Diet of Breeding Neotropic Cormorants at the Carrileufu

River, Patagonia: Is there Any Impact on Recreational Fish

Resources?
Author(s): Ricardo Casaux, Analía Ramón, María Alejandra Tartara, Verónica
Borrell and Romina Gonc
Source: Ardeola, 59(2):279-289. 2013.
Published By: Spanish Society of Ornithology/BirdLife
DOI: http://dx.doi.org/10.13157/arla.59.2.2012.279
URL: http://www.bioone.org/doi/full/10.13157/arla.59.2.2012.279

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Ardeola 59(2), 2012, 279-289

DIET OF BREEDING NEOTROPIC CORMORANTS

AT THE CARRILEUFU RIVER, PATAGONIA: IS THERE
ANY IMPACT ON RECREATIONAL FISH RESOURCES?

DIETA DEL BIGUÁ EN EL RÍO CARRILEUFU, PATAGONIA,

DURANTE LA REPRODUCCIÓN: ¿HAY IMPACTO
EN LAS PESQUERÍAS DE INTERÉS RECREATIVO?

Ricardo CASAUX 1, 2, 3 *, Analía RAMÓN 4, María Alejandra TARTARA2, 3,

Verónica BORRELL 2 and Romina GONC 2

SUMMARY.—Some cormorant species are perceived as negatively interacting with fish resources but
conclusive evidence supporting such suggestions are scarce. Here we studied the diet of the neotropic
cormorant and assessed its impact on recreational fish resources. A total of 33 pellets (regurgitated casts)
and 73 regurgitations produced by the Neotropic cormorant were collected between 19 October 2006
and 24 February 2007 at a colony on the Carrileufu River, Patagonia, Argentina. Fish (94.3%) were the
most abundant prey, followed by crustaceans (5.1%) and molluscs (0.6%). The Patagonian silverside
Odontesthes hatcheri was the most frequent fish prey, followed by the rainbow trout Oncorhynchus
mykiss and the creole perch Percichthys trucha. The contribution of fish species to the diet varied
throughout the breeding season. The fish intake estimate for the colony based on parameters drawn from
previous studies was 4.708 tons, which represents a consumption of 41,504 fish specimens. Native
species were the major component of the cormorant diet, comprising 75.7% of the specimens and 73.2%
of the biomass consumed during the course of the breeding season. Given fish availability and
productivity within the study area, the impact on recreational fish resources by Neotropic cormorants can
be estimated to be of minor importance.
Key words: diet composition, fish intake, impact on fish resources.

RESUMEN.—Varias especies de cormoranes son percibidas como perjudiciales para la explotación

de recursos pesqueros. Sin embargo, la información que sustente tales supuestos es escasa. En este tra-
bajo se estudió la composición de la dieta del biguá y se evaluó la magnitud del impacto sobre pobla-

1
Instituto Antártico Argentino, Cerrito 1248, (1010) Buenos Aires, Argentina.
2
Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET),
Av. Rivadavia 1917, (1033) Buenos Aires, Argentina.
3
Laboratorio de Investigaciones en Ecología y Sistemática Animal (LIESA),
Universidad Nacional de la Patagonia, Ruta 259 km 5, Planta de Aromáticas,
9200 Esquel, Chubut, Argentina.
4
Subsecretaría de Medioambiente, Municipalidad de Esquel, Mitre 524,
9200 Esquel, Chubut, Argentina.

* Corresponding author: pipocasaux@infovia.com.ar

280 CASAUX , R., RAMÓN , A., TARTARA, M. A., BORRELL , V. and GONC , R.

ciones de peces de interés deportivo. Mediante muestreos desarrollados cada 15 días entre el 19 de
octubre de 2006 y el 24 de febrero de 2007 en la colonia de biguás localizada en el río Carrileufu,
Patagonia, Argentina, se recolectó un total de 33 pellets y 73 regurgitados espontáneos. Peces (94,3%),
seguidos por crustáceos (5,1%) y moluscos (0,6), fueron la presas más abundantes. El pejerrey pata-
gónico Odontesthes hatcheri fue el pez más importante en la dieta, seguido por la trucha arco iris
Oncorhynchus mykiss y la trucha criolla bocachica Percichthys trucha. La contribución de las diferen-
tes especies de peces a la dieta varió a lo largo del período reproductivo. El consumo de peces estimado
para la colonia basado en los parámetros extraídos de estudios previos a lo largo del ciclo reproductivo
es de 4.708 toneladas, que representan el consumo de 41.504 peces. Los peces nativos representaron el
75,7% de los ejemplares y el 73,2% de la biomasa consumida a lo largo del ciclo reproductivo. Dada la
disponibilidad y productividad de peces dentro del área de estudio, el impacto en las especies de peces
de interés deportivo puede estimarse como de importancia menor.
Palabras clave: composición de la dieta, impacto sobre recursos pesqueros, ingestión de peces.

INTRODUCTION lake and they estimated that these birds had

The Neotropic cormorant Phalacrocorax
olivaceus has a widespread distribution, from
the south of the USA to Cape Horn, Patago-
nia (Orta, 1992). In Argentina it occupies a
wide variety of wetlands in fresh, brackish
and salt waters. Given that cormorants often
breed in large colonies, they can consume
great quantities of fish locally and there is well
established concern that these birds may have
a negative impact on recreational fisheries
and aquaculture. Consequently, as with other
cormorants elsewhere (Orta, 1992; Bildsoe
et al., 1998; Cairns et al., 1998; Frederiksen
et al., 2001; Barras, 2007), the neotropic cor-
morant is severely persecuted in Argentina, the
usual population control involving shooting
and harassment of colonies. Despite the abun-
dance of neotropic cormorants and their sup-
posed negative effects on fisheries, only a
few studies have studied their natural diet,
reproduction and demography in Patagonia,
and most of them were carried out in colonies
settled within marine environments (see
Daneri, 1960; Quintana et al., 2002 and 2004;
Forero et al., 2004; Frere et al., 2005). Re-
cently, Casaux et al. (2009) have provided the
first information on the diet of non-breeding
Neotropic cormorants at a west Patagonian
a negligible impact on a fish farm there.
The only currently active colony of
Neotropic cormorants at west Chubut is on
the Carrileufu River. The river and the
surrounding lakes are frequented by recrea-
tional fishermen targetting introduced rain-
bow trout Oncorhynchus mykiss and brown
trout Salmo trutta. Thus, the objective of this
study is twofold. Firstly, we provide the first
report on the breeding season diet of breeding
Neotropic cormorants from a West Patagonian
river based on the analysis of pellets and regur-
gitations collected throughout the breeding
cycle. We then estimate the possible impact
of this bird on recreational fishing stocks.
MATERIAL AND METHODS
The diet was estimated on the basis of 33
pellets and 73 regurgitations (produced spon-
taneously by cormorants when we approached
the colony). These were collected during
visits every two weeks between 19 October
2006 and 24 February 2007 at a colony with
62 active nests on the Carrileufu River
(42º 32.358’ S, 71º 34.112’ W), Chubut,
Patagonia, Argentina. During each visit all
the area under the nests was surveyed.

Ardeola 59(2), 2012, 279-289

 DIET OF THE NEOTROPIC CORMORANT
Nets were deployed below nests located in
branches over the water to prevent the loss
of samples. Despite this procedure, and
according to evidence found in nets and on
the ground, it is probable that some samples
were ingested by mammals and birds. Pellets
(4, 6, 22, 0 and 1 pellets recovered per month
from October to February) were the only sam-
ple type recovered in October whereas the
number of regurgitations found (0, 2, 9, 54
and 8 per month) increased from November
and, except for one pellet, were the only sam-
ple type obtained in January and February. It
has been proposed that stomach contents may
mainly reflect the chicks’ diet, the adults’
food being completely or partly digested
before their return to the nest (Wanless et al.,
1993; Favero et al., 1998), whereas pellets
may represent both the adult and chick diet
more completely (Harris and Wanless, 1993).
It is obvious that the food represented in
stomach contents during the pre-laying, laying
and incubation periods reflects the adult
diet. On the other hand, it has been observed
that the food carried to the nest by Antarctic
shags Phalacrocorax bransfieldensis during
early- and mid-rearing seemed to exceed the
chicks’ energy requirements (Casaux, 1998).
Also, according to Bowmaker (1963), the
durations of those foraging trips were shorter
than the time required by shags to digest their
own food. From this information it could be
inferred that the stomach contents collected
during these periods of the breeding season
represent not only the chicks’ diet but also,
at least partly, the adults’ diet, which validates
the comparison throughout the study period
of both sources of information. On the other
hand, Casaux et al. (1998) compared results
from pellets and stomach contents of the
Antarctic shag collected simultaneously and
they observed slight qualitative differences
(the trophic spectrum represented in pellets
was wider than that observed in stomach con-
tents) but important quantitative differences
between both methods. Those authors sug-
281
gested that such differences might be ex-
plained by the higher number of food loads
represented in pellets (four to seven, Casaux
et al., 1997) compared to the stomach con-
tents (one), which also supports the com-
parison of pellets and stomach contents in
this study.
After collection, pellets were dried at 60 ºC
in the laboratory and their contents sorted into
prey classes using a binocular microscope.
The otoliths present in the samples were iden-
tified to species, where possible, using our
own reference collection based on fish also
caught at West Chubut. The otoliths of each
fish species were sorted into right and left
and the most abundant was taken as the num-
ber of individuals per fish species present in
the sample. Otolith lengths were measured to
0.01 mm to estimate the size and mass of the
individuals, applying the equations below
estimated from fish caught in the study area
(Casaux et al., 2009; R. Casaux, unpublished
information). Given that fragile parts of
otoliths were intact, these estimates were not
affected by any degradation of otoliths during
the gastrintestinal process.
Patagonian silverside Odontesthes hatcheri
TL = 11.12642 * OL0.618657 (N = 109, r = 0.91)
M = 7.428144 * OL2.041558 (N = 109, r = 0.92)
Creole perch Percichthys trucha
TL = 3.64696 * OL0.776257 (N = 78, r = 0.96)
M = 8.483607 * OL1.325035 (N = 78, r = 0.93)
Large puyén Galaxias platei
TL = 7.6719 + 5.1024 * OL (n = 308, N = 0.27)
M = 4.152005 * OL3.029642 (n = 308, N = 0.73)
Rainbow trout
TL = 11.03561 * OL0.809850 (N = 13, r = 0.73)
M = 50.93738 * OL1.537905 (N = 13, r = 0.57)
where “TL” is the fish total length in centime-
ters, “OL” is the otolith length in millimeters
and “M” is the fish mass in grams.
Ardeola 59(2), 2012, 279-289
 TABLE 1 282

Diet composition of the Neotropic cormorant as reflected by the analysis of 33 pellets and 73 regurgitations collected at Carrileufu River, Patagonia,
Argentina, between October 2006 and February 2007. Percentage frequencies of occurrence (F%), number (N%) and mass (M%), and total length
(TL, in cm ± standard deviation, range and the number of specimens measured in parentheses) for each prey category.
[Composición de la dieta del biguá basada en el análisis de 33 egagrópilas y 73 regurgitaciones recogidas en el río Carrileufu, Patagonia, Argen-
tina, entre octubre de 2006 y febrero de 2007. Se presentan los porcentajes de presencia (F%), número (N%) y masa (M%), así como la longitud

Ardeola 59(2), 2012, 279-289

total (TL, en cm ± desviación típica, rango y número de ejemplares medidos en paréntesis) para cada categoría de presa.]

Pellets Regurgitations Overall

F% N% M% TL F% N% M% TL F% N% M% TL
Molluscs 3.0 1.0 — — — — — — 0.9 0.6 — —
Crustaceans
Insects
Coleopteran 6.1 6.3 — — — — — — 1.9 3.4 — —
Trychoptera
Parasericostoma
3.0 3.1 — — — — — — 0.9 1.7 — —
ovale
Fish 97.0 89.6 — — 100 100 — — 99.1 94.3 — —
Odontesthes
30.3 50.0 69.2 28.0 ± 5.7 (19.0 – 38.8, n = 43) 18.0 23.1 33.6 26.3 ± 6.4 (17.9 – 43.1, n = 18) 22.3 37.2 54.1 27.5 ± 5.9 (17.9 – 43.1, n = 61)
hatchery
Galaxias platei 3.0 1.2 0.4 19.0 (n = 1) 6.6 7.7 2.7 18.2 ± 0.6 (17.6 – 19.0, n = 6) 5.3 4.3 1.4 18.3 ± 0.6 (17.6 – 19.0, n = 7)
CASAUX , R., RAMÓN , A., TARTARA, M. A., BORRELL , V. and GONC , R.

Percichthys
12.1 25.6 16.2 14.1 ± 4.3 (10.0 – 23.0, n = 22) 14.8 12.8 12.6 17.6 ± 2.6 (14.0 – 23.0, n = 10) 13.8 19.5 14.7 15.2 ± 4.2 (10.0 – 23.0, n = 32)
trucha
Oncorhynchus
12.1 9.3 12.7 21.3 ± 3.0 (17.8 – 26.6, n = 8) 16.4 23.1 41.9 22.2 ± 2.6 (18.9 – 27.4, n = 18) 14.9 15.9 25.1 21.9 ± 2.7 (17.8 – 27.4, n = 26)
mykiss
Salmo trutta 6.1 3.5 1.4 14.8 ± 1.4 (13.3 – 16.0, n = 3) 18.0 24.4 9.2 13.0 ± 1.4 (10.0 – 15.5, n = 19) 13.8 13.4 4.7 13.3 ± 1.4 (10.0 – 16.0, n = 22)
Unidentified 27.3 10.5 — — 11.5 9.0 — — 17.0 9.8 — —
 DIET OF THE NEOTROPIC CORMORANT
Regurgitations were composed solely of
fish, and species were identified from the
external characters of undigested fish or by
identifying the otoliths recovered from di-
gested fish. Lengths of undigested specimens
were measured to 0.1 cm and their weights
recorded to 0.1 g using an electronic balance.
Lengths and masses of digested specimens
were estimated from their otolith length using
the equations above.
To estimate the fish mass intake (FMI,
equation 1) during the breeding season (BS,
105 days) of Neotropic cormorants at the
Carrileufu River colony we considered the
mean number of breeding cormorants (BC)
observed there during the study period (mean
114 individuals, range 94-124, N = 6 cen-
suses); data on daily food consumption (DFC)
estimated for non-breeding adults at Chas-
comús Lagoon, Argentina (229.5g, Padín,
1987); the energy requirements of chicks
(CER) from hatching to fledging (60 days,
R. Casaux, pers. obs.) estimated according to
Weathers (1992): considering a fledging mass
similar to the adult mass of 1,800 g (Orta,
1992); and the number of fledged chicks
(NFC), taking a fledging success of 1.7
chicks per nest and 57 nests (R. Casaux,
pers. obs.). To estimate the contribution of
different fish species to the diet by mass
(FSCM) we distributed the FMI between
fish prey species according to their overall
contribution to the diet by mass (see table 1).
These estimates should be taken with caution
and regarded as only tentative given that they
are based on several parameters extracted
from previous studies from different areas
and/or under different conditions. To estimate
the number of fish individuals ingested (NFI,
equation 2) throughout the breeding season,
we divided the FSCM corresponding to each
fish species by the estimated mean mass of
the individuals of that species (EMM) repre-
sented in the samples [Patagonian silverside
173.9 g (± 137.4, range 35.8 – 698.7, N = 62),
large puyén 38.7 g (± 4.4, range 33.8.6 – 43.2,
283
N = 7), creole perch 90.2 g (± 47.9,
range 35.0 – 200.0, N = 33), rainbow trout
189.3 g (± 45.1, range 126.9 – 287.3, N = 26)
and brown trout 41.9 g (± 11.7, range
21.0.6 – 65.0, N = 24)].
FMI = (BC * DFC * BS) + (CER * NFC)
(Equation 1)
NFI = (FSCMx/EMMx) + (FSCMy/EMMy) +
(FSCMz/EMMz) +
(Equation 2)
The c2-test was used to evaluate differences
in the contribution by mass of fish species
to the diet of cormorants as reflected by the
analysis of pellets and regurgitations. The
Mann-Whitney U-test was used to analyse
differences in the fish sizes represented in
pellets and regurgitations. The Kruskall-
Wallis test was used to analyse changes in the
sizes fish represented in the diet throughout
the study period. Spearman tests were used to
detect trends in the fish sizes ingested and
to analyse the relationship between the num-
ber and size of the Patagonian silversides
ingested throughout the breeding season.
RESULTS
Fish were by far the most frequent and
important prey by number, followed by
coleopterans and trichopterans (table 1).
Among the detected fish, the patagonian sil-
verside was the most frequent prey and it also
predominated by number and reconstructed
mass; the rainbow trout and the creole perch
followed in importance (table 1). The con-
tribution of different fish species to the
cormorant diet differed between pellets and
regurgitations ( c 24 = 67.9, p < 0.01 ) . The
Patagonian silverside, followed by the creole
perch, were the fish that most contributed
to the diet by mass in pellets, whereas the
rainbow trout, followed by the Patagonian
Ardeola 59(2), 2012, 279-289
284 CASAUX , R., RAMÓN , A., TARTARA, M. A., BORRELL , V. and GONC , R.

FIG. 1.—Changes throughout the breeding season in the contribution to the diet (as percentage of mass
consumed) of fish represented in pellets and regurgitations produced by Neotropic cormorants at the
Carrileufu River colony, Patagonia, Argentina.
[Cambios durante la temporada de reproducción del biguá en la contribución a la dieta (como porcentaje
de la masa ingerida) de los peces representados en las egagrópilas y regurgitaciones producidas por las
aves de la colonia del río Carrileufu, Patagonia, Argentina.]

silverside, did so in regurgitations (table 1).
The creole perch was the most important fish
prey in cormorant diet from nest building to
early incubation (mainly October), whereas
the Patagonian silverside was the fish that
most contributed to the diet during hatching
and brood rearing (mainly early November
to mid January). The rainbow trout predomi-
nated at the time of brood fledging (mainly
late January and February) (fig. 1).
The size of the individuals represented in
the samples ranged between 10 cm (creole
perch and brown trout) and 43.1 cm (patago-
nian silverside) in total length (table 1).
Except for the creole perch (Mann-Whitney
U-test, U = 54.5, p < 0.05), there were no
differences between the size of fish found in
pellets and regurgitations. The size of the fish
ingested fluctuated throughout the season
(Kruskall-Wallis, H = 33.3, df = 6, p < 0.001)
without any evident trend (Spearman test, ns),
although the overall fish length peaked both
in mid December and January (fig. 2). It is
interesting that the rainbow and the brown
trout presented opposite trends. The number
and size of the Patagonian silversides ingested
throughout the season were positively corre-
lated (Spearman test, P < 0.05, rs = 0.86).
According to estimates of fish intake based
on published information (see Material and
methods), during the course of the breeding
season Neotropic cormorants (adults and
chicks) at the Carrileufu River colony in-
gested 4.708 tons of fish (2.747 and 1.961
tons ingested by adults and chicks respec-
tively), which represents a consumption of

Ardeola 59(2), 2012, 279-289

 DIET OF THE NEOTROPIC CORMORANT 285

FIG. 1.—Changes throughout the breeding season in the size (total length in cm) of fish represented
in pellets and regurgitations produced by Neotropic cormorants from the colony at Carrileufu River,
Patagonia, Argentina.
[Cambios durante la temporada de reproducción del biguá en el tamaño (longitud total en cm) de
los peces aparecidos en las egagrópilas y regurgitaciones producidas por las aves de la colonia del río
Carrileufu, Patagonia, Argentina.]

41,504 fish specimens (creole perch 1.18 tons
and 17,842 individuals; large puyén 0.04 tons
and 1,164 individuals; patagonian silverside
2.22 tons and 12,413 individuals; rainbow
trout 1.02 tons and 4,849 individuals; brown
trout 0.24 tons and 5,236 individuals).
Overall, native species predominated in the
diet, contributing 75.7% of specimens and
73.2% of the mass consumed.
DISCUSSION
Neotropic cormorants from the Carrileufu
River foraged predominantly on fish and much
less intensively on insects and molluscs, as
reported elsewhere in South America (Jordán,
1967; Oliveros and Beltzer, 1983; Kalmbach
et al., 2001; Regidor and Terroba, 2001;
Fortuna et al., 2003; Barquette et al., 2008;
Casaux et al., 2009). However, given that
molluscs and insects were represented in
pellets but not in regurgitations, these inver-
tebrates may actually have been consumed
directly by the fish prey and not by cor-
morants. This hypothesis is also supported
by the fact that those molluscs and insects
are important dietary components of fish in
the study area (Casaux and Di Prinzio, 2007;
Di Prinzio and Casaux, 2012).
Eight fish species were available within
the foraging area (Liotta, 2005) and the
Neotropic cormorant preyed on a variety
of these, confirming that it is a generalist
(Telfair and Morrison, 1995; Barquete et al.,
2008). As observed by Casaux et al. (2009)

Ardeola 59(2), 2012, 279-289

286 CASAUX , R., RAMÓN , A., TARTARA, M. A., BORRELL , V. and GONC , R.
at Rosario Lake, a locality close to the
Carrileufu River, the Patagonian silverside
and the rainbow trout were important prey,
whereas the benthic-demersal and cryptic
large puyén scarcely contributed to the diet.
This seems to reflect a general pattern of fish
consumption by the Neotropic cormorant in
the study area, which may be related to its
foraging technique or to the conspicuousness
of potential prey (see Casaux et al., 2009).
As indicated above, the contribution of
fish species to the diet varied according to
the analyses of pellets or regurgitations. This
may be because pellets and regurgitations
may reflect the diet of chicks and adults
differentially (Harris and Wanless, 1993;
Wanless et al., 1993; Favero et al., 1998).
However, such difference s seem to be
mainly restricted to the later stages of chick
rearing (Casaux, 1998; see Material and
methods). In this study pellets and regurgi-
tations were mainly collected during the
initial and the final part of the sampling
period respectively. This suggests that the
differences in the fish represented in pellets
and regurgitations may also reflect changes in
the diet through the breeding season. Thus,
the creole perch was the most important fish
prey from nest building to early incubation,
whereas the Patagonian silverside was the
fish that most contributed to the diet from
hatching to fledging, when the rainbow trout
increased in importance.
It has been suggested that cormorants can
deplete fish stocks in waters close to their
colonies throughout the breeding season
(Birt et al., 1987; Leopold et al., 1998;
Casaux, 2003). The fish represented in the
samples have similar energy densities (Cian-
cio et al., 2007). Thus, it is probable that the
observed changes in the diet throughout the
breeding season reflect natural or cormorant-
related changes in prey availability in the
foraging area. It is interesting that the num-
ber of Patagonian silversides (the most im-
portant prey in terms of biomass) consumed
Ardeola 59(2), 2012, 279-289
throughout the breeding season was positive-
ly correlated with the size of the individuals
ingested, which may be evidence of prey
size preferences and temporal changes in the
abundance of different Patagonian silverside
length classes, perhaps related to predation
pressure.
Cormorants are perceived as harmful to
recreational fish stocks and fish farms (see
also Nettleship and Duffy, 1995; Suter, 1995;
Bildsoe et al., 1998; Lekouna, 1998; Barras,
2007; among others). In west Patagonia, this
assumption is supported by the fact that
Neotropic cormorants forage on introduced
salmonids, the target species for recrea-
tional fishing and fish farms, at recreational
fishing areas or close to pound nets. Hence,
Neotropic cormorants are usually persecuted
in order to displace them from their feeding
grounds. As a result, currently their breeding
colonies are really rare in west Chubut, and
only two colonies occur along the 350 kilo-
metres that separate Villa La Angostura,
Neuquén, and Río Pico, Chubut (R. Casaux,
unpublished information). According to
our fish intake estimates, during the entire
breeding season Neotropic cormorants from
the colony at Carrileufu River can ingest
4.71 tons and 41,504 individual fish, of
which native species account for 75.7%
and 73.2% of the number of individuals and
biomass respectively. Although our calcula-
tions must be taken with caution, according
to the maximum daily allowable catch per
fishermen for rivers and lakes within the
foraging area of cormorants (Anonimous,
2009), the Neotropic cormorants only take
the trout specimens that 15 fishermen could
have legally obtained throughout the fishing
season (180 days). Although there are no pre-
cise estimates for the Neotropic cormorants’
foraging area, approximately 10,000 fisher-
men visit west Chubut during the fishing sea-
son (R. Malerba, pers. com.). These estimates,
together with the fact that salmonids con-
sumed by Neotropic cormorants are smaller
 DIET OF THE NEOTROPIC CORMORANT
than those sought by fishermen, suggests
that these birds have only a minor impact on
recreational fish stocks. We also considered
the potentially detrimental effect of cor-
morants on fish recruitment. According to
Baigún (2001), the estimated annual fish
productivity for lakes surrounding the study
colony (without considering the rivers) is
46 tons year –1. Taking our estimates into
account, these cormorants consume during
the breeding cycle and during an entire year
roughly 10% and 30%, respectively, of the
annual fish production of the lakes within
their foraging range. Thus, these birds should
be protected as natural regulators of fish
populations instead of being regarded as
harmful to fish stocks.
ACKNOWLEDGEMENTS.—We thank Estancia
Los Murmullos for the facilitation of the access
to the sampling site and logistic support. This is
the contribution to the Laboratorio de Investiga-
ciones en Ecología y Sistemática Animal (LIESA)
N° 92.
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