Site by Bennett Web & Design Co.
2008 NORTHEASTERN NATURALIST 15(1):97–110
Aquatic Plant Communities in Waneta Lake and
Lamoka Lake, New York
John D. Madsen1,*, R. Michael Stewart2, Kurt D. Getsinger2,
Robert L. Johnson3, and Ryan M. Wersal1
Abstract - A point-intercept survey was implemented in August 2000 to determine
the distribution and richness of aquatic plant species present in Waneta Lake
and Lamoka Lake, NY. Myriophyllum spicatum (Eurasian watermilfoil) was the
most commonly observed species in Waneta Lake (25% of entire lake, 78% of littoral
zone) and Lamoka Lake (43% of entire lake, 77% of littoral zone). Eurasian
watermilfoil biomass (24.3 g DW/m2) was also significantly greater (p ≤ 0.001) in
Waneta Lake than native plant biomass. Our data suggests that Eurasian watermilfoil
is invading the native plant communities of Waneta Lake and Lamoka Lake,
thereby displacing native plants and limiting their growth to the shallow waters of
the littoral zone.
Aquatic plants are important to lake ecosystems (Madsen et al. 1996,
Wetzel 2001). These plants are essential in promoting the diversity of an
aquatic system (Carpenter and Lodge 1986). Aquatic plants in the littoral
zone may be responsible for a significant proportion of primary production
for the entire lake (Ozimek et al. 1990, Wetzel 2001); they produce
food for aquatic organisms and serve as the base of the food chain. Also,
these plants provide habitat for invertebrates, fish, and other aquatic or
semi-aquatic organisms (Cyr and Downing 1988, Madsen et al. 1996).
Littoral-zone habitats are prime areas for the spawning of most fish
species, including many species important to sport fisheries (Madsen
et al. 1996, Savino and Stein 1989). Aquatic macrophytes anchor soft
sediments, stabilize underwater slopes, remove suspended particles, and
remove nutrients from overlying waters (Barko et al. 1986, Doyle 2000,
Madsen et al. 2001). Reductions in littoral-zone species richness may
lead to decreases in fish production (Savino and Stein 1989) as well as
increased sediment resuspension, turbidity, and algal production that will
further exacerbate plant loss (Case and Madsen 2004, Doyle 2000, Madsen
et al. 1996, Wersal et al. 2006).
The introduction of non-native plants may alter the complex interactions
occurring in this habitat (Madsen 1998). Dense stands of non-native
1GeoResources Institute, Mississippi State University, Box 9652, Mississippi State,
MS 39762-9652. 2US Army Engineer Research and Development Center, Environmental
Laboratory, 3909 Halls Ferry Road, Vicksburg, MS 39180-6199. 3Ecology
and Evolutionary Biology, 150 Corson Hall, Cornell University, Ithaca, NY 14853.
*Corresponding author - firstname.lastname@example.org.
98 Northeastern Naturalist Vol. 15, No. 1
plants are often responsible for reduction in oxygen exchange, depletion
of dissolved oxygen, increases in water temperatures, and internal nutrient
loading (Madsen 1998). Myriophyllum spicatum L. (Eurasian watermilfoil)
is a non-native invasive species that, when present, has been associated with
declines in native-plant species richness and diversity (Madsen et al. 1991b).
Eurasian watermilfoil is a submersed, herbaceous, perennial aquatic plant
that typically grows in water depths of 1 to 3 m (Aiken et al. 1979). Vegetative
propagation is either by direct stem fragmentation (e.g., cutting by a
boat motor) or by autofragmentation, through the development of an abscission
layer in stem segments (Madsen et al. 1988). The production of these
stem fragments either by external forces or by autofragmentation allows for
widespread plant dispersal in littoral habitats and rapid infestation and establishment
of monotypic stands of Eurasian watermilfoil. Monotypic stands
of Eurasian watermilfoil directly reduce native-plant species richness and
diversity, and also indirectly reduce habitat complexity resulting in reduced
macroinvertebrate abundance (Keast 1984, Krull 1970), and reduction in fish
growth (Lillie and Budd 1992). Eurasian watermilfoil also poses nuisance
problems to humans in the form of increasing flood frequency and intensity,
impeding navigation, and limiting recreation opportunities (Madsen et al.
Waneta and Lamoka Lakes are used extensively for recreation and fishing.
Both water-bodies have plant communities that have become dominated
by Eurasian watermilfoil, and assistance was requested by the lake associations
on the design and implementation of measures to control this problem.
Prior to designing and implementing lake-wide management programs for
Eurasian watermilfoil suppression on these two lakes, preliminary site evaluations
were recommended to document the current distributions of Eurasian
watermilfoil and native plant species in the two lakes. For this purpose, we
performed a quantitative whole-lake study of plant communities to evaluate
plant distribution and abundance as well as to quantify the potential influences
of Eurasian watermilfoil on native-plant species richness, density, and
Field Site Description
Waneta Lake and Lamoka Lake are located in the Finger Lakes Region
of New York. Both lakes are surrounded by residential homes and support
extensive recreational activities, most notably fishing and boating. Lamoka
Lake is located in Schuyler County (42°24'59"N, 77°05'10"W). The lake
is 334.2 hectares in size with a mean depth of 5.2 m and a maximum depth
of 14.1 m. Lamoka Lake has a shallow basin, with an extensive shelf at a
depth range of 2.9 to 7.9 m. Lamoka Lake is one of the most biologically
productive lakes in central New York due to its diversity of plants and animals.
Mean dissolved oxygen in Lamoka Lake is approximately 3.5 ± 1.0
2008 J.D. Madsen, R.M. Stewart, K.D. Getsinger, R.L. Johnson, and R.M. Wersal 99
mg L-1, mean pH is 8.3 ± 0.2, mean Secchi depth is 136.0 ± 12.9 cm, and
chlorophyll-a content ranges from 24.0–57.0 μg L-1. Lamoka Lake is connected
on the north end to Waneta Lake via a 0.8-km long channel. Waneta
Lake is 329 hectares in size and is located in Schuyler and Steuben counties
(42°27'56"N, 77°06'17"W). The mean depth is 5.3 m, with a maximum depth
of 9.2 m. Mean dissolved oxygen in Waneta Lake is approximately 4.9 ± 1.1
mg L-1, mean pH is 8.0 ± 0.2, mean Secchi depth is 108.0 ± 14.6 cm, and
chlorophyll-a content ranges from 24.0–69.0 μg L-1.
To assess plant species distribution in Waneta and Lamoka Lakes,
a whole-lake point-intercept survey was conducted in August of 2000.
For each lake, a 50-m grid of sample points was developed using Map-
Info (MapInfo Corp., Troy, NY) (Figs. 1 and 2). Once on the lake, a
Figure 1. Map
of Waneta Lake
and survey sample
100 Northeastern Naturalist Vol. 15, No. 1
GeoExplorer II GPS (Trimble Corp., Santa Rosa, CA) with real time
correction was used to locate the sampling points (Madsen 1999). A total
of 303 points were visited on Waneta Lake, and 314 points were visited
on Lamoka Lake. At each point, species present in a 3-m by 3-m area
were identified and recorded. Floating plant species were identified and
recorded by visual observations. Submersed plant species were sampled
by deploying a plant rake at each point to sample species growing in
the water column (Case and Madsen 2004, Madsen 1999, Wersal et al.
2006). The plant rake was deployed from the boat to the lake bottom and
retrieved. Plants harvested by the rake were identified and recorded as
being present for that sample point. Water depth was also determined at
each sample point during the vegetation surveys. Voucher specimens of
all submersed aquatic plant species in each lake were taken and archived
at the US Army Engineer Research and Development Center, Lewisville
Aquatic Ecosystem Research Facility herbarium (Hellquist 1993). Differences
in distribution between native plant species and non-native
Figure 2. Map
of Lamoka Lake
and survey sample
2008 J.D. Madsen, R.M. Stewart, K.D. Getsinger, R.L. Johnson, and R.M. Wersal 101
plant species (mainly comprised of M. spicatum) were determined using
Statistical Analytical Software’s (SAS) McNemar’s test for dichotomous
response variables that assesses differences in the correlated proportions
within a given data set between variables that are not independent (Stokes
et al. 2000). For the purposes of this study, we used the McNemar’s test
to determine if there was a difference in the distribution of native and
non-native species by analyzing the differences in proportion of the distribution
frequencies represented by the two variables at every point. For
the purposes of these analyses, only the presence of rooted native plants
were compared to the presence of non-native species. An α = 0.05 was
used to determine statistical significance in these analyses.
Depth distribution of plants. The depth distribution of plant species
was estimated by categorizing water depth and the survey points corresponding
to those water depths into 30-cm intervals from depth 0.0 to the
maximum water depth observed during the surveys of Waneta Lake and
Lamoka Lake. Percent frequency of occurrence within a depth interval
for native species and Eurasian watermilfoil was estimated by dividing
the number of vegetated points in a given depth interval by the total
number of points in that interval. This relationship allows for a visual
representation of how plants are distributed within a lake in relation to
water depth. The depth distribution was used to estimate the littoral zone
(i.e., all survey points at or below the maximum observed depth of plant
growth was considered littoral zone) as well as the aerial coverage of native
species and Eurasian watermilfoil in Waneta Lake and Lamoka Lake.
Littoral-zone percent frequency of occurrence for native and non-native
species were estimated based on the number of points where plant species
were observed growing, in relation to the total number of points within
the littoral-zone boundary. Whole-lake species’ percent frequency of occurrence
was estimated for native and non-native species based on the
number of points where plant species were observed growing, in relation
to the total number of points sampled within a given lake.
Plant biomass collection
Aquatic plant abundance in each lake was measured in August 2000 by
harvesting plant biomass. The biomass samples were taken at 50 of the grid
points visited during the vegetation survey. The 50 biomass sample locations
were randomly selected from those points visited during the vegetation survey.
Samples were taken by a SCUBA diver using a 0.1-m2 quadrat frame
and harvesting the above-ground plant biomass of rooted plants at the sediment
surface (Madsen 1993). Samples were placed in cold storage until they
could be processed. Plant processing consisted of washing and sorting plants
by species and drying biomass at 105 °C until a constant mass was achieved.
Plant samples were then weighed to assess biomass. A one-way ANOVA
was used to analyze differences in biomass within each lake; a pairwise
102 Northeastern Naturalist Vol. 15, No. 1
comparison between plant species’ means within each lake was conducted
using a Bonferoni post hoc analysis. A linear regression analysis was used to
determine if a relationship exists between biomass of Eurasian watermilfoil
and biomass of native plant species. Analyses were conducted using Statistics
8.0 software, with an α = 0.05 threshold for statistical significance for
In Lamoka Lake, we observed a total of 20 plant species, with 16
being submersed species, 3 floating species, and 1 emergent species
Table 1. Aquatic plant species observed in Waneta Lake and Lamoka Lake during August 2000.
* denotes non-native species.
Lamoka Lake Waneta Lake
Entire Littoral Entire Littoral
lake zone lake zone
Scientific name Common name freq (%) freq (%) freq (%) freq (%)
Ceratophyllum demersum L. Coontail 108 (36.0) 108 (64.0) 42 (13.0) 42 (41.0)
Chara sp. Chara 2 (0.6) 2 (1.0) 4 (1.0) 4 (4.0)
Elodea canadensis Canadian elodea 89 (29.0) 89 (53.0) 17 (5.0) 17 (17.0)
Lemna trisulca L. Duckweed 3 (0.9) 3 (2.0) 0 0
Myriophyllum spicatum L.* Eurasian watermilfoil 130 (43.0) 130 (77.0) 80 (25.0) 80 (78.0)
Najas flexilis Willd. Bushy naiad 4 (1.0) 4 (2.0) 9 (3.0) 9 (9.0)
N. guadalupensis Spreng. Southern naiad 41 (14.0) 41 (24.0) 29 (9.0) 29 (28.0)
Nymphaea odorata Ait. White water lily 40 (13.0) 40 (24.0) 4 (1.0) 4 (4.0)
Nuphar advena Yellow pond lily 24 (8.0) 24 (14.0) 2 (0.6) 2 (2.0)
Potamogeton amplifolius Large-leaved 13 (4.0) 13 (8.0) 4 (1.0) 4 (4.0)
Potamogeton crispus L.* Curlyleaf pondweed 1 (0.3) 1 (0.6) 0 0
P. diversifolius Raf. Narrow pondweed 0 0 1 (0.3) 1 (1.0)
P. praelongus Wulf. Whitestem pondweed 8 (3.0) 8 (5.0) 2 (0.6) 2 (2.0)
P. pusillus L. Baby pondweed 0 0 2 (0.6) 2 (2.0)
P. robinsii Oakes Robbins’ pondweed 36 (12.0) 36 (21.0) 8 (3.0) 8 (8.0)
P. zosteriformis Fern. Flatstem pondweed 18 (6.0) 18 (11.0) 2 (0.6) 2 (2.0)
Ranunculus sp. Water buttercup 4 (1.0) 4 (2.0) 0 0
Typha angustifolia L. Narrowleaf cattail 3 (1.0) 3 (2.0) 0 0
Ultricularia vulgaris L. Common bladderwort 16 (5.0) 16 (9.0) 0 0
Vallisneria americana Wild celery 27 (9.0) 27 (16.0) 12 (4.0) 12 (12.0)
Zanichellia palustris L. Horned pondweed 2 (0.6) 2 (1.0) 0 0
Zosterella dubia (Jacq.) Water stargrass 33 (11.0) 33 (20.0) 2 (0.6) 2 (2.0)
Total species occurrence (mean ± 1 SE per point) 1.9 ± 0.1 3.6 ± 0.2 0.7 ± 0.1 2.2 ± 0.2
Native plant occurrence (mean ± 1 SE per point) 1.6 ± 0.1 2.8 ± 0.2 0.4 ± 0.1 1.4 ± 0.2
Non-native plant occurrence (mean ± 1 SE per point) 0.4 ± 0.0 0.8 ± 0.0 0.3 ± 0.1 0.8 ± 0.1
Depth (m) (mean ± 1 SE per point) 5.2 ± 0.1 1.6 ± 0.0 5.3 ± 0.2 1.8 ± 0.1
Total number of sites 302 169 316 102
2008 J.D. Madsen, R.M. Stewart, K.D. Getsinger, R.L. Johnson, and R.M. Wersal 103
(Table 1). Of these, Eurasian watermilfoil and Potamogeton crispus L.
(curlyleaf pondweed) were the only two non-native species. Dominant
species in the lake by frequency of occurrence were Eurasian watermilfoil
(77% of the littoral zone), Ceratophyllum demersum (coontail, 64%),
and Elodea canadensis (Canadian elodea, 53%). Comparing all vegetated
sites, the distribution of native plants versus Eurasian watermilfoil was
not statistically different (χ2 = 2.66, d.f. = 1.0, p = 0.102). Littoral-zone
plant diversity was relatively high with 3.56 species per point, 0.78 nonnative
species per point, and 2.79 native species per point. Plants were
present in 96% of the littoral zone samples, with native plants occurring
at 84% of the points in the littoral zone. Plants were widely distributed
in Lamoka Lake, particularly in the southern arm. Eurasian watermilfoil
(43% frequency of occurrence) was the most widely distributed species
and was observed growing along most shorelines. Coontail (36%) was the
dominant native plant species followed by Canadian elodea (27%).
Waneta Lake had a total of 16 plant species, with 14 submersed species,
and 2 floating species. Of these, only one non-native species, Eurasian watermilfoil,
was observed (Table 1). The dominant species observed in the lake
was Eurasian watermilfoil (78% of samples in the littoral zone), followed
by coontail (41%). Comparing all vegetated sites, the distribution of native
plants versus Eurasian watermilfoil was statistically different (χ2 = 6.736, d.f.
= 1.0, p = 0.013). Littoral-zone plant diversity in Waneta Lake was somewhat
lower than in Lamoka, with 2.16 plant species per point in the littoral zone.
Similar to Lamoka, 0.78 non-native species per point was observed. Native
species richness was 1.37 species per point. Waneta Lake plant cover was
89% in the littoral zone, with 63% of points in the littoral zone having native
plants. Whole-lake plant distribution was sparse in Waneta Lake; plants
were most common in the southern portions of the lake. Plant distribution was
very sparse along the eastern shore. This pattern was consistently observed
for coontail (13%), Eurasian watermilfoil (25%), and Najas guadalupensis
Spreng. (southern naiad, 9%). Canadian elodea (5%) was found predominantly
in the shallow southern end of the lake, while Vallisneria americana Michx.
(water celery, 4%) was scattered along all shores.
Depth distribution of plants. The actual observed maximum depth of
plant colonization for Lamoka Lake was less than 3.6 m, indicating that
plants occupied approximately 55% of the lake bottom. From lakeshore
to 2.0 m depth, almost 100% of the points were vegetated (Fig. 3). In Waneta
Lake, plants were observed to a maximum depth of 3.4 m, with plants
observed in 100% of the sample points at depths less than or up to 2.1 m,
more than 80% from 2.1 m to 3.0 m, and 40% of the sites 3.3 m to 3.4
m (Fig. 3). No plants were found at the three sites in the 3.1-m to 3.3-m
depth interval. A maximum depth of plant colonization out to 3.4 m indicates
that about 34% of the lake area is littoral zone, with 89% of this zone
being vegetated. Eurasian watermilfoil was observed at 70% of sample
104 Northeastern Naturalist Vol. 15, No. 1
points out to a depth of 3.0 m, with frequency of occurrence dropping to
40% at a depth of 3.4 meters.
Plant abundance by biomass
Plant biomass in Lamoka Lake was different among species (F = 2.8,
d.f. = 350.0, p = 0.009). Lamoka Lake was dominated by Canadian elodea
(50.8 g DW m-2), followed by coontail (24.3 g DW m-2) and Eurasian
watermilfoil (21.9 g DW m-2) (Fig. 4). Plant biomass in Waneta Lake
was also different among species (F = 7.74, d.f. = 249.0, p ≤ 0.001).
Waneta Lake was dominated by Eurasian watermilfoil (24.3 g DW m-2).
Total macrophyte biomass in Waneta Lake was 47.3 g DW m-2. There
was no relationship between biomass of native plants (total macrophyte
biomass minus Eurasian watermilfoil biomass) and biomass of Eurasian
Figure 3. Depth
of the percent
f r e q u e n c y
of occurrence of
aquatic plants in
data for nativeplant
grey bars represent
2008 J.D. Madsen, R.M. Stewart, K.D. Getsinger, R.L. Johnson, and R.M. Wersal 105
watermilfoil for Lamoka Lake (F = 2.62, d.f. = 49, p = 0.112) or Waneta
Lake (F = 0.31, d.f. = 49, p = 0.582) (Fig. 5).
Eurasian watermilfoil was the dominant species in Waneta Lake and was
co-dominant in Lamoka Lake as determined by frequency of occurrence and
biomass samples. Overall plant species richness was much lower in Waneta
Lake than in Lamoka Lake, a result of the increased presence of Eurasian
watermilfoil in Waneta Lake. Eurasian watermilfoil was able to colonize and
spread in deep-water habitats where it was observed growing in water depths
to 3.4 m. In Lamoka Lake, native plants were dominant to a water depth of
2.0 m, with suppression of these native plants in deeper waters where they
competed with Eurasian watermilfoil. Native species in Waneta Lake were
observed in depths out to 1.2 m, and were also suppressed in deeper areas.
Eurasian watermilfoil was commonly observed in deep-water habitat and
appeared to replace native plants in depths of 1.5 m to 3.6 m, indicating that
Eurasian watermilfoil was limiting native plant growth in deeper water. The
absence of native plants in deep-water habitat accounts for the difference
Figure 4. Plant biomass of the most abundant species in Lamoka Lake and Waneta
Lake during the time of biomass harvest, August 2000. Abbreviations: CD = coontail,
EC = elodea, HD = water stargrass, MS = Eurasian watermilfoil, NF = bushy naiad,
NA = yellow pondlily, VA = wild celery, and TOT = total macrophyte. Analyses were
conducted for each lake and not between lakes; different lower-case letters above
bars refer to differences (± 1 SE) in plant biomass within Lamoka Lake, and different
capital letters above bars refer to differences (± 1 SE) in plant biomass within
106 Northeastern Naturalist Vol. 15, No. 1
in distribution between Eurasian watermilfoil and native plants in Waneta
Lake. The difference in distribution between native plant species and Eurasian
watermilfoil in Waneta Lake suggests that Eurasian watermilfoil has
already displaced native vegetation throughout the majority of the lake. In
Lamoka Lake however, no differences in distribution were detected between
native plant species and Eurasian watermilfoil, indicating that Eurasian
watermilfoil is able to invade and inhabit the same locations as native plant
species. One can speculate that Eurasian watermilfoil may have invaded
Lamoka Lake after Waneta Lake and has not been present long enough to
displace the native species.
The suppression and displacement of native plants by Eurasian watermilfoil
has been observed in other New York lakes (Madsen et al. 1991a, b). Over
a three-year period (1987–1989) in Lake George, NY, Eurasian watermilfoil
Figure 5. Comparison
of Eurasian watermilfoil
biomass and native-
in Lamoka Lake
and Waneta Lake
during the time of
2008 J.D. Madsen, R.M. Stewart, K.D. Getsinger, R.L. Johnson, and R.M. Wersal 107
spread from 30% coverage to over 95% coverage at a monitoring site (Madsen
et al. 1991b). At this same location, it was empirically shown that the
native-plant density was significantly reduced from 5.5 species per quadrat
to 2 species (Madsen et al. 1991b). Native plant species occurrence per point
for Waneta Lake and Lamoka Lake was 0.4 and 1.6 respectively in the presence
of Eurasian watermilfoil, values much lower than in other studies.
The coverage of Eurasian watermilfoil in Waneta Lake and Lamoka Lake
was approximately 80% in the littoral zone. Madsen et al. (1991b) stated
that Eurasian watermilfoil coverage greater than 50% is considered a dense
bed. However, the overwhelming presence of Eurasian watermilfoil and its
subsequent biomass in the two lakes was not related to native-plant biomass,
though both lakes do show a general negative relationship between Eurasian
watermilfoil and native-plant biomass. It has been implied that there is an
inverse relationship between the abundance (biomass) of Eurasian watermilfoil
and native plant species (Madsen 1998) where lakes with more than 50%
Eurasian watermilfoil were found to have less than 60% native-plant cover
(Madsen 1998). The relationship has been quantitatively documented by
Madsen et al. (1991b) and reported as occurring in other systems (Aiken et
al. 1979, Grace and Wetzel 1978, Madsen 1998, Smith and Barko 1990). The
small number of samples used to compare Eurasian watermilfoil biomass to
native-plant biomass in this study limited the power of the analysis; had we
collected more samples for this portion of the study, a negative relationship
would have likely been found.
Although Eurasian watermilfoil was dominant in both lakes, there was
still a diverse native-plant community in each lake. Waneta Lake had 16
species of aquatic plants present; of these, only one was non-native (Eurasian
watermilfoil), and 13 of these species were native submersed plants
that directly competed with Eurasian watermilfoil. Similarly, Lamoka Lake
had 20 species of aquatic plants present, of these only two were non-native—
Eurasian watermilfoil and curlyleaf pondweed—and 14 were native
submersed plants. Coontail, Canadian elodea, and southern naiad were the
dominant submersed native plants in both Waneta Lake and Lamoka Lake.
Potamogeton spp. (native pondweeds) appear to be the most vulnerable to
invasion by Eurasian watermilfoil due their low presence compared to other
species of submersed plants during the survey. The pondweeds may have
been better adapted to grow in low light environments (deep-water habitats),
which were the first areas to be invaded by the more aggressive Eurasian
watermilfoil. Spence and Chrystal (1970a, b) demonstrated a greater photosynthetic
capacity in deep water of some pondweeds, and suggested that
shade tolerance was directly linked to the natural depth distribution of these
species. Madsen and Adams (1989) found that Stuckenia pectinata Börner
(sago pondweed) has photosynthetic characteristics that allow it to grow in
a broad range of environments. However, the pondweeds are primarily early
season dominants, and have a lower temperature optimum than species such
108 Northeastern Naturalist Vol. 15, No. 1
as coontail and Eurasian watermilfoil (Madsen and Adams 1989), which
would put the pondweeds at a disadvantage when competing with Eurasian
watermilfoil for light later in the growing season (the time of this survey).
Also, under low-light conditions, Eurasian watermilfoil reallocates its biomass
to the formation of a canopy which further reduces light availability to
native plants, resulting in reductions in native plant distribution and biomass
(Barko and Smart 1981, Madsen et al. 1991a).
The presence of Eurasian watermilfoil in Waneta Lake and Lamoka
Lake has caused a shift in the distribution of native plant species. The
growth of these native plants has been limited to shallow depths within
the littoral zone while Eurasian watermilfoil dominates the deeper
water. The dominance of Eurasian watermilfoil should be of concern
as its aggressive growth and ability to survive under adverse environmental
conditions could allow it to overtake the remaining native plant
communities in Waneta Lake and Lamoka Lake. Adequate diversity
and representation of native plant species already occur in Waneta and
Lamoka Lakes to revegetate or replace Eurasian watermilfoil after the
implementation of management techniques. Furthermore, light transparency
in both lakes is low, and additional efforts should be made to reduce
algal and particulate turbidity in these lakes. The reductions in turbidity
will further assist in native-plant restoration and establishment and reduce
the competitive advantage by Eurasian watermilfoil.
We thank Adam Way for assistance in field sampling and survey preparation.
We thank Lloyd Wetherbee from the Schuyler County Soil and Water Conservation
District, NY for supplying water-quality data. We thank Louis Wasson for
assistance with GIS software. This research was supported by the Aquatic Ecosystem
Restoration Foundation and was approved for publication as Journal Article
No. J-11026 of the Mississippi Agricultural and Forestry Experiment Station,
Mississippi State University. Permission was granted by the Chief of Engineers to
publish this information
Aiken, S.G., P.R. Newroth, and I. Wile. 1979. The biology of Canadian weeds. 34.
Myriophyllum spicatum L. Canadian Journal of Plant Science 59:201–215.
Barko, J.W., and R.M. Smart. 1981. Comparative influences of light and temperature
on the growth and metabolism of selected submersed freshwater macrophytes.
Ecological Monographs 51:219–235.
Barko, J.W., M.S. Adams, and N.S. Clesceri. 1986. Environmental factors and their
consideration in the management of submersed aquatic vegetation: A review.
Journal of Aquatic Plant Management 24:1–10.
Carpenter, S.R., and D.M. Lodge. 1986. Effects of submersed macrophytes on ecosystem
processes. Aquatic Botany 26:341–370.
2008 J.D. Madsen, R.M. Stewart, K.D. Getsinger, R.L. Johnson, and R.M. Wersal 109
Case, M.L., and J.D. Madsen. 2004. Factors limiting the growth of Stuckenia pectinata
(sago pondweed) in Heron Lake, Minnesota. Journal of Freshwater Ecology
Cyr, H., and J.A. Downing. 1988. Empirical relationships of phytomacrofaunal abundance
to plant biomass and macrophyte bed characteristics. Canadian Journal of
Fisheries and Aquatic Science 45:976–984.
Doyle, R.D. 2000. Effects of sediment resuspension and deposition on plant growth
and reproduction. US Army Corps of Engineers, Rock Island, IL, USA. Environmental
Report 28. 64 pp.
Grace, J.B., and R.G. Wetzel. 1978. The production biology of Eurasian watermilfoil
(Myriophyllum spicatum L.): A review. Journal of Aquatic Plant Management 16:
Hellquist, C.B. 1993. Taxonomic considerations in aquatic vegetation assessments.
Lake and Reservoir Management 7:175–183.
Keast, A. 1984. The introduced macrophyte, Myriophyllum spicatum, as a habitat for
fish and their invertebrate prey. Canadian Journal of Zoology 62:1289–1303.
Krull, J.N. 1970. Aquatic plant-invertebrate associations and waterfowl. Journal of
Wildlife Management 34:707–718.
Lillie, R.A., and J. Budd. 1992. Habitat architecture of Myriophyllum spicatum as
an index to habitat quality for fish and macroinvertebrates. Journal of Freshwater
Madsen, J.D. 1993. Biomass techniques for monitoring and assessing control of
aquatic vegetation. Lake and Reservoir Management 7:141–154.
Madsen, J.D. 1998. Predicting invasion success of Eurasian watermilfoil. Journal of
Aquatic Plant Management 36:28–32.
Madsen, J.D. 1999. Point- and line-intercept methods for aquatic plant management.
APCRP Technical Notes Collection (TN APCRP-M1-02), US Army Engineer Research
and Development Center, Vicksburg, MS. 16 pp.
Madsen, J.D., and M.S. Adams. 1989. The light and temperature dependence of
photosynthesis and respiration in Potamogeton pectinatus L. Aquatic Botany 36:
Madsen, J.D., L.W. Eichler, and C.W. Boylen. 1988. Vegetative spread of Eurasian
watermilfoil in Lake George, New York. Journal of Aquatic Plant Management
Madsen, J.D., C.F. Hartleb and C.W. Boylen. 1991a. Photosynthetic characteristics
of Myriophyllum spicatum and six submersed macrophyte species native to Lake
George, New York. Freshwater Biology 26:233–240.
Madsen, J.D., J.W. Sutherland, J.A. Bloomfield, L.W. Eichler, and C.W. Boylen.
1991b. The decline of native vegetation under dense Eurasian watermilfoil canopies.
Journal of Aquatic Plant Management 29:94–99.
Madsen, J.D., J.A. Bloomfield, J.W. Sutherland, L.W. Eichler, and C.W. Boylen.
1996. The aquatic macrophyte community of Onondaga Lake: Field survey and
plant-growth bioassays of lake sediments. Lake and Reservoir Management 12:
Madsen, J.D., P.A. Chambers, W.F. James, E.W. Koch, and D.F. Westlake. 2001. The
interactions between water movement, sediment dynamics, and submersed macrophytes.
110 Northeastern Naturalist Vol. 15, No. 1
Ozimek, T., R.D. Gulati, and E. van Donk. 1990. Can macrophytes be useful in biomanipulation
of lakes? The Lake Zwemlust example. Hydrobiologia 200/201:
Savino, J.F., and R.A. Stein. 1989. Behavior of fish predators and their prey: Habitat
choice between open water and dense vegetation. Environmental Biology of
Smith, C.S., and J.W. Barko. 1990. Ecology of Eurasian watermilfoil. Journal of
Aquatic Plant Management 28:55–64.
Spence, D.H.N., and J. Chrystal. 1970a. Photosynthesis and zonation of freshwater
macrophytes. I. Depth distribution and shade tolerance. The New Phytologist 69:
Spence, D.H.N., and J. Chrystal. 1970b. Photosynthesis and zonation of freshwater
macrophytes. II. Adaptability of species of deep and shallow water. The New
Stokes, M.E., C.S. Davis, and G.G. Koch. 2000. Categorical Data Analysis Using the
SAS® System. John Wiley and Sons, Chicago, IL. 648 pp.
Wersal, R.M., J.D. Madsen, B.R. McMillan, and P.D. Gerard. 2006. Environmental
factors affecting biomass and distribution of Stuckenia pectinata in the Heron
Lake System, Minnesota, USA. Wetlands 26:313–321.
Wetzel, R.G. 2001. Limnology: Lake and River Ecosystems, Third Edition. Academic
Press, San Diego, CA. 1006 pp.