with respect to the behavior of

the participating individuals. It does involve,

however, certain peculiarities of
position which are not characteristic of
individual challenging. In the most common
position the fish are about 1 inch
apart, broadside to each other, head to
tail, with their bodies flexed. The two
together resemble parentheses. They
usually remain in one place or drift slowly
backward, forward, or sideways. Rare-
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156 JAMES C. BRADI)OCK AND ZORA I. BRADDOCK
ly they revolve horizontally like a wheel
while maintaining the same position with
regard to each other. The position next
in frequency resembles the first, except
that the fish are head to head. They were
never observed to revolve in this position.
Other positions noted were a headon
figure with the bodies flexed in opposite
directions and a head-to-side arrangement,
resembling the letter T with
a tilted stem. Mutual challenges vary a
great deal in duration. The maximum
observed lasted 3 minutes and 30 seconds;
the minimum less than 15 seconds.
Also, they may be given in rapid succeschallenges
during the period prior to
fighting.
If mutual challenges are treated as
the individual challenges of two fish, i.e.,
added to the individual challenges of
each participant, the maximum number
of challenges per individual was 21, the
minimum 1, and the mean 9. For contact
pairs these figures were 35, 3, and
17, respectively. Some sort of challenging
occurred in all experiments and was exhibited
by every individual. The total
number of all challenges was 605
(Table 3).
These data contain no evidence of
TABLE 4
FREQUENCIES OF CHALLENGING IN PERIOD PRIOR TO FIGHTING (35 EXPTS.)
DURATION OF PERIOD (MINUTES)
CHALLENGES
0-2 2-4 4-6 6-8 8-10 10-12 12-14* 16-18 18-20
i no. of individual challenges/
pair .................... 3 4 6 7 4 10 4 4 17
Sno. of mutual challenges/
pair.................... 1 5 5 6 12 10 7 8 6
i no. of all challenges/pair (indiv.+
2X mutual+n)...... .5 13 15 17 28 29 18 20 29
* There were no periods having the time range of 14-16 minutes.
sion, or the interval of time between
them may be long. Behavior that starts
as an individual challenge by one fish
may become mutual challenging when
the pair-mate counterchallenges and the
instigator continues unchanged. As was
the case with individual challenges, the
mutual challenges continued throughout
the period prior to fighting and were not
more prevalent in one part of this period
than another. The total number of mutual
challenges for the 35 experiments was
199. They occurred in all the experiments.
The maximum per contact pair
was 15, the minimum 1, and the mean 6
(Table 3). It will be noted that the number
of mutual challenges was approximately
equal to the number of individual
correlation between the duration of the
period prior to fighting and the number
of challenges. Indeed, certain of the
shorter periods had more challenges than
the long ones (Table 4).
Counterchallenges are nothing more
than individual challenges given in response
to challenges by the pair-mate.
They are included among the individual
challenges mentioned above. Fifty-eight
of the 207 individual challenges recorded
for the periods prior to fighting were of
this type.
While challenging of one type or another
was by far the most characteristic
activity of the period prior to fighting,
other activities occurred and should be
recorded. Chasing was one of these. As
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AGGRESSIVE BEHAVIOR OF FEMALE BETTAS 157
the name implies, this involves rapid
pursuit around the aquarium of one pairmate
by the other. The pursuer may or
may not challenge while chasing. The total
number of instances of this activity
was 19. Ten of the seventy pair-mates
chased, and this form of behavior occurred
in 9 of the 35 fighting pairs. Considering
only those individuals that actually
chased, the maximum number of
instances per individual was 3, the minimum
1, and the mean 2. Corresponding
figures for contact pairs were 6, 1, and 2
(Table 3).
Retreating occurred as a response to
challenging, biting, chasing, or the approach
of the pair-mate without signs
of belligerence visible to the observer. It
was not necessarily rapid, and it merely
involved leaving the vicinity of the other
individual. The total number of retreats
during the period prior to fighting was
38. Fourteen of the seventy pair-mates
retreated, and the activity occurred in 12
of the 35 experiments. Among those individuals
that retreated, the maximum
number per individual was 11, the minimum
1, and the mean 3. For contact
pairs these figures were 15, 1, and 3
(Table 3).
Thirty-three of the seventy pair-mates
bit their opponents during the period
prior to fighting. The total number of
bites recorded for the period was 62. The
activity occurred in 22 of the 35 experiments.
Unlike all the other activities,
biting was especially characteristico f the
end of the period prior to fighting. In all
but two of the experiments where it was
observed, it occurred just before active
fighting began. Based only upon the data
recorded for those individuals that
actually bit, the maximum per fish was 5,
the minimum 1, and the mean 2. Corresponding
figures for contact pairs were
6, 1, and 3 (Table 3).
Exploring involves active swimming
into various regions of the aquarium,
poking the head into corners and the
gravel on the bottom, and maintaining
a generally alert posture. In contrast, the
fish often swim about in an apparently
aimless manner or rest quietly in one
place. The total number of instances of
exploratory activity was 28. Twenty-two
of the pair-mates explored. This type of
behavior was noted in 13 of the 35 experiments
where fighting later occurred.
Once again, considering only those cases
where the type of behavior actually took
place, the maximum number of instances
of exploring per individual was 4, the
minimum 1, and the mean 1. For contact
pairs these figures were 7, 1, and 2, respectively.
At intervals all individuals went to the
surface for air. Bettas are labyrinth
fishes, and this is part of their normal behavior.
When this happened, all social
activities ceased. No instances were recorded
where a fish was challenged,
bitten, or hindered in any other way
while this was taking place.
Reference to Table 3 indicates that
challenging of all types was by far the
most common activity during the period
prior to fighting. On the basis of the
number of participating pair-mates, biting
was second, followed by exploring,
retreating, and chasing in that order.
However, the large variability among
individuals and the small number of
cases observed make it unwise to attempt
to rank the relative frequencies of activities
other than challenging.
Various changes of color and pattern
occurred during the period prior to fighting.
Such changes are complicated and
should be the subject of further investigation.
One of the variables is the extent
of saturation of the color of the body
exclusive of stripes. Another is the
saturation of the color of the median
fins. The pectoral fins do not change
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158 JAMES C. BRADDOCK AND ZORA I. BRADDOCK
color but remain colorless at all times.
The pattern of the stripes which sometimes
adorn the body also varies. These
are always darker than the background
color and may be longitudinal, dorsoventral,
or plaid (a combination of the
other two). Finally, the saturation of the
color of the stripes themselves may vary.
Each of these will now be discussed separately
for the sake of clarity.
The background color of the body may
vary from pale to fully saturated.
Changes may occur rapidly and involve
passing through the intermediate stages.
An individual engaged in contact-pair
behavior may assume any phase of background
body color, depending upon the
nature of the behavior.
The degree of saturation of the color
of the median fins follows that of the
background color of the body in a general
way. Ordinarily, changes in one are accompanied
by similar changes in the
other. The saturation of fin color, however,
is slightly greater than that of the
body in all phases except that of full
saturation. A few cases were noted in
which there was little or no similarity
between the color phases of body and
fins. This suggests that they may be controlled
by separate mechanisms.
The patterns of striping are related to
a certain extent to the phases of the
background color of the body. Longitudinal
stripes occur only when the latter
is relatively pale. Dorsoventral stripes
have been recorded in combination with
all phases of saturation of background
color and are especially characteristic of
females in breeding condition. This is not
their sole significance, however. The
plaid effect has been noted only in combination
with very pale background
color, but it occurred so rarely that no
conclusions should be drawn concerning
the possibility that it may also accompany
other phases. It may be a transitional
stage between the two types of
stripe pattern. Any of the striping patterns
may change into any other without
passing through a third, and the changes
are sometimes very rapid. The color of
the stripes themselves generally tends to
become more intense as the background
color lightens, but such is not always the
case. The width of the stripes also varies.
In the absence of stripes the background
color of the body and the color of the
fins may be of any degree of saturation.
These facts would seem to indicate that
striping patterns and color phases of the
body and fins are all under the control of
separate mechanisms.