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Birding • august 2002
346
1,2 Long Point Waterfowl and Wetlands
Research Fund Bird Studies Canada
P. O. Box 160
Port Rowan ON N0E 1M0
1 spetrie@bsc-eoc.org
2 mschumme@uwo.ca
SP is the research director of the Long
Point Waterfowl and Wetlands
Research Fund. His work has focused
primarily on the ecology of waterfowl
in semi-arid environments and the
staging ecology of waterfowl in north-
temperate regions.
MLS is a Ph.D. candidate in Zoology
at the University of Western Ontario,
where he specializes in waterbird
research and the advancement of wet-
land conservation. His areas of inter-
est include waterfowl population
dynamics, socio-economic impacts on
wetland conservation, and landscape
ecology.
Waterfowl Responses to Zebra
Mussels on the Lower Great Lakes
Scott Petrie1 and Michael L. Schummer2
Greater Scaup, Buffleheads,
Long-tailed Ducks, and other
waterfowl species that con-
centrate in large numbers on the
Great Lakes each winter are in dan-
ger of being poisoned by a novel
food source. The problem is present-
ed by a small creature with a
remarkable ability to concentrate
toxic substances found in lake
water—the zebra mussel.
The Great Lakes are widely
acknowledged to be one of the most
beleaguered ecosystems in North
America. Water pollution and shore-
line development are two of the bet-
ter- known affronts on this ecosys-
tem, but some ecologists consider
exotic plants and animals to be an
even greater threat. For better or for
worse, virtually every native species
in the Great Lakes has been impact-
ed by exotics, and waterfowl are no
exception.
The introduction of the zebra mus-
sel (Dreissena polymorpha) and closely
related Quagga Mussel (D. bugen-
sis)—hereafter referred to collectively
as zebra mussels—is a fairly recent
phenomeon, but it has already had
dramatic impacts on plants, animals,
and ecosystem processes of the Great
Lakes. It is tempting to view this
mussel invasion as a positive develop-
ment for waterfowl. Zebra mussels
provide a novel food source easily
exploited by certain species of water-
fowl. But the short-term benefits of an
increased food supply may be out-
weighed by the threat of food con-
tamination: as zebra mussels feed,
they accumulate in their tissues tox-
ins that may be passed up the food
chain.
MIKE DANZENBAKER
Long-tailed Ducks are annual on the lower Great Lakes. They are uncommon at most times and at most places, but large concentra-
tions are sometimes noted both on migration and during the winter months. It is unknown whether Long-tailed Duck populations on the
lower Great Lakes are being affected by the zebra mussel invasion in the region.
waterfowl responses to zebra mussels 347
Great Lakes and their inhabitants.
Native mussels, which averaged ten
individuals per square meter prior to
zebra mussel colonization, have been
almost completely displaced by this
highly competitive, exotic species.
Influence on Diet and Distribution
In Europe, zebra mussels provide a
readily available source of food used
by certain species of waterfowl. These
waterfowl species are known to alter
movement patterns to take advantage
of zebra mussels, particularly in
recently invaded lakes. It is not sur-
prising that certain species of North
American waterfowl also have shifted
their dietary choices, as well as their
distributions, to take advantage of this
novel, yet easy-to-find, food source.
When a prey item becomes super-
abundant and is easily exploited,
waterfowl tend to concentrate their
foraging efforts on that food source.
Zebra mussels certainly fit this bill (no
pun intended). They now are much
more readily available than any other
invertebrate food source, their con-
sumption generally requires limited
search, and they are easy to eat.
From 1991 to 1995, the diets of 552
ducks of twelve species were sampled
from the Long Point, Lake Erie, water-
fowl check station to identify food
habits. Five of the twelve species ana-
lyzed had consumed zebra mussels,
but only Lesser Scaup, Greater Scaup,
and Buffleheads consistently incorpo-
rated zebra mussels in their diet.
Other species of diving ducks have
been reported to consume zebra mus-
sels: Common Goldeneye, Long-tailed
Duck, and White-winged Scoter,
according to research by D.J. Hamilton
and C.D. Ankney. Given their inability
Exotic Introductions
From the common carp (late 1800s),
to the Eurasian watermilfoil (1952),
and to one of the newest introduc-
tions, the round goby (about 1990),
the Great Lakes have endured their
share of exotic invasions over the past
200 years. Over 140 non-indigenous
species have invaded the Great Lakes
since the early 1800s. Some intro-
duced species compete strongly with
native fauna and flora, often resulting
in changes in community structure.
This competition ultimately can influ-
ence food web interactions and ecosys-
tem functioning. Few introductions,
however, have had a greater effect on
the lower Great Lakes ecosystem than
has the zebra mussel.
Zebra mussels are small (less than
one inch) clam-like invertebrates
native to European lakes. They were
introduced into Lake St. Claire, east of
Detroit, in 1986, apparently a result of
the discharge of larvae in ship ballast
water. Faced with limited competition,
zebra mussels rapidly increased in
numbers, expanding their range
throughout the lower Great Lakes.
They occur at densities from a few
hundred to ten thousand per square
meter on sandy, loamy, and vegetative
surfaces. In rocky areas, where the
mussels preferentially settle, densities
have reached 750,000 per square
meter.
The ability to expand rapidly and
colonize new areas (via free-swimming
larvae called veligers), coupled with
the extremely high densities at which
they can occur (they can adhere to
most surfaces, including each other),
has enabled zebra mussels to change
the entire dynamic of the lower Great
Lakes lake-bottom community.
Previously slow-growing, with limited
influence on ecosystem interactions,
that community now is dominated by
a single species that has transcendent
effects on the ecology of the lower
MICHAEL L. SCHUMMER
Exotic zebra mussels can reach astonishingly high densities in suitable environments in
the lower Great Lakes. On sandy, loamy, or vegetative substrates, they occur at densities
of a few hundred to ten thousand per square meter. On rocky substrates, densities as
high as three-quarters of a million per square meter have been recorded. In contrast,
native mussels average only ten individuals per square meter.
Birding • august 2002
348
to forage in deep water, and a normal
diet of vegetation during the non-
breeding period, it is not surprising
that dabbling ducks generally do not
consume zebra mussels.
At Long Point, there was a three-fold
increase in the number of waterfowl
staging there between 1986 (prior to
zebra mussel colonization) and 1997
(six years after colonization). Most of
this gain can be attributed to increased
numbers of Lesser and Greater Scaup
(Figure 1). In fact, the use of Long
Point by Lesser and Greater Scaup
together increased 92-fold between
1986 and 1997 (based on day-use cal-
culations), despite a substantial
decline in the North American popula-
tion of scaup during that time. Similar
increases in scaup use have been
reported for Lake St. Clair, as well as
for Points Pelee and Rondeau on Lake
Erie.
These increases in scaup numbers
are most likely a consequence of more
birds gathering in these areas during
migration and of those birds then
remaining longer to consume zebra
mussels. Not only has the presence of
the mussels influenced the diets of
certain species of diving ducks on the
lower Great Lakes, but it also has
caused dramatic shifts in their distri-
butions and relative abundance during
migration.
Although no dietary studies have
been performed on diving ducks win-
tering on the lower Great Lakes since
the arrival of zebra mussels, we do
know that there have been major
changes in the wintering populations
of waterfowl since zebra mussel colo-
nization. One of the better long-term
sources of information on wintering
Great Lakes waterfowl numbers is an
extensive ground-based midwinter sur-
vey of the Canadian shoreline of Lake
Ontario, compiled by Bill Edmunds of
the Toronto Ornithological Club.
0
20,000
1971
1975
1979
1984
1986
1988
1991
1992
1993
1994
1995
1996
1997
1998
1999
2000
40,000
60,000
80,000
100,000
120,000
140,000
PILGSOPULATION NCREASES: ESSER AND REATER CAUP
Spring
Fall
Year
Numbers of Birds
Figure 1. This chart shows a steep rise in Lesser and Greater Scaup populations at Long
Point, which is on the northern shore of Lake Erie. Similar increases have been reported
for Lake St. Clair, as well as for Points Pelee and Rondeau on Lake Erie.
Numbers of Common Goldeneyes wintering on the lower Great Lakes have increased
appreciably in recent years. These recent population increases are probably tied to the
fact that the species is a known consumer of zebra mussels—which are a potentially
major food resource for wintering waterfowl.
MIKE DANZENBAKER
waterfowl responses to zebra mussels 349
Numbers of Canada Goose and dab-
bling ducks (all species combined)
increased by 2.8- and two-fold, respec-
tively, between 1980 and 2000. But
numbers of diving ducks (all species)
increased nine-fold (Figure 2).
Actually, wintering diving duck num-
bers have been increasing exponential-
ly since 1991, around the time zebra
mussels arrived on Lake Ontario. This
substantial increase in overwintering
diving ducks can be attributed for the
most part to increased numbers of
Lesser Scaup, Greater Scaup, Long-
tailed Ducks, Common Goldeneye,
Buffleheads, White-winged Scoters,
and Common Mergansers, all of
which—with the exception of
Common Mergansers—are known to
consume zebra mussels. Higher winter
temperatures and reduced ice cover
associated with the recent warming
trend have in all likelihood con-
tributed to increased use of the lower
Great Lakes by these birds. However,
substantial increases in numbers of
staging and overwintering birds have
been primarily limited to those species
which consume zebra mussels, which
suggests that mussel availability is
quite possibly a primary factor influ-
encing these changes in distribution
and abundance.
Possible Adverse Impacts
Rather than consuming algae and
other food items that adhere to rocks
and plants on the lake bottom, zebra
mussels acquire their nutrition by fil-
tering suspended matter from the
water. Each zebra mussel filters several
quarts of water per day. It is this pro-
lific filtering capacity, combined with
the fact that the animals occur at such
high densities, that has enabled zebra
mussels to have such a profound effect
on lake ecology. This species has
increased water clarity on the lower
Great Lakes, diverted organic matter
from the water column to the bottom,
BRIAN E. SMALL
The Lesser Scaup, like other diving duck species, appears to have been a beneficiary of
the recent zebra mussel invasion of the lower Great Lakes. But the species may suffer in
the long run from eating too many zebra mussels. This hazard arises because of the fact
that zebra mussels bioaccumulate toxins that are passed up the food chain to predators
such as the Lesser Scaup. In particular, Lesser Scaup may be in danger of eating zebra
mussels that have been contaminated with cadmium, selenium, polychlorinated biphenyls,
and polynuclear aromatic hydrocarbons.
0
50,000
100,000
150,000
200,000
250,000
300,000
350,000
400,000
PIAWOPULATION NCREASES: LL ATERFOWL
1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000
Year
Numbers of Birds
Diving Ducks
Dabbling Ducks
Canada Geese
Figure 2. This chart shows across-the-board increases in waterfowl populations on the
Canadian shoreline of Lake Ontario. Note that numbers of diving ducks have been
increasing especially rapidly.
Birding • august 2002
350
altered foodweb interactions, and
influenced contaminant and nutrient
cycling. The uptake and subsequent
transfer of contaminants through the
food chain by these creatures is an
area of continuing research and con-
cern.
Because they filter particles indis-
criminately, zebra mussels incorporate
and accumulate into their tissues
water-associated contaminants—poly-
chlorinated biphenyls (PCBs), polynu-
clear aromatic hydrocarbons (PAHs),
and heavy metals (e.g., cadmium).
They do this much more readily than
do native Great Lakes bivalves. These
contaminants can subsequently be
passed up the food chain to waterfowl
that consume the mussels. This ulti-
mately could compromise waterfowl
reproductive output or survival. For
instance, reproductive success in cap-
tive Tufted Ducks fed contaminated
zebra mussels was 60 percent less than
that of individuals fed less-contaminat-
ed mussels, according to research by
W.C. de Kock and C.T. Bowmer.
Waterfowl surveys on the breeding
grounds indicate that the continental
population of Lesser and Greater
Scaup declined from a high of eight
million birds in 1972 to 3.7 million in
2001. However, the bulk of this
decline has occurred since the mid-
1980s, around the time zebra mussels
began colonizing the Great Lakes.
United States harvest data also suggest
that the proportion of juveniles and
adult females in the Lesser Scaup pop-
ulation has declined over the last two
decades. This implies that both female
survival and breeding or fledgling suc-
cess have declined. Furthermore, there
has not been an increase in the num-
ber of scaup taken by hunters, and
populations of most other species of
diving ducks have been increasing
during the period of scaup decline.
Considering the increased use of the
Great Lakes by scaup, plus a diet shift
toward zebra mussels, scaup could be
acquiring unhealthy contaminant bur-
dens.
Studies to date have indicated that
PCB and DDE burdens in staging and
wintering Lesser and Greater Scaup on
the Great Lakes are below the known
effect levels for waterfowl. However,
selenium has been detected in the ele-
vated-to-potentially-harmful range in
most birds collected from Lakes
Ontario, Erie, St. Clair, and Michigan.
Elevated selenium levels (more than
ten parts per million, dry weight) can
impair reproduction; yet higher levels
(more than 33 parts per million) actu-
ally can cause mortality, according to a
1996 study by G.H. Heinz. Selenium is
a semi-metallic trace element occur-
ring naturally in some soils; it is also a
byproduct of smelting operations and
other industrial activities. Although
selenium is nutritionally required by
birds in very small amounts, it is high-
ly toxic in greater quantities.
Selenium can rapidly increase in
aquatic organisms, particularly in filter
feeders such as zebra mussels. Field
studies show that bottom-zone inverte-
brates can accumulate 20 to 370 parts
per million of selenium and still main-
tain stable, reproducing populations.
These levels are somewhat alarming as
reproduction in Mallards is impaired at
a dietary concentration of just nine
parts per million. It has been recom-
mended by A.D. Lemly that three parts
per million is the toxic threshold for
selenium in aquatic food-chain organ-
isms consumed by fish and wildlife.
Selenium concentrations quickly build
up in tissues when birds are intro-
duced to a selenium-contaminated
diet. Selenium is also quickly excreted
from the body when birds are removed
from a selenium source. Females use
the egg as a route of selenium excre-
tion, and high selenium burdens can
impair reproduction.
It is difficult to speculate at the present time about the eventual impacts of the zebra
mussel on waterfowl populations that winter on the Great Lakes. Zebra mussels present
a superabundant resource for Buffleheads and other species. But zebra mussels have
greatly destabilized foodweb dynamics and contaminant cycling in the lower Great
Lakes, with potentially harmful long-term effects on waterfowl populations.
LARRY SANSONE
Hamilton, D.J. and C.D. Ankney. 1994.
Consumption of zebra mussels
Dreissena polymorpha by diving
ducks in Lakes Erie and St. Clair.
Wildfowl 45:159–166.
Heinz, G.H. 1996. Selenium in birds.
pp. 447–458 in: W.N Beyer, G.H.
Heinz, and A.W. Redmond-Norwood,
eds. Environmental Contaminants in
Wildlife: Interpreting Tissue
Concentration. SETAC Special
Publication Series. Lewis Publishers,
Boca Raton.
Herbert, P.D.N., B.W. Muncaster, and
G.L. Mackie. 1989. Ecological and
genetic studies on Dreissena polymor-
pha (Pallas): a new mollusc in the
Great Lakes. Canadian Journal of
Fisheries and Aquatic Sciences
46:1587–1591.
Lemly, A.D. 1996. Selenium in aquatic
organisms. pp. 427–445 in: W.N.
Beyer, G.H. Heinz, and A.W.
Redmond-Norwood, eds.
Environmental Contaminants in
Wildlife: Interpreting Tissue
Concentration. SETAC Special
Publication Series. Lewis Publishers,
Boca Raton.
Mitchell, C.A. and J. Carlson. 1993.
Lesser Scaup forage on zebra mussels
at Cook Nuclear Plant, Michigan.
Journal of Field Ornithology
64:219–222.
Petrie, S.A. and R.W. Knapton. 1999.
Rapid increase and subsequent
decline of zebra and quagga mussels
in Long Point Bay, Lake Erie: possible
influence of waterfowl predation.
Journal of Great Lakes Research
25:772–782.
Prince, H.H., P.I. Padding, and R.W.
Knapton. 1992. Waterfowl use of the
Laurentian Great Lakes. Journal of
Great Lakes Research 18:673–699.
Wormington, A. and J.H. Leech. 1992.
Concentrations of diving ducks at
Point Pelee, Ontario, in response to
invasion of zebra mussels, Dreissena
polymorpha. Canadian Field
Naturalist 106:376–380.
waterfowl responses to zebra mussels 351
Because of the rapid rate of uptake
and excretion, birds collected on the
Great Lakes that show elevated seleni-
um burdens quite possibly acquired
those burdens while foraging on the
Great Lakes. Even if selenium inputs
to the lower Great Lakes have not
increased substantially over the past
fifteen years, zebra mussels—through
filter feeding and bioaccumulation—
may have concentrated selenium in
their tissues, thereby increasing the
availability of this trace element to
waterfowl that consume them. Due to
the large numbers of scaup staging on
the lower Great Lakes and Mississippi
River, where zebra mussels are readily
available, selenium intake may be a
factor in the decline of these species.
For these reasons, the Long Point
Waterfowl and Wetlands Research
Fund began an intensive study of
dietary intake, contaminant burdens,
and body condition of spring and fall
staging Lesser and Greater Scaup on
Lakes Ontario, Erie, and St. Clair in
1999. Birds collected by the Canadian
Wildlife Service in 1985 (prior to
zebra mussel colonization), as well as
birds collected throughout the fall of
1999 and spring of 2000, plus the
zebra mussels themselves, are being
analyzed for burdens of heavy metals
and trace elements, including seleni-
um. This study will enable us to
determine conclusively if Lesser and
Greater Scaup are acquiring
unhealthy burdens of selenium (or
any other heavy metals or trace ele-
ments) while staging on the lower
Great Lakes, and if contaminants are
in fact acquired through zebra mussel
consumption.
We know that zebra mussels pro-
vide a readily available, easily
exploitable food source, and that cer-
tain species of waterfowl will change
their movement patterns to exploit it.
But we are as yet uncertain if this
novel food item is in fact toxic to its
dinner guests.
Further Reading
For further details on some of the
studies mentioned in the article, along
with additional information on zebra
mussels, we recommend the following
resources:
Brieger, G. and R.D. Hunter. 1993.
Uptake and depuration of PCB 77,
PCB 169, and hexachlorobenzene by
zebra mussels (Dreissena
polymorpha). Ecotoxicology and
Environmental Safety 26:153–165.
Custer, C.M. and T.W. Custer. 1996.
Food habits of diving ducks in the
Great Lakes after the zebra mussel
invasion. Journal of Field
Ornithology 67:86–99.
de Kock, W.C. and C.T. Bowmer. 1993.
Bioaccumulation, biological effects,
and foodchain transfer of contami-
nants in the zebra mussel (Dreissena
polymorpha). pp. 503–533 in: T.F.
Nalepa and D.W. Schloesser, eds.
Zebra Mussels: Biology, Impacts, and
Controls. Lewis Publishers, Boca
Raton.
Fisher, S.W., D.C. Gossiaux, K.A.
Bruner, and P.F. Landrum. 1993.
Investigations of the toxicokinetics
of hydrophobic contaminants in the
zebra mussel (Dreissena polymor-
pha). pp. 465–490 in: T.F. Nalepa
and D.W. Schloesser, eds. Zebra
Mussels: Biology, Impacts, and
Controls. Lewis Publishers, Boca
Raton.
Gillis, P.L. and G.L. Mackie. 1994.
Impact of the zebra mussel,
Dreissena polymorpha, on popula-
tions of Unionidae (Bivalvia) in Lake
St. Clair. Canadian Journal of
Zoology 72:1260–1271.
Griffiths, R.W., D.W. Schloesser, J.H.
Leach, and W.P. Kovalak. 1991.
Distribution and dispersal of the
zebra mussel (Dreissena polymorpha)
in the Great Lakes region. Canadian
Journal of Fisheries and Aquatic
Sciences. 48:1381–1388.