Bird Study

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The Significance of Some Behaviour Patterns of

Pigeons

Derek Goodwin

To cite this article: Derek Goodwin (1956) The Significance of Some Behaviour Patterns of
Pigeons, Bird Study, 3:1, 25-37, DOI: 10.1080/00063655609475836

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1956 IBERIAN PENINSULA AND MIGRATION 25

SCHNEIDER, w. 1 937. Beringungs-Ergebnisse an der mitteleurop-

áischen Schleiereule (Tyto alba guttata Brehm). Vogelzog, 8:
159-70.
scHUZ, E. & WEIGOLD, H. 1931. Atlas des Vogelzugs nach den
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Vogelforsch. 3.
sKOVGAARD, P. 1951. Ayes anilladas en Dinamarca y cobradas en la
peninsula ibérica. Bol. R. Soc. Esp. Hist. Nat., 494 1- 54.
SNOW, D. W., OWEN, D. F., & MOREAU, R. E. 1955. Land- and sea-
bird migration in north-west Spain, autumn 1954. Ibis. 97:
557 -7 1 .
SPAEPEN, J. 1953. De trek van de Boompieper Anthus trivialis (L.)
in Europa en Afrika. Gerfaut, 43 : 178-23o.
STRESEMANN, E. 1955. Die Wanderungen des Waldlaubsángers
(Phylloscopus sibilatrix). J. Orn. Lpz., 96: 1 53 -6 7.
TAIT, W. 1924. The Birds of Portugal. London.
TEN KATE, C. G. B. 1936. De purperreigers Ardea p. purpurea L., in
Ambt -Vollenhove. Org . Cl. Ned. Vogelk., 9: 68- 75.
THEMIDO, A. A. 1952. Ayes de Portugal. Coimbra.
THOMSON, A. L. 1 953. The migrations of the British warblers (Sylvi-
idae) as shown by the results of ringing. Brit. Birds, 46:441-50.
TINBERGEN, L. 1941. Over de trekwegen van Vinken (Fringilla
coelebs L.). Ardea, 30 :42-73.
VERHEVEN, R. 1954. Sur la migration de la population européenne
du Combattant. Vol. Juba Victor Van Straelen, 2:1013-24.
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Belgique. Gerfaut, 45 :5-25.

THE SIGNIFICANCE OF SOME BEHAVIOUR PATTERNS

OF PIGEONS
by
DEREK GOODWIN
Received i i September, 1 955
SUMMARY
1. Certain of the social behaviour-patterns of some pigeons are described
and their significance discussed.
2. Head shaking is shown on a physical level in response to mechanical
irritation of the head, but also as a response to stimuli that are (evidently)
psychologically displeasing.
3. Nodding is derived from nest-building movements. It is shown when the
bird intends to "hold its ground" and may be either hostile or appeasing in
context. The bill-opening of the Green Imperial Pigeon (Ducula aena) has a
similar meaning and derivation.
4. Displacement-preening occurs when the impulse to bill or copulate is
frustrated but no tendency to attack or to flee is present. It serves to indicate
sexual and peaceable intentions and perhaps also as an auto-erotic stimulus.
5. Caressing occurs only between mates or young birds that have a social
bond between them. It appears to be due to sublimated aggressive impulses.
On a physical level it serves to remove foreign bodies from the plumage of the
head and the facial skin.
 26 BEHAVIOUR OF PIGEONS 3(I)

6. Driving occurs as a response to the presence near the female (when she is
sexually receptive) of potential sexual rivals. Her mate tries to drive her away
from them. It does not occur if other males are not present. It functions to
prevent insemination of the female by males other than her own mate. Inter-
ference with copulating pairs is part of the same behaviour-complex as driving
and has a like causation and function.

INTRODUCTION
In this paper I propose to discuss and I hope to clarify a few of
the behaviour-patterns which pigeons show in various social con-
texts and which have been often neglected or misinterpreted. It is
based on observations of Rock, Feral and Domestic Pigeons
(Columba livia), Wood Pigeons (C. palumbus), Stock Doves (C.
oenas) and Turtle Doves (Streptopelia turtur) both in a wild state
and in captivity ; and on captive specimens of Speckled Pigeons
(Columba guinea), Red-eyed Doves (Streptopelia semitorquata)
and Barbary Doves (S. risoria). Many of the Barbary Doves have
been, allowed full liberty except during the winter. Briefer observa-
tions have been made on other species, chiefly on captives in the
London Zoo. Throughout the paper the term C. livia has been used
where the remarks apply equally to wild, feral and non-monstrous
domestic forms of that species. The line sketches are merely to
augment the written descriptions and have no pretension to art or
detailed accuracy.

HEAD-SHAKING
A sudden, quick, shaking or jerking of the head. The movement
appears chiefly lateral, but at times seems to involve a rotary move-
ment also, though this is not noticeable except when it is made
more slowly than usual.
This movement is shown immediately after drinking or as a re-
sponse to physical irritation of the head, especially the mouth.
Gently prodding or tickling the head of a pigeon (if it is sufficiently
tame not to be frightened by one's close proximity) with a grass
stem will at once elicit this reaction. The use of the movement under
these circumstances is clearly to remove drops of water or other
foreign objects adhering to the head or bill. It is very similar to,
and possibly homologous with, the head-shaking movements that
many water birds—grebes, gulls, guillemots, etc.—invariably show
after feeding, drinking or preening.
In many species this movement is also made under what at first
appear to be quite different circumstances. When a pigeon coos at
another the latter will head-shake if it is not aroused to overt
sexual, aggressive or fleeing behaviour. It is evidently the sound that
elicits this response, not the display which usually accompanies it.
Under the same conditions to those mentioned above, tame pigeons
will head-shake if their owner "coos" at them. I have see two
Speckled Pigeons (a male and female) head-shake when another,
completely hidden from their sight, gave the advertising coo.
Since the head-shaking of a pigeon that is displayed at by
another may be followed by, or interspersed with, movements of
avoidance or withdrawal, one is tempted to regard it as an appeas-
ing or submissive display. I have, however, seen nothing to sug-
I956 BEHAVIOUR OF PIGEONS 27

gest that it serves any such function. It seems rather that the
pigeon not in sexual or self-assertive mood finds the cooing dis-
pleasing ; that it is, in effect, the psychological equivalent of a
physical irritant and that the bird thus reacts to either stimulus with
the same movement. On what may be (but I think is not) a higher
plane of behaviour, we see the same phenomena in Man. Most of
the gestures of disgust, contempt or repudiation used by Homo
sapiens—of varying races—are essentially identical with those used
on the purely physical level to rid the mouth or body of irritating or
distasteful substances.
I think head-shaking is used in the above sense by all pigeons
and have seen it in the Speckled Pigeon, Wood Pigeon, White-
crowned Pigeon (Columba leucocephala), Barbary Dove, Red-eyed
Dove, Turtle Dove, Laughing Dove, Grayson's Dove (Zenaidura
graysoni), Wonga Pigeon (Leucosarcia picata) and Plumed Ground
Dove (Lophophaps ferruginea). C. livia uses it far less freely than
other species known to me. Indeed, whereas I have seen this ges-
ture hundreds of times from all my other captive pigeons, I have
seen it very seldom in my Domestic Pigeons. Perhaps this more
gregarious species seldom feels the emotional mood that elicits
head-shaking at the sound of another cooing.
The following incident seems worth giving, as it suggests that
head-shaking will be inhibited by any positive sexual feelings
towards the cooing bird. A pair of Wonga Pigeons in a Zoo had
been separated and placed in aviaries a few yards apart. The hen
was desperate to rejoin her mate. She alternately ran up and down
the wire trying to get to him or perched on a ledge and there uttered
her advertising coo. A cock Turtle Dove in the same aviary, who
was inhibited from cooing or displaying by an aggressive pair of
Red-eyed Doves, frequently showed a strong sexual response to
the Wonga Pigeon when she stood still calling on the ledge. He
would cautiously approach her, obviously torn between fear
(probably because of her much greater size) and desire to copulate
with her, and begin to make the intention-movements of mounting.
During the hour of so that I watched he made several such
attempts, but since, when he was near her, any mov ement of the
Wonga Pigeon caused him to flee a little from her, he never got
any further. The Wonga Pigeon showed no reaction at all towards
him. The Turtle Dove's tentative approaches alternated with
periods when he apparently lost interest and sat mopily on the far
end of the ledge. When thus sitting he shook his head at each
repeated fluting "coo" of the Wonga's call. During the periods
that he was approaching her, however, he gave no such reaction to
her cooing.
NODDING
A downward stroke of the head, at the lowest point of which the
bill is at an angle of about 85 to go degrees to the horizontal, is
given by C. livia under the following circumstances : (I) by both
members of a pair when at the nest-site or when coming together
again after a short parting, (2) by a female in response to sexual,
self-assertive, or aggressive behaviour from a male with whom she
28 BEHAVIOUR OF PIGEONS 3(I)

is eager to pair, (3) by a bird alone (normally with its mate nearby)
on the nest-site, (4) by a bird threatening (usually defensively)
another. In all these instances it is interspersed with various self-
assertive, aggressive or sexual behaviour patterns.
Thus the nodding is shown in what at first seem different situa-
tions. When given by a sexually aroused female in response to the
sometimes rather rough advances of a male, it gives the impres-
sion of being a submissive display. When given by a nest-calling
bird, that tilts its head to eye its mate whilst so-doing, it suggests
an eager beckoning. When given by a defending or threatening
bird its hostile character is at once obvious. If one tries to analyse
the precise difference between "friendly" and "hostile" nodding
the most one can say is that in the latter the movement tends to be
swifter and more forceful, but also more often incomplete. Yet few
pigeon-keepers, seeing a bird nod, are ever in doubt as to whether
the gesture is directed at mate or foe—an interesting, if not unique,
example of the superiority of man's innate perception to his
rational analytical faculties.
Nodding seems to be universal in pigeons and has been observed
in many species. I will here confine myself to a few in which I have
closely observed it on many occasions. In the Wood Pigeon, the
Stock Dove and the Speckled Pigeon the movement is nearly the
same as in C. livia. The sexually excited Wood Pigeon tends to
give short nods in which the head is only lowered a little. The
Speckled Pigeon, when on the nest-site, nods with the neck less
drawn in and at the lowest point of the movement the bill seems
usually to be at an angle of about 45 degrees from the horizontal.
In view, however, of the amount of variation shown even by the
same individual these apparent specific differences must not be over-
emphasised. In the Barbary Dove the movement is slight. Even
when on the nest this bird usually only moves its head forward and
downward through a small arc. In sexual and aggressive en-
counters away from the nest the movement is even less definite and
so interspersed with the "aiming" flight-intention movements as
to be difficult to separate from them. In the Red-eyed Dove the
nod, although similar to that of the Barbary, is more emphatic and
quite easily recognisable.
These species all use the nodding in the same situations as does
the Domestic Pigeon. In the Wood Pigeon and the Streptopelia
species it may sometimes seem to be the accompaniment of active
rather than defensive aggressiveness. The nodding of an attacking
bird is, however, shown in intervals between attacks, when it is,
for the moment, only "holding its ground". This in fact appears
to be the basic "meaning" of the movement. It always signifies
the bird's intention of holding its ground and refusing to be driven
from it. Probably the mood of the nodding bird is very different in
the different situations but it always embodies a determination to
remain in spite of opposition.
I was for some time unable to decide whether this movement
was a ritualised intention-movement of pecking, or of nest-building.
The frequent occurrence of actual pecking of the prospective mate
by courted Wood Pigeons and Stock Doves and the very obvious

1956 BEHAVIOUR OF PIGEONS 29

aggressiveness in the quick "spiteful" nods of C. livia and Speckled

Pigeons when angry suggested the former. On the other hand the
most intense form of the nodding movement in all these species
suggested the movement used when adding material to the nest. I
became convinced that the latter explanation was correct when I
first saw the nodding of the Bar-shouldered Dove (Geopelia
humeralis). In this species the nod culminates in the typical move-
ment of a pigeon fixing a twig into its nest. So far as I have been
able to observe from the two captive specimens at the London Zoo
this species nods in threat, when away from any possible nest-site.
even more freely than the others mentioned, although otherwise
nodding under like circumstances. It is indeed bizarre to see the
bird, with empty bill, continually lowering its head and making the
typical "shuddering" movement as if fixing an imaginary twig in
an imaginary nest. The male Wood Pigeon, when "inviting" the
female to join him on the nest-site, sometimes makes deep nods that
culminate in a slight "twig-fixing" movement.

Figure t.—"Friendly" nodding of male Wood Pigeon enticing female to

nest-site.

It seems probable that the nod has been derived from the nest-
building movement. It may originally have been used only at the
nest-site and in such form might be "translated" as the equivalent
of saying "Here I intend to nest", this "statement" being either
hostile or inviting according to circumstances. Its extension to
hostile and sexual encounters away from the nest-site is explicable
if during such the bird is always, to some extent, in the same mood
as one calling on or defending its nest-site. Since in pigeons fight-
ing and sexual behaviour are usually linked with possession of
territory, this seems highly likely. It could perhaps be suggested
that instead of having evolved from a completed building move-
ment the nodding is the original movement—of displacement or re-
directed pecking—shown when the impulse to attack is inhibited by
fear or sexual attraction and that the complete building movements
have been developed out of it by natural selection. This second
theory seems less likely, but by no means impossible.
The only member of the Treroninae that I have seen showing
similar behaviour is the Green Imperial Pigeon (Ducula aena).
This.species has a movement which is used in the same situations
as the nodding of Columba and is, I think, probably homologous
with it. It consists of a rapid opening and shutting of the mandibles.

30 BEHAVIOUR OF PIGEONS 3(I )

Usually only the terminal part of the bill seems to open but this may
well be an optical illusion. The head is held in its normal position
or slightly raised, but it may be lowered as in a nodding Columba.
This movement seems to be a ritualised intention-movement of seiz-
ing a twig, or taking one from the mate, for building purposes.
It thus derives from the same general behaviour-pattern--nest-
building—as does the nodding of other pigeons.

Figure 2.—Bill-opening of Green Imperial Pigeon in nest-site.

DISPLACEMENT-PREENING
The stereotyped movement in which a pigeon turns its head,
thrusts its bill between body and scapulars and quickly withdraws
it, is well known. It seems to be common to most species, at any
rate to those whose behaviour has been recorded. In C. livia (and
I think in other species of Columba and in the Streptopelia species)
it is given by the male when he is sexually aroused but unable to
copulate or commence the pre-copulatory billing either because the
female is not yet giving the appropriate response or because of his
own incapacity. That the same movement is shown in these appar-
ently different situations suggests that it is used whenever an im-
pulse to bill or copulate is being frustrated but no other positive
conflicting tendencies—such as to flee or to attack—are present.
It also corroborates Whitman's (Iglg) opinion that this act pro-
vides auto-erotic stimulation as well as serving as a signal to the
mate.
The female does not usually displacement-preen so much as the
male. This seems to be because she reacts to sexual unreadiness on
her mate's part by more intense soliciting of billing or coition, by
"switching over" to caressing him, or (if the male crouches and in-
vites her) by mounting him instead. She will, however, displace-
ment-preen freely if thwarted in her sexual desire and unable to find
outlet in the above-mentioned alternatives. Thus a female "paired"
to a human being will displacement-preen as much as, or more
than, a male normally does.
Some male C. livia (perhaps all, but I have seen it so seldom
that I think only some do it) also displacement-preen in the side of
the breast, as does the Black-headed Gull (Larus ridibundus). This
may alternate or be interspersed with the usual "behind the wing"
movement. More often, however, it seems to indicate a less intense
sexual feeling. I have not seen this movement in other species but
in view of its relative inconspicuousness and rarity in C. livia it is
quite likely that I have missed it in others.
1956 BEHAVIOUR OF PIGEONS 3I

The Red-eyed Dove shows a striking form of displacement-

preening. It lowers its head and neck till the bill is over the brood
patch, makes a rapid waggling movement of the head, and bill, and
resumes its normal stance. This movement is usually preceded and
immediately followed by the typical "behind the wing" displace-
ment-preening. At lower intensity it may merely touch the abdomen
lightly with its bill, or it may reach only to the side of the breast as
C. livia. There is a possibility that the "waggling" head movement

Figure 3.—Displacement-preening of Red-eyed Dove.

given at high intensity is actually a building or egg-turning move-

ment. The erection of the plumage in the area of the brood patch
suggests the latter, although these feathers are, in pigeons, as in
passerines, almost always erected (together with those of the rump)
in sexual excitement.
It is not always possible to draw an absolute line between the
displacement-preening behind the wing, and real preening in this
area. If a pigeon that is engaged in preening is very slightly
sexually aroused by the arrival or sexual behaviour of another, it
will preen behind its wing. It then uses normal preening movements
—not the stereotyped displacement form. Nevertheless its subse-
quent behaviour towards the other bird usually indicates that they
were sexual impulses—at low intensity—that caused it suddenly to
pay attention to this area.
The function of this displacement-preening is, as is well known,
to indicate the bird's sexual and peaceable intentions and to arouse
a similar mood in the partner.
CARESSING
The mutual caressing commonly indulged in by pigeons (at least
by all species of Columba and Streptopelia known to me) is a
behaviour-pattern that appears to have both psychological and
"utilitarian" significance. Before discussing these it will be neces-
sary to describe briefly what the caressing bird does and when it
does it.
The caressing consists of a gentle-looking nibbling movement
of the bill, which is thrust into, and moves about among, the
partner's feathers apparently in contact with the skin. Frequently
the bird will seize some small object in its bill tip and either swallow
it or (less often) cast it away. The rump, back, wings and breast
32 BEHAVIOUR OF PIGEONS 3(I)

of the partner may be treated very briefly in this manner, but in

general the caressing bird confines its attentions to the other's
head and nape. At least 99 per cent. of all caressing is done where
the bird cannot reach with its own bill. Approximately (but not ex-
actly) these are the areas above, and on the upper periphery of, the
display plumage on the neck, and the caressing bird often com-
mences by plunging its bill into the display plumage and then
working upwards. Although usually gentle, it may be rather rough
on occasions, and distinct though they are in their typical forms,
it is difficult, even on the observation level, to say where aggressive
pecking ends and caressing begins, or vice versa.
Mutual caressing by members of the pair occurs on the nest-
site, particularly by the female in the early stages when she joins
the male on his selected site. It may occur before or after copula-
tion, in such case usually initiated by the hen when more eager for
coition than her mate. It occurs very frequently and in prolonged
"sessions" between birds that are paired but whose nesting is de-
layed because they are not yet in full reproductive condition.
Whichever bird is more forward sexually usually does a major share
of the caressing. In the Barbary Dove and C. livia (probably also
in other species) both sexes, but more often the male, may caress
their young, particularly when these are fully fledged and beginning
to fly about. Juvenile birds, even when still in the nest, may caress
each other. Owing to the amount of caressing indulged in being so
much correlated with sexual readiness, or lack of it, it is rather
dangerous to try to draw specific distinctions. There seems little
doubt, however, that the Streptopelia species indulge in this
behaviour to a greater extent than do the European species of
Columba. If my breeding pair of Speckled Pigeons were typical,
then this species also does a great deal of caressing.
I think there can be no doubt that the function of caressing—
on the physical level—is the removal of ectoparasites, and perhaps
also other foreign bodies, from the mate's head. It is often shown
when sexual tendencies are thwarted, owing either to the bird or
(more often) its mate, not being ready to copulate. At other times,
as with a newly-mated pair on a nest-site, it seems likely that it is
aggressive, rather than sexual, impulses that are checked and find
an outlet in caressing. Although caressing is shown in situations
where aggressive or sexual impulses are inhibited, I do not think
it likely that the bird caressing its mate or young is usually in the
same mood as one about to attack or copulate. On the contrary, it
appears rather to be in a mood of affectionate tenderness in which
overt sexual and aggressive tendencies have been sublimated. As
has been mentioned, the caressing partner may, however, be quite
rough on occasions. In juvenile pigeons one often sees aggressive
behaviour "switch over" into caressing. Whitman (1919) gives an
example of a young C. livia which attacked and tried to drive away
a Passenger Pigeon (Ectopistes migratorius) caged with it. If the
latter fought back the young bird would "lovingly fondle" the
Passenger's head feathers. Tempting as it is to follow Whitman's
implication that this behaviour showed adaptability and intelligence
(directed towards maintenance of its position at the food dish) I

1956 BEHAVIOUR OF PIGEONS 33
doubt this interpretation. It seems more likely that when the young
C. livia met with a strong resistance, its mood changed to one of
submissive affection which inhibited further overt aggression, but
allowed the now sublimated aggressive impulses to express them-
selves as caressing.
Normally, in pigeons, caressing only takes place between mates,
or other birds that have an apparently affectionate social bond
between them. The bird actually being caressed often moves its
head about as if trying to avoid the other's attentions. This is par-
ticularly the case with young birds caressed by their parents.
Mutual preening between "casual acquaintances" such as takes
place in some of the weaver-finches, and deliberate solicitation of
caressing seem not to occur. The evidence suggests that in all situ-
ations in which caressing occurs, the caressing bird would have
become angry and shown aggression had there not been a bond of
affection between it and the recipient of its caresses.
DRIVING
This occurs regularly in Domestic Pigeons, kept in flocks, from
about a week or more before laying until the first egg has been
laid. It can also be seen among wild or feral populations of C. livia
wherever numbers of breeding birds are gathered at some feeding
ground. In its more intense forms, as seen in the pigeon loft or
some town square, it is familiar to all. The driving cock follows his
mate everywhere, often litePally "treading on her tail". He pecks,
usually in a gentle manner, but sometimes fircely, at her head. If
the hen takes wing he flies closely behind and above her. If she goes
to the nest (domestic birds) the male stops driving.
Most pigeon-keepers suppose that the male is trying to force
his mate to go to the nest. The Heinroths (1949) consider this the
case and that the function of driving is to impress (presumably on
the female) the whereabouts of the nest-site. As evidence for this
they state that driving is most intense in young pairs or in pairs
which are nesting in a new site. Whitman (i9I9) noted that driving
only took place when other pigeons were present and considered
that the driving male was impelled by feelings of jealousy or posses-
siveness towards the female, but he also concluded that "the pur-
pose of the act is to force the female to the nest." Levi (194o),
although subscribing to the popular theory, notes that sometimes
the male is contented with getting the female away on to a perch
by herself.
I have previously (Goodwin, 1955) expressed doubts as to the
correctness of the "driving to nest" hypothesis and am now satis-
fied that I was correct in so-doing. Driving does not take place—
even with young pigeons or pairs using a new nest-site--if there
are not others of their species nearby. Of course a male pigeon that
is reacting socially to man will often begin to drive his mate when
a human being approaches them closely. The male's behaviour
when driving shows a variable intermixture of aggressive and
sexual elements. If the female goes to a distance of about thirty
yards or more from other pigeons, he usually stops driving. If she
flies on to a ledge or roof `where there are no others he invariably
34 BEHAVIOUR OF PIGEONS 3(I)

stops. If the female approaches other pigeons—or they approach

her—the male begins to drive by getting between his mate and the
others, thus forcing her to move away from them. Occasionally
driving pairs are seen on the ground at a greater distance from
others of their species, and one may often see males driving in
flight when no others are anywhere near. I think in both cases this
is due to the male's not having realised that the pair are now
alone. In the first if the female stops for any reason the male will
look around and (apparently on seeing they are now alone together)
cease to drive. That the same explanation is true for the male
driving in flight is suggested by the number of times when I have
seen a pair fly up from some crowded feeding ground, and the male,
who was at first driving hard in flight, swing out beside or in front
of his mate as they got well away from others in the air, usually
going into display flight as he did so.
A newly-paired Domestic Pigeon (a light-print tippler) did not
drive his mate (a blue feral bird from London) at all during the first
two nesting cycles, for each of which they used nest-sites new to
both of them. When the third cycle commenced a male juvenile
(their own offspring) and an unpaired male (a silver tumbler) were
also present. The light-print did not now drive his mate, as I had
expected, but he frequently drove the silver away from her, chasing
him in a manner very similar to the most intense and aggressive
forms of driving. He showed no annoyance whatsoever at the
presence of his offspring who—unlike the silver—did not court the
blue hen or interfere with their copulation. In subsequent nesting
cycles of this pair, numbers of other pigeons were being kept and
the Light-Print drove his mate in the "normal" manner.
Immediately prior to driving, or after stopping (as a result of
the female now being away from others) the male usually shows
signs of sexual desire (swallowing, preening behind the wing, etc.).
His posture when driving could equally well be aggressive or sexual
in implication, but the usually gentle nature of the pecks delivered
are almost certainly due to aggressiveness being inhibited by sexual
feeling. When "driving hard" the male's aggressive impulses
often "get the better of him" and he may fairly run after the
harried female, pecking hard at her head. That this is a form of re-
directed aggression and that the male's mood is one of anger, caused
by thwarting, seems to me extremely probable, if unprovable. The
fiercest driving I have ever seen was by a male with swollen feet.
These appeared to cause him much pain as he limped awkwardly
after his mate on the hard pathway. Every few seconds he would
stop and rest a moment, then catch up his mate by taking a short
flight to her. Whenever he reached her he pecked her with the ut-
most ferocity. That pain or thwarting usually arouse feelings of
anger in Man is common knowledge, and the same would seem true
for Columba livia.
During the driving period, or most of it, the female is sexually
receptive. Should her mate be killed or removed she will usually
pair again at once with any male that offers. If her own mate is
sexually inactive she may desert him and, even when she does not,
will allow another male to tread her.

1956 BEHAVIOUR OF PIGEONS 35
From the above facts it is, I think, evident that the purpose of
driving—of which the male gives the impression of being quite
conscious—is to remove the female from the immediate neighbour-
hood of possible sexual rivals. It functions to prevent the female
from being fertilized by other males and to give the pair oppor-
tunity to copulate undisturbed.
That in a relatively crowded pigeon-loft or aviary the male does
not stop driving unless his mate goes to the nest is doubtless
because under these conditions his nest-box is the only place where
the male feels sufficiently secure from interference to be at ease
with his mate at this stage of the nesting cycle.
I have not been able to study driving to the same extent in any
other species. The Heinroths (1949) state that the Wood Pigeon
drives with a characteristic call ("Treiblaut"). I have never ob-
served driving in this species, nor has Cramp (personal communi-
cation) who has been studying it for some years. In the Stock
Dove I have seen driving, in the same manner as that of C. livia,
taking place in fields where many of the species were feeding. Un-
fortunately, I omitted at the time to note down precise details.
Since becoming seriously interested in this aspect of pigeon
behaviour, I have only twice seen driving in the Stock Dove. In
both instances a pair were on the ground within the nesting colony
area and at the close approach of another of his sex the male,
getting between them, drove his mate away from it. Whitman
(1919) gives instances of driving (always when others of the same
or allied species were present) in the Mourning Dove (Zenaidura
macroura), Common Bronzewing (Phaps chalcoptera) and Barbary
Dove. When I had several Barbary Doves flying at liberty (Good-
win, 1952) I often saw driving, always when other males were
present. In the winter of 1 954 -5 I placed two pairs of Barbaries in
an aviary about sixteen feet square and seven feet high. At first
one male dominated the rest, but by February the other male had
claimed a small territory in a corner where he and his mate were
nesting. Shortly after this his more courageous attitude resulted in
his being tolerated on the ground also. If he was on the ground with
his mate and the other male approached, he would get between
them and drive his mate away. When so-doing he used the typical
attacking posture but with little or no erection of the back and rump
feathers, and his pecks at the female's head were very gentle. He
never tried to drive his mate from the vicinity of the other female.
A pair at liberty in the spring of 1955 at first ignored the Domestic
Pigeons. Then the unpaired Silver Tumbler cock began to display
at the hen Barbary Dove. After this her mate would drive her away,
or attempt to do so, whenever she approached or was approached
by the Domestic Pigeons.
These observations suggest that driving in other species has the
same causation and serves the same function as in C. livia. In no
case is there any evidence to suggest that driving ever takes place
except as a response to the presence of sexual rivals, or creatures
so considered by the driving male. In a natural state it probably
occurs only rarely—if at all—except in species that nest colonially
and/or feed in flocks during the breeding season.

36 BEHAVIOUR OF PIGEONS 3(I)
INTERFERENCE WITH COPULATION
In C. livia copulating pairs are very frequently interfered with
by others. Whitman infers, no doubt correctly, that this may occur
in other species. I was for long unable to find any reasonable in-
terpretation of this behaviour. The Heinroths' (1949) suggestion
that the birds feel similar subjective emotions of indignation as do
some humans at the sight of others behaving sexually, may well be
correct, but hardly provides a functional explanation.
Attacks on copulating pairs are normally made only by males in
full sexual condition. The interfering bird runs or flies straight at
the mating pair, dislodges the mounted male, then at once begins to
peck at the female's head. If the female remains crouching, he may
mount and copulate with her, but this does not often occur. When
the female rises the male usually desists from his attack, and often
begins the bowing display. Rarely he may chase the female a short
distance, pecking at her head. If several males interfere simul-
taneously (as often happens in such places as Trafalgar Square
where large numbers of male birds are present) some or all of
them will often chase the female for some moments after, pecking
her head as they do so. From the above it will be seen that
throughout the period that his attention is focused on her, the
behaviour of the interfering male towards the female is identical
with that of a very aggressive driving male towards his own mate.
Interference with copulation is, in fact, part of the same behaviour-
complex as driving. As we have seen the latter functions to prevent
the female being fertilized by a male other than her mate and is
elicited by the presence of a potential sexual rival in her vicinity.
Since such driving is shown more intensely according to the degree
of sexual interest which other birds show in his mate, it is reason-
able to suppose that the highest possible stimulus for it would be
the sight of another male actually mounting her. In such a situation
it is obvious that only an instantaneous reaction could remedy
matters. A male who paused to make sure that it was his mate and
not another female who was involved could never interfere in time
to prevent insemination. Hence, it is not surprising to find that
selection has favoured those males that showed an instantaneous
response to the sight of any others of their kind copulating. That
this is the true explanation is shown by the behaviour of the inter-
fering male towards the female, his apparent "indignation", his
ferocious attempt to drive her quickly away, and his desisting as
soon as he "realises" that she is not his mate. The continued driv-
ing that may occur if more than one male interferes is due to the fact
that in such circumstances each is for some moments too occupied
in trying to get her away from the others to recognise her.
It may be objected that females have been recorded (Whitman,
1919) interfering with copulation. This, however, may be due to the
bird in question showing male behaviour. The only female that I
can remember seeing behave in this way was one that was homo-
sexually paired. I have kept watch on the interfering individuals in
scores of cases of interrupted copulations among free-living feral
C. livia. Invariably such birds were very soon seen to give the pre-
Ig5b BEHAVIOUR OF PIGEONS
37
dominantly sexual version of the bowing display, thus proving
themselves males in full sexual condition.
ACKNOWLEDGEMENTS
I am indebted for much helpful discussion and for information
on pigeon behaviour to Dr. G. Bodenstein, Miss T. Gompertz, Mr.
G. T. Kay, Mr. Kenneth Simmons, Dr. David Snow and Mr.
Bertram Stone. ,

REFERENCES
GOODWIN, D. 1952. Observations on Barbary Doves kept at semi-
liberty. Avic. Mag. 58:205-19.
GOODWIN, D. 1955. Notes on European wild pigeons. Avic. Mag.,
61:54-85.
HEINROTH, 0. & K. 1949. Verhaltensweisen der Felsentaube (Hause-
taube) Columba livia livia L. Z. Tierpsych., 6:153-201.
LEVI, W. M. 1941. The Pigeon. Columbia S.C.
WHITMAN, C. o. 1919. The Behaviour of Pigeons. Washington.

SEAWARD FLIGHTS OF SWIFTS AT DUSK

by
DAVID LACK
(Edward Grey Institute of Field Ornithology, Oxford)
Received 29 September, 1955
SUMMARY
7. On the east coast in summer, Swifts were seen setting out to sea at
dusk, either after feeding over a mere, when some, but not all, those present
headed seawards, or at the end of a coasting weather-movement, when the
birds stopped, circled and went out.
2. It is suggested that the birds were strangers to the district and that they
spent the night at sea.
3. A B.T.O. Enquiry is proposed to discover more about such movements.

INTRODUCTION
The following observations, made jointly with my wife, are pre-
liminary, but they are published now since the best way to advance
the subject further is with the help of others, and at the end of this
paper a proposal is made for a B.T.O. Enquiry on the subject. I am
most grateful to the Royal Society for the Protection of Birds for
permission to watch from the sea-wall at Minsmere. All times men-
tioned in this paper are G.M.T.
At 8.3o p.m. with an overcast sky on 2 July, 1952, as the light
failed, ten Swifts (A pus apus) flew high over us on the beach at
Minsmere, Suffolk, and went out to sea northeast. They were only
just visible to the naked eye but were picked up as they screamed
repeatedly. The next occasion when we were at the same place at
dusk was on 8 July, 1954• At 8.37 p.m., with a clear sky, 33 Swifts
that had been feeding over the marshes flew together to the coast,
screamed once or twice as they crossed it and set out to sea east-
wards. Visibility was excellent and they were still flying east when
lost to sight. At our request, Dr. M. C. Radford watched at the