Journal of

Animal
Ecology . . .
1995,
64, 747-757

A field test of ideal free distribution in flockfeeding common cranes

L.M. BAUTISTA, J.C. ALONSO and J.A. ALONSO*

Museo Nacional de Ciencias Naturales. Jos Gutirrez Abascal 2. 28006 Madrid. Spain; and
*Departamento de Biologa Animal. Facultad de Biologa. Universidad Complutense. 28040 Madrid.
Spain

Summary
1. The densities of common cranes Grus grus in 10 zones of a wintering area
were approximately proportional to the amounts of food resources, but some
overuse of the zones with highest food densities was observed, i.e., greater
numbers of birds than expected used these zones. The distribution resembled
ideal free distribution only after numbers of cranes had exceeded carrying
capacity. The seasonal pattern of settlement deviated from ideal free
distribution. During the early and late part of the season, when there were
fewer birds at the study site, cranes preferred to forage as close to the roost as
possible provided that there was enough food, instead of selecting the zones
further away with highest food densities.
2. However, 12 individually marked cranes differed in their competitive
ability and foraging area selection. Larger adult birds were dominant in
aggressive encounters, displacing subdominant cranes from good feeding
positions. Food intake rate of dominant cranes tended to be higher than the
flock average, the difference increasing with rank. Dominant cranes preferred
to forage in the zones with highest food densities and had higher absolute
daily food intakes. The relative pay-offs of different phenotypes changed
across zones with different food densities: subordinate birds could not
increase their intake rate at the richest zones as much as dominants.
3. The average daily food intake of a crane was thus positively correlated with
both the quality of the foraging zone and the dominance rank of the bird.
These results fulfil most assumptions and predictions of the interference
phenotype-limited distribution model, although truncation of phenotypes
between zones was imperfect.

Key-words: interference, dominance, foraging behaviour, common crane,

ideal free distribution.

Journal of Animal Ecology (1995) 64, 747-757

Introduction
In their review of the published tests of Fretwell & Lucas's (1970) ideal free
distribution (IFD) model, Parker & Sutherland (1986) realized that many of
the first studies involving continuous-input situations showed consistent
individual differences in pay-off due to their different competitive abilities
and thus violated the basic IFD assumption of equal ability and the prediction
of equal gain (e.g. Milinski 1979; Harper 1982; Godin & Keenleyside 1984).
With respect to interference studies, most of them demonstrated differences in
average gain between sites, contradicting the second basic IFD prediction (e.g.
Monaghan 1980; Sutherland 1983; Sibly & McCleery 1983; Goss-Custard,
Clarke & Durell 1984). Parker & Sutherland (1986) recognised the need to
incorporate individual differences and formally developed the subject of IFD
of unequal competitors, proposing testable assumptions and predictions for
various phenotype-limited models.
In this paper we present a field test of the interference model. We study the
distribution of flock-foraging common cranes Grus grus throughout two
winter seasons in relation to food resources. We also study interindividual
differences in competitive ability of 12 radiotagged birds in relation to their
foraging area selection. We applied the interference model because
interference, as defined by Goss-Custard (1980), was important in our system,
both at the flock level, reducing the individual's intake rate (Alonso & Alonso
1991, 1993) and at the population level, as a cause of increased flock dispersal
(Alonso, Alonso & Veiga 1987). The main assumptions of interference
models are that competitive differences exist between individuals, that payoffs to competitors are reduced by the presence of more competitors and that
relative pay-offs to competitors change between sites. Given such conditions,
the only stable distribution of competitors is a truncated phenotype
distribution, which predicts that most competitive individuals should be at the
highest quality sites, and that food intake should be correlated with both
competitive ability and site quality. With respect to numbers of individuals in
each site, the results of the model depend on the distribution and range of
phenotypes present in the population and the exact way in which relative
competitive ability affects interference. However, where interference is low,
as expected in a gregarious feeder, the model predicts an overuse of the richest
sites, i.e. greater numbers of birds than expected at these sites (Sutherland &
Parker 1985; Parker & Sutherland 1986; Milinski & Parker 1991).
In a previous paper (Alonso & Alonso 1991) we already discussed the
suitability of Pulliam & Caraco's (1984) equilibrium flock model to the

gregarious behaviour in common cranes. Here, the present results are

discussed in relation to the extension of Pulliam & Caraco's model of flock
size selection to habitat selection. This model also predicts an overuse of the
richest habitats, due to the benefits derived from flocking for the individual.
Finally, because depletion of the relatively unrenewable food resources (seed
patches) was also important regulating the carrying capacity of the area
(Alonso, Alonso & Bautista 1994), we also discuss some predictions of
Sutherland & Anderson's (1993) depletion model: that predators should
initially exploit the richest zone, gradually increasing the range of zones
occupied, overusing the richest zones and depleting all to the same critical
level.

Material and methods

STUDY AREA AND SPECIES

The study area was located in a lake basin of 53 637 ha, at Gallocanta, NE
Spain (40o 58' N, 1o 30' W). Over the last two decades, common cranes have
increasingly used it as a staging and wintering area (Bautista, Alonso &
Alonso 1992). Most cranes arrive in November, and stay in the area some
weeks. Between late November and mid-January they decide either to remain
to winter there or to migrate further south. By mid-February, cranes begin
spring migration to their breeding areas in northern Europe. In April most
cranes have already left Gallocanta. We recorded food abundance and number
of cranes from early October to early March in 1989-90 and from early
November to late February in 1990-91. The lake is endorreic, saline, and very
shallow and extends over 1400 ha. It is used as a roost from which cranes
disperse daily to forage on the surrounding fields, most of which (28 712 ha)
are sown with winter wheat and barley, and a few with sunflower. When
cranes arrive in autumn from the breeding areas, the fields have been
harvested, and cranes feed almost exclusively on the cereal and sunflower
seeds left on the stubble fields. In average years, ploughing activities and the
depletion of stubble fields by cranes, cause a decrease in food availability
throughout the winter, which is in part compensated by new sown cereal fields
(Alonso et al. 1994). Cranes shift progressively from feeding on waste seeds
on stubbles to feeding on recently sown seeds. No other animal species
consumes sown cereal, and only small numbers of other birds and microtine
may eventually take a negligible amount of seeds from stubble fields.
However, when rainfalls are abundant after harvest, farmers advance the
agricultural cycle, tilling up to 75% of the arable land (Alonso et al. 1994). In

these conditions, sown fields progressively occupy the majority of the study
area.
This system was appropriate for a field test of ideal free distribution for the
following reasons. First, the availability of roost and fresh water were not
limiting factors of the crane population, given (i) the size of the lake and its
shallow margins, which provided more than enough roosting sites for the
cranes and (ii) the presence of several fresh water channels and streams that
were regularly used by cranes to drink. Secondly, flat landscape facilitated
location of foraging birds and accurate counting of the crane population.
Thirdly, the simplicity of both the habitat structure and the crane's diet, and
the virtual absence of food competitors during winter, made it possible to
obtain accurate measures of food availability in each zone of the study site
and to relate crane's distribution directly with food abundance.

SAMPLING PROCEDURE

We divided the study site into 10 zones, according primarily to the phenology
of their farming cycle and to their distance from the lake, and excluding nonarable land and other areas not used by the cranes as feeding grounds (Fig. 1).
Due to farming cycle differences between zones, there were clear differences
in food density throughout the winter, which enabled us to test some of the
predictions. For example, zone 3 was ploughed later than zone 2 and thus had
more food (seeds left on stubble fields after harvesting). While delimiting the
zones we tried to keep their respective surfaces utilizable by the cranes to
forage as equal as possible. The apparently large size differences in Fig. 1
were mainly due to some zones including villages and other non-utilizable
areas. However, in our tests we divided food abundance by size of the zone to
control for size differences. We estimated food availability by a car transect
every two weeks throughout the study area, recording the types of ground at
both sides of the itinerary. Every fortnight we sampled ten zones, except in
early March in 1990-91, when we sampled six. The length of the transect was
76 km in both years, and each zone was sampled in proportion to its area. The
ground types identified were cereal stubbles, sunflower stubbles, ploughed
fields, recently sown cereal fields, sprouted cereal fields, pastures, and others.
We estimated the total surface occupied by each type of ground with their
frequency values obtained from the transect. We also sampled the density of
seeds on stubble fields each fortnight, counting all seeds in twenty randomly
selected quadrats. We threw a square metal frame, measuring 25 x 25 cm on
cereal stubbles and 50 x 50 cm on sunflower stubbles, while walking in a
straight line along the diagonal of each field. The number of stubble fields
sampled varied from 98 in early November 1989-1990 to 20 in late January

1989-1990. The number of seeds on stubble fields did not significantly differ
from zero after January (see Alonso et al. 1994).

Fig. 1. Sketch map of the study area, showing the lake

(shaded), the two main crane roosts (asterisks), and the 10
zones in which the area was divided for the study.

In the case of sown grounds, food availability is not equal to 100% of the
cereal seeds sown, since the cost of digging up progressively deeper seeds
from the ground is a positively accelerated function of time spent digging at
one site. Some limit will exist beyond which it is uneconomical for a crane to
continue digging up seeds. This limit represents the food availability for the
cranes, and by definition must be higher than actual seed consumption by the
birds. To estimate food availability on sown grounds, we used midwinter
sown seed consumption data obtained during an early crop damage study
(values varied between 01% and 46%, see Alonso, Alonso & Veiga 1984),
and data on food intake rate of 12 cranes radiotracked for a total of over 150
whole days (values varied between 23% and 35%, unpublished data). Given
such figures, it seems not unrealistic to take 5% as the limit beyond which
cranes cannot economically dig up more seeds. Therefore, we considered that
on average 5% of the 180 kg seeds sown per hectare was actually available to
the cranes. Sprouted cereal fields were not used by foraging cranes. We
calculated overall food abundance at each zone of the study site, multiplying
the total area occupied by each type of ground by the mean density of seeds
laying on the surface in the case of stubble fields, and by 5% of the average

amount of cereal in sown fields (180 kg ha-1). The mean dry weights of seeds
in stubbles were 0027 g (cereal) and 0064 g (sunflower), calculated from
samples of 3000 seeds each.
For each of the 10 zones considered, we calculated an Expected Carrying
Capacity, ECC = FA / (DFCd ) where FA is the total food availability in each
zone, DFC is the daily food consumption of a crane (Alonso & Alonso 1992),
and d is the number of days remaining until the peak spring migration passage
of cranes in early March. It thus expresses how many birds a zone can
potentially sustain until the end of the wintering season. ECC provided a good
estimation of the number of wintering in a previous study at the same area
(see Alonso et al. 1994).
Each fortnight we established the distribution of the crane population in the
study area, averaging the numbers of birds seen on each foraging zone during
weekly surveys of the whole area, completed with additional information
gathered during daily individual radiotracking (see below). During these
surveys we recorded the location, type of substrate and number of individuals
of each foraging flock between 6 and 12 hours, waiting enough time for the
birds leaving the roost to reach even the farthest zones and stabilise as
foraging flocks. Within this morning period intake rate is usually the highest
of the day and the distribution of the cranes is not affected by satiation
(Alonso & Alonso 1992). Throughout the winter, we repeated the same crane
survey and used the same points to count the cranes. To account for possible
errors in our crane distribution maps, each fortnight we made a detailed
census of the crane population, counting all cranes entering the roost at
evening, from 3-4 observation points around the lake. There were no
significant differences between the sum of crane numbers seen on the 10
foraging zones during the day and our evening total population censuses.

DATA ANALYSIS

We studied the correspondence between density of cranes and density of food

abundance within fortnights in each zone with the Spearman correlation
coefficient. Because the distance from each zone to the lake was not the same
and could influence the decision where to forage, we calculated the partial
correlations between cranes and food, including the distance from the central
point of each zone to the centre of the lake as a third variable. The distance to
the centre of the lake was also correlated with average patch quality (within
zones) in previous studies, due to lower depletion of fields at farther distances
from the roost (Alonso et al. 1987; Alonso et al. 1995). This could also affect
the distribution of the birds.

Despite the benefits of using ECC as a measure of resource abundance,

fortnightly correlations could be done both with food density or ECC, since
number of days and daily intake are constant within each fortnight. Using the
food density expressed as kg ha-1 and crane density expressed as birds ha-1,
we controlled for differences in the size of each zone. To compare between
fortnights, we expressed food and bird density as proportions of the sum of
densities in 10 zones (pi = di / Sdi, i = 1,...,10). The ideal free distribution
predicts that at equilibrium, the proportion of individuals in a habitat or site
will be equal to the proportion of resources in that habitat or zone if the degree
of interference is high (Sutherland 1983; Pulliam & Caraco 1984).
We performed linear regressions with both the proportions of bird and food
densities. Because of the low sample size (10 zones), and the possible nonlinear relationship between both proportions (Kennedy & Gray 1993), we
applied nonparametric regression (Hollander & Wolfe 1973; Davison &
McCarthy 1988) to calculate the values of each slope and constant. We
applied the sign test (Siegel 1956) to test if slopes of regressions (b) were
different from zero, different from one or greater than one. We applied the
sign test to examine if the constants of regressions (a) were greater, equal or
lower than zero.

BEHAVIOUR OF MARKED INDIVIDUALS

Between late December and early February we radiotracked 12 individuals

(five in 1989-90 and seven in 1990-91; eight were adults, two immatures, i.e.
second- or third-year birds, according to their iris colour and pattern of wing
feather moult (Glutz, Bauer & Bezzel 1973; Prange 1989) and two first-year
birds already independent from their parents) for periods ranging between 4
and 19 days (mean = 11, total = 131 days). Throughout the text, when we use
the term 'adults' we include the two immature birds. We performed all
analyses related to competitive ability both including and excluding the two
juveniles. Since significance results were identical in both cases, we only
present those including the juveniles. Furthermore, inclusion of juvenile birds
is reasonable because the general foraging behaviour of independent first-year
birds is indistinguishable from that of immature or adult birds, and non-adult
birds represent an important fraction of the total crane population. We
captured these birds with rocket nets or oral tranquillisers. After weighing and
measuring wing, tarsus and bill, we colour-banded and radio-tagged them. We
were able to sex three adult birds as males (birds A, C and J, see Table 1) by
their unison-call behaviour (Glutz et al. 1973; Prange 1989). We began
studying their behaviour al least one week after having been released into the
wild. One observer followed a radiotagged bird by car continuously during the
day, watching it with a 60-90x Questar telescope from distances of 500-1000

m. Cranes foraged in flocks, at distances of 1-25 km from the roost. Individual

cranes spent the morning usually on only one of the 10 zones in which we
divided the study area, obtaining there on average over 70% of their daily
food ration. This percentage increased with the quality of the morning
foraging area. Total daily accumulated food intake was thus highly correlated
with morning intake ( r= 087, n = 131, P < 00001), and therefore we defined
the zone used in the morning as the main foraging zone for each day. After
drinking at midday or early afternoon, birds frequently used a different
foraging area, usually closer to the roost, during late afternoon.
At 30-60 minutes-intervals, we tape-recorded the behaviour of the marked
individual over 5-min intervals and that of 7-10 other randomly selected adult
flock members over 1-min intervals. Time spent in different activities was
measured to the nearest second. We measured the rate of food intake, counting
the number of characteristic swallowing movements of the feeding birds,
which almost exactly correspond to the number of seeds ingested, according
to our observations of free-living and captive cranes. We calculated the
accumulated total food intake for both, the main foraging period during
morning, and the whole day. We also estimated the net food intake for both
periods, subtracting from total food intake the estimated cost of flying to each
foraging zone, transformed to grams of food, using Pennycuick's (1989)
values of 054g consumed per km flying. Net daily food intake was used in all
subsequent tests. We assumed 78% assimilation efficiency (Castro, Stoyan &
Myers 1989). We excluded incomplete radiotracking days and days with
heavy rain or snowfall. We also calculated the difference in intake rate (seeds
min-1) between the marked crane and the mean of the 7-10 adult flock
members for the sample of foraging patches used. We used paired Students t
-tests for each individual to see if these intaake differences were greater than
zero. A patch was defined as one or a few adjacent farms where the flock
including our marked bird spent some time foraging between two flights
(respectively from, and to another patch, drinking site, or roost). An average
flock size was calculated for each individual using the sample of mean flock
sizes for each foraging patch. We also recorded if the marked individual was
dominant or subordinate at aggressive encounters with other flock members.
Most agonistic interactions in crane flocks were kleptoparasitic attacks to take
food discovered by the attacked bird. In a typical kleptoparasitic attack the
dominant crane is looking for sown cereal seeds without much success and
spots a nearby subdominat flock mate that has discovered a patch of seeds.
Since seeds are sown in parallel lines, cranes have learned to follow such lines
while undigging the grain (personal observation). This means that once a seed
or patch of seeds is discovered, the probability of finding further seeds in the
immediate vicinity is relatively high. Thus, if the discoverer is a subdominant
bird, it pays the dominant to displace it and take up the place of the
subdominant. The aggressor almost invariably increases its intake rate by
taking the seeds discovered by the victim. The displaced crane suffers then a

significant temporary decrease in intake rate. In a different paper we study the

kleptoparasitism in cranes in more detail (unpublished). Since our sample of
aggressive encounters included highly variable situations with respect to site,
flock size and daytime, we assumed that the percentage of successful
encounters reflected the dominance rank of the individual with respect to the
population.
Finally, each fortnight we ordered the 10 zones from the richest (quality = 10)
to the poorest (quality = 1), using the food density data obtained during the
food availability transects. We defined the average quality of the foraging
zones visited by a marked crane as the mean of these fortnightly quality
values.

Results
NUMBER OF CRANES AND CARRYING CAPACITY

Figure 2 compares the seasonal changes in mean numbers of foraging cranes

and the mean carrying capacity figures per zone in the study area. After an
initial crane arrival period in October-early November, peak numbers were
reached in both years in late November. During this autumn peak migratory
period, the mean number of cranes per zone slightly exceeded mean carrying
capacity figures, but later the mean number of cranes tended to equal the
carrying capacity until late January, when both figures were practically
identical. During the spring staging period the mean number of cranes per
zone remained below the carrying capacity.

Fig. 2. The mean number of foraging cranes per zone () during 1989-90 and 1990-91 in
logarithmic scale. In both years the number of cranes in winter progressively adjusted to the
expected carrying capacity (ECC, O). ECC was calculated dividing the food availability in
each zone by both the daily food consumption of a crane and the number of days remaining
until the peak spring migration passage of cranes in early March (see methods). The units of
ECC are also number of cranes per day. Vertical bars are one standard error.

The seasonal decreasing pattern in food availability in 1989-90 represents the

usual pattern in our study area in years with dry autumns. The high ECC
values in October-November are due to the large amount of waste seeds left
after harvest on cereal stubble fields and to the absence of a significant
germination of these seeds during dry autumns (see Alonso et al. 1984,
Alonso et al. 1994). In dry years the ECC decreases throughout the season
mainly due to food depletion by the cranes (Alonso et al. 1994). On the other
hand, in years with rainy autumns, such as in 1990, farmers begin ploughing
the land earlier, considerably reducing the food availability on stubble fields.
However, in spite of a lower food availability in autumn 1990 than in the
previous autumn, the midwinter food availability was slightly higher due to a
higher sowing activity between October and December that year.
In 1989-90, the cranes arriving early in the season (350 birds in early October)
foraged in zone 6, the closest zone to the roost (see Fig. 1), in spite of this
being the zone with the lowest food density. During the second half of
October cranes used zone 6 less intensively and began using zone 5, the
second closest to the roost and fourth lowest in food density. By early
November the cranes used zones 6, 5 and 4 (in decreasing order of bird
density), although these zones were respectively, the tenth, fifth and sixth in
order of food density. During the second year cranes used all zones each
fortnight. The correlation between density of cranes and density of food was
negative and not significant at the beginning of the winter season, and
changed to positive with increasing significance as the season progressed (Fig.
3). The highest significance values were reached in January in both years.
Later, the correlation coefficients tended to decrease again to zero. We have
no data for October and March of the second year, but the data in Fig. 3
suggest a similar pattern for both years. Partial correlations between density of
cranes and distance to the roost, once the effect of food density was accounted
for, were only significant in the second and third fortnights of the first year,
with more cranes close to the lake than far from it.

Fig. 3. Change
in the Spearman
correlation
coefficients
between food
density and
crane density of
the study area
during both
winters. The
arrows show the
critical values
of the
correlations (P
< 005, n = 10
zones for each
fortnight). The
line represents
the best fitted
function: y =
-1147 + 0514x
- 0035x2.

The fortnightly regressions between proportions of cranes and proportions of

food for the 10 zones are shown in Fig. 4. In 1989-90, the constants and slopes
were not significant in October and early November, but pooling all non-used
zones in a single value clear negative relationships emerged for these three
fortnights (a > 0 and b < 0, P < 005 in both parameters). From late November
to late February, as carrying capacity was reached (see Fig. 2), cranes
distributed themselves according to food resources, both using the 10 zones as
independent data or pooling the non-used zones in a single value. In March,
when cranes were already leaving the area, the slope of the regression was not
different from zero. Although the slope of the regression was not significantly
higher than one in 1989-90, in most December-February fortnights the values
of crane use of the preferred zones were above the diagonal. In 1990-91 the
pattern was similar, but the four fortnights with a slope significantly higher
than one indicate that during this winter season the distribution of cranes
clearly overmatched that of resources.

Fig. 4. Seasonal changes in

the relationship between
crane distribution and food
distribution in both winters.
Each point represents one
zone. Parametric
regressions are shown at
the same scale to enable
visual comparisons. Nonparametric slopes (b) and
constants (a) were tested
vs. the expected values of a
= 0, b = 0 and b = 1 at P <
005. Filled dots indicate
proportions greater than 05
(respectively 100 in
October, 070 in November
and 091 in January 198990).

COMPETITIVE ABILITY OF MARKED INDIVIDUALS

The weights and measurements of the twelve cranes studied are shown in
Table 1. The four birds with lowest weight were two juveniles and two
immatures. There was considerable individual variation in weight, the largest
bird weighing a 46% more than the smallest immature. Individual differences
in wing length and tarsus length were smaller, but still signifcant, with

extreme values differing respectively by a 24% and 26%. Weight and both
measurements were correlated (weight = -1948 + 126 wing, r2 = 062, P =
0002; weight = 1968 + 219 tarsus, r2 = 037, P = 003; wing = 1675 + 17
tarsus, r2 = 061, P = 0003).

The 12 individuals differed in their mean food intake, during both the main
foraging period in the morning, and across the whole day (Table 2). Individual
means for morning intake and whole day intake were correlated (r = 090, P <
0001). Thus, individuals ingesting lower average amounts of food during the
morning were unable to compensate this by feeding more in the afternoon.
Most individuals visited several zones during the winter, although each crane
seemed to prefer one or two (on average, each crane foraged on 62% of the
days in its most preferred zone during the morning, SE = 60, n = 12 birds).
The high correlation between the average quality of the foraging zone visited
by an individual and both, the mean morning intake and mean daily intake
(respectively, morning intake = 153 + 175 zone quality, r2 = 086, and daily
intake = 518 + 199 zone quality, r2 = 079, both P < 0001) suggests that the
selection of foraging zone had a major influence on daily food intake.

Individuals also differed in their degree of success in aggressive encounters

(mostly kleptoparasitic) with other flock members (Table 2). Dominance
ranks varied from highly dominant birds (adults C and J, both males) to
clearly subdominant birds (immature D and both juveniles, Y and Z). The
rank was also correlated with both, morning and total daily food intakes
(respectively, morning intake = 753 + 076 rank, r2 = 052, P = 0008; daily
intake = 1220 + 082 rank; r2 = 043, P = 002). Dominant birds tended to
select foraging zones with higher food density (zone quality = 398 + 003
rank; r2 = 031, P = 005).
Thus, two variables could have potentially determined the interindividual
differences in food intake, namely, foraging zone quality and dominance rank.
We performed a stepwise multiple regression analysis of both variables on
morning food intake. The resulting model was highly significant (F2,9 = 572,
P < 0001, 93% of the variance explained), with zone quality as the first
variable (P < 0001) and rank as the second (P = 0024). Identical result was
obtained by a two-way ANOVA of all daily data grouped in two ranks classes
and two zone quality classes. The model included significant effects of both

factors (P < 0001 for zone quality and P = 0021 for dominance rank). The
interaction factor was also significant (P = 0016), suggesting that the relative
intake rates of individuals with different competitive abilities were nonconstant across zones with different food densities (see Fig. 5).

Fig. 5. Changes in mean food intake during the morning

(mean foraging period of the day) with zone quality and
dominance rank. Dominants were those individuals with a
percent success in aggressive encounters of > 50% (see Table
2). Good zones were those ranking > 5 in food density during
the radiotracking period (late December-early February).
Vertical bars are one standard error. ANOVA showed
significant effects for zone quality (F1,135 = 1150, P = 001)
and rank (F1,135 = 548, P = 002); the interaction term was
also significant (F1,135 = 597, P = 001).

Dominant cranes tended to feed at higher rates than the average rate of the
flock, while subordinates did not. Although only two birds showed clear
significant differences in intake rate with respect to the flock average (Table
2, adults C and G, the latter with marginal significance), most dominant
cranes (five of a total of seven being successful in > 50% of encounters) had a
higher mean intake rate than their flock members, while most subordinate
cranes (three of a total of five being successful in < 50% of encounters) had a
lower intake rate than their flock members (see Table 2). Moreover, there was
a positive significant correlation between the dominance rank and the intake
rate difference with respect to the average intake rate in the flock (Fig. 6).

Fig. 6. Correlation
between the
dominance rank (%
success in
aggressive
encounters) and the
difference in intake
rate of each marked
crane with respect
to the mean flock
intake rate (see
Table 1). The
correlation was
also significant
excluding the two
juveniles, Y and Z
(r = 076, P = 001).
The regression
equation is y =
-447 + 009x.

Finally, dominance rank was positively correlated with the size of the crane
(rank = -2053 + 0046 weight; r2 = 056, P = 0005, see Fig. 7; rank = -2737
+ 054 wing length, r2 = 030, P = 006; the correlation with tarsus length was
positive, r = 026, but non-significant).

Fig. 7. Correlation
between the
dominance rank (%
success in
aggressive
encounters) and the
weight of the crane.
The correlation was
also significant
excluding the two
juveniles, Y and Z
(r = 067, P = 003).

Discussion

Our results agreed with most assumptions and predictions of the interference
phenotype-limited model proposed by Parker & Sutherland (1986). First, there
were clear individual differences in competitive ability, as shown by the
differences in success in aggressive encounters for food. These rank
differences were probably at least in part due to age and size differences
between individuals. Secondly, the relative pay-offs of different phenotypes
changed across different food densities. The data in Fig. 5 suggest that
subordinate birds probably could not increase their intake rate at the richest
zones as much as dominants. Additional costs of flying to these zones, which
were the farthest from the roost, made them umprofitable for many low rank
individuals. Thus, it may pay juveniles, subadults or adults with lower
competitive ability to stay in poorer foraging zones, if the benefit of moving to
the richest zones is comparatively low to make the effort profitable. Thirdly,
the more competitive cranes tended to be found in those foraging zones with
highest food densities. Fourth, average intake rates were highest in the richest
zones (see also Alonso et al. 1995). Finally, in a previous study we tested a
further assumption of the interference model, namely that pay-offs of
individuals decreased with increasing competitor density. Although intake rate
slightly increased with number of birds for a range of small flock sizes (up to
c. 50 birds) due to a reduction in individual vigilance time (Alonso & Alonso
1991), in very large flocks it was lower for equal seed densities (unpublished
data). The intake rate decrease was still significant even without considering
obvious kleptoparasitic attacks, which means that the decrease in intake rate
was mostly due to subtle threats, evasive actions and distractions
imperceptible for an observer.
Our results strongly suggest that (i) the quality of the foraging zone and (ii)
the dominance rank of the individual, were important in determining each
individual's daily food intake. The dependence of intake rate on food density
at a site was not surprising. We have shown that this correlation is strongly
positive, either considering a flock feeding in a small food patch (Alonso &
Alonso 1991, Alonso et al. 1995) or a number of flocks in a larger foraging
zone (present paper). But the influence of dominance rank on food intake (see
Fig. 6) was not so intuitive, particularly in a flock-foraging species. The
positive correlation between rank and difference in intake rate with respect to
flock members suggests that dominant individuals can benefit from
kleptoparasitic attacks to subordinate flock members, slightly increasing their
intake rate above the average of the flock. Such increase could theoretically
range from 40g less to 395g more than the average daily food intake,
respectively in the most subordinate and most dominant cranes, assuming
dominance status would be maintained throughout a whole day (see regression
line in Fig. 6: from -448 to +442 seeds min-1; given 003g per seed and 62%
time feeding during 8 hours daytime, see Alonso & Alonso 1992). Such intake
differences represent a significant proportion of the daily food intake
(respectively -230% and +227% of the 174g mean daily food intake of the 12

cranes studied), suggesting that this behaviour may substantially increase the
fitness of dominants.
Some previous results on behaviour and distribution of cranes in our study
area provide further support to the phenotype-limited distribution. The poorest
zones in our study area have a significantly higher percentage of crane
families than the better zones (Alonso et al. 1984, 1987), which may be
explained by the lower competitive ability of juveniles. Elsewhere (Alonso &
Alonso 1993) we show that juvenile cranes (i) had a lower intake rate in
flocks than in isolated families, probably due to interference with other flock
members, (ii) increase throughout their first winter their foraging efficiency
and success in aggressive encounters, and (iii) were involved in more
kleptoparasitic encounters with increasing flock size.
In spite of the general agreement of our results with the interference
phenotype-limited model, the distribution of crane phenotypes in our study
area was far from being perfectly truncated. As already stated, the twelve
individuals studied visited several of the ten zones defined. Also, there were
always some juveniles in all zones and even in any flock within each zone.
Imperfect phenotype-truncated distributions have also been observed in two
other field studies, with herring gulls Larus argentatus (Monaghan 1980) and
oystercatchers Haematopus ostralegus (Goss-Custard et al. 1984). In the first
study, gulls feeding at two areas of a refuse tip differing in quality represented
a highly competitive situation more likely than our case to fit these
predictions. This study is probably one of the best examples of truncated
phenotype distributions, but it was based more on proportions of each sex and
age class than on detailed behavioural observations (dominance and intake
rate) of individually marked birds. The study by Goss-Custard et al. (1984) is
a detailed account of individual behaviour and population distribution of
oystercatchers at two mussel Mytilus edulis beds. The aggression rates of
oystercatchers were only slightly higher than those of our cranes, but the
biomass stolen per attack represented a much higher proportion of the daily
food ration. Thus, individual dominance differences were probably more
important in oystercatchers than in cranes as a strategy to increase fitness. As
in our study, Goss-Custard et al. recognised the difficulty of separating
aggressiveness, experience in feeding and bird quality, but also suggested, as
we do, that intake rate was at least in part causally related to aggressiveness
(see also Ens & Goss-Custard 1984). Parker & Sutherland (1986)
acknowledged that perfect truncation is probably never met in the wild, and
proposed several possible explanations: disrupting by some naive individuals,
temporal changes in the relative qualities of patches, kleptoparasitism leading
to continuous switching of phenotypes between patches, and heterogeneous
food distribution within patches, which may cause a phenotype truncation
within each patch (see also Milinski & Parker 1991). Although all of these
processes may probably be relevant in our case, the last agrees particularly

with the observed tendency of crane families or independent juveniles to

occupy the periphery more often than the centre of large flocks foraging on
rich patches within high quality zones, or even to forage isolated to avoid
attacks to their offspring, but still close to these large flocks in the same zone.
Eventually, as their intake rate decreases, they may leave and go to a different
lower-quality zone (Alonso et al. 1984, 1987, Alonso & Alonso 1993).
Therefore, even for some dominant cranes, if they have dependent juveniles, it
may pay to occupy lower-quality patches and zones.
As for matching between bird and food densities, as long as quality
differences between zones are large and/or interference is low, a higher
number of birds at the richest zones may be expected, since it may pay many
of the lower competitive individuals to stay on the richest zones as long as
their intake rate there is higher than on the worst (Sutherland & Parker 1985,
1992; Parker & Sutherland 1986). Our data indeed showed the predicted
overuse of the richest foraging zones, contrasting with the pattern of underuse
observed in most ideal free distribution studies (review in Kennedy & Gray
1993).
This overuse of the richest sites is also predicted, and probably enhanced, in
gregarious species, according to Pulliam & Caraco (1984). These authors
applied Fretwell's (1972) 'Allee principle' which posits that patch quality
increases, up to a point, as density of predators increases, and thereafter
declines. The food intake rate of cranes indeed follows a curvilinear
relationship with respect to flock size (see above). Combining their flock
model with IFD theory, Pulliam & Caraco (1984) concluded that equilibrium
group sizes would usually be larger than optimal size, and frequently of the
maximum size allowed by patch quality. They suggested that their model
could be extended to describe the distribution of flock-foragers among zones
of different qualities, with an equilibrium distribution showing overuse of the
richest sites.
An alternative model which we also considered was the depletion model with
no interference developed by Sutherland & Anderson (1993) and Sutherland
& Allport (1994). This model also predicts an overuse of the richest zones.
However, our data did not fulfil the assumption of negligible interference. In
addition, although the seasonal crane distribution pattern globally agreed with
the assumption that predators gradually exploit an increasing number of zones
as they deplete them to a critical value, there were slight differences between
model and field data. First, cranes did not deplete all foraging zones exactly to
the same prey density, leaving more resources unexploited in the richer zones
(see Alonso et al. 1995). Secondly, cranes did not always exploit the richest
zone first. Our results indeed show, in apparent agreement with the depletion
model, that matching between cranes' distribution and food distribution was a
gradual process beginning when the number of cranes exceeded the carrying

capacity expected for the rest of the winter. Early and late in the season,
respectively before mass arrivals of migrating birds and prior to spring
departures, the distribution of birds did not correlate with food distribution.
The limiting effect of carrying capacity was probably the factor determining
the observed increase in the number of zones used, and the consequent
increase in the matching between cranes' and food distributions as the season
progressed. However, although cranes present in the area early in the first
winter season could have maximized food intake by foraging on the richest
zones, as predicted by the depletion model, they preferred those adjacent to
the roost, probably because they saved travel costs and still had enough food.
This interpretation is supported fact that the partial correlation between crane
numbers and distance to the roost in the foraging zones, once food density
differences were excluded, was only significant in late October and early
November of the first year, being not relevant later in that season, nor in the
second one. In contrast to the first year, in the second year the low initial food
availability forced the cranes to use all zones from earlier in the season. But
the higher midwinter food availability enabled a higher number of birds to
spend the winter in the area.
In spite of these interannual differences, the process of adjustment of bird
numbers to food availability was similar in both years. Many birds were
forced to leave the area between late November and January, when the
population stabilised at a level closely corresponding to that of the carrying
capacity (Alonso et al. 1994). With respect to the end of the season, matching
between cranes' and food distributions tended to disappear again, probably
due to the increase in food availability above the needs of the population. In
addition, previous results suggested that during spring migration food
availability was not a primary factor determining crane numbers and
distribution (Alonso et al. 1990). We conclude that depletion is perhaps more
important in limiting the carrying capacity of the whole area (Alonso et al.
1994), while interference mainly influences distribution of phenotypes
between zones (this study).
The problem with current depletion models, like that of the earliest IFD
models, is that they do not consider individual differences, and this is an
unrealistic simplification of natural situations. We think that in our system
interference and depletion were probably both important. In fact, interference
may be caused or enhanced by depletion. Since both processes are not
mutually exclusive (see Goss-Custard 1980; Sutherland & Goss-Custard
1991), we do not find it justified to distinguish distribution models based on
one or the other. Rather, we agree with Sutherland & Allport's (1994)
statement that a future challenge is to explore these two approaches including
individual differences and apply them to field data (see e.g. Sutherland &
Dolamn 1994).

Acknowledgements
We are grateful to C. Bernstein, A. Kacelnik, W.J. Sutherland, and two
anonymous referees for helpful comments on earlier versions of the
manuscript. This is a contribution to Project PB91-0081 of the Direccin
General de Investigacin Cientfica y Tcnica.

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