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Aggressive and territorial
behaviour in captive
Zebra Finches
S.M. Evans
Published online: 22 Jun 2009.
To cite this article: S.M. Evans (1970) Aggressive and territorial behaviour in
captive Zebra Finches, Bird Study, 17:1, 28-35
To link to this article: http://dx.doi.org/10.1080/00063657009476252
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Aggressive and territorial behaviour in captive
Zebra Finches
by S. M. Evans
THE ZEBRA FINCH Taenopygia guttata is an Australian grassfinch
which has been kept and bred by aviculturists in many different
countries. It is a social bird which normally lives in flocks of about
50-100 individuals (Immelmann 1965). In captivity it also behaves
socially and, like other social estrildine finches (Crook 1961), the
behaviour of a flock consists of alternating cycles of activity and
rest. During their active periods, the birds fly about, feed or sit
in an alert and attentive manner on the perches; during the periods
of inactivity, they preen, sit drowsily on the perches or even go to
sleep. Their behaviour is, in fact, even more precisely synchronised
than this because, at any one time, all of the birds tend to be
engaged in exactly the same activity : they tend to feed together,
preen at the same time and so on (Evans 1968).
Aggressive encounters frequently occur when several birds are
housed together in the same cage. An individual that is about to
attack another usually adopts a horizontal posture on the perch
with its feathers sleeked against its body and beak pointing towards
its rival (Morris 1954). Subordinate birds normally react by fleeing,
and sometimes supplanting attacks occur in which the aggressor
pursues its rival from one perch to another. Occasionally the
subordinate bird does not fly away but adopts a submissive posture.
Usually the body of the submissive individual is held vertically on
the perch (Morris 1954) but a fluffed posture, in which the bird
squats and fluffs out its feathers, and a posture which resembles
the food-begging response in fledglings, have also been described.
When two birds are equally matched they may beak-fence, by
stabbing at one another with closed beaks. These contests are
rarely prolonged and never seem to result in physical damage to
either of the combatants.
In flocks of some birds, such as Canaries (Shoemaker 1939;
Tsuneki 1960) and Bengalee Finches (Masatomi 1958), aggression
leads to complex dominancesubordinance relationships between
individuals and sometimes to the establishment of peck-orders.
It is also important in territorial defence, which usually occurs
when birds defend the area around their nests. In the present
investigation aggressive behaviour has been studied in com-
munities of captive Zebra Finches, with particular reference to
the establishment of peck-orders and territorial behaviour.
METHOD AND MATERIALS
The Zebra Finches used in this study were obtained from the
Toddington Bird Farm, Bedfordshire. Individuals were ringed
with and identified by coloured plastic rings. The following
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BEHAVIOUR IN CAPTIVE ZEBRA FINCHES
abbreviations, based on these rings, are used: RM (red male);
yM (yellow male); PIM (purple male); GM (green male); PkM
(pink male); RF (red female); YF (yellow female); PIF (purple
female); BF (black female); GF (green female).
Four communities were observed during the investigation. Three
of these were kept in cages measuring 4 X 2 x 2 feet, of similar
design to those used by Morris (1954); these birds were not per-
mitted to breed. The fourth flock was kept in an aviary measuring
9 x 4 x 8 feet; wooden nest-boxes, with open fronts and dried grass
as nesting material, were provided so that these birds could breed.
One of the non-breeding flocks consisted of five cocks and four
hens : they were introduced into a cage on 24 June, and were
observed daily until 6 July, 1966. The other two flocks of non-
breeding birds consisted of individuals of the same sex. One was
made up of five males and the other of five females (Tables I and II):
both groups were introduced into cages on 20 September 1966, and
were observed daily from 3-7 October inclusive. They were kept
in the same room and could hear but not see one another. The
breeding flock consisting of four cocks and three hens (Table III),
was introduced into the aviary on 5 June 1967, and was observed
regularly until 20 August.
All the birds were fed on a mixture of canary and millet seeds
given to themin earthenware bowls on the floors of the cages and
aviary. Water was provided in similar containers. Cuttlefish bone
and some green food, such as groundsel and lettuce, was also
given to the birds from time to time. Hides were not constructed
for the purpose of observing the birds because they did not appear
to be disturbed by a human observer. Most recording sessions
lasted an hour or more.
RESULTS
(a) Non-breeding birds
Aggression encounters were frequent in all three flocks of non-
breeding birds. They were most common when the birds were
actively flying about the cages and occurred less often when the
birds were inactive and when they were feeding. Several individuals
would feed within the small area around the food-bowl without
interfering with one another. Such closeness was not usually
tolerated; each Zebra Finch has an individual territory of about
5-15 cm. around itself (the `individual distance') in which other
birds, except for mates and young ones, are not normally per-
mitted to settle. Invasion of the individual distance either leads
to aggression by one of the birds or, alternatively, adjustment of
the positions of one or both of them so that the distance between
them is increased. However, birds often attack others that are at
greater distances from them so that by no means all aggression is
involved in maintaining the individual distance. In fact, encounters
between birds which are initially some way apart are often more
intense than those resulting fromthe encroachment of the individual
distance, and frequently give rise to supplanting attacks by the
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BIRDSTUDY
aggressor. But even these contests rarely lead to actual fighting and,
indeed, physical damage resulting from it has not been observed.
Most encounters were settled more by ferocity than actual fighting
ability.
Individuals in all three of the flocks were involved in at least
some encounters with other birds. However, some of them were
involved far more often than others. In the flock of hens, for
example, BF was involved in 453 (85 per cent) of the 533 observed
encounters and, in the flock of males, YM took part in 315 (57
per cent) of 553 contests. Conversely some birds took part in
relatively few aggressive encounters : RF was involved in only 64
(12 per cent) and P1M in 107 (19.3 per cent) in their respective
flocks.
The birds in the single-sex flocks can be ranked in orders of
dominance (peck-orders) based on the results of their encounters
with one another (Table I). In the mixed flock, however, there is
insufficient data for a reliable estimate of peck-order.
TABLE I-RECORDOF AGGRESSIVE ENCOUNTERS IN THE FLOCK OF FIVE MALES ANDTHE
FLOCK OF FIVE FEMALES
Encounters with other birds
Male Flock
Position
in peck GM YM RM Pk/v1 PIM
order Individual wins loses wins loses wins loses wins loses wins loses
5 GM 2645 1349 1222 512
4526 3 YM 4497 4822 1221
9744 1 RM 4913 599 2311
959 4 PIcM 2212 2248 1013
1310 2 PIM 125 2112 1123
Female Flock
BF GF RF P1F YF
wins loses wins loses wins loses wins loses wins loses
1 = BF 511 311 118 214 316
151 4 GF 81 610 117
18 5 RF 131 211 19
112 1= P1F 214 118 106 68
86 1= YF 631 171 91
In the flock of males and, as far as can be seen, the mixed
flock, the orders are linear when records for the whole period are
compared. That is to say, each bird tends to dominate all birds
below himin the order but is subordinate to all those above. In the
female flock there is a triangular relationship between BF, P1F and
YF (BF dominated P1F but was subordinate to YF, while P 1F
dominated YF) but otherwise the order is linear. However, there
were frequent changes in the orders from one recording session
to the next. In most cases the dominance relationship between a
particular pair of birds was clear-cut for any particular recording
session, although it often changed in different sessions. For example,
YM dominated RM during two of the five recording sessions for
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BEHAVIOUR OF CAP TIVE ZEBRA FINCHES
the male flock but finished below himin the overall order (Table 1).
The actual record of wins and losses was as follows (YM first) :
15-6; 17-71; 4-11; 8-0; 0-9. Some times there was no clear
dominance of one bird over another and occasionally there was a
sudden change in the dominance relationship between two birds
during the course of a recording session. In the male flock, for
example, GM defeated YM in four successive encounters on one
occasion but was then defeated in the next seven encounters with
the same bird.
TABLE II-CHANGES IN P ECK ORDER DURING DAILY RECORDING SESSIONS FOR THE FLOCK
OF FIVE MALES AND THE FLOCK OF FIVE FEMALES
Position in peck-order
Male f lock
Individual
3.10.66 4.10.66 5.10.66 6.10.66 7.10.66
GM
5 5 3 5 4=
YM
3 2 2 2 4=
RM
1 1 1 3 2
P1cM
4 4 4= 4 3
PIM
2 3 4= 1 1
Female f lock
BF 1
1 2 3 2
GF
4 2 4= 4= 4
RF 5 4= 4= 4= 5
P1F 3 3 1 1= 1
YF 2 4= 3 1= 3
Birds that were near the top of the order were sometimes par-
ticularly aggressive towards one another. The most obvious
example is in the female flock which the majority (62.3 per cent) of
encounters were between BF and P1F, which were ranked first equal.
In the mixed flock, the top two birds RM and YM were frequently
aggressive towards one another and, in the male flock, these two
birds were involved in 25.5 per cent of all the aggressive encounters,
although they were ranked 1 and 3 in this order.
Conversely, there was only occasional aggression between some
birds. In the mixed flock pair-bonds were formed between RM-YF,
GM-P1F, YM-GF, and a weak bond between PIM-RF, and there was
an average of only 1.5 aggressive encounters per pair between these
birds compared with an average of 3.2 for other birds. These pairs
could be recognised because they clumped together (i.e. sat in
bodily contact) and the males often preened (allopreened) their
mates; occasionally the females reciprocated by allopreening the
males. In the flocks of males and females, some pairs of birds
formed homosexual pair-bonds. These birds clumped together and
allopreened, like normal heterosexual pairs, and were infrequently
aggressive towards one another. In the flock of females the two
pairs, RF-GF and YF-P1F, which regularly clumped together, were
aggressive towards one another on only five and 14 occasions
respectively compared with an average of 64.3 encounters per pair
for the rest of the flock. In the male flock, PkM regularly clumped
with both GM and PIM and was involved in hostility with them
c
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BIRD STUDY
on only 34 and 23 occasions respectively, compared with an
average of 62.0 for the rest of the flock.
There is little doubt that sexual arousal has some effect on the
behaviour of males and if a hen is introduced into a cage con-
taining two males, there is a marked increase in the aggression
between them. In the mixed flock, the introduction of an un-
familiar female, BF, had an interesting effect on the behaviour of
the only unpaired male PkM. Before her introduction PkM was
subordinate to all of the other birds, winning only one in a total
of 30 encounters. He was persistently chased by both YM and RM.
Most attacks were made on himwhen he moved and, consequently,
he spent most of his time sitting quietly on a perch, usually near
RF and PIM, which were also low in the peck order. However,
when BF was introduced into the community he became more
active and more aggressive and subsequently won 11 out of 20
encounters with other birds. He even defeated RM on three
occasions and YM twice. BF was courted promiscuously by all
the males, except PIM, but she was attacked by the other females
and replaced PkM at the bottom of the peck order.
(b) Breeding Birds
Three pairs of birds, which had already formed pair bonds,
(GF-YM; PIF-GM; YF-RM) and one unpaired male (PIM) were
introduced into the aviary on 5 June, 1967. They started to nest
almost immediately and, within 24 hours, each pair had taken
over one of the nest-boxes and had started to construct a nest in
it. Eggs were laid by all three hens and incubated by the male
and female in turn. Those of GF-YM failed to hatch, but P1F-GM
reared three young and YF-RM reared four young. The first of
these broods left their nests on 18 and 14 June respectively. Two
of the offspring of P1F-GM were white, a domestic variety lacking
all feather pigmentation.
The birds behaved territorially when they were breeding. Each
pair defended the area around its nest against intruders and even
previously dominant birds were chased fromit. Aggression was par-
ticularly intense when the trespassers came near the nest; it was
less intense near the boundaries, where beak-fencing, a mild form
of aggression (Morris 1954), was sometimes observed.
Because each bird tends to win encounters inside its own terri-
tory but lose those outside it, the concept of peak-order becomes
meaningless. In fact, only 34 (16.8 per cent) out of a total of 202
observed aggressive encounters were not concerned with territorial
defence. Several of these involved PIM which did not behave terri-
torially (see below) and eight were on the floor of the aviary. Here,
there was no evidence of territorial behaviour and, in fact, little
aggression. Several birds often fed peacefully together and wandered
beneath their rivals' nests without eliciting hostile reactions.
PIM, the only unpaired bird in the community, was chased out
of the territories of all the other birds whenever he entered them.
Sometimes he would flee from one territory into another and was
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BEHAVIOUR OF CAPTIVE ZEBRA FINCHES
chased successively by different birds. However, he did dominate
other birds in 19 encounters, although these were either outside
the territories of his rivals or near to their boundaries.
TABLE III-RECORD OF AGGRESSIVE ENCOUNTERS BETWEEN BIRDS IN THE BREEDING FLOCK
OF 4 MALES AND 3 FEMALES
Encounters with other birds
GM
Individual wins loses
PIP
wins loses
YM
wins loses
GF
wins loses
RM
wins loses
YF
wins loses
Ph %
wins loses
GM 0 0 4 5 0 1 3 29 4 5 13 7
PiF00 2 1 1 0 2 4 9 1 28 1
1 2 YM5 4 0 0 5 10 4 3 15 5
0 0 GF1 0 0 1 0 4 0 0 2 0
4 0 RM29 3 4 2 10 5 0 0 20 1
0 0 YF5 4 1 9 3 4 0 0 8 5
5 8 PIM7 13 1 28 5 15 0 2 1 20
NOTE: GM-PIF, YM-GF and RM-YF were the breeding pairs.
Like non-breeding birds, there was a reduction in aggression
between the pairs that had formed pair-bonds. Indeed, no aggres-
sion at all was observed between mated birds during this investiga-
tion (Table III). Hens were involved in less aggression towards
rivals than their mates and were apparently more tolerant of
trespassers near the nest.
For about the first three weeks after leaving their nests, young
birds were involved in little aggression. They spent much of their
time clumping with one another or with their parents near the
nest. Occasionally they strayed into the territories of other breed-
ing birds and although they were usually attacked, aggression
against them was relatively mild. Within 3-4 weeks they had
moulted into adult plumage; at the same time, their beaks changed
colour from black to red, the beak colour of mature birds. Con-
comitantly there was a change in their behaviour. They became
more aggressive towards other individuals and conversely other
birds were more aggressive towards them. Previously they had
roosted in their parents' nests but now they roosted in extra
boxes provided for them. Some of them also formed pair-bonds
with other birds. One of the white hens paired with P1M; they
took over an uninhabited nest-box, built a nest in it and defended
the area around it against intruders. One egg was laid before the
experiment ended on 20 August. A second brother-sister pair also
made a crude attempt to build a nest in another of the nest-boxes.
DISCUSSION
Aggressive acts frequently occur among captive Zebra Finches
housed together, although they usually involve no more than the
adoption of threatening postures or one bird chasing another from
one part of the cage to another. Actual fighting is rare.
Individuals behave territorially when they are breeding and
defend the areas around their nests. Indeed, most of the aggression
that occurs in a breeding community is concerned with territorial
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BIRD STUDY
defence. This behaviour probably prevents other birds frompilfer-
ing nesting materials or even from stealing the nest site itself.
Furthermore it allows one member of the pair to carry out
activities, such as building the nest, incubating eggs and so on,
without interference from other birds. The establishment of terri-
tories usually leads to the dispersal of an animal community over
a larger area (Lorenz 1966). This effect can even be seen in captive
Zebra Finches. A flock of breeding birds tends to become dispersed
over the whole of the area available to it, whereas non-breeding
birds tend to congregate in part of their cage, even though they
maintain individual distances between one another. There is prob-
ably a similar dispersal of breeding birds in the wild. Zebra Finches
usually occur in flocks of about 50 to 100 birds when they are not
breeding but pairs nest in separate bushes as long as sufficient nest-
sites are available (Immelmann 1965). The survival value of this
dispersal is not clear. One possibility is that the spacing of nests
affords some degree of protection from predators. Alternatively,
it may prevent over-crowding of birds and thereby reduce com-
petition for food. However, if this is so, it is curious that captive
birds only occasionally fight over food and will even permit others
to feed in the area beneath their nests.
Although captive Zebra Finches only behave territorially when
they are breeding, there is still a good deal of aggression between
non-breeding birds. Some of it is involved in maintaining individual
distances but other aggressive encounters occur for no obvious
reason. They do not apparently confer any advantage, such as the
acquisition of food or a mate, to the winner. However, aggression
does lead to the establishment of peck-orders of dominance,
although in communities of Zebra Finches these are only semi-
permanent. According to Tordoff (1954), orders of dominance are
advantageous to a community as a whole because subordinate
individuals give way to dominant ones in competitive situations,
thereby reducing the amount of fighting that takes place. He
found that his flock of captive Crossbills Loxia curvirostr fed in
order of rank when there was competition for food and there was
evidence that highly ranked birds obtained the favoured roosting
sites. Similarly in flocks of Canaries Serinus canarius, dominant
birds usually feed before subordinate ones (Tsuneki 1960). This
explanation may also apply to the function of peck orders in
Zebra Finches. Detailed recordings were not made but on three
occasions when a dish containing bathing water was provided for
the flock of five males, they bathed one at a time and in an order
that bore a close resemblance to the peck order.
Captive Zebra Finches breed extremely readily and will attempt
to do so at any time of the year in Britain. Brood after
brood is reared by some pairs and there is a report of one that
reared nineteen successive broods (Weston 1930). Young also mature
rapidly and in the present investigation birds less than three months
old were attempting to breed. These are almost certainly adapta-
tions to the need for rapid reproduction in the wild. In parts of
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Australia breeding only occurs after rains, when seeding grasses
become abundant. But although the birds may not breed for months
or even years at a time in dry weather, breeding is prolific when
the opportunity to do so occurs (Immelmann 1965).
ACKNOWLEDGEMENTS
I should like to thank Professor W. H. Thorpe, F.R.S., who has supervised
this research, for his kind help and the Royal Society's Committee on
Scientific Research in Schools for financing it. My best thanks are also due
to Mr. P. G. Caryl for an extremely valuable discussion when this work first
started and, subsequently, for several suggestions which led to improvements
in this paper.
Several senior pupils at this school have spent many hours carefully record-
ing the behaviour of Zebra Finches and, although I amsure that they have
benefited fromthe experience, I should like to acknowledge their help.
Mr. G. R. Patterson and Mr. M. R. Thompson, in particular, deserve thanks.
SUMMARY
1.
Aggressive behaviour has been studied in captive communities of the
Zebra Finch Taenopygia guttata. Peck-orders of dominance are established
in non-breeding flocks but they are only semi-permanent and may change
fromone day to the next.
2. In both mixed flocks of cocks and hens and homosexual flocks, there is
reduced aggression between birds that pair with one another. Homosexual
pairs behave like normal heterosexual pairs in the sense that they clump
together and allopreen one another.
3. Breeding birds behave territorially and the majority of aggression is
in defence of the area around the nest. Young birds mature rapidly and may
attempt to breed when they are less than three months old.
REFERENCES
CROOK, I. H. 1961. The basis of flock organisation in birds. In Current Prob-
lems in Animal Behaviour. Eds. W. H. Thorpe and 0. L. Zangwill.
EVANS, S. M. 1968. The study of bird behaviour in schools (in preparation).
IMMELMANN, K. 1965. Australian Finches. Angus and Robertson.
LORENZ, K. 1966. The Nature of Aggression. London
MASATOMI, H. 1960. Attacking behaviour in homosexual groups of the Bengalee,
Uroloncha striata var. domestica Flower. J. Fac. Sci. Hokkaido Univ., VI,
Zool., 14:234-251.
MORRIS, D. j. 1954. The reproductive behaviour of the Zebra Finch (Poephila
guttata) with special reference to pseudo-female behaviour and displacement
activities. Behaviour, 6:271-322.
SHOEMAKER, H. 1939. Social hierarchy in flocks of canaries. Auk, 56:381-406.
TORDOFF, H. B. 1954. Social organisation and behaviour in a flock of captive
and non-breeding crossbills. Condor, 56:346.
TSUNEKI, K. 1960. Social organisation in flocks of canaries. Jap. J. Ecol.,
10:177-189.
WESTON, n. 1930. Foreign finches that are easy to breed. Avicult. Mag.,
8:272-273.
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