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: ellen.marsden@uvm.edu (J.E. Marsden),

Rich.Langdon@state.vt.us (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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