Siamese fighting fish, Betta

splendens, has long been known

for its highly aggressive behavior.
It is easily bred and readily available.
The species is relatively hardy. Specimens
are maintained in the laboratory
without difficulty. These factors recommend
it to students of aggressive behavior
among animals. Although there is
available a certain amount of literature
concerning B. splendens, much of it is of
a popular or semipopular nature and
often errs as to details. Furthermore,
most accounts of the behavior of the
species concern themselves exclusively
with the fighting behavior of the males
or with reproductive behavior (Smith,
1947; Noble and Borne, 1941). The late
G. K. Noble (1939) stated that the females
form hierarchies, but presented
no details.
The work reported here was undertaken
as a survey of the aggressive behavior
of the females. Its specific purpose
was to record in detail the various
activities that occur when two females
are placed together in a small aquarium.
Once known, such material might form
the basis for further investigations of a
more detailed nature.
MATERIAL AND METHODS
The observations reported here were
all made in a laboratory located on the
third floor of the Natural Science Building
of Michigan State College. The room
has three large windows facing north and
receives additional light from a battery
of fluorescent lamps on the ceiling. Other
fluorescent lamps are attached to the
racks upon which the smaller aquaria are
placed and are located directly over the
aquaria. All lights were turned on at
8:00 A.M. and off at 5:00 P.M. each day
during the observation period.
The laboratory is heated by the standard
radiators of the building. These are
located under the windows and well
away from the aquaria used in the experiments.
In addition, an electric heater located
high on the north side of the west
wall and controlled by a separate thermostat
prevents the room temperature from
falling below 250 C. A recording thermometer
indicated that the air-temperature
range during the period when these
observations were taken was 270 A 10 C.
The regular tap water of the building
was used after 2 weeks of aging and aeration
in 20-gallon crocks. This water originates
in deep wells maintained by the
college.
The fish were housed in containers of
two types. Except when actually under
observation and for a short period immediately
thereafter, they were kept separately
in wide-mouthed, 1-gallon food
jars, two-thirds full of water. Each of
these contained gravel, one snail, and a
stalk of Myriophyllum approximately 20
cm. long. Similar containers were used
during the recovery period after fighting
but contained only a weak, aqueous solution
of methylene blue as a precaution
against infection of the wounds with
fungus. Fish actually under observation
152
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AGGRESSIVE BEHAVIOR OF FEMALE BETTAS 153
were placed in an aquarium whose dimensions
were as follows: 30.5 X 26.5 X
26.5 cm. This was filled to within 31 cm.
of the top. It contained gravel and one
clump of Valissneria. All jars and the
aquarium were arranged on racks along
the south and west sides of the room and
down the center, running east and west.
Each morning the fish were given as
much grated frozen shrimp as they would
eat. This diet was supplemented by an
occasional ration of laboratory-reared
Drosophila.
All observations were made between
March 6 and June 18, 1953.
The fish were all adult females obtained
from a commercial distributor in
Detroit. Their ages were not known, but
they appeared to be young, probably
not more than nine months old, as suggested
by their general vigor, the high
rate of survival during and after the experimental
period, and the excellent level
of health maintained at all times. Sizes
were rather uniform; the six individuals
measured averaged 3.2 cm. from the anterior
end to the base of the tail fin. It
was not found advisable to measure
every individual because of the generally
uniform size and in view of the possibility
of injury.
When fish were to be paired, their relative
sizes were first recorded by means of
symbols. The categories were: similar,
where there was no apparent difference;
greater by 1, where one individual was
clearly larger than the other but the difference
was slight; greater by 2, where the
difference was somewhat more than
slight; and greater by 3, the largest degree
of difference that occurred. The
largest difference measured was 9 mm.
These criteria were not exact but were
useful in making rough comparisons.
Forty-five individuals were used in the
experiments. Fifty-one pairings were
made. Ten individuals were paired only
once. Twenty-four were paired twice.
Five were paired three times; three, four
times; one, five times; and two, six
times. No fish was paired more than
once with the same opponent. All were
isolated physically and visually for 7
days prior to pairing.
Before two individuals were placed together
in the aquarium, the jars containing
them were moved to a table close by.
Visual isolation was maintained while the
jars were left undisturbed on the table for
15 minutes in order to allow the fish to
recover from any disturbance resulting
from their being moved. They were then
rapidly netted, one after the other, and
placed in the aquarium. This never required
more than 30 seconds, and usually
the time consumed was a fraction of
that. Observations were started as soon
as both fish were in the aquarium and
continued until the fight was over, except
in those cases in which no fight occurred.
The latter were observed for at least 45
minutes. As far as possible, a written
record was kept of all activities observed.
These records of the activities of the
fifty-one pairings were later tabulated
and analyzed in various ways and form
the basis for the observations to be presented
here.
At the conclusion of each observation
period the fish were isolated in methylene
blue solution in order to guard against
infection with fungus. They were kept
thus from 3 to 7 days, depending upon
the extent of the injuries sustained in
combat. Thereafter they were returned
to their home jars, where they remained
until paired again, if such was to be their
fate. Most individuals were treated in
precisely this manner. A few that neither
fought nor sustained injury were returned
directly to their home jars. No
individual was paired again before its
wounds were completely healed, although
evidence of previous fighting
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154 JAMES C. BRADD)OCK AND ZORA I. BRADDOCK
often remained in the form of incompletely
regenerated fin membranes. As has
been previously noted, every fish spent
at least 7 days in isolation in its home
jar prior to each pairing.
For convenience in reporting results
the sequence of events in each experiment
was divided into the following sections:
1. Activities prior tofighting.-This period
included all activities from the moment
the two fish were placed together in
the aquarium until actual fighting began.
Cases where no fights occurred are
to be treated in another report. Conse-
TABLE 1
DURATIOONFSP ERIODS
PRIORTO F IGHTING
(35 EXPTS.)
Time No. of
(Min.) Expts.
0-2............. 1
2-4 ............ 9
4-6............. 9
6-8............. 9
8-10............. 2
10-12............. 2
12-14 ........... 1
14-16............. 0
16-18............. 1
18-20............. 1
quently, this paper is primarily concerned
with the 35 experiments in which
the pair-mates actually fought.
2. The fight proper.-In all cases this
began with the first rapid exchange of
bites and terminated with the surrender
of one of the fish. Sixteen of the fifty-one
pairings did not exhibit this phase.
3. Activities after surrender.-Each experiment
was observed for at least 5
minutes after fighting ceased. It was
thought necessary to place the fish in
methylene blue solution quickly, to avoid
the possibility of losing them through
fungus infection; therefore, no attempt
was made to observe the postfight period
for a longer time.
ACTIVITIEPSR IORT OF IGHTING
Active fighting does not begin immediately
after two female Bettas are
placed together in a small aquarium. It is
preceded by a period during which the
fish react to each other and to the physical
environment around them. In this
series of observations the period prior to
actual fighting varied in duration (Table
1). The maximum was 20 minutes, the
minimum 2 minutes, and the mean, 6.6
minutes In 28 of the 35 cases in which
fights occurred, the length of the period
prior to fighting was 8 minutes or less.
Activities characteristic of the period
prior to fighting are: reactions to being
moved, individual and mutual challenging,
counterchallenging, chasing, retreating,
biting, exploring, going to the surface
for air, and changes in color and
pattern. These will now be taken up in
order.
In 18 of the 35 experiments, one or
both fish of a pair reacted to the other as
soon as the second individual was placed
in the aquarium. In the other 17 experiments
a short period of time elapsed before
reactions occurred between them.
This appeared to be due to the experience
of being moved. Of the 17 pairs
whose reactions to one another were not
instantaneous, 3 pairs showed such reactions
at 15 seconds and 5 between 16 and
30 seconds. No special behavior was
noted during these short periods of time.
In the other 9 pairs, all but two of the
individuals remained stationary. The
two exceptions explored the aquarium.
They were not pair-mates. The durations
of this first phase varied from a
maximum of 180 seconds to a minimum
of 0 seconds. The mean was 33.4 seconds
(Table 2).
In every case the first reaction observed
between the pair-mates took the
form of challenging. In 22 of the 35 ex-
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AGGRESSIVE BEHAVIOR OF FEMALE BETTAS 155
periments the first challenges were individual,
i.e., given by only one of the two
fish. Individual challenging is a stereotyped
reaction. The aggressor approaches
the other fish, usually at an angle, and
stops at a distance of less than 1 inch.
It then erects its gill covers, holds its
body rigidly in a flexed position, erects
all its median fins, and rapidly vibrates
the pectoral fins. The color of the body
and fins becomes more intense, and striping
tends to disappear. Individual challenges
may be almost instantaneous or
prolonged for 30 seconds or more. They
may occur as a rapid series or with long
intervals between successive challenges.
This type of behavior continued throughout
the period prior to fighting, and the
In the 13 experiments in which the
first reaction between pair-mates was not
individual challenging, it was mutual
challenging. This occurs when two fish
challenge each other simultaneously. It
may have no significance apart from in-
TABLE 2
TIME ELAPSED BEFORE FIRST
SOCIAL REACTION
(35 EXPTS.)
Time No. of
(Sec.) Expts.
0-15........... 21
16-30........... 5
31-45........... 1
46-60.......... 2
61-75............ O
76-90........... .1
106-20...........
3
166-80............ 2
TABLE 3
FREQUENCIES OF VARIOUS ACTIVITIES IN THE PERIOD PRIOR TO FIGHTING (35 EXPTS.)
Individual
Mutual All
al
Chal- Chal- Chasing* Retreats* Biting* Explor- Chal- ig
lenges lenges lenges ing*
Totals for all expts. 207 199 605 19 38 62 28
No. pair-mates involved 60 70 70 10 14 33 22
No. expts. involved ..... 33 35 35 9 12 22 13
Max./indiv............ 15 15 21 3 11 5 4
Min./indiv.......... 0 1 1 1 1 1 1
Mi/indiv...1.
.... 3 6 9 2 3 2 1
Max./expt.. . 19 15 35 6 15 6 7
Min./expt 1 3 1 1 1 1
v/contact pair.......... 6 6 17 2 3 3 2
* Maximum, minimum, and mean figures apply only to those experiments where activity occurred.
data contain no evidence that it was
more prevalent early or late in the period.
The total number of individual
challenges for the 35 experiments was
207. Both pair-mates exhibited individual
challenging in 27 experiments, one
only in 6, and neither in 2. The maximum
number of individual challenges per fish
during the period prior to fighting was
15, the minimum 0, and the mean 3. For
contact pairs these figures were 19, 0, and
6, respectively (Table 3).