Ornithology Research

https://doi.org/10.1007/s43388-023-00140-6

ORIGINAL ARTICLE

Trophic ecology of Geranoaetus polyosoma and Tyto furcata

in an Atacama Desert agricultural landscape
Pablo Valladares‑Faúndez1 · Katherine Ramírez‑Meza2,3 · Sergio Alvarado Orellana4 · Robert Langstroth5

Received: 30 December 2022 / Revised: 5 July 2023 / Accepted: 9 July 2023

© The Author(s), under exclusive licence to Sociedade Brasileira de Ornitologia 2023

Abstract
The diets of the variable hawk (Geranoaetus polyosoma) and the barn owl (Tyto furcata) have been previously reported for
the Atacama Desert of northern Chile. However, their ecological interactions and their role in the control of invasive species
are unknown. In this work, the diet and competitive interaction of both raptors in an agricultural landscape were evaluated.
Both species consume a high proportion of rodents (26.3% and 63.3% respectively), particularly Rattus sp. (20.8% and 30.0%
respectively), which represent the highest contribution to the biomass consumed (67.9% and 60.2% respectively). Geranoae-
tus polyosoma consumes reptiles such as Microlophus yanezi (11.7%) and invertebrates (42.7%), mainly Coleoptera (30.7%)
and Decapoda (7.11%). On the other hand, Tyto furcata consumed native rodents as Oligoryzomys flavescens (7.13%) and
Auliscomys boliviensis (6.22%) and marine birds nesting in the desert, as Oceanodroma markhami (0.55%). Both raptors
hunt over wide geographic ranges from the coast to the highlands, in wetlands, agricultural and desert areas. Despite sharing
44% of the prey consumed, there is only moderate interspecific competition. We discuss the possible role of both raptors as
controllers of the exotic invasive Rattus sp.

Keywords Agriculture landscape · Arid ecosystem · Invasive rats · Raptors · Trophic ecology

Introduction barn owl Tyto furcata. Geranoaetus polyosoma is a spe-

Our understanding of the trophic ecology of Atacama
Desert raptors has increased in recent years, particularly
for the variable hawk Geranoaetus polyosoma and the
Communicated by Caio Graco Machado (Associate Editor)
* Pablo Valladares‑Faúndez
pvalladares@academicos.uta.cl
1 phic (Brown and Amadon 1968), of medium size, with a
Laboratorio de Zoología Integrativa, Departamento de
Biología, Facultad de Ciencias, Universidad de Tarapacá,
General Velásquez, 1775 Arica, Chile
2
Programa de Magíster en Áreas Silvestres y Conservación
de la Naturaleza, Facultad de Ciencias Forestales y
Conservación de la Naturaleza, Universidad de Chile,
Avenida Santa Rosa, 11315 Santiago, Chile
3
Fundación ANKA, Conservación y Educación, Macul,
2349 Arica, Chile
4
Laboratorio de Ecología de Vida Silvestre, Facultad de
Ciencias Forestales y de la Conservación de la Naturaleza,
Universidad de Chile, Avenida Santa Rosa, 11315 Santiago,
Chile
5 geophytes and annual species adapted to these conditions
Independent Researcher, South Riding, VA, USA
cies of raptor of the Accipitridae family, which has a wide
distribution in South America; it has a northern distri-
bution from the central mountain range of Colombia to
Cabo de Hornos in the south, and from the coasts of Chile
to Argentina (Brown and Amadon 1968; Cabot 1991; De
Lucca 2011), reaching the Malvinas Islands in the Atlantic
(Cabot 1991) and the Juan Fernández Islands in the Pacific
(Blake 1977). It is considered one of the most studied rap-
tors in Chile (Raimilla et al. 2012); it is sexually dimor-
wingspan of 1100 mm, body length between 450 and 530
mm for males and 485 and 630 mm for females, but adult
males between 690 and 1134 g and adult females between
876 and 1417 g (Jiménez 1995). The diet of variable hawk
in Chile was first studied by Schlatter et al. (1980) and
Jiménez (1995). In the Atacama region, Valladares-Faún-
dez et al. (2015, 2018) first analyzed its diet and then com-
pared in areas of “absolute desert” and “blooming desert.”
The “blooming desert” refers to an area of the southern
Atacama Desert where there are occasional austral spring
rainfalls that produce episodic massive flowering events of

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(Gutiérrez 2008). Ponce et al. (2017) analyzed its diet in
the Tarapacá region of Chile. Finally, the diet of this raptor
from Tacna region, southern Peru, is mainly composed of
Coleoptera (76.22%) and reptiles, such as Microlophus sp.
(11.06%) and Liolaemus sp. (5.75%). However, the bio-
mass contributed by Microlophus sp. represents 64.6%,
and Coleoptera only 1.22% (Valladares-Faúndez et al.
2021).
The nocturnal raptor Tyto furcata is a member of the
Strigidae family, a raptor species which is one of the most
widespread in South America. In Chile, this raptor is found
across the entire country, inhabiting a large diversity of envi-
ronments (Carmona and Rivadeneira 2006). The diet of barn
owl in northern Chile has been studied in the regions of
Coquimbo (Schamberger and Fulk 1974), Atacama (Valla-
dares-Faúndez et al. 2016), Antofagasta (Jaksic et al. 1999),
and Tarapacá (Carmona and Rivadeneira 2006), where in
each case rodents, particularly phyllotines, were the pri-
mary source of food. However, Ramírez et al. (2000) also
reported the consumption of lizards in Tacna, Peru, and sug-
gested that the owls engage in diurnal hunting. In the Medi-
terranean ecosystems of central Chile, Jaksic et al. (1981)
found a food-niche overlap (i.e., Pianka’s index) of 0.537
between our focal species, which they considered to indicate
a medium level of overlap.
While both focal species predate a range of prey species,
there are no reports of predation by variable hawk on inva-
sive exotic species in arid ecosystems of Chile. However,
Cabot-Nieves et al. (2020) reported high levels frequencies
(26.7%) and biomass (56.3%) of Rattus spp. in pellets of
the closely related Puna hawk Geranoaetus poecilochrous
in the high Andes of the Antofagasta region. Carmona and
Rivadeneira (2006) reported a low percentage (0.9%) of
Rattus rattus for barn owl in Tarapacá. Also, Ramírez et al.
(2000) reported house mice (Mus musculus) as the primary
item in barn owl pellets in Tacna, both in number of indi-
viduals and biomass. In Mediterranean Chile, Jaksic et al.
(1981) reported R. rattus as prey for B. virginianus but not
for variable hawk, barn owl, or any other raptor.
Although the number of studies on raptor diets in arid
ecosystems has increased in recent years, these studies have
been limited to the description of diets and preferences for
certain prey species and there have been no studies of the
interspecific relationships in raptor assemblages, nor of the
roles of raptors as controls of invasive vertebrate species.
Therefore, we report here on our research to evaluate (1)
the diversity and abundance of prey species in the diets of
variable hawk and barn owl coexisting in an Atacama Desert
ecosystem with a strong influence of agricultural production;
(2) the importance of invasive exotic species in the diet of
the two raptor species, particularly Rattus sp.; and (3) the
overlap of trophic niches to the infer the intensity of compe-
tition between the two species of raptors.
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Ornithology Research
Methods
Study area
This study was carried out in the Quebrada de Camarones
valley, Arica y Parinacota region, Chile (Fig. 1), located in
the Atacama Desert. The Camarones river creates a rela-
tively high level of vegetative cover compared to other parts
of the desert and supports a high level of human activity,
primarily crop cultivation, and industrial-scale poultry pro-
duction. There are two main human settlements in the val-
ley: Camarones (19°00′40.49″S; 69°51′32.40″W) and Cuya
(19°09′40.42″S; 70°09′47.43″W), some 33.6 km apart.
Between these localities, there are a series of perches fre-
quently used by each of the two raptor species studied.
Pellet collection and dietary composition
We collected pellets of the variable hawk and barn owl
between October 2017 and May 2020, particularly in repro-
ductive seasons; this is approximately May and October for
both species. All the pellets were collected under nests or
perches that they frequently use, and that we have been able
to identify for each of the species analyzed. Each pellet was
collected in a paper bag and assigned a numerical code.
The pellets were transported to the Universidad de Tara-
pacá, Laboratorio de Zoología Integrativa in Arica, Chile,
where they were individually hydrated for an hour prior to
the separation of the remains and the taxonomic determina-
tion of the prey species. For rodents, we used the available
regional molar keys (Muñoz-Pedreros and Gil 2009; Yáñez
et al. 2009), along with reference collections for rodent,
lizard, and bird species that inhabit that Camarones valley
and adjacent areas of extreme northern Chile. The remains
were examined under an Olympus XTL-2310 binocular
microscope.
Data analysis
Analyses of the frequency, abundance, and biomass of prey
were performed. As measures of the diversity of prey con-
sumed, we calculated species richness (S) and Simpson’s
diversity ­(D1), which is the complement to the original
Simpson’s index (D) (Simpson 1949) and represents the
probability that two randomly selected individuals belong
to the same species (McCune and Grace 2002). Simpson’s
evenness index (E) represents the degree to which individ-
uals are divided between species with low values, which
indicate that one or a few species dominates, and high val-
ues which indicate that most of the species is represented
by a relatively similar number of individuals (Morris et al.
Ornithology Research
Fig. 1  Locations of pellet
collection in both species of
raptors. (A) Chile, (B) Arica,
and Parinacota region, and (C)
squares indicate pellet collec-
tion of Tyto furcata and circles
to Geranoaetus polyosoma
2014). We also calculated Shannon’s index (H′), which is
equally sensitive to rare and abundant species. We also cal-
culated the Berger–Parker dominance index (BP), where the
proportional abundance of species can be incorporated in
indices that represent diversity and report the proportional
abundance of only the most abundant species (Berger and
Parker 1970).
To evaluate trophic niche overlap, we calculated Pianka’s
index (Pianka 1974), where 0 indicates no overlap and 1
indicates complete overlap of resource use. We also calcu-
lated the Morisita-Horn index, which considers the similar-
ity of species composition as well as the similarity of their
abundances (Morisita 1959), where 0 indicates no similarity
and 1 indicates complete similarity. Finally, we performed
rarefaction analysis using the Chao-1 index to evaluate reli-
ability of the pellet sampling. Statistical analyses were per-
formed using PAST 4.03 (https://​www.​softp​edia.​com/​get/​
Scien​ce-​CAD/​PAST.​shtml).
Results
We analyzed a total of 1046 pellets: 509 for variable hawk
and 537 for barn owl, resulting in a total of 1982 identi-
fied prey, of which the majority was identified to species
or genus, except for arthropods. Tables 1 and 2 present the
main results on the abundance and biomass of prey and the
calculated indices.
A total of 27 prey taxa and 1435 prey were identified for
variable hawk, of which mammals represent 26.76% of the
number of preys consumed but 79.8% of the biomass. Of
these, rodents accounted for 26.3% of the prey and 79.6%
of the total biomass. The second most consumed group of
vertebrates were birds, which represent 16.3% of the prey,
but represent only 4.3% of the biomass, which explains
why the birds hunted by these raptors are small. As in other
populations of variable hawk reported from northern Chile,
reptiles (lizards) are also an important element of the diet,
representing 13.0% of the prey and 14.6% of the biomass.
Invertebrates comprised 42.9% of the prey but only 1.3%
of the biomass. Of these, Coleoptera are the most abundant
order with 30.7% of the total prey and river shrimp Cryphi-
ops caementarius another 7.1%.
A total of 16 prey taxa as identified for barn owl and 547
prey were identified, of which rodents accounted 63.3% of
the total prey and 79.0% of the biomass. Birds were the sec-
ond most abundant group overall, comprising 25.8 of the
prey and 20.7% of the biomass. In contrast to variable hawk,
we detected no reptile or invertebrate prey items in the barn
owl pellets.
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Table 1  Abundance and biomass of prey consumed by variable number of individual preys identified, nf number of pellets in which
hawks (Geranoaetus polyosoma) and American barn owls (Tyto fur- the prey is found, g grams
cata) in the Camarones valley, Arica y Parinacota region, Chile. n

Prey Geranoaetus polyosoma Tyto furcata

Abundance Frequency Biomass Total biomass Abundance Frequency Biomass Total biomass
n % nf % (g) (g) % n % nf % (g) (g) %

Order Rodentia
Andinomys edax 2 0.14 2 0.16 57 114 0.13 - - - - - - -
Akodon albiventer 3 0.21 3 0.23 51 153 0.17 - - - - - - -
Abrothrix andinus - - - - - - 2 0.37 1 0.19 20 40 0.07
Abrothrix olivaceus - - - - - - 14 2.56 15 2.87 35 490 0.90
Auliscomys cf boliviensis 22 1.53 20 1.56 54.6 1201.2 1.36 11 2.01 9 1.72 54.6 600.6 1.10
Auliscomys cf pictus 6 0.42 5 0.39 72.7 436.2 0.49 34 6.22 31 5.93 72.7 2471.8 4.54
Phyllotis cf limatus 10 0.70 9 0.70 68.5 685 0.78 21 3.84 20 3.83 68.5 1438.5 2.64
Eligmodontia hirtipes 2 0.14 2 0.16 28.5 57 0.06 6 1.10 37 7.08 28.5 171 0.31
Oligoryzomys flavescens - - - - - - 39 7.13 4 0.76 37 1443 2.65
Neotomys ebriosus 1 0.07 1 0.08 57.9 57.9 0.07 3 0.55 2 0.38 57.9 173.7 0.32
Cavia tschudi 24 1.67 22 1.71 320 7680 8.71 9 1.65 9 1.72 320 2880 5.29
Rattus sp. 299 20.84 291 22.66 200 59800 67.85 164 29.98 153 29.31 200 32800 60.20
Unidentified rodents 9 0.63 9 0.70 - - - 43 7.86 43 8.23 - - -
Total Rodentia 378 26.34 - 70184.3 79.63 346 63.25 - 42508.6 78.02
Order Chiroptera
Demosdus rotondus 2 0.14 2 0.16 30 60 0.07 1 0.18 1 0.19 30 30 0.06
Order Didelphimorphia
Thylamys pallidior 4 0.28 4 0.31 14.9 59.6 0.07 34 6.22 32 6.13 14.9 506.6 0.93
Total Mammals 384 26.76 - 70303.9 79.76 381 69.65 - 43045.2 79.01
Order Passeriformes
Sicalis uropygialis 56 3.90 53 4.13 15.04 842.24 0.96 - - - - - - -
Zonotrichia capensis 42 2.93 41 3.19 20.06 842.52 0.96 45 8.23 44 8.42 20.06 902.7 1.66
Spinus sp. 16 1.11 16 1.25 16.8 268.8 0.30 - - - - - - -
Phrygilus fruticeti fruticeti 6 0.42 5 0.39 41.8 250.8 0.28 - - - - - - -
Pyrocephalus rubinus 21 1.46 20 1.56 16.1 338.1 0.38 - - - - - - -
Sturnella bellicosa albipes 4 0.28 4 0.31 78.5 314 0.36 - - - - - - -
Order Columbiformes
Columba livia 2 0.14 2 0.16 360 720 0.82 25 4.57 25 4.79 360 9000 16.52
Zenaida auriculata 2 0.14 2 0.16 125 250 0.28 11 2.01 11 2.10 125 1375 2.52
Order Procellariiformes
Oceanodroma markhami - - - - - - - 3 0.55 3 0.57 53 159 0.29
Unidentified birds 85 5.92 85 6.62 - - - 57 10.42 57 10.91 - - -
Total Aves 234 16.31 - 3826.46 4.34 141 25.78 - 11436.7 20.70
Order Squamata
Liolaemus sp. 19 1.32 10 0.78 37.4 710.6 0.81 - - - - - - -
Microlophus sp. 168 11.71 136 10.59 72.6 12196.8 13.84 - - - - - - -
Total Reptiles 187 13.03 - 12907.4 14.64 - - - - - - -
Unidentified vertebrates 14 0.98 14 1.09 - - - 25 4.57 25 4.79 - - -
Total Vertebrates 805 57.07 - 87037.76 98.75 547 100.00 - - -
Class Insecta
Order Orthoptera 37 2.58 20 1.56 0.2 7.4 0.01 - - - - - - -
Order Hymenoptera 3 0.21 2 0.16 0.2 0.6 0.00 - - - - - - -
Order Coleoptera 440 30.66 377 29.36 0.5 220 0.25 - - - - - - -
Order Odonata 7 0.49 6 0.47 0.2 1.4 0.00 - - - - - - -

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Table 1  (continued)
Prey Geranoaetus polyosoma Tyto furcata
Abundance Frequency Biomass Total biomass Abundance Frequency Biomass Total biomass
n % nf % (g) (g) % n % nf % (g) (g) %

Total Insecta 487 33.94 - 229.4 0.26 - - - - - - -

Class Arachnida
Order Scorpiones 27 1.88 25 1.95 1.5 40.5 0.05 - - - - - - -
Class Malacostraca
Cryphiops caementarius 102 7.11 96 7.48 8.15 831.3 0.94 - - - - - - -
Total Invertebrates 616 42.93 - 1101.2 1.25 - - - - - - -
Total prey 1435 100.00 100 - 88138.96 100.00 547 100.00 - - - 54481.9 99.65

Table 2  Interpretation of the ecological index in the diet of G. polyosoma and T. furcata in the Arica and Parinacota region, Chile
Index Ecological interpretation G. polyosoma T. furcata

Richness (S) Number of species/taxa present in the diet of each species. 27 16

Simpson’s diversity (­ D1)Considers the number of prey species/taxa present in the diet 0.8235, (0.8356), 0.8472 0.845, (0.8659), 0.8818
of each species, as well as the relative abundance of each
species, where 1 represents infinite diversity and 0 represent
no diversity. As species richness and evenness increase, so
diversity increases.
Simpson’s evenness (E) Low values indicate that diet is dominated by a few species; 0.3289 0.5803
high values indicate that relatively equal numbers of items
belong to each prey species, where 1 indicates that one or a
few species are more numerous in the diet and 0 indicates that
all prey species are equally consumed.
Shannon index (H′) Measures the information content by species in samples 2.23, (2.3), 2.35 2.31, (2.4), 2.47
obtained at random from an extensive sample of which the
total number of species is known (S), where a value of H′ = 0
indicates a community of only a single species.
Berger-Parker index Reports the proportional abundance of only the abundant prey 0.2822, (0.3066), 0.3296 0.2614, (0.2998), 0.3382
species in the population of prey species.
Pianka index Estimates the degree of trophic niche overlap between the pair 0.4992
of raptor species, where 0 is no overlap and 1 is complete
overlap.
Morisita-Horn index (M-H) It is a statistical measure of dispersion of individuals in a popu- 0.4917
lation. It is used to compare overlap among samples. where 0
is no overlap and 1 is complete overlap.

In terms of invasive species, our sampling and analyses
demonstrate both variable hawk and barn owl to be impor-
tant predators of exotic rats Rattus sp. in agricultural mosaic
landscape in the Atacama Desert. Rattus were the single
most abundant prey item for the barn owl (30.0% of all prey)
and the most important vertebrate for the variable hawks
(20.8% of all prey).
Trophic level and niche amplitude
As shown by Table 2, the Simpson’s diversity indices were
similar for both species, yet slightly higher in barn owl
(0.866 vs 0.836). Simpson’s eveness was higher in barn owl
(0.580 vs 0.329), indicating a more uniform diet in terms
of the relative number of each prey species consumed in
barn owl while variable hawk demonstrates stronger pref-
erences for certain prey species. Trophic diversity, as esti-
mated by Shannon-Wiener indices, was 2.3 for the hawk
and 2.4 for the owl, where values greater than 1 indicate
sensitivity to the more abundant prey species (Jost 2007).
Similarly, the Berger-Parker dominance index was similar in
barn owl (0.300) and variable hawk (0.307), which indicates
the dominance of a few species in the diet of each species,
in this case Rattus sp. While the Morisita-Horn (0.492) and
Pianka (0.499) indices suggest only an intermediate level
of dietary overlap between the focal species, the contrasted
overlap and similarity values are likely explained by the data
that indicate that while Rattus sp. is dominant in the diet of

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both barn owl and variable hawk each raptor also consumes
several unique preys not preferred by the other. For exam-
ple, variable hawks consume far more passeriform birds and
lizards than do barn owls, while barn owls show a greater
preference for rock doves Columba livia and certain rodents
such as Abrothrix, Auliscomys, and Oligoryzomys genus,
which were not identified in variable hawk pellets (Table 1).
The rarefaction analysis indicates that our pellet sample
sizes were more than adequate to characterize the diet of
each target species. For variable hawk, the number of prey
species asymptotes at around 20–25 species after 300 sam-
ples, and for barn owl, the prey species curve levels out at
around 15–20 species after 150 samples. The Chao-1 index
indicates the predicted number of prey species at 30 for vari-
able hawk and 19 for barn owl, values coinciding with the
results of the rarefaction analysis.
Discussion
The diets of variable hawks and barn owls in the Camar-
ones valley agricultural landscapes generally reflect the pat-
terns previously reported for Atacama Desert ecosystems
in Chile (Table 3). While variable hawks were found to
consume large numbers of coleoptera, the biomass of these
prey is insignificant compared to that of the vertebrate prey,
especially rodents and lizards. The barn owl diets observed
here are like those reported previously, not only in arid eco-
systems, but also in other latitudes where rodents dominate
the diet (see Valladares-Faúndez et al. 2016). However, the
prey species richness observed in the Camarones valley is
greater than that reported in other localities of southern Peru
and northern Chile, likely because our sample landscape is
largely agricultural with higher vegetation cover and more
diverse habitats that in most of the Atacama Desert. Also,
the raptors of the Camarones valley have opportunities to
forage in habitats ranging from the Pacific coast to the edge
of the Altiplano, which provides them with access to a wide
range of prey species. For example, the presence of bats
in the pellets is related to the high number of caves that
exist on the coasts of the region, while rodents of the genera
Abrothrix, Cavia, and Rattus, and various passeriform and
columbiform species indicate the importance of the central
valley. In contrast, rodents of the genera Andinomys, Aulis-
comys, Eligmodontia, and Neotomys found in the sampled
pellets are characteristic of the higher Andean region.
Novel prey items reported here for variable hawk include
native guinea pigs Cavia tschudii, vampire bats Desmodus
rotundus, lizards of the genus Microlophus, and the fresh-
water river shrimp C. caementarius. We observed hawks
actively foraging for these shrimps by overturning small
stones and vegetation along small watercourses. The barn
owl diet includes Ausliscomys boliviensis and Auliscomys
pictus, Phyllotis limatus, Eligmodontia hirtipes, Neotomys
ebriosus, C. tschudii, Oligoryzomys flavescens, Demos-
dus rotondus, and Markham’s storm petrel Oceanodroma
markhami. This storm petrel is an IUCN-listed Near
Threated nocturnal migratory sea bird with inland desert
nesting colonies, one of which is at Pampa Camarones (Bar-
ros et al. 2019; Medrano et al. 2019). In other hand, sev-
eral skulls and mandibles belonging to rodents of the O.
flavescens were identified in the barn owl pellets, species
recently identified for Chile (Quiroga-Carmona et al. 2023).
Similarly, N. ebriosus is another Andean species poorly rep-
resented in Chilean collections (Revollo-Cadima et al. 2021)
but was identified in pellets of both raptors. The presence

Table 3  Comparison of statistical values and ecological indices for the diets of Geranoaetus polyosoma and Tyto furcata in arid ecosystems of
northern Chile. AD absolute desert, BD “blooming desert.” (†) This study, (‡) Carmona and Rivadeneira (2006), and (§) Ponce et al. (2017)
Geranoaetus polyosoma Tyto furcate
Camarones Pampa del Atacama region (§) Camarones Pampa del Atacama
valley (†) Tamarugal (‡) valley (†) Tamarugal (‡) region
AD BD (§)

Pellets sampled 509 63 417 566 537 66 149

Prey taxa 27 4 8 8 16 12 11
Total prey 1435 52 1460 1839 547 118 275
Abundance (%) Mammals 26.76 40.38 8.15 10.17 69.65 76.2 78.5
Birds 16.31 - - - 25.78 3.4 17.8
Reptiles 13.03 45.28 70.34 56.6 - 5 0.4
Invertebrates 42.93 13.46 21.5 33.22 - 15.1 3.3
Biomass (%) Mammals 79.76 30.86 16.6 11.06 79.01 - 96.4
Birds 4.34 - - - 20.7 - 3.5
Reptiles 14.64 68.76 82.71 86.8 - - 0.1
Invertebrates 1.25 0.39 1.13 2.13 - - 0.1

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of native guinea pigs in the diet of both species is of inter-
est given that the closest known locality for this species is
at Quebrada de Tana, 43 km to the south of the Camarones
valley (Valladares-Faúndez et al. 2020).
The Pianka index of 0.499 suggests a moderate or inter-
mediate level of competition between variable hawks and
barn owls in the Camarones valley. While the hunting activi-
ties of the two species are temporally segregated, there is
some overlap in the early evening or early morning periods
of crepuscular activity. The two species share about 50% of
their prey species and Rattus is the most frequent prey for
each, at 20.8% for the variable hawks and at 30.0% for the
barn owls. In terms of biomass consumed, Rattus comprised
67.9% of the hawks’ diet and 60.2% of the owls’ diet. While
not encountering any owl remains in the hawk pellets, we
found the remains of five barn owls very near an active vari-
able hawk nest, suggesting that competition for prey is not
the only interaction between these species.
The high frequencies of Rattus sp. in the diet of both barn
owls and variable hawks could demonstrate their importance
as controls of exotic invasive species, especially when con-
sidering that these rats are among the most widespread
and damaging invasive species, being highly aggressive.
Its species have generalist habits, consuming both native
animal and plant materials, leading to the displacement or
extirpation of local species, and whose eradication is key
to the recovery of many species of conservation concern
(see details in Barney et al. 2021). In Chile, the only avian
predator previously shown to frequently predate invasive
species is G. poecilochrous in the Altiplano and deserts
of the Antofagasta region, where exotic species of Rattus
and Mus were approximately 42% of its diet (Cabot-Nieves
et al. 2020). This may be explained by the high abundance of
these species in the Loa River valley near the city of Calama
(22°27′45″S; 68°55′38″W).
Information on the density of species prey in the desert
and highland rodents is very scarce in Chile. It is only avail-
able from Cofré and Marquet (1999) for some species of
rodents, such as Andinomys edax (398 ind/km2), Akodon
albiventer (394 ind/km2), Abrothrix olivaceus (1862 ind/
km2, but all of country), A. boliviensis (153 ind/km2), N.
ebriosus (56 ind/km2), and C. tschudi (102.8 ind/km2). We
do not have information on the abundance of the rest of the
prey species, some as relevant as rats, birds, or reptiles. This
information is relevant since it can help us understand what
the reason for the higher proportion of some species in the
diet is, a problem that will be considered in the future.
The barn owl and variable hawk nests encountered in
the Camarones valley are used for one or two clutches
annually. We observed two chicks in each barn owl brood
and consistently four chicks in each variable hawk brood.
Other raptors we observed in the Camarones valley are
black-chested buzzard-eagles Geranoaetus melanoleucus,
Harris’s hawks, American kestrels Falco sparverius, aplo-
mado falcons Falco femoralis, peregrine falcons Falco
peregrinus, Magellanic horned owl Bubo magellanicus,
and burrowing owls Athene cunicularia, even without find-
ing nests or perches to collect pellets.
Acknowledgements We thank the collaboration of Alexander Clifford
in the field and especially thank Jorge Claudio Fuentes for identifying
hawk nest’s locations in the Camarones valley. We thank Dr. Ángel
Spotorno (Universidad de Chile) for facilitating access to rodent skull
and mandible reference collections. We thank Dra. Marcela Vidal
(Universidad del Biobío) for her help with Microlophus biomass data.
Finally, we thank the company Ariztía S.A. for providing access to their
properties in the Camarones valley.
Funding This work was financed by grant UTA Mayor 4725/21 from
the Universidad de Tarapacá.
Declarations
Conflict of interest The authors declare no competing interests.
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