpation of the female with the antennae. It
may be, of course, that this palpation with the
antennae, as well as with the proboscis, produces
chemotactic sensations in a situation where the
usual olfactory stimuli from the female are likely
to be weak or incomplete. In the European
satyrid, Eumenis semele, movements of the an-
tennae form an integral part of courtship (Tin-
bergen et al., 1943).
Disoriented fanning, in which the male may
face in any direction with respect to the female,
appears to be a disintegration of the normal pat-
tern, resulting from a breakdown in the usual
sequence of male-female responses. Subsequent-
ly the pattern often collapses altogether, the
remaining energy being channelled into displace-
ment-feeding or displacement-flight.
1957]
Crane: Imaginal Behavior in Butterflies of the Family Heliconiidae
143
In all irrelevant behavior appearing during
courtship, the non-responsiveness of the female,
for whatever reason, appears to be the major
factor. Males that are not in a physiological con-
dition to complete courtship simply break it off
at an early stage.
As stated in earlier pages, it is now certain
that females normally mate only once. Each
develops a specifically characteristic odor, given
off by the abdominal glands, which becomes ap-
parent to the human observer about an hour or
more after the butterflies have separated. It is
the same odor given off by mated females when
seized. It seems likely that it is one of the de-
terrents leading to irrelevant behavior when
males court mated females and do not follow
through to copulation even though the females
are making, to the human eye, all the visible
responses characteristic of the species. Another
deterrent in such cases seems to be the failure
of the female to fold the wings, which have been
more or less flattened during the courtship flut-
tering, above her back when the male alights
beside her in Stage III. It is impossible for him
to reach her abdomen with his harpes so long
as her wings are flattened.
Use of the proboscis by H. sara, when egg-
laying is apparently thwarted, is no part of the
normal egg-laying procedure of any of the mem-
bers of the family that have been studied, and
perhaps should be termed displacement be-
havior. Unlike members of some other families
(Use, 1955 and refs.), none of the heliconiids
drum the foodplant with their feet before laying.
Finally, it may be re-emphasized that almost
all these types of irrelevant behavior occur in the
lives of all individuals of the appropriate sexes
and ages that have been studied. No reliable
interspecific differences have yet been observed.
VIII. Summary
Six species of heliconiid butterflies from Trini-
dad, B.W.I., were reared in the laboratory and
their post-emergence behavior studied in insec-
taries. Their social behavior patterns are briefly
compared.
Social responses, including courtship, vary
with age. In females the full courtship pattern
is elicited only on the second and third days,
although they can mate successfully when both
younger and older. Younger females not only
give little or no overt response but draw scarcely
any pre-copulatory behavior from males. Older
females show progressively fewer specifically
characteristic responses.
Brief chasing of males or unreceptive females
by males of any age and by old females is a com-
mon form of activity. It apparently represents
simply a fragment of the appetitive portion of
the courtship pattern. Neither territorial defense
nor inter-male threat behavior seems to be in-
volved.
Irrelevant actions of a number of kinds are
frequent. Most occur in males that are thwarted,
usually by the unresponsiveness of the female,
after an advanced stage of courtship has been
reached. Two kinds of irrelevant actions, a dis-
oriented type of wing-fanning and palpation
with the antennae, are not known in any normal
behavior pattern of the species. Other types of
irrelevant behavior appear to be true displace-
ment activities, since they occur as appropriate
actions in other fields of the insects’ behavior,
such as in feeding and flight.
IX. References
Armstrong, E. A.
1950. The nature and function of displacement
activities. Symposia of the Society for Ex-
perimental Biology, No. 4. Physiological
mechanisms in animal behavior. Academic
Press, N. Y., 361-384.
Bastock, M., & A. Manning
1955. The courtship of Drosophila melanogaster.
Behaviour, 8: 85-111.
Bastock, M., D. Morris & M. Moynihan
1953. Some comments on conflict and thwarting
in animals. Behaviour, 6: 66-184.
Beebe, W.
1952. Introduction to the ecology of the Arima
Valley, Trinidad, B.W.I., Zoologica, 37:
158-184.
Buckingham, E.
1910. Division of labor among ants. Proc. Amer.
Acad. Arts & Sci., 46: 425-507.
Crane, J.
1949. Comparative biology of salticid spiders
at Rancho Grande, Venezuela. Part IV. An
analysis of display. Zoologica, 34: 159-214.
1952. A comparative study of innate defensive
behavior in Trinidad mantids (Orthoptera,
Mantoidea). Zoologica, 37: 259-293.
1955. Imaginal behavior of a Trinidad butterfly,
Heliconius erato hydara Hewitson, with
special reference to the social use of color.
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1957. Basic patterns of display in fiddler crabs
(Ocypodidae, Genus Uca). Zoologica, 42:
69-82.
Crane, J., & H. Fleming
1953. Construction and operation of butterfly
insectaries in the tropics. Zoologica, 38:
161-172.
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Fabre, J. H.
1879. Souvenirs Entomologiques. Paris, Dela-
grave. (Ed. def. illustree, 1914-1924. Vol..
2).
Ford, E. B.
1945. The New Naturalist: Butterflies. Collins,
London, xiv -f 368 pp.
Gordon, H. R. S.
1955. Displacement activities in fiddler crabs.
Nature, 176 (4447): 356-357.
Hagen, K. S.
1953. A premating period in certain species of
the genus Opius (Hymenoptera: Eurytom-
idae). Proc. Hawaiian Ent. Soc., 15(1):
115-116.
Hinde, R. a.
1953. The conflict between drives in the court-
ship and copulation of the chaffinch. Be-
haviour, 5: 1-31.
Iersel, J. J. a. van
1953. An analysis of the parental behaviour of
the male three-spined stickleback {Gas-
terosteus aculeatus L.). E. J. Brill, Leiden.
159 pp.
Ilse, D.
1955. Behaviour of butterflies before oviposition.
Jour. Bombay Natural Hist. Soc., 53:
486-488.
Lorenz, K. Z.
1950. The comparative method in studying in-
nate behaviour patterns. Symposia of the
Society for Experimental Biology, No. 4.
Physiological mechanisms in animal be-
haviour. Academic Press, N. Y., 221-268.
Mayr, E.
1942. Systematics and the origin of species,
Columbia University Press, N. Y., viii -f
334 pp.
Morris, D.
1954. The reproductive behaviour of the zebra
finch {Peophila guttata), with special ref-
erence to pseudofemale behaviour and
displacement activities. Behaviour, 6:
271-322.
Moynihan, M.
1953. Some displacement activities of the black-
headed gull. Behaviour, 5: 58-80.
Nielsen, E. T.
1945. Moeurs des Bembex. Spolia Zool. Mus.
Haun. Copenhagen 7: 1-174.
Nielsen, A., & E. T. Nielsen
1952. Migrations of the Pieride butterfly Ascia
monuste L. in Florida. Ent. Meddel.,
26(5): 386-391.
Pardi, I.
1947. Beobachtungen fiber das interindividuelle
Verhalten bei Polistes gallicus (Unter-
suchungen fiber die Polistini no. 10). Be-
haviour, 1(2): 138-172.
Rand, A. L.
1943. Some irrelevant behavior in birds. Auk,
60: 168-171.
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1956. Metamorphosis: a physiological interpre-
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Rosch, G. a.
1925. Untersuchungen fiber die arbeitsteilung
im bienenstaat. I. Teil: Die Tatigkeiten
im normalen bienenstaate und ihre bezie-
hungen zum alter der arbeitsbienen.
Zeitschr. Vergleich. Physiol., 2: 571-631.
SCHNEIRLA, T. C.
1953. Basic problems in the nature of insect be-
havior. (In: Insect Physiology, edited by
K. D. Roeder; John Wiley & Sons, Inc.,
New York, xiv -[- 1100 pp.) . pp. 656-684.
Steiner, A.
1932. Die arbeitsteilung der feldwespe Polistes
dubia K. Zeitschr. Vergleich. Physiol., 17:
101-152.
Tinbergen, N.
1951. The study of instinct. Oxford, Clarendon
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1952. “Derived” activities; their causation, bio-
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27: 1-32.
Tinbergen, N., B. J. D. Meeuse, K. K. Boereme
& W. W. Variosseau
1943. Die balz des sampfalters, Eumenia
(Satyrus) semele (L.). Zeitschr. Tierpsy-
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1957]
Crane: Imaginal Behavior in Butterflies of the Family Heliconiidae
145
EXPLANATION OF THE PLATE
Plate I
Figs. 1-6, incl. Species and subspecies of butterflies
discussed in this contribution. Photograph by Sam
Dunton, New York Zoological Society.
Fig. 1. Dryas julia julia. General color; Orange.
Fig. 2. Heliconius melpomene euryades. General
color: Black; forewing band scarlet.
Fig. 3. Heliconius erato hydara. Color as in H.
melpomene.
Fig. 4. Heliconius ricini insulana. General color:
Black; forewing bands pale yellow; central
portion of hindwing scarlet.
Fig. 5. Heliconius isabellaisabella. General color:
Reddish - brown to yellowish - brown;
marked with black.
Fig. 6. Heliconius sara rhea. General color:
Black; forewing bands pale yellow; basal
half of hindwing with dark blue irides-
cence.
Fig. 7. Irrelevant courtship behavior in Dryas
julia: Sedentary phase, Stage II, showing
poor orientation in fanning. In normal
fanning the flying male faces in exactly
the same direction as the sedentary female
(Table II and p. 138). Note that female
has forewings almost closed as required in
Stage III. However, the hindwings are still
fluttering, as is characteristic of earlier
courtship stages, but atypical here. Photo-
graph by M. Woodbridge Williams, © Na-
tional Geographic Society.
Fig. 8. Irrelevant courting behavior in Heliconius
erato, corresponding to stage shown in
Fig. 7. Photograph by Rosemary Kenedy.
CRANE
PLATE 1
FIG. 7
FIG. 8
13
Habits, Palatability and Mimicry in Thirteen
Ctenuchid Moth Species from Trinidad, B.W.Id
William Beebe & Rosemary Kenedy
Department of Tropical Research,
New York Zoological Society, New York 60, N. Y.
(Plates I & II)
[This paper is one of a series emanating from the
tropical Field Station of the New York Zoological
Society, at Simla, Arima Valley, Trinidad, British
West Indies. The Station was founded in 1950 by the
Zoological Society’s Department of Tropical Re-
search, under the direction of Dr. William Beebe.
It comprises 200 acres in the middle of the Northern
Range, which includes large stretches of undisturbed
government forest reserves. The laboratory of the
Station is intended for research in tropical ecology
and in animal behavior. The altitude of the research
area is 500 to 1,800 feet, with an annual rainfall of
more than 100 inches.
[For further ecological details of meteorology and
biotic zones see “Introduction to the Ecology of the
Arima Valley, Trinidad, B. W. I.,” William Beebe.
(Zoologica, 1952, Vol. 37, No. 13, pp. 157-184.)]
Contents
I. Introduction 147
II. Material and Methods 148
III. Observation and Results 148
Pseudosphex kenedyae Fleming 148
Saurita clusia (Druce) 150
Histiaea cepheus (Cramer) 151
Macrocneme thyra intacta Draudt 151
Calonotos crane ae Fleming 152
Aethria carnicauda (Butler) 152
Episcepsis hypoleuca Hampson 152
Napata walkeri (Druce) 152
Agyrta dux (Walker) 153
Eucereon maia Druce 153
Eiicereon obscurum (Moschler) 154
Correbidia assimilis (Rothschild) 154
Ctenucha andrei Rothschild 155
IV. Discussion 155
V. Summary 156
VI. References 157
^Contribution No. 980, Department of Tropical Re-
search, New York Zoological Society.
I. Introduction
The highly specialized moths of the family
known as the Ctenuchidae (Syntomidae,
Amatidae, Euchromiidae) are among the
most interesting and colorful of the Lepidoptera.
In general, they are medium to small in size;
many are brightly colored, some with metallic
luster on wings and abdomen. More than 2,000
species are known, the great majority being re-
stricted to the American tropics. Although they
are commonly referred to as “day-flying” moths,
many species fly also, or even exclusively, at
night. Some species “mimic” other insects, and
the larvae of some show remarkable adaptations
(see especially Beebe, 1953).
The taxonomy of this group in Trinidad has
been discussed by Kaye & Lamont (1927) and
Fleming (1957), but relatively few observations
have been reported on these moths in life : flight
pattern, position at rest, frequency, time of flight
and seasonal occurrence. Also, although it is
well known that this family includes species
which are obvious wasp and beetle “mimics,”
little experimental work has been done on the
palatability of these moths to possible predators.
For this paper field observations and palata-
bility tests were limited to 13 of the more than
80 species so far recorded in Trinidad. The
choice of species was based on the following
criteria: most are relatively common, therefore
more readily available for observation and col-
lection; they include both day- and night-flying
species; they include genera from the most high-
147
148
Zoologica: New York Zoological Society
[42; 13
ly specialized (Pseudosphex) to the most gener-
alized {Eucereon, Ctenucha); they include
species which show great differences in wing
scaling, i.e., completely hyaline (Pseudosphex),
completely scaled and having bright patches of
color {Histiaea), cryptic fuscous {Episcepsis)
and metallic lustrous (Macrocneme); some spe-
cies which are obvious “mimics” are included,
and some which are drawn to Heliotropium in-
dicum and others which apparently are not at-
tracted to it.
This attractant is reported in detail by Beebe
(1955 and refs.) but a brief summary is perti-
nent here. Heliotropium indicum Linnaeus is a
common weed which, when uprooted and al-
lowed to shrivel, exercises a remarkable selective
attraction for several lepidopteran families
including the Ctenuchidae. For some unknown
reason, however, only certain species appear to
be attracted to it, while other species common in
the same locality ignore it completely.
The authors are indebted to Mr. Henry Flem-
ing for identification and range of the ctenuchid
species used in this study.
II. Material and Methods
Nearly all observations and collections were
made within a radius of approximately a quar-
ter of a mile from the Simla field station, the
majority within 100 yards of the laboratory. At
night they were made at several incandescent
electric light bulbs outdoors and at a spotlighted
sheet, by day along trails, roadside or in the
laboratory itself where a large screened window
trapped many of the day-flyers which entered
at an open end of the room. H. indicum was
used both night and day as a collecting and
observing site. The best results were had with
this attractant when a clump of the weed was
suspended from a low tree branch along a shaded
trace.
Abundance records used in this paper were
kept during the three-year period 1953-1955 but
cover only the months of December to June, the
extreme limits of our stay at the field station.
Rest position was usually observed on a cap-
tured uninjured moth placed in a large glass jar
or terrarium. Most flight observations were
made on free-flying moths at night lights or at
H. indicum, or inside a screened insectary mea-
suring 12 X 18 feet.
As shown in Table 1, the relative acceptability
of moths to various vertebrates and invertebrates
was tested. In experiments with ants, the ctenu-
chids were either freshly dead or completely in-
capacitated. No chemicals were used for killing.
With all the other testers, living active specimens
were used. There is an occasional record of a
mantid or a spider capturing a ctenuchid in the
field, but all experiments with bird, lizards,
frog, tarantula and with most mantids were done
with the testers in captivity. The Polychrus lizard
and the bird, a Moriche Oriole, were tamed ,
specimens which readily accepted food held by
forceps in the hand.
In each test, with the exceptions just men-
tioned, an active moth was placed in the ter-
rarium or cage with the tester and the response
of the latter was noted. Each test was controlled,
i.e., a negative result was fisted only if another
lepidopteran specimen, usually a pyralid moth
or a riodinid butterfly of comparable size, was
accepted immediately afterwards.
The results of the tests were classified as
follows:
A— Moth eaten entirely (with occasionally the ,
obvious exception of wmgs or genitalia) . 1
B— Moth eaten partially or with apparent dis- J
taste, i.e., mouthed (by bird or mantid), sucked
for short time (spider), partially eaten only
(ants, mantid, bird), or eaten with apparent \
distaste (frog). i
C— Moth originally noticed (antennaed or
dragged by ants, seized by mantid, spider, frog, 1
lizard or bird) but eventually released without |
being injured.
D— Moth ignored completely or fled from.
The numeral following each of the above
classifications in Table 1 indicates the number
of tests performed with that result.
III. Observation and Results
Pseudosphex kenedyae Fleming
(Plate I, Figure 1; Plate II, Figure 14)
Range.— Trinidad.
Field Name.— Dusky-winged Wasp-mimic.
Field Characters.— Thordix. yellow with dark
brown markings; abdomen yellow banded with
brown, constricted at base; wings hyaline with
fuscous streak along costal margin of forewing.
In flight, cannot be distinguished from another
Trinidad ctenuchid, Sphecosoma trinitatis Roth-
schild, nor from its vespid wasp “models” (see
below) . Sexes differ by presence of ventral valve
in $.
Frequency.— This species was not seen by us
until dead Heliotropium indicum was used as an
attractant. Since then it has been very common
on the weed. The attraction is almost limited to
53, as out of 114 individuals seen or taken, only
one was 5.
Active Period.— One, of the most decidedly
Palatability of Ctenuchid Moths
1957]
Beebe & Kenedy: Habits, Palatability & Mimicry in Ctenuchid Moths
149
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150
Zoologica: New York Zoological Society
[42: 13
diurnal of the ctenuchids, flying, often with its
model, from early morning (6:00 to 6:15 A.M.)
to nearly dusk. In a terrarium it is invariably
positively phototropic. Observed every month
from December to June.
Rest Position.— Wings 75° back, at times ver-
tical. When walking, wings 45° back, 45° up.
Antennae in constant motion, often alternately.
When pausing between steps, the abdomen is
frequently twitched up and down several times.
Flight.— Quick, jerky darts, wholly wasplike;
sometimes remains suspended before landing,
motionless except for the wings, the hindlegs
extended downward and slightly backward. It
has been seen to alight directly on the underside
of a leaf without any hesitation and without
alighting on the upperside flrst.
Palatability.— (Table 1). In every one of the
22 tests made, this species proved to be unac-
ceptable to lizard, bird and three species of ants.
In 15 tests the moth was ignored completely; in
only seven cases was it approached or noticed.
Army ants dragged a living moth by one leg for
a few seconds before releasing it, uninjured. In
another instance, ants approached a freshly dead
specimen, touched it with their anteimae for a
short time but ignored it thereafter. In five tests
the tame Moriche Oriole started to approach a
living moth but backed away almost imme-
diately; in one case it pecked at the forceps but
not at the insect held by it. In five other tests
with the same bird the moth was ignored com-
pletely.
Mimicry.— There exists a remarkably exact
resemblance between this day-flying moth and
several of the common Simla wasps. In our first
search for this mimicry we captured seven of the
wasps and a single ctenuchid. The latter was
resting head down on a dry leaf of Heliotropium
indicum, while the former were either flying or
resting on leaves or inside the laboratory. This
particular hymenopteran was Stelopolybia palli-
pes var. anceps (Saussure).
The two field characters by which the “model”
and “mimic” may best be distinguished are the
head, that of the wasp being wider and bearing
a pair of jaws rather than a proboscis, and
secondly, the antennae. The solidity of the an-
tennae in the case of the wasp or the pectination
in the case of the moth is made more difiBcult
for differentiation by the fact that in the moth
the pectination comes to an abrupt ending about
two-thirds from the base, the distal end being
reduced to a slender, easily overlooked filament.
Other insects bearing a close resemblance to
this moth are Oriodes in Syrphidae and one of
the Stratiomyidae, both dipterans.
The force of the mimicry is enhanced by the
following characters: Size: In total length the
wasp is 12 mm., the moth 10 mm. Shape: The
abdomen of the moth is proximaUy constricted
to a most unmothlike degree. Color and Pattern : |
Basic amber color the same, also the thoracic i
and abdominal pattern. Antennae: Nervous j
twiddling. Walk and Flight: Character, speed I
and details of manoeuverability similar in moth |
and wasp. When the moth is feeding, its abdo- J
men is lifted into a curve and occasionally raised I
and lowered, emphasizing the wasplike shape
and pattern. Wings: As in the wasp, the wings
are held 45° up and out, and the costal area is
darker than the remainder of the forewing. An-
other remarkable wing character was noted by
Kaye (1913) in Pseudosphex spp. He observed
that they “curve the forewing while the hind-
wing is folded up, so that the wings appear to
be as narrow as the completely folded wings of
the Vespidae and Eumenidae.” This character
applies also to the Trinindad species, and is
shown to some extent in Plate II, figure 14.
On the whole, the present species of ctenuchid
appears to offer an almost perfect example of
Mullerian mimicry: the moth is definitely dis-
tasteful, and its extremely close hymenopteran
counterpart possesses an efficient sting.
Saurita clusia (Druce)
(Plate I, Figure 2)
Range. — French Guiana and the Amazon
Region.
Field Name.— Black-tipped Whitewing.
Field Characters. — Wings yellowish-white,
dusky at base with large apical spot. Five red
spots, paired on shoulders and collar and one at
base of dorsal abdomen. Hindwings short and
rounded. $$ with more extensive black scaling
at apex and base of forewing and antennae less
heavily pectinated, almost threadlike.
Frequency.— One of the most common species
at Simla, but of the 156 individuals seen or
taken, only five have been $?.
Active Period. — Strictly nocturnal. Taken
every month, December to June.
Rest Position.— Wings 60° back, flat, tips clear
of surface. When walking the wings are raised
45° and held well apart. Thorax and narrow
anterior segment of abdomen exposed, revealing
the five scarlet spots. Antennae 45° apart.
Flight.— At first, forewings are waved slowly,
then all four are whirred, before taking off.
Flight is flylike, usually more or less straight
upward, the insect finally alighting and walking
at once. When liberated from a hand-held con-
1957]
Beebe & Kenedy: Habits, Palatability & Mimicry in Ctenuchid Moths
151
tainer in an insectary, the moth seeks shelter
beneath fingers or in a dark crevice in the
rafters, as if to avoid the light. Twice found rest-
ing beneath a leaf.
Palatability— 1). This species appears
to be acceptable to mantids, moderately so to
spider and frog. A specimen was seized by a
tarantula but immediately released; later the
same specimen was captured and the abdomen
sucked for several minutes, but it was eventually
released alive and able to walk. Two specimens
were eaten by frog with apparent distaste. Two
were refused by Moriche Oriole. But as the moth
is strictly nocturnal, these negative results with
the bird appear to have little or no survival value.
Mimicry —1^0 definite models have been ob-
served in association with this species. It is flylike
in many respects but is strictly nocturnal.
Histiaea cepheus (Cramer)
(Plate I, Figure 3)
Venezuela to Surinam.
Field Name.— Large Red-and-yeUow-spotted
Brown.
Field Characters. — Large (expanse 2 Vi
inches), dark brown; forewings with three buffy
yellow spots, hindwings with two large, irregu-
lar, scarlet spots; basal segments of dorsum with
three incomplete yellow bands. Sexes very simi-
lar; abdomen of ? slightly broader.
Frequency.— Common. Of 81 moths seen or
taken, two-thirds were 35.
Active Period. — Strictly nocturnal, usually
taken at lights before 10 P. M. Twelve imagoes,
reared from larvae (Kenedy, unpub.), were
placed in an insectary for observation. They
were inactive during the day, usually resting,
singly, beneath a large leaf. From 6:00 to 6:30
P.M. they began to be active, flying and walking
on the screening of the roof. Seen or taken all
months, December to June.
Rest Position.— Wmg?, 50° back, tips not touch-
ing surface. Abdominal bands exposed. The
moth has often been observed to wave its wings
up and down, through a rather large arc, when
standing still or walking, or to whirr its wings
rapidly.
Flight.— In an insectary, it took off with moth-
like flight, the slow beat of the wings rendering
them visible. Liberated in the open, it rose swift-
ly and smoothly, circled twice in mid-air, then
flew straight over the valley.
Palatability.— (Table 1 ) . Thirty-four tests were
completed, 26 of which demonstrated that this
species could be considered unpalatable. A
rather unique phenomenon has been observed
in this species. When a specimen is immobilized
suddenly, whether captured by a caged tester
such as a mantid or held by observer, it has been
seen in many cases to exude droplets of thick
yellow fluid from the thorax. Sometimes the
droplets are bubbly and almost completely cover
the dorsal thorax. This seems to serve as an
effective defense against at least some predators.
In two cases mantids bit into the dorsum but
dropped the moth when the yellow froth came
in contact with their eyes and mouthparts; a
lizard five times threw a captured but uninjured
insect from its mouth with violent shaking of its
head. In the single test with the Moriche Oriole
it ate the interior of the thorax and abdomen,
leaving the head, genitaliaand entire exoskeleton.
In this case, no drops of fluid were observed on
the insect’s thorax. A tarantula ignored two
specimens, and captured but immediately re-
leased two others undamaged.
Macrocneme thyra intacta Draudt
(Plate I, Figure 4; Plate II, Figure 15)
Range.— Trinidad and Colombia.
Field iVa/ne.— White-bellied Greenwing.
Field Characters.— Wings black, basal half
with metalhc blue-green sheen; dorsal base of
abdomen with four white spots; hind tarsi white
tipped; S white below on thorax and abdomen,
$ venter with white spots.
Frequency.— Abundant. S $ dominate in num-
ber: of 326 seen or taken, only 10 were 99.
Active Period.— ChieQy night flyers. Of 326
individuals, 14 (nine $$, five 99) were netted
by day. Seen or taken every month, December
to June.
Rest Position.— W'mgs 20° back, flat, the fore-
wings overlapping the hindwings so that the
latter are invisible. Antennae 45° apart. When
walking, movements are nervous and jerky,
wasplike; wings held up and back 30° to 45°;
antennae in motion.
F/ig/it.— Wasplike, direct. The long hindlegs
are held down and backward in flight. When
several captured SS were released they spiralled
upward fairly slowly or circled several times
before flying away. Half of the total 99 collected
were netted while they were flying slowly along
roadside or near stream, in bright sunshine.
Palatability.— (Table 1). Fifty-two tests were
made with this species. In 29 the moths were
ignored completely and in eight they were origi-
nally noticed but uneaten. In five tests with the
Moriche Oriole, the bird would not approach the
insect; in one case the bird appeared to jump
agitatedly around its cage during the several
152
Zoologicar N ew York Zoological Society
[42: 13
minutes the moth was crawling inside. Ants
ignored freshly dead specimens in all 14 tests.
Mantids, however, ate 75 per cent, of the speci-
mens offered to them, and frog and Polychrus
lizard occasionally accepted the moths.
Mimicry— A% noted by Kaye (1913), mem-
bers of the genus Macrocneme seem to be ex-
cellent mimics of fossorial or pompilid wasps,
especially the genera Salius and Pepsis. M. thyra
intacta is wasplike both in facies and in behavior.
The wings and abdomen possess a metallic blue-
green sheen; the long hindlegs, more heavily
scaled than is usual in ctenuchids, are extended
downward and backward in flight. Kaye also
noted that they wave their antennae and vibrate
their wings rapidly when alighted, and that they
settle usually, like fossorial wasps, on ground,
bank or leaf.
Calonotos craneae Fleming
(Plate I, Figure 5)
Trinidad.
Field Name.— White-spotted Green-stripe.
Field Characters.— Wings black, with two or
three white spots on forewing ($$ have two and
sometimes a smaller third; $$ have three); a
single spot on hindwing; abdomen black with
three longitudinal, iridescent, pale green stripes.
Frequency.— Common, 78 having been re-
corded or collected, with $9 slightly outnumber-
ing $$.
Active Period.— This species has been taken
at Simla only at night lights, chiefly before 10
P.M. However, one specimen was captured by
Fleming in central Trinidad in daytime, in palm
and immortelle forest. Have been taken Decem-
ber to June, with March, April and May the
time of heaviest flight.
Rest Position.— W'mgs 50° back, 30° raised.
Hindwings concealed. Black abdomen exposed,
showing conspicuous streaking of iridescent
green. Antennae 45° up and out.
Relatively slow, direct. On one occa-
sion at night light a moth was seen to “play
dead” when taken.
Palatability. -{Tahie 1). Specimens refused
or rejected by bird, frog and tarantula. Ants ate
interior of two moths, ignored two others. Eaten
by mantids.
Aethria carnicauda (Butler)
(Plate I, Figure 6)
Venezuela and Brazil.
Field Va/ne.— Red-tailed Clearwing.
Field Characters.— with red-tipped ab-
domen; wings hyaline with margins and veins
black. with palpi, shoulder spot, forecoxae
and ventral base of abdomen white; 99 lack
white. This species may be mistaken for Dinia
mena (Hiibner) but the latter has the scarlet
abdominal fringe extended well up the sides.
Frequency.— A common species; 66 were ob-
served or collected; 99 slightly outnumbered $$.
Active Period.— Conhned to daylight, from
8:30 A.M. to 4:30 P.M. Observed from Decem-
ber to June. The moths are more numerous at
the end of the dry season, in March and April.
Rest Position.— Wings 60° back, flat, inner
edges not quite touching. In walking the wings
are raised 40°, spread so that the hindwings are
exposed; wings in slight but constant motion.
When the moth is walking about on the attract-
ant H. indicum the antennae are constantly in
play, tapping the surface.
Apparently more swift and direct than
is usual in the group. A moth liberated five times
in an insectary flew straight upward toward the
light.
Palatability.— (Table 1). This species was test-
ed with ants and bird only. Refused in single
test with Moriche Oriole; ants ate five of 16
specimens.
Episcepsis hypoleuca Hampson
(Plate I, Figure 7)
Range.— Central America.
Field Vame.— Red-collared Brownwing.
Field Characters.— W'mgs unicolored snuff
brown; forecoxae red; dorsal abdomen metallic
blue. 33 with white at base of ventral abdomen.
Frequency.— Common. Total recorded 128, of
which 82 were 33, 34 99 and 12 undetermined.
Active Period.— ChieQy nocturnal, both at
lights and H. indicum. Approximately 20 per
cent, were taken in the daytime; the sex ratio
remained constant. Seen December to May, with
heaviest flight in December and January.
Rest Position.— W'mgs 70° back, not touching
surface, overlapping at anal angle, meeting me-
dially.
Flight.— Slow, slightly wavering until full
speed is attained, then direct, not very rapid.
Fairly sluggish on H. indicum, crawling slowly
in and out of dried branches, sometimes drop-
ping instead of flying off when approached.
Palatability.— (Table 1). Refused or rejected
by bird in three tests, by house gecko in a single
test; frog ate one, rejected another; eaten by
mantids; ants ate four, ignored six.
Napata walker i (Druce)
(Plate I, Figure 8; Plate II, Figure 16)
1957]
Beebe & Kenedy: Habits, Palatability & Mimicry in Ctenuchid Moths
153
Mexico, Central America.
Field Name— Barred-tip Yellow.
Field Characters— Wings orange-yellow; apex
of forewings white barred with black; palpi, legs
and ventral surface black and white. Sexes indis-
tinguishable in the field.
Frequency.— Taken occasionally at Simla. Of
a total of 25 specimens, 22 were $$, three 99.
Active PmW.— Nocturnal. Taken from De-
cember to June, with the exception of May.
Rest Position.— Wings 70° back, just meeting
at midline over dorsum; tips not touching sur-
face.
Flight.— Wavering and not rapid. At end of
flight may seek shelter under leaf.
Palatability.— Because of the comparative
rarity of this species, only one specimen was
tested, and that inadvertently. It was a rare 9
which was being kept in captivity in the hope
of obtaining eggs. When freshly dead, ants
swarmed over it and ate it before it could be
salvaged for the collection.
M/m/cry.— This orange-yellow ctenuchid is
quite unlike other members of its family, but
there are several species of other families which
in color and pattern might be considered as
models. Among these are the riodinid butterfly,
Mesene semiradiata, and the zygaenid moth,
Malthaca radialis. A pyralid moth, Mapeta xan-
thomelas, fulfills all the demands in numbers
and appearance of a model, although it is a day-
flying species. At rest the ctenuchid and the
pyralid are remarkably similar, and not until
one is close enough to see the backward angle
of the antennae and the prominent palpi of the
pyralid can they be distinguished (see Plate II,
fig. 17) . In flight, however, the barred hindwings
of the pyralid are quite visible; the ctenuchid
lacks these markings.
Two of the pyralids were tested for palata-
bility with ants and were eaten except for the
shell of the thorax. However, two specimens
were ignored by the Moriche Oriole.
Agyrta dux (Walker)
(Plate I, Figure 9)
Range.— Central America to Venezuela and
Brazil; St. Lucia.
Field Vume.— Six-windowed Black.
Field Characters. — Relatively large moth,
black shot with blue; wings with six large, ir-
regular hyaline areas, two in forewing, one in
hindwing, crossed by black veins; wing bases,
thorax and abdomen iridescent blue; palpal base
and tongue scarlet; narrow dorsal abdominal
streak and venter white. Sexes similar except for
slightly shorter antennal pectinations in 9. Dif-
fers from closely related A. micilia by hyaline
forewing area arising near base.
Frequency.— OccasionaBy recorded at Simla.
A total of 23 consisted of 19 three 99 and
one undetermined.
Active Period.— Exclusively diurnal, taken
from early morning to late afternoon, with the
majority seen in late afternoon. December and
January have been the best months for recording
this species, but they have also been taken in
March, April and May.
Rest Position.— Wings 20° to 40° back, meet-
ing over back. When walking, wings are waved
through a 45° arc; antennae in constant motion,
moving in unison.
Flight.— Fairly rapid, darting, low, circling
before alighting. It was seen to alight on dead
H. indicum three times at the same spot. At once
it began to search eagerly with its tongue, within
a small area. After feeding on a panicle it liter-
ally ran through the dense, dry foliage, its wings
rubbing against the leaves in the narrow places.
Its speed was remarkable in the deeper inter-
stices. The wing iridescence and the deep red
collar and tongue could be plainly seen when the
rest of the insect was invisible.
Palatability .-{Table 1). This species is ap-
parently palatable to ants. A small ponerine ant
was once seen carrying off a 9 specimen, holding
it at the tip of the abdomen. The insect was
rescued, but died. The Moriche Oriole twice
picked up a specimen hesitantly, but dropped it
without injury.
Mimicry.— An unidentified geometrid moth is
a close model for the species except for its smaller
size, spreading only 28 mm. whereas A. dux
extends 40 mm. between wingtips. The rest posi-
tion of the geometrid is similar to that of the
ctenuchid and it also is a day-flyer. Both in
facies and flight, A. dux appears more butterfly-
like than is usual in the group.
Eucereon maia Druce
(Plate I, Figure 10)
Range.— Mexico, Central America, Venezuela,
British Guiana.
Field Name.— Red-bodied Brown.
Field Characters.— Tborax and forewings light
brown, mottled with darker; hindwings pale; ab-
domen red above, dotted with black, whitish
below. Forewings of 9 paler.
Frequency.— of this species are abundant.
Total of 260 seen or taken consisted of 196
12 99 and 52 undetermined.
154
Zoologica: New York Zoological Society
[42: 13
Active Per/oci.— Nocturnal, flying throughout
the night, appearing at dusk and seen also at
dawn with records every hour to 3 A.M. The
peak of the flight seems to be 10:30 to 11:30
P.M. In an insectary the moth is negatively pho-
totropic. Recorded every month from December
to June, with December and January the months
of heaviest flight.
Rest Position— 70° back, overlapping,
completely covering abdomen.
Flight.— Rapid and direct, occasionally cir-
cling once or twice. When disturbed, the moth
often drops to the ground, beetlelike, before tak-
ing flight. Or it may crawl along the ground
instead of flying, sometimes hiding under a low-
growing leaf. The habit of dropping is so usual
that a specimen can almost always be collected
merely by placing the open end of a vial be-
neath it.
Palatability.— (Table 1). Ants ate seven speci-
mens, ignored five; spiders sucked three; one was
captured and eaten by a centipede on H. indicum.
In majority of tests the frog captured but spit
out this species. The Moriche Oriole ate one,
ran two others through its bill, then flicked them
away, injured.
Eucereon obscurum (Moschler)
(Plate I, Figure 11)
Mexico and Central America south
to the Amazon.
Field Name.— Blue-spangled Bronzewing.
Field Characters.— Abdomen black, the termi-
nal segments shot with blue; forewings bronze-
black, faintly spangled and lined with bluish-
white; hindwings semi-hyaline, pale brown. Sexes
indistinguishable in the field.
Frequency.— A common species. Of 87 speci-
mens 58 were $$, 10 52 and 19 undetermined.
Active Exclusively nocturnal, like
others of the genus. It first appears in early
evening and has been found as late as 3 : 30 A.M.
Has been taken from December to June, with
the heaviest flight in December and February.
Rest Position.— Wings 80° back, almost par-
allel; inner edges overlapping, concealing body;
tips resting on surface.
Flight.— A rapid fluttering, but rather slow
flight. The moth circles slowly, hovers before
alighting. When released in insectary, it invari-
ably flew to the ground and hid among grass.
Palatability.— (Table 1). Eaten by ants, man-
tids and frog; Moriche Oriole ate head and
thorax of one, seized four others but dropped
them immediately.
Correbidia assimilis (Rothschild)
(Plate I, Figure 12; Plate II, Figures 18, 19)
Venezuela, British Guiana, Surinam,
Brazil.
Field ATamc.— Yellow-banded Beetle Mimic.
Field Characters.—Small moth. Wings black,
crossed in center by a wide, pale yellow band.
Sexes indistinguishable in the field except by
slightly shorter antennal pectinations in 2.
Frequency.— The most abundant ctenuchid at
Simla. Total recorded 337, of which 205 were
55, 30 22 and 102 undetermined.
Active Period.— More than 95 per cent, were
seen or taken at night, being drawn chiefly to
lights but also to uprooted H. indicum. Occa-
sionally taken during the day and then usually
in the early morning or late afternoon, within
two hours of dawn and dusk. Seen from Decem-
ber to May. The numbers drop off in April and
May.
Rest Position.— Wings 80° back, outline
curved, overlapping medially, only thorax ex-
posed. In this position the light yellow midwing
markings form a continuous transverse band.
Antennae 45° forward and out, tips slightly
curved. When walking the moth waves its an-
tennae alternately up and down. This species
appears to be unusually lethargic and slow to
take flight.
Flight.—Slow and fluttering.
Palatability.— (Table 1). Ants ate three speci-
mens, ignored two; eaten by mantids, spider,
frog and lizard; Moriche Oriole ignored three,
approached one hesitantly but would not take it.
M/m/cry.— Several authors, especially Kaye
(1913), have commented on the remarkable
resemblance that exists between members of this
genus and lycid beetles, such as the genus Lyco-
morpha in North America. Lycid beetles are
general in the Neotropics and are known to be
protected (Forbes, 1930, p. 27). There are three
Simla insects which might be considered as
models for C. assimilis: a small arctiid moth,
Lycomorphodes aracia, and two lycid beetles,
a smaller and a larger species (Plate II, figs. 18,
19). The arctiid and the smaller beetle are noc-
turnal, the larger beetle is a day-flyer.
The smaller lycid is common and frequently
found associated with the ctenuchid. The larger
beetle and C. assimilis are similar in color and
pattern and in their slow, lethargic movements.
They also resemble one another in appearance
and movements of the antennae, and in the
mutual habit of slowly lifting and lowering the
forewings (elytra in the case of the beetle) as
1957]
Beebe & Kenedy: Habits, Palatability & Mimicry in Ctenuchid Moths
155
they walk. The average in both is seven waves
in five seconds.
During the same length of time 54 arctiids,
37 lycids and 130 specimens of the ctenuchid
were recorded.
Kaye (1913) made several interesting obser-
vations on mimicry in Correbidia which we have
also noted in C. assimilis in Trinidad: they are
sluggish in the early morning and, beetlelike,
drop to the ground when disturbed, drawing
their legs in; the shape and color of wings are
like a lycid beetle. To heighten the deception,
the abdomen of the moth is slightly flattened,
the legs are short and the heavy pectinations of
the antennae are carried to the tip.
To check palatability, the two beetles and the
arctiid were tested with ants. Three specimens
of the arctiid were eaten; one large and two small
beetles were untouched.
Ctenucha andrei Rothschild
(Plate I, Figure 13)
Range.— been recorded also from British
Guiana.
Field Name.— Large White-banded Black.
Field Characters.— A good-sized moth; fore-
wings blue-black with broad, oblique, white
band; hindwings steel blue. $$ with white on
venter, ?S black.
Frequency.— $$ common, $$ rare. Of 89 speci-
mens, 63 were S3, 2 59 and 24 undetermined.
Active Period.— A diurnal species, flying at
any hour of the day from dawn to dusk. A single
exception was a $ taken at the night light in
early evening. Observed from December to June.
Rest Position.— Wings 60“ back, meeting over
dorsum; white bands on forewings not meeting.
When walking the moth waves its wings slowly
through a 45° arc. Antennae 70° forward, 20°
apart.
Flight.— Rather slow and fluttering. In flight
the wings become a black haze, with curved ring
of white above and below. Wings move through
an arc of 130°.
Palatability. — {Table 1). This species was
eaten by mantid, tarantula, frog and lizard. In
60 per cent, of the tests with ants the moth was
eaten. Army ants ignored one specimen, killed
but did not eat another. In two tests the Moriche
Oriole took the moth but dropped it imme-
diately; it ignored four others.
Mimicry.— The most reasonable “model” for
this species is Cecropterus bipunctatus (Hes-
periidae). In abundance, pattern and color, gen-
eral motions and habitat this approximates the
ctenuchid. However, this skipper was found to
be acceptable to the Moriche Oriole.
IV. Discussion
Table 2 is a recapitulation of data, most of
which are already given in the text under in-
dividual species, with the addition of positive or
negative results with the use of Heliotropium
indiciim as an attractant. These results bring out
several interesting problems: although the at-
traction of the weed is highly selective, being
almost exclusively limited in the Heterocera to
the family Ctenuchidae, yet some of the most
common Trinidad ctenuchids, such as Macroc-
neme thyra intacta, have never been found on it.
Pseudosphex kenedyae is a common visitor to
FI. indicum, but it has not been seen elsewhere
nor had we ever seen a specimen before using
the attractant. Other ctenuchids are common
both at lights and on the weed, such as Corre-
bidia assimilis and Eucereon maia; however, an-
other member of the same genus, E. obscurum,
is fairly common at lights but extremely rare
on H. indicum.
Frequency.— The species are listed in order
of abundance in Table 2. The sex ratio of 11
species shows a predominance of males. How-
ever, females slightly outnumber males in
Calonotos craneae and Aethria carnicauda. As
the latter flies by day and as the larval foodplant
is a low-growing sedge (see Beebe, 1953) and
as eggs are laid singly (Kenedy, unpub.), it is
perhaps reasonable to suppose that ovipositing
females would be seen and netted more often
than males. But in the case of C. craneae all the
females seen or taken at Simla were attracted to
night lights.
Active Period.— Six species were found to be
exclusively nocturnal and four exclusively di-
urnal; three species were active both by day and
at night, although chiefly nocturnal.
Rest Position.— The backward angle of fore-
wings at rest ranges among the 13 species from
20° in Macrocneme thyra intacta to an almost
parallel 80° in Eucereon obscurum and Corre-
bidia assimilis. The wasp “mimics,” M. thyra
intacta and Pseudosphex kenedyae, hold their
wings at a slightly upward angle when walking;
the other species hold them either parallel to the
surface or angled slightly downward so that the
apex is almost touching the surface.
Flight.— There is great variation in habits of
flight, from slow and wavering {Correbidia
assimilis, Episcepsis hypoleuca) to rapid and
direct {Aethria carnicauda, Eucereon maia).
Palatability and Mimicry.— In Table 1 the re-
sults of palatability tests show that the only
156
Zoologica: New York Zoological Society
[42: 13
Table 2. FREQtrENCT, Active Period and Attraction to Heliotropium indicum
Species are listed in order of abundance. Totals are for three-year period 1953-1955, principally during
January to April. Extreme limits of observation and collection were December to June.
Species
Frequency
Active Period
Attraction
to
H. indicum
Total
Seen
or
Taken
%
Sex Ratio
%
9 9
%
Un-
determined
%
Diurnal
%
Nocturnal
Conebidia assimilis
337
61
9
30
5
95
+
Macrocneme tkyra Intacta
326
97
3
4
96
-
Eucereon maia
260
75
5
20
100
-1-
Saurita clusia
156
97
3
100
-
Episcepsis hypoleuca
128
64
27
9
20
80
-1-
Pseudosphex kenedyae
114
>99
<1
100
+
Ctenueha andrei
89
71
2
27
100
-1-
Eucereon obscurum
87
67
11
22
100
-f (rare)
Histiaea cepheus
81
68
22
10
100
-
Calonotos craneae
78
45
49
6
100
-
Aethria carnicauda
66
45
55
100
-t-
Napata walkeri
25
88
12
100
+ (rare)
Agyrta dux
23
83
13
4
100
-1-
species which appears to be well protected from
potential enemies is the vespid wasp “mimic”
Pseudosphex kenedyae and therefore is an ex-
cellent example of Miillerian mimicry. Although
tests with the other species are inconclusive, it
seems that they are at least partially protected
against possible predators.
V. Summary
Field characters, sex differences, frequency,
active period, rest position and flight are re-
corded in 13 of the more common species of
ctenuchid moths in Trinidad.
Interesting habits include extremely wasplike
flight as well as facies in two species, and in two
others a beetlelike dropping to the ground when
disturbed.
In two species only, more females were seen
or taken than males; in the others, males far
outnumbered females.
There is great variation in habits of flight and
in rest position among the 13 species.
Their palatability to various vertebrates and
invertebrates was tested under controlled condi-
tions. They appear to be at least partially un-
acceptable to possible predators.
It is suggested that a fluid exuded from the
thorax of Histiaea cepheus serves as an effective
defense.
Mimicry in several species is discussed.
Pseudosphex kenedyae appears to be an almost
perfect example of Mullerian mimicry.
1957]
Beebe & Kenedy: Habits, Palatability &. Mimicry in Ctenuchid Moths
157
VI. References
Beebe, W.
1952. Introduction to the ecology of the Arima
Valley, Trinidad, B.W.I. Zoologica, 37:
157-184.
1953. A contribution to the life history of the
euchromid moth, Aethria carnicauda But-
ler. Zoologica, 38: 155-160.
1955. Two little-known selective insect attraa-
ants. Zoologica, 40: 27-32.
Fleming, H.
1957. The Ctenuchidae (moths) of Trinidad,
B.W.I. Zoologica, 42: 105-130.
Forbes, W. T. M.
1930. Insects of Porto Rico and the Virgin
Islands. Sci. Surv. Porto Rico and Virgin
Isl. (New York Acad. Sci.), 12 (1): 1-171.
Kaye, W. J.
1913. A few observations on mimicry. Trans, ent.
Soc. London, 1913: 1-10.
Kaye, W. J., & N. Lamont
1927. A catalogue of the Trinidad Lepidoptera
Heterocera (moths). Memoirs of the De-
partment of Agriculture, Trinidad and
Tobago. No. 3.
158
Zoologica: New York Zoological Society
[42: 13: 1957]
EXPLANATION OF THE PLATES
Plate I
Fig.
1.
Pseudosphex kenedyae.
Fig.
2.
Saurita clusia.
Fig.
3.
Histiaea cepheus.
Fig.
4.
Macrocneme thyra intacta.
Fig.
5.
Calonotos craneae.
Fig.
6.
Aethria carnicauda.
Fig.
7.
Episcepsis hypoleuca.
Fig.
8.
Napata walkeri.
Fig.
9.
Agyrta dux.
Fig. 10.
Eucereon maia.
Fig. 11.
Eucereon obscurum.
Fig. 12.
Correbidia assimilis.
Fig. 13.
Ctenucha andrei.
Plate II
All photographs are of living specimens in natural
rest position, taken in a confined area.
Fig. 14. Pseudosphex kenedyae feeding on Helio-
tropium indicum seed panicle. Note wasp-
like pattern, constriction of abdomen and
curve of forewings; also reduction of pec-
tinations on distal third of antennae.
Fig. 15. Macrocneme thyra intacta, showing long,
heavily scaled hindlegs and white tarsi.
Fig. 16. Napata walkeri.
Fig. 17. Mapeta xanthomelas (Pyralidae).
Fig. 18. Correbidia assimilis and small lycid beetle.
Fig. 19. Same, with large lycid beetle.
BEEBE a KENEDY
PLATE I
FIG. 3
FIG, 4
FIG. 5
FIG. 6
FIG. 7
FIG. 9
HABITS, PALATABILITY AND MIMICRY IN THIRTEEN
CTENUCHID MOTH SPECIES FROM TRINIDAD. B.W.I.
BEEBE a KENEDY
PLATE II
FIG. 14
FIG. 16
FIG. 15
FIG. 17
FIG. 18 FIG. 19
HABITS, PALATABILITY AND MIMICRY IN THIRTEEN
CTENUCHID MOTH SPECIES FROM TRINIDAD, B.W.I,
14
Serological Relationships among Members of the Order Carnivora^
Ludwig K. Pauly- & Harold R. Wolfe
Department of Zoology, University of Wisconsin,
Madison, Wisconsin
The serological technic has been used in
taxonomic studies for more than fifty years.
The discovery of precipitins by Krause in
1897 and the publication in 1904 of Nultall’s
book applying the new technic to problems of
animal relationships gave taxonomists an ap-
proach which showed great promise of clarify-
ing disputed or undetermined relationships. The
precipitin technic has been used to a limited ex-
tent for the latter purpose, but has been quite
extensively applied for verification of existing
relationships based on morphological criteria.
The usefulness and trustworthiness of the tech-
nic have been shown by many workers. Studies
have chiefly been made with vertebrates but
there has also been research with a few of the
invertebrate phyla. The following is a brief list
of some representative papers in the field:
Boyden (1926, 1934, 1943), Boyden & Noble
(1933), Wolfe (1936), Brown & Helfron
(1928), Eisenbrandt (1938), Wilhelmi (1940),
Martin & Cotner ( 1934) , Baier & Wolfe (1942) ,
Gemeroy (1943) and Leone & Pryor (1954).
It seems to us that the greatest value of the sero-
logical technic now lies in its possibility of set-
tling questionable animal or plant relationships.
Perhaps no other order of the Class Mam-
malia contains such a diversified group of ani-
mals as does the Order Carnivora. This diversi-
ty has been responsible for considerable conflict
and uncertainty in attempts at classification. The
problem lies not so much in the over-all picture
of Carnivora classification as in the grouping of
related forms in the suborders and superfamilies.
Most authors seem to agree that the order should
^Supported in part by the Research Committee of
the University of Wisconsin Graduate School from
funds supplied by the Wisconsin Alumni Research
Foundation.
^Present address: University of Wisconsin, Milwaukee.
be divided into two suborders: the Fissipedia
and the Pinnipedia. There are disagreements as
to the relationships and inter-relationships
among the families of the Fissipedia and the re-
lationships of these families to the Pinnipedia.
Winge (1923-24) and Scott (1937) both seem
to favor combining the Ursidae, Canidae and
Procyonidae into one group, with the Musteli-
dae as a distinct but associated family, and the
Viverridae and Hyaenidae into another group,
with the Felidae as a distinct but associated
family. Winge associates the Pinnipedia with the
Ursidae and Canidae. Beddard (1902), Romer
(1933) and Simpson (1945) favor combining
the Canidae, Ursidae, Procyonidae and Mus-
telidae into one superfamily and the Viverridae,
Hyaenidae and Felidae into another superfamily.
Beddard apparently believes the Pinnipedia
closely associated with the Mustelidae, whereas
Romer seems to indicate that the Pinnipedia are
most closely associated with the Ursidae and
Canidae. Other authors have opinions varying
slightly from the above.
Since their discovery in the late Nineteenth
Century, little use has been made of serological
reactions in the taxonomic study of the Carni-
vora. Nuttall (1904) summarized the results of
his flocculation tests with the sera of 56 different
species of Carnivora. In general his results
agreed with the systematic position of the species
tested. With this lone exception no other exten-
sive work has been done on the serological
taxonomy of the Carnivora. Brief mention of
them is made in several papers: Boyden (1926,
1942), Boyden & Gemeroy (1950) and Wolfe
(1936).
An extraordinary study on the taxonomy of
the Carnivora was included in the book by
Reichert & Brown (1909) in which the rela-
tionships between various members were deter-
159
160
Zoologica: New York Zoological Society
[42: 14
mined by the resemblances and differences of
hemoglobin crystals formed from the different
species. On the basis of their study Reichert &
Brown claimed that the Ursidae and Mustelidae
showed a closer relationship to the Pinnipedia
than did any of the other Fissipedia. A peculiar
finding was that the hemoglobin crystals of the
skunk more nearly resembled those of the Pro-
cyonidae, which in turn did not resemble those
of the Pinnipedia. The hemoglobin crystals of
the Canidae resembled those of the Pinnipedia
less, while the crystals of the Felidae and Viver-
ridae resembled those of the Pinnipedia least
of all.
The classification and nomenclature used in
this paper follow those of Simpson (1945) as
much as possible.
Materials and Procedures
Many of the blood sera used as antigens were
received from the New York Zoological Park
through the courtesy of Dr. L. J. Goss and from
the San Diego Zoo and the Serological Museum
of Rutgers University through the courtesy of
Drs. C. R. Schroeder and Alan A. Boyden
respectively. Table 1 presents a list of animals
from which sera were obtained.
Both chickens and rabbits were used for the
production of antisera. Three different technics
of precipitin testing were employed. These were
the ring (inter facial) test, the photronreflectom-
eter method of Libby (1938) and the micro-
densitometer method of Baier (1943). The
latter two are turbidimetric methods; the pho-
tronreflectometer measures scattered light pro-
duced by the flocculating particles and the
microdensitometer measures transmitted light.
The authors found it advisable to use different
injection procedures in order to obtain antisera
of different precipitating ability, for it was neces-
sary to have quite heavy precipitates when the
microdensitometer was used, and weaker pre-
cipitating sera when the photronreflectometer
was employed.
With one exception, all antisera used in the
ring tests were produced in chickens. Each
chicken was given a single intravenous inocula-
tion of 1 ml. of a 2 per cent, solution of blood
serum (the antigen). This injection procedure
is the best for production of antiserum of low
precipitating power, a high interfacial titer
and good specificity (Wolfe, 1936). The birds
were bled eight to ten days after the injec-
tion. The antisera were allowed to stand for
at least seven days in the refrigerator before
use, as in vitro changes occurred in the serum
upon such standing (Wolfe, 1942). The one
rabbit used was treated in the same manner
Table 1. Animals Used in Study
Order CARNIVORA
Suborder Fissipedia
Family Canidae
Canis familiaris (dog)
Canis lupus (timber wolf)
Vulpes fulva (red fox)
Family Ursidae
*Ursus americanus (black bear)
■• Thalarctos maritimus (polar bear)
Family Procyonidae
Procyon lot or (raccoon)
*-'Potos caudivolvuliis (kinkajou)
**Nasua narica (coati-mundi)
Family Mustelidae
Mustela furo (ferret)
Mephitis mephitis (skunk)
Taxidea taxus (badger)
Mustela vison (mink)
■•■Tayra sp. (tayra)
Family Felidae
■•'Felis concolor (mountain lion)
Felis doinesticus (house cat)
Panthera pardus (leopard)
*Panthera tigris (tiger)
*Acinonyx jubatus (cheetah)
Family Hyaenidae
Hyaena hyaena (striped hyaena)
Suborder Pinnipedia
Family Otariidae
■■■**Eiimetopias jiibata (Steller’s sea lion)
***Zalophus californianus (California sea
lion)
'^Zalophus californianus (California sea
lion)
Family Odobenidae
*Odobenus rosinanis (walrus)
Family Phocidae
*-'*Phoca vitulina richardii (harbor seal)
Order ARTIODACTYLA
Family Bovidae
Bos taurus (cattle)
Bison bison (American bison)
Order PRIMATES
Family Hominidae
Homo sapiens (man)
*Sera furnished by Dr. L. J. Goss.
**Sera furnished by Dr. A. A. Boyden.
* * •^■'Sera furnished by Dr. C. R. Schroeder.
All other sera collected from local sources, in-
cluding Madison Zoo.
1957]
Pauly & Wolfe: Serological Relationships among the Carnivora
161
described for production of antisera used in the
photronreflectometer method.
Both chiekens and rabbits were used in the
production of antisera for the photronreflectom-
eter studies. The rabbits were given a single
series of three intravenous injections on alter-
nating days, totaling 3 ml. of undiluted serum;
the first injection was 0.5 ml., the second 1 ml.
and the third 1.5 ml. The chickens received
the same number of injections and on similar
days but the solution was a 2 per cent, solution
of the antigen rather than undiluted serum. The
rabbits and chickens were bled on the seventh
day after the last injection.
The antisera used in the microdensitometer
studies were produced in chickens and rabbits.
The increased precipitating power of the anti-
sera needed in such studies was produced by
increasing the amount of antigen inoculated into
the animals. The rabbits were given two more
series of three injections each at an interval of
approximately 30 days. Each series consisted
of a total of 3 ml. of undiluted serum. The chick-
ens were given only one injection series, totaling
3 cc. of undiluted serum in three injections. The
rabbits and chickens were bled seven days after
the last injection. It should be emphasized that
the above injection procedures did not always
result in antisera of sufficient potency, and
rather than reinject the animals that were poor
antibody producers they were discarded. Chick-
ens were found to be much better producers of
antibody than the rabbits.
The chickens and rabbits were starved for
18 to 24 hours before bleeding. The blood was
removed by cardiac puncture and allowed to clot.
The serum was removed after centrifugation
and stored in the refrigerator.
The ring test was performed in a 1.8 per
cent, saline solution when chicken antiserum
was used and in a .9 per cent, buffered saline so-
lution when rabbit antiserum was employed.
Serial dilutions of the antigen were made from a
2 per cent, solution which in turn was made
from the undiluted antigen. One-tenth of a
milliliter of antisera was layered below the anti-
gen solutions (.5 ml.). Readings were made at
5, 10, 20, 30 and 60 minutes, but only the 60-
minute readings are recorded in this paper.
The test antigens employed in the photron-
reflectometer and microdensitometer studies
were also serially prepared. The final reaction
mixtures of antigen and antisera were approxi-
mately .9 per cent, for the rabbit system and
8 per cent, for the chicken system, since
Goodman, Wolfe & Norton (1951) showed
that 8 per cent, was the optimum for the chicken
antiserum system. In order to conserve anti-
serum, only alternate dilution tubes were used
in the microdensitometer and photronreflectom-
eter tests.
The photronreflectometer tests were conduc-
ted according to a procedure modified slightly
from that outlined by Baier (1947). The cells
used were standardized as to thickness. This
means that the light beam passes through the
same distance in the liquid of each set of cells.
This is important in measuring the light-scat-
tering effect of particles in a suspension. Com-
parative results cannot be obtained if this dis-
tance varies among the individual cells of each
set. The microdensitometer tests were conducted
according to the procedure outlined by Baier
( 1 947 ) . The tubes used in these tests were stand-
ardized according to diameter and transmission
of light beams.
The photronreflectometer and microdensitom-
eter tests were made with antigen dilutions so
chosen that the final readings for the highest
and lowest dilutions (antibody excess and anti-
gen excess) equalled the control reading at those
two points. This was not always possible at the
antigen excess region because of the small
amount of antigen that was available in several
cases.
Values of the relationships are expressed in
percentage. The homologous reaction is con-
sidered to be 100 per cent, and the heterologous
reactions are related to this. The galvanometer
readings were summated for all the antigen
concentrations used when the photronreflectom-
eter and microdensitometer technics were used.
Results
Table 2 presents a summary of the results
obtained with 15 different antisera using the
ring test technic. The two anti-Mustelidae sera
(anti-mink and anti-ferret) gave high cross-re-
actions not only with other mustelids but also
with the dog, fox, black bear and raccoon. The
degree of cross-reactions with members of the
Felidae were considerably lower in two of the
three tests made and the reactions with Bovidae
and Hominidae were very low.
Only one Canidae antiserum was produced.
The anti-red fox serum reactions indicated that
the Ursidae and Procyonidae were more closely
related to the Canidae than to the Mustelidae or
Felidae.
The anti-black bear serum was a very specific
serum and gave a high cross-reaction only with
the raccoon serum. Much weaker reactions oc-
curred with other families of Fissipedia and
also with man.
162
Zoologica: New York Zoological Society
[42: 14
Table 2. Serological Relationships among the Carnivora — Ring Test Technic.
Relationship Values Expressed in per cent.
Vh
<L>
yi
C
<
Antigen Source
Mink-67
(12800)*
Ferret- 107
(51200)
Fox-165
(25600)
Black bear-162
(12800)
Cat-65
(51200)
Leopard-139
(25600)
Mountain lion-111
(25600)
Kinkajou-PC-10
(51200)
Coati-mundi-PC- 1 0
(51200)
Raccoon-C-1416
(51200)
Steller’s sea lion-C-1 55
(51200)
Steller’s sea lion-C- 1 63
(51200) }
Walrus-PC-88
(6400)
Harbor seal-C-165
(25600)
Harbor seal**
12800
Suborder Fissipedia
Family Procyonidae
Kinkajou
100
6.3
6.3
Coati-mundi
100
6.3
Raccoon
50
100
25
25
6.3
2.3
50
50
100
12.5
0
25
0
0
Family Ursidae
Polar bear
12.5
50
6.3
12.5
25
0.4
Black bear
100
25
100
3.1
6.3
100
50
6.3
25
0.8
Family Canidae
Red fox
50
25
100
1.6
0.4
12.5
25
0
25
0
Dog
25
50
50
3.1
6.3
1.6
0.8
0
Family Mustelidae
Ferret
100
100
12,5
4.7
1.6
25
Skunk
50
50
3.1
3.1
12.5
3.1
0.4
6.3
12.5
50
12.5
1.6
12.5
0
Mink
100
100
3.1
1.6
1.6
Badger
50
50
3.1
3.1
3.1
3.1
1.6
Family Felidae
Cat
9.4
18.8
3.1
1.6
100
12.5
50
6.3
0
Leopard
3.1
3.1
100
Mountain lion
25
6.3
100
50
100
Suborder Pinnipedia
Family Otariidae
Steller’s sea lion
50
100
25
100
100
50
3.1
25
Family Odobenidae
Walrus
25
50
25
50
100
3.1
25
Family Phocidae
Harbor seal
25
50
25
50
100
100
Order Artiodactyla
Cattle
6.3
3.1
0
1.2
0.8
3.1
0.8
0
0
Bison
6.3
0
1.6
6.3
0
Order Primates
Man
6.3
0.9
0.8
3.1
3.1
2.3
* Ring test titer.
** Produced in rabbit; all others produced in chickens.
Three anti-Felidae sera were tested. All of
these gave strong cross-reactions with the cat,
leopard and mountain lion, but a low order of
reactions with the members of other families.
The intra-family results were very peculiar in
one test. The reactions of the anti-leopard serum
indicated that the leopard was more closely re-
lated to the mountain lion than to the house
cat. Such unexpected results warrant further
investigation.
The three Procyonidae sera were all quite
aspecific in their cross-reactions. All these sera
gave large reactions with the Pinnipedia and the
anti-kinkajou serum indicated that these Pin-
nipedia were more closely related to the kin-
kajou than were the Mustelidae, Canidae and
Ursidae. The one test made with a representa-
tive of the Felidae showed the cat to be more
distantly related to the raccoon than to other
Fissipedia.
1957]
Pauly & Wolfe: Serological Relationships among the Carnivora
163
Table 3. Serological Relationships among the Carnivora-Microdensitometer and
Photronreflectometer Technics. Relationships Expressed in per cent.
Microdensitometer
Photronreflectometer
ci
u
o
C
c
Antigen Source
*Steller’s sea lion-157
*Steller’s sea lion-CU
Steller’s sea lion-3698-3796
*Black bear-3659-3660
1
*Raccoon-CX
1
Steller’s sea lion-5
UJ
Oh
tJL
"o
pq
C/2
*
*Cat-PA
Suborder Fissipedia
Family Procyonidae
Kinkajou
Raccoon
11.6
42
23.4
49.2
34.9
65.7
100
18.2
16.7
0
1.37
0.7
Family Ursidae
Polar bear
Black bear
34.9
26.7
39
35.9
55.6
59.5
93
100
53.6
26.9
15.7
18.4
21.6
18.8
15.9
11.2
0
22
Family Canidae
Red fox
Dog
Timber wolf
20.5
13.4
47.3
38.4
18.6
2.9
52.5
88.1
100
5.
0
0
1.3
Family Mustelidae
Skunk
Ferret
Tayra
30.7
55.3
45.3
37.3
20.1
13.1
12.1
100
11.3
15.1
2
1
Family Felidae
Cat
Tiger
Mountain lion
Cheetah
6.1
0
1
3.2
100
50.4
47.8
76.6
Family Hyaenidae
Hyaena
0
18.3
Suborder Pinnipedia
Family Otariidae
Steller’s sea lion
Cal. sea Uon
100
100
100
45.3
32.7
100
82.6
9.3
6
0
.9
Family Odobenidae
Walrus
45.5
8.9
8.1
Family Phocidae
Harbor seal
81.5
52.4
76.6
44.2
47.4
9
10.3
Order Artiodactyla
Cattle
14.4
17.7
5.2
4.5
* Antisera produced in chickens; other produced in rabbits.
Five anti-Pinnipedia sera were tested. They
all gave distinct suborder reactions. One of the
two anti-sea lion sera was more specific than
the other and its reactions indicated that the
Ursidae were more closely related to the sea
lion than was the raccoon, fox or skunk. The
anti- walrus sera was aspecific; it gave fairly dis-
tinct subordinal reactions but the degree of reac-
tions with representatives of the Fissipedia
showed a similarity of the relationships of all
four families of this suborder to the Pinnipedia.
The reactions of the two harbor seal antisera
were interesting. The antiserum (C-165) pro-
duced in the chicken was highly specific and its
164
Zoologica: New York Zoological Society
[42: 14
reactions with the blood of the sea lion and
walrus were very low. It gave no reactions with
the raccoon, fox or cat and only slight reactions
with the polar bear and black bear. On the other
hand, the antiserum produced in the rabbit gave
reactions with the kinkajou and coati-mundi but
not with the raccoon, dog or cat. No explanation
suggests itself for the differences shown by these
two antisera.
Table 3 records the results secured with the
turbidimetric technic. Nine antisera were used
and of these four were produced against the sea
lion. These four antisera gave definite subordinal
reactions and the three that were tested by the
microdensitometer method indicated that the
Ursidae were probably more closely related to
the Pinnipedia than were the red fox or raccoon.
On the other hand reactions with antiserum-CU
were slightly higher for the kinkajou than with
the Ursidae and antiserum 3698-3796 gave reac-
tions with the skunk sera that were of similar
magnitude to that of the Ursidae.
The black bear antiserum gave distinct famil-
ial reactions and the cross-reactions with the
Pinnipedia were slightly higher than with the
raccoon, red fox and skunk. The high degree of
cross-reaction with the polar bear could indicate
that a closer relationship exists than the present
classification of these animals indicates. It would
be unsafe to argue this question on the evidence
of but one antiserum.
The anti-raccoon serum results showed the
expected close relationship between the raccoon
and the kinkajou. The Ursidae, Mustehdae and
Pinnipedia showed a closer relationship to the
Procyonidae than did the Canidae. In this par-
ticular series of tests the great difference between
the polar bear result and the black bear result is
strange considering the close relationship shown
between these two animals in other tests. The
authors cannot account for this variation but
future tests may give a satisfactory answer.
The anti-wolf serum results gave an excellent
example of intra-family relationships. The dog
showed an 88 per cent, relationship to the wolf,
while the red fox showed only a 52 per cent,
relationship. Of the families tested, the Ursidae
and Procyonidae seemed to be more closely re-
lated to the Canidae than were the mustelids.
The representatives of three families of Pinni-
pedia showed a remarkably similar relationship
to the wolf, while the cat (Felidae) was most
distantly related.
The anti-skunk serum had relatively low pre-
cipitating power and was highly specific. The
ferret and the tayra are members of the sub-
family Mustelinae while the skunk belongs to
the subfamily Mephitinae. The ferret and tayra
showed no closer relationship to the skunk than
did the Ursidae. The Pinnipedia, Canidae and
Felidae showed distant relationship while the
raccoon and hyaena gave no cross-reactions.
The anti-cat serum results showed strong in-
tra-family relationships but extremely weak in-
ter-family relationships with all but the Hyaeni-
dae. The intra-family results were peculiar. The
cheetah showed a 76 per cent, relationship to
the house cat while the tiger showed a 50 per
cent, relationship and the mountain hon showed
a 47 per cent, relationship. From these limited
data one gets the impression that the tiger and
mountain lion might belong to a different genus
than the house cat. These results warrant much
more research.
Discussion
The findings set forth above are at least a
beginning in the serological study of the rela-
tionship of the Carnivora. The species used in
the study were chosen primarily because of their
availability.
As indicated previously, authorities disagree
about the relationship of the families of the
Fissipedia to each other and to the Pinnipedia on
the basis of comparative anatomy and paleon-
tology. This has resulted in great variations and
confusion in the classification and nomenclature
of the Order Carnivora. On the basis of the sero-
logical data presented in this study, the following
tentative conclusions can be drawn:
1. The Pinnipedia are more closely related
to the Ursidae, Canidae, Mustelidae and Pro-
cyonidae than to the FeUdae and Hyaenidae.
2. The closest serological resemblance exists
between the Ursidae and Pinnipedia.
3. The Felidae and Hyaenidae are more
closely related to each other than to any other
family.
4. The polar bear seems to be quite closely
related to the black bear and possibly should be
included in the same genus.
If additional data support present findings,
it might mean that the nomenclature of members
of the family Felidae should be revised. Simpson
in his work on the classification of mammals
came to this same conclusion in his discussion
of the taxonomy of the Felidae.
To attempt a serological classification of the
Carnivora on the basis of the results presented
in this paper might be premature. However,
certain indications are given by these somewhat
limited data. The tendency to classify the Cani-
dae, Ursidae, Procyonidae and Mustelidae to-
gether into one superfamily (Canoidea) is per-
fectly valid serologically. Whether the Felidae,
1957]
Pauly & Wolfe: Serological Relationships among the Carnivora
165
Hyaenidae and Viverridae can be classified to-
gether into another superfamily serologically
will have to be determined in the future. There
are very strong indications that the suborders
Fissipedia and Pinnipedia do not exist serologi-
cally. Future research may show that there are
three or even four serological suborders of the
Carnivora.
The authors realize the shortcomings of this
paper as well as other serological research in
which animal relationships have been consid-
ered. The shortcomings are due to the lack of
materials to make more complete studies. It
would be much more preferable to concentrate
on fewer species but with larger numbers of
specimens so that a number of antisera could
be produced against each species and tested
against several members of each studied. In this
way it might be possible to make a statistical
analysis of the data and one that might yield
significant results rather than mere indications.
Summary
1. Serological tests were carried out using the
blood sera of 23 different species from nine fam-
ilies of the Order Carnivora, two sera of the
Order Artiodactyla, and human serum (Order
Primates) .
2. The ring test, the Baier microdensitometer
and the Libby photronreflectometer were all used
in performing these tests.
3. Where comparable reactions were made,
the results of the microdensitometer and pho-
tronreflectometer tests paralleled those of the
ring test.
4. A serological basis for the classification
of the Carnivora was indicated.
The authors are especially grateful to Dr.
Joseph G. Baier of the University of Wisconsin
in Milwaukee for the use of his microdensi-
tometer.
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1947. An analysis of photoelectric instruments
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1942. Quantitative serologic relationships within
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Boyden, A. A. & D. Gemeroy
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1933. The relationship of some common Am-
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1938. On the serological relationship of some
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Gemeroy, G. G.
1943. On the relationship of some common
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1951. Precipitin production in chickens. VI. The
effect of varying concentrations of NaCl
on precipitate formation. Joum. Immun.,
66: 225-236.
Leone, C. A. & C. W. Pryon
1952. Serological correspondence among three
species of Penaeid Crustacea. Journ. Elisha
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Libby, R. L.
1938. The photronreflectometer— an instmment
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372-379.
Nuttall, G. H. F.
1904. Blood Immunity and Blood Relationship.
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Scott, W. B.
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[1957]
Zoologica: Index to Volume 42
167
INDEX
Names in bold lace indicate new
genera, species or subspecies,- num-
bers in bold face indicate illustra-
tions; numbers in parentheses are
the series numbers ol papers con-
taining the plates listed immediately
following.
A
Achlya ambisexualis, 132
bisexualis, 132
glomerate, 132
Klebsiana, 132
sp., 132
Sparrowii, 132
Acinonyx jubatus, 160
Aethria carnicauda, 152, (13) PI. I
Agyrta dux, 153, (13) PI. I
Anlichloris trinitatis, 109
Aphanomyces laevis, 132
Astridia, 108
Autochloris trinitatis, 109
B
Bathygobius soporator, 99, (9) Pis. I,
II
Bison bison, 160
Bos laurus, 160
Bufo boreas boreas, 63, 65
boreas halophilus, 63, 65
C
Calonotos, 127
chalcipleurus, 128, 129
craneae, 128, (10) Pis. I, II, 152,
(13) PI. I
helymus, 127
hoffmannsi, 109
tiburlus, 127, (10) PI. Ill
tripunotatus, 129, (10) PI. Ill
Calyplralegnia achlyoides, 132
Canis familiaris, 160
lupus, 160
Chrostosoma, -116
viridipunctatum, 116, (10) PI. Ill
Collybus drachma, 52, (4) PI. Ill
Correbidia assimilis, 154, (13) Pis. I,
II
Cosmosoma, 117
achemon, 118, (10) PI. Ill
anoxanthia, 118, (10) PI. Ill
gemmata, 112
klagesi, 118, (10) PI. Ill
melalhoracia, 117, (10) PI. Ill
pytna, 118
remote, 118
rubriscapulae, 117, (10) PI. Ill
subfiamma subflamma, 117, (10)
PI. Ill
Cricosaura, 85
Ctenucha andrei, 155, (13) PI. I
D
Dixophlebia, 118
holophaea, 118, (10) PI. Ill
Dryas julia julia, 135, (12) PI. I
Dycladia, 122
basimaculata, 122, (10) PI. Ill
correbioides, 122
E
Episcepsis hypoleuca, 152, (13) PI. I
Eucereon maia, 153, (13) PI. I
obscurum, 154, (13) PI. I
Euchromiinae, 106
Eumetopias jubala, 160
Eupemphix pustulosus, 63, 64
F
Felis concolor, 160
domeslicus, 160
H
Heliconius Isabella Isabella, 135, (12)
PI. I
erato hydara, 135, (12) PI. I
melpomene euryades, 135, (12)
PI. I
ricini insulana, 135, (12) PI. I
Sara rhea, 135, (12) PI. I
Heliotropium indicum, 148, (13) PI. II
Histiaea, 123
cephus, 123, (10) PI. Ill, 151, (13)
PI. I
meldolae, 123, (10) PI. Ill
Homoeocera, 108
magnolimbata, 108
Homo sapiens, 160
Hyaena hyaena, 160
Hyla crepitans, 63, 64
crucifer, 63, 65
maxima, 63, 64
I
Isanthrene, 108
tryhanei, 108
Isoachlya monilifera, 132, (11) PI. I
K
Klauberina, 83, 86
I
Lepfodactylus typhonius, 63, 64
Leucotmemis, 116
albigutta, 109
hoffmannsi, 109
nexa, 116
thoracica, 109
Loxophlebia, 113
bisigna, 113, (10) PI. Ill
diaphana, 113
klagesi, 114
postflavia, 114, (10) PI. Ill
M
Macrocneme, 123
alesa, 126
eacus, 126
nigritarsus, aberration Irinitatensis,
126
plumbea, 123, (10) PI. Ill
spinivalva, 125, (10) Pis. I, II
thyra intacla, 124, (10) PI. Ill, 151,
(13) Pis. I, II
thyra, 124
vittala, 126, (10) PI. Ill
Mapeta xanthomelas, (13) PI. II
Mephitis mephitis, 160
Mesothen, 114
aurantegula, 114
desperate, 114
endoleuca, 114 (10) PI. Ill
pyrrha, 114, (10) PI. Ill
Methysia, 110
Mustela furo, 160
vison, 160
Mydropastea chrysonota, 109
N
Napala walkeri, 152, (13) Pis. I, II
Nasua narica, 160
Nyridela, 116
chalciope, 116
O
Odobenus rosmarus, 160
P
Panthera pardus, 160
tigris, 160
Paramya chrysonota, 109
? hoffmannsi, 109
Phaeo, 110
Pheia, 115
beebei, 115, (10) Pis. I, II
Phoca vilulina richardii, 160
Phoenicoprocla, 109
astrifera, 112
atrapennis, 112
aurantipatagiata, 112
auriflua, 112
biformata, 112
capistrata, 112
chrysorrhoea, 109
cubana, 112
exima, 113
flavipicta, 112
haemorrhoidalis, 112
insperata, 112
intermedia, 112
jamaicensis, 113
latimarginata, 113
lydia, 113
melachrysea, 109
mexicana, 112
168
Zoologica: Index to Volume 42
[1957]
nigricoxa, 112
nigri venter, 112
nigropeltata, 109
paucipunctata, 112
punicea, 112
rubriventer, 109
sanguinea, 112
schreiteri, 113
selecta, 112
sieboldi, 113
teda, 113
thomae, 112
trinitatis, 109
vacillans, 109, (10) PI. Ill
variabilis, 112
Pleurosoma, 108
trinitatis, 108, (10) PI. Ill
Poliopastea, 124
Potos oaudivolvulus, 160
Procyon lotor, 160
Prostherapis trinitatis, 63, 64
Protoachlya paradoxa, 132
Psarocolius decumanus, 87, 92, 93,
(8) Pis. I, II
virens, (8) PI. II
Pseudomya, 118
melanthus, 119, (10) PI. Ill
sanguiceps, 119
Pseudosphex, 106
deceptans, 107
kenedyae, 106, (10) Pis. I, II, 148,
(13) Pis. I, II
laticincta, 106, 107
melanogen, 107
Psoloptera, 122
leucosticta, 122
R
Rana catesbeiana, 33, 36-43, 48
clamitans, 63, 65
pipiens, 33, 36-43, 48
sylvatioa, 63, 65
Rhynohopyga, 119
flavicollis, 119, (10) PI. Ill
S
Saprolegnia delica, 132
ferax, 132
megasperma, 132, (11) PI. I
mixta, 132
parasitica, 131, 132
Saurita, 119
afflicta, 121, (10) PI. Ill
arimensis, 120, (10) Pis. I, II
Cassandra, 119, (10) PI. Ill
clusia, 119, (10) PI. Ill, 150, (13)
PI. I
concisa, 122
lacteata, 120
nox, 120
perspicua, 121
salta, 121
temenus, 121
Sphecops, 108
aurantiipes, 108, (10) PI. Ill
Steinegeria rubescens, 52
Syntomeida, 122
melanthus, 122
T
Taractes asper, 56
breevoorti, 56
longipinnis, 52, (4) Pis. I-III
miltonis, 56
platycephalus, 56, (4) PI. Ill
princeps, 56
raschi, 56, (4) PI. II
saussuri, 56, (4) PI. Ill
steindachneri, 56
Taxidea taxus, 160
Tayra, sp., 160
Thalarctos maritimus, 160
Thraustotheca clavata, 132
primoachlya, 132
Tursiops truncatus, 11, 13, (2)
Pis. I-III
U
Uca annulipes, 78, (6) PI. I
batuenta, 78
beebei, 78
cumulanta, 78
deichmanni, 78
dussumieri, 78, (6) PI. I
festae, 78
gaimardi, 78
galapagensis, 78
heterochelos, 78
heteropleura, 78
inaequalis, 78
insignis, 73, 78
inversa, 78
ischnodactyla, 78
lactea, 75, 76, 78
lalimana, 78
latimanus, (6) PI. I
leptodactyla, 78
limicola, 78
longidigitum, 78
manii, 79
maracoani, 79
marionis, 79, (6) PI. I
minax, 79
mordax, 79
oerstedi, 79
olympioi, 79
panamensis, 79
princeps, 79
pugilator, 79
pugnax, 79
rapax, 75, 79
rathbunae, 79
rhizophorae, 72, 76, 79
rosea, 79
saltitanta, 79
signata, 72, 79
speciosa, 79
stenodactyla, 79
stylifera, 79
terpsichores, 79
letragonon, 79
Ihayeri, 79
zamboangana, 72, 76, 79
Ursus americanus, 160
V
Vulpes fulva, 160
X
Xantusia, 86
riversiana, 83, 84, 85
Xiphophorus maculatus, 133
Z
Zalophus calif ornianus, 160
Zygaena parthenii, 112
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