Parental Care of the Endangered Chaco Eagle (Buteogallus

coronatus) in Central Argentina

Author(s): Maximiliano A. Galmes José H. Sarasola and Juan Manuel Grande
F. Hernán Vargas
Source: Journal of Raptor Research, 52(3):316-325.
Published By: The Raptor Research Foundation
https://doi.org/10.3356/JRR-16-82.1
URL: http://www.bioone.org/doi/full/10.3356/JRR-16-82.1

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J. Raptor Res. 52(3):316–325
Ó 2018 The Raptor Research Foundation, Inc.

PARENTAL CARE OF THE ENDANGERED CHACO EAGLE

(BUTEOGALLUS CORONATUS) IN CENTRAL ARGENTINA
MAXIMILIANO A. GALMES1
Centro para el Estudio y Conservación de las Aves Rapaces en Argentina (CECARA), Facultad de Ciencias Exactas
y Naturales, Universidad Nacional de La Pampa, Avda. Uruguay 151, 6300 Santa Rosa, La Pampa, Argentina
and
The Peregrine Fund, 5668 West Flying Hawk Lane, Boise, ID 83709 USA

JOSÉ H. SARASOLA AND JUAN MANUEL GRANDE

Centro para el Estudio y Conservación de las Aves Rapaces en Argentina (CECARA), Facultad de Ciencias Exactas
y Naturales, Universidad Nacional de La Pampa, Avda. Uruguay 151, 6300 Santa Rosa, La Pampa, Argentina
and
Instituto de las Ciencias Ambientales y de la Tierra de La Pampa (INCITAP), Universidad Nacional de La Pampa–
CONICET, Avda. Uruguay 151, 6300 Santa Rosa, La Pampa, Argentina

F. HERNÁN VARGAS
The Peregrine Fund, 5668 West Flying Hawk Lane, Boise, ID 83709 USA

ABSTRACT.—Sexual differences in parental care investment may affect individual survival or performance,
and also demography. We investigated parental-care behavior of the endangered Chaco Eagle (Buteogallus
coronatus) to evaluate implications for population conservation in semiarid habitats of central Argentina.
From 2004 to 2012, we monitored three breeding attempts using video recording (275 hr) during the
incubation period, and nine pairs during the nestling-rearing period by means of video recording (1087 hr)
and focal observations at nests (232 hr). Chaco Eagles showed a strong division of parental care by sex.
Females contributed significantly more than males to incubation, shading/brooding, and feeding of the
nestling, whereas males provided prey to females during incubation and to nestlings during the early
nestling-rearing period. Foraging males, which spend more time away from the nest, may be more exposed to
anthropogenic activities and negative interactions with humans. If males are more vulnerable to such effects,
this could decrease the survival of breeding males and potentially increase the recruitment of immature
males to the breeding population. In this event, the ability of the inexperienced immature males that replace
the adult eagles to cope with the food and care demands of the mate and offspring is critical for ensuring
Chaco Eagle population maintenance.

KEY WORDS: Chaco Eagle; Crowned Eagle; Crowned Solitary Eagle; Buteogallus coronatus; Argentina; breeding; nest;
nestling rearing; parental care.

CUIDADO PARENTAL DE LA ESPECIE EN PELIGRO DE EXTINCIÓN BUTEOGALLUS CORONATUS EN

EL CENTRO DE ARGENTINA

RESUMEN.—Las diferencias sexuales en el cuidado parental pueden afectar la supervivencia o el desempeño

individual y también la demografı́a. Investigamos el comportamiento de cuidado parental de la especie en
peligro de extinción Buteogallus coronatus para evaluar las implicaciones en la conservación poblacional en
hábitats semiáridos del centro de Argentina. De 2004 a 2012, utilizando el registro en video, monitoreamos
tres intentos reproductivos durante el periodo de incubación (275 horas), nueve parejas durante el perı́odo
de crı́a de pollos (1087 horas) y realizamos observaciones focales en nidos (232 horas). B. coronatus evidenció
una fuerte división por sexo del cuidado parental. Las hembras contribuyeron significativamente más que los
machos en la incubación, empollando o dando sombra, y en la alimentación de los pollos, mientras que los
machos proveyeron de presas a las hembras durante la incubación y a los pollos durante el periodo inicial de

1
Email address: [email protected]

316
SEPTEMBER 2018 CHACO EAGLE PARENTAL CARE 317

crı́a. Los machos en busca de alimento, que pasan más tiempo lejos del nido, pueden estar más expuestos a
actividades antropogénicas y a interacciones negativas con los humanos. Si los machos son más vulnerables a
estos efectos, esto podrı́a disminuir la supervivencia de los machos reproductivos e incrementar
potencialmente el reclutamiento de machos inmaduros en la población reproductiva. En este caso, la
habilidad de los machos inmaduros poco experimentados que reemplazan a las águilas adultas en la
obtención de alimento y en el cuidado de la pareja y los pollos es crı́tica para asegurar el mantenimiento de la
población de B. coronatus.
[Traducción del equipo editorial]

For altricial species, the survival of newborn parental care behavior and the roles of pair
individuals is influenced by the parents’ ability to members during breeding.
provide food and care to the young. Most bird The division of labor between the sexes during
species, and particularly raptors, share parental breeding and its possible effects on adult survival
duties between both adults (Newton 1979, Clutton- may have implications for the conservation of long-
Brock 1991, Cockburn 2006, Webb et al. 2010). lived bird species with deferred maturity, such as the
Although shared parental care maximizes fitness in Chaco Eagle. In other eagle species, a high
some species (Emlen and Oring 1977), different proportion of immature individuals in the breeding
parental-care strategies also exist. Among some population may be an indicator of a population
decline (Balbontı́n et al. 2003, Ferrer et al. 2003).
raptors that exhibit little or no sexual-size dimor-
Underlying the relationship between the replace-
phism (e.g., vultures, condors), the division of roles
ment of one of the members of a pair by a subadult
is minimal (Margalida and Bertran 2000), whereas in
individual and possible negative population trends is
other species the division of duties during breeding,
the inability of a young and inexperienced parent to
including the hatching, nestling-rearing, and fledg- cope with food and care demands of the mate and
ing periods, is more marked (Newton 1979). offspring. Thus, subadult individuals that attempt to
The Chaco Eagle (formerly known as the Crowned breed are often unsuccessful, which may reduce
Eagle or Crowned Solitary Eagle; Buteogallus corona- productivity, particularly in small populations (Pen-
tus) is one of the largest birds of prey in South teriani et al. 2003); in those cases, nesting failure
America. It inhabits open semiarid forests from could be higher than expected, with more severe
southern and central Brazil, Bolivia, and Paraguay to consequences on population demography of those
northern Patagonia in Argentina (Ferguson-Lees small or threatened populations.
and Christie 2001). It is classified as endangered by Here we examine the parental-care behavior of the
the International Union for Conservation of Nature, endangered Chaco Eagle in breeding territories of
with a global population estimated at ,1000 central Argentina. We evaluated the roles of males
reproductive individuals (BirdLife International and females during the incubation and nestling-
2016). Mortality factors appear to include human rearing periods to assess whether a marked division
persecution (Sarasola and Maceda 2006, Sarasola et in parental roles occurs and to evaluate future
al. 2010, Barbar et al. 2016), electrocution at power implications for population conservation.
lines (Galmes et al. 2017), and consequences of STUDY AREA
habitat loss (Bellocq et al. 2002, Fandiño and
We studied Chaco Eagles in the southernmost part
Pautasso 2013).
of their range in western La Pampa province, central
The ecology of the Chaco Eagle is poorly known.
Argentina (Fig. 1) (approx. 378S, 668W). The study
The species seems to have low productivity, laying a area included approximately 18,000 km2, with an
single egg per breeding attempt (Giai 1952, Maceda elevation ranging between 220–340 masl, within the
2007, Carvalho Filho et al. 2009, Berkunsky et al. Espinal and Monte Desert ecoregions (Brown et al.
2012, Fandiño and Pautasso 2013). In addition, 2006). Vegetation types within the Espinal ecoregion
Chaco Eagles probably do not breed until 3–5 yr of include deciduous xerophytic forests of caldén
age, as is true for other large eagles (Newton 1979, (Prosopis caldenia), grassy savannahs, grassy steppes,
Del Hoyo et al. 1994), and the species may breed and bushy steppes. Typical vegetation in the Monte
only every 2 or 3 yr (Maceda 2007, Berkunsky et al. Desert ecoregion is represented by communities of
2012). There is no information about Chaco Eagle high shrub-steppe (characterized mainly by Larrea
318 GALMES ET AL. VOL. 52, NO. 3

Figure 1. (A) Location of Chaco Eagle nests monitored with video cameras (B) and focal observations at our study area in
(C) La Pampa Province, central Argentina, from 2004 to 2012.

spp.), with isolated trees of the genus Prosopis. In 200–300 m from the nest using 10 3 50 binoculars
both ecoregions, the climate is temperate arid, with and a 20 3 60 spotting scope. Observations were
high temperatures in summer (up to 448C) when conducted daily from dawn-to-dusk (about 0630 H
most of the scarce rainfall occurs; annual rainfall to 2200 H). At each nest, we recorded the time and
ranges between 80–300 mm in the Monte Desert and duration of each activity performed by each member
300–550 mm in the Espinal (Fernandez and Busso of the pair.
1997). Although Chaco Eagles are not sexually dimorphic
in plumage, like most raptor species they are sexually
METHODS dimorphic in size, with females being larger than
We searched for Chaco Eagle nests in our study males (Newton 1979, Ferguson-Lees and Christie
area from mid-September to mid-February in 2004– 2001). Thus, we determined sex visually, based on
2005 and 2007–2012 (Fig. 1). Eagle nests were relative size. We carefully recorded particular char-
located during field surveys by car and on foot, and acteristics of each individual (e.g., plumage features
by soliciting information during interviews with local such as molt or incomplete adult plumage) to
people. facilitate identification when only one member of
We conducted focal observations of individuals the pair was present. When sex identification was
during the 2007 and 2008 breeding seasons. Two unclear, we recorded the sex of the bird involved as
observers recorded behavior from a hide located unknown.
SEPTEMBER 2018 CHACO EAGLE PARENTAL CARE 319

To record behavior we installed video cameras on interaction effects of the explanatory variables, the
branches of the nest tree (1–2 m from the nest). least significant variables (P . 0.05) were successive-
During 2004 and 2005 breeding seasons we used an ly eliminated until a suitable minimum model was
analog camera connected to a time-lapse VHS video obtained in which all the retained variables im-
recorder (VCR Panasonic AG-1070 DC, Kadoma, proved the adjustment level. We tested for differ-
Osaka, Japan), and from 2009–2012 we used a digital ences between models including a variable and
camera (IP Vivotek 7330, San José, CA, USA) those not including it using ANOVA; if the
connected to a 1TB hard disk (Network Attached difference was significant we excluded that variable.
Storage, LaCie, d2 Network2, Cupertino, CA, USA). Only significant effects (P , 0.05) were retained in
All recorders were housed in a waterproof compart- the final model. We checked for overdispersion in all
ment. Monitoring systems were powered by a 12 V models.
battery connected to a solar panel that allowed In the different models, the sex of the adult
power autonomy of the equipment. The VCR was performing the behavior was included as a two-
programmed to record during two 8-hr time blocks category factor (male or female). Daily time effect
(0600–1400 H, 1400–2200 H) and the digital camera was included in the models as a three-category factor
was programmed to record in a continuous fashion (one for each of three time blocks). The age of
from 0600–2100 H, at which time the camera was nestlings was included as an explanatory variable
turned off until the next day. with six levels in the brooding/shading and nest-
During the incubation period, we recorded the material delivery models, and with nine levels for the
number and length of incubation bouts per individ-
other models, with each level corresponding to one
ual (male vs. female) per day. We also recorded
week of the breeding season. To evaluate whether
deliveries of nest material (branches and grasses)
there was variation between the sexes in food
made by each adult eagle to rebuild or repair nests.
provisioning to the nestling, we tested for an
We assumed that nocturnal incubation was per-
interaction of sex of the food provider and age of
formed by the last member of the pair identified
the nestling.
incubating at dusk.
To assess for variation in the timing of prey
During the nestling-rearing period, we recorded
deliveries throughout the day, and sex differences in
the number and length of bouts of brooding/
shading of nestlings performed by each adult. We that timing, in addition to time blocks and sex, we
also recorded deliveries of nesting material (branch- incorporated the type of prey as a factor, grouping
es and sprigs) to the nest for this period. Finally, we most prey deliveries into two classes (snakes or
recorded the number of prey deliveries to the nest by mammals). To assess those variations, we tested all
each member of the pair, the arrival time at the nest, two-way interactions among the three variables. To
as well as the type (snake or mammal) and size assess variation in size of prey delivered to the nest by
(small: 250 g; medium: between 251–500 g, and sex or date, we included prey size as a three-level size
large: .500 g) of food items. We estimated these class in the models (see above); we also included all
values visually based on our experience of weights of two-way interactions among the three variables.
prey taxa in the area. To explore the daily timing of Generalized linear mixed models were run with
prey deliveries, we divided daily video recording the lme4 package (Crawley 2007, Zuur et al. 2009) of
periods into three time blocks (0600–1100 H, 1101– R statistical software version 3.0.2 (R Development
1600 H, and 1601–2100 H). We recorded behavior Core Team, R Foundation for Statistical Computing,
until the young left the nest, at about 10 wk old. Vienna, Austria, www.R-project.org) with the pair/
Statistical Analyses. We used generalized linear nest identity as a random factor in all tests. We
mixed models (GLMMs) to test for sexual differenc- present values as mean 6 SD except when stated
es in the number of incubation bouts, brooding/ otherwise.
shading bouts, nest-material deliveries, prey deliver-
RESULTS
ies, and daily number of feeding bouts to nestlings
during the nestling-rearing period. Models were From 2004 to 2012, we collected information on
built using Poisson distribution errors and a log-link parental-care behavior during 12 breeding attempts
function. For modelling, we followed a backward by Chaco Eagles at 11 different nests. The mean
stepwise procedure. From a full model, which minimum distance between studied nests was 32.3
included the main effects as well as all the two-way km (range: 16.1–69.2 km).
320 GALMES ET AL. VOL. 52, NO. 3

Incubation Period. We used cameras to monitor coefficient of nestling age ¼0.38 6 0.12, where age
three breeding attempts during the incubation is expressed in weeks; v2 ¼ 9.86, df ¼ 1, P , 0.01).
period: one in the 2005–2006 breeding season (28 We recorded 238 prey deliveries to the nest, 158
Dec–6 Jan) and two in the 2010–2011 season (7–19 (66.4%) by males, 66 (27.7%) by females; in 14 cases
Dec 2010 [when the camera stopped working], and (5.9%) the sex of the provider could not be
23 Oct–17 Nov 2010). Adults incubated 71% of the identified. Parent eagles delivered 2.33 6 1.52 prey
time in 256 incubating bouts (n ¼ 275 hr); eggs were items per d (range ¼ 1–10). Males provided more
left exposed 29% of the time. The length of daily food items per day than females (males: 1.60 6 1.23,
mean incubation bout was 47.0 6 24.4 min (range ¼ range ¼ 0–6, n ¼ 98; females: 0.66 6 0.85, range ¼ 0–
8–126, n ¼ 256). All three nests failed before 4, n ¼ 98; v2 ¼ 39.3, df ¼ 1, P , 0.01) during the
hatching. entire nestling-rearing period; the difference was
Incubation was shared by both members of the reduced after the nestling reached 4 wk old
pair, although not equally. Females performed most (interaction of the sex of the parent 3 nestling
of the daily incubation (93.2 6 17.1% of the time vs. age: v2 ¼ 5.70, df ¼ 1, P ¼ 0.02). Females rarely
6.9 6 17.1% for males; n ¼ 39, v2 ¼ 3460.6, df ¼ 1, P provided food to the nest in the first 3 wk of the
, 0.01) and throughout all nights recorded. nestling’s life (Fig. 2).
Females performed more incubation bouts per day The size of food items delivered by the adults did
(6.9 6 7.2 bouts, range ¼ 1–31, n ¼ 40) than males not change over the nestling-rearing period (v2 ¼
(0.6 6 1.4 bouts, range ¼ 0–6, n ¼ 40; v2 ¼ 249.85, df 3.98, df ¼ 2, P ¼ 0.14) and food size was not related to
¼ 1, P , 0.01). the sex of the parent providing the prey (v2 ¼ 1.59,
During incubation, adults brought nest material df ¼ 2, P ¼ 0.45). Eagles brought more small and
19 times, at a rate of 0.68 deliveries/d with a medium-sized prey items than large ones to the nest
maximum of two deliveries in a day (n ¼ 28 d). (v2 ¼ 48.47, df ¼ 2, P , 0.01).
Females made most deliveries (89%) although males Both parents delivered food items to the nest
also contributed to this task (11%). Although throughout the day, from early morning (0645 H)
nesting material was delivered throughout the day, until dusk (2045 H), with a marked peak from 0800–
most deliveries were in the morning. 1000 H and a lesser peak from 1800–2000 H (Fig. 3).
Nestling-rearing Period. We monitored nine pairs The number of food items delivered to the nest did
during the nestling-rearing period for a total of 1319 not differ by sex of parent through the day (sex
hr. Most observations were obtained by video effect: v2 ¼ 1.86, df ¼ 1, P ¼ 0.17) or with the age of
monitoring (1087 hr) with a smaller contribution the nestling (week: v2 ¼ 0.61, df ¼ 1, P ¼ 0.44), but
from focal observations (232 hr). did change with the type of prey (v2 ¼ 5.61, df ¼ 1, P
Adult eagles brought nest material 32 times during ¼ 0.02), with snakes being delivered mainly in the
the nestling-rearing period at a rate of 0.76 morning and mammals in the afternoon.
deliveries/d with a maximum of four deliveries in a Females did more feeding of the nestlings than
single day. The provision of nesting material differed did males (females: 2.22 6 1.65 feeding bouts/d,
between males and females during the nestling- range ¼ 0–8; males: 0.20 6 0.47 bouts/d, range ¼ 0–
rearing period (v2 ¼ 3.79, df ¼ 1, P ¼ 0.05); as during 2; n ¼ 69; v2 ¼ 136.52, df ¼ 1, P , 0.01). The number
incubation, females contributed more (72% of nest of feeding bouts by adults decreased over time as the
material provisions) to the maintenance of the nest growing nestling began to feed itself (v2 ¼ 5.40, df ¼
than males (28%) in this period, especially in the 1, P ¼ 0.02; Fig. 4). The nestling started feeding itself
first weeks of life of the nestling. during the fifth week of age and feeding bouts by the
Nest attendance (brooding/shading) was done male ceased entirely after the nestling reached 7 wk
almost exclusively by females (96.5% of the brood- old.
ing/shading bouts, n ¼ 57), with males brooding or
shading only twice. The length of brooding/shading DISCUSSION
bouts was 110.4 6 110.5 min (range ¼ 1.2–514.1, n ¼ Our results indicated a strong sex-based division of
57). Adult eagles brooded/shaded the nestling for roles in the parental care of Chaco Eagles. As in
2.1 6 1.5 bouts/d (range ¼ 1–6, n ¼ 27). We did not other monogamous birds of prey (Newton 1979,
record shading events after the nestling reached an Everett 1981, Collopy 1984, Margalida et al. 2007),
age of 7 wk. As the nestling aged, the number of females carried out most activities at the nest,
shading bouts/d decreased (intercept ¼ 3.01 6 0.45; including incubation, shading/brooding, and feed-
SEPTEMBER 2018 CHACO EAGLE PARENTAL CARE 321

Figure 2. Mean (6SD) daily prey deliveries to nests by Chaco Eagles during the nestling-rearing period.

Figure 3. Hourly number of prey deliveries to Chaco Eagle nests in La Pampa Province, central Argentina.
322 GALMES ET AL. VOL. 52, NO. 3

Figure 4. Mean daily number of feeding bouts to the nestling performed by females, males, and nestlings (6SD) during
the nestling-rearing period.

ing of the nestling, while males performed most During the nestling-rearing period, as in the
activities away from the nest. Males provided prey incubation period, parental-care tasks were not
both to the female during incubation (M. Galmes equally shared by the sexes. Females were closely
unpubl. data) and to the nestling during early associated with the nest during the first weeks of
nestling-rearing, and females joined males in hunt- nestling rearing, while males hunted and provided
ing and providing food progressively as the nestling food. The nestlings hatched at the end of spring or
grew. Most diurnal and all nocturnal incubation was in summer, when temperatures in our study area can
done by the female, with males incubating only reach up to 448C (Casagrande and Conti 2004).
briefly, presumably while the female consumed prey Therefore, during the first weeks after hatching,
delivered by the male. Therefore, as in other eagle when nestlings are unable to thermoregulate,
species (Collopy 1984, Margalida et al. 2007), female regular protection from the sun by the adults is
essential. In fact, direct sun exposure during mid-
Chaco Eagles remained at the nest to incubate, and
and late-afternoon has been suggested as an
self-fed from food delivered by the male.
important cause of nestling mortality in Golden
During incubation, Chaco Eagles spent extended
Eagles (Aquila chrysaetos; Beecham and Kochert
periods away from the nest (up to 29% of daylight
1975). In Chaco Eagle pairs, most brooding and
hours) compared with other large raptor species in shading of the nestling was provided by the female,
which the eggs are left exposed for no more than with only occasional assistance by the male. This
10% of the time (e.g., 3.7 6 0.4% [Collopy 1984]; behavior ended when the nestling was around 7 wk
4.2 6 4.48% [Margalida et al. 2007]). However, the old. Similar sharing of duties during the nestling-
roles of males and females during incubation and rearing period has been described for other eagle
the differential rates of nesting-material deliveries species (Ellis 1979, Collopy 1984, Margalida et al.
during the incubation period were similar to those 2007, Watson 2010, Bakaloudis and Vlachos 2011).
documented in other eagle species (Collopy 1984, The adult male provided most food for the
Margalida et al. 2007), suggesting they are probably nestling and in some cases for the female as well.
representative of the behaviors and roles of adult This sex-based difference in food provisioning was
Chaco Eagles. greatest during the first weeks of the rearing period,
SEPTEMBER 2018 CHACO EAGLE PARENTAL CARE 323

with .90% of prey items delivered by males. The the nest by males in the first weeks of the nestling-
rate of prey deliveries by the female increased rearing period is shared by both the eaglet and the
gradually, as the necessity for brooding/shading female. In contrast, at the end of that period, the
decreased. Differences between the adults in prey nestling eats alone and vigorously defends the prey,
delivery rates decreased by the middle of the rearing which is rarely shared with an adult.
period when the young was approximately 40 d old, Most of the feeding bouts were carried out by the
and remained fairly constant until the end of the female, especially during the first weeks of the
rearing period (wk 10), when the contribution by rearing period when she was closely associated with
the female reached a maximum of 45%. Similarly, the nest. Although the male sometimes fed the
male Golden Eagles delivered over 80% of prey nestling, he did so only occasionally and never after
during the first 2 wk of the rearing period, but the nestling reached 7 wk of age. This behavior by
females reached a maximum of 47% of prey the male is likely related to the beginning of self-
deliveries by wk 9 (Collopy 1984). feeding by the nestling. During its first 7 wk of life,
The daily rate of prey delivery (2.36 deliveries) was the nestling was apparently mostly dependent on its
similar to that of the Short-toed Snake-Eagle parents for handling of food. After that, the nestling
(Circaetus gallicus; Bakaloudis and Vlachov 2011), was able to feed itself from food left at the nest by
which has the same brood size (one young) and a adults.
similar diet based on snakes (Fergusson-Lees and Conservation Implications. The role division
Christie 2001). However, we did not observe exhibited by Chaco Eagles suggests it would be
variation during the nestling-rearing period as unlikely for a single adult to raise a nestling
described by Collopy (1984) for Golden Eagles. successfully if the other pair member died during
Prey deliveries to the eaglet suggest a bimodal incubation or shortly after hatching. Some mortality
distribution with a marked peak of deliveries in early factors for the species are related to direct human
morning and a smaller peak in late afternoon. These persecution and electrocution (Sarasola and Maceda
results are consistent with the crepuscular habits of 2006, Galmes et al. 2017). The strong attachment of
the Chaco Eagles (Canevari et al. 1991, Collar et al. females to the nests may reduce their risk of such
1992). This bimodal pattern was constant through- mortality during the nesting season, relative to the
out the rearing period and was independent of the risk incurred by males, which hunt for prey over
sex of the provisioning adult. However, reptiles were larger areas of the landscape, potentially encounter-
delivered more often in the morning and mammals ing more anthropogenic threats. Conversely, the
more often in the evening. Although we did not female is consistently found at the nest, and thus
investigate hunting strategies, our results suggest would be more vulnerable to shooting by a poacher
that Chaco Eagles exhibit hunting activity consistent aware of the nest location. If there is a sexual
with the daily activity of its main prey. In particular, difference in vulnerability to mortality factors, this
snakes should be vulnerable to capture in the would potentially result in recruitment of less-
morning when they are more active (Bakaloudis experienced immatures of the more vulnerable sex
2010), as suggested for the Short-toed Snake-Eagle into the breeding population. In this context, the
(Bakaloudis and Vlachov 2011). Conversely, mam- ability of nonbreeding adults (‘‘floaters’’) or imma-
mals such as the pichi (Zaedyus pichiy), the main tures to enter the breeding population to replace
mammalian prey of the Chaco Eagle (Sarasola et al. adult eagles may be important for ensuring popula-
2010, Pereyra Lobos et al. 2011), are taken late in the tion maintenance. We recommend additional re-
day when the temperature starts to decrease (Abba search to assess which sex is more vulnerable to
et al. 2009). mortality factors, and the potential effect of breed-
In theory, the increasing requirements of food by a ing experience in this species.
growing nestling could be met by an increasing
number or size of prey delivered to the nest, or both ACKNOWLEDGMENTS
(Newton 1979, Collopy 1984). However, we did not We thank Joaquı́n Cereghetti, Fernando Urquiza,
note such patterns. Although the parents delivered Facundo Barbar, Anabel Gallardo, Marcos Reyes, Claudina
small prey items more frequently than larger prey, Solaro, Milena diBenito, Marı́a del Mar Contaldi, Laura
Beinticinco, Elena Brandoni Bescos, Nayla Massara, Melina
this pattern was consistent throughout the rearing Martı́n, Emanuel Leizica, Leela Pahl, Morgan Peters, Clare
period, even after the female began providing food. Staveley, Jay Carslile, José Marı́a Galea, and Juan José
A likely explanation could be that prey delivered to Maceda for their assistance in fieldwork, and Verónica
324 GALMES ET AL. VOL. 52, NO. 3

Salvador for help with data analysis. We thank the many Ambiental Argentina 2005. Fundación Vida Silvestre,
local landowners from La Pampa province that gave us Buenos Aires, Argentina.
permission to access their ranches and provided informa- Canevari, M., P. Canevari, G. R. Carrizo, G. Harris, J.
tion on the location of Chaco Eagle nests and perch sites. Rodrı́guez Mata, and R. J. Straneck (1991). Nueva Guı́a
We thank Germán Almeira who designed the digital
de las Aves Argentinas. Fundación Acindar, Buenos
cameras equipment and Lorenzo Sympson and the
University of California at Berkeley for providing the VHS Aires, Argentina.
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Aves Argentinas/AOP (Argentina), IDEA WILD (USA), the diurnas do Parque Estadual do Rio Doce, Minas Gerais,
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Constructive comments provided by Antoni Margalida and
Pampa /Universidad Nacional de La Pampa (Editors).
an anonymous referee improved an earlier version of this
work. Inventario Integrado de los Recursos Naturales de la
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