BRIEF COMMUNICATIONS
COMMENTS ON REDIRECTION WITH EXAMPLES OF AVIAN
COPULATIONS WITH SUBSTITUTE OBJECTS
By MILLICENT S . FICKEN AND WILLIAM C . DILGER
Laboratory of Ornithology, Cornell University, Ithaca, New York
Animals sometimes direct activities toward an
object or animal other than the one which
normally releases a particular behaviour. This is
termed redirection and has been defined by
Moynihan (1955) as follows : "autochthonous
activities of a drive directed toward an object or
animal other than the one releasing and usually
directing them (although the releasing object or
animal remains available, or partly available, as a
potential goal at the time) ." Incidentally, recent
work by Andrew (1956) and Van lersel & Bol
(1958) strongly suggests that the term "alloch-
thonous" may no longer be valid, thus rendering
the term "autochthonous" unnecessary . Further-
more, until more information is accumulated
bearing on the causal factors for phenomena
such as redirection and displacement these terms
should be used in a purely descriptive sense .
Our experiences with redirection (examples of
which follow) have led us to feel that these actu-
ally may be best considered as reactions to sub-
normal stimuli rather than a separate pheno-
menon .
There are numerous reports of redirected
aggression in the ethological literature, but pub-
lished examples of redirection involving other
tendencies are relatively rare . However, there are
scattered references in the ornithological liter-
ature which seem to be examples of redirected
copulations .
A male Agapornis fischeri (Fischer's Lovebird)
was placed with a female of the same species .
The male came into sexual condition earlier
than did the female and demonstrated intense
precopulatory behaviour which initially achieved
little in the way of overt sexual responses from
the female . She, in fact, seemed to largely
"ignore" the prolonged and intense displays of
the male (females of the Agapornis species are
commonly somewhat aggressive toward sex-
ually behaving males) . The male frequently
tried to mount the female . She responded to
this by being aggressive or by merely sidling out
from under him while he still had one foot on
the perch . At this point the male would scratch
his bill (a common and more or less ritualized
act, depending on the species, of Agapornis) .
219
After a few days of intermittent activity of this
type the male began to culminate his bouts of
precopulatory behaviour with copulatory move-
ments on the perch as close to the female as he
could get. The movements were normal in every
respect and would last from two to four minutes
at a time (such prolonged copulations are
normal to Agapornis) . These copulations were
made immediately after the male demonstrated
intention movements to mount the female and
was thwarted because she did not assume the
soliciting posture or because she avoided and/or
threatened him . Copulation intention move-
ments of varying degrees of completeness were
made almost anywhere on the single perch (a
piece of tree branch about three-quarters of an
inch thick and two feet long) . The female, in
retreating along the perch from the male's ad-
vances, usually ended up against the far side of
the cage . About three inches from this end of the
perch was a fairly large protuberant knot . It
was upon this knot that the male performed
the complete copulations while crowding against
the "indifferent" female .
The female clearly released and directed the
normal precopulatory displays and perhaps even
released the copulatory response but she failed,
at least, to provide the proper stimuli for copula-
tion itself (the soliciting posture) . She also
commonly attacked him if he persisted in his
attempts to copulate . Hence the male responded
to the convenient knot . The female was perhaps
utilized only to the extent of providing close
contact with the male's side as he copulated .
A male Hylocichla mustelina (Wood Thrush)
was placed in a large aviary and a female of the
same species was in an adjacent aviary . The two
birds were thus in full view and hearing of one
another and were separated only by the wire
netting . The male displayed both visually and
vocally to the female . He made repeated at-
tempts to get at the female through the netting .
The early attempts to do this were clearly stimu-
lated by his attack tendency but this gradually
changed until it appeared that he had become
primarily stimulated to behave sexually . Various
activities, presumably displacement activities,
220 ANIMAL BEHAVIOUR, VIII, 3-4
occurred as a result of this thwarting of the
sexual and aggressive drives in the male . These
were primarily bill-wiping and various preening
movements . Finally, he began to make frequent
copulation intention movements while on the
ground and as close to the female as possible .
When his tendency to copulate was particularly
strong he conducted perfectly normal copulatory
motor patterns (except of course he could not
lower his tail the usual amount) on a smooth
rounded stone about an inch high and about
three inches wide by four inches long . This and
the more frequent intention movements were
performed many times over a period of days .
Again, in this case, the female provided the
external stimuli releasing and directing the
normal precopulatory displays but since she
was not available (intervening wire netting) the
male responded to another object (a convenient
stone) ; as did the male Agapornis to the knot .
A male Setophaga ruticilla (American Red-
start) was kept in the same cage with a female,
both of which were hand-raised from the age of
six days . At the age of about three months he
began to sing the advertising song frequently and
to direct "courtship" displays to the female .
These displays usually culminated in copulation
attempts . He often got as far as placing both feet
on the female's back, but never brought his tail
down far . On several occasions he dismounted
without any detectable aggression from the
female . The female's attitude was sometimes
merely "indifferent" but in other instances she
either moved away along the perch or threatened
with open gape at the male . After these un-
successful copulations, he would chase the
female around the cage for a few seconds or
merely resume feeding . Bill-wiping was observed
immediately after two of these attempted
copulations .
After three months of sporadic behaviour of
this type he was seen to attempt copulation with a
pile of dried droppings about one inch high
which had accumulated on a perch . Unlike the
observations on Agapornis this was never
observed to follow an unsuccessful copulation
attempt with the female, but generally occurred
after a period of no overt sexual activity . On one
occasion he immediately attempted to copulate
with the female after the redirected copulation .
He always sang before mounting the pile of
droppings and often got as far as placing both
feet on it with wings fluttering, pecking it, and
then dismounting . He continued these redirected
copulations for a month and largely gave up the
copulation attempts with the female, although
he still directed "courtship" displays to hei .
When copulation was thwarted, the male
Redstart performed other activities such as
feeding and chasing the female which would seem
to indicate a lowering of the tendency to behave
sexually or the increase of other tendencies
relative to sex . Presumed displacement activities
were rarely seen to follow this sexual thwarting .
However, after a period of thwarted sexual
activity another object in the cage became a
source of stimuli directing, or possibly also re-
leasing, copulatory activity and he largely gave
up copulatory attempts directed to the female
and responded instead to the pile of droppings .
That this male was a young, inexperienced
bird probably contributed to the fact that he was
able to utilize the substitute object with such
facility. It may even have been possible that the
pile of droppings gradually, through learning,
came to have releasing (as well as directing)
properties of its own.
Young (1949) observed a male Robin, Turdus
migratorius, which tried to copulate with a
mound of earth after being threatened by a
female that he was approaching. The male then
approached the female again and was rejected .
He then tried to copulate with a crumpled
newspaper . Finally, he approached the female
once more and she squatted and copulation took
place . The female then attacked and chased him .
The male in this instance was thwarted by the
fact that the female was not sexually receptive
when he initially approached (in fact she actually
threatened him) . He then attempted copulation
with the nearby substitute objects-the mound
of earth and the newspaper before the final
copulation with the tardily receptive female .
Simon (1940) found that strutting cock Sage
Grouse, Centrocercus urophasianus, frequently
tread a pile of earth as if they were copulating
with it. In fact this is of such frequent occurrence
that there is a legend among Wyoming "old
timers" that the females of this species are
fertilized externally. The females frequently feed
in the cocks' strutting ground and it was believed
that they ate the semen which the males de-
posited and thus fertilized themselves !
Discussion
In all three of the cases that we observed the
females released at least some of the pre-
copulatory behaviour of the males but were un-
available as copulation partners. In Agapornis,
Turdus, and Setophaga the females were not
 FICKEN & DILGER : COMMENTS ON REDIRECTION
available because they did not provide stimuli
directing copulation to themselves (or actively
prevented it through their aggression) . In
Hylocichla the female was unavailable because
of the intervening wire netting.
All of the substitute objects used seemed to be
non fear-provoking protuberances contrasting
with the rest of the immediate surroundings . It
seems that although copulation is a comparative-
ly complex act the substitute partners may be
very simple with but a few critical properties
similar to those characteristic of soliciting
females .
The males behaved somewhat differently in
approaching the substitute objects than when
approaching the females . There were no court-
ship displays by Turdus or Setophaga directed
toward the substitute object . The Agapornis
male regularly performed the usual bill scratch-
ing displacement activity upon being prevented
from performing the act of copulation . Later,
when he began performing copulations with the
unresisting knot, the displacement activities
understandably disappeared in this context .
The male Wood Thrush likewise abandoned dis-
placement behaviour after beginning his re-
directed copulations .
The problem arises as to the difference be-
tween reactions to subnormal stimuli and what
has been termed redirected activities . In con-
sidering the nature of the stimuli which are
operating in redirection, one must attempt to
look at them from the animal's "point of view" .
Stimuli which appear subnormal to us (by their
very form) may not "appear" this way to the
animal . In the above cases of redirected copula-
tions in Agapornis and Setophaga the females
rendered themselves unavailable to the males by
their behaviour . It is probable that the females,
.under the circumstances described, presented
stimuli less optimal for promoting copulation
than did the substitute stimuli which were finally
utilized . The best method for determining
whether one set of stimuli is less optimal than
another is to give the animal a choice between
them . The redirected copulations described are
actually natural experiments of this type . The
results would suggest that an unreceptive or
actually aggressive female may be, under certain
conditions, a less optimal object than is a stone,
knot, or a pile of droppings .
The only apparent distinction between re-
direction and reactions to subnormal stimuli is
the fact that in redirection the usual stimulus
situation initiates the behaviour (often appetitive
22 1
behaviour) . A situation might arise, for instance,
where an animal stimulates another to react,
but then becomes unavailable (e.g ., leaves)
as an adequate stimulus for the continuation of
the usual course of events, and the first animal
then continues to react to some substitute object .
This, however, would not be redirection accord-
ing to Moynihan's definition " . . . . (although the
releasing object or animal remains available, or
partly available, as a potential goal at the time)" .
It seems to us, however, that the difference be-
tween what has been termed redirection and re-
actions to suboptimal stimuli is virtually non-
existent . Therefore, we suggest that re-direction
be considered behaviour in which an act of overt
behaviour is initiated by the normal stimulus (i)
or stimulus situation and is then directed to a
substitute object or animal upon the subsequent
unavailability of the original source of stimu-
lation .
An animal normally experiences little diffi-
culty in finding a suitable object to which
aggression may be redirected . In birds, for in-
stance, pecking is an almost universal aggressive
act and any environment will usually provide
substitute objects for pecking (other animals,
the substrate, vegetation, etc .) . This is probably
one of the reasons why redirected aggression is
so much more common than are other types . If
redirection is to occur in an animal with a
thwarted sex drive, the animal must find an
object close by which will serve as a substitute
sexual partner . Such an object is necessarily
more complex and other individuals are not
likely to co-operate . The relationships among re-
actions to subnormal stimuli, redirection, and
displacement are not clear . Apparently, if the
animal cannot perform an activitated behaviour,
either to the normal situation or to an available
subnormal situation (redirection) the next most
available (not necessarily the next strongest)
drive will be disinhibited (see Van lersel & Bol,
1958) and displacement will result .
Summary
1 . Redirected copulation is described in the
Fischer's Lovebird, Agapornis fischei ; Wood
Thrush, Hylocichla mustelina ; and the American
Redstart, Setophaga ruticilla .
2 . Redirected copulation occurs when the
male is thwarted sexually because the female
(providing the initial stimulation) becomes un-
available for some reason during the normal
course of a precopulatory sequence .

222 ANIMAL tEHAVIOUR, VIII, 3-4
3 . The substitute object resembles a soliciting
female in that both are non fear-provoking pro-
tuberances in the male's immediate surroundings .
4 . In one case, that of the Redstart, the sub-
stitute object apparently came, through learning,
to have releasing (as well as directing stimuli)
of its own .
5. Apparently redirection rather than dis-
placement may be performed if the immediate
surroundings provide an adequate substitute
for the suddenly unavailable usual object or
situation .
6 . Redirection is considered to be a special
type of reaction to "subnormal" stimuli, al-
though the activity is initiated by " normal "
stimuli .
REFERENCES
Andrew, R. J . (1956). Some remarks on behaviour in
conflict situations, with special reference to
Emperiza Spp. Brit . J. anim. Behav ., 4, 41-45 .
Morris, D . (1956). The function and causation of courtship
ceremonies. L'Instinct dans le comportement des
animaux et de l'homme. Paris : Mason et Cie .
Moynihan, M . (1955) . Remarks on the original sources of
displays. Auk., 72, 240-259 .
Simon, J. R. (1940) . Mating performance of the Sage
Grouse . Auk ., 57, 467-471 .
Van lersel, J . J . A . and Bol, A. C . A. (1958) . Preening of
two tern species. A study on displacement activit-
ies . Behaviour, 15, 1-88 .
Young, H . (1949) . Atypical copulatory behaviour of a
Robin . Auk., 66, 94 .
Accepted for publication 16th May, 1960.