IDENTIFICATION, ECOLOGY AND CONTROL OF...

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IDENTIFICATION, ECOLOGY AND CONTROL OF NUISANCE FRESHWATER ALGAE KENNETH J. WAGNER, Ph.D, CLM WATER RESOURCE SERVICES

Transcript of IDENTIFICATION, ECOLOGY AND CONTROL OF...

IDENTIFICATION, ECOLOGY AND CONTROL OF NUISANCE

FRESHWATER ALGAE

KENNETH J. WAGNER, Ph.D, CLMWATER RESOURCE SERVICES

PART 1: THE PROBLEMS

ALGAL PROBLEMS

Algal problems include:• Ecological imbalances

• Physical impacts on the aquatic system

• Water quality alteration

• Aesthetic impairment

• Taste and odor

• Toxicity

ALGAL PROBLEMSEcological Imbalances

High algal densities:• Result from overly successful growth processes

and insufficient loss processes

• Represent inefficient processing of energy by higher trophic levels

• May direct energy flow to benthic/detrital pathways

• May actually reduce system productivity (productivity tends to be highest at intermediate biomass)

ALGAL PROBLEMSAesthetic Impairment

High algal densities lead to:

• High solids, low clarity

• High organic content

• Fluctuating DO and pH

• “Slimy” feel to the water

• Unaesthetic appearance

• Taste and odor

• Possible toxicity

ALGAL PROBLEMSTaste and Odor

• At sufficient density, all algae can produce taste and odor by virtue of organic content and decay

• Some algae produce specific taste and odor compounds that are released into the water

• Geosmin and Methylisoborneol (MIB) are the two most common T&O compounds; these can induce T&O at very low concentrations

ALGAL PROBLEMSTaste and Odor

T&O by blue-greens• Anabaena, Aphanizomenon, Microcystis, and

Oscillatoria are most common T&O producers, but many other genera produce T&O as well

• Geosmin and MIB often produced• Odors usually include musty, grassy, and

septic• May be produced by planktonic or benthic

growths

ALGAL PROBLEMSTaste and Odor

T&O by greens• Chara, Cladophora, Chlorococcales

(Dictyosphaerium, Scenedesmus, Pediastrum, Hydrodictyon), Volvocales (Chlamydomonas, Volvox), and desmids (Staurastrum, Closterium, Cosmarium, Spirogyra) are primary greens causing T&O, but almost any genus can induce T&O at high density

• Main odors are fishy, skunky, musty, grassy and septic (wide range, species-specific)

• Not usually as severe as with blue-greens

ALGAL PROBLEMSTaste and Odor

T&O by diatoms• Melosira/Aulacoseira, Stephanodiscus,

Cyclotella, Asterionella, Fragilaria/Synedra, and Tabellaria are primary diatoms causing T&O, usually only at high density

• Main odors are geranium, spicy and fishy, with occasional musty or grassy scent

• Not as severe as with blue-greens unless diatom density is very high

ALGAL PROBLEMSTaste and Odor

T&O by goldens• Synura, Dinobryon, Mallomonas, Uroglenopsis,

and Chrysosphaerella are major T&O producers, others impart odor at high density Main odors are cucumber, violet, spicy and fishy

• Can produce substantial T&O even at low density

ALGAL PROBLEMSTaste and Odor

T&O by other algae• Dinoflagellates can produce a

fishy or septic odor at elevated densities

• Euglenoids can produce a fishy odor at elevated densities

• Major die-off of high density algae may produce a septic smell

• Actinomycetes, a non-photosynthetic bacterium, can also produce geosmin and MIB

ALGAL PROBLEMSTaste and Odor

Control of T&O• Minimizing algal density and specific T&O

producers is desirable for T&O control• Activated carbon removes most T&O, but

effective use is slow and increases treatment cost

• Aeration/mixing may oxidize/volatilize T&O compounds, but not completely effective

• Possible toxic link, but no severe direct health effects known from T&O; aesthetic issues can be major

ALGAL PROBLEMSToxicity-Cyanotoxins

• Cyanobacteria are the primary toxin threats to people from freshwater; acute toxicity is rare, but chronic effects may be significant and are difficult to detect.

• Research (e.g., 3 papers in Lake and Reservoir Management in September 2009) indicates widespread occurrence of toxins but highly variable concentrations, even within lakes.

• Water treatment in typical supply facilities is adequate to minimize risk; the greatest risk is from substandard treatment systems and direct recreational contact.

• Some other algae produce toxins - Prymnesium, or golden blossom, can kill fish; marine dinoflagellates, or red tides, can be toxic to many animals and humans.

ALGAL PROBLEMSToxicity-Cyanotoxins

• Dermatotoxins– produce rashes and other

skin reactions, usually within a day (hours)

• Hepatotoxins– disrupt proteins that keep

the liver functioning, may act slowly (days to weeks)

• Neurotoxins– cause rapid paralysis of

skeletal and respiratory muscles (minutes)

ALGAL PROBLEMSToxicity-Which Blue-Green Has Which

Toxin?Blue-Green Algae Microcystin Nodularin Cylindro-

spermopsin Anatoxin Saxitoxin Dermato-

toxin Anabaena X X X X Anabaenopsis X Aphanizomenon X X X Cylindrospermopsis X Cylindrospermum X X Hapalosiphon X Lyngbya X X Microcystis X X Nodularia X X Nostoc X Oscillatoria X X X Schizothrix X Umezakia X

More forms found each year – methods or actual increase?

ALGAL PROBLEMSToxicity-Analytical Methods

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ELISAPhosphatase Assay

ALGAL PROBLEMSToxicity- Key Issues

• Acute and chronic toxicity levels -how much can be tolerated?

• Synergistic effects - those with liver or nerve disorders at higher risk

• Exposure routes - ingestion vs. skin

• Treatment options - avoid cell lysis, remove or neutralize toxins

ALGAL PROBLEMSRecommended Toxicity Precautions

• Monitor algal quantity and quality

• If potential toxin producers are detected, increase monitoring and test for toxins

• For water supplies, incorporate capability to treat for toxins (PAC or strong oxidation seem to be best)

• For recreational lakes, be prepared to warn users and/or limit contact recreation

• Avoid treatments that rupture cells

PART 2: ALGAL FORMS

Algal FormsAlgal “Blooms”

♦ Water discoloration usually defines bloom conditions

♦ Many possible algal groups can “bloom”

♦ Taste and odor sources, possible toxicity

♦ Potentially severe use impairment

Algal FormsAlgal Mats

♦ Can be bottom or surface mats -surface mats often start on the bottom

♦ Usually green or blue-green algae

♦ Possible taste and odor sources

♦ Potentially severe use impairment

Algal Types: Planktonic Blue-greens

Aphanizomenon

Microcystis

Anabaena

Coelosphaerium

Algal Types: Planktonic Blue-greensPlanktolyngbya Planktothrix

(Oscillatoria)

Algal Types: Planktonic Blue-greensGloeotrichia

Algal Types: Planktonic Blue-greensCylindrospermopsis

•A sub-tropical alga with toxic properties is moving north.

•Most often encountered in turbid reservoirs in late summer, along with a variety of other bluegreens.

Algal Types: Mat Forming Blue-greens

Oscillatoria

Lyngbya/Plectonema

Algal Types: Planktonic GreensPediastrum

Scenedesmus

Lagerheimia Dictyosphaerium

Schroederia

Oocystis

(Order Chlorococcales)

Algal Types: Planktonic Greens

Chlamydomonas

Pyramichlamys

Carteria

Eudorina

Volvox

(Order Volvocales)

Algal Types: Mat Forming Greens

Cladophorales, with Cladophora (above), Rhizoclonium and Pithophora

Zygnematales, with Spirogyra (above), Mougeotia and Zygnema

Hydrodictyon, an unusual type of Chlorococcales

Oedogoniales, with Oedogonium and Bulbochaete

Algal Types: Mat Forming GreensZygnematales - Unbranched filaments, highly

gelatinous

MougeotiaSpirogyra

Zygnema

Mats trap gases and may float to surface

Cladophorales - Large, multinucleate cells, reticulate chromatophores, filamentous forms

Pithophora

RhizocloniumCladophora

Algal Types: Mat Forming Greens

Algal Types: Mat Forming Greens

Hydrodictyon

Oedogonium

Bulbochaete

Algal Types: Planktonic Diatoms

Aulacoseira

Asterionella

Cyclotella

Algal Types: Planktonic Diatoms

Fragilaria NitzschiaTabellaria

Algal Types: Plankton Goldens

Dinobryon

Synura

Chrysosphaerella

Prymnesium

Algal Types: Mat Forming GoldensTribonema

Vaucheria

Algal Types: Other Plankton

Ceratium

Peridinium

Euglena

Trachelomonas

Phacus

(Euglenoids)

(Dinoflagellates)

PART 3: THE ECOLOGICAL BASIS FOR ALGAL CONTROL

ALGAL ECOLOGYKey Processes Affecting Abundance

Growth Processes• Primary production – controlled by light

and nutrients, algal physiology• Heterotrophy – augments primary

production, dependent upon physiology and environmental conditions

• Release from sediment – recruitment from resting stages, related to turbulence, life strategies

ALGAL ECOLOGYKey Processes Affecting Abundance

Loss Processes

• Physiological mortality – inevitable but highly variable timing – many influences

• Grazing – complex algae-grazer interactions

• Sedimentation/burial – function of turbulence, sediment load, algal strategies

• Hydraulic washout/scouring – function of flow, velocity, circulation, and algal strategy

ALGAL ECOLOGYKey Processes Affecting Abundance

Annual variability in growth/loss factors in dimictic temperate lakes

• Winter –– Possible ice cover, reverse stratification

– Under ice circulation is an important factor

– Low light and temperature affect production

– Variable but generally moderate nutrient availability

– Possibly high organic content

– Grazer density usually low

ALGAL ECOLOGYKey Processes Affecting Abundance

Annual variability in growth/loss factors in dimictic temperate lakes

• Spring/fall –– Isothermal and well-mixed

– Relatively high nutrient availability

– Light increases in spring, decreases in fall

– Temperature low to moderate

– Stratification setting (spring) or breaking down (fall)

– Grazer density in transition (low to high in spring, high to low in fall)

ALGAL ECOLOGYKey Processes Affecting Abundance

Annual variability in growth/loss factors in dimictic temperate lakes

• Summer –– Potential stratification, even in shallow lakes

– Often have low nutrient availability

– Light limiting only with high algae or sediment levels

– Temperature vertically variable – highest near surface

– Vertical gradients of abiotic conditions and algae

– Grazer densities variable, often high unless fish predation is a major factor

ALGAL ECOLOGYPhytoplankton Succession - Winter

• Under ice, mainly cryptophytes, chrysophytes, diatoms, naked dinoflagellates

• Some non-nitrogen fixing blue-greens, but also possibly Aphanizomenon

ALGAL ECOLOGYPhytoplankton Succession – Early Spring

• Most lakes experience rapid increase in algal density

• Diatoms, cryptophytes, and chrysophytes tend to dominate

• Temperature is a primary control factor

ALGAL ECOLOGYPhytoplankton Succession – Late Spring

• Diatoms tend to dominate, often with Chlorococcalean greens and cryptophytes

• Chrysophyte blooms possible

• The later the spring maximum, the less likely that diatoms will dominate

• Overwintering benthic colonies may be recruited to the plankton community

ALGAL ECOLOGYPhytoplankton Succession – Late Spring

• Increasing light cues spring blooms where nutrients are available

• Grazing and algal settling increases during spring with rising water temperature

• Clear water phase often results from loss processes overshadowing growth in

late spring

ALGAL ECOLOGYPhytoplankton Succession – Early Summer• Increasing light and temperature• Nutrient availability may decrease as internal sources

are isolated by stratification - N, P, Si• Grazing, sinking and metabolism all increasing• Composition will depend upon resource gradients -

low N:P favors blue-green Nostocales and green Volvocales, high N:P favors other chlorophytes, chrysophytes and dinoflagellates

• N:P ratio at sediment:water interface can be more important than ratio in epilimnion

• High variability among lakes and within lakes among years

ALGAL ECOLOGYPhytoplankton Succession – Early Summer

• Greens increase, especially Volvocales and Chlorococcales

• Some blue-greens appear at increased density

ALGAL ECOLOGYPhytoplankton Succession – Early Summer

• May get metalimneticblooms of cryptophytes, chrysophytes, dinoflagellates or blue-greens, especially Planktothrix

ALGAL ECOLOGYPhytoplankton Succession – Late Summer

• High bloom potential -most often blue-greens, but also thecate dinoflagellates, large desmids

• Temperature is a major factor in blooms

ALGAL ECOLOGYPhytoplankton Succession – Late Summer

• Nitrogen-fixing blue-greens especially common bloomers

• May get blooms of non-N fixing blue-greens if N is high

• May get population oscillations between N-fixers and non-N-fixers

ALGAL ECOLOGYPhytoplankton Succession – Late Summer

• May also get green or blue-green mats floating to the surface

• Most often Cladophorales or Oscillatoriales

• Mats tend to start on bottom, floating to surface after trapping gases

ALGAL ECOLOGYPhytoplankton Succession – Late Summer

• A variety of other algae may be mixed in, but dominance by one taxon or a few taxa is typical in fertile lakes

• Growth often nutrient limited, but dense surface growths may create light limitation below

• Grazing may be high, may be selective

ALGAL ECOLOGYPhytoplankton Succession - Autumn

• “Leftover” blue-greens may remain abundant well into autumn, exception is Cylindrospermopsiswhich often blooms late summer into mid-fall

• Metalimnetic growths often reach surface upon mixing

• Diatoms often assume dominance after turnover

ALGAL ECOLOGYPhytoplankton Succession - Autumn

• Desmids also respond positively to mixing

• As the water cools, the oil-laden Botryococcus may become abundant

• Nutrients tend to be abundant after turnover

ALGAL ECOLOGYPhytoplankton Succession - Notes

• Biomass can vary by 1000X from winter minimum to late summer maximum in temperate to polar waters

• Primary productivity and biomass may not correlate due to time lags, cell size and nutrient or light limitations

• Highest productivity normally at intermediate biomass (Chl a = 10 ug/L)

• High pH during blooms may trigger resting cyst formation

ALGAL ECOLOGYTrophic Gradients

• Based on decades of study, more P leads to more algae

• More algae leads to lower water clarity, but in a non-linear pattern

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ALGAL ECOLOGYTrophic Gradients

• As algal biomass rises, a greater % of that biomass is cyanobacteria. So more P = more algae + more cyanobacteria.

From Canfield et al. 1989 as reported in

Kalff 2002

ALGAL ECOLOGYTrophic Gradients

• High P also leads to more cyanobacteria, from considerable empirical research. Key transition range is between 10 and 100 ug/L

(10 ug/L) (100 ug/L)From Watson et al. 1997 L&O 42(3): 487-495

Three step process:– Don’t lose your head– Get ducks in a row– Don’t bite off more than you can chew

PART 4: METHODS OF ALGAL CONTROL

Algal Control: Watershed Management

Understand the watershed and

manage it!

Algal Control: Watershed Management♦ Source controls• Banning certain high-impact actions• Best Management Practices for minimizing risk of

release

♦ Pollutant trapping• Detention• Infiltration• Uptake/treatment• Maintenance of facilities

Watershed management should be included in any successful long-term algal management plan

Algal Control: In-Lake ManagementData needs include:♦ Algal types and quantity (planktonic and benthic)♦ Water quality (nutrients, pH, temp., oxygen,

conductivity, and clarity over space and time)♦ Inflow and outflow sources and amounts, with

water quality assessment♦ Lake bathymetry (area, depth, volume)♦ Sediment features♦ Zooplankton and fish communities♦ Vascular plant assemblage All of which facilitate a hydrologic and nutrient

loading analysis and biological assessment essential to evaluating algal control options

Algal Control: In-Lake Management

In-lake detention and wetland

systems to limit nutrient inputs

Fulfilling a watershed

function in the lake

Algal Control: In-Lake ManagementDredging

♦ Dry (conventional)

♦ Wet (bucket/dragline)

♦ Hydraulic (piped)

♦ Removes nutrient reserves

♦ Removes “seed” bank

♦ Potential mat control

Algal Control: In-Lake Management

Harvesting

♦ Not feasible for many algal nuisances

♦ Possible to collect surface algal mats

Algal Control: In-Lake Management

Dilution and Flushing

♦ Add enough clean water to lower nutrient levels

♦ Add enough water of any quality to flush the lake fast enough to prevent blooms

Algal Control: In-Lake ManagementSelective withdrawal

♦ Often coupled with drawdown

♦ May require treatment of discharge

♦ Best if discharge prevents hypolimnetic anoxia

Algal Control: In-Lake ManagementDye addition

♦ Light limitation -not an algaecide

♦ May cause stratification in shallow lakes

♦ Will not prevent all growths, but colors water in an appealing manner

Algal Control: In-Lake ManagementSonication

♦ Disruption of cells with sound waves – may break cell wall or just dissociate plasma from wall

♦ Used in the lab to break up algal clumps

♦ Won’t eliminate nutrients, but may keep algae from growing where running all the time

♦ Varied algal susceptibility

Algal Control: In-Lake ManagementAlgaecides

♦ Relatively few active ingredients available

♦ Copper-based compounds are by far the most widely applied algaecides

♦ Peroxides gaining acceptance

♦ Some use of endothall

♦ Effectiveness and longevity are issues

Algal Control: In-Lake ManagementAbout copper

♦ Lyses cells, releases contents into water♦ Formulation affects time in solution and

effectiveness for certain types of algae♦ Possible toxicity to other aquatic organisms♦ Long term build up in sediment a concern, but no

proven major negative impacts♦ Resistance noted in multiple nuisance blue-greens

and greens♦ Usually applied to surface, but can be injected

deeper by hose♦ Less effective at colder temperatures

Algal Control: In-Lake ManagementAbout peroxide

♦ Lyses cells, but more effective on thin walled forms; less impact on most diatoms and greens

♦ Degrades to non-toxic components; adds oxygen to the water, may oxidize some of the compounds released during lysis

♦ Typically applied to surface, but may reach greater depth with adequate activity (slower release/reaction rate)

♦ No accumulation of unwanted contaminants in water or sediment

♦ Considerably more expensive than copper

Algal Control: In-Lake ManagementProper Use of Algaecides

♦ Prevents a bloom, not removes one♦ Must know when algal growth is accelerating♦ Must know enough about water chemistry to

determine most appropriate form of algaecide♦ May involve surface or shallow treatment where

nutrients are fueling expansion of small population♦ May require deep treatment where major migration

from sediment is occurring♦ May require repeated application, but at an

appropriate frequency - if that frequency becomes too high, recognize that the technique requires adjustment or will not be adequate for the long-term

Algal Control: In-Lake Management

Phosphorus inactivation - anti-fertilizer

treatments

Algal Control: In-Lake Management♦ Iron is the most

common natural binder, but does not hold P under anoxia

♦ Aluminum is the most common applied binder, multiple forms, permanent results, toxicity issues

♦ Calcium used in some high pH systems

♦ Used for water column or sediment P

Algal Control: In-Lake Management

Factors in Planning LakeTreatments:

• Existing P load, internal vs external

• Sources and inactivation needs – field and lab tests

• System bathymetry and hydrology

• Potential water chemistry alteration - pH, metals levels, oxygen concentration

• Potentially sensitive receptors - fish, zooplankton, macroinvertebrates, reptiles, amphibians, waterfowl

• Accumulated residues - quantity and quality

Algal Control: In-Lake Management

Lake Water Column Treatment:

• Doses vary - need 5-20 times TP conc.

• Can achieve >90% P removal, 60-80% more common

• Effects diminish over 3-5 flushings of the lake

Algal Control: In-Lake ManagementLake Sediment

Treatment:

• Can reduce longer-term P release

• Normally reacts with upper 2-4 inches of sediment

• Dose usually 25-100 g/m2 - should depend upon form in which P is bound in sediment

Algal Control: In-Lake ManagementSediment Dose Calculation

1. Oldest approach is to treat at maximum level that maintains pH >6; may need to do jar tests and repeat application over time to achieve goal

2. Alternative older method is to measure accumulated hypolimnetic P during stratification, or change in lake P over a dry period, or release from incubated sediment, translate into sediment P concentration (per square meter), then treat with dose at least 10 times that level on areal basis – buffer if necessary to control pH

3. Most current approach is to determine available P in sediment (Rydin and Welch method), treat aliquots of sediment slurry with aluminum, re-test for P availability, graph results and evaluate in context of target P level, diminishing returns, and cost; dose of 10 to 100 times available sediment P level expected

Algal Control: In-Lake ManagementMethods for Minimizing Aluminum Toxicity

Aluminum dose at any one time should be <10 mg/L, preferably <5 mg/L

When buffering alum with aluminate, use a 2:1 ratio of alum to aluminate, by volume, to avoid pH change (can be adjusted – 1.8 to 2.1 common)

Treat defined areas of the lake in a pattern that minimizes contiguous area treated at once (patchwork with adjacent blocks not treated sequentially)

Apply aluminum at enough depth to create a surface refuge (can even treat below thermocline)

Secchi Disk Transparency in Hamblin Pond, 1992-2003

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Hamblin Pond ExampleCape Cod, MA

P levels dramatically reduced, water clarity substantially increased

Long Pond ExampleCape Cod, MA

P levels dramatically reduced, but only for a year; clarity remains high through 3 years

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Algal Control: In-Lake ManagementSelective nutrient

addition♦ Addition of nutrients

(most likely N or Si) to shift ratio (N:P:Si) to favor more desirable algae

♦ Sometimes practiced in association with fertilization for fish production, but rare in water supplies or recreational lakes

Algal Control: In-Lake Management

Aeration/Mixing

Algal Control: In-Lake ManagementAeration/mixing can work by:

♦ Adding oxygen and facilitating P binding while minimizing release from sediments

♦ Physical mixing that disrupts growth cycles of some algae

♦ Alteration of pH and related water chemistry that favors less obnoxious algal forms

♦ Turbulence that neutralizes advantages conveyed by buoyancy mechanisms

♦ Creation of suitable zooplankton refuges and enhancement of grazing potential

Algal Control: In-Lake ManagementKey factors in aeration:

♦ Adding enough oxygen to counter the demand in the lake (usually about 75% from sediment) and distributing it where needed in the lake

♦ Maintaining oxygen levels suitable for target aquatic fauna (fish and invertebrates)

♦ Having enough of a P binder present to inactivate P in presence of oxygen

♦ Not breaking stratification if part of goal is to maintain natural summer layering of the lake

Algal Control: In-Lake ManagementDestratifying aeration:

Lake is mixed, top to bottom, input of oxygen comes from bubbles and interaction with lake surface

Algal Control: In-Lake ManagementNon-destratifying aeration:

Bottom layer is aerated, but top layer is unaffected; oxygen input comes bubbles (can be air or oxygen)

Algal Control: In-Lake ManagementKey factors in mixing:

♦ Moving enough water to prevent stagnation; may mix whole lake or just the top layer (if any)

♦ Fostering greater homogeneity in mixed zone and greater interaction with the atmosphere (oxygen and pH effects may be large)

♦ Getting enough motion or change in water quality to disrupt target algal species; moving algae to dark zone helps, but may be possible to disrupt with only surface layer mixing

Algal Control: In-Lake ManagementMixing systems:

Algal Control: In-Lake ManagementBarley straw as an algal

inhibitor

♦ Decay of barley straw appears to produce allelopathic substances

♦ Bacterial activity may also compete with algae for nutrients

♦ Limited success with an “unlicensed herbicide” in USA

Algal Control: In-Lake ManagementViral controls were

attempted without much success in the

1970s, but recent research suggests renewed potential.

Virus SG-3 in tests at

Purdue Univ.

Algal Control: In-Lake ManagementBacterial additives

♦ Many formulations, details proprietary

♦ Seem to focus on limiting N; competitive with algae

♦ Claims of sediment reduction

♦ Often paired with aeration/mixing

♦ Variable results, inadequate scientific documentation

Algal Control: In-Lake ManagementBiomanipulation -

altering fish and zooplankton

communities to reduce algal

biomass

Algal Control: In-Lake ManagementAlewife and other planktivore control - cascading effects

Algal Control: In-Lake Management“Rough” fish

removal -limiting nutrient

regeneration

Algal Control: In-Lake Management

Rooted plant assemblages as algal inhibitors

Algal Control: In-Lake ManagementTechniques that didn’t quite make the list

Algal Control: In-Lake ManagementTechniques that didn’t quite make the list

Algal Control: In-Lake ManagementTechniques that didn’t quite make the list

Algal Control: In-Lake ManagementTechniques for algal control with highest

probability of success♦ Watershed management (where external load is high)

♦ Phosphorus inactivation (for internal load or inflow)

♦ Aeration/mixing (possibly with dyes or bacteria)

♦ Dredging (where feasible, especially for mats)

♦ Algaecides (with proper timing, limited usage)

♦ Sonication (for susceptible algae, ltd nutrient control)

♦ Biomanipulation (but with high variability)

♦ Other techniques as scale and circumstances dictate (do not throw away any tool!)

The End

QUESTIONS?

After that, we might need another one of these…