Ecology
of the Beaver
Note:
This online review is updated and revised continuously, as soon as results
of new scientific research become available. It therefore presents
state-of-the-art information on the topic it covers.
The
beaver inhabits freshwater streams and lakes of the Northern Hemisphere.
In addition, it has been introduced by humans into southernmost South America
(Lizarralde et al. 2004). Two species are recognized: Castor fiber
of Eurasia and Castor canadensis of North America.
Ecologists
call beaver "ecosystem
engineers" because these animals physically alter habitats by cutting
down trees, building dams, digging canals and building lodges.
In doing so, beaver change the distribution and
abundance of many other animals and plants, mostly by indirect
interactions. In this series of reports, we review the ecology of
beaver and the many
diverse effects that beaver engineering has on other
organisms.
When
beavers dam a stream they slow the movement of water. Behind the
beaver dam, a pond of still water is formed. This pond (impoundment)
is then colonized
by animals and plants that typically live in lakes rather than
streams. Organisms dependent on fast moving water die out
in the beaver pond, or move
to parts of the stream where the flow of water has not been slowed by the
beaver dam. After a beaver dam has existed for ten years
or more, the pond it created usually has an abundance of submersed and
emergent vegetation, along with the many animals that live in such
vegetation.
The
forest beside the stream also changes after beaver occupation. When beavers cut down trees for food and for building their dams and
lodges, they
select the species of trees that they prefer, and leave other tree species
standing. Consequently, after many years, the forest beside a beaver
pond is usually dominated by different tree species than it was before beaver
occupation, and in
the gaps where the beavers removed trees, bushes and saplings now grow and with them the animal species that live in the early stages of forest
regeneration (Barnes and Dibble 1986; Johnston and Naiman 1990; Pastor and Naiman
1992; Donkor et al. 2000). In addition, when
the beaver pond is formed by the dam, water floods and covers the roots of
trees that formerly stood along the stream bank. These flooded trees
die because the standing water prevents their roots from getting air.
When the flow of
water in a stream is slowed by the beaver dam, soil and organic sediment
carried in the water usually settle to the bottom of the beaver pond. When beaver subsequently abandon a locality, their dam eventually breaks
and the pond drains leaving a large open space. A meadow usually
grows on the nutrient-rich soils that once formed the bottom of the pond.
These "beaver meadows" usually have more light penetration, higher soil
moisture, more nitrogen and a different vegetation than the adjacent riparian
forest (Johnston et al. 1995; Wright et al. 2002).
Beaver engineering also includes two other activities: lodge making and canal digging. The construction of lodges by beaver adds coarse woody debris to the beaver pond which some fish
species use for cover (France 1997; Collen and Gibson
2001). Canals are usually about "30 to 60 cm wide
and 20 to 35 cm deep, and can extend hundreds of yards into the forest
(Stocker 1985; Collen and Gibson 2001). After a beaver has dug a canal, it can float branches from trees it has cut and move them to safer
feeding locations.
Effects
of beaver engineering on wildlife
Beaver
engineering alters the
distribution and abundance of so many organisms, that we cannot mention
them all on this page. Therefore, we have produced separate
reviews for different groups of animals and plants affected by beaver
engineering. Click
the following links to learn more about the effects of beaver engineering on specific species of
plants and animals:
birds
amphibians
reptiles
invertebrates trees
For effects of beaver engineering on fishes, see Collen and Gibson (2001).
Salmon stop using certain rivers whose streams are dammed by beaver
because the dams block salmon movements upstream. In the arctic,
Beluga Whales (Delphinapterus leucas) which feed on salmon, also stop
entering the same dammed rivers (Huntington
and Myrmin 1996; Pierotti and Wildcat 2000).
Effects
of beaver engineering on humans
In
Wyoming, a survey showed that owners of private lands believed that they benefited from
beaver engineering because it elevated water tables, increased the area of
riparian vegetation on their lands, and also increased livestock watering
opportunities (McKinstry and Anderson 1999). However, these same
landowners regarded beaver as pests when these rodents girdled timber,
blocked irrigation ditches and culverts with wood, and flooded roads, railroads, crops and timber (McKinstry and Anderson 1999).
In
New York state, beaver plug highway culverts with wood, creating
"roadside impoundments that damage and sometimes flood the
roadbed" (Jensen et al. 2001). However, oversized
culverts were less likely to be plugged by beaver, so it was recommended
that oversized culverts be installed (Jensen et al. 2001). Although
such oversized culverts are more expensive, over the long run they are
regarded as more cost-effective than trapping or debris removal.
In
some places, such as the southeastern United States, beaver cause
extensive damage to valuable timberland by flooding bottomland forests
and eating tree seedlings (Bhat et al. 1993; Conner at al.
2000). Although trapping can control beaver populations, low pelt
prices often fail to provide a stimulus to professional trappers, and the
landowner is often left with the cost of removing the nuisance
beavers.
Unfortunately, after one landowner traps beaver on his
property, surplus beaver from neighboring properties often invade the now
vacant habitat and the landowner is right back where he or she
started. Bhat et al. (1993) argue that all landowners in an area
must therefore cooperate together to manage beaver. They propose a
long-term trapping program with increased trapping in the initial
years. This results in fewer beavers in the total area, less
trapping required in subsequent years, and a smaller number of beaver
being killed over the long-term. A weakness of this proposal is that
it requires all landowners in an area to cooperate. Landowners whose
own economic or recreational interests are benefited by beaver and beaver engineering
may be reluctant to cooperate.
Relationships with
Predators
Many large predators occasionally prey on beaver, however only the wolf (Canis lupus)
does so regularly and to the extent that it can significantly reduce
numbers of beaver
(Shelton and Peterson 1983). For example, in southeastern Alaska, 31% of
wolf feces contained the remains of beaver (Kohira and Rexstad (1997).
In Belarus, the frequency of beaver remains in wolf feces ranged from 6%
to 22% over a ten-year period (Sidorovich et al. 2003). Frequent
wolf predation on beaver has also been documented in Latvia, Ontario, Minnesota
and Alaska (Voigt et al. 1976; Fuller 1989; Thurber and Peterson
1993; Andersone and Ozolins 2004).
On the other hand, wolves
benefit beaver indirectly by killing and scaring away potential competitors of beaver.
For example, fear of reintroduced wolves caused wapiti (Cervus elaphus)
in Yellowstone National Park, to avoided some riparian zones where they had
previously over-browsed and eliminated young willow and cottonwood trees.
The resulting relief from wapiti over-browsing allowed tree populations in
these areas to
recover, providing food for beaver which promptly colonized the area
(Ripple and Beschta 2004).
Effects
of beaver engineering on biodiversity
In the Appalachian Plateau region of New York, active beaver impoundments
contained "significantly more bird species and a greater average
number of bird species than abandoned beaver
ponds and control sites with no record of beaver occupation (Grover and
Baldassarre 1995)."
In
the Upper Piedmont of South Carolina, the abundance, richness and
diversity of reptiles were significantly higher at beaver impoundments
than at unimpounded streams, however the "richness, diversity and
evenness of amphibians was significantly higher at unimpounded streams
than at beaver ponds (Metts et al. 2001).
In
the Adirondack region of New York, Wright et al. (2002) found that beaver
engineering increased species richness of plants at the landscape scale,
because beaver created patches
of habitat (beaver ponds and meadows) had a combination of conditions that
were not present elsewhere in the landscape, and some plant species that
lived in these beaver-modified habitats were not present in habitats
unmodified by beaver.
References
Andersone A, Ozolins J
(2004) Food habits of wolves Canis lupus in Latvia.
Acta Theriologica 49: 357-367
Barnes WJ, Dibble E (1986) The effects of beaver in
riverbank forest succession. Canadian Journal of Botany
66:40-46
Bhat
MG, Huffaker RG, Lenhart SM (1993) Controlling forest damage
by dispersive beaver populations - centralized optimal management
strategy. Ecological Applications 3: 518-530
Collen
P, Gibson RJ (2001) The general ecology of beavers (Castor
spp.) as related to their influence on stream ecosystems and riparian
habitats, and the subsequent effects on fish - a review. Reviews
in Fish Biology and Fisheries 10: 439-461
Conner WH,
Inabinette LW, Brantley EF (2000) The use of tree shelters in
restoring forest species to a floodplain delta: 5-year results. Ecological
Engineering 15: S47-S56, Supplement 1
Donker NT, Fryxell JM (2000) Lowland boreal
forests characterization in Algonquin Provincial Park relative to beaver (Castor
canadensis) foraging and edaphic factors. Plant Ecology
148: 1-12
Forbes GJ, Theberge JB (1996)
Response by wolves to prey variation in central Ontario. Canadian
Journal of Zoology 74: 1511-1520
France
RL (1997) The importance of beaver lodges in structuring
littoral communities in boreal headwater lakes. Canadian Journal
of Zoology 75: 1009-1013
Fuller TK
(1989) Population dynamics of wolves in north-central Minnesota.
Wildlife Monographs 105: 1-41
Grover
AM, Baldassarre GA (1995) Bird species richness within beaver
ponds in South-central New York. Wetlands 15: 108-118
Huntington
HP, Myrmin NI (1996) Traditional ecological knowledge of
beluga whales: an indigenous knowledge project in the Chukchi and northern
Bering Seas. Inuit Circumpolar Conference, Anchorage, Alaska,
USA
Jensen
PG, Curtis PD, Lehnert ME, Hamelin DL (2001) Habitat and
structural factors influencing beaver interference with highway
culverts. Wildlife Society Bulletin 29: 654-664
Johnston
CA, Naiman RJ (1990) Browse selection by beaver: effects on
riparian forest composition. Canadian Journal of Forest Research
20: 1036-1043
Johnston CA, Pinay G, Arens C, Naiman RJ (1995)
Influence of soil properties on the biogeochemistry of a beaver meadow
hydrosequence. Soil Science Society of America Bulletin 59:
1789-1799
Kohira M,
Rexstad EA (1997) Diets of wolves, Canis lupus, in
logged and unlogged forests of southeastern Alaska. Canadian
Field-Naturalist 111: 429-435
Lizarralde
M, Escobar J, Deferrari G (2004) Invader species in Argentina: A review
about the beaver (Castor canadensis) population situation on Tierra
del Fuego ecosystem. Interciencia 29: 352+
Metts
BS, Lanham JD, Russell KR (2001) Evaluation of herpetofaunal
communities on upland streams and beaver-impounded streams in the upper
piedmont of South Carolina. American Midland Naturalist 145:
54-65
McKinstry
MC, Anderson SH (1999) Attitudes of private- and public-land
managers in Wyoming, USA, toward beaver. Environmental Management
23: 95-101
Pastor
J, Naiman RJ (1992) Selective foraging and ecosystem processes
in boreal forests. American Naturalist 139: 690-705
Pierotti
R, Wildcat D (2000) Traditional ecological knowledge: the
third alternative (commentary). Ecological Applications 10:
1333-1340
Ripple WJ,
Beschta RL (2004) Wolves and the ecology of fear: can
predation risk structure ecosystems? Bioscience 54: 755-766
Shelton
PC, Peterson RO (1983) Beaver, wolf and moose interactions in
Isle Royale National Park, USA. Acta Zoologica Fennica 174:
265-266
Sidorovich VE, Tikhomirova LL, Jędrzejewska
B (2003) Wolf Canis lupis numbers, diet and damage to
livestock in relation to hunting and ungulate abundance in northeastern
Belarus during 1990-2000. Wildlife Biology 9: 103-111
Stocker
G (1985) The beaver (Castor fiber L.) in Switzerland -
Biological and ecological problems of re-establishment. Swiss
Federal Institute of Forestry Research Reports. 242: 1-149
Thurber JM,
Peterson RO (1993) Effects of population density and pack size
on the foraging of gray wolves. Journal of Mammalogy 74:
879-889
Voigt DR,
Kolenosky GB, Pimlott DH (1976) Changes in summer foods of
wolves in central Ontario. Journal of Wildlife Management 40:
663-668
Wright
JP, Jones CG, Flecker AS (2002) An ecosystem engineer, the
beaver, increases species richness at the landscape scale. Oecologia
132: 96-101
Information
about this Review
Author: Dr. Paul D. Haemig (PhD in Animal Ecology)
Photographer of the double beaver dam: Victor Maltby (Canada)
The
proper citation is:
Haemig
PD 2012
Ecology
of the Beaver. ECOLOGY.INFO 13.
If
you are aware of any important scientific publications that were omitted
from this review, or have other suggestions for improving it, please
contact the author at his e-mail address:
director {at} ecology.info
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