397

HUNGER-DEPENDENCY OF HIDING BEHAVIOUR AFTER A

PREDATOR ATTACK IN DOMINANT AND SUBORDINATE
WILLOW TITS
KARl KOIVULA, SEPPO RYTKONEN & MARKKU ORELL

ABSTRACT We studied rank-specificity of predation risk taking in captive

Willow Tits Parus montanus. Level of risk taking was expressed by dura-
tion of hiding after a simulated predator attack. By manipulating available
food, we separated the effects of hunger and other rank associated factors
on accepted predation risk. As expected, hungry birds took more risks than
satiated ones. When both the dominants and the subordinates were hungry
or satiated, no rank-related differences existed in the duration of hiding.
When the dominants had consumed more food than the subordinates, they
also hid longer. Similarly, the subordinates hid longer when they were sati-
ated and dominants hungry. We conclude that incautiousness of subordi-
nates, reported earlier with regard to variety of antipredatory behaviours of
flocking birds, probably reflects a strategy to cope with high risk of starva-
tion resulting from restricted access to food.
Department of Biology, University of Qu1u, Linnanmaa, FIN-90570 Qu1u,
Finland. Fax: +358-81-5561227; E-mail: ktkoivu1@cc.ou1u.fi. Correspon-
dence to Karl Koivu1a.

INTRODUCTION such as foraging. When the tits Parus spp. detect

Obtaining food and avoiding predators are often
considered as conflicting goals for animals. In
many instances this trade-off has been converted
into a time allocation problem. Especially in
flocking birds the relationship between group size
and time devoted to foraging and scanning for
predators has been studied a lot (see e.g. Elgar
1989 and references). Optimality models predict
that when compromising between predation and
starvation risks, hungry animals should take gre-
ater risks in terms of predator avoidance than the
ones near satiation (e.g. McCleery 1978, Krebs
1980, Krebs & Kacelnik 1991).
Dominance secures free access to essential re-
sources such as food or shelter (e.g. Kaufmann
1983). Because of restricted access to food, prob-
ability of starvation can be higher among subordi-
nates. To cope with increased starvation risk sub-
ordinates may be forced to decrease time devoted
to antipredator activities.
One aspect of predation risk emerges when a
predator appears and interrupts other activities
a predator, they usually rapidly fly into a hide,
dense bushes or trees where they freeze (e.g.
Morse 1973, Ficken & Witkin 1977). After the
predator has disappeared, birds have to decide
when to resume feeding. The probability that the
predator could still be in the vicinity obviously
decreases as time passes. Therefore, hesitation
can increase chances of survival i.e. be less risky
than ceasing from hiding earlier. On the other
hand, starvation risk increases in the course of
time used in hiding.
Regner (1985) found that subordinate captive
Blue Tits Parus caeruleus returned to a feeder
earlier than dominants after presentation of Spar-
rowhawk Accipiter nisus model. De Laet (1985),
Waite & Grubb (1987) and Rogstad (1988a) have
obtained similar results from other Paridae.
There are several alternatives, although not mutu-
ally exclusive, how this kind of rank-dependence
could evolve. First, it is possible that because of
their free access to food dominants could have
better nutritional status than subordinates, who
therefore could be willing to take more predation

Received 29 December 1993, accepted 2 March 1995 ARDEA 83:397-404

398 ARDEA 83(2), 1995
risks to avoid starvation. Second, if dominants
can exclude subordinates from food sources, sub-
ordinates could be forced to feed during unsafe
periods when dominants are not using the feeding
sites. A third alternative is that the higher risk tak-
ing among subordinates might be caused by rank-
correlated factors not attached with food access.
For example, subordinates could be younger and
less experienced, and hence, less cautious than
dominants.
If risk taking in terms of predation is hunger
mediated, subordinates should take more risks
than dominants only when they are hungrier than
dominants. On the other hand, if accepted high
predation risk reflects a strategy to cope with re-
stricted time in access to food, subordinates could
risk more regardless of their hunger levels. Simi-
larly, if the factors that determine the social status
are correlated with risk taking, differences in hun-
ger levels should not have an influence in them.
To test these predictions, we measured risk
taking and manipulated hunger levels of captive
Willow Tits Parus montanus, which is a forest-
living, year around resident passerine that forms
dominance-structured flocks during the non-
breeding season.
METHODS
The birds
The experiments were conducted in Oulu,
northern Finland (65°N 25°30'E) during two suc-
cessive winters 1989-1991. We used captive birds,
because it is nearly impossible to individually
manipulate the hunger levels of birds in natural
conditions. The birds for the experiments were
captured at seed-baited feeders using mist-nets.
To get the birds habituated to sunflower seed diet,
the feeders were maintained for several weeks be-
fore catching the birds. Netting always took place
before noon. The birds were immediately trans-
ported indoors. They were marked and their wing,
tail and tarsus lengths were measured using meth-
ods recommended by Svensson (1984). The birds
were sexed by dimorphism in wing and tarsus
lengths (for details see Koivula & Orell 1988).
Two age-classes were separated: adults (born be-
fore the previous summer) and juveniles (born in
the previous summer). Ageing was based on dif-
ferences in tail feather shape and abrasion (Laak-
sonen & Lehikoinen 1976).
The birds were habituated to captivity condi-
tions for at least one week before the experiments.
They were kept separately in wire-meshed cages
(size: 50*80*50 cm). The cages were covered
with white clothes to avoid disturbance and visual
contact to other individuals. Furthermore, the
birds were maintained on ad libitum diet of sun-
flower seeds and water.
The experiments
The experimental cages consisted of two
parts: inner and outer compartments. The inner
cage was similar to the ones in which the birds
were housed during habituation. On the other
side, there were two perches and a thick hiding
place that was built using small birch and spruce
branches. Food and water dishes were situated on
the open side. The floor was wire-meshed and did
not touch the platform surface. Therefore, once
dropped, the seeds were not available for the birds
any more and therefore the food dish was the only
available seed source. The amount of food offered
was controlled by an observer who supplied food
through a plastic tube which pierced the walls of
both cages and ended in the food dish. The walls
of the outer cage were cloth-walled and higher
than those of the inner cage. Therefore, the birds
could not see the observer sitting outside. A loose
piece of cloth formed a ceiling which was re-
moved on the day before the predator presenta-
tions. The birds were observed through a small
hole in the wall of the outer cage.
Two birds of the same sex were introduced at
the same time into the experiment cage on the day
before the first trial. The birds came from differ-
ent winter flocks and were therefore not familiar
to each other. In the evening the food dishes were
emptied and the lights were switched off. The
darkness lasted for 17 hours (16.00 - 09.00),
which represents the approximate light conditions
 Koivula et al.: RISK TAKING IN WILLOW TITS
of midwinter in northern Finland. The tempera-
ture of the room was constantly kept near +20 dc.
In the morning, after the lights were switched on,
the birds were deprived of food for two hours be-
fore the experiments were conducted.
Dominance
When measuring the dominance order, sun-
flower seeds were given to the birds one by one.
The next seed was provided only after the bird
that picked the former one had consumed it. The
individual which first had picked and consumed
10 seeds was considered to be the dominant one.
Usually, subordinate individuals got no seeds.
The dominants, according to this method, were
the winners in all the observed agonistic interac-
tions, too.
Predator presentation and risk variables
A mounted male Tengmalm's Owl Aegolius
funereus, a realistic predator for Willow Tits
(Korpimaki 1981), was used as a predator model.
The owl was mounted to a standing position with
its head facing downwards. It was placed by the
observer on the upper frame of the outer cage at
the same end where the food dishes were situated
in the inner cage. The birds did not see the ob-
server during the placement. Model presentation
lasted for 15 seconds. Food dishes that had been
empty for the preceding two hours were filled
during the predator presentation. When the model
emerged, the birds rapidly moved into the hide,
froze and occasionally gave alarm calls.
Two time variables were measured after the
removal of the owl: the time that passed before
the bird started to move inside the hiding place
("scanning period", SP) and the time between the
removal and the moment when the birds left the
thicket ("hiding period", HP). After leaving the
cover, the birds usually came to the food dish im-
mediately.
Treatments
Experiment 1 Three kinds of nutritional situa-
tions were arranged for each dyad: (1) neither
bird got food before the model presentation; (2)
399
both birds were satiated by giving them 40 seeds
at the same time; (3) the dominant bird got more
seeds than the subordinate. Last of the treatments
was arranged by offering the seeds to the birds
one by one as when measuring dominance. In this
treatment the subordinate sometimes succeeded
to get some (1-4) seeds. In such cases we gave as
many additional seeds as needed, so that finally
the dominant had got 10 seeds more than the sub-
ordinate. We altered the sequence of treatments to
avoid errors due to possible habituation to preda-
tor model. After the trial the birds got food in ex-
cess. Only one nutritional treatment and predator
presentation trial was conducted in one day for
each dyad.
A total of 13 dyads were tested. One pair,
however, was excluded due to the death of the
subordinate counterpart on the day the last trial
was conducted.
Experiment 2 Cage arrangements of the first ex-
periment did not allow a treatment where the sub-
ordinates would have been more satiated than the
dominants. Therefore, the design was changed by
building a cage consisting of two compartments.
A tunnel with a trap door was built between them.
Closing the door when the birds were in different
cages enabled us to control the hunger levels of
both birds. Two treatments were performed: dom-
inants satiated and subordinates hungry and vice
versa. Also here the birds were deprived of food
before the experiments and the sequence of treat-
mentswas altered. The bird that was made sati-
ated got 10 sunflower seeds and the other one got
none. After this the trap door was opened and clo-
sed again immediately when the birds were in the
same compartment. Predator presentation and the
variables measured were the same as in experi-
ment 1. A total of 12 dyads were tested here.
Statistics
Despite the transformation attempts, durations
of scanning and hiding periods were not normally
distributed. Therefore, non-parametric methods
were used when comparing risk-taking between
400 ARDEA 83(2), 1995

hunger levels and dominance classes. All the tests

were two-tailed. Critical values were obtained
180 o
from Rohlf & Sokal (1981). 150

~ 120
Ol
<:
RESULTS "c 90
<:
Cll
0
(J) 60
The duration of motionless scanning and time
30
spent inside the cover varied greatly between the
individuals and treatments. The shortest scanning 0
period lasted only for 3 seconds while the most
hesitating bird spent 11 minutes inside the hide.
180 ®
150
However, the frequency distributions of both
scanning and hiding were skewed towards short
periods. Most of the birds started to move and left
~
.e
Ol
120
/***
<: 90
the cover within the first minute. The birds scan-
ned on the average for 60.5 seconds (SE = 6.5,
~
..c:
60 / /***
n = 120). The mean duration of hiding period was 30 ~ 0 expenment 1
76.6 seconds (SE = 8.4, n = 120). • experiment 2
0
hungry satiated

Predation risk and hunger levels
The birds had significantly shorter scanning
and hiding periods when they were hungry than
when satiated. When no seeds were given before
the predator presentation, the average duration of
both scanning and hiding periods was approxi-
mately 30 seconds (Fig. 1). When the birds were
satiated, scanning lasted on the average for 1 min-
ute and hiding nearly for 2 minutes (Fig. 1). In ex-
periment 1, one individual scanned equally long
after both treatments, but 20 birds out of 24 (83%)
scanned more and stayed inside the cover longer
when they were satiated (Wilcoxon's matched-
pairs signed-ranks. tests; SP: T = 30, n = 23,
p =0.001, HP: T = 29, n = 24,p =0.0005, Fig. 1).
Similarly in experiment 2, 83% of the birds had
longer SP's and HP's when they were satiated
(SP: T = 23, n = 24, p = 0.0003, HP: T = 28.5,
n = 24, p = 0.0005, Fig. 1).
Predation risk and dominance
When the dominants got more food they were
more cautious than the subordinates. They both
scanned and remained inside the hide signifi-
Figure 1. Time stayed motionless (SP) and time
spent inside the cover (HP) in different stages of hun-
ger. Means with SE-bars. n = 24 in all the cases. Wit-
hin-individual comparisons between treatments with
Wilcoxon's matched-pairs signed-ranks tests (2-tailed).
*** = p < 0.001.
cantly longer than the subordinates in both ex-
periments (experiment 1: SP: T = 8, n = 12,
p = 0.012, HP: T = 8, n = 12, p = 0.012, Fig. 2c;
experiment 2: SP: T = 10, n = 12, p = 0.021, HP:
T = 8, n = 12, p =0.012, Fig. 3a). In experiment 2,
however, dominants took more risks than subor-
dinates when they were hungrier; in 11 out of 12
dyads they had shorter scanning and hiding peri-
ods than subordinates (SP: p = 0.023, HP:
p = 0.023, Fig. 3b). No such differences existed in
experiment 1 when the hunger levels were equal.
When both the birds were hungry, dominants
were the first to move and leave the cover in 6 out
of 12 cases (SP: T = 37, n = 12, p = 0.88, HP:
T = 33.5, n = 12, p = 0.67, Fig. 2a). Similarly,
when both the birds were satiated dominants had
shorter scanning and hiding periods in 50% of the
trials (SP: T = 39, n = 12, P = 1, HP: T = 37,
 Koivula et al.: RISK TAKING IN WILLOW TITS 401

180 180
150 150
120 120
~ ~
Ql 90 Ql 90
E
:;::; ~
60 n.s. n.s. 60
30 30
0 0

180 ® 180
®
150 150
120 120
~ ~
Ql 90 Ql 90
E E
:;::;
:;::; 60 60
30 30 dominants
subordinates
0
O:=~~~=============~ scanning hiding
180
DOMINANT SATIATED
©
Figure 3. Lengths of scanning and hiding periods in
150 experiment 2 when (a) dominant was satiated and sub-
120 ordinate hungry and when (b) dominant was hungry
~ and subordinate satiated. Means with SE-bars. n = 12
Ql 90 for each treatment. Difference between dominance cat-
.§ egories tested with Wilcoxon's matched-pairs signed-
60
ranks test (2-tailed). * = p < 0.05.
30 dominants
subordinates
0

Figure 2. Mean lengths (with SE-bars) of scanning
and hiding periods of dominants and subordinates in
experiment 1 when (a) both the birds were hungry, (b)
both were satiated, and (c) dominants were satiated and
subordinates hungry. n = 12 for each treatment. Differ-
ences between dominance categories tested with Wil-
coxon's matched-pairs signed-ranks test (2-tailed).
n.s. = p > 0.10, * = p < 0.05.
n = 12, p = 0.88, Fig. 2b). The results show that
dominance without differences in satiation has no
impact on a taken predation risk. Subordinates
can take more risks than dominants in terms of
predation, but only when they are hungrier than
the dominants.
DISCUSSION
Recent research suggests that varying predation
pressure is the key factor behind the rank-depen-
dent survival observed in certain birds, among
others in the Willow Tit (e.g. Ekman et al. 1981,
Ekman 1987, Desrochers 1989). Additionally, for-
aging time seems to be critical for survival in
temperate tits (e.g. Jansson et al. 1981, Ekman
1987). In fact, rank-dependent time allocation pat-
tern forms an evident link between rank and sur-
vival (e.g. Ekman 1990).
It is known that because of presence of domi-
nants, subordinate Willow Tits use more open or
otherwise unsafe microhabitats than dominants
(Ekman & Askenmo 1984, Rogstad 1988 a,b). As
402 ARDEA
with foraging habitat, exposure to predation may
vary with foraging time. We assumed that the
time passed after the appearance of the predator
predicts its probability to be nearby, which in tum
equals to a predation risk. In this sense the risk
taking of Willow Tits was consistently dependent
on their hunger levels. Thus, Willow Tits seem to
be capable of adjusting their foraging behaviour
to balance with predation and starvation risks.
The results are in line with Krebs (1980) who
found risk taking in Great Tits Parus major to be
state-dependent: tits allocated more time for scan-
ning when approaching satiation. Similar hunger-
dependency of predation risk taking has also been
reported with regard to other aspects of foraging
and antipredator behaviour (Milinski & Heller
1978).
Subordinate Willow Tits were less cautious
than the dominants, but only when being more
hungry than them. There are results of four differ-
ent Parus species showing that subordinates are
less cautious and resume feeding earlier than
dominants after the appearance of a natural pred-
ator (De Laet 1985) or a model of it (Hegner 1985,
Waite & Grubb 1987, Hogstad 1988a). The obser-
vation that subordinates take more risks than
dominants has also been reported in studies con-
cerning the openness of foraging habitat. Subor-
dinates use more feeding sites far from protective
cover (Schneider 1984, Hogstad 1988a, Piper
1990, Koivula et at. 1994). Hunger levels, how-
ever, were not controlled in these studies. In fact,
De Laet (1985) reported that dominant Great Tits
visited the feeders more often than subordinates
before the predator appearance. Therefore, their
nutritional status could also have been better.
Similarly, dominant Blue Tits in Hegner's (1985)
laboratory study tended to spend more time in
feeders before the presentation of the predator
model. In captive Tufted Titmice Parus bicotor
Waite & Grubb (1987) tried to minimize the effect
of hunger levels, although they did not equalize
the food provision in a quantitative manner. Re-
sults of Waite & Grubb (1987) suggest that domi-
nants could behave more cautiously than subordi-
nates even if there were no differences in hunger
83(2), 1995
levels. In our study the absence of differences in
risk taking, when the dominant and subordinate
Willow Tits were both hungry or both satiated,
suggests that dominance per se does not cause
any differences in risk taking. Thus, it is not prob-
able that subordinates in natural flocks risk more
to acquire rank-limited foraging time in the ab-
sence of dominants. In fact, this possibility is re-
alistic only if the food sources are so highly clum-
ped that simultaneous feeding by many group
members is not possible. Food at the feeders can
provide such circumstances, but as far as natural
food is concerned, this is hardly the case in boreal
tits.
Our experiments consisted of unisexual dy-
ads. Therefore, we could exclude the possible
sex-specificity in antipredatory behaviour. Simi-
larly, with one exception, the birds in dyads of ex-
periment 1 were matched with respect to their
age, although in captivity age does not correlate
with rank among birds of matching status of resi-
dency (Koivula et at. 1993). Nevertheless, there is
evidence for sex- or age-related predation risk
taking, for example with respect to scanning for
predators (e.g. Heinsohn 1987, Elgar 1989), giving
alarm calls (Alatalo & Helle 1990), or responding
to alarming (Breitwisch & Hudak 1989). There-
fore, we cannot rule out the possibility that corre-
lates of rank such as sex and age are partly re-
sponsible for the reported associations between
rank and predation risk. Further experiments are
needed to solve their role. Our results, however,
show that hunger level can be a correlate of dom-
inance playing an important role in shaping those
associations. The fact that the dominants were
less cautious than the subordinates when they
were more hungry, strengthens this interpretation.
The type of risk taking that we handled can
have direct survival consequences. In addition,
there is a hypothetical indirect mechanism that
could augment the rank-related differences in pre-
dation pressure. When the subordinates resume
feeding first, the predator can attack them if it is
still in the vicinity. Consequently, the dominants
can have a possibility to get information whether
it is safe for them to resume feeding or not. In the-
 Koivula et al.: RISK TAKING IN WILLOW TITS
ory, using subordinates as decoys, dominants can
by small loss of foraging time reach a low level of
predation risk, which without incautiousness of
subordinates would have required a substantially
longer hiding time, and hence, a higher risk of
starvation.
ACKNOWLEDGEMENTS
We thank Kimmo H. Kumpulainen and Kimmo Lahti
for helping us in catching the birds and maintaining
them in the laboratory. The study was financially sup-
ported by the Finnish Cultural Foundation, the Acad-
emy of Finland and the University of Oulu.
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SAMENVATTING
Van Matkopmezen werd in gevangenschap bekeken
hoe individuen van verschillende sociale status in een
groep reageerden op het aanbod van voedse1 in re1atie
tot de aanwezigheid van een predator. Ret nemen van
risico werd hierbij uitgedrukt als de duur waarin ge-
schuild werd na een aanva1 van een gesimuleerde pre-
dator. De aangeboden hoeveelheid voedsel werd gema-
nipuleerd, om uit te zoeken tot welk niveau risico aan-
vaardbaar werd geacht door de vogels. Zoals verwacht
namen uitgehongerde individuen meer risico dan ver-
zadigde soortgenoten, ongeacht hun socia1e status. Er
werd geen verschil gevonden in schui1gedrag tussen
dominante en ondergeschikte vogels bij een gelijk aan-
bod van voedsel. De auteurs concluderen dan ook dat
het tijdens eerdere onderzoeken gevonden roeke1oze
gedrag van ondergeschikte vogels in een groep in re1a-
tie tot diverse antipredator-gedragingen te maken heeft
met het hoge risico van uithongering als gevolg van een
beperkt voedselaanbod. - GOK