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The aggressive behaviour between territorial Pseudocrenilabrus multicolor (pisces, cichlidae) with unlimited or only visual contact
Behavioural
Processes,
13 (1986) 353-366
353
Elsevier THE
AGGRESSIVE
MULTICOLOR
BEHAVIOUR
(PISCES,
TERRITORIAL
BETWEEN
WITH
CICHLIDAE)
PSEUDOCRENILABRUS
UNLIMITED
OR
ONLY
VISUAL
CONTACT
Anders
Fern6
Department
of
1839, N-5011
(Accepted
15
Fisheries Bergen,
July
University
Biology,
of Bergen,
P.O. Box
Norway
1986)
ABSTRACT behaviour Fern6, A., 1987. The aggressive between Pseudocrenilabrus multicolor (Pisces, Cichlidae) with only visual contact. Behav. Processes, 13: 353-366.
territorial unlimited or
between development 0 f aggression territorial fish has experiments using visual been studied in laboratory only contact. In this comparisons of the aggressive behaviour study, were made between conditions of unlimited and only visual contact. Fighting cichlids in direct contact between two territorial decreased over time in a way similar to the aggressive activity between subjects permitted only visual contact. However, there were also important differences between the two conditions of contact. Low-intensity aggression with a distance between the subjects but was generally not occurred frequently in free interaction observed visual contact. presence of under conditions of The non-territorial fish increased the aggression between subjects with visual contact had no influence on fighting and but conversely decreased low-intensity between interacting aggression freely subjects. In addition, the low level of aggression after some time of unlimited contact was not transferred to a subsequent period of visual contact. indicate that the decrease of The results aggression visual contact is caused under conditions of by habituation, whereas in a interaction a balance is also free developed between attack and escape tendencies. The
mainly
Key words:
aggression,
habituation,
territory,
cichlid
fish
INTRODUCTION
The the
aggressive
expenditure
behaviour of an animal's
it to the risk of physical territorial
0376-6357/86/$03.50
animal
to
0 1986 Elsevier
in
defence
of a territory
time and energy
injury.
It is therefore
behave
in
Science
Publishers
such
involves
as well as exposing important
for
a way as to maximize
B.V. (Biomedical
Division)
a the
354
benefits
of the aggressive
behaviour
1964).
Especially
crucial
towards
territorial
conspecifics.
neighbours enemy
is generally
effect
In
(Wilson,
field
aggression
(Lowe-McConnel,
some
In
on
frequent (Fernald
and Hirata,
sites
(McKaye,
1977),
found
behind
waning and
been
has
observed result
between of
Peeke,
fish
has,
were
contact
sexual
breeding
fish has been where
fish
tube generally
wanes
(e.g.
Peeke
et.
and Peeke,
and Veno, to a
it has
another
cichlids
1972;
Peeke
et
1979),
al.,
1970) and
the
1976). The response
process
of
habituation
that the low level of aggression territorial
reinforcement
aggression however,
fish
and Molen,
found
between
rarely
establishment
is
often also
1969; Peeke
a and
physical
1980).
than one with
contact unlimited
may
interacting
investigated.
Siamese
conditions
between
freely
been
in the
(Cain and Baenninger,
without
situation
In the
fighting
of unlimited It was proposed be associated
contact
fish,
and only that
with
and consequently
a
less more
lead to habituation.
There
is
one
conditions
convict
the decrease contact.
laboratory
of unlimited
(Peeke and Peeke,
pairs
of
of dominance
differences
of
or
towards
glass
(see e.g. Assem
development
context
both
e.g.
1978)
territorial
1966; Peeke
interacting
habituation
territorial
easily
a for
been attributed
freely
resource,
is
1973).
The
visual
in
al.,
suggested
1984).
fish
experiments,
displayed
1969; Peeke
(e.g. Peeke,
has generally
it
between
or
et
1977; Holzberg,
and Colgan,
evident.
aggression
partition
of
observed
territorial
in laboratory
(e.g. Baenninger,
stickleback
Loiselle,
for a limited
of aggression
Gallagher
anabantoids
dear
a low level
often
1983; Grant
has been demonstrated
1971;
al.,
is
between
is often
the
glass This
to territorial
the so called
1977; Kodrick-Brown,
in more detail that
a
rapidly.
1968;
and competition
development
(Brown,
of aggression
in fish,
neighbours
aggression
mates
been
territoriality
1979; McKaye,
more
The
to strangers,
Albrecht,
et. al.,
investigated
the costs
regulation
The responsiveness
territorial
species
the
1975).
1956;
1978; Colgan,
and minimize
be
less than
studies
between
may
study
1982). The aggression
cichlids
with
unlimited
was less pronounced
Aggression
decreased
between
in stable
on
and conditions
the
freely but
of only
between
contact
than under
periods
territorial
fish visual
territorial
decreased conditions interacting
increased
with
over of
under contact pairs
time but visual
territorial transitions
355
bet.ween
stages
explained
as an interaction 1970,
There
in
fish. Aggression
of individuals
There
may,
as
is thus
and also
between
presence
or absence
natural
conditions.
on the aggressive was also
aggression situation
MATERIAL
found with
reducing
Groves
and
to
been
1985). between
of habituation
unlimited
visual
and
compare
the development
contact
and to examine
visual
of
information
influence
of a period a
subsequent
contact
was derived
fish which
in order
unlimited
also
species.
experiments
investigated
the
Recognition
may occur
The importance
and visual
in
has
1979; Myrberg,
of
territorial
changing
1979).
aggression
conditions
behaviour
between
that habituation fish.
different
from
during
and
that habituation
after
et. al.,
(e.g. Thresher,
of non-territorial
visual
e.g.
indicating
(Colgan
Additional
The
(see
increased
was undertaken
situation.
were
(habituation)
of aggression
evidence
of conclusions
free contact
contact
males
between
study
studies
sunfishes
in both unlimited
aggression transfer
field
territorial
differ
The present
processes
results
The
decremental
1983).
factor
some
interacting
however,
contact
a
for in damselfish
freely
between
regulation
the
between
of individual
appearance
claimed
some
also
cycle.
reproductive
and Peeke,
are
involved
is
the
(sensitization)
incremental Thompson,
in
may
vary
of unlimited period
of
to
from
was
transferred
a the
under contact visual
to see if the attenuation
contact
of the
to
of a
contact.
AND METHODS
cichlid Pseudocrenilabrus The small, mouth-brooding fish multicolor (Schoeller, formerly Haplochromis multicolor, Hilgendorf chosen for or Hemihaplochromis multicolor, see Wickler, 1963) was can be established in this study as several territories growth of middle-sized aquaria. The distribution, migration and this East African cichlid have been investigated in the field conducted (Welcomme, 1969), but only laboratory studies have been on the 1937; reproductive and territorial behaviour (Peters, Reinboth, establish 1956; Kuenzer, 1975; Mrowka, 1982). The males territories and are then generally brightly coloured with a bluish body and black eye-bars whereas females are more cryptically coloured. There is no pair-bondi_ng and after spawning the female leaves the male's territory with the fertilized eggs in her mouth. The fish in this study came from a population of P.multicolor originating from a local dealer and held The at the laboratory. territorial varied in length 5.7 cm males between 4.0 and (including the tail fin). The two subjects in each trial had a mean length difference of than 2 mm and were isolated from each less other at least three weeks prior to contact in the trial. Aquaria of 200 1 (100x50~40 cm) were used. The floor of the tanks was covered by sand and devoid of plants. A hide made by flower-pots and stones was placed at where the each end wall
356
subjects centered their territories. The aquaria were illuminated from above by three 15 W bulbs with a photoperiod from 11 a.m. to 11 p.m. The temperature was held at 25 '1.0" C. The fish were fed boiled liver or frozen fish flesh at the beginning and end of the light period. opaque partitions. Each aquarium was bisected with glass and One subject, along with four or five smaller conspecific males and aquarium and allowed to females, was placed in each half of the establish a territory. After five days the opaque partition was removed and, in Exp. 2, the glass partition was also removed. were made with only visual contact In Exp. 1, six trials fish and the opaque between the test subjects. Non-territorial time t partition were removed five min before (around 1 p.m.). Observations were made for one hr at time t and for 30 min at time 30 min at tf3 hrs during the first day of visual contact and for time t the following six days. In two trials, non-territorial fish placed in each half 0 f the (two males and two females) were aquarium after the observation on Day 7, and observations were made for 30 min at time t the following five days. Two of the trials were In Exp. 2, nine trials were conducted. one of the subjects lost his not included in the material as the trial was terminated. territory to the other subject before The subjects were first studied for a period with unlimited contact in the presence of non-territorial fish. (In trials l-3 this period lasted 14 days but as no change over time was found after the first five days in trials few days of contact., this period was set to 4-7, and only data from Days I-5 will be presented for all trials). time t and On Day 1, the partitions were removed five min before subjects were observed for one hr at time t and for 30 min at the were made for 30 min at time t + 3 hrs. On Days 2-5, observations time t. fish was then included in A period without non-territorial Exp. 2. The day after the period with non-territorial fish present, observations these fish were taken away five min before time t and were made for 30 min at time t for five days. periods without nonwith and The sequence of the territorial fish was thus not the same in Exp. 1 and Exp. 2. The reason for inital absence of non-territorial fish in Exp. 1 the usually include such was that visual contact experiments do not variable situation. In Exp. 2, fish because of the resulting absent,one subject pretests showed that if non-territorials were partition was removed. always lost his territory when the unlimited contact in Exp. 2, The day after the periods with the glass partition was reintroduced in the middle of the aquarium observed for seven days five min before time t and the subjects visual contact with the same procedure as in under conditions of Exp. 1. by one observer, Observations were made from behind a screen spoken into a tape recorder and later transcribed. The following behaviour patterns were recorded: Both unlimited
and only
visual stroke
contact
Tail-beat
A strong
with
the tail.
Low-intensity
subject or both subjects made a aggression One towards (approach), away distinct movement and away (approach--retreat, (retreat), or towards frontal in relati.on to the other subject. display) (The frontal displays were few and of short duration and were therefore included in low-intensity aggression). In free interaction, both subjects
351
generally at a distance from the territorial were border. Under conditions of visual contact, low-intensity aggression occurred when the was territorial border of one or both subjects not in contact with the glass partition. Sexual
behaviour Quivering often quivering motion.
Fluttering Specific
A swimming unlimited
to
movement
followed
along
by
the walls
swimming
with
of the aquarium
contact
Attack
A rapid
Fight
Various combinations of circling, tail-beating, attacks. lateral threat, mouth-fighting and The types 0 f behaviour in a fight were not recorded. A fight consisted often of long series of attacks and more than one reciprocated attack was defined as a fight.
Specific
to
only
thrust
visual
towards
another
fish.
contact
towards the glass partition A subject swam swimming up and down the glass partition oriented towards the other male and often bit at the glass panel.
Aggressive
these behaviour patterns by The frequency of performance of the subjects towards each other was recorded as well as the time Attacks and spent fighting, aggressive swimming and fluttering. sexual behaviour patterns towards non-territorial fish were also recorded. When the subjects did not exhibit any of the behaviour patterns described either swimming relatively above, they were were passively, digging or staying within the hide. The subjects brightly coloured during all the defined behaviour patterns except for fluttering. The size of a subject's territory was recorded to the nearest five cm in each observation as the distance from the end wall of the subjects during aquarium to the position bet.ween the low-intensity aggression (assuming that the territorial border ran parallel with the end walls). RESULTS Only
visual
The
contact
aggressive
swimming
displayed
by both
swimming
decreased
Trend
previous
Lehman,
of
Biting
aggressive
swimming,
downward
contact
and
trend
no bites
Experiments
the glass
occurred
the same
in Exp.l,see
Fig-l).
observed
in
1). In Exp.
swimming
tail-beating
9
out
of
aggressive
2,
with
the subjects, and
a
(p
during
seven days the
13
a
there
subsequent
frequently
tendencies After
was usually
1,
to a low on Day 7 (p
between
and showed
were
1 and 2.
of aggressive
Exp. 1
on Day
contact
partition
In
1975, see Fig.
unlimited
increase
decrease.
the
towards
simultaneously.
from a high mean
test,
period
was an initial
subjects
the for
of visual trials
of
358
15
AGGRESSIVE SWIMMING
* 1\
EXP.l EXP. 2 o----o
,y?
0' I
BITE
', \
-7
FLUTTERING
LOW-INTENSITY
1
2
3
4
5
6
TAIL-BEAT
7 DAY
frequency of different behaviour duration or mean Fig.1 The made during observation patterns per territorial male and 30 min were periods contact. (Two observation visual seven days of In Exp. 2, the males had a previous period of conducted on Day 1). Significant differences between observations contact. unlimited fluttering in both and were found for aggressive swimming, bite in Exp. 1 and low-intensity agggression in Exp. 1 and 2, tail-beat denote significant Asterisks
359
Experiments
When
and tail-beating
p
the
Exp.
and
biting
swimming
tail-beating
when
in Exp.
2 had more on Day
difference
3 than
Exp.
maximum
were
1
or
in Exp.
(Day
2
was no
biting
activity
Exp.
U
aggressive
There
swimming,
and
1)
Day
2 (Mann-Whitney
1 (p < 0.05).
with
Day
swimming
on Day 2 and more
aggressive
the observations period
period
1 than Exp.
biting
in either
observation
the aggressive
observation
first
frequent
< 0.01).
significant
(first
in more
significantly test,
1 and 2 are compared,
2)
1
were
compared. The decrease increase
in
in
time
difference
was
significant
upward
0.001, In
test). 1,
with
either
only the
the glass
first
aggression and
aggression
with
the glass
one
subject
subject
after
infrequently
the usual
trial.
No
subjects
for
the
The addition
attack
rate
and sexual
marked
increase
(mean
aggressive
observation marked
change
increase
aggression several fish
over
in aggression
of the variable
time).
the
towards
30
Fluttering
these
situation
been
of
the other
added
observed
fluttering only
days
the
a
never
in a
subjects
with
occurred.
neighbours
(see Discussion).
was high
of additional
observation
almost
border
fish but also
between
the territorial
test)
in contact
fish were
in not
min
in
1 and 2.
then
five
1 and
than
from the glass
whereas
days
activity
per
between
had
resulted
during
male
Exp.
towards
in the aggressive
3.6 min per
behaviour
contact
Trend
territorial
cm distance
2,
Low-intensity
territory
in both
activity
swimming
0.01).
1, non-territorial
in Exp.
of other
frequent
claimed
Sexual
observed
trials
common.
result
the trial).
in
1
and
7)
of Day
0.001;
the
in
1). In Exp.
territory
more
a
of Day
Day
2 (p <
IO-15
at
until ( Fig.
<
but
territories
observation
(p
(in one trial
two
between
1
in Exp.
generally
however,
throughout
was
In
partition
observed claimed
Exp.
significant
observation
occur
the first
time
subjects
was,
by the first
an
1 and 2 (p <
defended
significantly
of
over
from Day 2 both
in
No
by
observations Experiments
generally
seldom
were
observation
in both
subject
partition
decreased
partition
was
1).
(Fig.
this did not
one or neither glass
found
partition
where
aggression
low-intensity the
was
accompanied
consecutive
subjects
in one trial
low-intensity
with
the
was
fluttering
between
trend
Trend
(except
spent
found
Exp.
contact
swimming
aggressive
no The
may be a
360
ATTACK OTHER
'*i
t t2
50
/
f-%h
I I
SEXUAL
E IL
i
BEHAVIOUR
, I
10
IL*
LOW-INTENSITY
*m g* t
I
1
1
OTHER
I
I
I
2
3
4
FISH
PRESENT
I
I
I
12 OTHER
I
1
I
3
4
5 DAY
FISH
ABSENT
or frequency of different behaviour duration mean The Fig.2 during observation 30 min and male territorial patterns per The initial five days were in the presence of contact. unlimited these nondays five subsequent while in the conspecifics, were periods removed. (Two observation were fish territorial Significant fish). with other Day 1 in the period conducted on lowfight, for found were observations between differences other intensity aggression and sexual behaviour in the period with fish (Friedman's test, p <0.05). Asterisks as in Fig. 1.
361
Unlimited
When
contact
the
partitions
were
removed
first
swam randomly
subjects first
time
Trend
of Day
was
hand
period
the aquarium 2, all fish at
fights
between
a few minutes.
the
During
the
up to 5.3 min occurred.
the
the first
or
subsequent
The
observations
aggression
and second
(p <
increased
observation
signed-ranks
other
period
or
subjects
the
these
upward
The territorial than
the
except
often
not
rate
(Wilcoxon days
2-5
however,
2).
for their
near
occurrence
the
and sexual
activity
in
territorial
activity
(p < 0.05,
with
the middle
from 25 to 75 cm.
and
seldom
changed
five cm from one day to another.
DISCUSSION
The
development
territoral be compared multicolor activity visual
of
fish coming in this males
contact.
study.
decreased
in contact
with
There
between
the
occurred
frequently
the
aggressive
into unlimited Fighting over
time,
the glass
were,
two conditions during
also
of contact. free
freely
similar
partition
however,
behaviour
and only visual between
interaction
to
a
Fig. 2).
around
in size
stable
by the
observation
located
ranged
was relatively
(p <
did
during
low in the first
was generally
time
fish
observed.
observed
for sexual
no but
over
increased,
see Fig.
infrequently
but territories
border
mean
aggression
The aggressive
border
aquarium,
on
or attack
(p < 0.05,
fish was
trend
decrease
time
observed,
fish were
The territorial the
removed
surface.
towards
significant
test
was also
Non-territorial border
the fighting
was seldom
Fluttering
slight
fish yielded observations
of non-territorial
Low-intensity
fish were
Tail-beating fights.
non-territorial consecutive
a
absence
matched-pairs
the two periods).
without
showed
time
presence
influence
more
in
of
in Exp.
actual
after
lasting
between
significantly
of
halves
2). Low-intensity
between
difference
fighting
O.O5).The
when
and
until
reduced
see Fig.
following
significant
in
fro
and
the
of a trial
1.
The
the
to
long fights
test,
other
the
beginning
not observed
were
fighting
separating
the
observation,
0.01, on
at
between
contact
can
interacting the
between
subjects
important Low-intensity and showed
p.
aggressive with
differences aggression a
certain
362
increase
initially
separated
by a glass
fish
also
Response a
waning
(Peeke with
behind
a
glass
indicated
by
only
A
cichlid
and
with
differently
in the
subjects
with
not involved
The changes
over
explained
if
development
an
in
and
territorial fighting
and
attack-escape
assumes
Vodegel, in another
been
there
activated
lead to
retreated
retreated
presented
therefore
in
be
better
addition,
attack
and
by
border.
stimulation to the
decrease
aggression. e.g.
in or
and the opponent
the escape
too close
observed
low-intensity indicated
the
when
is, the
the
and
opponent
decreasing
however,
or coming
and
with
and as
P.multicolor.
by the aversive
crossing
of is
in the
to the territorial
approached
increased
in the present
interacting
between
approached
subjects
(Ferns,
by each other
can,
aggression
increased
fish
waning
involved
freely
low-intensity
removal
from other
stimulated
of
The
as that
1975;
incidents
not influence
pattern
found
in relation
could
increase
(see also Ferno,
may
study,
(Rowland,
that
when
equilibrium
1977)
this
readiness
in the reponse
equilibrium
This
in
a behaviour
isolation
have
may become
border.
1975; Bols,
in
fish
system
between
attacks
response
stimulus
rapid.
the subjects
tendencies
fights
a
to
attack
in this situation
one
of
tendencies
time
from
has been
attributed
An avoidance
of aggression
were visually
also
may
stimulus
study,
generally
of fluttering,
long time
aggression
is
this
(Rhoad et al.,
was relatively
of
in
1973).
decreased
after
Habituation intensity
were
were
non-territorial
between
feed-back
a withdrawal
as the subjects
the decrease
Peeke,
inappropiate
general
species
and
in the decrease
1978) is probably
and
the subjects
of
aggression-eliciting found
in fish
the increase
may correspond
an as
partition
be involved
1978).
towards
demonstrated
connection
Escape
when
behaviour
to an increase
partition,
habituation
of
the aggressive
leading
glass
repeatedly
study
observed
The presence
contact.
behind
also
partition.
influenced
two situations, visual
but was seldom
the
which
An frequent
one one
where
of
subject subject
was oriented
laterally
1987). of non-territorial
the fighting
low-intensity visual other
was,
in a different
added.
on
comparable,
this
the other
did
the subjects between
Aggression
Although
sequence
interaction
rate between
aggression.
contact
fish were
not strictly
fish in a free
time or attack
hand,
the two
markedly
conditions
in the two experiments
and
observation
the
stresses
363
difference
between
of visual
the two conditions other
contact,
fish
aggressive
response
habituation
to take place
be
a
balance
independent
between
by
subjects
the
devoted
again.
attack
aggression
in
significantly trials
following
different
way.
intensity
wall
glass
in the aquarium
explanation
is
restricted
received
aggression
in
contact
explanation
transfer
to
defended in
which end
the of
territories territory The period
is
border the
were
influence
unequal,
of a period
study.
made
when
However, the
contact in three
glass
first
of visual
a
after
trials
of the aquarium whereas
with
contact
removed
increase.
in
the
the glass
2
could
of the subjects
partition
with
time,
result,
observation
subject
while
in the
fights
should
contact,
in Exp.
was
and
that
partition over
As
on
was not systematically
partition
of
glass
tendency
neither
the glass
the
the
glass
partition
in contact
a
contact,
contact
decreased
in the middle
trials
with
of visual
unlimited
only
a territory
of unlimited
this
with
increase
from direct
partition.
the
of
if one assumes
from the
which
with
was located
period
defended
by in
a
increased.
from the attacks
contact, in contact
the
in contact
the glass
with
in
of the reduced
the placing
of the escape
glass
the
supported
visual
territories
the
stimulation
through
of the
before
aggression
strength
low not
delay
arose
a distance
of
level
a
decreased
by
observation
with
level
This
partition
of aggression
fish. was not
(dishabituation).
interpreted
Low-intensity
as aversive
be
This
aggression
to
and then
which
the
was initially
interaction,
the glass
first
The
lead to an immediate
better
level
the
contact.
contradicted
due to a decreasing
subjects not
the
occurred.
frequently probably
be
these
contact
was the cause
changed
equilibrium,
during
low-intensity
with
could
attack-escape
still
should
therefore
in contact
partition
free
situation
the
result
The
contact
actually
because
in the beginning
level
alone
during
with
observations
the
habituation
when
aggression
from
was
perhaps
partition
for
could
in the subjects
unlimited
of visual
the glass
by unlimited
If
of aggression
aggression
the
with
in
the
fish,
to interacting
period
of the
there
aggression
during
contact
not preceded
similar
time
conditions
may be too variable
tendencies
other
of aggression
to a subsequent
increased
of
considerable
transferred
but
and escape
presence
Under
a dishabituation
In free interaction,
Low-intensity
fish.
attenuation
The
of contact. produce
and the new situation
of other
decreased
could
a
largest
partition. subsequent
investigated
observations after
at
when
the
in were
trial.
364
Fighting
was then lost
subjects observation
that
transferable Siamese
This
territorial
It seems,
however, between
balance
experiments
with
response
between
attack
patterns the
most
attack
aggressive
response
unlimited
experiments
and
using
be transferred be presented
A similar
interaction
visual
visual
contact
to free interaction. in another
paper
aggression
with
situation.
(Vodegel,
a In
1978),
strength
decreased
of certain
behaviour
probably
with
determines
a different The
therefore
Further
(Ferns,
did
by an interaction
contact.
should
of
in this zebra
whose
in El. multicolor
only
not
in the
1980).
interacts
be explained
tendencies
is
found
in free interaction.
time and by the performance
(e.g. biting).
the the
stimulation
habituation
tendencies
could
of
supports
(Lobb and McLain,
et. al.,
Pseudotropheus
models
escape
and
combats
that habituation
and escape
the cichlid
over
physical
can not take place
likely
been
to visual
that
one
This
has also
habituation
suggest
trials day.
in one situation
This
1976; Meliska,
fish
towards
spontaneously
between
not
all one
of aggression
in actual
and Meliska, does
within
situation.
fish, where
aggression
paper
between
the
a decrease
fighting
1976; Meliska
seen and in
territory
to another
reduce
not
frequently
his
outcome
results
from
not uncritically
evidence
for this will
19871.
ACKNOWLEDGEMENT
I
am
Radesater Karin
indebted for
Pittman
to
Dr.
constructive for valuable
Nance
Vodegel
criticism. remarks
and
professor
I also wish
Tommy
to thank
Miss
on the manuscript.
REFERENCES Albrecht, H., 1968. Freiwasserbeobachtungen an Tilapien (Pisces. Cichlidael in Ostafrika. 2. Tiernsvchol., 25, 377-394. J. van Assem, den and Molen, J.N. van der, 1969. Waning of the aggressive three-spined response in the stickleback upon exposure to a conspecific. constant I. A preliminary analysis of the phenomenon. Behaviour, 34, 286-324. of aggressive motivation in Betta Baenninger, R., 1966. Waning solendens. Psvchon. Sci., 3, 241-242. R.J., 1977. Bols, Display reinforcement in the Siamese fighting fish, Betta spendens: Aggressive motivation or curiosity? JCamp. phvsiol. Psvchol., 91, 233-244. Brown, J.L.,1964. The evolution of diversity in avian territorial systems. Wilson Bull., 76, 160-169. and the Cain, N.W., and Baenninger, R., 1980. Social organization maintenance of aggressive behaviour in community - housed male Siamese fighting Learn. and fish (Betta splendens). Anim. Behav. 8 171-176. -1-1 Colgan, P.W., Nowell, W.A., Gross, M.R. and Grant, J.W.A., 1979.
365
rim Aggressive habituation and circling in the social organization of bluegill sunfish (Lepomis macrochirus). Env. Biol. Fish., 4, 29-36. Field study of Haplochromis Fernald, R.D. and Hirata, N.R., 1977. burtoni: Quantitative behavioural observations. Anim. Behav., 25, 964-975. Fernij, A., 1978. The effect of social isolation on the aggressive and sexual behaviour in a cichlid fish Haplochromis burtoni. Behaviour, 65, 43-61. between Fernd, A. 1987 The aggresive behaviour territorial Astatotilapia burtoni (Pisces, Cichlidae) with unlimited or only visual contact emphasizing low-intensity aggression during free interaction. Behav. Processes Gallagher, J.E., Herz, M.J. and Peeke, H.V.S., 1972. Habituation of aggression: the effects of visual social stimuli on behaviour between convict cichlids (Cichlasoma adjacently territorial nisrofasciatum). Behav. Biol., z, 359-368. Grant, J.W.A. and Colgan, P.W., 1984. Territorial behaviour of the male johnny darter, Etheostoma nisrum. Environmental Biology of fishes 10 261-269. Groves, PyM?nd Thompson, R.F., 1970. Habituation: a dual-process theory. Psvchol. Rev., 77, 419-450. behaviour Holzberg, S., 1978. A field and laboratory study of the and ecology of zebra (Boulenger), an endemic Pseudotropheus Malawi (Pisces, Cichlidae). 2. Zool. Svst. u. cichlid of Lake Evolutionsforsch., 16, 171-187. of breeding Kodric-Brown, A., 1978. Establishment and defence Anim. territories in a pupfish (Cvorinodontidae: Cvprinodon) Behav., 26. 818-834. der ausl6senden Reizsituationen fiir die Kuenzer, P., 1975. Analyse Anschwimm und Fluchtreaktion junger Eindring Hemihaplochromis multicolor (Cichlidae). 2. Tierpsvchol., 2, 505-545. Lehmann, 1975. Non-parametrics. Statistical methods based E.L., on ranks. MC Graw - Hill International Book Company. INew York Lobb, M.L. and McCain, G. Procedurally related differences 1976. Anim. in the aggressive behaviour of Betta splendens (Rerranl. Learn. and Behav., 4, 367-373. Colonial breeding by an African substratum Loiselle, P.V., 1977. Biolosv of spawning cichlid fish, Tilapia zillii (Gervais). behaviour, 2, 129-142. Lowe-McConnel R.H., 1956. breeding behaviour of Tilapia The species (Pisces. Cichlidae) in natural waters: Observations on karomo variabilis Boulenger. T. Poll and T. Behaviour, 9, 140-163. MC Kaye, K.R., 1977. Competition for breeding sites between the 291-302. cichlid fishes of Lake Jiloa, Nicaragua. Ecolosv 58, MC Kaye, Ecology and breeding behaviobr of a cichlid K.R., 1983. fish, Cvrtocara eucinostomus, lek in Malawi, on a large Lake Africa. Env. Biol. Fish, 8, 81-96. Meliska, J.A. and Meliska, C.J., 1976. Effects of habituation on threat establishment in Siamese display and dominance the fighting fish, Betta solendens. Anim. Learn and Behav., 4, 167-171. Meliska, C.J., Meliska, J.A. and Peeke, H.V.S., 1980.Threat display and combat aggression in Betta following visual salendens exposure to conspecifics and one-way mirrors. Behavioural and Neural Biolosv a, 473-486. Mrowka, der i982. Untersuchungen zum Enstehen w. I Brutpflegebereitschaft des Kleinen Maulbriiters Pseudocrenilabrus multicolor (Cichlidae, Pisces). Z. Tieresvchol., s, l-24. Myrberg, A.A. Acoustically mediated and Riggio, R.J., 1985.
366
reef individual recognition by a coral fish (Pomacentrus partitus). Anim Behav., 33, 411-416. Peeke, H.V.S., 1969. Habituation of conspecific aggression in the three-spined stickleback (Gasterosteus aculeatus L.). Behaviour, 35, 137-156. Peeke, H.V.S. and Habituation of conspecific Peeke, S.C., 1970. aggressive response in the Siamese fighting fish (Betta selendens). Behaviour, 36, 232-245. Peeke, H.V.S., M.J. and Gallaqher, J.E., 1971. Chanqes in Herz, adjacently aggressive interaction in territorial convict cichlids (Cichlasoma A study of habituation. nisrofasciatum): Behaviour, 40, 43-54. Peeke, H.V.S. and Peeke, S.C., 1973. Habituation in fish with special reference to intraspecific aggressive behaviour. In: Habituation: studies Behavioural (Peeke, H.V.S. and Herz, M.J. eds.) Academic Press New York, Vol.1, pp. 59-83. Peeke, H.V.S. and Veno, A., 1976. Response independent habituation of territorial three-spined stickleback aggression in the (Gasterosteus aculeatus). 2. Tierpsvchol., 40, 53-58. Peeke, H.V.S., H.H. and Peeke, S.C., 1979. Motivational Avis, variables and the sensitization and habituation of aggression in the convict cichlid Cichlasoma nisrofasciatum). zTierpsvchol., u, 363-379. Peeke, H.V.S. and Peeke, S.C., 1982. Parental factors in the sensitization and habituation of territorial aggression in the cichlid nisrofasciatum). J.comp. phvsiol. convict (Cichlasoma Psvchol. 96, 955-966. sensitization and redirection of Peeke, H.V.S., 1983. Habituation, aggression and feeding behaviour in the three-spined stickleback (Gasterosteus aculeatus L.) J. camp. Psvcholosy; 9l, 43-51. Peters, H., 1937. Experimentelle Untersuchungen iiber die Brutplege von Haplochromis multicolor, einem maulbriitende Knochenfish. z_ Tierpsvchol., 1 201-218. Reinboth, R., Untersuchungen zur Maulbrutplege von 1956. 2001. Jb. (Phvsiol), 66, Haplochromis multicolor (Hilgendorf). 217-272. Rhoad, K.D., Kalat,J.W. and Klopfer, P.H., 1975. Aggression and artificial avoidance by splendes towards natural and Betta stimuli. Anim. Learn. and Behav., 3, 271-276. Rowland, W.J., 1975. System interaction of dummy-elicited behaviour bimaculatus in the jewel cichlid Hemichromis Gill. Behaviour, 53, 171-182. Thresher, R.E., 1979. The role of individual recognition in the territorial behaviour of the threespot damsel fish Eupomacentrus planifrons. Mar. Behav. Phvsiol., 6, 83-93. Vodegel, N., 1978. A causal analysis of the behaviour of Pseudotropheus (Pisces, Cichlidae). Doct. zebra (Boulenger) University of aiss. Zoological Laboratory, Groningen, Netherlands. Welcomme, R.L., 1969. The biology aca ecology of the fishes of a small tropical stream. J. 2001. Lona., 158, 485-529. Wickler, W., 1963. Zur klassification der Cichlidae, am Beispiel der Gattungen Tropheus, Petrochromis, Haplochromis una Hemihaplochromis n. gen. Senckenberq (Pisces, Perciformes). -biol. -44 I 83-96. Wilson, E.O., 1975. The synthesis. The Sociobiolosv. new Belknap press of Harward University Press, Cambridge.
Processes,
13 (1986) 353-366
353
Elsevier THE
AGGRESSIVE
MULTICOLOR
BEHAVIOUR
(PISCES,
TERRITORIAL
BETWEEN
WITH
CICHLIDAE)
PSEUDOCRENILABRUS
UNLIMITED
OR
ONLY
VISUAL
CONTACT
Anders
Fern6
Department
of
1839, N-5011
(Accepted
15
Fisheries Bergen,
July
University
Biology,
of Bergen,
P.O. Box
Norway
1986)
ABSTRACT behaviour Fern6, A., 1987. The aggressive between Pseudocrenilabrus multicolor (Pisces, Cichlidae) with only visual contact. Behav. Processes, 13: 353-366.
territorial unlimited or
between development 0 f aggression territorial fish has experiments using visual been studied in laboratory only contact. In this comparisons of the aggressive behaviour study, were made between conditions of unlimited and only visual contact. Fighting cichlids in direct contact between two territorial decreased over time in a way similar to the aggressive activity between subjects permitted only visual contact. However, there were also important differences between the two conditions of contact. Low-intensity aggression with a distance between the subjects but was generally not occurred frequently in free interaction observed visual contact. presence of under conditions of The non-territorial fish increased the aggression between subjects with visual contact had no influence on fighting and but conversely decreased low-intensity between interacting aggression freely subjects. In addition, the low level of aggression after some time of unlimited contact was not transferred to a subsequent period of visual contact. indicate that the decrease of The results aggression visual contact is caused under conditions of by habituation, whereas in a interaction a balance is also free developed between attack and escape tendencies. The
mainly
Key words:
aggression,
habituation,
territory,
cichlid
fish
INTRODUCTION
The the
aggressive
expenditure
behaviour of an animal's
it to the risk of physical territorial
0376-6357/86/$03.50
animal
to
0 1986 Elsevier
in
defence
of a territory
time and energy
injury.
It is therefore
behave
in
Science
Publishers
such
involves
as well as exposing important
for
a way as to maximize
B.V. (Biomedical
Division)
a the
354
benefits
of the aggressive
behaviour
1964).
Especially
crucial
towards
territorial
conspecifics.
neighbours enemy
is generally
effect
In
(Wilson,
field
aggression
(Lowe-McConnel,
some
In
on
frequent (Fernald
and Hirata,
sites
(McKaye,
1977),
found
behind
waning and
been
has
observed result
between of
Peeke,
fish
has,
were
contact
sexual
breeding
fish has been where
fish
tube generally
wanes
(e.g.
Peeke
et.
and Peeke,
and Veno, to a
it has
another
cichlids
1972;
Peeke
et
1979),
al.,
1970) and
the
1976). The response
process
of
habituation
that the low level of aggression territorial
reinforcement
aggression however,
fish
and Molen,
found
between
rarely
establishment
is
often also
1969; Peeke
a and
physical
1980).
than one with
contact unlimited
may
interacting
investigated.
Siamese
conditions
between
freely
been
in the
(Cain and Baenninger,
without
situation
In the
fighting
of unlimited It was proposed be associated
contact
fish,
and only that
with
and consequently
a
less more
lead to habituation.
There
is
one
conditions
convict
the decrease contact.
laboratory
of unlimited
(Peeke and Peeke,
pairs
of
of dominance
differences
of
or
towards
glass
(see e.g. Assem
development
context
both
e.g.
1978)
territorial
1966; Peeke
interacting
habituation
territorial
easily
a for
been attributed
freely
resource,
is
1973).
The
visual
in
al.,
suggested
1984).
fish
experiments,
displayed
1969; Peeke
(e.g. Peeke,
has generally
it
between
or
et
1977; Holzberg,
and Colgan,
evident.
aggression
partition
of
observed
territorial
in laboratory
(e.g. Baenninger,
stickleback
Loiselle,
for a limited
of aggression
Gallagher
anabantoids
dear
a low level
often
1983; Grant
has been demonstrated
1971;
al.,
is
between
is often
the
glass This
to territorial
the so called
1977; Kodrick-Brown,
in more detail that
a
rapidly.
1968;
and competition
development
(Brown,
of aggression
in fish,
neighbours
aggression
mates
been
territoriality
1979; McKaye,
more
The
to strangers,
Albrecht,
et. al.,
investigated
the costs
regulation
The responsiveness
territorial
species
the
1975).
1956;
1978; Colgan,
and minimize
be
less than
studies
between
may
study
1982). The aggression
cichlids
with
unlimited
was less pronounced
Aggression
decreased
between
in stable
on
and conditions
the
freely but
of only
between
contact
than under
periods
territorial
fish visual
territorial
decreased conditions interacting
increased
with
over of
under contact pairs
time but visual
territorial transitions
355
bet.ween
stages
explained
as an interaction 1970,
There
in
fish. Aggression
of individuals
There
may,
as
is thus
and also
between
presence
or absence
natural
conditions.
on the aggressive was also
aggression situation
MATERIAL
found with
reducing
Groves
and
to
been
1985). between
of habituation
unlimited
visual
and
compare
the development
contact
and to examine
visual
of
information
influence
of a period a
subsequent
contact
was derived
fish which
in order
unlimited
also
species.
experiments
investigated
the
Recognition
may occur
The importance
and visual
in
has
1979; Myrberg,
of
territorial
changing
1979).
aggression
conditions
behaviour
between
that habituation fish.
different
from
during
and
that habituation
after
et. al.,
(e.g. Thresher,
of non-territorial
visual
e.g.
indicating
(Colgan
Additional
The
(see
increased
was undertaken
situation.
were
(habituation)
of aggression
evidence
of conclusions
free contact
contact
males
between
study
studies
sunfishes
in both unlimited
aggression transfer
field
territorial
differ
The present
processes
results
The
decremental
1983).
factor
some
interacting
however,
contact
a
for in damselfish
freely
between
regulation
the
between
of individual
appearance
claimed
some
also
cycle.
reproductive
and Peeke,
are
involved
is
the
(sensitization)
incremental Thompson,
in
may
vary
of unlimited period
of
to
from
was
transferred
a the
under contact visual
to see if the attenuation
contact
of the
to
of a
contact.
AND METHODS
cichlid Pseudocrenilabrus The small, mouth-brooding fish multicolor (Schoeller, formerly Haplochromis multicolor, Hilgendorf chosen for or Hemihaplochromis multicolor, see Wickler, 1963) was can be established in this study as several territories growth of middle-sized aquaria. The distribution, migration and this East African cichlid have been investigated in the field conducted (Welcomme, 1969), but only laboratory studies have been on the 1937; reproductive and territorial behaviour (Peters, Reinboth, establish 1956; Kuenzer, 1975; Mrowka, 1982). The males territories and are then generally brightly coloured with a bluish body and black eye-bars whereas females are more cryptically coloured. There is no pair-bondi_ng and after spawning the female leaves the male's territory with the fertilized eggs in her mouth. The fish in this study came from a population of P.multicolor originating from a local dealer and held The at the laboratory. territorial varied in length 5.7 cm males between 4.0 and (including the tail fin). The two subjects in each trial had a mean length difference of than 2 mm and were isolated from each less other at least three weeks prior to contact in the trial. Aquaria of 200 1 (100x50~40 cm) were used. The floor of the tanks was covered by sand and devoid of plants. A hide made by flower-pots and stones was placed at where the each end wall
356
subjects centered their territories. The aquaria were illuminated from above by three 15 W bulbs with a photoperiod from 11 a.m. to 11 p.m. The temperature was held at 25 '1.0" C. The fish were fed boiled liver or frozen fish flesh at the beginning and end of the light period. opaque partitions. Each aquarium was bisected with glass and One subject, along with four or five smaller conspecific males and aquarium and allowed to females, was placed in each half of the establish a territory. After five days the opaque partition was removed and, in Exp. 2, the glass partition was also removed. were made with only visual contact In Exp. 1, six trials fish and the opaque between the test subjects. Non-territorial time t partition were removed five min before (around 1 p.m.). Observations were made for one hr at time t and for 30 min at time 30 min at tf3 hrs during the first day of visual contact and for time t the following six days. In two trials, non-territorial fish placed in each half 0 f the (two males and two females) were aquarium after the observation on Day 7, and observations were made for 30 min at time t the following five days. Two of the trials were In Exp. 2, nine trials were conducted. one of the subjects lost his not included in the material as the trial was terminated. territory to the other subject before The subjects were first studied for a period with unlimited contact in the presence of non-territorial fish. (In trials l-3 this period lasted 14 days but as no change over time was found after the first five days in trials few days of contact., this period was set to 4-7, and only data from Days I-5 will be presented for all trials). time t and On Day 1, the partitions were removed five min before subjects were observed for one hr at time t and for 30 min at the were made for 30 min at time t + 3 hrs. On Days 2-5, observations time t. fish was then included in A period without non-territorial Exp. 2. The day after the period with non-territorial fish present, observations these fish were taken away five min before time t and were made for 30 min at time t for five days. periods without nonwith and The sequence of the territorial fish was thus not the same in Exp. 1 and Exp. 2. The reason for inital absence of non-territorial fish in Exp. 1 the usually include such was that visual contact experiments do not variable situation. In Exp. 2, fish because of the resulting absent,one subject pretests showed that if non-territorials were partition was removed. always lost his territory when the unlimited contact in Exp. 2, The day after the periods with the glass partition was reintroduced in the middle of the aquarium observed for seven days five min before time t and the subjects visual contact with the same procedure as in under conditions of Exp. 1. by one observer, Observations were made from behind a screen spoken into a tape recorder and later transcribed. The following behaviour patterns were recorded: Both unlimited
and only
visual stroke
contact
Tail-beat
A strong
with
the tail.
Low-intensity
subject or both subjects made a aggression One towards (approach), away distinct movement and away (approach--retreat, (retreat), or towards frontal in relati.on to the other subject. display) (The frontal displays were few and of short duration and were therefore included in low-intensity aggression). In free interaction, both subjects
351
generally at a distance from the territorial were border. Under conditions of visual contact, low-intensity aggression occurred when the was territorial border of one or both subjects not in contact with the glass partition. Sexual
behaviour Quivering often quivering motion.
Fluttering Specific
A swimming unlimited
to
movement
followed
along
by
the walls
swimming
with
of the aquarium
contact
Attack
A rapid
Fight
Various combinations of circling, tail-beating, attacks. lateral threat, mouth-fighting and The types 0 f behaviour in a fight were not recorded. A fight consisted often of long series of attacks and more than one reciprocated attack was defined as a fight.
Specific
to
only
thrust
visual
towards
another
fish.
contact
towards the glass partition A subject swam swimming up and down the glass partition oriented towards the other male and often bit at the glass panel.
Aggressive
these behaviour patterns by The frequency of performance of the subjects towards each other was recorded as well as the time Attacks and spent fighting, aggressive swimming and fluttering. sexual behaviour patterns towards non-territorial fish were also recorded. When the subjects did not exhibit any of the behaviour patterns described either swimming relatively above, they were were passively, digging or staying within the hide. The subjects brightly coloured during all the defined behaviour patterns except for fluttering. The size of a subject's territory was recorded to the nearest five cm in each observation as the distance from the end wall of the subjects during aquarium to the position bet.ween the low-intensity aggression (assuming that the territorial border ran parallel with the end walls). RESULTS Only
visual
The
contact
aggressive
swimming
displayed
by both
swimming
decreased
Trend
previous
Lehman,
of
Biting
aggressive
swimming,
downward
contact
and
trend
no bites
Experiments
the glass
occurred
the same
in Exp.l,see
Fig-l).
observed
in
1). In Exp.
swimming
tail-beating
9
out
of
aggressive
2,
with
the subjects, and
a
(p
during
seven days the
13
a
there
subsequent
frequently
tendencies After
was usually
1,
to a low on Day 7 (p
between
and showed
were
1 and 2.
of aggressive
Exp. 1
on Day
contact
partition
In
1975, see Fig.
unlimited
increase
decrease.
the
towards
simultaneously.
from a high mean
test,
period
was an initial
subjects
the for
of visual trials
of
358
15
AGGRESSIVE SWIMMING
* 1\
EXP.l EXP. 2 o----o
,y?
0' I
BITE
', \
-7
FLUTTERING
LOW-INTENSITY
1
2
3
4
5
6
TAIL-BEAT
7 DAY
frequency of different behaviour duration or mean Fig.1 The made during observation patterns per territorial male and 30 min were periods contact. (Two observation visual seven days of In Exp. 2, the males had a previous period of conducted on Day 1). Significant differences between observations contact. unlimited fluttering in both and were found for aggressive swimming, bite in Exp. 1 and low-intensity agggression in Exp. 1 and 2, tail-beat denote significant Asterisks
359
Experiments
When
and tail-beating
p
the
Exp.
and
biting
swimming
tail-beating
when
in Exp.
2 had more on Day
difference
3 than
Exp.
maximum
were
1
or
in Exp.
(Day
2
was no
biting
activity
Exp.
U
aggressive
There
swimming,
and
1)
Day
2 (Mann-Whitney
1 (p < 0.05).
with
Day
swimming
on Day 2 and more
aggressive
the observations period
period
1 than Exp.
biting
in either
observation
the aggressive
observation
first
frequent
< 0.01).
significant
(first
in more
significantly test,
1 and 2 are compared,
2)
1
were
compared. The decrease increase
in
in
time
difference
was
significant
upward
0.001, In
test). 1,
with
either
only the
the glass
first
aggression and
aggression
with
the glass
one
subject
subject
after
infrequently
the usual
trial.
No
subjects
for
the
The addition
attack
rate
and sexual
marked
increase
(mean
aggressive
observation marked
change
increase
aggression several fish
over
in aggression
of the variable
time).
the
towards
30
Fluttering
these
situation
been
of
the other
added
observed
fluttering only
days
the
a
never
in a
subjects
with
occurred.
neighbours
(see Discussion).
was high
of additional
observation
almost
border
fish but also
between
the territorial
test)
in contact
fish were
in not
min
in
1 and 2.
then
five
1 and
than
from the glass
whereas
days
activity
per
between
had
resulted
during
male
Exp.
towards
in the aggressive
3.6 min per
behaviour
contact
Trend
territorial
cm distance
2,
Low-intensity
territory
in both
activity
swimming
0.01).
1, non-territorial
in Exp.
of other
frequent
claimed
Sexual
observed
trials
common.
result
the trial).
in
1
and
7)
of Day
0.001;
the
in
1). In Exp.
territory
more
a
of Day
Day
2 (p <
IO-15
at
until ( Fig.
<
but
territories
observation
(p
(in one trial
two
between
1
in Exp.
generally
however,
throughout
was
In
partition
observed claimed
Exp.
significant
observation
occur
the first
time
subjects
was,
by the first
an
1 and 2 (p <
defended
significantly
of
over
from Day 2 both
in
No
by
observations Experiments
generally
seldom
were
observation
in both
subject
partition
decreased
partition
was
1).
(Fig.
this did not
one or neither glass
found
partition
where
aggression
low-intensity the
was
accompanied
consecutive
subjects
in one trial
low-intensity
with
the
was
fluttering
between
trend
Trend
(except
spent
found
Exp.
contact
swimming
aggressive
no The
may be a
360
ATTACK OTHER
'*i
t t2
50
/
f-%h
I I
SEXUAL
E IL
i
BEHAVIOUR
, I
10
IL*
LOW-INTENSITY
*m g* t
I
1
1
OTHER
I
I
I
2
3
4
FISH
PRESENT
I
I
I
12 OTHER
I
1
I
3
4
5 DAY
FISH
ABSENT
or frequency of different behaviour duration mean The Fig.2 during observation 30 min and male territorial patterns per The initial five days were in the presence of contact. unlimited these nondays five subsequent while in the conspecifics, were periods removed. (Two observation were fish territorial Significant fish). with other Day 1 in the period conducted on lowfight, for found were observations between differences other intensity aggression and sexual behaviour in the period with fish (Friedman's test, p <0.05). Asterisks as in Fig. 1.
361
Unlimited
When
contact
the
partitions
were
removed
first
swam randomly
subjects first
time
Trend
of Day
was
hand
period
the aquarium 2, all fish at
fights
between
a few minutes.
the
During
the
up to 5.3 min occurred.
the
the first
or
subsequent
The
observations
aggression
and second
(p <
increased
observation
signed-ranks
other
period
or
subjects
the
these
upward
The territorial than
the
except
often
not
rate
(Wilcoxon days
2-5
however,
2).
for their
near
occurrence
the
and sexual
activity
in
territorial
activity
(p < 0.05,
with
the middle
from 25 to 75 cm.
and
seldom
changed
five cm from one day to another.
DISCUSSION
The
development
territoral be compared multicolor activity visual
of
fish coming in this males
contact.
study.
decreased
in contact
with
There
between
the
occurred
frequently
the
aggressive
into unlimited Fighting over
time,
the glass
were,
two conditions during
also
of contact. free
freely
similar
partition
however,
behaviour
and only visual between
interaction
to
a
Fig. 2).
around
in size
stable
by the
observation
located
ranged
was relatively
(p <
did
during
low in the first
was generally
time
fish
observed.
observed
for sexual
no but
over
increased,
see Fig.
infrequently
but territories
border
mean
aggression
The aggressive
border
aquarium,
on
or attack
(p < 0.05,
fish was
trend
decrease
time
observed,
fish were
The territorial the
removed
surface.
towards
significant
test
was also
Non-territorial border
the fighting
was seldom
Fluttering
slight
fish yielded observations
of non-territorial
Low-intensity
fish were
Tail-beating fights.
non-territorial consecutive
a
absence
matched-pairs
the two periods).
without
showed
time
presence
influence
more
in
of
in Exp.
actual
after
lasting
between
significantly
of
halves
2). Low-intensity
between
difference
fighting
O.O5).The
when
and
until
reduced
see Fig.
following
significant
in
fro
and
the
of a trial
1.
The
the
to
long fights
test,
other
the
beginning
not observed
were
fighting
separating
the
observation,
0.01, on
at
between
contact
can
interacting the
between
subjects
important Low-intensity and showed
p.
aggressive with
differences aggression a
certain
362
increase
initially
separated
by a glass
fish
also
Response a
waning
(Peeke with
behind
a
glass
indicated
by
only
A
cichlid
and
with
differently
in the
subjects
with
not involved
The changes
over
explained
if
development
an
in
and
territorial fighting
and
attack-escape
assumes
Vodegel, in another
been
there
activated
lead to
retreated
retreated
presented
therefore
in
be
better
addition,
attack
and
by
border.
stimulation to the
decrease
aggression. e.g.
in or
and the opponent
the escape
too close
observed
low-intensity indicated
the
when
is, the
the
and
opponent
decreasing
however,
or coming
and
with
and as
P.multicolor.
by the aversive
crossing
of is
in the
to the territorial
approached
increased
in the present
interacting
between
approached
subjects
(Ferns,
by each other
can,
aggression
increased
fish
waning
involved
freely
low-intensity
removal
from other
stimulated
of
The
as that
1975;
incidents
not influence
pattern
found
in relation
could
increase
(see also Ferno,
may
study,
(Rowland,
that
when
equilibrium
1977)
this
readiness
in the reponse
equilibrium
This
in
a behaviour
isolation
have
may become
border.
1975; Bols,
in
fish
system
between
attacks
response
stimulus
rapid.
the subjects
tendencies
fights
a
to
attack
in this situation
one
of
tendencies
time
from
has been
attributed
An avoidance
of aggression
were visually
also
may
stimulus
study,
generally
of fluttering,
long time
aggression
is
this
(Rhoad et al.,
was relatively
of
in
1973).
decreased
after
Habituation intensity
were
were
non-territorial
between
feed-back
a withdrawal
as the subjects
the decrease
Peeke,
inappropiate
general
species
and
in the decrease
1978) is probably
and
the subjects
of
aggression-eliciting found
in fish
the increase
may correspond
an as
partition
be involved
1978).
towards
demonstrated
connection
Escape
when
behaviour
to an increase
partition,
habituation
of
the aggressive
leading
glass
repeatedly
study
observed
The presence
contact.
behind
also
partition.
influenced
two situations, visual
but was seldom
the
which
An frequent
one one
where
of
subject subject
was oriented
laterally
1987). of non-territorial
the fighting
low-intensity visual other
was,
in a different
added.
on
comparable,
this
the other
did
the subjects between
Aggression
Although
sequence
interaction
rate between
aggression.
contact
fish were
not strictly
fish in a free
time or attack
hand,
the two
markedly
conditions
in the two experiments
and
observation
the
stresses
363
difference
between
of visual
the two conditions other
contact,
fish
aggressive
response
habituation
to take place
be
a
balance
independent
between
by
subjects
the
devoted
again.
attack
aggression
in
significantly trials
following
different
way.
intensity
wall
glass
in the aquarium
explanation
is
restricted
received
aggression
in
contact
explanation
transfer
to
defended in
which end
the of
territories territory The period
is
border the
were
influence
unequal,
of a period
study.
made
when
However, the
contact in three
glass
first
of visual
a
after
trials
of the aquarium whereas
with
contact
removed
increase.
in
the
the glass
2
could
of the subjects
partition
with
time,
result,
observation
subject
while
in the
fights
should
contact,
in Exp.
was
and
that
partition over
As
on
was not systematically
partition
of
glass
tendency
neither
the glass
the
the
glass
partition
in contact
a
contact,
contact
decreased
in the middle
trials
with
of visual
unlimited
only
a territory
of unlimited
this
with
increase
from direct
partition.
the
of
if one assumes
from the
which
with
was located
period
defended
by in
a
increased.
from the attacks
contact, in contact
the
in contact
the glass
with
in
of the reduced
the placing
of the escape
glass
the
supported
visual
territories
the
stimulation
through
of the
before
aggression
strength
low not
delay
arose
a distance
of
level
a
decreased
by
observation
with
level
This
partition
of aggression
fish. was not
(dishabituation).
interpreted
Low-intensity
as aversive
be
This
aggression
to
and then
which
the
was initially
interaction,
the glass
first
The
lead to an immediate
better
level
the
contact.
contradicted
due to a decreasing
subjects not
the
occurred.
frequently probably
be
these
contact
was the cause
changed
equilibrium,
during
low-intensity
with
could
attack-escape
still
should
therefore
in contact
partition
free
situation
the
result
The
contact
actually
because
in the beginning
level
alone
during
with
observations
the
habituation
when
aggression
from
was
perhaps
partition
for
could
in the subjects
unlimited
of visual
the glass
by unlimited
If
of aggression
aggression
the
with
in
the
fish,
to interacting
period
of the
there
aggression
during
contact
not preceded
similar
time
conditions
may be too variable
tendencies
other
of aggression
to a subsequent
increased
of
considerable
transferred
but
and escape
presence
Under
a dishabituation
In free interaction,
Low-intensity
fish.
attenuation
The
of contact. produce
and the new situation
of other
decreased
could
a
largest
partition. subsequent
investigated
observations after
at
when
the
in were
trial.
364
Fighting
was then lost
subjects observation
that
transferable Siamese
This
territorial
It seems,
however, between
balance
experiments
with
response
between
attack
patterns the
most
attack
aggressive
response
unlimited
experiments
and
using
be transferred be presented
A similar
interaction
visual
visual
contact
to free interaction. in another
paper
aggression
with
situation.
(Vodegel,
a In
1978),
strength
decreased
of certain
behaviour
probably
with
determines
a different The
therefore
Further
(Ferns,
did
by an interaction
contact.
should
of
in this zebra
whose
in El. multicolor
only
not
in the
1980).
interacts
be explained
tendencies
is
found
in free interaction.
time and by the performance
(e.g. biting).
the the
stimulation
habituation
tendencies
could
of
supports
(Lobb and McLain,
et. al.,
Pseudotropheus
models
escape
and
combats
that habituation
and escape
the cichlid
over
physical
can not take place
likely
been
to visual
that
one
This
has also
habituation
suggest
trials day.
in one situation
This
1976; Meliska,
fish
towards
spontaneously
between
not
all one
of aggression
in actual
and Meliska, does
within
situation.
fish, where
aggression
paper
between
the
a decrease
fighting
1976; Meliska
seen and in
territory
to another
reduce
not
frequently
his
outcome
results
from
not uncritically
evidence
for this will
19871.
ACKNOWLEDGEMENT
I
am
Radesater Karin
indebted for
Pittman
to
Dr.
constructive for valuable
Nance
Vodegel
criticism. remarks
and
professor
I also wish
Tommy
to thank
Miss
on the manuscript.
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