Journal of Zoology.

Print ISSN 0952-8369

Influence of watercourse depth and width on dam-building

behaviour by Eurasian beaver (Castor fiber)
G. Hartman & S. Törnlöv
Department of Conservation Biology, Swedish University of Agricultural Sciences, Uppsala, Sweden

Keywords
Castor fiber ; beaver; dam building; dams;
Sweden.
Correspondence
Göran Hartman, Department of
Conservation Biology, Swedish University
of Agricultural Sciences, Box 7002, 750 07
Uppsala, Sweden. Tel: +46 (0)18 672266;
Fax: +46 (0)18 673537
Email: Goran.Hartman@nvb.slu.se
Received 8 November 2004; accepted 7 April
2005
Abstract
Dam-building behaviour is a distinctive feature of beavers; yet they often occupy
streams or sections of streams where they do not construct dams. This study was
designed to determine how stream depth and stream width influence dam-building
behaviour. Stream depth and stream width were measured at 39 beaver lodges,
which did not have dams associated with them and at 74 dams. The mean water
depth at the lodges was 1.16 m (range = 0.55–2.0 m), while the mean water depth at
the dam sites was 0.36 m (range = 0.10–0.85 m). The stream width was greater at
lodge sites (mean = 11.15 m, range= 1.0–40 m) than at dam sites (mean = 2.5 m,
range= 0.5–6.0 m) A discriminant analysis classified 93% of the sites correctly as
either lodge or dam sites.

doi:10.1111/j.1469-7998.2005.00025.x

Introduction
Both extant beaver species, the Eurasian beaver Castor fiber
and the North American beaver C. canadensis, are semi-
aquatic animals that occur in all kinds of inland waters.
They are well known for their ability to change their
surroundings to fit their needs and have therefore been
considered as a prime example of an ‘ecological engineer’,
i.e. an organism that strongly affects its surroundings by its
activity rather than by its mere presence (Jones, Lawton &
Schachak, 1994). ‘More than any other animal except hu-
mans, beavers geomorphically alter the landscape through
their dam building and related activities’ (Butler, 1995).
The beaver-created impoundments increase the retention
of sediments and organic matter, alter the riparian zone and
modify the aquatic habitat by changing local conditions
from lotic to lentic. Beaver dams can increase the hetero-
geneity of the landscape by creating a mosaic of aquatic and
semiaquatic elements, causing short-term as well as irrever-
sible changes (Naiman, Johnston & Kelley, 1988). The scale
of impact may be substantial. For example, in Quebec,
beaver dams affect as much as 30–50% of the total length
of second-to fourth-order streams (Naiman & Melillo, 1984)
while in Poland Zurowski (1989) reported 24 dams in a
1.3 km reach.
Beavers may occur in all kinds of lentic and lotic environ-
ments, from small seepages and ponds to large rivers and
lakes. However, beavers build dams only in small streams,
meaning that many beaver settlements lack dams. In a study
by Zahner (2001), only 19% of all beaver colonies in Bavaria
had dams. Nevertheless, dam-building behaviour is so
closely associated with beavers that habitat selection and
dam-site selection sometimes have been regarded as syno-
nyms (McComb, Sedell & Buchholz, 1990; Barnes & Mallik,
1997; Suzuki & McComb, 1998). Habitat selection studies
have actually shown that beavers prefer deeper waters,
which meet their demands without alteration (Beier &
Barret, 1987; Dieter & McCabe, 1989; Hartman, 1996).
Several studies indicate that Eurasian beavers settle on
smaller tributaries later in a colonizing process, when all
preferred sites on larger streams and rivers are occupied
(Semyonoff, 1951; Curry-Lindahl, 1967; Zurowski & Kas-
perczyk, 1986; Pupinnikas, 1999; Gorshkov et al., 2002). It
has also been reported that C. fiber engage less in construc-
tion behaviour, including dam building, than C. canadensis
(Danilov & Kanshiev, 1983).
The observed difference in the presence of dams among
sites occupied by beavers will immediately lead to the
question: when do beavers consider a site suitable without
damming? To date, the information available has been is
expressed using the rough classification of streams into
orders (Horton, 1945; Strahler, 1957). Beaver impound-
ments are usually found in low-order streams, 1–4 (Naiman
et al., 1988), 1–3 (Snodgrass, 1997), but streams of the same
order may vary considerably in size and a single stream will
change along its course. Considering the impact that a
beaver dam will have on its surroundings, and the possible

Journal of Zoology 268 (2006) 127–131

c 2006 The Zoological Society of London 127
Influence of watercourse depth and width on beaver dam building
damage it may cause to for example roads and commercial
forests, it is of great importance to have more detailed
information on which physical features influence beavers in
terms of dam construction.
This study was designed to determine how water depth
and stream width influence dam-building activity.
Study area
Beaver dams and lodges were surveyed in two study areas.
One area is in the southern boreal zone within the Grimsö
Wildlife Research Area in south central Sweden (59140 0 N,
15125 0 E). The watercourses surveyed are a part of river
Sverkestaån and some of its tributaries. The river
Sverkestaån varies in width from a few metres to several
hundred meters when it forms lakes. The topography is
fairly flat and about 100 m above sea level. Overall, stream
gradients are 0.2–0.5%. The second area is in the subalpine
zone in the vicinity of the town Idre (61150 0 N, 12150 0 E) in
the southern most part of the Scandinavian mountain range.
The topography is hilly and varies between 450 and 650 m
above sea level. The waters surveyed are small tributaries to
the River Dalälven. The surveyed tributaries varied in width
from less than half a metre to about 15 m. The overall
stream gradients were 1–2%. The vegetation is similar in
both areas although richer in the southern area. Coniferous
forest dominates (Scots pine, Pinus silvestris, Norwegian
spruce, Picea abies). Bogs are numerous and farmland are
scarce. The dominating deciduous species is birch Betula
pendula, B. pubescens. Other species are less common i.e.
willows Salix spp., aspen Populus tremulus and alder Alnus
glutinosa, A. incana.
The beaver was exterminated in Sweden during the 19th
century, and was reintroduced in the 1920s. The population
is still expanding its range. The first two beaver colonies
appeared in the Grimsö area in the beginning of the 1980s.
The Idre area was most likely recolonized during the 1960s
or 1970s. The beaver populations in the areas have been
relatively stable in recent years.
Methods
With the previous knowledge that beavers build dams only
in smaller streams, we focused on determining stream depth
and width thresholds where beavers would cease to con-
struct dams. We compared stream depth and width at dams,
with depth and width at lodges not associated with dams.
Lodges without dams were used to reflect beaver sites that
are suitable without damming. Both active and inactive
dams and lodges were included, as our aim was to determine
original stream characteristics. Watercourses were surveyed
by foot and followed from their sources to where they joined
the larger river (Sverkestaån, Dalälven). A 48 stream-km in
the Idre area was surveyed in June 1998 and 1999 and a 52
stream-km in the Grimsö area was surveyed in August 1999.
Surveys were conducted in June and August to standardize
flow conditions and eliminate water-level variation caused
by spring and autumn floods and the normal summer low.
128
G. Hartman and S. Törnlöv
Stream width and depth were measured at all dams
and lodges that were unaffected by a downstream dam.
The stream by a dam or lodge was classified as unaffected
if shallow sections of stream with fast-flowing water were
found between the site and any downstream dam.
Stream depth was measured 1 m outside lodges, taking care
to avoid any beaver-excavated channels associated with
the lodge exit hole, and stream width was measured at the
lodge site.
At dams, stream depth and width were measured on the
downstream side and as close as possible to the dam.
Because water-overflowing dams may erode the streamside
and create a depression in the river bottom, measurements
were made where the river topography was unaffected by
the dam (within a few metres). Stream depth was measured
in the centre of the stream to reflect the maximum depth.
Additionally, we measured raising of the water level at
active, maintained dams to determine at what achieved
depth beavers would cease to increase water depth. Raising
of the water level was defined as the vertical distance
between water surfaces in front and behind dams. The
achieved depth was defined as the raising of water level
+depth downstream the dam. Measures were made by
using a rope with a weight to throw across the stream and a
folding rule.
Data were analysed using the Mann–Whitney U-test,
Pearson correlation and discriminant analysis.
Results
Exclusion of eight dams and six lodges, where site condi-
tions were affected by a downstream dam resulted in a total
of 74 dams and 39 lodges; 58 dams and 20 lodges from the
Idre area; and 19 lodges and 16 dams from the Grimsö area.
No evidence of beavers living in bank burrows was ob-
served, which was because of the flat banksides and/or rocky
soils in the study areas. All lodges were bank lodges (not
surrounded by water).
Data from both areas were pooled in the analysis since
the results overlapped substantially. The average stream
depth at lodges was 1.16 m (SD: 0.44, range: 0.55–2.00 m,
n = 39). The average stream depth at dams was 0.36 m (SD:
0.14, range: 0.10–0.85 m, n = 74). Water depth differed
significantly between dams and lodges (Mann–Whitney
U-test, Z=8.54, P= 0.0001). The average stream width
at lodges was 11.5 m (SD: 10.4, range: 1–40 m, n= 39). The
average stream width at dams was 2.5 m (SD: 1.1, range:
0.5–6.0 m, n =74). Watercourse width differed significantly
between dams and lodges (Mann–Whitney U-test,
Z =6.95, P= 0.0001).
Plotting depth and width at dam and lodge sites in the
same graph showed a clear separation (Fig. 1). Three
percent of the dams were built at sites with an initial water
depth of 0.7 m or more, and 5% of all lodges unaffected by a
downstream dam were built where the water depth was less
than 0.7 m. No dams were found at sites where the water
course was more than 6 m wide, and 96% of them were built
where it was less than this. In a discriminant analysis, 93%
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c 2006 The Zoological Society of London
G. Hartman and S. Törnlöv Influence of watercourse depth and width on beaver dam building

3 10
9
8
Watercourse depth (m)

7
2
6

Count
5
4
3
1
2
1
0
.2 .4 .6 .8 1.0 1.2 1.4
0 Water depth (m)
0 5 10 15 20 25 30 35 40
Watercourse width (m) Figure 3 Frequency distribution of achieved water depth (m), (initial
depth + raising of water level) by beaver dams (n = 48).
Figure 1 Watercourse depth and width by beaver dams (), (n = 74)
and beaver lodges unaffected by a downstream dam (), (n =39).
n = 48), and 68% of the dams resulted in a water depth
between 0.7 and 1.0 m (Fig. 3).
1.2
Discussion
1.0 Considering that the maximum depth where beavers had
built a dam (0.85 m) was equal to the average achieved depth
Raising of water level (m)

0.8
0.6
0.4
0.2
0.0
0.0 0.2 0.4 0.6
Initial water depth (m)
Figure 2 Relationship between initial water depth and raising of water
level at beaver dams (n = 48).
of the sites were correctly classified as either dam or lodge
sites. The analysis misclassified six of the lodge sites and two
of the dam sites. Weightings for lodge sites were: depth:
14.210, width: 0.031, and for dam sites: depth: 4.689, width:
0.024.
Watercourse depth and width were positively correlated
at dam sites as well as lodge sites (Pearson correlation,
r2 = 0.215, d.f. = 73, P= 0.0001, r2 = 0.162, d.f. =38,
P= 0.006).
When measuring raising of water level, 26 dams had to be
excluded because they were judged as not having been
maintained. The average raising of the water level was
0.46 m (SD: 0.21, range: 0.1–1.0 m, n = 48). Water raising
was negatively correlated to original depth (Pearson correla-
tion, r2 = 0.165, P= 0.0025, d.f.= 47, n = 48), (Fig. 2). The
achieved depth averaged 0.84 m (SD: 0.20, range: 0.4–1.28 m,
at dam sites (0.84 m), and only 5% of all lodges unaffected
by a downstream dam were built where the water depth was
less than 0.7 m, we interpret the results of this study as that
most beavers seem to consider a depth of roughly 0.7–1 m as
sufficient. The larger overlap in stream width than in stream
depth between dam and lodge sites indicates that a sufficient
water depth is most probably more important than the
achieved pond area. Although no beavers built a dam where
the stream width was greater than 6 m, this result is more
complicated to interpret because of the positive correlation
between watercourse depth and width.
0.8 1.0
According to the results of this study, beavers that settle at
sites with water characteristics below the above-mentioned
limits may be expected to build dams. This information may
be valuable when assessing damage risks as well as positive
impacts on the biodiversity of beaver impoundments.
It was not within the scope of this study to pursue the issue
of why beavers build dams and we did not collect data with
this in mind. Nevertheless, the results have some relevance to
the question, and we believe that in future studies the
proposed multipurpose role of dams must be taken into
consideration. It is hardly controversial to say that the overall
purpose of dam constructions is to improve local habitat
suitability and thereby increase the range of inhabitable
areas. It is, however, more difficult to pinpoint exactly how
beavers benefit from their dam constructions. Different dams
may have different purposes and therefore may be con-
structed in various ways and places. There is a consensus on
the purpose of a dam that elevates the water level at the lodge.
This so-called primary dam (Hodgdon & Hunt, 1966) keeps
the water, level high and stable, keeping the lodge entrance
under water and ensures sufficient water volume to hold a

Journal of Zoology 268 (2006) 127–131

c 2006 The Zoological Society of London 129
Influence of watercourse depth and width on beaver dam building
functional winter food cache (Hartman & Axelsson, 2004).
However, considering that a beaver colony will often main-
tain a number of dams within their territory, most dams will
not affect the water level at the lodge and obviously serve
other purposes. In a study from Minnesota, there were on an
average three dams per colony (Longley & Moyle, 1963) but
in other areas there may be up to 40 dams on one site (Müller-
Schwarze & Sun, 2003). Possibly, a secondary dam down-
stream the main dam serves to strengthen the main dam by
backing up water against it, but this has not been proved
(Longley & Moyle, 1963). In our study, only 13% of the
lodges had to be excluded from the analysis because they had
a dam affecting the water level, and only 10% of the dams
were excluded for the same reason. These data suggest that
the majority of dams in our study areas serve another
purpose. Secondary dams have also been proposed to in-
crease food availability by moving the shoreline closer to
stands of palatable trees. A study by Breck, Wilson &
Andersen (2003) comparing beaver herbivory between a
flow-regulated and a free-flowing river supports this explana-
tion. Possibly, this is the role of most dams in our study areas,
although this conclusion is contradicted by our notion that
water depth is more important than water width.
There are several explanations regarding why beavers do
not always impound waters that they inhabit. The first and
most obvious is that they simply do not have the capacity to
impound very large bodies of water. The second is that there
is an upper limit in the river size/stream gradient where the
force of the water will be too strong to allow the construc-
tion of a dam or that it would run a great risk of being
washed out by freshets (Naiman et al., 1988). In a study
from France, dam building ceased when the stream gradient
exceeded 1% (Erome, 1983) but in Germany beavers built
dams at a gradient of 2.5% (Schulte, 1989). In Oregon, dams
were found at gradients up to 4% (McComb et al., 1990).
The third explanation would be based on the view that
there is substantial cost involved in dam building, and
beavers would be expected to only build dams when they
find it absolutely necessary, i.e. if the water volume is
insufficient to meet their needs. Considering the possible
differences in the purpose of dams, sufficient water volume
and/or pond size might vary. This should be taken into
account in future studies of beaver dam-building behaviour.
Acknowledgements
We thank Peter Busher, Peter Collen and Frank Rosell for
providing constructive comments and suggestions for im-
proving the manuscript.
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