Italian Journal of Zoology

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Eurasian otter (Lutra lutra) prey selection in

response to a variation of fish abundance

L. Remonti , C. Prigioni , A. Balestrieri , S. Sgrosso & G. Priore

To cite this article: L. Remonti , C. Prigioni , A. Balestrieri , S. Sgrosso & G. Priore (2010)
Eurasian otter (Lutra lutra) prey selection in response to a variation of fish abundance, Italian
Journal of Zoology, 77:3, 331-338, DOI: 10.1080/11250000903229809

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 Italian Journal of Zoology, September 2010; 77(3): 331–338

Eurasian otter (Lutra lutra) prey selection in response to a variation

TIZO

of fish abundance

L. REMONTI, C. PRIGIONI*, A. BALESTRIERI, S. SGROSSO, & G. PRIORE

Otter prey selection

Department of Animal Biology, University of Pavia, Pavia, Italy

Abstract
In 2001 and 2002 we studied Eurasian otter (Lutra lutra) fish selection in southern Italy by comparing otter diet (805
analysed spraints) with fish availability, estimated by electrofishing. The surveyed river was dominated by cyprinids, partic-
ularly by chub (Leuciscus cephalus), roach (Rutilus rubilio) and bleak (Alburnus sp.). Roach and chub were the most common
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prey recorded in spraints. Otters ate mainly small prey (total length < 90 mm). Chub were widely consumed according to
abundance, whilst roach and bleak consumption was not proportionate to their availability. Otters selected roach both in
2001 and 2002 and responded to a decrease in roach availability (–27.3%) by increasing their frequency of consumption
rather than switching to an alternative prey. Despite their high abundance, bleak seemed to be markedly avoided by otters.
We hypothesized that differences in microhabitat selection may influence the susceptibility of cyprinids to otter predation.
The possible effects on fish community were briefly discussed.

Keywords: Lutra lutra, diet, prey selection, size selection, southern Italy

Introduction susceptibility to predation (depending on a variety of

behavioural mechanisms; review in Lehtiniemi
Fish are known to be the preferred prey of Eura-
2005). Handling time may be influenced by the
sian otters (Lutra lutra), although they can also rely
external protections of the prey (exoskeletons,
on alternative prey, such as crabs, crayfish,
spines, etc.) and by the ontogeny of the predator
amphibians and insects (Mason & Macdonald
hunting behaviour. As an example, crustaceans are
1986; Ruiz-Olmo & Palazón 1997; Jedrzejewska et
considered to require long handling times by otters
al. 2001; Clavero et al. 2003; Prigioni et al. 2006a;
(Watt 1993). Predation rate may be density-depend-
Remonti et al. 2008, 2009; Smiroldo et al. 2009).
ent, because prey density affects profitability; never-
Accordingly, the otter may be considered a gener-
theless, in other cases, predators may consume prey
alist aquatic predator, even though only a few
regardless of its abundance (Hughes & Croy 1993;
studies have simultaneously compared otter diet to
Prough 2005). The effects on prey communities are
food availability (Lanszki & Körmendi 1996; Carss
different: density-dependent predation leads to prey
et al. 1998; Lanszki et al. 2001; Ruiz-Olmo et al.
stability, whereas the disproportionate use of a
2001).
resource by a predator (Krebs & Davies 1993) may
Generalist predators feed opportunistically
destabilize prey communities and produce cyclical
according to prey profitability, which is the net
predator–prey dynamics (Hanski et al. 1991).
energy intake after the pursuit and handling of vari-
In addition to fish abundance, otter feeding
ous prey items (Stephens & Krebs 1986). When the
behaviour is expected to be influenced by fish size:
profitability of the main prey falls below a critical
large fish provide more energy, but probably need
threshold, a generalist predator is expected to switch
longer handling time, while small-sized fish are
to alternative prey items (Pyke et al. 1977; Stephens &
probably easily caught, but assume a lower energetic
Krebs 1986; Krivan 1996). The time spent by a
intake.
predator to catch prey may depend on prey density
We investigated the possible influence of fish
(conditions being equal, those prey which are more
abundance and fish size on otters’ feeding behaviour
abundant require lower search time) and prey

*Correspondence: C. Prigioni, Dipartimento di Biologia Animale, Università degli Studi di Pavia, Piazza Botta 9, I-27100, Pavia, Italy. Tel: +39 0382
986304. Email: prigioni@unipv.it
ISSN 1125-0003 print/ISSN 1748-5851 online © 2010 Unione Zoologica Italiana
DOI: 10.1080/11250000903229809
 332 L. Remonti et al.
over a two-year period, in a river of southern Italy
where fish were the main otter prey (Remonti et al.
2008). We compared otters’ diet (assessed by ana-
lysis of spraints) with both fish availability and fish
size (estimated by electrofishing) and we tested if:
(i) otters preyed on fish (both species and size-class)
in proportion to their abundance, or alternatively if
(ii) otters’ predation rate was unrelated to the
relative abundance of prey items, and otters
selected some species or size-classes and avoided
some others.
Moreover, we examined the variation in fish avail-
ability between the two years and tested for variation
in otter prey selection and whether otters switched
prey according to variation in fish abundance. The
possible effects on the fish community were briefly
discussed.
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Materials and methods
Study area
The study was conducted on the upper and middle
course of the River Sinni, a section of which runs
through the Pollino National Park (Figure 1). The
park is located in southern Italy between Basilicata
and Calabria (40°00’ N; 16°30’ E) and covers an
area of 1930 km2, with altitude ranging from 200 m
a.s.l. in the lower valley of the River Sinni to over
2000 m a.s.l. The River Sinni (101 km in length)
forms a wide east–west oriented valley in the north-
ern portion of the park and flows into the Ionian
Sea. The monitored stretch was between 15 and
53 km from the source and was 2–15 m wide and
0.5–2 m deep. Water flow peaks in winter and
decreases in late summer. Besides seasonal
Figure 1. Study area showing sampling sites.
variation, it is affected by dams, and ranges from
0.08 m3/s to 2.18 m3/s. Riparian woods consist
mainly of poplar (Populus alba, P. nigra), alder
(Alnus glutinosa) and willow (Salix spp.). In the
River Sinni, fish form the bulk of otter diet, whilst in
the surrounding rivers otters show a great adaptabil-
ity, preying on fish, amphibians and crustaceans
(Remonti et al. 2008). In the Pollino National Park
and surrounding areas, spraint distribution (Prigioni
et al. 2005) and genetic typing of fresh faeces
(Prigioni et al. 2006b) revealed a stable and relatively
abundant otter population (0.18–0.20 individuals/
km of watercourse).
Spraint collection and analysis
We gathered otter spraints on a monthly basis,
from January 2001 to December 2002, along six
river stretches (mean length = 675 m, S.D. =
151.4) almost uniformly distributed along the
river (mean distance between sampling reaches =
6.8 km, S.D. = 2.0). We washed the collected
spraints through a sieve of 0.5 mm mesh, gently
breaking them up by hand. Using a binocular
microscope, we identified fish remains from their
vertebrae, jawbones and scales, using personal col-
lections and identification keys by various authors
(Webb 1976; Wise 1980; Camby et al. 1984;
Prigioni 1997). We identified reptiles and amphib-
ians by the keys by Di Palma and Massa (1981),
whilst the undigested remains of insects (wings,
legs and cuticle parts) were compared with per-
sonal collections. Mammal hairs were compared at
20× and 40× magnifications by using the keys pub-
lished by Debrot et al. (1982) and Teerink (1991).
We considered each identified prey class as an
occurrence and expressed the overall diet as per-
cent frequency of occurrence (%FO = number of
occurrences of a prey type/number of analysed
spraints × 100). The raw frequency data were
compared by the χ2 test. Because of the presence
of repeated tests on related data, the sequential
Bonferroni technique was used to adjust the level
of significance (Rice 1989).
In the river stretches where spraints were col-
lected, fish were sampled by electrofishing in July
and November 2001 and 2002. We assumed that
our electrofishing sessions allowed a reliable estim-
ate of yearly fish availability. We surveyed a section
of each sampling reach (mean length = 82 m, S.D.
= 32.9) by moving upstream with a portable, bat-
tery-powered electrofishing apparatus. The length of
each section was at least 10 times the river width
(Verneaux 1981). We did not limit the sections with
nets. In streams and small rivers our electrofishing

procedure provides, under most circumstances, a
useful estimate of fish relative abundance at lower
costs than removal sampling (Fièvet et al. 1999;
Ross et al. 2001), even if small-sized fish and turbid
waters may reduce capture efficiency (Zitek et al.
2004).
Collected fish were identified to species or genus
level according to Gandolfi et al. (1991). They were
measured, weighed and released at the capture site.
The estimated fish biomass (B, g/100 m2) was calcu-
lated. Because of roach/chub hybridization, recently
verified by Amplified Fragment Length Polymor-
phism (AFLP) technique (Maldini et al. 2006),
roach (Rutilus rubilio), chub (Leuciscus cephalus) and
their hybrid forms were visually classified into two
functional groups of ‘chub-like’ fish and ‘roach-like’
fish, simply named as chub and roach.
We calculated fish availability (FA = yearly
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number of individuals of each fish species in elec-
trofishing samples/yearly number of all sampled
fish), and tested for variations in FA between 2001
and 2002 by the χ2 test. FA was compared to fish
Figure 2. Linear regression between the maximum length of jawbones and the maximum fish length for a sample of four cyprinid species
captured in the study area.
Otter prey selection 333
use (FU = yearly number of occurrences of a fish
species in otter spraints/yearly number of fish-con-
taining spraints) by the χ2 test, using Bonferroni’s
confidence intervals for the proportion of use (Neu
et al. 1974). We assumed that a fish species was
selected by otters when FU significantly exceeded
FA and, conversely, that a fish species was avoided
when FA significantly exceeded FU. Moreover, to
test for variations in otters’ food selection between
years, we compared the annual estimates of otter
selection ratio for each fish species (FU/FA) by the
χ2 test, as described in Manly et al. (2002).
The total length of cyprinids preyed by otters was
estimated by the length of jawbones found in otter
spraints. For this purpose, the maximum length of
the jawbone of a sample of four cyprinid species
(chub, roach, bleak Alburnus sp. and barbel Barbus
sp.) captured in the study area, was related to max-
imum fish length by a linear regression (Figure 2).
To test for size selection we compared, by the χ2 test
with the Bonferroni’s confidence intervals for the
proportion of use (Neu et al. 1974), the frequency of
 334 L. Remonti et al.

occurrence of six length-classes (50–70 mm, 71–90 Table II. Variation of fish availability (FA = yearly number of
mm, 91–110 mm, 111–130 mm, 131–150 mm, individuals of each fish species in electrofishing samples/yearly
number of all sampled fish) in 2001 and 2002 (***P < 0.001).
>150 mm) of fish preyed by otters in the whole
study period (roach: N = 109, chub: N = 108), with Species 2001 2002 χ2 1d.f.
the frequency of occurrence of the same length-
Brown trout 0.070 0.049 4.88
classes in sampled fish (roach: N = 688, chub: N =
Eel 0.001 0.001 0.05
778). When a low number of estimated prey lengths Roach 0.293 0.210 23.54***
was available, we compared the mean lengths of Chub 0.310 0.294 0.81
preyed and sampled fish by the t test. Bleak 0.271 0.343 15.88***
Barbel 0.037 0.047 1.79
Perch 0.010 0.008 0.20
Results Spined loach 0.008 0.047 42.71***

We collected and analysed a total of 805 otter

spraints, 490 in 2001 and 315 in 2002. Fish were
the most common prey items recovered in spraints
in both years (Table I), followed by amphibians
(Rana sp.). Crustaceans (Potamon fluviatilis and
Austropotamobius pallipes), birds, insects and reptiles
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were food sources of minor importance.

Electrofishing provided a total of 2663 fish,
belonging to eight taxa (brown trout Salmo trutta, eel
Anguilla anguilla, chub, roach, bleak, barbel, perch
Perca fluviatilis and spined loach Cobitis tenia).
Cyprinids dominated the fish assemblage, represent-
ing 94% of sampled fish (Table II). Brown trout
predominated only in the most upstream site (site 1,
Figure 1); brown trout and eels joined cyprinids in
site 2 (Figure 1). The cyprinid community was
dominated by chub, roach and bleak (Table II).
Overall fish availability decreased from 654 g/100
m2 in 2001 to 374 g/100 m2 in 2002, and otters
responded by decreasing fish consumption and by
increasing predation on amphibians (Table I).
Roach and chub were the commonest fish recorded
in spraints both in 2001 and 2002 (respectively,
72.5% and 81.0% of consumed fish). Chub were
widely consumed throughout the study period,
according to their frequency and without significant
differences between the two years (Table II, Figure 3).
Otters selected roach both in 2001 and 2002
(Figure 3), despite the decline of roach frequency in Figure 3. Otter fish selection (fish availability vs. fish use, see
Methods) in 2001 and 2002 (*P < 0.05, **P <0.01, ***P <
0.001).
Table I. Percent frequency of occurrence (%FO) of the main
food items in otter diet (in brackets, the number of spraints
examined) and comparison between 2001 and 2002 (*P < 0.05,
****P < 0.0001).
the second year (–27.6%). Otters responded to the
Overall 2001 2002 2002 decrease in roach availability by increasing
Food items diet (490) (315) χ2 1d.f. their predation effort, as showed by the increasing
difference between FU and FA (Figure 3), by the
Insects 1.52 1.63 1.28 2.3
Crustaceans 5.93 5.31 7.23 2.0 strong increase of the selection ratio for roach
Fish 81.66 85.10 74.47 249.9**** (Table III) and by the significant increase in roach
Amphibians 31.03 26.73 40.00 7.9* FU (χ2 = 4.7 P = 0.03; Figure 3). In contrast, otters
Reptiles 1.52 1.84 0.85 4.5 strongly avoided bleak (Figure 3) even when bleak
Birds 0.28 0.41 0.00 2.0
frequency increased significantly (Table II). The
 Otter prey selection 335

Table III. Comparison between annual estimates of otter N.S.) and barbel (sampled: N = 110, mean = 115
selection ratios (FU/FA; *P < 0.05, **P < 0.01, ****P < 0.0001). mm S.D. = 52.7; preyed: N = 31, mean = 99 mm,
Species 2001 2002 χ2 1d.f.
S.D. = 25.3, t = –1.69 N.S.).

Brown trout 1.09 0.19 83.9****

Eel 25.13 10.61 20.4**** Discussion
Roach 1.26 2.17 47.1****
Chub 1.15 1.21 0.6
Detecting food preference in wild animals is often
Bleak 0.23 0.18 6.0* difficult. Even when the environmental availability
Barbel 1.76 1.30 8.6** of different trophic resources has been carefully
Perch 3.91 5.30 1.9 assessed, only a rough picture of the relative abun-
Spined loach 2.09 0.10 251.2**** dance of each prey can still be drawn (Chanin
1985). As a consequence, most studies on otter food
selection have been carried out through feeding
experiments on captive animals (Erlinge 1968;
Fumagalli et al. 1995). Results of these investiga-
tions do not always agree, suggesting that many eco-
logical factors may affect the exploitation of a
particular resource and that local results can hardly
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Figure 4. Fish size selection (frequency of sampled fish vs. fre-
quency of preyed fish, see Methods) for roach and chub (*P <
0.05, **P <0.01, ***P < 0.001).
increasing selection toward roach showed by otters
in the second year was accompanied by a significant
decrease of the selection ratio for brown trout, eel,
bleak, barbel and spined loach (Table III).
Otters mainly ate small roach and chub (Figure 4),
selecting 71–90 mm long individuals and avoiding the
length class 91–110 mm. No significant difference
emerged between sampled and preyed fish for bleak
(sampled: N = 418, mean = 66 mm S.D. = 28.6;
preyed: N = 7, mean = 68 mm S.D. = 33.9, t = 0.14
be generalized.
The accuracy of the study can also be influenced
by the methods chosen to represent otter diet and
some authors have suggested that the use of the rela-
tive frequency of occurrence (RFO) or of FO may
produce a biased picture of otter diet (Carss & Par-
kinson 1996). Nevertheless, the general picture of
otter diet drawn by FO is often quite similar to what
is obtained by estimating the bulk or the biomass of
otter prey (Jacobsen & Hensen 1996; Jedrzejewska
et al. 2001; Clavero et al. 2004; Lanszki & Sallai
2006). The estimation of the bulk of food items may
be useful when otters largely rely on prey with a
large quantity of indigestible parts, such as crusta-
ceans (Remonti et al. 2008).
Our results only partially support the hypothesis
of a correspondence between fish species abundance
and predation rate by otters, suggesting a selective
feeding behaviour.
Concerning the main otter prey, roach were
selected, even when their frequency decreased in the
second year. Possibly roach availability did not reach
the critical threshold beyond which it would become
advantageous for otters to switch towards alternative
fish prey. This result agrees with reports by
Breathnach and Fairley (1993), who documented a
prompt and intensive predation by otters on intro-
duced roach (R. rutilus) in western Ireland. On the
contrary, otters avoided bleak, although they were
abundant, syntopic with widely eaten similar
cyprinids and their mean size fell into the range of
otters’ preferred prey-size.
Various hypotheses can be formulated in order
to explain why otters maximize profitability by
selecting roach, avoiding bleak and preying chub
in proportion to their abundance in the fish
community.
 336 L. Remonti et al.
Bleak can react to the chemical cues emitted by
predators, to predator images or to chemical signals
produced by injured conspecifics by altering their
activity and habitat choice (Jachner 1995a, 1995b,
1996). However, several cyprinid species show
behavioural alterations in response to the presence
of a predator (Reed 1969; Lima & Dill 1990;
Magnhagen & Forsgren 1991) and no evidence sup-
ports the hypothesis that bleak would be less suscep-
tible to predation by otters than roach or chub, by
virtue of a more efficient anti-predator behaviour
than chub or roach.
The predatory pressure of diurnal predators, such
as birds, can induce fish to feed in open waters dur-
ing the night, when a decreased light intensity
reduces the risk of predation even though this makes
search for food less effective (Winfield 1990). In our
study area, fish are exposed to pressure from aquatic
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predators (e.g. perch, large size chub), piscivorous
birds (e.g. herons) and otters, and each fish species
may ‘choose’ a different behavioural pattern, some-
times becoming more vulnerable to one predator by
escaping another.
Moreover, the diel dynamics of fish behaviour and
habitat use are not only predator-driven, reflecting a
complex trade-off between predator avoidance,
resource availability and inter-specific competition.
Bleak are known to be specialized open-water feed-
ers, foraging primarily on zooplankton (Politou et al.
1993; Herzig 1994; Vinni et al. 2000), while roach
rely primarily on aquatic macrophytes (92.5% of the
overall roach diet in our study area; Balestrieri et al.
2006). In a fish pond in Hungary, this difference in
food habits and consequently in habitat use influ-
enced otter predation rates: bleak, tending to remain
in open-water areas, were avoided by otters, while
roach, preferring shoreline aquatic plants, were
selected (Lanszki et al. 2001).
Although our study was conducted in a relatively
narrow river, we hypothesize that temporal or spatial
differences in the feeding activity or, more generally,
differences in microhabitat selection among fish spe-
cies influence their susceptibility to otter predation
and, consequently, their profitability as preys.
Concerning the food items of a minor importance
for otters, spined loach are small fish (up to 12 cm),
which spend most of their time inside the sediment
forming the riverbed, and their profitability to otters
is probably limited by the disproportionate time
costs for pursuing them. In 2002, the sharp decrease
of otter selection ratio for trout and eel may result
from a bias due to the low number of salmonid sam-
pling stations.
Otter selection for small-sized cyprinids con-
firmed reports from several studies carried out in
slow-running freshwaters, mainly populated by
Cypriniformes and/or Perciformes polyspecific assem-
blages (Brzezinski et al. 1993; Lanszki & Körmendi
1996; Lanszki et al. 2001; Clavero et al. 2004).
Young cyprinids are gregarious and, especially from
May to October, when many streams dry up in the
Mediterranean area, they are forced to assemble in
small, scattered pools (Prenda et al. 2001; Magal-
hães et al. 2002), thereby representing an easily
exploitable resource for otters.
Although otters did not switch to alternative fish
prey when the abundance of the most used fish spe-
cies declined, the decrease of the overall fish availa-
bility in 2002 produced an increase in amphibian
consumption, as previously reported for surround-
ing rivers (Remonti et al. 2008).
The selective feeding behaviour shown by otters
might destabilize the fish community: the predation
rate on chub, adding density-dependent mortality to
the dynamics of the chub population, will dampen
fluctuations and increase stability (Hanski et al.
1991). On the contrary, otter predation on roach
was not density dependent and, if confirmed, could
destabilize roach population, possibly driving it to a
progressive decline within the fish community. The
opposite effect could act on bleaks, leading to an
increase in their frequency. The impact of predators
on prey populations is however hard to assess, and
further long-term studies are surely needed to clarify
the role of otters on cyprinid dynamics.
Acknowledgements
This study has been promoted and financially sup-
ported by the Pollino National Park. We wish to
thank Chiara Misin, Monica Viapiana, Sabina Spada
and Roberta Anania for their help in field research.
We sincerely thank P. Chanin and D.J. McCafferty
for their useful revision of the manuscript.
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