Mammalian Biology 80 (2015) 385389

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Mammalian Biology
journal homepage: www.elsevier.com/locate/mambio

Short Communication

How harbor seals (Phoca vitulina) pursue schooling herring

Meike Kilian, Guido Dehnhardt, Frederike D. Hanke
University of Rostock, Institute for Biosciences, Sensory and Cognitive Ecology, Albert-Einstein-Str. 3, 18059 Rostock, Germany

a r t i c l e

i n f o

Article history:
Received 11 December 2014
Accepted 25 April 2015
Handled by Luca Corlatti
Available online 2 May 2015
Keywords:
Hunting strategy
Visual hunting
Prey
Foraging
Pinniped

a b s t r a c t
Only a few reports have described how pinnipeds hunt schooling sh. However, we had the opportunity
to systematically observe captive harbor seals hunting a school of herring in a shallow enclosure during
daylight. The seals actively pursued the sh, mostly changing from one side of the school to the other,
swimming in a supine position close to the water surface. They were often found swimming rapidly
down to the sh out of this position. When hunting in the vertical, the seals mostly adopted a dorsal
body posture relative to the school. They swam and attacked the school in a supine orientation when
approaching from the water surface and swam in a prone orientation when approaching from below.
They even maintained this relative body position during turning movements. These phenomena suggest
that the seals were constantly keeping the school in their large dorsal visual eld, which favors visual
hunting during daylight and in clear waters. When interacting with the school, the school mostly split
asymmetrically, and the seals were following preferably a smaller number of sh afterwards. Successful
prey catch was only observed, when a small group or a single herring had been separated, thus the seals
probably avoided the confusion effect of the school. In conclusion, we provide detailed insight in seals
pursuing schooling prey, thereby extending previous reports and conrming speculations as well as brief
observations.
2015 Deutsche Gesellschaft fr Sugetierkunde. Published by Elsevier GmbH. All rights reserved.

Harbor seals hunt benthic as well as pelagic sh individually and

in schools (Bowen et al., 2002; Marshall, 1998). From the few studies available, we know that, when hunting schooling sh, harbor
seals attack the edges of the school most likely in an attempt to
separate single sh or small subunits (Bowen et al., 2002; Zamon,
2001). Seals often seem to be engaged in active pursuit, during
which they constantly keep contact with the school and shift from
one side of the school to the other (Bowen et al., 2002). Zamon
(Zamon, 2001) document behavioral differences of adults and pups;
adult harbor seals ingest a mouthful of sh from the edges of a
school whereas pups swim through the school and are seen darting on single sh that were separated from the school. Harbor seals
also hunt schooling sh close to the sea oor which seems to constrain sh to escape (Olsen and Bjorge, 1995). The same effect could
be achieved by chasing sh towards the water surface as described
for separated sand lance (Ammodytes dubius) (Bowen et al., 2002).
These few insights in harbor seal foraging behavior were
obtained with the help of animal-borne video systems (Bowen and
Harrison, 1996; Bowen et al., 2002; Marshall, 1998), developed in

Corresponding author. Tel.: +49 381 666971914; fax: +49 381 666971935.
E-mail addresses:
meike kilian@web.de (M. Kilian), guido.dehnhardt@uni-rostock.de (G. Dehnhardt),
frederike.hanke@uni-rostock.de (F.D. Hanke).

the 1990s (Davis et al., 1992, 1999; Marshall, 1998), or by observations from vessels and land (Middlemas et al., 2005; Wright et al.,
2007; Yurk and Trites, 2000; Zamon, 2001). However, these documents are rare and allow only glimpses on the foraging behavior.
This is due to two main challenges associated with studying foraging in marine mammals. One challenge is that seals hunt under
water, an environment, which is still relatively inaccessible to
humans despite advances in modern technology. The other challenge refers to seals being highly mobile species. To successfully
monitor their foraging behavior thus requires techniques that do
not impose spatial constraints and preferably allows observing the
seal-school-unit in total.
In the present study, we had the unique opportunity to characterize the foraging behavior of harbor seals hunting schooling sh
in greater detail. We could systematically observe a group of nine
captive harbor seals (327 years), which are kept for scientic reasons under semi-natural conditions in a large seawater enclosure
(36 m depth, 60 m in length and 30 m in width) separated from
the open sea only by a net (mesh size 5 5 cm) at the Marine Science Center, Rostock, Germany. All seals were born in zoos and had
only come into contact with living sh such as atsh, eel and herring after moving into the seawater enclosure in summer 2008. In
spring 2009, a school of herring (Clupea harengus) with more than
1000 sh entered the seals enclosure and stayed for four weeks
from mid-April to May. During this time, the seals were observed

http://dx.doi.org/10.1016/j.mambio.2015.04.004
1616-5047/ 2015 Deutsche Gesellschaft fr Sugetierkunde. Published by Elsevier GmbH. All rights reserved.

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M. Kilian et al. / Mammalian Biology 80 (2015) 385389

Table 1
Ethogram used to analyze the video recordings listing all behaviors documented and their denitions as well as the results listing the frequency with which the behavior
occurred as number of total events N and in % (within a category).

Behavioral components of prey pursuit

Absolute body position
Prone
Supine
Side
Relative body position
Dorsal
Ventral
Lateral
Turning movements
Unchanged
Changed
Effect of an approaching seal on school
Splitting
Symmetrical
Asymmetrical
Following
Following smaller subunit
Following larger subunit
Following a single sh
Not following
Acceleration
Acceleration on small subunit
Acceleration in large subunit
Acceleration on single sh
Acceleration when others accelerated
Acceleration with no apparent reason
Prey capture

Denition
Body position absolute in the water column
Back pointing towards the water surface
Belly pointing towards the water surface
A ipper pointing towards the water surface
Body position relative to the school
A seals back pointing towards the school
A seals belly pointing towards the school
A seal pointing with one side, tail or snout towards the school
A change of absolute and/or relative body position during ascent and descent
A seal keeping its relative body position during turning
A seal changing its relative body position during turning

The two resulting parts of the school were approximately the same size
A different number of individuals was estimated in two or more subunits
A seal following sh after splitting
A seal following the smaller sh subunit after splitting
A seal following the larger sh subunit after splitting
A seal following a single sh after splitting
A seal did not follow sh after splitting
A seal was accelerating
A seal was accelerating when following a small sh subunit
A seal was accelerating when following a large sh subunit
A seal was accelerating when following a single sh
A seal was accelerating when another seal accelerated
A seal was accelerating with no apparent reason
A seal was successfully catching a sh

while hunting throughout the day. Two of the seals exclusively fed
on herring captured from the school. All other seals occasionally
hunted but also participated in the scientic experiments and routine training activity, during which they received sh rewards. The
behavior of the hunting seals during daylight hours was recorded
from a top view video camcorder (Canon XL1S; Canon Deutschland GmbH, Krefeld, Germany) that captured the seal as well as the
school. On the basis of an ethogram (Table 1) that was developed
during preparatory observations and that focused on behavioral
components of pursuing the sh and on the effect an approaching
seal has on the school, we analyzed 75 min of high quality video
footage during which six of the altogether nine seals and the school
of sh could be observed throughout the entire water column of
the enclosure. The depth within the water column, at which a specic behavior occurred, was documented and categorized as at the
water surface in cases where at least one part of the seals body was
seen out of the water, down to approximately 1 m depth when
the seal was below the water surface, and deeper than approximately 1 m. The 1 m transition point was estimated on the basis of
the height of a seal and took some uncertainty in the determination
of the water depth from above into account.
In contrast to previous studies, we were able to continuously
record six of the nine hunting seals during daylight hours and to
systematically document the foraging behavior over a large area
with full view on the seals and the school together. Our goal
was to generally extend the existing knowledge of harbor seals
hunting schooling sh and to specically describe the behavioral
sequence occurring during the pursuit of a school of herring. We
were interested in how the seals were positioning themselves absolutely in the water column and relatively to the sh, which might
allow assessing the sensory modality the seals were predominantly
using. Our hypothesis was that the seals mainly hunt visually in
the shallow enclosure in which light penetrates up to the bottom. Furthermore we wanted to test the hypothesis that the school
breaks into subunits by an approaching seal and that the seals are
predominantly following a smaller number of sh afterwards. In
this context, we hypothesized that attacks occur mostly on small

In %

609
1066
179

32.8
57.5
9.7

329
41
1484

17.8
2.2
80.0

327
138

70.3
29.7

91
318

19.3
80.7

176
54
29
59

55.0
17.0
9.0
19.0

87
43
22
9
48
0

41.2
20.4
10.4
4.3
22.7
0

subunits and single shes, which would document the adaptive

value of breaking a school into subunits or separating single sh.
Our analysis revealed that, during hunting, the six recorded seals
were moving through or with the school and only rarely remained
stationary. Their absolute body position was signicantly correlated with water depth (2 = 11,770.5, df = 8, p < 0.0001, Fig. 1).
Overall, the seals were swimming in a supine orientation in 57.7%
of the time (n = 1066, 2 = 636.7, df = 2, p < 0.0001). However, when
100
90
80

N=303
N=709

70
N=294

frequency (%)

Documented behaviors

60
50
40
30

N=123
N=183

20
10

N=63
N=100

N=42

N=37

0
at surface

< 1m

> 1m

absolute body position depending on depth

Fig. 1. Orientation of the seals in the water. Absolute body positions adopted by
the seals in the water column depending on water depth. The absolute body position (prone, supine, or side) was documented depending on the depth (at
the water surface, down to approximately 1 m and deeper than approximately
1 m), at which it was performed within the water column. Frequency is plotted as
percentage of total events, absolute numbers are indicated at the respective bars.
Black bars indicate the frequency of adopting the body position prone, gray bars
the frequency of the position supine and white bars the frequency of the position
side.

M. Kilian et al. / Mammalian Biology 80 (2015) 385389

387

N=914

50
45
40

frequency (%)

35
30
N=460

25
20
15
10

N=146

N=126

N=110

N=57
N=23

N=12

N=6

0
prone

supine

side

absolute and relative body position to school

Fig. 2. Orientation of the seals in the water and relative to the school. The absolute body position of the seal within the water column as well as its orientation relative to
the school are documented. Concerning the relative body position the school could be either ventral (black bars), dorsal (gray bars) or lateral (white bars) of the seal.
Absolute body positions are as in Fig. 1. Insets give examples of the respective absolute and relative body positions.

the seals were swimming deeper than approximately 1 m within

the water column, they changed their absolute body position into a
prone orientation in 75.2% of the time (n = 303). If the body position of the seals relative to the school was analyzed, it appeared
that in 80% of the cases the seals swam slightly lateral orientated
to the school (n = 1484, 2 = 1887.4, df = 2, p < 0.0001, Table 1). In
the majority of cases being laterally orientated, the snout was
pointing towards the school. When the absolute as well as the relative body position were taken into consideration, the seals swam
in 49.3% (n = 914, 2 = 3487, df = 8, p < 0.0001, Fig. 2B) of the cases
in a supine orientation slightly lateral relative to the school.
When interacting with the school, the seals swam constantly
ascending and descending but always kept contact with the school
during these turning movements. In 70.3% of the time, the seals kept
their relative body positions unchanged towards the school of herring by changing their absolute body positions (n = 327, 2 = 76.8,
df = 1, p < 0.0001, Table 1). Analyzing the turning movements, during which the seals did not change their body position relative to
the school, in detail, the seals mostly turned from a supine to a
prone position during descent (98.3%, n = 232) and from a prone
to a supine position during ascent (98.9%, n = 90), indicating that
the seal was orientated dorsal relatively towards the school of
sh.
An approaching seal initiated a splitting of the school (Table 1).
We documented an asymmetrical split of the school in 78% of the
cases (n = 318), thus signicantly (2 = 126, df = 1, p < 0.0001) more
often than a symmetrical splitting. When the school split asymmetrically, a seal was signicantly more likely to follow a smaller
sized group (55.0%, n = 176) than a bigger sized group (17.0%, n = 54)
or a single sh (9.0%, n = 29; 2 = 143.3, df = 2, p < 0.0001). Only
on a few occasions, the seal stopped interacting with the school
after it had split (19%, n = 59). We could observe the seals accelerate while interacting with the school of herring. They started
accelerating from right below the water surface (59.7%, n = 126)
and from deep in the water (38.9%, n = 82). In 41.2% (n = 87) of
all documented acceleration instances, a seal accelerated when

following a small group of sh, thus signicantly (2 = 89, df = 4,

p < 0.0001) more often than when following a big sh unit, a single sh, when another seal accelerated or without apparent reason
(Table 1). On the video footage, we did not record a single successful prey catch event. From observations that took place besides
recording we know, however, that the seals were catching prey,
especially when a small group of sh or a single herring had been
separated. Obvious systematic cooperative feeding such as prey
herding was not documented; however, we recorded opportunistic
behaviors such as kleptoparasitism, stealing sh caught by conspecics.
The analysis of the seals foraging behavior revealed that, when
the seals were interacting with the school of herring, they were
in most instances in motion and actively pursued the sh. They
always kept contact with the sh and moved from one side to
the other or from below the school to above, comparable to what
has been briey reported for wild seals hunting sand lance before
(Bowen et al., 2002). Our result of the seals being mostly lateral
of the school illustrates that they most often shifted from one side
to the other during active prey pursuit. During these movements,
a seal adopted a supine absolute body position in the majority
of the cases and was found mostly slightly below the water surface. We often observed the seals attacking the school out of this
position at the water surface. Yurk and Trites (Yurk and Trites,
2000) also observed seals to wait for salmon at the water surface in a supine position, however, they did not describe how
the seals behaved when attacking prey. In contrast, when hunting
sand lance (Bowen et al., 2002), seals were actively ascending rst
and then descending on the school. Active pursuit also took place in
the vertical with the seals ascending and descending continuously.
During these instances, the majority of the sh was mostly dorsal of the seal irrespective if the seals were residing at or slightly
below the water surface or deeper in the water column. This was
achieved by a change of the absolute body position in correlation
with water depth; while swimming close to the water surface, the
seals swam in a supine position whereas they adopted a prone

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M. Kilian et al. / Mammalian Biology 80 (2015) 385389

position at several meters depth. The relative body position dorsal

of the school was maintained during turning movements; the seals
mostly kept their body position unchanged relative to the school
irrespective of whether they were ascending or descending.
The way the seals were pursuing the sh in the horizontal
or vertical harmonizes with visual prey detection and pursuit
as hypothesized. Harbor seals possess a large dorsal visual eld
(Hanke et al., 2006). Consequently, the seals can permanently
observe the sh when being supine close to or at the water surface or when generally keeping the sh dorsal of the body. Visual
prey pursuit in seals has been proposed by various authors before
(Bonnot, 1932; Davis et al., 1992, 1999; Hobson, 1966; Lavigne
et al., 1977; Levenson and Schusterman, 1999), which was, however, solely based on the observation of seals attacking prey from
below when hunting pelagic sh (Bonnot, 1932; Davis et al., 1999;
Hobson, 1966), a behavior that we could also document in our
study. This way, the seals seem to silhouette their prey against the
bright water or under-ice surface. It is important to mention that
in all the studies we have referred to, including our own, light was
richly available enabling the seals to rely on vision while hunting.
However, seals often encounter conditions when diving deep, in
turbid waters or at night, which do not enable them to use their
visual system. This lead many authors to question visual hunting.
Especially as it was previously shown that visual acuity is drastically reduced by the presence of dissolved particles in the water
(Weiffen et al., 2006) or by decreased ambient luminance (Hanke
and Dehnhardt, 2009; Hanke et al., 2009; Schusterman and Balliet,
1971). Under these circumstances, harbor seals might rely on their
hydrodynamic or acoustic sense (Dehnhardt et al., 1998). Generally, hunting has to be considered a multimodal process, but our
results underline, that vision is indeed involved in this process.
The seals, we observed hunting herring, were constantly moving with or diving through the school and changed their behavior
between breaking the school and following the trailing edge continuously. In contrast, Bowen et al. (2002) reported two tactics that
seals used while foraging on sand lance: in most cases, the seals
seem to have attempted at separating small number of shes and
were attacking isolated shes. In a few instances, the seals moved
with the school and tried to catch sh from the trailing edge. In
our study, we could not clearly distinguish between these two tactics. We do not think that this difference is due to the different
sh species being hunted as overall the seals behaviors hunting
sand lance or herring are similar. Instead this difference might be
explained by the fact, that we could oversee a large area, including
the whole school and the seal, in contrast to the more restricted
visual eld of a critter cam, which might render the interpretation
of behavioral sequences more difcult.
Recording the school-seal-unit in total allowed us to obtain data
showing that the school indeed splits mostly asymmetrically by an
approaching seal. And, moreover, the seals in return followed the
smaller subunit in most instances. Accelerations occurred mainly
when following a small group of sh in line with our hypothesis.
By decreasing the number of individuals, the seals avoid or at least
reduce the confusion effect of the whole school (Krause and Ruxton,
2002), which is probably the prerequisite for successful prey capture. And indeed, only when chasing a small number of sh, the
seals were capturing sh. Unfortunately, we could not record prey
capture events but we observed the seals catching sh in various instances especially in a small enclosure in which a subunit
of the school used to settle. The reasons why the seals could not be
recorded catching sh in the main foraging area remain unknown.
The fact that most seals, except for two individuals, continued to
work and thus were fed besides hunting might have reduced their
motivation to actually catch a sh. However, we could not nd any
systematic differences between the seals that got additional sh
and those exclusively feeding on wild herring. In this context, it

might also be relevant to recall the fact that the seals observed
during the course of this study had only come into contact with living sh the year before the school entered the enclosure and thus
might not have been expert hunters although again two seals seem
to have been efcient enough to rely solely on sh intake from the
school.
The question arising now is whether our results are representative for seals in general, meaning if the behaviors we documented
could also be observed in seals in the wild. There are several apparent differences between the conditions encountered in our study
and wild seals hunting in their natural habitat. First, the seals we
observed in this study had limited experience in catching living sh
as just mentioned. Thus, they might be less experienced in hunting
than wild seals of the same age, which could result in different foraging strategies (Zamon 2001). Experience and learning might to
some extent be involved in hunting as Bowen et al. (1999) observed
that harbor seal pups were accompanying their mothers on foraging dives. The authors speculated that the pups learnt where and
what the mother was foraging and thus maybe also learnt how
to pursue and capture sh. Second, the seals hunted in very shallow waters of less than 6 m depth thus almost all behaviors can
be considered as surface behaviors. Wild harbor seals also perform
shallow dives during feeding, sometimes even shallower than 4 m
(Lesage et al., 1999), however, in most foraging dives, they reach
greater depths (Bjorge et al., 1995; Boness et al., 1994; Gjertz et al.,
2001; Tollit et al., 1998). However, harbor seals have to hunt under
diverse conditions, and we consider the conditions encountered
in our facility as one specic example out of many. It is evident
that every condition poses different demands on the seals. As an
example, the seals might use different senses for prey detection and
capture depending on the foraging depth that correlates with visibility in most waters. In return, the usage of different senses could
lead to different hunting strategies. Nevertheless we are condent
that the behaviors we observed belong to the natural repertoire
of the foraging behavior of harbor seals as the seals we observed
showed behaviors that have already been documented for wild
seals, however hunting different prey or only reported anecdotically, before. It is generally remarkable that the foraging behavior
of seals hunting in shallow waters shows some similarities to when
they hunt in deeper water.
In conclusion, we were able to obtain valuable information
about the foraging behavior in shallow waters of a group of harbor
seals living under semi-natural conditions. By continuously lming
seals hunting herring from a global perspective, we could extend
existing knowledge of harbor seal hunting behavior. Two aspects,
the seals relative body position to the school and their preferred
position before attacking, could be added in favor of visual hunting under light rich conditions. Moreover, we showed clearly that a
school indeed breaks into subunits by an approaching seal and that
the seals are more likely to follow a smaller subunit afterwards.
Acknowledgements
We express our gratitude to T. Coppack and J. Krieger commenting on a draft of this paper. This study was supported by a grant
of the VolkswagenStiftung to GD. The experiments are in line with
the current German law on the protection of animals. We thank the
subject editor and an anonymous reviewer for their helpful reviews.
The funding sources were not involved in any stage of the study
starting with study design, collection, analysis and interpretation
of the data up to the writing of the manuscript.
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