Ecography 31: 519
524, 2008

doi: 10.1111/j.2008.0906-7590-05477.x
# 2008 The Authors. Journal compilation # 2008 Ecography
Subject Editor: Kai Norrdahl. Accepted 4 March 2008

The beaver as an ecosystem engineer facilitates teal breeding

Petri Nummi and Anna Hahtola
P. Nummi (petri.nummi@helsinki.fi) and A. Hahtola, Dept of Forest Ecology, P.O. Box 27, FI-00014 Univ. of Helsinki, Finland.

Positive interactions are important in influencing communities. Facilitation is one such influence in which a species
makes the environment more suitable for others species. Ecological engineers are powerful facilitators as they physically
modify habitats in a way that alters resource availability. We studied the effect of beaver flooding on breeding success of a
dabbling duck, the teal. Our long-term (1988
2001) study was conducted in a boreal area where dabbling ducks face
resource limitation. We focused on two aspects of facilitation: resource enhancement and habitat amelioration. We found
that pond use by teal broods systematically increased upon beaver flooding. Beaver ponds harboured more resources, i.e.
aquatic invertebrates than undisturbed waters, and invertebrate abundance was reflected as enhanced teal brood density.
Beaver inundated shores were shallower than those of other waters, making their habitat structure more favourable to
ducklings. Consequently, teal brood mortality was lower in beaver ponds than in waters unaffected by beaver. In boreal
areas beaver facilitate teal breeding by enhancing both resource abundance and habitat structure.

For a long time, ecologists have emphasized factors such as

competition, predation and physiological stress, which
negatively affect species distributions and abundances
(Bertness and Callaway 1994, Machicote et al. 2004).
However, more recent research indicates that facultative or
positive interactions may be as important as other factors in
influencing populations and structuring communities (van
der Wal et al. 2000, Stachowicz 2001). One kind of positive
interaction, facilitation, occurs when one organism makes
the local environment more favourable for another species
(Bruno et al. 2003). Facilitation has mainly been shown in
plant communities, plant-animal interactions or intertidal
communities, and also recently among terrestrial vertebrates
(Machicote et al. 2004). Terrestrial vertebrates may facilitate other species in at least two ways: by behaviors
increasing e.g. feeding efficiency, and by physical modification and enhancement of the environment (Dickman 1992,
Jones et al. 1994). One form of physical modification is
ecosystem engineering (Jones et al. 1994, 1997).
Ponds and lakes in the vast area of boreal forest are
numerous, and so are their waterfowl (Mack and Morrison
2006). However, the per capita brood production is
typically low in boreal shield, although not necessary in
all boreal habitats world-wide (Nudds and Cole 1991,
Nummi and Poysa 1995a). Long term studies have revealed
that even if many ponds may sometimes contain pairs, a
great majority is devoid of broods year after year (Sjoberg
et al. 2000, Nummi et al. 2005). Moreover, experiments
have confirmed that many boreal ponds are indeed poor
duckling habitat, especially for dabbling ducks (Nummi
et al. 2000, Sjoberg et al. 2000, Gunnarsson et al. 2004).

Orians and Wittenberger (1991) stated that a suitable

habitat may need to contain a mixture of patches that
provides opportunities for all of the activities required for
successful reproduction. Ducks living in many boreal
habitats seem to lack highly suitable habitat during brood
rearing, so poor brood habitat is one potentially limiting
factor for boreal duck populations.
The lack of sustainable brood habitat applies especially
to boreal landscapes which lack one of their original
features, the beaver. Beavers were exterminated over a
century ago from most of their original range, especially in
Eurasia and also from Finland (Lahti and Helminen 1974,
Pollock et al. 1995, Nolet and Rosell 1998). Beavers act as
ecosystem engineers by creating and modifying ponds with
their dams (Jones et al. 1994). Via engineering, they can
directly or indirectly control the availability of resources to
other organisms, and increase habitat heterogeneity in
riparian landscapes (Jones et al. 1997, Wright et al.
2002). Earlier the effect of beaver activities was largely
unknown as modern research on different kinds of waters
began after the beavers were hunted out. Much of our
understanding of how creek and pond systems
especially
their littoral zones
function is thus often based on research
on the unnatural state of these habitats (Naiman et al.
1988).
Beavers have now either returned or are returning to
many parts of their original range, where they have been
absent for a century or more (Nolet and Rosell 1998). With
their dam building, beavers modify stream morphology and
hydrology thereby strongly influencing the structure and
dynamics of their habitats. These activities result in changes
519

in the function of the riparian zone and the stream channel,

and ultimately influence plant and animal community
composition and diversity (Naiman et al. 1986). It is of
special interest, therefore, to study how the engineering
resulting from returning beavers affect wetland ecosystems
and their species.
From the point of view of littoral animal species, the
essential features of beaver flowages include a wide area of
shallow water, the breakdown of inundated vegetation and
the increase in input of falling tree leaves. Vegetation
decomposition results in a release of nutrients that form a
base for a food web consisting of detritivores such as
chironomids and isopods (McDowell and Naiman 1986,
Nummi 1989). The increased production of insects and
other invertebrates benefit many vertebrate groups including fish, amphibians, birds and mammals (Grover and
Baldassarre 1995, Collen and Gibson 2001, Rosell et al.
2005).
Beaver ponds are a habitat much used by duck broods
(Renouf 1972, Nummi 1992), and teal pairs respond
promptly to beaver flooding (Nummi and Poysa 1997).
However, the processes behind this habitat use pattern are
not well known, and particularly there is a lack of
comparisons between beaver flowages and non-beaver
influenced ponds. Therefore, new information is needed
to adequately index breeding success and to more rigorously
test the full impact of beaver activity on duck production.
First, we studied the effect of beaver on pond occupation
by teal broods. Second, we explored factors affecting brood
habitat use, such as invertebrate abundance and habitat
structure. Finally, we examined how beaver affects the
success of teal broods. This is important since mere habitat
use is only an indicator of habitat quality. In order to
qualify habitats from the point of view of individuals one
needs to measure factors independent of bird numbers.
Breeding success is suitable for that because it measures
individual fitness (Fretwell and Lucas 1970, Wiens 1989).
Therefore we compared mortality of teal broods in beaver
flowages with that of undisturbed water bodies.

Methods
Site description
The study area was situated in a 39 km2 boreal watershed in
southern Finland (61810?N, 25805?E). All lakes and small
ponds with permanent water for the whole summer were
included, sole exeption being one near the Evo Game
Research Station which was subject to obvious human
disturbance. The shore types of the lakes range from
oligotrophic bog and forest without emergent plants to
more eutrophic types with lush stands of Equisetum spp.
and Typha spp. Apart from beaver disturbance, the water
conditions have only minor year to year variation (Nummi
and Poysa 1995a). The beaver species in the area is
introduced Castor canadensis which plays a similar ecological role as C. fiber although it may be a slightly more active
builder of lodges (Danilov and Kanshev 1983). Beaver
flowages of the area typically are formed by building a dam
at the outlet of a natural pond.
520

Brood data
To study the use of a pond by broods before and during
beaver settlement, duck data of 51 lakes were collected
during the breeding seasons 1988
2001. Of these, beavers
dammed nine during the study (one of these was a dammed
up stream). A survey was taken of broods four times per
season over the early-June to late-July period, one survey
approximately every second week. During each survey, a
point count was first made from the shore and after that the
lake was circled with a round count (Nummi and Poysa
1995a). The age of broods was determined following the
classification by Pirkola and Hogmander (1974). The
individual broods could be recognized based on the age,
and by the fact that in boreal wetlands the brood density is
low, and so is the number of broods in each wetland. The
averages of teal brood numbers for two years before beaver
occupation were compared to the brood numbers during
the two first years of beaver inundation. In a few cases only
data from one before or during year were available. For
each of the nine beaver ponds, there was one control pond
of a similar habitat type. The controls were the nearest
neighbours of the beaver ponds in a gradient representing
the structure of shore vegetation (Nummi and Poysa 1997).
The gradient was defined by a principal component analysis
based on vegetation descriptions of all the water bodies
(n 51) in the long term study area (Nummi and Poysa
1995a). Changes in brood numbers in each of the controls
were studied using exactly the same years that were used in
the appropriate beaver ponds.
For the brood mortality measurements, data were
collected from five extra beaver flowages in 2001 in
addition to the 51 lakes. The mortality of ducklings was
calculated as daily mortality rate (Mayfield 1961, Ringelman and Longcore 1982), which takes into account both
the brood size and the duration of the period over which the
mortality is calculated. The mortality rates can be obtained
only from relatively sedentary broods from which at least
two observations can be made. For comparison purposes,
one brood was used as a unit. Because mortality decreases as
ducklings get older we included only broods which were
seen for the first time at the age class I (B20 d of age).
Because mortality rate may vary even within the age class I,
we checked the average age of broods when seen for the first
time in beaver flowages and other waters; we found that
there were no difference between these groups (Z 
0.166, p 0.87, Mann-Whitney U-test). For comparison
of brood sizes in beaver flowages with other waters we used
almost fully grown ducklings (IIc-IIIa) which generally are
subject to low mortality (Orthmeyer and Ball 1990, Mauser
et al. 1994).
Invertebrates
Invertebrates were sampled during July with activity traps
(Murkin et al. 1983). The traps were made from 1-l glass
jars and funnels with an external diameter of 100 mm and
an internal diameter of 23 mm. Traps were placed in the
water on the bottom for 48 h, and 10 activity traps were
placed in each wetland. Traps containing fish were not used
in analyses (Elmberg et al. 1992). Traps were set parallel to

the shore, 1 m from the waters edge or where the water

depth ranged between 25 and 75 cm (details in Elmberg
et al. 1993). In total 11 beaver flowages and 26 undisturbed
ponds were sampled for both inverterates and duck broods
(1990: 2 flowages, 6 ponds; 1995: 2, 9; 1997: 2, 5; 2001: 5,
6). Different years could be used because from a 12 pond
set trapped yearly we know that apart from beaver
influenced variability, the permanent ponds retain their
characteristics from year to year (Nummi and Poysa 1995a,
Poysa et al. 2000).
Food abundance for each lake was indexed by using the
mean number of invertebrates per trap multiplied with their
respective length (Elmberg et al. 1993). Prey were assigned
to four size classes (0
2.5; 2.6
7.5; 7.6
12.5, and 12.5
mm) according to Nudds and Bowlby (1984, Table 2) with
small modifications according to Elmberg et al. (1993).
Food abundance in each of the 11 beaver flowages and
26 undisturbed ponds were compared to teal brood density
of the corresponding wetland. For these comparisons, the
brood density (broods/shoreline km) of the same year when
the invertebrate samples were taken was used. From this
data also average brood density for beaver flowages and
undisturbed ponds were calculated.
Shore configuration
Shoreline depth profiles was described simply by measuring
depths on half meter from the shore line from 10 points
chosen randomly around each pond. Depths were attained
from five beaver occupied flowagess, and from 27 unoccupied ponds and lakes. The 27 water bodies had shore
lines between 200 and 2000 m, which is the size class
normally affected by beaver, and only waters with no
human influence were included.
Data analysis
The comparison of the use of ponds by teal broods before
and during beaver occupation (and their controls) was made
using the Wilcoxons matched pairs signed ranks test
(Nummi and Poysa 1997). The association of brood density
and invertebrate abundance index was tested by the Spearman rank correlation. All the other comparisons were made
using the Mann-Whitney U-test.

Figure 1. Change in teal brood numbers per census in beaver

flowages and in control ponds expressed as brood number during
beaver occupancy years minus brood number before beaver
occupancy years. The data from control ponds are from the same
years as the data from beaver flowages.

vertebrate abundance (Spearman rank correlation, rS 

0.661, p0.000, Fig. 3).
Daily mortality rate of teal ducklings was three times
higher in waters without beaver flowages than with beaver
flowages (Mann-Whitney U-test, Z 3.353, p 0.001,
Fig. 4). The mean brood size of older ducklings tended
(Z 1.820, p 0.07) to be higher in beaver flowages
than in non-beaver influenced wetlands (mean 5.2 [SE 
1.93], n 16) versus 3.3 [SE 1.50]), but only 4 broods
were detected on that latter wetlands.

Results
The number of teal broods clearly increased upon beaver
flooding, (T 2.670, p 0.008, Wilcoxons matched
pairs signed ranks test), but no change was found in control
ponds (T 0.0, p1.0, Fig. 1). The average brood density
of teals in 11 beaver flowages was much higher (Fig. 2) than
in the 26 undisturbed ponds and lakes (Mann-Whitney Utest, Z 4.734, p 0.000).
Abundance of invertebrates was also higher in beaver
flowages than in other water bodies (Z 4.353, p 
0.000, Fig. 2). Isopods, cladocerans, Diptera larvae, corixids
and dytiscids were the most abundant invertebrates in
beaver flowages. Teal brood density correlated with in-

Figure 2. Density of teal broods per one shore line km and

invertebrate abundance index in beaver ponds (n 11) and nonbeaver ponds (n 26). Teal density is expressed per census,
invertebrate abundance is expressed per trap, and it takes into
account both abundance and average size of each taxon. Bars
denote SE.

521

Figure 3. Correlation between teal brood density and invertebrate

abundance index in 11 beaver flowages (black squares) and 26
non-beaver ponds (grey circles).

Average depths within 0.5 m from the shore were lower

in the five beaver impoundments than in the 27 undisturbed lakes and ponds (23.4 cm [SE 5.2] versus 64.2 cm
[SE 26.2], Mann-Whitney U-test, Z 3.402, p 
0.001). The shallow depths indicates how easily the
ducklings can reach for benthic invertebrates for food.

Discussion
In their conceptual model of facilitation, Bruno et al.
(2003) described three factors that can lead to recruitment
enhancement: 1) resource enhancement, 2) habitat amelioration, and 3) predation refuge (see also Dickman 1992,
Stachowicz 2001). The resource enhancement found in this
study
increased invertebrate production caused by beaver
ecosystem engineering
is beneficial for ducks since protein
is essential during the breeding season for egg production
and growth of ducklings (Scott 1973, Sedinger 1992, Krapu
and Reinecke 1992). This is especially so in this case since
ducklings seem to be limited by food in boreal lakes
(Nummi et al. 2000, Sjoberg et al. 2000).

Figure 4. Mortality of teal broods in beaver flowages and nonbeaver ponds expressed as daily mortality rate. n 29 for beaver
ponds, 10 for non-beaver ponds. Bars denote SE.

522

For teal ducklings, the amelioration of habitat structure

by beaver engineering is important, too, because it increases
the availability of the resources. The slope of the shores of
boreal ponds normally are very steep. Ducklings and often
not even adult ducks can not reach for the bottom lying
benthic food by dabbling. The difference in average depths
of beaver ponds and undisturbed waters (23 vs 64 cm) is
crucial since the maximum depth that teals reach by upending is 26 cm (Poysa 1983). Deep shores of boreal ponds
also have only a narrow belt of emergent vegetation. Beaver
inundation creates a flooded littoral zone with a wider
margin of shallow water suitable for foraging (Beard 1953,
Patterson 1976). In our study, the average depth in one
typical boreal mire pond lined with Sphagnum moss
changed from 94 cm before beaver occupation to 18 cm
during beaver flooding. The width of the flooded area
around pond ranged between 20 and 50 m. In contrast, the
foraging habitat available for ducklings in undisturbed
Sphagnum ponds was only the 10
20 cm wide edge zone
between water and land. In a pond with 500 m of shore line
this will provide 50
100 m2 of foraging habitat. With
flooding by beaver, there was 10 000
25 000 m2 of foraging
habitat giving at least a hundredfold increase in the
duckling foraging habitat.
The shallow flooded area contains a wide belt of
emergent plants and twigs, especially during early stage of
inundation (Beard 1953, Nummi 1989). Emergent plants
harbour abundant emerging insects (Danell and Sjoberg
1982), which are important for young ducklings which can
not readily forage under water (Chura 1961, Pehrsson
1979). When chased, insects escape by flying off. Therefore
a duckling will benefit from bating the maximum number
of insects from an undisturbed feeding patch (Nummi
1992).
The structural characteristics of the flowage are also
protective against predation. Ducklings can avoid terrestrial
predation because they do not have to forage right beside
the shore line where mammalian predators may ambush
(Pehrsson 1979). A thicket of flooded bushes will also
hamper aerial attacks by raptors (Hepp and Hair 1977).
Furthermore, dense vegetation cover is suboptimal for pike
Esox lucius (Casselman and Lewis 1996), which can be an
efficient duckling predator in the boundary zone between
aquatic vegetation and open water (Paasivaara and Poysa
2004). In new flowages the density of pikes and other fish is
low because it takes some time until the populations build
up in newly formed aquatic habitat.
During spring, teal pairs are the first ducks to colonize
the patches created by beavers (Nummi and Poysa 1997).
Moreover, teals not only colonize these beaver patches
rapidly but also use them succesfully for brood rearing
(Orians and Wittenberger 1991). This reinforces earlier
findings by Poysa et al. (2000) which showed that breeding
mallards Anas platyrhynchos could forecast brood-stage food
resources. Teal also seemed to show little delay in their
response of appearance at a good habitat. In this respect
there could be a slight difference in teal and mallard,
because mallards might fill good habitats after some delay,
possibly because of breeding site philopatry (Krapu et al.
1983, Poysa 2001). The teals diet shows adaptation to early
colonization of flowages: the bird eats a lot of cladocerans
(Nummi 1993) which are among the first invertebrates to

poliferate in beaver impoundments (Nummi 1989, Krylov

2002).
Theoretically, positive interactions are assumed to be
more common in stressful situations (Bertness and Callaway
1994, Choler et al. 2001). In boreal ponds, ducks face a
stressful situation because resources for reproduction are
limiting (Poysa et al. 2000, Sjoberg et al. 2000, Gunnarsson
et al. 2004). Therefore they can benefit from beaver
modified patches where the amount and availability of
food is higher than in undisturbed patches.
The population level effect on ducks caused by the
beaver depends on beaver density. In our study area, the
beaver density was low (0.08 colonies km2, Hyvonen and
Nummi 2008) and, hence, the effect on teal population was
probably not very strong. Nevertheless, over 60% of young
teal broods were observed in flooded habitats which
comprised only 7% of the total lake shore line (Nummi
and Poysa 1995b). In some instances the density of beavers
and their ponds can be considerably higher. Colony density
may reach 3 km 2, and in Minnesota the number of
aquatic habitats increased by 440% when beaver population
recovered after the period of overexploitation. As a result,
beaver flowages covered 13% of the total area of Kabetogama peninsula (Naiman et al. 1988, Johnston et al. 1993).
Geographically the ranges of beavers and teal (or greenwinged teal Anas crecca carolinensis) overlap in extensive
areas in North America and northern Europe, therefore a
large number of teal are potentially affected by beaver
ponds.
Apart from teal, beaver engineering beneficially affect
many organisms from plants and insects to various
vertebrates. The term keystone facilitator (Stachowicz
2001) therefore well describes the ecosystem function of the
beaver. It has been noted that facilitations include both
tightly coevolved, obligate relationships and much looser
facultative interactions (Bruno et al. 2003). Teal-beaver
relationship is a loose, facultative one beacause teals live in
other habitats as well. But it is intriguing to think that
among the numerous relationships that beaver has with
other species, there also might be some with a more obligate
nature.
Acknowledgements
We are greatful to Petri Timonen and other
field workers at Evo for their assistance in duck surveys. M.
Milonoff, H. Parker, J. Pellikka, H. Poysa, V.-M. Vaananen and
one anonymous referee provided valuable comments on the ms.
Esa Pienmunne kindly prepared the figures, and Alisdair McLean
checked the language.

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