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Minnesota Department of Natural Resources

Special Publication 160, 2004

THE ROLE OF SUBMERSED AQUATIC VEGETATION
AS HABITAT FOR FISH IN MINNESOTA LAKES,
INCLUDING THE IMPLICATIONS OF NON-NATIVE PLANT INVASIONS
AND THEIR MANAGEMENT

Ray D. Valley, Timothy K. Cross, and Paul Radomski.

Minnesota Department of Natural Resources
Division of Fish and Wildlife
500 Lafayette Road
St. Paul, MN 55155-4020

Executive Summary

This review updates the Division of Fish and Wildlife’s understanding of the role of
submersed aquatic vegetation (SAV) in providing fish habitat in Minnesota lakes. Below, are
several generalizations and recommended approaches for aquatic plant management.

1. Many fish, such as sunfish, largemouth bass, northern pike, and muskellunge, de-
pend on SAV for food and shelter. Nongame fish such as darters, minnows, and
killifishes depend primarily on nearshore emergent and submersed vegetation.
2. The presence of SAV tends to promote higher water clarity.
3. Black bullhead and common carp often dominate turbid lakes with little to no SAV.
Carp are an invasive non-native species that contributes to the loss of native SAV by
dislodging rooted plants and resuspending sediments.
4. Generally, conditions for game fish deteriorate when the percentage of a basin that is
covered with SAV falls below 10% or exceeds 60%. This range does not consider
basin morphometry (i.e., shallow versus deep) which ultimately controls how much
vegetation naturally grows within a lake.
5. Studies show native plants provide higher quality habitat for desirable fish than in-
vasive non-native plants such as curly-leaf pondweed or Eurasian watermilfoil.
However, these non-native plants provide better habitat than little or no SAV.
6. Minnesota lakes infested with curly-leaf pondweed or Eurasian watermilfoil have
not seen large declines in game fish populations.
7. Lake productivity and initial plant conditions appear to greatly affect selective
whole-lake herbicide’s (such as fluridone) effect on fish habitat. Whole-lake studies
in infested, moderately-productive (mesotrophic) Michigan lakes with abundant na-
tive plants, showed neutral to positive effects of fluridone on fish habitat.
13. Fluridone applications in infested productive (eutrophic) Minnesota lakes with low
cover of native SAV can have dramatic negative effects on SAV habitats, water clar-
ity, and fish communities.

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14. Aquatic plant management policies should reflect a precautionary approach where it
is understood that any alteration to SAV will invariably have some effect on a lake’s
fish community. Therefore, policies should ostensibly be conservative with the in-
tent to minimize habitat degradation.
15. Limiting the cumulative amount of SAV removal may be the most prudent approach
towards precautionary management. However, thresholds should be dependent on
lake type. The current 15% rule (maximum treatment area within the 15 foot depth
zone) for chemicals and 50% rule for mechanical harvesting may be reasonable for
some lakes (e.g., small eutrophic lakes); stricter thresholds may be needed for others
(e.g., soft water lakes, large or deep lakes).
16. Overall, whole-lake aquatic plant treatment is risky. Significant biological risks as-
sociated with large-scale manipulations include excessive removal of fish habitat
and thus decline of fish populations, loss of sensitive plant species, declines in water
clarity and potential long-term cumulative effects of multiple treatments, since
eradication of non-native plant species is highly unlikely.
17. Vegetated, nearshore habitat is critical for fish recruitment. Any removal should be
viewed as habitat loss, and efforts should be made to minimize this loss. It follows
that 100 feet of removal is worse than 50 feet of removal even if the removal is of a
non-native species.
18. Mechanical harvesting may be the best alternative for managing nuisance surface
growth of vegetation. Although this requires perpetual maintenance, harvested boat
lanes through surface-growing vegetation represents a positive benefit for recrea-
tional access and fish habitat (harvested strips of SAV increases edge and may
benefit game species).

Introduction Tonn 1990; Schupp 1992). In Minnesota, we
see a great diversity of lake types, each dis-
The role of submersed aquatic vegeta- playing its own physical and biological
tion (SAV) in structuring lake ecosystems, and ‘signature’ (Figure 1, Table 1). Environ-
in particular fish communities, has been the mental filters, hierarchical in space and time
source of much research during the last 25 (i.e, continental → regional or watershed →
years. Several published papers comprehen- lake-type → within-lake), determine these sig-
sively review these roles and demonstrate that natures and constrain fish communities within
the relationship between SAV and fish is a defined range (Tonn 1990). Herein, we fo-
highly complex and variable (Dibble et al. cus primarily on the lake type and within lake
1996; Weaver et al. 1996; Diehl and Kornijów effects of SAV on fish populations. However,
1998; Petr 2000). Nevertheless, several gen- the following generalizations should be inter-
eralizations have emerged from this collective preted as nested within the context of these
body of research. They are discussed here in larger filters operating on any particular wa-
the context of Minnesota lakes. terbody of interest.
The relationship of SAV and fish var- Aquatic plants serve many ecosystem
ies among lake types, thus we must first functions including primary production, stabi-
establish a framework on which these gener- lizing sediments, maintaining water clarity,
alizations may be based. The quality of lakes and providing habitat for zooplankton, macro-
as habitat for fish, depend on their glacial his- invertebrates, and numerous fish species
tory, climate, water chemistry, productivity, (Carpenter and Lodge 1986; Dibble et al.
and morphometry (Tonn and Magnuson 1982; 1996; Diehl and Kornijów 1998). Many
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L a k e S ize ( A c r es ) < 500 > 500 > 80 % < 80 % L a k e D e pt h > 80 % < 80 % ( % < 1 5 f t) < 100 > 100 < 100 > 100 < 100 > 100 < 100 > 100 T o t a l A lka lin it y ( m g /L C aC O 3) L a k e Ty p e 1 L a k e Ty p e 3 L a k e Ty p e 5 L a k e Ty p e 7 L a k e Ty p e 2 L a k e Ty p e 4 L a k e Ty p e 6 L a k e Ty p e 8 Figure 1. Separa- tion of large and small lakes is arbitrary and relative to the size range of lakes in Minnesota. Physical classification of Minnesota Lakes using a reduction or ‘lumping’ of Schupp’s (1992) lake classification groupings. 3 .

e Fish species richness in this lake class depends on the productivity and the SAV richness in the lake. deep. shallow. soft 685 N. 408 Central – SW high to very high Low Low to High √ common Low hard water 3 Small. deep. Characteristics of the eight distinct types of Minnesota Lakes identified in Figure 1. d Alternate stable states refers to the dynamic characteristic of shallow lakes whereby abundant vegetation and relatively clear water occupy one state and turbid water with no submersed vegetation occupies another mutually-exclusive state. Relative Plant Spe.Table 1. deep. High Low Low to High √ common Low hard water Central 7 Large. hard 1. hard 340 N. b Relative plant species richness was determined by DNR vegetation survey species lists. and algal concentra- tion (Chlorophyll a). shallow. shallow. soft 102 NE Low to moderate No Data Low rare moderate water 8 Large. Central – Moderate to high High Low rare Low water NE 2 Small. shallow. Central Moderate to high High High More than rare. phosphorus concentration (total phosphorus). c Statistical analyses (cluster analysis) to determine relative abundance of aquatic vegetation was performed on recent vegetation survey data from 1. soft 34 NE High High Low rare Low water 6 Large. Radomski.173 Central Moderate to high Moderate Low to High √ common Low to Highe water 5 Large. highly productive lakes with species poor SAV communities have low fish species richness. 190 Central – S. 4 . unpublished data). soft 991 N. Central – Moderate High Low rare High water NE 4 Small.410 lakes collected by Minnesota DNR Fisheries field personnel (P. deep. Alternate Number cies Relative SAV Stable Non-native Plant Fish Species Lake Description of Lakes Location Productivitya Richnessb Abundancec Statesd Infestationsc Richness Type 1 Small. less high water than common a Productivity is based on Trophic State Index (Carlson 1977) and is a combination of water clarity (secchi).

Many fish species depend on Crowder and Cooper (1979) developed a SAV for nest building and spawning.e. dance.. particularly for large- in open pockets to ambush fish prey (Killgore mouth bass and bluegill L. Wrenn et al. crappie vegetation covers less than 10% of a water- Pomoxis spp. ceeds 40-60% of the entire waterbody (i. 5 . Other species such as smallmouth bass Mi. ceina 1996. body (e. several experimental studies macroinvertebrates living on SAV. At low plant abundances. should nursery habitat for young age classes of fish be highest at intermediate levels of plant abun- and small nongame fish (Poe et al. and reducing op- found in SAV habitats in the Upper Missis. are adapted to open dependent species are scarce and production water environments. Carp directly vival. Studies consistently the sediment and nutrients in the water col- demonstrate that fish abundance is greater in umn. Valley and Bremigan Esox maquinongy. Miranda and Vegetated Minnesota lakes support a Pugh 1997). sup- cropterus dolomieu. Lammens 1999. vegetation attracts an abundance of inverte. they increase their vegetated habitats than in unvegetated habitats competitive advantage over most game fish (Dibble et al. and SAV is of less im. when flavescens. result (Colle and Shireman 1980. (Breukelaar et al. In general.S. et al. yellow bullhead Ameiurus na. turbid reservoirs).. white bass Morone chrysops. portunities for reestablishment by increasing sippi River Basin. merous minnow and shiner species (Family: shallow vegetated lakes or bays). Ma- 1996..g. Bettoli et al. rates or poor growth by predators commonly talis. macrochirus. Submersed kos et al. Cross and McInerny 2001). 1992. (1984). of largemouth bass and bluegill is low portance for their populations (Dibble et al. fisheries game fish and nongame fish species (Keast managers have sought to identify the level of 1984). Less work because they are tolerant of the low oxygen has been done in north temperate lakes. Maceina 1996. and 1992. at least in part. 1996. In fact. have documented a reduction in predator feed- In Minnesota lakes.. 1996. Trebiz et al. This suggests that trast. the probability of finding a quality largemouth native carp Cyprinus carpio are often associ. bass or bluegill population is highest between ated with turbid lakes with little to no SAV 10% and 60% total cover of SAV. This attracts larger. refuge and foraging. low feeding Cyprinidae). In con. In doing so. yellow perch Perca ported this theory. and thus growth and production. 1996. ming barriers created by dense vegetation Aquatic vegetation is consumed by some ani. Cross and McInerny 2001. Wiley et al. Dibble et al.species of fish depend on SAV for their sur. SAV for spawning. When vegetation coverage ex- diversity of nongame species including nu. 1982. some darter species Etheostoma spp. vegetation- and catfish Ictalurus spp. predatory game aquatic plant abundance that supports high fish that patrol the edge of plant beds or wait game fish production. SAV is important ing rates as vegetation becomes more dense to game fish such as sunfish Lepomis spp. Wrenn et al. 1986. visual and swim- species including ducks and wading birds. Indeed. Nearshore scarce because of a lack of predator refuges. prey are Bryan and Scarnecchia 1992). reduce the ability of predators to capture fish mals while other animals consume the prey. Near. Anderson 1984. 1984. Given the importance of SAV for fish brates that provides prey for many juvenile populations and productive fisheries. banded killifish Fundulus diaphanus. vegetation also is critical for many wildlife At high plant abundances. lakes and reservoirs have. (Durocher et al. and muskellunge 1987. 1989). walleye Sander vitreus. brook silverside Labidesthes sicculus. 2003). (1996) noted that affect the abundance of SAV by dislodging 112 fish species representing 19 families were rooted plants while feeding. never- environments typically associated with these theless. Miranda and Pugh 1997). lakes (Drake and Pereira 2002). Gotceitas and Colgan northern pike Esox lucius. Numerous studies from southern U. model that suggested that predator feeding shore vegetation is particularly important rate. black bullhead Ameirus melas and non. Savino and Stein largemouth bass Micropterus salmoides. (Crowder and Cooper 1982. These species depend on 2002a). 1994. Bettoli et al. Diehl 1988. Par- Pratt and Smokorowski 2003).

for a variety of fish species. vegetation cover in shallow lakes im. Hinch 1991. We use the term ‘heterogene- structuring management decisions (Hoyer and ity’ to reflect what is commonly referred to as Canfield 2001). It is widely recognized that holding all • Lake size. heterogeneity will be highest in of SAV. 1988). Hoyer and Canfield form (arrangement of stems and leaves) of 1996. and high substrate diversity (i. abundance Chara spp. water chemistry. variable studies to our knowledge show reducing slopes. high water clarity with moderate nutrient lev- proves fish growth. and productivity influence 1996. individual plant species and the spatial distri- Vestergaard and Sand-Jensen 2000) bution and species composition of plant beds • Aquatic plant architecture and patchiness creates a patchy littoral landscape. shape. and nested within. 1996. ‘Heterogeneity’ simply refers to a overlooked when relating coarse estimates of collection of diverse micro-habitats suitable SAV cover to fish. patchiness is positively related to fish diversity For example. sand. patches whole-lake cover of SAV will never be of clay. unpublished data) to. In deep lakes. question the appropriate. • Lake depth controls the maximum cover Ultimately. plant species assemblages (Figure 2. Weaver et al. Lodge et al. substrate composition. Dibble et al.’ ‘Quality’ is subjective and difficult characteristics and features of lakes that are to measure. and numerous points. Vestergaard and Sand-Jensen 2000). Downing and Plante 1993. Naturally. such as spawning substrates (Annet et al. Diversity in the architectural growth and McIvor 1994. Weaver et demonstrate that a diversity of plant types and al. the above habitat factors constant. depth. Schneider (2000). shape. No published complex basin morphometry (i. Duarte Weaver et al. Cheruvelil and N. Studies (Chick and McIvor 1994. and current studies such as Eurasian watermilfoil Myriophyl- in Michigan (K. 1997. thereby providing habitat for a the effect of a range of SAV cover solely diversity of fish species. quality fish trophic status.. affecting the total habitat heterogeneity of a ness of using a defined percentage in particular lake. other physical forces that taken together. abundance. 1997. These condi- excessive (Hoyer and Canfield 2001). (Weaver et al. and gravel).. gravel substrates). 1997. within the littoral zone. lum spicatum.(1997). Weaver et al. and islands). 1996. Below. Michigan State University. Factors such as • Availability of other critical microhabitats lake size.g. tions between littoral and pelagic habitats gether suggest a preferred range of 18% – (Diehl and Kornijów 1998. 40% total cover of SAV. as well as control the greater than the scale of analysis (Tonn composition. Chick munities.e. 1996. 1996. the distribution of non-macrophyte habitats • Ecological constraints operating at scales (e. Weaver et al. substrates (Downing et al. muck. tions provide habitats for a diversity of aquatic Most published studies have not explored plant species. lakes with > 60% large lakes with convoluted shorelines and cover are shallow lakes. percent els. 100% cover of a low. water chemistry. 1996). we list important ‘quality. Valley and Bremigan 2002a). Pursuit of this elusive ‘optimal’ per- cent coverage of SAV for littoral fish species Habitats created by plants are affected has revealed numerous complexities of lakes by. provides vastly different habi- tat than 100% cover of a canopy species 6 . habitat is associated with diverse plant com- 1990. Nichols 1992. Miranda and Dibble and Kalff 1990. Pratt and growing macrophyte species such as chara Smokorowski 2003). Nate. Department of • Composition of the food web and interac- Fisheries and Wildlife. 1996.e.. and distribution of 1990. 2002). depth. Maceina 1996.

Lake Productivity Oligotrophic Mesotrophic Eutrophic Hyper- eutrophic Lake Size Small Large Basin Morphology and Shoreline Shallow Round Convoluted Bowl depth and shoreline Substrate Homogeneous Heterogeneous Figure 2. and muskellunge respond to foraging abilities (Valley and Bremigan patchiness at larger scales. King 1993. shallow lakes (lake types 1. and 6) deserve special attention. thereby providing a feedback to habitat heterogeneity. Therefore. larger. Essing. In our consideration of the diversity of ton and Kitchell 1999. shallow lakes inherently have low (1 x 1 x 1 m volume of water close to shore) is habitat heterogeneity. Pratt and Smokorowski Minnesota lakes. plant composition at small scales suggests. 2. Di- spawning. Multiple age-classes and species of mosaic becomes increasingly important to fish depend on a diversity of habitats for their their success (Cross and McInerny 2001). 1996. 1996. and a lake’s patch 2002a). (Killgore et al. fish communities are depauperate 7 . Conceptual heterogeneity “Equalizer” with gray bars demonstrating levels of highest habitat heterogeneity. Environmental factors perhaps more important than the spatial ar. Weaver et al. Werner et al. north- Pratt and Smokorowski 2003) and predator ern pike. Annet heterogeneity at multiple spatial scales. for small vegetation dwell. and refuge needs (Chick verse native plant communities produce and McIvor 1994. 1997). more mobile In general. Chick and McIvor 1994. 1977. In contrast. Patchiness is highly scale-dependent SHALLOW LAKES – UNIQUE IN THEIR and is perceived differently by species of fish STRUCTURE AND FUNCTION depending on their size and home range (Kotliar and Weins 1990. species such as adult largemouth bass. vegetation. 1989. these characteristics generally lead to diverse macro- phyte assemblages. Collectively. 2003). Figure 2 ing species. foraging. such as oxygen depletion in winter further rangement and size of large offshore beds of contributes to poor habitat conditions for fish. 5. et al.

biomanipulation. aquatic plants can colonize spread to 159 lakes as of summer 2004. as well as composition and abundance of macrophytes. macrophyte detailed life history of Eurasian watermilfoil.e. dominated state can be difficult to achieve consult Smith and Barko (1990). foraging environment in the sub-canopy that tant to a conversion to a turbid state because may confine predators to small open pockets the hypolimnion in stratified pelagic zones act or bed edges (Killgore et al. 1993. Carpenter and Cottingham 1997. or low abundance of plank. Moss et al. and Bremigan 2002a). 1996.g. first found in Minnesota in 1987 in Lake Min- Jeppesen et al.in shallow. Nevertheless. eutrophic Minnesota lakes (Drake EFFECTS OF INVASIONS OF NON- and Valley in review). techniques include water level drawdowns. age-classes of largemouth bass are more effi- ham 1997). Lillie and Budd 1992. Management Eurasian watermilfoil grows rapidly. petually high nutrient loading from the Field studies suggest EWM depresses watershed (Cross and McInerny 2001). SAV in shal. lake users. Given low external and netonka. Bronmark and Weisner 1992. NATIVE PLANTS ON FISH HABITAT low Minnesota lakes serves a greater role in providing habitat for waterfowl and other wet. Shallow. Engel these deeper lakes. and the fer et al. Some fisheries only when it forms extensive. being within the Twin Cities metro area. ing or direct removal (e. Eurasian watermilfoil and curly-leaf pond- or one characterized by turbid water and little weed Potamogeton crispus are a concern to to no vegetation (Blindow et al. This results in an inhospitable 2004). Rather. and pH (Carpenter and Lodge 1986. 1997). Madsen 1997). Deeper lakes are generally more resis. Eurasian watermilfoil–Eurasian wa- thivorous fishes (“top-down” forces). Minnesota Department of Natural Resources internal mixing (i. High external nutrient loads. Madsen 1997. homo- of the same techniques used to rehabilitate geneous beds throughout the littoral zone shallow lakes may lead to improvements in (Keast 1984. These dense canopies reduce sub-canopy light. 1985. 1995). Young as a sink for nutrients (Carpenter and Cotting. increase termilfoil (EWM) was introduced into the U. high abundance of zooplanktivorous or ben. 1989). typically forming extensive homogeneous sur- substantial reductions to nutrient inputs and face canopies that displace native macrophytes control of planktivores through predator stock. In Minnesota lakes. For a Return from a turbid to a clear. local units of government. interfere with recreation.S. eutrophic lakes typically occupy one the establishment of non-native invasive spe- of two alternative states: one characterized by cies of submersed aquatic plants such as clear water and abundant aquatic vegetation.. without substantial intervention. the probability that phytoplankton and algae in the 1940s and has since spread throughout will dominate the water column (Carpenter et the nation (Couch and Nelson 1985). Hanson and Butler 1990. most littoral areas and maintain a clear-water state. 1991. Areas infested by large monospecific beds of EWM tend to have less abundant fish 8 . Herwig et al. prevent its proliferation and spread. Invasion by non-native species of land-associated species. Figure 3). (Madsen et al. Despite numerous measures to internal loading. submersed aquatic plants may displace native Hosts of unique biological and physi. oxygen. Schef.. deep lakes in cient feeders in native plant assemblages heavily developed or agricultural watersheds compared with dense EWM canopies (Valley often occupy a turbid-water state due to per. EWM has tivorous fish. 1993). “bottom-up” forces) or a (DNR). plant species and reduce the suitability of cal processes within shallow lakes affect the habitat for certain species of fish. It was al.

presence of native plants reduces the probabil- tivelyaffected (Weaver et al. Olson et al. with dominance highest in mesotrophic scribed for EWM. Canada (percent cover within the littoral zone) in 300 waterweed Elodea canadensis. 1991). He found lake trophic watermilfoil Myriophyllum sibiricum can form status was the best predictor of EWM domi. and invertebrates than do areas with diverse then. 2001). hypereutrophic condition (> 30 µg L-1 TP). mulative native plant cover.. 1998. naiads Najas spp. ity that EWM will dominate the littoral zone. EWM in-lake spreading can be EWM can be beneficial to fisheries if it occurs expected despite current control methods. Madsen (1998) investigated the corre. suggesting the fish populations generally are not nega. Nevertheless. decreased rapidly as lakes approached a plants (Keast 1984. Schematic representation of a canopied macrophyte monoculture (A) and a diverse macrophyte community (B). Dominance. 1997. 15 years. Figure 4). native lation between physical and chemical macrophyte species such as coontail Cerato- characteristics of lakes and EWM dominance phyllum demersum. Valley and Bremigan 2002b). and thus may affect ( >10 µg L-1 Total Phosphorus) to eutrophic fisheries in similar ways (Frodge et al. In fact. Nichols and Lathrop 1994). and northern lakes across the US. with in lakes that typically do not support much some displacement of native plants (Madsen et growth of native submersed species (Engel al. lakes ( < 30 µg L-1TP. Figure from Madsen (1997). A B Figure 3. 1990). 9 . however. In non-infested eutrophic lakes. if EWM is part of a diverse Madsen (1998) also documented that EWM plant community or if it grows in patches dominance was inversely proportional to cu- where open pockets permit fish movement. Nevertheless. Cheruvelil et al. monospecific canopies similar to those de- nance. displacement of native 1995) because more fish and invertebrates are plants may not be permanent and EWM domi- found in areas with EWM than areas devoid of nance may decline over time (approximately SAV (Pratt and Smokorowski 2003).

it can form dense mats that for these fish during winter and spring (J. habitats. which suggests that this plant provides habitat By late spring. may interfere with recreation and limit the Laurer. Prior to senescence. lakes. including juvenile game fish. Curly-leaf pondweed is currently algal blooms. CLP usually mersed vascular plant that was first noted in senesces. CLP remains green understanding is mostly anecdotal. As a response to warm water pondweed (CLP) is a perennial. soft-bottom areas (Nichols 1992). MN DNR Fisheries Biologist. it tures small fish. From Madsen (1998). sub. Curly-leaf pondweed –Curly-leaf Dobson 1985). rooted. fish sampling in stands of CLP cap- (Wehrmeister and Stuckey 1978). 1994) and 1945). CLP plants known to occur in 65 of the 87 Minnesota form vegetative propagules called turions counties (Exotic Species Program 1997). personal growth of native aquatic plants (Catling and communication. tween fish and CLP is sparse and thus present Unlike most native plants. 10 . therefore.Figure 4. is often the first plant to appear after ice-out. Miranda and Pugh 1997). temperatures in mid-summer. (hardened stem tips) from which new plants Curly-leaf pondweed can grow in a variety of sprout in the fall (Catling and Dobson 1985). but its most prolific growth occurs in Published data on the relationship be- shallow. During and viable under thick ice and snow cover spring. leading to increases in concentra- Minnesota circa 1910 (Moyle and Hotchkiss tions of phosphorus (Bouldan et al.S. Relationship between Eurasian watermilfoil dominance (percent littoral cover) and total phosphorus for 25 U.

tion in SAV cover. Removal of CLP Horseshoe Lake may have altered their distri- (or any other plant for that matter) without bution in the littoral area. but limited spawning found effects on the fish community shifting it habitat in spring.. MN DNR Fisheries Texas reservoir).4-D. effort has been devoted and black crappie were significantly positively to their management (Madsen 1997). higher growth. catfish. R. to alter fish habitat.100 ha bonne. Lake than the abundance of SAV. Nuisance growth of CLP often whole-lake cover prior to treatment) and 60% occurs in shallow. Despite continued absence of SAV in 1993 and 1994. mechanisms underlying these observations 1992). Biological control with grass carp– structure of sunfish populations are common Grass carp Ctenopharyngodon idella elimi- in some lakes dominated by CLP (B. such as shad Dorosoma spp. based on summer air total plant abundance and patchiness. 1995). area. Control methods can influence on fish dynamics in Little Horseshoe target CLP because it grows under ice in win. Naturally. we discuss cover. This repre- ter and is abundant early in the spring before sents an example where a larger physical force many native macrophytes. observed growth onstrate that the magnitude of effect is was neither consistently higher nor lower than dependent on the degree to which APM alters the predicted growth. Growth indices of bluegill. personal communications). Al- evaluations of the effects of aquatic plant though slower growth generally occurred after management (APM) on fish populations dem. or high lake cies such as largemouth bass and bluegill to productivity of infested lakes are potential fac. not to mention impair rec. In addition. the aquatic herbicide endothall and promelas. and size. removed all SAV (approximately 50% commersoni. one dominated by pelagic and river species tors. Field correlated with summer air temperature. in habitat. nated all vegetation in Lake Conroe (8. (i. More studies have been more than the denuded regions of the littoral conducted with Eurasian watermilfoil and hy. Below. reation and aesthetics. Ramsell. This sirable for native Minnesota fish. and At the end of spring when CLP white bass (Bettoli et al. Grass carp are senesces or dies back. carp. complete increased algal blooms due to the release of elimination of aquatic plants would not be de- phosphorus into the water column. changes in largemouth bass. 1993). remov. which was 44% hydrilla Biologists. 11 . However.. and northern pike abundance DIRECT AND INDIRECT EFFECTS OF and growth could not be attributed to reduc- AQUATIC PLANT MANAGEMENT tions in SAV abundance. Ner. from a community dominated by littoral spe- niles after CLP senescence. high predation upon juve. If bluegill were using shallow emergent drilla Hydrilla verticillata. temperature. bluegill in Little cant challenge to managers. Elimination of all vegetation had pro- have not been studied. fish growth Given the ability of non-native plants and abundance did not significantly change. perhaps by using replacement by native plants compromises fish floating-leaf and emergent vegetation cover and wildlife habitat. largemouth bass. eutrophic basins (lake types of the floating-leaf vegetation in 1992 2 and 6) where native SAV has been lost due (Radomski et al. fathead minnows Pimephales type 4).e. dramatic shift in habitat conditions may favor Whole-lake chemical control–In Little disturbance-tolerant fish species such as black Horseshoe Lake (Crow Wing Co. chemical treatment of SAV. climate) exerts a greater effect on the fish ing CLP from shallow eutrophic lakes and populations and overrides within-lake changes promoting native species represents a signifi. after the elimination of SAV.Also low abundance. MN. Despite a large reduc- to the loss of water clarity and carp. bluegill. the decrease in indiscriminate herbivores and feed on native submersed vegetation often is followed by plant species as well. foraging success of largemouth bass some relevant findings. The prior to grass carp stocking (Bettoli et al. and white sucker Catostomus 2. Little is known regarding the effects Water temperature (which varied greatly of alternative control methods of CLP on fish among years) apparently had a more profound habitat and populations. lake bullhead.

before and after these treatments were lost as a porarily experienced greater growth rates after result of past treatments in these lakes (J. but less than 20 ppb was specu. Preliminary analyses in these lated.. within the littoral zone (Figure 5). chemical control of EWM by fluridone (> ment (Getsinger et al. Schutz Lake Carver Co.. these lakes were treated. Valley Zumbra Lake were not sampled during the and Bremigan 2002b. Crowell cases. 1999). 2001). Concentration and contact time was type 4] and Eagle Lake. Detailed hy- wild celery Vallisneria americana (common droacoustic assessments were conducted in ‘pioneer’ species in mesotrophic lakes and Schutz Lake during 2002 and 2003 using resilient to fluridone) rapidly recolonized de. low- Schneider (2000) evaluated the indi. [lake EWM. biovolume In another Michigan study. plant species (e. speculated that some fluridone-sensitive native tion with little recovery of native SAV (whole. Largemouth bass and bluegill tem. methods described by Valley et al. lake type community that was present prior to the treat- 4). In most plant biomass and water clarity (W. and largemouth lilies) was 37% just prior to fluridone applica- bass were too small to determine whether tion (early June 2002. chara and personal communication).[lake type not evaluated.. 2001. there effects did not last into the post-treatment was no effect of the treatments on whole-lake year.and northern pike may not have changed ap. average littoral biovolume declined to populations. Sample sizes for umn occupied by vegetation. However.g. eight was 3% (Figure 5C). 6. mesotrophic Michigan lakes were also treated 12 . Nevertheless. Average proved significantly in many lakes after littoral biovolume (percent of the water col- treatment with fluridone. (in press). presumably because Madsen. Elodea canadensis and lake extent was not documented. However. post-treatment year (Pothoven 1996). nuded littoral areas and presumably increased This analysis demonstrated a drastic decline in the habitat heterogeneity in these lakes. Several nongame species treatment on fish and invertebrate populations present prior to the fluridone treatments in were detected (Cheruvelil et al. In June 2003. Carver Co. As a the total abundance of submersed vegetation result. lectively remove EWM. Hanson 2001). excluding water yellow perch. diverse plant (Parkers and Zumbra. Mississippi State University. Figure 5A). However. per- of greater prey availability. fluridone initially removed nearly all Minnesota Department of Natural Resources. Carver Co. with low doses (5 – 7 ppb) of fluridone to se- preciably with the elimination of SAV. northern pike. 2]) in 2001.5% (Figure 5B). it is 10 ppb) greatly reduced the cover of vegeta. despite continuing declining vegetation cover of SAV and no negative effects of the in Zumbra Lake. most sonal communication). In August fluridone had a significant effect on these 2002. dose fluridone applications (5 ppb target con- rect effects of whole lake treatments of the centration) were applied to three eutrophic herbicide fluridone on game fish populations lakes infested with EWM (Crooked Lake. These treatments did In two eutrophic Minnesota lakes not impact the fluridone-hardy. Schneider (2000) observed large lakes demonstrate large reductions in total changes in the plant community. in 11 mesotrophic Michigan lakes infested by Hennepin Co. submersed vegetation. In a recent study in Minnesota. bluegill and crappie size structure im. Pothoven et Ceratophyllum demersum) that were absent al.

June 2003 (C). and August 2003 (D). Extensive floating-leaf vegetation was present during both August sampling periods. 13 . A Figure 5. Distribution of vegetation biovolume (percent of water column occupied by vegetation) in Schutz Lake just prior to 5 ppb fluridone applications during June 2002 (A). August 2002 (B).

B white and yellow waterlilies Figure 5. Continued 14 .

Continued 15 . C Figure 5.

Continued 16 . White and Yellow waterlilies Figure 5.

especially in large or primarily by EWM in several lakes. 1997). littoral biovolume declined to Mechanical harvesting can uninten- 0. 1997. 1999. CONCLUSIONS deed. me. 1995. pike and sunfish populations in MN lakes with ment lakes. Radomski and Goeman 2001). abundance that is optimal for Minnesota fish In Wisconsin. slow) their tect significant effects if they did occur spread among lakes. lake type 4) on largemouth bass. This recognizes that non-native plants do 17 . we must growth for some age-classes of largemouth accept that eradication of heavily infested bass and bluegill. further exacerbating turbidity. thus potentially benefiting game foil or curly-leaf pondweed beds could benefit fish such as largemouth bass and bluegill game fish populations because this type of (Smith 1993. with ap. and trations designed to selectively remove EWM.11% vegetation removal) in nearshore SAV and emergent vegetation (typi- two Minnesota lakes (Mary and Ida lakes. tionally kill juvenile fish. and amounts of EWM from the water column fa. Hal- fluridone had negative indirect effects on total ler et al. (Carpenter et al. the extent and distribution of manipulated ar- chanical control of vegetation with harvesting eas. Within infested lakes. domski and Goeman (2001) documented amined. Trebitz et al.In August 2003. 1997). however. Carp inhabited all three treatment mowing the surface (Unmuth et al. consequently. prevent (or more realistically. cordingly. populations. they reported increased first reduced biomass and mean size of northern year growth of largemouth bass in the treat. we cannot rates and size structure of bluegill and large. probability of spread is high regardless of any Olson et al.. However removing tiple plots (6% . (1997) determined approximately 20% . Olson et al. amphibians. Removing vegetation will have vary- ments. Despite low target concen. cally used as nursery areas) can have adverse Wright Co. plants returned to lakes is not a realistic management goal. however. deep lakes where SAV cover is limited. Booms (1999) reported losses of 406 per hec- vored algal growth rather than recolonization tare of harvested EWM. effects on fish diversity and game fish produc- bluegill. turtles (Haller et al. Jennings et authors did not detect any significant effects of al. Harvesting and maintaining deep-cut can be used to increase edge and vegetation channels through offshore Eurasian watermil- patchiness. Ac- pretreatment densities after one growing sea. cess without significant losses of fish habitat. this management strategy manage the nuisance to allow recreational ac- requires harvesting each year. 1998). 1998).6% (Figure 5D). and northern pike populations. Trebitz et al. manipulation was not large enough to effect detectable changes in population metrics. (1980) reported losses of 460 juvenile plant cover because the removal of large fish per hectare of harvested hydrilla. In. actions should include those that son. For now. Cross et al. responsibly prescribe a uniform level of SAV mouth bass populations. Ra- the harvesting on the population metrics ex. Cross et al. Booms 1999). lakes and may have resuspended bare sedi. fewer of barren areas by the few remaining native fish are killed by deep-cutting compared with species. Nevertheless. Extensive pilot work was done plant species. (1998) har. ing effects on fish populations depending on Mechanical control–In theory. high coverage of vested many deep-cut channels perpendicular SAV within the littoral zone is highly impor- to shore throughout littoral zones dominated tant to many fish species. Trebiz et al. Apparently. (1992) speculated the little emergent or floating-leaf vegetation. proximately 20% total vegetation removal for With respect to non-native aquatic each lake. (1998) documented increases in current control technique. our primary goal should be to prior to this study to ensure the ability to de. The tion (Brian and Sarnecchia 1992.40% of a fully vegetated Given the diversity of Minnesota lakes littoral zone would need to be removed in and the unique fish and habitat relationships patchy fashion to affect population growth that are often found in these lakes. Olson et al. (1992) examined the effect of harvesting mul. manipulation increases edge (Trebitz et al. 1980.

lakes. dominated. socio-economic. lake types 2. life perspective. the context of APM. are threatened by rapid development. 6. a precautionary ment will not succeed (Rosenberg 2002). 15 feet may be treated with chemicals or no Vegetated or woody nearshore habi. A fundamental principle of being used in Atlantic salmon management the precautionary approach requires proof that (NASCO 2000). community and the onus should be on indi- sider the lake type (i. recreational access must be balanced fish habitat. and 8) harm than to later repair it. tion will not have significant adverse affects foil or curly-leaf pondweed into lakes will lead on the resource (FAO 1996). ary framework. From a fisheries and wild. For example. Current limits for SAV control that risks of repeated treatments are unknown and state no more than 15% of the area less than must be considered.g. regulation for APM where risks are larger). A desirable than no SAV. Figure 1. enforcement. Richards and successful precautionary management Maguire 1998. but rather habitat. precautionary manage- To achieve this end. managers should carefully con. there is little evidence proposed resource exploitation or manipula- to suggest introductions of Eurasian watermil. 4. For mesotrophic lakes program management plan should include with diverse plant communities. in to dramatic declines in fisheries. However. nently altered many lakes within agricultural Current APM thresholds may be safe for some areas of Minnesota and the Twin Cities metro. program and lake management plans. prevents undesirable outcomes.serve as habitat.g. Restrepo et al. to uncertainty. lakes needed for soft water lakes and large. This decision analysis dem- 18 . paid to these forces. more use and SAV removal practices have perma.e. transparency. commercial. action in advance of formal proof and shifts and political questions are raised and influence the burden of proof to those challenging pre. compromise lake. management approach has been encouraged Simplicity. by mass. Finally. Large manipulations significantly harm fish populations (FAO such as whole-lake herbicide treatments in 1996).. socio-economic and political ity is the key objective and recreation and considerations must be built into a precaution- commercial benefits must be regulated to pro. overly abundant SAV is more monitoring. In fact. a precautionary approach requires that protecting and managing Minnesota’s aquatic management be evaluated. but regulatory frameworks are key components of globally as well (FAO 1996. In others parts of the state. removal. however. the cumulative long-term habitat. and recreational benefits to the and monitors and addresses non-compliance. and their aggregate impact on fish less risky. hard water lakes where SAV is more limited. stricter thresholds may be politan area. 1999. and evaluation. by the target species can Important elements of the precaution- lead to large declines in total plant abundance ary approach are the rules controlling SAV and water clarity. Auster (Rosenberg 2002). Table 1) and viduals or groups proposing manipulation to associated risks prior to making aquatic plant habitats to prove that their activities will not management decisions. and flexibility of not only in Minnesota (Welling 2001). deep. Building decision frame- 2001). the direction of policy decisions are currently cautionary policy.. with the long-term sustainability and integrity It is precautionary to have different of the lake ecosystem. The Minnesota DNR is charged with Also. The precautionary approach advocates works where key biological. It follows. within any particular define limits that when exceeded.. alterations to lakes in- When managing nuisance aquatic variably has some effect on the lake’s fish plant species. Accordingly. Without significant attention tect long-term sustainability. more than 50% of this area for mechanical tats are especially important for fish harvesting should not be viewed as some de- populations and any removal is a loss of fish sirable level of plant removal. whole-lake mechanisms to monitor and control shoreline treatments with selective herbicides may be alterations. Long-term sustainabil. Past and ongoing land thresholds for different methods (e. Evaluations should resources and associated fish communities for attempt to determine if management is robust their intrinsic values and long-term ecological. that it is easier to prevent eutrophic lakes (e. people of Minnesota.

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S. R. Perleberg. Enger. Anderson. Edited by: Paul J. Fisheries Research Supervisor 25 . Nerbonne. W. M. B. C. and D. Fish and Wildlife Research Manager Charles S. Wright. Drake. Pereira. Anderson. Wingate. D. Norrgard. ACKNOWLEDGMENTS Many people offered helpful comments and suggestions on earlier drafts of this manu- script and include: J. D. Welling. Crowell.