thehistoryandfutureoflakechamplain's fishesand sheries and langdon...
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The history and future of Lake Champlain's fishes and fisheries
J. Ellen Marsden a,⁎, Richard W. Langdon b,1
a Rubenstein School of Environment and Natural Resources, Aiken Center, 81 Carrigan Dr., University of Vermont, Burlington, VT 05405 USAb Vermont Department of Environmental Conservation, 103 South Main St., Waterbury, VT05671-0401, USA
a b s t r a c ta r t i c l e i n f o
Article history:
Received 11 January 2011
Accepted 7 June 2011
Available online 29 November 2011
Communicated by Doug Facey
Keywords:
Habitat fragmentation
Zoogeography
Management
Degradation
Restoration
In the last two centuries, physical, chemical, and biological alterations of Lake Champlain have resulted in the
loss of two species, addition of 15 fish species, and listing of 16 species as endangered, threatened or of spe-
cial concern. The lake currently supports 72 native fish species; lake trout (Salvelinus namaycush) and Atlantic
salmon (Salmo salar) were extirpated by 1900, American eel (Anguilla rostrata) and lake sturgeon (Acipenser
fulvescens) populations are extremely low, andwalleye (Sander vitreum) are declining. Dams on several rivers,
and ten causeways constructed in the mid 1800s to early 1900s, cut off access to critical spawning areas and
may have limited fish movements. Siltation and sediment loading from agricultural activity and urban growth
have degraded substrates and led to noxious algal blooms in some bays. A commercial fishery targeting
spawning grounds of lakewhitefish (Coregonus clupeaformis), lake trout, andwalleye probably reduced numbers
of these species prior to its closure in 1912. Non-native species introductions have had ecosystem-wide impacts.
Sea lamprey (Petromyzon marinus) populations were very high prior to successful control, possibly as a conse-
quence of ecological imbalance and habitat changes. A paucity of historic survey data or accurate species
accounts limits our understanding of the causes of current fish population trends and status; in particular, the
effects of habitat fragmentation within the lake and between the lake and its watershed are poorly understood.
Holistic, ecosystemmanagement, including pollution reduction and examination of habitat impacts, is necessary
to restore the general structure of native biological assemblages.
© 2011 Published by Elsevier B.V. on behalf of International Association for Great Lakes Research.
Introduction
Following the European discovery of Lake Champlain in 1609 by
its namesake, Samuel de Champlain, the ensuing 400 years brought
substantial physical changes to the watershed, lake sediments, and
hydrological connections within the lake. Two fish species, lake
trout2 and Atlantic salmon, were extirpated, 15 fish species were
added, and 16 fish species have been listed as endangered, threat-
ened, or of special concern/susceptible. Chemical inputs from land
use and industries have caused algal blooms and have contaminated
fish tissue. These changes in the biological, physical, and chemical char-
acteristics of the lake present practical and philosophical challenges to
management: to what extent have ecosystem services been compro-
mised, is restoration possible, and should restoration, rather than ac-
ceptance of an altered system, be the goal? Herein, we review the
history of biological and physical changes in the lake and the subse-
quent changes in the stability and distribution of fish populations. We
then discuss the consequences for management of the fisheries and
protection of fish populations and communities.
Description of Lake Champlain
Lake Champlain is a long (193 km), narrow (20 km at its widest
point) lake that lies on the border between New York and Vermont,
extending into Quebec at the north (Fig. 1). The lake averages
19.5 m depth, with the deepest portion (122 m) in a narrow trench
immediately south of the main basin. Three long islands split the
northern third of the lake into eastern and western arms; causeways
constructed among these islands and between the islands and the
mainland have further divided the lake. Currently five distinct basins
are recognized: Missisquoi Bay at the north is shallow (4.3 m maxi-
mum depth) and highly eutrophic, the Northeast Arm (locally called
the Inland Sea) and Malletts Bay to the east are moderately deep
(48 and 30 m, respectively) and mesotrophic; the Main Lake, com-
prising the broad lake and northwestern arm, is largely deep and oli-
gotrophic, and the South Lake is eutrophic and largely riverine
(Marsden et al., 2010, Fig. 1). The watershed is large (21,326 km2)
in relation to the lake area (1130 km2), so that anthropogenic uses
of the landscape have the potential to significantly impact the lake.
Vermont, New York, and Quebec contain 56%, 37%, and 7% of the wa-
tershed, respectively; 62% of the lake surface area is in Vermont, 34.5
in New York, and 3.5% in Quebec. The lake receives input from
Journal of Great Lakes Research 38 (2012) 19–34
⁎ Corresponding author. Tel.: +1 802 656 0684; fax: +1 802 656 8683.
E-mail addresses: [email protected] (J.E. Marsden),
[email protected] (R.W. Langdon).1 Tel.: +1 802 734 6498.2 All scientific, current, and historic common names of fish species present in Lake
Champlain and mentioned in this paper appear in Table 1.
0380-1330/$ – see front matter © 2011 Published by Elsevier B.V. on behalf of International Association for Great Lakes Research.
doi:10.1016/j.jglr.2011.09.007
Contents lists available at SciVerse ScienceDirect
Journal of Great Lakes Research
j ourna l homepage: www.e lsev ie r .com/ locate / jg l r
numerous tributaries; the 11 major rivers each drain from 2252 to
3500 km2 of watershed. The outlet to the lake is the Richelieu River,
which flows into the St. Lawrence River from the north end of the
lake. The Chambly Canal, opened in 1843, bypasses the rapids on
the Richelieu River. The Champlain Canal, opened in 1823, connects
the lake to the Hudson River drainage and to the Great Lakes via the
New York State Canal System.
Physical history of Lake Champlain
Beginning about 18,000 years ago, melting of the retreating
Wisconcinan glacial ice sheet, the last in a series of glaciations, created
vast proglacial water bodies across the North America (Dyke and
Prest, 1987). In thewake of the receding glacier,fishes and other aquatic
populations began populating glacial melt waters through connections
to glacial refugia located to thewest, south and east of the shrinking gla-
cier. Proglacial Lake Vermont filled the Champlain Valley with various
shorelines up to 183m higher in elevation than present day (Fig. 2;
Chapman, 1937). In the Midwest, species originating from the rich
Mississippian refugium diffused northward and eastward into the pro-
glacial Great Lakes (Schmidt, 1986). Lake Vermont was connected to
the outlet of the Great Lakes that ran to the Atlantic Ocean, first through
the Mohawk and Hudson Valleys, then through the St. Lawrence Valley
(Fig. 2; Langdon et al., 2006). Fishes also entered meltwater rivers and
lakes to the south from unglaciated areas along the mid-Atlantic Coast
into Lake Vermont via the connection with the Hudson River Valley
(Schmidt, 1986) and possibly via the outlet of Lake Winooski into
the Connecticut Valley (Langdon et al., 2006). Between 13,000 and
10,000 years ago, migrations via freshwater connections from the St
Lawrence Valley were interrupted by the incursion of the Champlain
Sea (Cronin et al., 2008).
Following the eventual freshening of the Champlain Sea, access to
the lakes by Midwestern fauna was temporarily reestablished via the
Great Lakes outlet, the St. Lawrence River, that led to the Atlantic
Ocean (Dadswell, 1972). This avenue was available for fish movement
until glacial rebound of the Champlain Valley floor created abrupt
changes in gradient of the Richelieu River outlet at the north end of
the lake. The resulting waterfalls created barriers that prevented
Fig. 1. Lake Champlain, showing major rivers, lake segments, and towns mentioned in the text. The two Vermont dams, at Milton and Swanton, are indicated with stars. Inset
indicates the location of the two dams on the Richelieu River, and the canals that link Lake Champlain with the Hudson and Mohawk rivers to the south, and bypass the rapids
on the Richelieu River to the north.
20 J.E. Marsden, R.W. Langdon / Journal of Great Lakes Research 38 (2012) 19–34
natural access of fishes to the Champlain Valley from the St. Lawrence
and Richelieu Rivers, leaving the existing native fauna that we see
today. The completion of canals at the southern and northern ends
of the lake during the early 1800s once again opened dispersal corri-
dors to fish species, though fishes accessing Lake Champlain via these
routes are now regarded as non-native (Langdon et al., 2006).
Currently, the principal fall line runs north–south on the Vermont
side of the lake at an elevation of approximately 46 m. The fall line is
characterized by precipitous drops in elevation of major tributaries;
the resulting falls are barriers to most fish species. The area below
the falls in these tributaries provides needed spawning habitat for
many species such as walleye, lake sturgeon, and three redhorse spe-
cies. In addition, these areas provide unique habitat for smaller spe-
cies, many of which are rare, such as the eastern sand and channel
darters, mottled sculpin, and stonecat. No similarly abrupt fall line is
present on the New York side of the valley, where tributaries descend
more quickly and steadily to lake level.
Changes in the watershed
Most of the Lake Champlainwatershedwas forestedprior to European
colonization; in the late 1800s up to 60% of the landscapewas deforested,
with additional land cleared at various times. Currently, the watershed is
largely reforested in the AdirondackMountains to thewest and the Green
Mountains to the east, with extensive agricultural areas in Vermont and
Quebec. The human population in the basin, estimated to be 571,000 in
2000, is largely concentrated in the two cities of Burlington, Vermont,
and Plattsburgh, New York (Lake Champlain Basin Program, 2004).
Urban and developed areas within the basin now cover 6% of the water-
shed; 16% is in agricultural use, 64% is forested, and 14% is wetland or
open water (Lake Champlain Basin Program, 2004).
Anthropogenic changes in Lake Champlain include increased trib-
utary loadings of sediment and nutrients, industry inputs of sawdust
and contaminants, shoreline alteration, and construction of barriers
and causeways. The periods of deforestation resulted in high, erosive
storm flows that washed nutrients and sediments into tributaries,
which in turn carried them into the lake. Increased stream flows
also reduced fish habitat in streams by removing large woody debris
and other structures (Thompson, 1824). Organic material in the
form of sawdust, grain wastes, and cloth fibers (from fulling mills)
were deposited into streams (Public Health Service, 1951; Thompson,
1853). Increased sediment loads would have negatively impacted
spawning gravels of stream-spawning species, including Atlantic
salmon, brook trout, white suckers and redhorse. Increased sedimen-
tation may have also altered benthic invertebrate communities, and
created habitat for native mussels and larval lampreys. Sediment
accumulation rates increased in the Main Lake between the 1930s
and 1980s, and in Missisquoi Bay between the 1970s and 2000s
(Schwarting, 2011). Nitrogen and carbon inputs increased in both
areas during the 1970s to 1980s, and the carbon:nitrogen ratio in
Missisquoi Bay reflect inputs of terrestrial organic and sediment ma-
terial (Schwarting, 2011). Elevated phosphorus levels in Missisquoi
Bay, St. Albans Bay, and the South Lake have caused these areas to become
highly eutrophic, and frequent nuisance blooms of blue-green algae occur
(Smeltzer et al., 2012). Upgraded wastewater treatment facilities have
reduced phosphorus effluents and federal and state programs are cur-
rently targeting agricultural sources, but phosphorus levels remain high
in the sediments (Smeltzer et al., 2012).
Habitat fragmentation — rivers
Dams were constructed on most of the major rivers in the 1800s,
including the Great Chazy, Little Chazy, Salmon, Little Ausable, Ausable,
Fig. 2. Geographic extent of A) Lake Vermont (12,500 years ago) and B) the Champlain
Sea (10,000 years ago), relative to C) the present-day lake and lake basin.
21J.E. Marsden, R.W. Langdon / Journal of Great Lakes Research 38 (2012) 19–34
Boquet, Winooski, Lamoille, Missisquoi, and Otter Creek. Smaller rivers
and streams were also substantially affected by various types of dams;
in the early 1800s, Zadock Thompson estimated there were a total of
786 sawmills, 373 grist mills, and 252 fullingmills on Vermont streams
(Thompson, 1824). Currently there are 463 standing dams in the
Champlain drainage in Vermont; fewer dams are present on the New
York side due to the steepness of the shoreline and relative paucity of
farming and industry in the early days of colonization. On the Vermont
side of the lake only two dams have been built downstream of a natural
barrier to upstreammovement: Swanton dam on the Missisquoi River,
built in 1797, and Peterson dam on the Lamoille River, built in 1948
(Fig. 1). Both dams cut off fish access to necessary habitat for critical
life stages. Affected species include lake sturgeon and Atlantic salmon,
which were historically abundant in the Lamoille River, and suckers,
redhorses, and lake whitefish that move into the Missisquoi River at
different times of the year. Hydroelectric dams have been built along
the fall line of the remaining larger tributaries in Vermont, potentially
precluding upstream movement of Atlantic salmon in search of
spawning areas.
Habitat fragmentation — lake
Since the mid 1800s, the connectivity of the lake itself has been con-
siderably altered by the construction of several causeways between the
mainland and the two largest islands, South Hero (Grand Isle) and
North Hero (Fig. 3, Table 2). Historical descriptions of these causeways
are sometimes confusing due to the use of the term ‘bridge’, which in
early writings could mean ‘bridge of land’ rather than a bridge over
water. The first of these land bridges was the Sandbar Causeway, con-
structed in 1850 between the mainland and the southeastern corner of
Grand Isle, isolating the Northeast Arm from the Main Lake (Fig. 3).
Prior to its construction, a natural sandbar existed which, during low
water periods, was used to bring people, horses, and carriages to and
from the island; however, when the lake level was higher, or storms oc-
curred, this passage was flooded and presumably fishes were able to
pass over the bar (Stratton, 1980). The lack of an opening in this ‘bridge’,
and potential effects on fish movements, caused considerable concern
among fishermen; concerns of boaters were not reported. Vermont
State Fish and Game Commission reports dating from 1894 suggested
that an opening be constructed through the causeway, but it was not
until 1907 that the state of Vermont opened a 25 m sectionwith a bridge.
A second gap, 54 m wide, was subsequently opened. In the meantime, a
railroad bridge was constructed at Rouses Point in 1851, and a railroad
causeway at Larabees Point in 1871; the latter included a 91 m gap with
afloating bridge. Causewayswere built from Isle LaMotte to themainland
in 1882, betweenNorthHero andAlburg in 1886, and betweenGrand Isle
andNorth Hero in 1862; the latter two each incorporated a 60 mopening
with a drawbridge (Stratton, 1980). The most extensive causeway-
building occurred in 1899 with the construction of the Island Line, a rail-
road that ran from the Vermont side of the lake, through Burlington,
Grand Isle, and North Hero, to the New York side of the lake at Rouses
Point. This line entailed construction of causeways enclosing the west
side of Malletts Bay (5.2 km with two openings), at Pelots Point on the
west side of Carry Bay, and across the Alburg Passage (Fig. 3, Table 2).
Finally, in 1938, Missisquoi Bay in the north was mostly isolated from
the Northeast Arm by construction of a 1463m causeway.
These causeways transformed open embayments into essentially
isolated basins: Malletts Bay, now closed on all sides and isolated
from the Main Lake and the Northeast Arm; the Northeast Arm; the
Gut, which had previously been a passage between two islands; and
Missisquoi Bay. Carry Bay was also closed off at its western and north-
ern ends (Fig. 3). The lake has been transformed from a large body of
water, with islands in the center, into five separate basins. The extent
towhich these barriers have obstructedfishmovement, altered seasonal
migrations, and potentially fragmented fish populations is unknown.
The smaller bays (the Gut, Carry Bay) are currently highly vegetated in
summer, possibly as a consequence of reduced flows, increased deposi-
tion of sediment, and retention of nutrients. The Missisquoi Bay cause-
way, in particular, has been highly controversial for many years, as it
was considered to be partly responsible for the elevated phosphorus
levels and consequent eutrophication of the bay. In 2004 the Interna-
tional Joint Commission convened a task force to examine effects of
the causeway on water quality and water levels in the bay, and the
causeway was subsequently partly dismantled and replaced with a
modern bridge.
The restricted openings between bays focus water movement into
flows that, at times, can achieve substantial velocities and volumes,
such that fish passage would be either obstructed or facilitated,
depending on direction of travel. For example, 86% of the Missisquoi
Bay volume flows outward, mostly through the Alburg Passage and
Carry Bay; 84–88% of the volume of the Northeast Arm drains west-
ward through the Gut and Carry Bay, with the flow reversing during
periods of south wind; Malletts Bay flows approximately equally
into or out of the Northeast Arm to the north, depending on wind di-
rection; net flow between Malletts Bay and the Main Lake to the west
is 71% outward in calm winds, but 99% outward in a north wind, and
99% inward in a south wind (Myer and Gruendling, 1979). The effect
of these flows can be seen in the four-year delay between zebra mus-
sel (Dreissena polymorpha) colonization of the Main Lake, including
the northwest arm, and their appearance in the Northeast Arm and
Mississquoi Bay (Stangel and Shambaugh, 2005). Similarly, invasive
aquatic plants spread rapidly from the Northeast Arm into the Main
Lake, but slowly in the reverse direction (Countryman, 1975).Fig. 3. Enlarged view of the northern half of the lake showing causeways, bridges, and
major islands and landmarks mentioned in the text.
22 J.E. Marsden, R.W. Langdon / Journal of Great Lakes Research 38 (2012) 19–34
Shoreline alterations
Development of the Lake Champlain shoreline has increased the
amount of nearshore rocky substrate, in the form of retaining walls,
and on the lake bed, in the form of breakwalls and rip-rap covering
water intake lines. Standing and ruined breakwalls create potentially
valuable substrate for fishes; lake trout, in particular, use these struc-
tures extensively for spawning (Ellrott andMarsden, 2004). However,
this added substrate also has the potential to alter fish behavior and
distributions in ways that could lead to higher predation or fishing
pressure (e.g., Marsden and Chotkowski, 2001). Draining and filling
of wetlands along the shoreline has likely reduced spawning and lar-
val habitats for some species; an estimated 35-50% of the wetlands in
the basin have been lost (Lake Champlain Basin Program, 2004).
Chemical changes in the lake
Toxic chemical contamination has been minor, relative to other
major lakes and waterways in North America, due to the lack of large
industries in the basin. However, there is a paucity of published studies
on effects of contaminants on biota in the lake. Locations of elevated
contaminants in sediments include Cumberland Bay (PCBs, PAHs, cop-
per, and zinc), St. Albans Bay (copper from copper sulfate used to con-
trol nuisance algal blooms), Malletts Bay (arsenic and nickel), the barge
canal in Burlington (lead, mercury, silver, zinc, and PAHs), and the
South Lake near Ticonderoga (Myer and Gruendling, 1979; McIntosh,
1994; Gao et al., 2006). Elevated tissue levels of PCBs and mercury in
some larger, older piscivorous species have resulted in consumption
advisories for walleye and lake trout (Lake Champlain Basin Program,
2004). Ongoing chemical treatments for sea lamprey control, initiated
in 1990, pose some risk for rare and state listed non-target fishes and
mussels. Among the most sensitive fishes are channel darter, stonecat,
lake sturgeon, American brook lamprey, and eastern sand darter. Most
treatments have resulted in little or no observed mortality among
these species thus far.
Fishes of Lake Champlain
Fish surveys and monitoring
The earliest accessible records of the fishes in Lake Champlain
were the natural history writings of Zaddock Thompson (1853),
who recorded 65% of the current species, and the more rigorous sur-
vey by DeKay (1842). The most thorough and systematic survey was
conducted by the New York Department of Environmental Conserva-
tion under Emmeline Moore (Moore, 1930). Moore's survey encom-
passed the entire lake, and fish were sampled primarily using seines,
fyke nets, and gillnets. The work included a synoptic survey of fishes
in the lake, studies focused on smelt, gar, fish food habitats, and fish
parasites. Subsequently, periodic surveys offish communitieswere con-
ducted by the Vermont Department of Fisheries andWildlife (VTDFW)
in the 1950s and 1970s (Halnon, 1954; Anderson, 1978). Halnon (1954)
surveyed fish populations throughout the lake except the northwest
arm and Missisquoi Bay in 1953 and 1954 using gillnets, seines, trap-
nets, and rotenone; remarkably, most of this work, including deep-
water gillnet sets in the main lake, was conducted by a crew of four
(including two recent high school graduates) in a “14′ metal boat”
with either a 5 or 10 hp motor (Halnon, 1954). Unfortunately he did
not report the dimensions of his nets, though duration of some of the
gear sets was reported. Anderson (1978) fished throughout the lake
using a trap-net and graded-mesh gillnets, set vertically (2×145 m)
and horizontally (2.4×30m); he reported catch-per-unit-effort and av-
erage length and weight of each species in each catch. In addition, he
reported diet data for yellow perch, walleye, northern pike, chain pick-
erel, and smallmouth bass, and age and growth data for several species.
A long-term smelt monitoring program using trawling and, more
recently, hydroacoutics in the Main Lake, Malletts Bay, and Inland Sea
began in the early 1990s and is conducted annually by VTDFW; walleye
were monitored annually by VTDFW by seining in Missisquoi Bay from
1953 to 1966, then every five years since 1985, and are monitored an-
nually in the south lake; lake trout are monitored at two spawning
sites in the Main Lake using electroshocking and, since 2009, trapnets;
Atlantic salmon are alsomonitored at the same sites using electroshock-
ing, and at fish passages on the Boquet, Great Chazy, and Winooski
rivers; assessment of northern pike age and growth began in 2008; lar-
val sea lamprey and spawning adults have been monitored in multiple
tributaries and six deltas starting in 1982 (Marsden et al., 2003, 2010).
Results of these monitoring programs appear in annual reports of the
Lake Champlain Fisheries Technical Committee.
Unlike coastal and Great Lakes waters, historic data on commercial
fishery harvest in Lake Champlain are extremely scarce. Trioreau
(1985) provided data for lake whitefish harvest in the Quebec waters
of Mississquoi Bay, and Halnon (1963) reviewed historic documents
(newspapers, warden reports, town records, hatchery logbooks, and
biennial reports of the Fish and Game Commissioners) that include
some data on harvest of sturgeon, lake whitefish, and walleye in the
late 1800s and early 1900s. Sport harvest of several species is moni-
tored via angler diaries.
Fish species in the lake
Lake Champlain supports 72 native fish species; an additional 12
species inhabit the tributaries up to the first barrier. Carlson and Daniels
(2004) reported a total of 92 species for the entire New York portion of
the Champlain drainage, 16 of which were non-native. Their numbers
differed from thosewe report for several reasons. First, their tally includ-
ed non-extant species as well as stocked hybrids, where we report only
extant species. Second, they included the entire drainage whereas we
consider only the lake proper plus the tributaries up to the first physical
barrier (fall line). Third, their data set is limited to 2004; we have added
tench and alewife, non-native species not known to have occurred in
the lake prior to 2005. Finally, some species not recognized by those
two authors were left out of their original data set and have since
been added (Doug Carlson, personal communication). Carlson and
Daniels also list a total of 61 species from the lake proper in 2004,
where we list 75 currently. Their updated and corrected records appear
to have brought the New York State distribution database more in line
with our listing.
In surveys in the 1920s and 1953–54, deep-water communities
consisted of cisco, smelt, and burbot; these surveys occurred prior
to the return of salmonids into the lake by stocking (Moore, 1930;
Halnon, 1954). In shallower waters, the most common species were
yellow perch, walleye, smallmouth bass, rock bass, and pumpkinseed.
Currently, warmwater fish communities, characterized by largemouth
bass, pumpkinseed, rock bass, and white and black crappies, dominate
in the South Lake and Mississquoi Bay, and in the shallow wetland
areas that are mostly located along parts of the Vermont shoreline
(Langdon et al., 2006). Coolwater communities, including smallmouth
bass, northern pike, walleye, and yellow perch, are found throughout
the lake. Coldwater fishes, including lake trout, Atlantic salmon,
brown trout, steelhead trout, and rainbow smelt, are most abundant
in theMain Lake, with populations of some of these species also present
in the Northeast Arm and Malletts Bay (Fig. 1).
Writers in the 1800s and early 1900s distinguished between fish of
potential value to commercial fisheries and anglers (salmonids, lake
whitefish, walleye, and ‘cullfish’, which were primarily yellow perch,
bullheads, and suckers), and species considered to be ‘undesirable’,
‘destructive’, or ‘vermin’ (Thompson, 1853; Moore, 1930; Halnon, 1963;
Fish Commissioner reports). The fish commissioners of Vermont sought
authority to eliminate destructive fish from Lake Champlain (Titcomb
and Bailey, 1896); as late as 1930, Greeley (1930) stated that burbot,
bowfin, and gar were vermin and reported the “general opinion that
23J.E. Marsden, R.W. Langdon / Journal of Great Lakes Research 38 (2012) 19–34
Table
1
CurrentandhistoricaldesignationsforextantLakeChamplainfishspecies.Specieslistedarefromthelakeproperandtributarysectionsdownstreamofthefirstbarriertoupstreamfishmovement.
“Sa
me”initalicsisusedwherescientific
nameisthesameasiscurrentlyrecognized.“Same”withoutitalicsisusedwherecommonnameisthesameasiscurrentlyrecognized.Sources:1.Greeley(1930)2.EvermannandKendall189613.Thompson,1842,18534.Williams18092.
ScientificandcommonnameuseisaccordingtoNelsonetal.,2004.StatusindicateslistingbyVermont(VT),NewYork(NY),orCanada(C)asendangered(E),threatened(T),ofspecialconcern(SC),ornon-native.
Family
Currentcommonandscientificname
Commonandscientificnamesofhistoricaccounts
Status
Petromyzontidae
Northernbrooklamprey(Ich
thyomyzonfossor)
1–4.Notreported
VT-E,C-SC
Silverlamprey(Ich
thyomyzonunicuspis)
1.I.concolor,same2.Sam
e,same,mudlamprey3.A
mmocoetes
concolor,mudeel,blindeel34.Notreported
VT-SC
Americanbrooklamprey(Lampetra
appen
dix)
1–4.Notreported
VT-T
Sealamprey(Petromyzonmarinus)
1.Sa
me,lakelamprey,lampreyeel2.Sa
me,greatsealamprey,bluelamprey3.P.nigricans,bluelamprey4.Notreported
Acipenseridae
Lakesturgeon(A
cipen
serfulvescens)
1.Sa
me,same2.A.rubicundus,same,rocksturgeon.3.Sa
me-rock-,sharp-nosed-,sturgeon,
(regardedyoungofA.r.asA.oxy
rhynhus-round-nosedsturgeon)4.A.sturio,sturgeon
VT-E,NY-T
Lepistosteidae
Longnosegar(Lep
isosteu
sosseu
s)1.Sa
me,long-nosedgar,billfish,garpike2.Sa
me,long-nosedgar,billfish.3.L.oxy
uras,commonbillfish
(actuallyadultofL.osseu
s),L.linea
tus,stripedbillfish(actuallyyoungofL.osseu
s)4.Notreported
Amiidae
Bowfin(A
mia
calva)
1.Sa
me,same,scaledling,dogfish2.Sa
me,bowfin,mudfish3.A.ocellicauda,same4.Notreported
Anguillidae
Americaneel(A
nguilla
rostrata)
1.A.bostoniensis,same2.A.ch
rsypa,commoneel3.Mureanavulgaris,commoneel4.Muraen
aangulla,eel3
VT-SC
Clupeidae
Gizzardshad(D
orosomaceped
ianum)
1–4.Notreported
non-native
Bluebackherring(A
losa
aestivalis)
1–4.Notreported
non-native
Alewife(A
losa
pseudoharengus)
1–4.Notreported
non-native
Mooneye(H
iodontergisus)
1.Same,same,whiteshad2.Same,moon-eye,wintershad3.H
.clodalis,wintershad4.Notreported
NY-T
Salmonidae
Steelheadtrout(O
ncorhynch
usmykiss)
1–4.Notreported
non-native
Browntrout(Salm
otrutta)
1.S.
fario,same,Germanbrowntrout.2–4.Notreported
non-native
Atlanticsalmon(Salm
osalar)
1.S.s.salar,same-referredtosearun,believedextinct,(S.s.sebagoreferredtolandlockedvarietystockedfromMaine)
2.Same,commonAtlanticsalmon,notreported3.Same,salmon4.Salmo,salmon
Brooktrout(Salvelinusfontinalis)
1–4.Notreported
Laketrout(Salvelinusnamaycu
sh)
1.Cristivomer
namaycu
sh,same,notreportedbutlistedaspresent.2.Sa
me,same,notreportedbutlistedaspresent
3.Sa
lmonamaycu
sh,same,longe,togue4.Sa
lmosalar,salmontrout
Ciscoorlakeherring(C
oregonusarted
i)1.Leucichthys
artedi,same2.A
rgyrosomusartedi,same3.C.clupeaform
is,herringsalmon4.Notreported
Lakewhitefish(C
oregonusclupea
form
is)
1.Same,shad,commonwhitefish2.Same,commonwhitefish,andpossibly
C.labradoricu
s(musquawwhitefish)
3.C.albus,whitefish,lakeshad4.Notreported
Osmeridae
Rainbowsmelt(O
smerusmordax)
1.Sa
me,same,icefish2.Same,smelt,ice-fish3.O.esperlanus,smelt4.Notreported
Umbridae
Centralmudminnow(U
mbra
limi)
1.Mudminnow,U.limi,2.Same,same,mudfish3.Hydrargyra
fusca,mudfish4.Notreported
Esocidae
Redfinpickerel(Esoxamericanus)
1.E.vermiculatus4,little-,grass-,pickerel2–4.Notreported
Chainpickerel(Esoxniger)
1.Same,chain-,eastern-,grass-pickerel2.E.reticulatus,pickerel3–4.Notreported
Northernpike(Esoxlucius)
1.Same,same,pickerel2.Luciuslucius,commonpike,pickerel3.E.estor,commonpike4.Same,pike,pickerel
(didnotdistinguishfrommuskellunge)
Muskellunge(Esoxmasquinongy)
1.E.m
.masquinongy,muskalunge2.Luciusmasquinongy,maskallonge,muskallonge,mascalonge3.E.nob
ilior,masquallonge
4.Esoxlucius,muschilongoc(seenorthernpike)
VT-SC
Cyprinidae
Commoncarp(C
yprinuscarpio)
1.Sa
me,same,Germancarp.2–4.Notreported
Non-native
Goldfish(C
arassiusauratus)
1–4.Notreported
Non-native
Cutlipsminnow(Exo
glossum
maxillingua)
1.Sam
e,cut-lipsminnow,2.Sam
e,cutlipminnow3.Exoglossum
nigrescens,nonegiven4.Notreported
Longnosedace(R
hinichthyscataractae)
1.Sa
me,long-noseddace2–4.Notreported
Blacknosedace(R
hinichthysatratulus)
1–4.Notreported
Bluntnoseminnow(P
imep
halesnotatus)
1.Hyborhynusnotatus,blunt-nosedminnow2–4.Notreported
Fatheadminnow(Pim
ephalespromelas)
1–4.Notreported
Northernredbellydace(Phoxinuseo
s)1.ReportedinthedrainageinNew
Yorkbutitisunclearastoitsdistributionbelowfirstbarriertolake.2–4.Notreported
Finescaledace(Phoxinusneo
gaeu
s)1–4.Notreported
Fallfish(Sem
otiluscorporalis)
1.Leucosomuscorporalis,same,windfish,silverchub2.Sa
me,fallfish,silverchub3.Leuciscuspulchellus,commondace4.Notreported
Creekchub(Sem
otilusatromacu
latus)
1.Sa
me,horneddace,chub2–4.Notreported
Pearldace(M
argariscusmargarita)
1.M
.m.n
atchtrebi,reportedinatributaryofGreatChazyRiverpossiblynearmouth2–4.Notreported
Rudd(Scardiniuserythrophthalm
us)
1–4.Notreported
Non-native
Goldenshiner(N
otemigonuscrysoleucas)
1.N
.c.c.,same2.A
bramuscrysoleucas,shiner,roach3.Leuciscuscrysoleucas,shiner4.Notreported
Easternsilveryminnow(H
ybognathusregius)
1.Sa
me,same.2–4.Notreported
Brassyminnow(H
ybognathushankinsoni)
1–4.Notreported
VT-SC
Tench(Tinca
tinca)
1–4.Notreported
Non-native
Commonshiner(Luxiluscornutus)
1.Sa
me,andN.c.ch
rysocephalus-same,red-finshiner,(almostcertainlywascommonshiner)2–4.Notreported
Rosyfaceshiner(N
otropisrubellus)
1–4.Notreported
Emeraldshiner(N
otropisatherinoides)
1.Sa
me,lakeshiner,emeraldminnow2–4.Notreported
Spottailshiner(N
otropishudsonius)
1.Sa
me,spot-tailedminnow2–4.Notreported
24 J.E. Marsden, R.W. Langdon / Journal of Great Lakes Research 38 (2012) 19–34
Spotfinshiner(C
yprinella
spilopterus)
1.N
.whipplii
spilopterus,satin-finnedminnow,silverfin.2–4.Notreported
Mimicshiner(N
otropisvolucellus)
1.N.v.volucellus,nocommonnamegiven.2–4.Notreported
Sandshiner(N
otropisstramineu
s)1.N.deliciosusstramineu
s,straw-coloredminnow2–4.Notreported
Blackchinshiner(N
otropisheterodon)
1.Sa
me,nocommonnamegiven.2–4.Notreported
VT-SC
Bridleshiner(N
otropisbifrenatus)
1.Sa
me,Cayuga-,bridled-minnow2–4.Notreported
VT-SC,C-SC
Blacknoseshiner(N
otropisheterolepis)
1–4.Notreported
Catostomidae
Quillback(C
arpoides
cyprinus)
1.Sa
me,carpsucker,quillback2.Carpoides
thompsoni,lakecarp,buffalo,carpsuckerdrum3.Sa
me,carpsucker4.Notreported
VT-SC
Longnosesucker(C
atostomuscatostomus)
1.Sa
me,fine-scaled-,red-striped-,long-nosed-,sucker,(reportedintributaryclosetolake)2–4.Notreported
Whitesucker(C
atostomuscommersonii)
1.Sam
e,common-,whitesucker,mullet2.Sam
e3.C
.teres,sucker4.Sucker5
Silverredhorse(M
oxo
stomaanisurum)
1.Same,white-nosed-,red-finsucker,red-finmullet2–4.(mayhavebeenreportedasshortheadredhorse)
VT-SC
Shortheadredhorse(M
oxo
stomamacrolepidotum)
1.M
.aureolum,short-headed-,redfin-sucker,red
finmullet2.M
.aureolum,redhorsesucker,mullet
3.Catostomusoblongu
s,lakemullet,mullet4.Notreported
Greaterredhorse(Moxo
stomavalencien
nesi)
1–4.Mayhavebeenreportedasshortheadredhorse
VT-SC
Ictaluridae
Stonecat(N
oturusflavus)
1–4.Notreported
VT-E
Channelcatfish(Ictaluruspunctatus)
1.Sam
e,spottedcatfish,channelcat2.A
meiuruslacustris,GreatLakescatfish3.Pim
elodussp.4.Notreported
Yellowbullhead(A
meiurusnatalis)
1–4.Notreported
Brownbullhead(A
meiurusneb
ulosus)
1.Same,commonbullhead,bullheadcatfish,hornedpout2.A
meiurusvu
lgaris,bullpout3.Pim
elodusvu
lgaris,bullpout4.Silu
risfelis(?)pout
Blackbullhead(A
meirusmelas)
1–4.Notreported
Percopsidae
Trout-perch(Percopsisomiscomaycu
s)1.Sa
me,same2.P.guttatus,same3.Sa
lmoperca
pellucida,same4.Notreported
Gadidae
Burbot(Lota
lota)
1.L.m
aculosa,ling,skinling,eel-pout,lawyer,burbot,grodgeon2.L.m
aculosa,ling,methycusk3.Lotamaculosa,ling,methy4.Notreported
Fundulidae
Bandedkillifish(Fundulusdiaphanus)
1.F.d.m
enom
a,same,barredkillifish,graybackminnow2–4.Notreported
Atherinidae
Brooksilverside(Labidesthes
sicculus)
1–4.Notreported
Non-native
Gasterosteidae
Brookstickleback(C
ulaea
inconstans)
1–4.Notreported
Cottidae
Mottledsculpin(C
ottusbairdi)
1.Sam
e,sculpin,millersthumb(identifiedinfishstomachs)2–4.Notreported
Slimysculpin(C
ottuscognatus)
1.ReportedinthedrainageinNew
Yorkbutitisunclearastoitsdistributionbelowfirstbarriertolake.2–4.Notreported
Moronidae
Whiteperch(M
oroneamericana)
1–4.Notreported
Non-native
Centrarchidae
Whitecrappie(Pomoxisannularis)
1–4.Notreported
Non-native
Blackcrappie(Pomoxisnigromacu
latus)
1.Pomoxissparoides,same,calicobass2–4.Notreported
Non-native
Rockbass(A
mbloplitesrupestris)
1.Sa
me,same,goggle-eyebass2.Sa
me,same3.Cen
trarchusaneu
s,same4.notreported
Smallmouthbass(M
icropterusdolomieu)
1.Sam
e,small-mouthedblackbass,blackbass2.Sam
e,small-mouthedblackbass,blackbass3.C
entrarchusfasciatus,blackbass
(notstatedifinlakeorbelowbarriersintributaries)
Largemouthbass(M
icropterussalm
oides)
1.Aplitessalm
oides,large-mouthedblackbass2–4.Notreported
Non-native
Bluegill(Lep
omismacroch
irus)
1.Helioperca
incisor,bluegillsunfish2–4.Notreported
Non-native
Pumpkinseed(Lep
omisgibbosus)
1.Eupomotisgibbosus,commonsunfishpumpkinseed.2.Sa
me,3.Potm
otusvulgaris,sunfishpondperch,pumpkinseed,bream
4.(listed“bream”butgavescientificnameofamarinespecies).
Percidae
Yellowperch(Perca
flavescens)
1.Sa
me,same2.Sa
me,same3.P.serrato-granulata,commonperch4.P.fluviatalis,redperch
Walleye(Sander
vitreus)
1.Stizostedionvitreum,wall-eyed-,yellow-pike2.Stizostedionvitreum,wall-eyedpike,pike3.Lucio-perca
americanus,Americanpike-perch
4.Perca
lucioperca
Sauger(Sander
canaden
se)
1.Stizostedioncanaden
se,same,sandpike2.Stizosted
ioncanaden
se,sauger,groundpike-perch3.Lucioperca
canaden
sis,groundpike-perch
4.Notreported
Easternsanddarter(A
mmocrypta
pellucida)
1–4.Notreported
VT-T,NY-T,C-T
Logperch(Percinacaprodes)
1.Sa
me,same2.Sa
me,logperch,hogfish3.Etheo
stomacaproides,hogfish4.Notreported
Channeldarter(Percinacopelandi)
[VT.endangered,Canadathreatened],1.Cottagaster
copelandi,Copeland'sdarter2–4.Notrecorded
Tessellateddarter(Etheo
stomaolm
sted
i)1.Johnny,tessellateddarter,Boleostomanigrum
olm
sted
i,2.tessellateddarter,sameas1.3–4.Notreported.Note:Noclearhistorical
distributionsforE.olm
sted
iandE.nigrumareavailablebecauseoftheirmorphologicalsimilarityandresultingconfusioninaccurate
identification.
Fantaildarter(Etheo
stomaflabellare)
1.Catonotusflabellaris,fan-taileddarter2–4.Notreported
Sciaenidae
Freshwaterdrum(A
plodinotusgrunniens)
1.Same,Sheepshead,fresh-waterdrum2.Sa
me,sheepshead,fresh-waterdrum3.Corvinaoscula,sheep'shead
1.EvermannandKendall(1896)reportedlargelyupdatedscientificnamesoffishesfromThompson's(1853)collections.TheysampledonlytwolocationsinLakeChamplainandreportedrecordsandobservationsfromothers.
2.Williamslistedafewfishesas“menow”,“sucker”,and“dace”,whichundoubtedlycomprisedseveralcurrentlyrecognizedspeciesinCyprinidaeandCatostomidaewhichnowinhabitLakeChamplain.Additionally,heincluded“shiner“
(Perca
nobilis),“chub”(P.p
hiladelphia),and“bream”(P.chrysoptera)aspresentinVermontwatersingeneral.Thesethreelaterscientificnamesactuallyreferredtomarinespecies.
3.Thenamecommonname“blind”eelcoupledwiththelocationofrecordsofthisspeciesindicatethatThompsonwasdescribingalarvalform
(noeyesandresidesalongriverbanks)ofanyoffourcurrentlyacknowledgedlampreyspecies.
4.Esoxamericanusisthecurrentnamegivenforthespecieswhichisknownasredfinpickerel.E.vermiculatusisasubspeciesandisknownasgrasspickerel.ItisnotknowntowhichsubspeciesGreeleywasreferring.
25J.E. Marsden, R.W. Langdon / Journal of Great Lakes Research 38 (2012) 19–34
Table 2
Timeline of major habitat alterations, biological changes, and management activities in Lake Champlain and its watershed.
Year Habitat changes Biological changes Management activities and regulations
1800s Period of major dam construction - Great Chazy,
Little Chazy, Salmon, Little Ausable, Ausable, Boquet,
Winooski, Lamoille, Missisquoi, and Otter Creek and
Poultney River
1823 Champlain Canal opened reduction in stream fishes due to increased
stream flows and loss of structure
1838 Last record of Atlantic salmon, Ausable
River
1843 Chambly Canal opened
1846 St. Ours dam constructed
1849 St. Ours canal constructed
1850 Sandbar causeway constructed between Malletts
Bay and Northeast Arm, 1.6 km long
1851 Rouses Point railroad bridge constructed, 2.4 m long,
61 and 152 m openings
1855 Quebec enacts first fishery regulations
(gear and season restrictions)
1871 Larrabees Point railroad bridge built, with 91 m opening
1878 VT fall lake whitefish seining closed until 1883
1881 VT bans most commercial fishing gear except seines
1882 Isle LaMotte bridge constructed
1885 VT and NY prohibit seining
1886 Causeway constructed between Grand Isle and North Hero
(west side of the Gut), 300 m long, with 60 m
opening Bridge constructed across Alburg Passage
between North Hero and Alburg
VT reopens commercial fishery
1890 Funds committed for construction of first state
hatchery at Roxbury
1892 Bridge constructed between Grand Isle and North Hero
(east side of the Gut)
1896 Chambly dam built, with fish passage
1899 Island Line constructed, including causeways between Main
Lake and Malletts Bay ( 5.25 km long, with 24 and 53 m
openings); Alburg Passage ~300 m long with a 60 m opening;
along west side of the Gut, 610 m long with a 55 m opening
Period of fluctuating commercial fishery
closures in VT
1900 native lake trout no longer seen in lake
1903 Walleye hatchery in operation in Missisquoi Bay
1907 26 m opening in Sandbar causeway created
1911 St. Ours dam bypass constructed
1912 VT commercial fishery closed
1918 Quebec commercial fishery closed
1919-28 Smelt stocked from Cold Spring Harbor
1923 Seine fishing permitted in Mississquoi Bay by Quebec
1929 Biological survey of basin by NY
1938 Second bridge at Rouses Point constructed, 2.4 km long;
Missisquoi Bay causeway built, 1.5 km long with a
0.25 km wide opening
1950s Sporadic stocking of lake trout by NYSDEC and
VTDFW; Quebec commercial walleye fishery open
1964 Quebec commercial lake whitefish fishery reopened
1965-9 St-Ours, Chambly dams refurbished; fish passage not
replaced on Chambly
1967 St. Ours dam bypass no longer functional Lake sturgeon fishery closed
1971 Quebec commercial walleye fishery closed
1973 Beginning of high salmonid wounding
by sea lamprey
Sustained stocking of lake trout and Atlantic salmon
begins; walleye commercial fishery closed in 1970s
1982 Commercial fishery for American eel authorized in VT
using electroshocking and baited pots
1984 White perch first sighted in lake
1987 New Alburg-Rouses Point bridge constructed;
old causeway partially removed
1990 8-yr experimental control of sea lamprey begins
1993 Zebra mussels first sighted in lake
1997 Eel ladder added to Chambly dam
1998 Eel fishing in Quebec closed due to harvest decline
2001 Fish ladder and eel ladder added to St. Ours dam Long-term sea lamprey program begins
2002 VT eel permits repealed
2003 Alewife first sighted in lake, in Missisquoi Bay
2004 Lake whitefish in Missisquoi Bay no longer
commercially viable
2005 Missisquoi Bay bridge constructed, 1.5 km long Eel stocking begins by Quebec
2008 First major alewife dieoff in South Lake
26 J.E. Marsden, R.W. Langdon / Journal of Great Lakes Research 38 (2012) 19–34
the lake would be better off without them”. He goes on to suggest,
however, that even these speciesmight have a role in the balance of spe-
cies in the lake, indicating a shift in attitudes towards non-game species
and a broader understanding of ecology.
Biological changes in the lake
Species additions
In the 1800s, fish stocking was perceived as a vital role of fisheries
management agencies, and numerous non-native fish species were de-
liberately stocked at various times into Lake Champlain. The height of
stocking non-native species occurred between 1864 and 1892 when
common carp, Chinook salmon (Oncorhynchus tshawytscha), Atlantic
salmon, brown trout, steelhead trout, largemouth bass, bluegill, and pos-
sibly black and white crappies, and black bullhead, were present in Lake
Champlain and its lower tributaries; additional stocked species include
grayling (Thymallus arcticus), kokanee salmon (Oncorhynchus nerka),
and American shad (Alosa sapidissima) (Anderson, 1978, Langdon et
al., 2006). One million white perch fry were stocked into Near St. Albans
Bay in 1912 (Titcomb, 1912). All of these species except Chinook salmon
are still present in the lake, although black bullhead appear only inter-
mittently in one tributary (Langdon et al., 2006; Marsden and Hauser,
2009). Of these species, only brown trout and steelhead trout are still
stocked, and are considered to be an established component of the
fish community (Marsden et al., 2010). Both species naturally reproduce
in at least some tributaries.
Tench and alewife were introduced into the basin through unauthor-
ized aquaculture and unauthorized stocking by anglers, respectively;
rudd were probably introduced via a bait bucket, and goldfish appear
periodically in the basin from aquarium discards. The remaining non-
native species, gizzard shad, blueback herring, brook silverside, and
white perch, all appeared at the south of the lake first, and likely entered
from the Hudson River via the Champlain Canal (Marsden and Hauser,
2009). Walleye were reared in the Swanton hatchery beginning in
1903, with an annual egg take of 20 to 383 million eggs (Halnon,
1963). Modern stocking began in 1988 with annual stocking of 1.0–-
8.6 million fry and 12–182 thousand fingerlings (Marsden et al., 2010).
Annual or semi-annual stocking of lake trout began in 1958, Atlantic
salmon in 1962, steelhead in 1971, and brown trout in 1977 (Anderson,
1978). A coordinated program of sustained annual stocking of salmonids
began in 1972 (Marsden et al., 2010).
Few studies provide data on the effect of most of the non-native
fish species in Lake Champlain. Non-native fishes can impact native
species through predation, or competition for food or other re-
sources. Brown trout and steelhead trout may compete with
stocked lake trout for forage. Introductions of Micropterus spp.
into smaller lakes in the Adirondacks and south-central Ontario
have been reported to result in the loss of cyprinid and other spe-
cies (Findlay et al., 2000; Jackson, 2002). However, the impact of lar-
gemouth bass in Lake Champlain, where it was probably introduced as
early as the 1870s, is unknown. Largemouth bassmay competewith na-
tive predators by preying on small, mostly soft-finned species that favor
shallow water and vegetation, including golden and common shiner,
fatheadminnow, brook stickleback, and banded killifish; all of these spe-
cies are still widespread in the lake. White perch, which are now the
dominant species in Mississquoi Bay, are significant predators of fish
eggs, particularlywalleye eggs (Roseman et al., 2006), andmay compete
with yellow perch and cyprinids for forage (Parrish and Margraf, 1990,
1994). Predation by white perch on Daphnia in Mississquoi Bay may re-
duce phytoplankton grazing and contribute to the frequent nuisance
blooms of blue-green algae (Couture andWatzin, 2008). The three clu-
peid species and brook silverside are all also planktivores, with the po-
tential to compete with a variety of native planktivores and the young
of native species. Alewife, in particular, can reach extremely large popu-
lation sizes very rapidly. In Lake Champlain this species appeared first in
Mississquoi Bay in 2003, and by 2008, the first major die-off was seen in
the southern portion of the lake (Marsden andHauser, 2009). The sheer
numbers of this invasive planktivore threaten to alter zooplankton com-
position throughout the lake, thereby altering the food base for early life
stages of native fishes. However, the dramatic effects of alewife in the
Great Lakes occurred in the absence of native predators and the presence
of planktivore fish populations that had been severely depleted by com-
mercial fishing; in contrast, Lake Champlain piscivorous fish popula-
tions, although supported by stocking, were intact in Lake Champlain
when alewife arrived, and native rainbow smelt populations were ro-
bust.Whether alewife, under these conditions, will cause themagnitude
of problems seen in the Great Lakes remains to be seen.
Other non-native species have contributed to substantial physical
and trophic alterations to the lake. Effects of many of these species
have been well documented elsewhere (see Marsden and Hauser,
2009). For example, the zebra mussel (Dreissena polymorpha) fouls
benthic spawning habitats, suffocates native mussels, and has profound
impacts on benthic communities and nutrient cycling. Since their arriv-
al in Lake Champlain in 1993, native mussels have been virtually elimi-
nated frommuch of theMain Lake and seven unionid species have been
added to the list of Vermont threatened and endangered species; cur-
rently there are no mussel species in the lake on the New York threat-
ened and endangered list. Non-native, invasive macrophytes such a
Eurasian water milfoil (Myriophyllum spicatum) and water chestnut
(Trapa natans) are abundant in the lake, with the latter forming exten-
sive mats in shallow areas of the South Lake. When growing in dense,
monospecific expanses, both species provide generally poor fish habitat
(Valley et al., 2004; Hummel and Kiviat, 2004).
Commercial and sport fisheries
From the earliest period of European colonization, fishes were
harvested from Lake Champlain using shoreline seines, trap nets,
pound nets, fyke nets, hand lines, set lines, spears, and grappling
irons. Fishing was largely from shore, or in small boats; offshore gill-
nets appear not to have been used at all in the lake, although lake
sturgeon were fished with gillnets in rivers until 1888 (Brainerd
and Atherton, 1890). A study to determine the feasibility of using gill-
nets to commercially harvest lake whitefish concluded that it was too
difficult, due to the hardships imposed by weather and hauling nets
from deep water (Titcomb, 1912). This opinion is curious, given the in-
tensive deepwater, offshore gillnet fishery that was flourishing at the
time in Lake Ontario on the other side of New York state. Trawls were
prohibited in both states, and the Canadian waters of Missisquoi Bay
were unsuitable for this gear. Fishing was concentrated in Missisquoi
Bay, the South Lake, and the Lamoille and Winooski rivers, and near
the towns of Swanton, Alburg, and Burlington (Smith, 1898). Fall
harvest focused on lake whitefish and lake trout on their spawning
grounds, while the spring fishery focused on walleye, yellow perch,
and basses; other harvested species included bullheads, channel catfish,
American eel, northern pike, chain pickerel, rock bass, rainbow smelt,
Atlantic salmon, and lake sturgeon. Between 1893 and 1904, 62–94
licenses were issued per year in Vermont, and the fishery yielded up
to 70,000 fish annually (Halnon, 1963); unfortunately, comprehensive
catch records are not available for most of these species. Some of the
catch was shipped to New York and Boston, but much of the early fish-
ery appeared to be very localized, and the harvest was used by farmers
who fished, or was sold to neighbors (Thomas and Davis, 1904).
Lake whitefish, historically referred to as shad, shad waiter, or lake
shad in Lake Champlain, was one of the most important commercially
fished species, being harvested primarily in Mississquoi Bay, the
Northeast Arm, and the southern lake during fall when it aggregated
near shore for spawning. Lake whitefish were not considered to be a
sport fish and were regarded as a species that could not be caught
with hooks (e.g., Titcomb and Warren, 1892; Titcomb and Bailey,
1898, 1900; Titcomb et al., 1902). Curiously, this view continues to
27J.E. Marsden, R.W. Langdon / Journal of Great Lakes Research 38 (2012) 19–34
the present day, with only a few anglers who are aware even of the
presence of this species in the lake. Different writers have stated
that there were two species of whitefish in the lake, the second
being Maine or Labrador whitefish (Coregonus labradorensis) which
is currently not recognized as a valid species (Nelson et al., 2004).
In the late 1800s, a third species, the menominee or round whitefish
(Prosopium cylindraceum) was noted, but this species has not been
found in the lake since then, though it is present elsewhere in
Vermont.
Other major targets of commercial fishing were Atlantic salmon
and walleye. Atlantic salmon were once so abundant and so large
that, from a wagon driven into a shallow tributary, men could
“spear the salmon with pitchforks, and thus obtain in a few minutes
all the fish needed for consumption. Many of the salmon taken…
would reach twenty pounds in weight” (Watson, 1876). The same
document notes that horses and wagons, while fording a stream,
would be impeded by throngs of salmon. A single seine haul near Port
Kendall was recorded as netting about 1500 lb of salmon (Watson,
1869). They ranged throughout the northern lake, but did not spawn
south of the Boquet River in New York and Otter Creek in Vermont
(Edmunds, 1876; Greeley, 1930).
Walleye were harvested by seining in spring on their spawning
grounds in Mississquoi Bay and West Swanton. Harvest reported in
Fish and Game Commission biennial reports, converted to numbers
and weight of fish by Halnon (1963), indicate that the average weight
of walleye was 0.7 kg, and the harvest between 1893 and 1904 aver-
aged 38,584 fish annually (Table 3). In the 1950s, an average of
30,000 walleye were harvested annually in Mississquoi Bay (Table 4;
Halnon, 1963). Spring seining for walleyewas legalized again in Quebec
at some print prior to the mid-1950s, and a quota of 7000 fish was
imposed in 1961 (Table 3; Halnon, 1963). This commercial fishery
continued until 1971, when it was closed due to concerns about declines
in the harvest; the Vermont daily creel limit was reduced in 1978 for the
same reason.
Thefirstfishery regulations on the lakewere enacted by the Province
of Canada in 1855, and implemented seasonal closures for Atlantic
salmon, muskellunge, and ‘trout’ during winter (Oct. 1 to Feb. 1), and
gear restrictions (no stake or barrier nets, or use of lights, or nets with
meshes smaller than 2 inches). Early reports of the Fish Commissioners
of the state of Vermont advocated for similar regulations in Vermont
(Hager and Barrett, 1867), and by 1881 all pound-net, trap-net, gill-
net, set-net, set-line, and fyke net fishing was prohibited, and seining
was restricted to October 1 through November 15, with a minimum
mesh of 1.5 in. (3.7 cm) (Cutting and Brainerd, 1882). By the late
1800s, there was an ongoing debate as to whether the commercial fish-
ery should continue. Vermont commercial fishermen and legislators
advocated for closure of the commercial harvest, arguing that the state
was better served by providing fishes for tourist angling, thus bringing
a high economic benefit. Fishermen at this time were also concerned
about fish population declines, and lobbied for regulation/closure.
“The revenue from commercial fishing benefits comparatively few indi-
viduals who are inhabitants of Vermont, and the net value of any com-
mercialfisheries of the past is very small in comparisonwith themarket
value of the fish taken by anglers” (Titcomb, 1912). “The seining bene-
fits only a small number of men, who are not in any sense professional
fishermen, but generally farmers who seek through this means to add
something to their income” (Wakeham and Rathbun, 1897). The author
goes on to note, however, that while the catch was very small, and clo-
sure would not result in particular hardship, the fishery appeared to do
no damage and therewas no reasonwhy it should not continue— espe-
cially as Canada particularly desired a fishery. During the periods of clo-
sure, poaching for lake whitefish, walleye, and probably other species
continued to occur; illegally harvested fish and illegal seines were reg-
ularly seized (Halnon, 1963).
Fall seining for lake whitefish was closed in Vermont from 1878 to
1883, and in 1885, Vermont and New York entered into a treaty to
prohibit seining; the New York fishery did not re-open. For a few
years, the discovery of seine fishing by Canadians in Missisquoi Bay,
by fishermen who had been issued a license by Canadian authorities
in direct contravention of the previous agreement, caused consterna-
tion on the part of Vermont fishermen and resulted in the reopening
of the fishery after some conflict between the two countries (Halnon,
1963). The commercial fishery in Vermont reopened in 1892 and was
closed again in 1899. After a period of fluctuating reopenings and pro-
hibitions, the fishery was finally closed permanently in 1912; seining
in Quebec was prohibited in 1918 (Thomas and Davis, 1904; Halnon,
1963). The Quebec whitefish fishery reopened in 1964, closed in 1970
due to concerns about mercury in fish flesh, and reopened in 1971;
this fishery remains open, although the harvest and number of
licenses have declined steadily and fishing apparently ceased in the
mid-2000s due to small harvests (Fig. 4; Trioreau, 1985; KennyMiller,
commercial fisherman, personal communication).
Despite the absence of a commercial fishery, other forms of exploita-
tion continued to impact fish populations. Van Oosten (1933) expressed
the opinion that angling and poaching for Atlantic salmon and lake
trout in Lake Champlain was sufficient to prohibit attempts to restore
these species; he noted in a letter that yellow perch were also severely
depleted by angling (Van Oosten, 1933). The lake sturgeon fishery was
closed in 1967, and commercial fishing for American eel, briefly
reopened in 1982, was closed in 1998 (Marsden et al., 2010).
Consequences of changes in Lake Champlain
By the end of the 19th century, Lake Champlain had experienced
habitat fragmentation (due to dams and causeways), physical habitat
degradation (siltation and shoreline alteration), and localized eutro-
phication. In addition, populations of several fish species had been
Table 3
Commercial harvest of walleye in spring seining in Quebec waters of Mississquoi Bay,
Lake Champlain, from 1893 to 1904 and 1954 to 1961. A quota of 7000 walleye was
imposed in 1961. Na — data not available. Data from Halnon (1963).
Year Number of walleye Total weight (kg)
1893 22,200 17,663
1896 13,200 8782
1897 47,775 31,734
1898 45,150 30,037
1901 32,500 21,655
1902 68,500 45,605
1903 50,850 33,829
1904 28,500 18,960
1954 42,720 na
1955 28,344 na
1956 33,571 na
1957 32,053 na
1958 23,728 na
1959 22,394 na
1960 24,800 na
1961 7000 na
Table 4
Species reported historically in Lake Champlain that are not currently recognized in
lake or its tributaries up to the first barrier.
Current species designation Name given and source
Iowa darter
(Etheostoma exile)
Poecilchthys exilis (Greeley (1930)
Round whitefish
(Prosopium cylindraceum)
Evermann and Kendall (1896) had “no doubt” of its
presence in Lake Champlain but did not collect it.
(Greeley, 1930) stated it was present by citing an
1894 record without naming the source, but did not
collect it
Not currently recognized Lake catfish Vallaris lacustris (Greeley, 1930)
Not currently recognized Labrador whitefish or Maine whitefish Coregonus
labradorensis (Titcomb and Bailey, 1896)
28 J.E. Marsden, R.W. Langdon / Journal of Great Lakes Research 38 (2012) 19–34
reduced by harvesting. The effects of habitat change and exploitation
were seen first, and most severely, among the coldwater, pelagic, pi-
scivorous species. In contrast, warmwater fish populations have
remained healthy and, in recent years, have begun to attract increas-
ing numbers of fishing tournaments (82 during 2008). The potential
impact of this increased angling may be a future cause for concern.
Eutrophication in areas such as Missisquoi Bay may benefit some
nonnative species, such as white perch (Hawes and Parrish, 2003).
Habitat fragmentation
Fragmentation of fish habitats in the Lake Champlain basin is a re-
sult of barriers in the lake proper (i.e., causeways) and barriers that
impact species using tributaries for spawning (i.e., dams). Whereas
dams create highly visible barriers to fish migrations, the effect of
causeways on fish movements in Lake Champlain is barely under-
stood and even less apparent. On one hand, a study of sea lamprey
movements indicated that they traveled through one or more cause-
ways in their 18-month lake residence as parasites; this movement
likely reflects the movements of their host fish while the lamprey
are attached (Howe et al., 2006). In contrast, rainbow smelt population
abundances in Malletts Bay, the Northeast Arm, and theMain Lake vary
independently in some years, which suggests that the populations are
not readily intermixing (Fisheries Technical Committee, 2009).Walleye
apparently moved along the eastern shore of the lake between Grand
Isle and the mainland, to reach spawning areas in the Lamoille and
Mississquoi rivers and in Mississquoi Bay; the Sandbar causeway was
believed to block their access to the Lamoille River, and necessitate a
longer, westward journey to reach Mississquoi Bay (Halnon, 1963).
The effects of dams are, in some cases, obvious — both Atlantic
salmon and lake sturgeon declined in part because access to their
spawning sites was blocked. For example, lake sturgeon harvest was
never substantial, but their abundance declined steadily through the
1900s, presumably as a result of decreased reproduction in addition
to harvest (Halnon, 1963). Prior to 1913, annual commercial harvest
of lake sturgeon averaged over 100 fish. In 1895, 6975 pounds of
lake sturgeon were harvested by spears and grapples, presumably
during spawning migrations in rivers. By the mid-1900s less than
15 fish were caught per year, and the fishery was closed in 1967.
The decline of lake sturgeon populations is probably attributable
largely to degradation of spawning substrate and dams that blocked
access to spawning areas. Similarly, declines in the number of Atlantic
salmon were seen by the early 1800s and were attributed variously to
dams, pollution from forestry and sawdust, deforestation that re-
duced stream volume, increased run-off volume and velocity, in-
creased stream temperatures, reduced large woody debris used for
refuge, seining and spearing near river mouths during spawning,
and disturbance from boats and other human activities (Thompson,
Fig. 4. A) Commercial harvest of lake whitefish in Mississquoi Bay, Lake Champlain, from 1885 to 2004, in kg; note break between 1913 and 1965 during closure of the fishery.
B) Commercial harvest of lake whitefish per license, as an approximation of catch per unit effort, from 1965 to 2004; consistent license data are not available prior to 1965.
Data from Halnon (1963) and Trioreau (1985).
29J.E. Marsden, R.W. Langdon / Journal of Great Lakes Research 38 (2012) 19–34
1824; Edmunds, 1876; Evermann and Kendall, 1896; Greeley, 1930;
Van Oosten, 1933). The last salmon reportedwas caught in the Ausable
River in 1838 (Van Oosten, 1933).
Of even less certainty is the potential effect on Lake Champlain
fishes of the St. Ours and Chambly dams on the Richelieu River, con-
structed in the 1800s. American eel was very likely impacted both by
the dams and the targeted eel fishery associated with the dams in the
Richelieu River; eel harvest in the Richelieu River between 1920 and
1980 averaged 34.6 mt. Rebuilding of both dams in the 1960s was
rapidly followed by a dramatic decline in American eel populations
in the lake, and commercial harvest declined to 4.7 mt by 1987
(LaBar and Facey, 1983; Verdon et al., 2003). Early authors speculat-
ed that both Atlantic salmon and rainbow smelt were largely, or in
part, supported by anadromous populations. Reports of salmon mi-
grations into the lake via the Richelieu River are largely unsubstan-
tiated, often second-hand, and may be based on misinterpretations
of place names; for example, Webster (1982) cites Clinton (1822)
as saying salmon ascended the “Champlain River”, which may have
meant the Richelieu or Great Chazy River. Follett (1932) stated that
salmon were “abundant in the Upper St. Lawrence and its tributaries.
They entered Lake Champlain and its tributaries, the Saranac River at
one time being famous for the abundance of salmon.” — however,
writing almost 100 years after the disappearance of this species
from the lake, it is unclear from where he drew this information.
Williams (1809) thought salmon entered Lake Champlain in spring,
being found in tributaries from early May to the middle of June,
and returned to the ocean in the end of September; these salmon
weighed as much as 35 or 40 lb (16–18 kg). Certainly, salmon
spawned in tributaries of the St. Lawrence River; whether they
ever ascended through the Richelieu to Lake Champlain and then
sought tributaries of the lake in which to spawn may never be
known.
Rainbow smelt occurred historically in Lake Champlain as two
‘races’, a normal and a giant race (Webster, 1982). A. N. Cheney
reported that smelt at Port Henry were “over 1 ft in length and
weighing ½ lb each” and he was told that even larger ones were
caught (Evermann and Kendall, 1896) Several writers have suggested
that smelt were anadromous in Lake Champlain: Zadock Thompson
(1853) describes smelt as anadromous, and an occasional visitor to
the lake; Murray (1890) stated he was “informed that they were
comparatively new to Lake Champlain”; Edmunds (1876 ) noted
that smelt move up the Richelieu River and at the Chambly rapids
“are taken in great abundance in their midwinter journey to the
lake”; the 1876 report of the US Commissioner of Fish and Fisheries
stated that smelt were marine, and almost unknown to the local fish-
ermen, “but in late years, it is often taken in vast quantities through
the ice, while in some season it is rarely seen”. Cheney presumed
they were anadromous, entering the lake regularly via the Richelieu
River, noting that Dr. Hugh Smith wrote, “the specimens of ice-fish re-
cently sent to us from Lake Champlain were the salt-water smelt
(Osmerus mordax)” (Evermann and Kendall, 1896). Unlike smelt in
most other areas, smelt in Lake Champlain do not ascend tributaries
to spawn in spring, but spawn in waters at least 15 m deep, and prob-
ably deeper (Plosila, 1984; JEM, unpubl. data). It is possible that the
lake once contained both land-locked (normal race) and anadromous
(giant race) smelt. However, 59 million smelt were stocked from the
Cold Spring Harbor hatchery in New York between 1919 and 1928
(Greene, 1930), and were presumed by some to be the origin of the
small race (Greene, 1930; New York Forest, Fish, and Game Commis-
sion, 1906). In the last several decades the proportion of giant smelt
has apparently steadily decreased. Carlander (1969) reported that
“slower-growing” smelt comprised 30% of the population in 1929,
but declined to 4% by 1950, and giant smelt have been rarely noted
in recent trawls. If the larger race were, in fact, anadromous, their
gradual disappearance could be attributable in part to the two dams
and inactive fishways on the Richelieu River.
Habitat degradation
The areas most severely impacted by physical and chemical
habitat change are Mississquoi Bay and the South Lake; both
areas are now highly eutrophic and have substantial accumulations
of silt. In addition, as of the late 1990s, substrates in the South Lake
have been heavily colonized by zebra mussel, such that in some
areas the soft sediment is largely comprised of dead zebra mussel
shells to depths down to 15 cm or more (JEM, pers. obs.). Lake
whitefish may have been one of the first species to be affected by
these changes. Historically, commercial fishing during the fall sea-
son was focused in Mississquoi Bay, the South Lake, and near
Alburgh and Swanton, presumably because fish were aggregated
nearshore for spawning, and these areas were particularly accessi-
ble for seining (Titcomb and Bailey, 1898; Van Oosten and Deason,
1939; Trioreau, 1985). Fall fishing grounds were also documented
in other areas of the lake, including Keeler Bay, Knights Island and
Butler Island in the Northeast Arm, Isle La Motte, and south of
Otter Creek (Thomas and Davis, 1904). Recent research indicates
that lake whitefish currently spawn extensively along the Vermont
shoreline of the Main Lake and the west side of Grand Isle (Herbst,
2010). However, no evidence of spawning has been found since
2006 in Mississquoi Bay and the South Lake, and few larvae were
found in the southern half of the Northeast Arm, including Keeler
Bay (Herbst, 2010). Physical conditions in these areas, i.e., dense
silt and organic material in the substrate, would likely no longer
support successful egg incubation. Causeways may have also con-
tributed to lack of movement between the Main Lake and spawning
sites in the Northeast Arm.
Changes with multiple or unknown causes
Understanding the effect of the alterations in Lake Champlain on
additional fish biota is hampered by an imperfect record of historical
native species abundance and distribution. For example, virtually
nothing is known about lake trout prior to their disappearance from
the lake by the late 1890s. The commercial fishery was focused in
fall when lake trout aggregated to spawn close to shore but, given
the variety of locations where lake trout spawn currently, including
sites that are inaccessible from shore, it seems unlikely that this
small, nearshore fishery could have extirpated the entire species. Na-
tive lake trout may have also spawned in deep water, but these sites
appear to have been degraded by anthropogenic inputs — the deep-
water and offshore reefs that have been explored recently are heavily
covered with silt (Ellrott and Marsden, 2004, unpubl. obs.). Currently,
stocked lake trout spawn at multiple sites along the lake shore, on
natural reefs and artificial structures, but there is virtually no recruit-
ment to the juvenile or adult populations (Ellrott and Marsden, 2004,
unpubl. data). The addition of alewife to the diet of lake trout and
Atlantic salmon has led to thiamine-deficiency in eggs collected for
hatchery rearing, and consequent earlymortality syndrome in hatchery
fry (Fitzsimons et al., 1999; unpublished data); the effect on naturally-
produced fry has not yet been examined.
Declines and changes in coolwaterfish populations are also puzzling.
Assessment of the walleye population in Mississquoi Bay using spring
seines shows a substantial and sustained decline in catch-per-unit-
effort since the 1960s (Fig. 5). Factors contributing to this decline, in
addition to harvest, likely include degradation of spawning habitat
and, more recently, competition with and larval predation by non-
native species such as white perch. Muskellunge were present in the
northern and central portions of the lake, but disappeared by the late
1970s. Sauger were common in the southern end of Lake Champlain
in the 1953–54 (Halnon, 1954) and Anderson (1978) collected them
in all areas of the lake except the Main Lake in 1971–1977, but this
species has not been seen in the lake in the last decade.
30 J.E. Marsden, R.W. Langdon / Journal of Great Lakes Research 38 (2012) 19–34
Current status of the fishery and non-game fishes
Sport and commercial fishes
Lake Champlain currently supports an active sport fishery, with a
small number of charter fishing boats. Commercial fishing is licensed
only in the Quebec portion of Missisquoi Bay, and has not been active
since the mid-2000s; in U.S. waters, angler-caught fishes and seined
bait fishes comprise a small market fishery. A survey in 1991 indicat-
ed that 8.2% of Vermont anglers and 18.7% of New York anglers
(among a total of 6,175 anglers) sold 338,767 kg of fish; the majority
(84.2%) were yellow perch, 10.2% were rainbow smelt, and the re-
mainder were centrarchids (Doug Facey, Saint Michael's College,
unpublished data). The coldwater fishery is now supported almost
entirely by annual stocking of 68,000–90,000 yearling lake trout,
78,000 steelhead, 68,000 brown trout, 240,000 Atlantic salmon
smolts and 450,000 Atlantic salmon fry. Up to 8.6 million walleye
fry and 182,000 fingerlings are also stocked annually (Marsden
et al., 2010). Nearshore angling focuses on yellow perch, white
perch, northern pike, basses, and sunfishes, and smelt are fished
under the ice in early spring. Popularity of bass fishing, in particular,
has increased dramatically, with increasing numbers of angling tour-
naments held on the lake each year. Sea lamprey control, initiated in
1990, resulted in wounding rates decreasing substantially after 2006,
with concurrent increases in size of angled lake trout and Atlantic
salmon (Marsden et al., 2010).
Non-game fishes
Records documenting the status of small and tributary fish species
are sparse; not surprisingly, many minnow species and other small
species went unrecognized until the 1900s and some were not recog-
nized until the last 30 years. Of the 72 currently knownnative species in
the lake, Thompson (1853) recognized a total of 33 species and Greeley
(1930) recognized 63 species. Some species noted in historic documents
are no longer recognized, such as Labrador whitefish, or have been con-
strued to be synonymous with other species (Table 4). Of the 23 known
native cyprinid species in Lake Champlain and its tributaries below the
first barrier, Greeley (1930) recognized 20 and Thompson (1853) only
four. Lesser-known fishes tend to be less than 15 cm in length as adults
and are either rare in the lake or not targets of commercial or recreational
fishing. Fish taxonomywas still developing in the late 1800s andmany of
these small species were not identified until the early 1900s. Many
species may have simply been confused with similar-looking species. Ex-
amples are blacknose dace, brassy minnow, channel darter, eastern sand
darter, mimic and sand shiners, slimy sculpin, northern redbelly dace and
American brook and northern brook lampreys. Some cyprinid species
were just regarded as shiners or minnows, without specific species
being identified (e.g., Halnon, 1954). Although redhorse suckers, genus
Moxostoma, are less obscure, as they commonly grow to lengths
exceeding 60 cm, the three species currently found in Lake Champlain
(shorthead, greater and silver) have not, until recently, been consistently
distinguished as individual species by most workers.
Because so little was recorded of lesser known, smaller species, lit-
tle can be said of their historic and current trends in distribution and
abundance. Not until Scott and Crossman (1973) and Smith (1985)
was there any semblance of a basic distribution published for the
darters and sculpins in Lake Champlain. Many of the rare species
are now state listed as threatened or endangered; all of these species
except lake sturgeon and sauger are small as adults, occur in the east-
ern edge of their range, and are considered to be intolerant to chem-
ical and physical habitat degradation. Increased sediment loading
and chemical runoff from historic manufacturing and current urban
stormwater and agricultural pesticides may have reduced abundance
of these species, and may continue to stress tributary populations of
minnows and darters. It may be, then, that following European colo-
nization in the late 1700s, abundance of some of these species has
dwindled and/or their distribution has become more restricted with-
in the lake and the lower reaches of its tributaries. However, new
populations of some species, including channel darter, yellow bullhead,
and stonecat, have been found in recent years. These new records are
probably a result of focused sampling in appropriate habitats with
equipment that targets small fishes, rather than expansion of their
ranges.
Sea lamprey
The sea lamprey became a serious fisheries management issue
soon after the beginning of the salmonid stocking program in 1972.
High wounding rates on stocked lake trout and Atlantic salmon
necessitated a control program, which was initiated experimentally
in 1990 (Marsden et al., 2003). Wounding rates in Lake Champlain
have ranged from 31 to 98 types AI–AIII wounds per 100 lake trout;
in the Great Lakes, except in areas influenced by the St. Mary's River
lamprey population prior to control, wounding rates rarely exceeded
20 wounds/100 lake trout (e.g., Heinrich et al., 2003; Lavis et al.,
2003).
There is no unanimity on the issue of the origin of sea lamprey in
Lake Champlain. Like rainbow smelt and Atlantic salmon, this lam-
prey had natural access from Atlantic Ocean to the lake since post-
Fig. 5.Walleye catch per unit effort (# walleye per seine haul) from 1953 through 2010 at Sandy Point, Mississquoi Bay, Lake Champlain. Data courtesy of Bernie Pientka, Vermont
Department of Fisheries and Wildlife.
31J.E. Marsden, R.W. Langdon / Journal of Great Lakes Research 38 (2012) 19–34
glacial times via the Richelieu River outlet. This connection provided
an especially easy route in the early stages of Lake Champlain before
isostatic rebound resulted in the formation of falls and cascades on
the Richelieu River. Recent genetic studies suggest that the popula-
tion is native to Lake Champlain (Bryan et al., 2005; Waldman et al.,
2006). However, the absence of historical recorded identification of
sea lamprey, coupled with the opening of the Champlain Canal in
1823 that provided a theoretical access from the Hudson River, has
been used as a rationale for a non-native status, similar to the argu-
ment used for their non-native status in Lake Ontario (Eshenroder,
2009, rebutted by Waldman et al., 2009). While the weight of evi-
dence for the sea lamprey's native status is strong, a designation of
status does not currently affect the ability of fisheries agencies to re-
ceive the required permits for chemical control in Vermont or New
York.
The assumption that the sea lamprey is native raises some intriguing
questions. First, is the current predator–prey relationship (characterized
by seemingly high wounding rates) a result of human-induced ecologi-
cal changes or is it simply natural?With no historic accounts of lamprey
scarring rates on coldwater species, this distinction is difficult to deter-
mine. Assessment data indicate that wounding is high relative to other
areas where sea lamprey are either native or introduced. Variation in
ecosystem condition and history among these lakes minimizes the use-
fulness of any meaningful comparison, however. We also caution that
the yardstick currently used to assess impact to target populations
may be based more on coldwater fishery performance expectations
than on purely ecological terms.
A follow-up question would then be: if one concedes that wound-
ing rates are indeed abnormally high, then what environmental fac-
tors could be responsible for the apparent imbalance between
lamprey and their prey?We propose three explanations. First, current
conditions in the larval habitat in tributaries probably differ greatly than
those during and before the mid 1800s. With settling of the land, agri-
cultural and urban development has led to increased sedimentation in
the tributary rivers, providing additional suitable substrate for ammo-
coetes. Also, tributary nutrient levels have undoubtedly increased over
the last 100 years, providing greater levels of suspended material
that provide the filtering ammocoetes more nourishment, increasing
growth rates and possibly shortening the length of time in the larval
stage. The construction of on-stream ponds and impoundments may
also increase the available organic material downstream, adding to
the food base.
Second, predation on ammocoetes may have decreased. Larval
lamprey are susceptible to predation during their 4- to 6-year resi-
dence in stream sediments, and during their downstream migration
to the lake as swimming transformers. Little is known about freshwa-
ter predators of sea lamprey; however, any predator that spends
at least part of its life in streams has the potential to affect larval
densities. Potential predators include lake sturgeon, which forage by
sieving substrate, and American eel, which are capable of burrowing
into substrates and have been observed capturing American
brook lamprey ammocoetes in a laboratory experiment (Perlmutter,
1951). Both species declined substantially in Lake Champlain prior
to the increase in sea lamprey. In addition, Atlantic salmon, which
do not generally feed while ascending streams to spawn, were
historically present in streams in reportedly high densities during
the period when sea lamprey transformers are descending toward
the lake. Predation on the vulnerable out-migrating transformers
would certainly have been an energetically beneficial strategy for
salmon. Spawning lamprey may also be vulnerable to predation
during their upstream migrations. Potential predators include
northern pike, walleye, and several fish-eating mammals and birds
(Scott and Crossman, 1973); of these, in Lake Champlain, walleye
and osprey have experience marked reductions in population size,
as have muskellunge. Predation by these species could also have
taken place in during out-migration as well.
A third possibility is that current lake trout stocking rates are sup-
porting an artificially high parasite population and maintaining an un-
natural host-parasite relationship. The original naturally-reproducing
lake trout population may have been historically less abundant, or
the adults may have been less likely to be attacked by lamprey. If sea
lamprey are native, then the native strain of lake trout co-evolved
with them, and would have developed either avoidance strategies,
depth preferences that led to lower spatial overlap with sea lamprey,
or resistance to sea lamprey-induced mortality, as seen in the Seneca
Lake strain (Schneider et al., 1996; Bergstedt et al., 2003). Although na-
tive strains of lake trout have been extirpated from Lake Champlain,
recent analysis of sea lamprey wounding data in Lake Champlain sug-
gests that sea lamprey growth and lethality of attacks on lake trout
were lower than in Lake Huron; this may be a characteristic of a histor-
ic stable parasite:host relationship (Madenjian et al., 2008). Most likely,
some of the aforementioned possibilities could have acted simulta-
neously to have caused the current phenomenon.
Management implications
The aquatic communities of Lake Champlain are substantially
changed from those of pre-European colonization. There may have
been fish species present then that were never recorded, and have
disappeared without our knowledge. The 72 extant native species in
the lake and its tributaries up to the first barrier have been joined
by 15 non-natives, increasing the total to 87 species. The presence
of water chestnut, Eurasian water milfoil, zebra mussel, white
perch, alewife, and other established non-native species has incurred
largely unknown biological changes on the native biota. Chemical and
physical changes to the lake, of which some are irreversible, at least in
the short term (decades), include phosphorus accumulations in the
sediments and sediment loading both in tributaries and in the lake it-
self. Nevertheless, some signs of fish population recovery have been
noted. Collection of eggs and larvae indicates that lake sturgeon are
spawning in the Winooski, Lamoille, and Missisquoi Rivers; stocking
of elvers in the Richelieu River since 2005 has resulted in substantial
numbers of American eel appearing in warm-water fisheries assess-
ments; lamprey wounding of lake trout and Atlantic salmon has de-
creased substantially (Marsden et al., 2010).
Very little is known about the population status of a large number
of native and non-native species in the lake, including some predators
such as burbot, bowfin, smallmouth bass, largemouth bass, and the
four esocid species. Currently, substantial management effort is di-
rected toward restoring the native game species (lake trout and
Atlantic salmon), lake sturgeon, and American eel, protecting habitat
(reduction of phosphorus and sediment inputs), and reducing the
risks from existing and new non-native species (water chestnut con-
trol, bait fishes and plant quarantine regulations, changes in stocking
practices to avoid spreading disease vectors, discussion of a biological
barrier on the Champlain Canal). Additional activities that are possi-
ble, but are politically, socially, or economically infeasible, include
removing dams, removing causeways, and removing non-native sal-
monids by discontinuing stocking.
Ultimately, as managers struggle with maintaining a coldwater
fishery and improving water and habitat quality in the lake, people
are also making choices (directly and indirectly) about what our biot-
ic community will look like; which species will we favor for recrea-
tional purposes, and which species will we attempt to limit or put
at risk in that effort?What importance will we attribute a more holistic
management approach in this process?
In the past 20 years, traditional fisheries management practices
have begun to be integrated with concerns over the ecological bal-
ance of aquatic systems. The concept of ecosystem management
has evolved in association with that of biodiversity and biological
integrity (e.g., Larkin, 1996). Increasingly, state and federal water
quality standards are being refined to more specifically address
32 J.E. Marsden, R.W. Langdon / Journal of Great Lakes Research 38 (2012) 19–34
biological criteria, with the premise that the near-native or natural
condition is desirable (Shelton and Blocksom, 2004). Givenmyriad bio-
logical, physical and chemical changes in Lake Champlain, it seems un-
likely that the original structure and function of the lake's communities
can ever be fully restored. Nevertheless, managing native species can be
less costly and provides a higher level of, and more predictable, ecosys-
tem services (e.g., Holmlund and Hammer, 2004). While altered from
the original assemblage of fishes, the lake still represents a viable, sus-
tainable system that we believe could still best support native species.
Therefore we suggest that long-term management of this resource be
primarily aimed at restoring native species.
However, due to insufficient resources and ecological knowledge,
recent restoration efforts tend to be approached in a piecemeal,
species-by-species basis without significant regard for the ecosystem
as a whole. To truly apply ecosystemmanagement, we must go beyond
what is implied by the term “fisheries management” and address a
broader range of objectives, not only fisheries related, but alsowith con-
sideration of the intrinsic value of native assemblages. Developing and
sustaining this approach would necessarily include a broader range of
expertise than fisheries managers.
Most efforts to control pollution from excessive nutrients, toxins,
and sediment improve conditions for all aquatic species and thus ad-
dress fisheries interests as well as ecologically based values. However,
some current management efforts that are aimed at improving the rec-
reational fishery are at odds with the objectives of ecological balance.
For example, the control of sea lamprey with the use of chemicals im-
pacts native lamprey species and may negatively affect other rare and
state listed species. Managing the lake exclusively for fisheries or exclu-
sively towards some sort of ecological balance would result in one or
the other concern being ignored. Managers consequently need to bal-
ance the risks to listed species with the need for a successful coldwater
fishery, or, in a larger context, manage to balance all of the values asso-
ciated with the resource as a whole under an ecosystem management
approach.
The difficult challenge for resource managers is to find a defensible
guiding set of priorities that balance the fisheries and ecological health
goals in proportion to thewishes of Vermont, NewYork andQuebec cit-
izenry. While this suggestion may sound simple and trite, many man-
agement activities will result in unfavorable compromises for one of
the two interests. The recent Strategic Plan for Lake Champlain Fisheries
strives to establish such guiding principles, while recognizing the ten-
sions that exist between the different groups involved in management
and use of the lake (Marsden et al., 2010). To facilitate informed discus-
sions between angling interests and those more concerned with
ecosystem-level management, resource managers and scientists are
working to provide a clear picture to the public of what the current sta-
tus of the resource is and clearly statewhatwill be gained andwhatwill
be lost by implementing every future management action for Lake
Champlain.
Acknowledgments
We thank Albert Joy and his colleagues at the Bailey-Howe Library
at the University of Vermont, and the State library in Montpelier for
assistance in finding resources and access to historic documents.
We are grateful to Leslie Morrissey at the University of Vermont and
Stephanie Strouse at the Lake Champlain Basin Program for producing
the maps of Lake Champlain.
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