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CSIRO PUBLISHING
www.publish.csiro.au/journals/emu Emu, 2009, 109, 260–264

Red-backed Hawks supply food to scavenging

Chimango Caracaras

Alejandro V. Baladrón A,B,D, Laura M. Biondi A,B, María S. BóA, Ana I. Malizia A
and Marc J. Bechard C
A
Laboratorio de Vertebrados, Departamento de Biología, Universidad Nacional de Mar del Plata, Funes 3250,
Mar del Plata, B7602AYJ, Argentina.
B
Consejo Nacional de Investigaciones Científicas y Técnicas – CONICET, Rivadavia 1917, Buenos Aires,
C1033AAJ, Argentina.
C
Raptor Research Center, Department of Biology, Boise State University, 1910 University Drive, Boise,
ID 83725-1515, USA.
D
Corresponding author. Email: abaladro@mdp.edu.ar

Abstract. Positive interactions between species of terrestrial vertebrates occur frequently and it is common among birds.
However, such interactions have generally been overlooked in raptor field studies. We analysed interactions between
Red-backed Hawks (Buteo polyosoma) and Chimango Caracaras (Milvago chimango) in order to determine the
characteristics of the association between these two South American raptors. We observed 171 bouts of activity, 40
when Red-backed Hawks were feeding and 131 when the Hawks were undertaking other activities. Chimango Caracaras
were more often present when Red-backed Hawks were feeding (81%) than during other activities of Red-backed Hawks
(69%), but they attacked Red-backed Hawks more frequently during other activities of the Hawks (66%) than when the
Hawks were feeding (43%). In addition, the rate of attacks by Chimango Caracaras was higher during other activities of the
Hawks (mean = 2.21 attacks per Caracara) than when the Hawks were feeding (mean = 0.68 attacks per Caracara). Duration
of feeding activities was not affected significantly by the presence of Chimango Caracaras or by attacks by the Caracaras.
During 16 of 40 feeding bouts of Hawks, an average of 1.81 ! 0.23 (s.e.) Caracaras fed on prey discarded by the Hawks. Thus,
Red-backed Hawks indirectly supplied food to Chimango Caracaras, which, in turn, reduced their agonistic behaviour
towards Hawks. Our findings represent one of few cases reported about facilitative associations among raptors, but it seems
likely that it represents a common strategy among these birds.

Introduction higher rate of energy intake in heterospecific associations than

Interactions among species have traditionally been categorised
depending upon the effect – positive, negative, or neutral – of
two or more species on each other (Dickman 1992). In recent
times, positive or facilitative interactions have begun to be
considered as one of the most important organising forces in
natural communities (Stachowicz 2001; Oliver et al. 2009).
These are interactions between organisms that benefit at least
one of the participants and cause harm to neither, including
commensal and mutualistic associations (Bruno et al. 2003).
Likewise, it has been observed that the strength of the benefits of
positive interactions become increasingly important to species
persistence with increasing levels of stress (Stachowicz 2001;
Eränen and Kozlov 2008).
Commensal and mutualistic interactions occur frequently
between species of terrestrial vertebrates (Dickman 1992).
They are common among birds, such as waterbirds, which
often associate in mixed-species flocks (Burger 1988), but it
has been rarely reported among raptors with the exception of
succession in scavenger’s guilds (Travaini et al. 1998; Selva and
Fortuna 2007). In both groups, individual birds often achieve a
! Royal Australasian Ornithologists Union 2009
when foraging alone or with conspecifics (Rand 1954; Dickman
1992). Such interactions enable a species to exploit a greater
portion of available resources, both by increasing the range of
food types in the diet and by exploiting leftover food obtained by
other species, thus increasing the utilisation of the fundamental
niche space (Bruno et al. 2003).
During a study of the trophic ecology of the Red-backed Hawk
(Buteo polyosoma; also known as the Variable Hawk) in the
pampas of Argentina, several cases of agonistic interactions
with Chimango Caracaras (Milvago chimango) were observed
(see Baladrón et al. 2006). Chimango Caracaras frequently
attacked Red-backed Hawks, probably as an innate reaction of
a territorial bird of prey to repel other larger, raptor species
(Newton 1979; Gaibani and Csermely 2007). These attacks
often ended with Caracaras feeding on prey remains discarded
or left behind by the hawks. This observation might be indicative
of a possible secondary function of the agonistic behaviour
showed by Chimango Caracaras toward Red-backed Hawks.
In the present study, we explore the behavioural characteristics
of the agonistic relationship between these two raptors, taking into
10.1071/MU09024 0158-4197/09/030260
Hawks supply food to caracaras
account how each interacting species is affected, and the costs
and benefits of this association. In particular, we expected that
if it responds to a facilitative association: (1) the behavioural
patterns of Red-backed Hawks should not be affected by the
interaction with Chimango Caracaras; (2) Red-backed Hawks
manipulating or eating prey should attract larger numbers of
Chimango Caracaras than Hawks undertaking other activities;
and (3) Chimango Caracaras should reduce the intensity of their
aggressiveness when Red-backed Hawks are feeding compared
with Hawks performing other activities.
Materials and methods
Study area
We conducted our study at Mar de Cobo, south-eastern
Buenos Aires Province, Argentina (37"580 S, 57"340 W). The
landscape is dominated by temperate grasslands, in which the
original gramineous vegetation has been highly modified by
agricultural activities (Bilenca and Miñarro 2004). The study
area (~60 ha) comprises a small coastal village (<10 m above sea
level), which is surrounded by wild areas (marshes, coastal
dunes, natural halophytic grasslands and native forests) and
agroecosystems (pastures, cultivated fields and cultivated
forests) (Stutz 2001). The dominant species of rodent of the
dune fringe is the Tuco-tuco (Ctenomys talarum), a subterranean
rodent highly abundant along coastal grasslands where it digs
its burrows (mean density = 30 individuals ha–1; Malizia et al.
2001).
Study species
The Red-backed Hawk is a medium-sized raptor (mean
weight = 950 g; Thiollay 1994) endemic to South America. In
Argentina, it is resident of a broad area along the Los Andes
mountain range to sea level in Patagonia (Canevari et al. 1991).
During winter, Red-backed Hawks migrate to the lowland
plains of northern and central Argentina (Narosky and Di
Giácomo 1993). In the study area, they are often seen perched
on tall utility poles near dune fringes as they search for their
main prey, the Tuco-tuco (Baladrón et al. 2006). Tuco-tucos
comprise >55% of the biomass in their diet and Hawks are
frequently seen carrying or feeding on this rodent. Usually,
Hawks discard the head and viscera when feeding on Tuco-
tucos (Baladrón et al. 2006).
The smaller Chimango Caracara (mean weight = 300 g; White
et al. 1994) also endemic to South America and it is considered
the most abundant raptor in central Argentina (Canevari et al.
1991). It shows an extraordinary behavioural plasticity associated
with its opportunistic mode of feeding and broadly based diet.
Although insects are the main prey of Chimango Caracaras
in the study area (frequency > 95%), it is very common to
see them feeding on garbage or cattle carrion (Cabezas and
Schlatter 1987; Biondi et al. 2005). Chimango Caracaras are
both solitary and gregarious, and can be seen in pairs defending
territories throughout the year, yet congregating in large mixed-
age feeding groups when a source of food is discovered, as well
as for resting and breeding (Fraga and Salvador 1986; Ferguson-
Lees and Christie 2001; Biondi et al. 2005). In addition,
interspecific and intraspecific kleptoparasitism seems to be a
common foraging strategy for this raptor (Copello and Favero
Emu 261
2001; Ferguson-Lees and Christie 2001), and it also utilises
resources generated by human activities, such as agricultural
practices, fishing and camping (Cabezas and Schlatter 1987;
Biondi et al. 2005).
Wintering Red-backed Hawks arrive in the study area in small
groups (up to five individuals) in mid-April and leave in late
August. Chimango Caracaras are permanent residents of the study
area, breeding in the surrounding grasslands and woodlands
(Isacch et al. 2001).
Behavioural sampling
We quantified agonistic interactions between Chimango
Caracaras and Red-backed Hawks during the non-breeding
period (April to August) of 2002–04. We employed a
continuous recording method (Martin and Bateson 1993), by
means of direct observation with 8 # 50 binoculars. Red-backed
Hawks used the study area as a foraging site each day. We
searched for individuals from their arrival in the morning
until they left in the afternoon and recorded observations of
behaviour from sighting until they were lost from sight.
Each day, the number of individuals of each species, their
activities, and the interactions between them (approaches and
attacks) were registered using a manual voice recorder (Olympus
WS-320M Digital Voice Recorder, Melville, NY, USA) and data
were transcribed later using a stopwatch.
We divided bouts of activity into two types: feeding activities
(FED) and other activities (OTH). We considered that a FED
started when a Hawk captured a prey item and finished when it
left after feeding. OTH were mainly activities related to prey
searching (passive searching from a perch, changes of perches,
and short movements between foraging patches) and, to a lesser
extent, comfort and flight activities. OTH bouts started when a
Red-backed Hawk was found, and finished when the Hawk
was lost from sight, or when a FED bout began. During each
OTH bout, we recorded the number of Chimango Caracaras that
approached Red-backed Hawks, as well as the number of attacks
by Caracaras. During each FED bout, we recorded the same
variables as in OTH, but also quantified the number of Caracaras
that obtained food by eating scraps of prey discarded by Hawks.
We calculated the rate of attacks for each interaction as the total
number of attacks divided by the number of Chimango Caracaras
(rate of attacks per Caracara).
Statistical analysis
For all analyses we used R software, Version 2.7.0
(R Development Core Team 2008). We used generalised linear
mixed models (GLMM) and binomial tests to assess the
differences in number of Chimango Caracaras and number of
attacks, and the rate of attacks per Caracara (all responses
variables) between types of activity (explanatory variable).
Since the number of Caracaras and number of attacks were
count data, errors followed a Poisson distribution with a log-
link function. In the case of rate of attacks per Caracara, we
log-transformed the variable and used a Gaussian distribution
family with an identity link function, including the individual
identity of Hawks (ID) as a random factor (Crawley 2007). We
used Spearman correlation to explore the relation between the
number of Chimango Caracaras and the number of attacks during
262 Emu A. V. Baladrón et al.

each activity type, as well as to evaluate the relationship between Table 1. Duration of feeding (FED) and other activities (OTH) by
rate of attacks per Caracara and the duration of each type of Red-backed Hawks and characteristics of interactions between Hawks
activity. In addition, we employed GLMM to compare duration of and Chimango Caracaras during these activities
Results are given as means with standard error in parentheses
FED in relation to the presence of Chimango Caracaras and
occurrence of attacks, using three categories: (1) Red-backed Event type FED (n = 40) OTH (n = 131)
Hawks feeding alone (without Chimango Caracaras present);
(2) Hawks feeding in the presence of non-attacking Caracaras; Mean duration (min) 18.35 (1.73) 33.84 (2.63)
and (3) Hawks feeding in the presence of Caracaras that were Mean number of Caracaras 2.14 (0.38) 1.61 (0.15)
attacking. In this case, duration of FED was the response Mean number of attacks 1.14 (0.35) 3.31 (0.58)
Rate of attacks per Caracara 0.658 (0.21) 2.21 (0.39)
variable and the categories of presence of Chimango Caracaras
and occurrence of attacks the explanatory variables (with three
levels). We log-transformed FED duration data and used a Table 2. Generalised linear mixed models (GLMM) comparing
Gaussian distribution family with an identity link function, differences in (a) number of Caracaras and (b) number of attacks
including the individual identity (ID) as a random factor. between the two types of activity bouts (feeding bouts and other
When a Gaussian family distribution was used, the activity bouts)
significance of fixed effects was tested with a likelihood ratio Individual identity (ID) was used as the random factor
test (Crawley 2007). Data are presented as mean ! s.e.
Fixed effect: Random effect: ID
type of activity
Results
(a) Number of Caracaras
The aggressive interactions of Chimango Caracaras followed a Estimate –0.32 Variance 0.24
consistent pattern during activities other than feeding (OTH). s.e. 0.14 s.d. 0.48
Aggressive interactions began when Caracaras flew past a z value –2.37
perched Hawk after which they typically returned in a fast P 0.017
swoop on the Hawk with their talons extended. After the (b) Number of attacks
swoop, they usually flew past a short distance but quickly Estimate 0.87 Variance 1.38
returned to swoop at the Hawk again. In contrast, when Red- s.e. 0.16 s.d. 1.17
backed Hawks were feeding (FED), the Chimango Caracaras z value –5.46
showed a different behavioural pattern. Initially, they made only P <0.001
one or a few swoops, before quickly perching near the Hawk.
After this, they would vocalise, attracting other Caracaras. When
other Caracaras arrived, they also performed one or a few swoops, significantly higher during OTH (GLMM: t = 2.64, P = 0.008).
and then perched. Often they attacked and displaced one another Finally, the duration of each activity type was not significantly
in order to have better access to a perch. Sometimes the Caracaras related with the rate of attacks per Caracara (OTH: rs = 0.19,
approached the Hawk very closely and perched a few centimetres P = 0.07; FED: rs = 0.17, P = 0.33).
away on the same post on which the Hawk was perched. Although When Red-backed Hawks fed on Tuco-tucos at a perch, they
the Red-backed Hawks were aware of the Chimango Caracaras, usually discarded both the head and viscera of the prey. After the
they were mostly tolerant of them. Hawk dropped these remains to the ground, usually a single
We recorded 171 bouts of activity by Red-backed Hawks, 131 Chimango Caracara (generally the nearest individual) flew down,
of which were OTH and 40 were FED. The percentage of events took the food from the ground, and fed on it at a nearby perch.
where Chimango Caracaras approached the Red-backed Hawks After feeding, the Red-backed Hawk stayed briefly on the feeding
was higher in FED (81%) than during OTH (69%), though post and, as soon as the Hawk departed, the Caracaras flew to the
the difference was not significant (binomial test, c21 = 2.24, feeding perch where they ate small pieces of food or fluids that
P = 0.133). Conversely, the percentage of events in which were left behind. During 48% of FED bouts, a total of 29
Chimango Caracaras attacked Red-backed Hawks, as a Chimango Caracaras fed on prey remains and scraps discarded
proportion of the total number of FED and OTH in which by the Hawks (1.81 ! 0.23 Chimango Caracaras per event). In
Caracaras participated, was significantly higher during OTH most cases, only one (n = 7 cases) or two (n = 4) Caracaras
(66%) than during FED (43%; binomial test, c21 = 3.96, obtained the most important prey remains (head or viscera),
P = 0.04). In addition, the number of Chimango Caracaras that whereas up to four Caracara fed on scraps left behind on the
approached Red-backed Hawks was correlated significantly with feeding post.
the number of attacks during OTH (rs = 0.26, P = 0.01) but was We found that the number of Caracaras feeding showed a
not significant correlated during FED (rs = 0.27, P = 0.12). significant relationship with the maximum number of Caracaras
The number of Chimango Caracaras that approached a Red- present (rs = 0.45, P = 0.008), but it was not significantly related
backed Hawk was higher during FED than during OTH, but the with rate of attacks per Caracara (rs = 0.21, P = 0.26). In addition,
number of attacks performed by Caracaras on Hawks was lower the comparison of the duration of FED bouts among the three
during FED than during OTH (Table 1). The GLMM showed a categories (without Caracaras, with Caracaras and without
significant difference in the number of Caracaras and attacks attacks, and with Caracaras and attacks) did not show
between both activity types (Table 2). Similarly, the rate of attacks significant differences (GLMM: tWITHOUT CHIMANGOS = 0.08,
per Caracara also differed between activity types, being tWITH ATTACKING CHIMANGOS = 1.53, P = 0.257). Thus, the
Hawks supply food to caracaras
duration of feeding activity by the Hawks appeared was not to be
affected by the presence of Caracaras or of Caracaras performing
attacks.
Discussion
Commensal and mutualistic associations among terrestrial
vertebrates are clearly dynamic, and form and dissolve under
different conditions of predation risk, resource levels or
competition (Dickman 1992). In particular, top predators can
have positive effects on smaller species by increasing the
availability of carrion (Stahler et al. 2002; Wilmers et al. 2003;
Daleo et al. 2005; Selva and Fortuna 2007) whereas negative
components for facilitators are minimised when scavenger
species obtain by-product benefits from them (Connor 1996). In
this work, we found that feeding Red-backed Hawks often
supplied food indirectly to Chimango Caracaras by discarding
prey remains on which Caracaras scavenged. Correspondingly,
Caracaras responded differently to the behaviour of Hawks,
reducing aggressiveness and agonistic behaviour depending on
the chance of gaining food benefits from Hawks.
We postulated that if this interaction responds to a facilitative
association, patterns of feeding behaviour of Red-backed
Hawks should not be affected by the interaction with
Chimango Caracaras. Our results indicate that the costs for the
Red-backed Hawks were probably low, because attacks by
Chimango Caracaras did not influence the Hawks’ feeding
behaviour, and the duration of feeding events was not
affected by attacks by Caracaras. If these costs are negligible
to the Red-backed Hawks, this relationship is an example of
facilitation, or positive interaction. Nevertheless, if the costs
for the Red-backed Hawks were significant, it would represent
a case of negative interaction, likely positioned in some
intermediate point between facilitation and contramensalism,
and probably linked to kleptoparasitism (Ellis et al. 1993;
Stahler et al. 2002).
We found that when Red-backed Hawks were feeding,
Chimango Caracaras congregated in higher numbers and
they showed a reduction in aggressiveness, in comparison with
non-feeding activities, thereby reinforcing to some extent
the facilitative hypothesis (our original second and third
predictions; see Introduction). Food is often considered an
important source of aggression among raptors (Gaibani and
Csermely 2007), but we found that Chimango Caracaras were
actually more aggressive when Hawks were doing activities
other than feeding. We observed that Caracaras performed
few attacks and rapidly perched near feeding Hawks, often
vocalising and attracting other Caracaras. Newly arriving
Chimango Caracaras approached the Hawks but did not attack
them, reducing the overall rate of attacks per Caracara. The
different behaviour when Hawks are feeding might indicate
that Caracaras benefit by not being aggressive, which could
ultimately be associated with the aim of obtaining food left by
the Hawks.
The more passive behaviour of Chimango Caracaras toward
Red-backed Hawks could be explained through a cost–benefit
trade-off. The discarding of head and viscera is a regular practice
when these Hawks feed on Tuco-tucos (Baladrón et al. 2006)
and this high-quality food source is usually unavailable to
Emu 263
Chimango Caracaras in any other way than by scavenging or
kleptoparasitism owing to size constraints. Theft of prey seems to
be a common strategy for Chimango Caracaras in the study area
(Copello and Favero 2001). However, we did not observe any
case of direct pilfering on the Red-backed Hawks during our
study. The strategy of waiting for food to be discarded would be
less costly to Chimango Caracaras as Caracaras are three times
smaller than Red-backed Hawks and a direct pilfering attempt
could result in injuries to the Caracaras. Also, not attempting to
steal prey may have avoided disturbance to the Hawks that could
have resulted in the Hawk taking flight and carrying the food
away. On the other hand, owing to their small size, Caracaras
might fulfil a high percentage of their dietary energetic needs just
with the consumption of Tuco-tuco remains discarded by the
Hawks (Aprile and Bertonatti 1996). In this sense, our results
showed that this passive strategy seems to be profitable since, in
almost half of feeding activities by Hawks, Caracaras gained
access to the food remains left by the Hawks.
We suggest that Chimango Caracaras could strike a balance
between directly stealing food (kleptoparasitism) and waiting for
discarded prey remains (facilitation), choosing the latter strategy
mainly owing to the high cost of stealing from a larger species and
the low probability of success of such behaviour. In view of the
habitat overlap of these two raptors over most of their range, it can
be argued that this relationship between Chimango Caracaras
and Red-backed Hawks is not local. Moreover, as combined with
their opportunistic and innovative behaviour (Biondi et al. 2008),
facilitative associations may be a common foraging strategy for
Caracaras, not only with Red-backed Hawks and other larger
raptors, but also with other top predators, such as scavengers and
seabirds with which Caracaras sometimes associate (Travaini
et al. 1998; Copello and Favero 2001).
Frequently, small-scale interactions, such as those we describe
here, are nested within larger scale positive interactions, which
may have a strong influence on the organisation of terrestrial
communities (Dickman 1992; Stachowicz 2001). Similar
strategies of opportunistic species that obtain by-product
benefits has often been observed among other bird groups
(Brockmann and Barnard 1979), such as corvids (Stahler et al.
2002; Wilmers et al. 2003) and seabirds (Burger 1988). Our
findings represent one of the few reported cases of facilitative
associations among raptors, but it seems likely that it represents a
common strategy among this group of birds.
Acknowledgements
We thank C. Godoy and L. Scenna for field assistance. J. P. Isacch, A. Vassallo
and three anonymous referees provided helpful and constructive comments on
an earlier version of the manuscript. Funding for this study was provided
through a grant from the Universidad Nacional de Mar del Plata (15-E238) and
CONICET fellowships to A. Baladrón and L. Biondi.
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Manuscript received 21 March 2009, accepted 3 July 2009