The effect of the river area and salmon

( Oncorhynchus nerka)
availability on the variation of the
proportion of bears (Ursus arctos) and their
hunting techniques
Introduction

Hunting behaviour in bears has been found to be affected by many different factors such as
foraging behaviour, food availability, and competition between individual bears. As foraging
opportunities change across landscapes, it is critical for individual bears to gain an understanding
of these changes and adjust their behaviours accordingly to maximize their energy intake. This
adjustment of behaviours can be a major advantage for ecologists to be able to make predictions
regarding the behaviour of bears. For example, assuming that brown bears (Ursus arctos) are
aware of the foraging opportunities across the landscape and adjust their behaviours based on the
availability of salmon in their area, one of the predictions could be as follows: Brown bears will
show fidelity to stream neighbourhoods due to the presumed benefits that follow with familiarity
with stream characteristics and the salmon dynamics (Wirsing et al. 2018). Different hunting
techniques are used by brown bears to acquire salmon. Brown bears compete to occupy the best
cites until they catch fish (Stringham 2012). This type of stand and wait technique can be
classified as a passive way of hunting. Bears can also walk or run through creeks to catch fish
(Stringham 2012). This type of technique can be classified as an active way of hunting.
Similarly, competition between individual bears can follow one of the two competition patterns:
exploitative competition or interference competition (Amarasekare, 2003). Exploitative
competition occurs when individuals remove resources from a common pool while interference
competition is when the individual directly prevents the foraging of others (Amarasekare, 2003).
It is predicted that the abundance and diversity of prey are affected by the inter and intraspecific
competition in the consumer’s environment such that the low abundance of prey is intensified by
the effects of competition

(Chesson, 2000).

Changes in salmon availability and intraspecific competition between individual brown bears for
the intake of food are among the factors that affect the hunting behaviours of the brown bears.
Factors such as the need to accumulate fat in brown bears leads them to hunt lipid and protein-
rich salmon that are located in frequent salmon streams (Ben-David et al. 2004).
OneadvantageofstudyingtherelationshipbetweenBrownbearsandsalmonisthatit can be easily
observed in less typical areas such as McNeil River that is a migratory bottleneck area for the
salmon where large quantities of bears come together and feed on them (Wirsing et al. 2018).

Studies have shown that female brown bears did not consume salmon while being accompanied
by their young due to high risks of infanticide while aggregating along salmon streams (Ben-
David et al. 2004). Intraspecific competition between male brown bears is what causes
infanticide, as killing the female’s offspring will increase the chance for males to father the
female’s next offspring (Steyaert et al. 2013). Therefore, female brown bears are faced with a
trade-off in systems where meat resources are spatially aggregated in the late summer and fall
(Ben-David et al. 2004).
The conduction of this study will help with identifying the specific interactions seen in brown
bears and the effects they have on their hunting behaviour. Additionally, the effects that the
abundance of salmon has on the close relationships between brown bears will be addressed.

In order to analyze how hunting behaviour is affected by the variation in salmon availability, in
this study we hypothesize that brown bears will alter their hunting behaviours when there is high
salmon availability due to lower intraspecific competition. We predict that the proportion of
Brown bears (Ursus arctos) in each river area will follow an equal distribution pattern when
salmon availability is high due to a decrease in competitive

interactions and hunting techniques will change from active to passive when salmon availability
is high.

Methods

Site description: The study was carried out in Brook Falls (58.5550° N, 155.7915° W) located in

Katmai National Park and Preserve, Alaska, USA. The topography of the area is rolling hills

forests and freshwater fall. Local vegetation includes forested areas and wet soils near the water

source. The salinity of the water is less than 1000ppm. The data used in the study was photos

collected three times a week, one from each morning(5am-10am), midday(10am-3pm), and

evening(3pm-8pm), between late June and early November from 2015 to 2020 respectively. The

average weekly temperatures ranged from -0.8C–18.3C. The average daily precipitation ranged

from 0.0mm–9.9mm and the average weekly wind ranged from 2.0mps-6.9mps. The study site
was previously used for hunting, fishing, and studying natural resources. Brooks Falls is the

water body located on Brooks River. The camera is set up directly beside a small waterfall (size

of area) that acts as a temporary barrier to migrating Sockeye salmon (Oncorhynchus nerka)

which leads to a high density of fish during spawning.

Study design: Three photos of Brown bears (Ursus arctos) were taken each week, one from each

morning, midday, and evening. The independent variable was the annual escape of Sockeye

salmon in each of the study areas (falls, ripples, rapids, flow, pool). The proportion of Brown

bears in each area implementing certain hunting techniques was the dependent variable. The

number of replicates was 70 photos (randomly selected from 300 database). 14 photos were then

randomly selected using a random number generator. Each year was one block therefore a fully

randomized block sampling design was used.

Sampling procedure: The sample size was 300 photos. A single photo was one observational

unit. A LiveCam was set up at a viewing platform by the waterfall. By default, the samples

collected included water bodies but depending on the specific photo, divisions could be observed

regarding falls, ripples, rapids, flow, and pools. Samples were collected in the morning, midday,

and evening.

Sample processing: The photos were taken by volunteers. The organization then archived these

photos, which were selected by the BIOL313 teaching team, and randomly selected by Group33

Bears. The BIOL313 teaching team used certain criteria such as: at least one adult bear present in

each photo, three photos per week one from each: morning, midday and evening, sufficient

context to determine location of bear, and be clear enough to evaluate bear condition and

behaviour.

The total annual escapement was the unit of measurement for salmon.
Different fishing styles of bears include standing on top of Brooks Fall and waiting for salmon to

jump high enough to catch in their mouth, sitting underneath the fall and waiting for salmon to

swim to them, dash and grab (pin salmon to river bottom), snorkeling (look for fish under water),

begging from others, and undetermined.

Data analysis: Summary statistics of this study reports annual variance, standard deviation, and

standard error from mean proportion of variation of bear’s location and hunting techniques with

respect to annual salmon escapement.

Results

In this section, trends and patterns between how salmon (Oncorhynchus nerka) availability will

impact the foraging behaviour of Brown bears (Ursus acrtos) will be stated. The first prediction

stated that the proportion of Brown bears in each river area would follow an equal distribution

pattern when salmon availability is high due to a decrease in the competitive interactions

between bears. In Fig.1-A, as the total annual salmon escapement (TASE) increased, the mean

proportion of bears (MPB) initially increased and peaked when TASE was 2218824. After this

point a gradual decline was observed in the MPB. In Fig.1-B, the MPB initially decreased as the

TASE increased, with the minimum point being 2218824 TASE, followed by a gradual increase.

In Fig.1-C, as TASE increased, the MPB increased. The peak point was 2218824 TASE,
followed by a sharp decrease and then an increase. In Fig.1-D, a general logistic decrease in the

MPB was observed as the TASE increased. Exceptions to this trend was at 2218824 and

4025802 TASE where an increase was seen in the MPB. In Fig.1-E, a general constant pattern

was maintained when the MPB was around 20-30%. A slight decrease can be observed in 2018

as the proportion decreases, showing the shape of an upside-down bell curve. Fig.1-D and Fig.1-

E are similar as they reach a 20% distribution as TASE is observed to reach a maximum, while

the remaining graphs do not follow such pattern.

From Fig.1-A,B,C as TASE increased, fluctuation was seen in the MPB. In these areas, the MPB

did not follow an equal distribution.

In Fig.1-D and Fig.1-E, as TASE increased, MPB approached 20% which resembles an equal

distribution.

The second prediction stated that the proportion of brown bears utilizing hunting techniques to

obtain salmon will change from active (dash and grab,stand and wait, and snorkeling) to passive

(sit and wait and begging), when salmon availability is high. There was an inconsistent trend

observed across different hunting techniques with no discernible pattern (Fig.2-A,B, C, D, E, and

F). Hunting techniques changed from active to passive as there was an increase in begging

(Fig.2-B) with a peak of 2830212 and decrease in snorkeling (Fig.2-C) when TASE increased,

with a minimum of 1677228. However, stand and wait MPB increased (Fig.2-D) with a peak of

2830212 while dash and grab and sit and wait (Fig.2-E and Fig.2-A) stayed fairly constant,

showing an opposite pattern as to what was predicted.

A

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Figure 1. the mean proportion of Brown bears (Ursus arctos) in various locations (A: falls, B:

Rapids, C: Ripples, D: Flows, E: Pools) as a function of total annual salmon (Oncorhynchus

nerka) escapement between the years 2016 and 2020. Bars represent +/- standard error of mean

proportion of bears annually and the data is composed of 14 replicates per year.
A
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Figure 2. Relationship between the total annual salmon (Oncorhynchus nerka) escapement

between 2016-2020 and the mean proportion (%) of bears (Ursus arctos) implementing various

hunting techniques (A: sit and wait, B: begging, C: snorkeling, D: stand and wait , E: dash and

grab, F: undetermined). Bars represent +/- standard error of mean proportion of bears annually

and the data is composed of 14 replicates per year.

Discussion

In the flow and pool areas, a fluctuating trend in which mean proportion of bears approached

20% as total annual salmon escapement increased was observed. In the rapids, ripples, and falls

the mean proportions were not converging to an equal distribution pattern of 20% as salmon

escapement increased. Instead, there was a fluctuating pattern between the proportions of 5%-
35% (Fig. 1). The hunting techniques of sit and wait and dash and grab had a general constant

mean proportion while begging and stand and wait showed an increase in mean proportion of

bears as annual salmon escapement increased. Also, a sharp decrease in the mean proportion was

observed for snorkeling (Fig. 2).

The ideal free distribution theory (IFD) predicts the distribution of animals in patches that

compete for resources (Fretwell and Lucas 1970). If all competitors can move without cost

between patches, have equal ability to acquire food, and have detailed enough information of

their competitors’ distribution and the food resources available to them, each competitor is

expected to go to a patch providing it with the highest energy intake (Smith 1982). Under

continues input conditions where a resource is consumed right after it is inserted into the patch

(Parker and Sutherland 1986), the IFD predicts that the number of competitors in each patch

would be proportional to the resource input rate (Hakoyama 2003). This prediction almost held

true for the river areas in this study. First, the fluctuation of yearly mean proportion of bears with

respect to annual salmon escapement showed a sine shape variation for all areas (except rapids

and falls that are bell shape curves). For example, the mean proportion for pools started at 26%,

then increased to 28%, followed by decreasing to 18%, and then increased again to 32%, and

ended up to 20% for the years 2016,2017,2018,2019, and 2020 respectively, which simulates a

sine shape variation (Fig1).

On the other hand, in the flow and pool areas, equal distribution in the mean proportion of bears

was observed as salmon escapement increased. However, for other areas it was hard to derive

such a conclusion as the extent of the variation per year are more scattered specifically for falls

and rapids. Alternative factors such as social dominance can be affecting the mean proportion of

Brown bears in river areas. It was found that in each small stream occupied by a socially
dominant bear, subdominant bears would directly be displaced either aggressively or by

avoidance. Additionally, to minimize social interactions, the subdominant bears use a smaller

fraction of each stream and carry their prey to a further location for consumption (Gende and

Quinn 2004). Bear size also influences fishing locations as bigger bears can fish in waist-deep

rushing water that can carry smaller bears away (Luque and Stokes 1976).

The pairs of total mean proportion of (active, passive) hunting techniques per year as

cumulatively derived from the graph (Fig. 2, parts a, b, c, d, and e) was as follows: {(45,32),

(32,23),(25,35),(28,33),(41,34)}. This trend did not show any meaningful relation between the

fluctuation of hunting techniques per annual salmon escapement. Thus, the proportion of bears

utilizing hunting techniques did not change from active to passive when the salmon availability

increased.

Classification of hunting techniques was difficult because of using still pictures. Additionally,

various factors (for instance no distinction made between the behaviour of male or female bears)

not considered in this study could be a source of error. In previous studies, it was found that

females with cubs were highly intolerant of other bears (Egbert and Stokes 1976) which in turn

can affect the location selection of adolescent males and females for the means of reducing

disturbance. Age of bears is an important factor affecting hunting techniques. A study on the

fishing behaviour of Alaska brown bears showed that bears used 9 to 28 techniques and old bears

used fewer techniques when compared to the younger (Luque and Stokes 1976).

While had the greatest mean proportion, the undetermined hunting technique was a source of

error as it could include any of the other five hunting techniques that were undetected.

For determining how variation in salmon availability affects hunting behaviours, the mean

proportion of bears in each area and changes in hunting techniques from active to passive, both
when salmon availability was high, was examined. An equal distribution pattern was only seen in

the areas of pools and flows. Additionally, there was no relationship observed between the

transition of hunting techniques from active to passive.

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