Phytoplankton Species in Lake Barkley

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Phytoplankton Species in Lake Barkley Author(s): Gary L. Jarrett and Joe M. King Source: Castanea, Vol. 56, No. 2 (Jun., 1991), pp. 90-98 Published by: Southern Appalachian Botanical Society Stable URL: http://www.jstor.org/stable/4033587 Accessed: 05/10/2010 16:08 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=sabs. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected].

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CASTANEA 56(2):90-98. JUNE 1991

Phytoplankton Species in Lake Barkley GARY L. JARRETT and JOE M. KING

ABSTRACT This paper presents a list of 244 taxa identified during a three-year study of the phytoplankton communities in Lake Barkley. Taxa belonged to Cyanophyceae (8.6%), Chlorophyceae (28.7%), Euglenophyceae

(6.1%), Dinophyceae (2.0%), Cryptophyceae

(1.2%), and Bacillariophyceae (53.4%). Additionally, seasonal distributions of phytoplankton standing crops in two embayments and at one main channel site are shown. These data indicate that Cyanophyceae and Bacillariophyceae dominate the phytoplankton in Lake Barkley..

INTRODUCTION Lake Barkley, a mainstream reservoir on the CumberlandRiver in western Kentucky and Tennessee, is the largest in surface area of Cumberland River impoundments. Lake Barkley was formed in 1966 by the U.S. Army Corps of Engineers primarilyfor flood control, hydroelectricpower generation, and navigation in the CumberlandRiver Valley. Additionally, Lake Barkley is used extensively for recreation and commercial fishing, and tourism on this reservoir is vital to the region's economy. Even though Lake Barkley is a major surface water resource in western Kentucky, relatively little is known about the composition of its phytoplankton communities. Past inventories (Craven et al 1983, Hiatt et al 1978, Williams 1972) of these algae are based on collections made during short-term studies of the main channel of this reservoir and identifications were often made to the generic level. Thus, sufficient data is not available for comparison of the phytoplankton communities in Lake Barkley to those in other aquatic ecosystems or for assessing long-term changes that might occur in the composition of these communities. Therefore,a study was undertaken to provide a more detailed inventory of phytoplankton species in Lake Barkley. Samples for identification of algae were collected at least seasonally from 1984 to 1986 from the main channel, major tributaries, and several embayments of this reservoir. Additionally, phytoplankton standing crops were determined monthly for one year in two embayments and at one main channel site. SITE DESCRIPTION The main channel of Lake Barkley extends along a northwest to southeast axis for 189.9 km from Barkley Dam at CumberlandRiver Mile (CRM) 30.6 near Grand Rivers, Kentucky to Cheatham Dam (CRM 148.7) near Ashland City in north-centralTennessee. Lake Barkley receives drainage from a 90

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4,557,900 ha watershed used primarily for silviculture and other types of agriculture (Kentucky Natural Resources and Environmental Protection Cabinet 1984). Besides Cumberland River, Lake Barkley has four major tributaries: Red River, Yellow Creek, Little River and Eddy Creek. Numerous small streams also flow into this reservoir and some of these form relatively large embayments. The major outflow from Lake Barkley is the Cumberland River below Barkley Dam, but water also flows through a canal connecting Lake Barkley to an adjacent reservoir, Kentucky Lake. Lake Barkley is bordered by private lands to the east and by public lands to the west that are part of the Tennessee Valley Authority's Land-Between-the-Lakes National Recreation Area. Lake Barkley is a variable volume reservoir with distinct winter and summer pool elevations. Winter pool elevation is maintained at 108 m above sea level from December to April. The surface area of the reservoir at this time is about 19,140 ha. The water level is allowed to rise to the summer pool elevation of 109.4 m above sea level, which is usually achieved by the first of May. Lake Barkley has a maximum depth of 22.9 m, mean depth of 4.6 m, maximum surface area of 23,490 ha and maximum volume of 1.08 x 1013 cubic meters (Kentucky Natural Resources and Environmental Protection Cabinet 1984). METHODS Eighteen sampling sites in Lake Barkley were used during this study (Figure 1). Seven of these were located along the main channel, four in major

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91

tributary embayments, and seven in minor embayments (Table 1). Embayments and tributary sites were located at least 640 m upstream of the main channel in order to avoid mixing zones between the two environments. Nine sites (5 main channel, 2 embayments, 2 major tributaries) were sampled at about biweekly intervals from May 1984 to August 1986. The other nine sites were sampled on a seasonal basis. Samples were collected at monthly intervals between May 1985 and May 1986 to determine temporal changes in phytoplankton standing crops at Eddy Creek (major tributary embayment), Donaldson Creek (embayment) and Cumberland River Mile 41.5 (main channel). Table 1. Locations of phytoplankton collection sites in Lake Barkley Collection Sites

General Location

CRM 31.5* CRM 41.5 Eddy Creek Mammoth Furnace Creek CRM 58.2 Little River Crooked Creek Donaldson Creek Fords Creek CRM 71.0 Saline Creek Nevells Creek Hickman Creek CRM 100.1 Yellow Creek CRM 124.0 Red River CRM 133.1

Mainstem, near Barkley Dam Mainstem Eastern Embayment Western Embayment Mainstem Eastern Embayment Western Embayment Eastern Embayment Western Embayment Mainstem Eastern Embayment Western Embayment Westernr Embayment Mainstem Western Embayment Mainstem Western Embayment Mainstem, near Cheatham Dam

* CRM = Cumberland River Mile

Phytoplankton samples were collected with a Kemmerer bottle from the photic zone at each site. Aliquots of each sample were placed in 250-ml screw cap plastic bottles and preserved with 4 ml of M3 fixative (American Public Health Association 1985). All algae, except diatoms, were identified through direct microscopic examination of wet mounts. Diatoms were identified from permanent HyraxTM mounts (American Public Health Association 1985). Identifications were made using an Olympus BH microscope with 60X (1.4 numerical aperature) and 10OX oil immersion objectives. Observations were made under both bright field and Nomarski differential interference contrast. Observations of diatoms were also made with a JEOL JSM-T330 scanning electron microscope after specimens were coated with a gold-palladium mixture in an Anatech Ltd. Hummer VI sputter coater. Verification of taxonomic assignments of many of the diatoms was made by comparison to type material 92

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(holotype, isotype and/or lectotype) available at The Academy of Natural Sciences of Philadelphia, and the Great Lakes Research Division of the University of Michigan. Voucher specimens are on deposit in the Aquatic Botany Laboratory, Department of Biological Sciences, Murray State University. Phytoplankton species were enumerated by procedures presented by the American Public Health Association (1985). The relative abundance of diatom species was determined by counting at least 400 valves on HyraxTM prepared slides. These numbers were used along with total absolute cell counts of diatoms to calculate cell numbers of each diatom species. RESULTS AND DISCUSSION Phytoplankton taxa identified in this study are listed alphabetically under the divisions and classes proposed by Prescott (1978). Of the 244 taxa identified, 53.4% (130) are in the Bacillariophyceae, while 28.7% (70) are Chlorophycean algae. Cyanophyceae, Euglenophyceae, Dinophyceae, and Cryptophyceae contributed only 8.6% (21), 6.1% (15), 2.0% (5), and 1.2% (3), respectively, of the total taxa identified. Following genera not identified to species is a number indicating an estimated number of species present. CYANOPHYTA CYANOPHYCEAE Anabaena circinales (Kutz.) Rabh. Anabaena sphaerica Bornet & Flah. Anabaenopsis circularis (West & West) Miller Aphanocapsa pulchra (Kutz.) Rabh. Chroococcus dispersus (Keissl.) Lemm. Chroococcus limneticus Lemm. Chroococcus minor (Kutz.) Nag. Gloeothece linearis Nag. Merismopedia convoluta Breb. Merismopedia elegans S. Braun Merismopedia punctata Meyen Merismopedia tenuissima Lemm. Microcystis aeruginosa Kutz. Microcystis incerta Lemm. Oscillatoria acuminata Gom. Oscillatoria angutissima West & West Oscillatoria geminata Menegh. Oscillatoria lacustris (Kleb.) Geit. Oscillatoria limnetica Lemm. Phormidium tenue (Menegh.) Gom. Raphidopsis curvata Fritsch & Rich

CHLOROPHYTA CHLOROPHYCEAE Actinastrum hantzschii Lag. Ankistrodesmus convolutus Corda Ankistrodesmus falcatus (Corda) Ralfs Ankistrodesmus falcatus (Corda) Ralfs var.

mirabilis (West & West) G.S. West Carteria sp. 1 Chlamydomonas sp. 1 Closteriopsis longissima Lemm. Closterium gracile Breb. JUNE 1991

Closterium setaceum Ehr. Coelastrum cambricum Arch. Coelastrum microporum Nag. Cosmarium circulare Reinsch. Cosmarium subcrenatum Hantz. Cosmarium tinctum Ralfs Cosmarium turpinii Breb. Crucigenia crucifera (Wolle) Collins Crucigenia fenestrata Schmid. Crucigenia irregularis Wille Crucigenia tetrapedia (Kirch.) West & West Crucigenia truncata G.M. Smith Dictyosphaerium ehrenbergianum Nag. Dictyosphaerium pulchellum Wood Euastrum ciastonii Racib. Euastrum turmerii West Eudorina elegans Ehr. Golenkinia paucispina West & West Golenkinia radiata (Chod.) Wille Gonium pectorale Muell. Kirchneriella elongata G.M. Smith Kirchneriella lunaris (Kirch.) Moebius Lagerhemia longiseta (Lemm.) Printz var.

major G.M. Smith Micractinium pusillum Fres. Nephrocytium limneticum G.M. Smith Oocystis crassa Witt. Pandorina morum (Mull.) Borg. Pediastrum biradiatum Meyer Pediastrum boryanum (Turp.) Menegh. Pediastrum duplex Meyen var. clathratum (A.

Braun)Lag. Pediastrum duplex Meyen var. gracillimum Pediastrum obtusum Lucks 93

Pediastrum simplex (Meyen) Lemm. var. simplex Pediastrum simplex (Meyen) Lemm. var. duodenarium (Bailey) Rab. Pediastrum tetras (Ehr.) Ralfs Polyedriopsis spinulosa Schmid. Pteromomas angulosa Lemm. Scenedesmus acuminatus (Lag.) Chod. Scenedesmus arcuatus Lemm. var. platydisca

G.M. Smith Scenedesmus armatus (Chod.) G.M. Smith var. armatus Scenedesmus armatus (Chod.) G.M. Smith var.

Chodat bicaudata(Guglielmetti-Printz) Scenedesmus bijuga (Turp.) Lag. Scenedesmus brasiliensis Bohlin Scenedesmus denticulatus Lag. Scenedesmus dimorphus (Turp.) Kutz. Scenedesmus incrassatulus Bohlin Scenedesmus quadricauda (Turp.) Breb. Schroederia setigera (Shroed.) Lemm. Selenastrum bibraianum Reinsch. Selenastrum convolutus Corda Selenastrum minutum (Nag.) Collins Selenastrum westii G.M. Smith Staurastrum achne West & West Staurastrum cuspidatum Breb. Staurastrum dickii (Ralfs) var. maximum West

& West Staurastrum hexacerum (Ehr.) Witts. Staurastrum leptocladum Nordst. var. insigne

West & West Staurastrum tetracerum Ralfs Tetraedron heteracanthum (Nordst.) Chod. Tetraedon minimum (A. Braun) Hansg. Tetraedon trigonum (Nag.) Hansg. Treubaria setigera (Arch.) G.M. Smith

EUGLENOPHYTA EUGLENOPHYCEAE Euglena acus Ehr. Euglena caudata Hubner Euglena elastica Prescott Euglena intermedia (Klebs) Schmitz Euglena proxima Dang. Phacus longicauda (Ehr.) Dujardin Phacus pleuronectes Muller Trachelomonas abrupta Defl. Trachelomonas ensifera Daday Trachelomonas euchlora (Ehren.) Lemm. Trachelomonas hexangulata Swir. Trachelomonas hispida Perty Trachelomonas lacustris Drezcp. Trachelomonas superba (Swer.) Defl. Trachelomonas volvocina Ehr.

PYRRHOPHYTA DINOPHYCEAE Ceratium hirundinella (O.F. Muell.) Duj.

94

Glenodinium quadridens (Stein) Schil. Hemidinium nasutum Stein Peridinium cinctum (Muell.) Ehr. Peridinium inconspicuum Lemm.

CRYPTOPHYTA CRYPTOPHYCEAE Chroomonas norstedtii Hansg. Cryptomonas erosa Ehr. Cryptomonas ovata Ehr.

CHRYSOPHYTA BACILLARIOPHYCEAE Achnanthes Achnanthes Achnanthes Achnanthes Achnanthes var. Achnanthes

affinis Grunow clevei Grunow exigua Grunow hungarica (Grunow) Grunow lanceolata (de Brebisson) Grunow lanceolata lanceolata (de Brebisson) Grunow

var. dubia (de Brebisson)Grunow Achnanthes linearis (W. Smith) Grunow Achnanthes microcephala (Kutz.) Grunow Achnanthes minutissima Kutz. Amphora submontana Hustedt Anomoeoneis vitrea (Grunow) Ross Asterionella formosa Hassall Caloneis bacillum (Grunow) Cleve Capartogramma crucicula (Grunow ex Cleve)

Ross Cyclostephanos

invisitatus

(Hohn & Heller-

man)Theriot,Stoermerand Hakansson Cyclostephanos tholiformis Stoermer, Hakans-

son and Theriot Cyclotella Cyclotella Cyclotella Cyclotella Cyclotella Cyclotella Cyclotella

atomus Hustedt comensis Grunow comta (Ehr.) Kutz. meneghiniana Kutz. michiginiana Skovrtzow pseudostelligera Hustedt stelligera (Cleve & Grunow) Van

Heurck Cymatopleura linearis W. Smith Cymbella affinis Kutz.

Cymbella minuta (Hilse ex Rabenhorst)var. minuta Cymbella minuta Hilse ex Rabenhorst var. si-

lesiaca (Bleischex Rabenhorst)Reimer Cymbella prostrata (Berk.) Cleve var. prostrata Cymbella triangulum (Ehr.) Cleve Cymbella turgida Greg. CymbeUa turgidula Grunow Diatoma vulgare Bory var. vulgare Fragilaria capucina Desz. Fragilaria crotonensis Kitt. Fragilaria vaucheriae (Kutz.) Petersen Frustulia rhomboides (Ehr.) De Toni var. amphipleuroides (Grunow) Cleve Gomphonema angustatum (Kutz.) Rabh. var. angustatum

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Gomphonema olivacoides Hustedt. var. denestriata Foged Gomphonema parvulum (Kutz.) Kutz. Gyrosigma obtusatum (Sullivant & Wormley)

Boyer

Hustedt

Gyrosigma scalproides (Rabenhorst) Cleve Gyrosigma spencerii (Quekett) Griffith & Henfrey Gyrosigma wormleyi (Sullivant) Boyer Melosira ambigua (Grunow) Mull. Melosira distans (Ehr.) Kutz. var. alpigena

Grunow Melosira granulata (Ehr.) Ralfs var. granulata Melosira granulata (Ehr.) Ralfs var. angutissima Mull. Melosira granulata (Ehr.) Ralfs var. angutissima Mull. forma curvata Mull. Melosira italica (Ehr.) Kutz. Melosira varians C.A. Agardh Meridion circulare (Greville) C.A. Agardh Naviculk aerophila Krasske Navicula anglica Raifs var. subsalsa (Grunow)

Cleve Navicula Navicula Navicula Navicula

arvensis Hustedt atomus (Kutz.) Grunow auriculata Hustedt capitata Ehr. var. hungarica (Grunow)

Ross Navicula Navicula Navicula Navicula

capitoradiata Germain citrus Krasske confervacea (Kutz.) Grunow contenta Grunow var. biceps (Arnott)

Van Heurck Navicula cryptocephaka Kutz. var. cryptocephala Navicula cryptocephala Kutz. var. veneta

(Kutz.) Rabenhorst Navicula Navicula Navicula Navicula Navicula Navicula

graciloides Mayer gysingensis Foged hustedtii Krasske impexa Hustedt ingenua Hustedt menisculus Schumann var. upsalien-

sis (Grunow)Grunow Navicula minuscula Grunow Navicula modica Hustedt Navicula mutica Kutz. var. nivalis

Cleve Nitzschia microcephala Grunow Nitzschia palea (Kutz.) W. Smith var. palea Nitzschia palea (Kutz.) W. Smith var. debilis

(Kutz.)W. Smith Nitzschia palea (Kutz.) W. Smith var. tenuirostris Grunow Nitzschia paleacea Grunow Nitzschia pura Hustedt Nitzschia pusilla Grunow Nitzschia romana Grunow Nitzschia rostellata Hustedt Nitzschia subtubicola Germain Nitzschia tryblionella Hantzsch var. victoriae

Grunow Opephora martyi Heribaud Skeletonema subsalsum (Cleve) Bethge Skeletonema potamos (Weber) Hasle Stephanodiscus alpinus Hustedt Stephanodiscus binderanus (Kutz.) Krieger Stephanodiscus hantzschii Grunow Stephanodiscus hantzschii Grunow forma tenuis

(Hustedt) Hakansson & Stoermer Stephanodiscus Stephanodiscus

niagarae Ehr. niagarae Ehr. var. magnifica minutulus

(Kutz.) Cleve &

Moeller (Ehr.)

(Gregory)Grunow Navicula radiosa Kutz. var. tenella (de Brebis-

son ex Kutz.) Grunow Navicula rhynchocephala Kutz. var. rhyncocephla Navicula rhynchocephala Kutz. var. germainii

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acicularis (Kutz.) W. Smith agnewii Cholnoky amphibia Grunow dissipata (Kutz.) Grunow filiformis (W. Smith) Schutt fonticola Grunow frustulum (Kutz.) Grunow fruticosa Hustedt gracilis Hantzsch hantzschiana Rabenhorst intermedia (Hantzsch ex Grunow)

Fricke

pelliculosa (de Brebisson ex Kutz) Hilse perlucida Hustedt pupula Kutz. var. elliptica Hustedt pupula var. pupula Kutz. pupula Kutz. var. rectangularis

(Wallace)Patrick

Nitzschia Nitzschia Nitzschia Nitzschia Nitzschia Nitzschia Nitzschia Nitzschia Nitzschia Nitzschia Nitzschia

Stephanodiscus

Hustedt Navicula Navicula Navicula Navicula Navicula

Navicula seminulum Grunow Navicula subminuscula Manguin Navicula ventralis Krasske Navicula viridula (Kutz.) Ehr. var. linearis

Stephanodiscus parvus Stoermer & Hakansson Stephanodiscus vestibulus Hakansson, Stoer-

mer & Theriot Synedra acus Kutz. Synedra delicatissima W. Smith var. angutissima Grunow Synedra filiformis Grunow var. exilis Cleve-

Euler Synedra minuscula Grunow Synedra parasitica (W. Smith) Hustedt var. subconstricta (Grunow) Hustedt Synedra pulchelUa Ralfs ex Kutz. Synedra socia Wallace Synedra ulna (Nitz.) Ehr. var. ulna

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Synedra ulna Nitz. var. amphirhynchus (Ehr.)

Surirella ovata W. Smith var. pinnata

Grunow

(W.

Smith) Hustedt

Surirella angusta Kutz. Surirella biseriata de Brebisson

Thalassiosira

weissflogii

(Grunow) Fryxell &

Hasle

Monthly phytoplankton standing crops at Cumberland River Mile 41.5 (CRM 41.5), Donaldson Creek, and Eddy Creek are presented in Figures 2-4. In general, phytoplankton were more abundant in the embayments of Donaldson and Eddy creeks than at CRM 41.5. This variation in spatial abundance cannot be explained on basis of data available in this study. However, it is within reason to assume that flow rates may have been higher in the main channel, thus preventing the accumulation of phytoplankton at CRM 41.5.

Cumberland River Mile 41.5

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Figure 2. Phytoplankton concentration versus month at CumberlandRiver Mile 41.5 (May 1985 - May 1986). Phytoplankton communities at the three sites were dominated in summer and early fall by Anabaena circinales (Kutz.) Rabh., Merismopedia tenuissima Lemm., Oscillatoria acuminata Gom. and Raphidopsis curvata Fritsch. & Rich. Diatoms, especially Melosira ambigua (Grunow) Muell., Stephanodiscus hantzschii Grunow and S. parvus Stoermer & Hakansson, dominated the phytoplankton in winter and early fall. During other months, Cyclotella meneghiniana Kutz., C. pseudostelligera Hustedt, Melosira distans (Ehr.) Kutz. var. alpigena Grunow and Stephanodiscus vestibulus Hakansson, Stoermer & Theriot were the most abundant diatoms. High euglenophycean standing crops at Eddy Creek and CRM 41.5 during June, 1985 resulted from increased cell concentrations of Euglena caudata Hubner. This alga was pres96

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Donafldson Creek 1500

12000 15000

9000

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10000

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2000

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30000

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Figure 3. Phytoplankton concentration versus month in Donaldson Creek embayment (May 1985 - May 1986).

Eddy Creek 30000 27000 24000

3D000 21000O

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c

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Figure 4. Phytoplankton concentration versus month in Eddy Creek embayment (May 1985 - May 1986).

JUNE 1991

97

ent in Donaldson Creekbut its populationdensity remainedlow in comparison to the other two sites. The high numbers of E. caudata Hubner at CRM 41.5 may have resulted from downstream transport of this alga from Eddy Creek rather than its growth at this main channel site. Chlorophyceanstanding crops were relatively low throughout the year. The most prevalent members of this group were Ankistrodesmus falcatus Nag., Scenedesmus ehrenbergianum (Corda) Ralfs, Dictyosphaerium acuminatus (Lag.) Chod., S. bijuga (Turp.) Lag. and S. quadricauda (Turp.)

Breb. Cryptophyceae and Dinophyceae were minor constituents of phytoplankton assemblages in Lake Barkley. This paper provides the most comprehensivecompilation of phytoplankton species in Lake Barkley and, it is hoped, will serve as baseline data for future studies of the ecosystem. Variations shown in the standing crops of these algae indicate a need to further examine their distribution in relation to physicochemical and hydrologicparametersin this. surface water resource. ACKNOWLEDGMENTS This research was supported by a contract from the Nashville District of the U.S. Army Corps of Engineers. We thank Ms. Jean Lynch for her assistance with preparationof this manuscript. Use of the herbariumat the Academy of Natural Sciences of Philadelphiaby the senior author was supported by the Jessup-McHenryAward. Dr. Eugene Stoermerhas our appreciationfor use of the diatom herbarium,University of Michigan. LITERATURE CITED AMERICAN PUBLIC HEALTH AssoCIATION.1985. Standard methods for the

examination

of water and wastewater, 16th ed. American Public Health Association, Washington, D.C. 1,268 p. CRAVEN, T.M., D.L. DYCUS,and D.C. WADE. 1983. Responses of selected aquatic biota to thermal discharges from Cumberland Steam-Electric Plant: Barkley Reservoir, Tennessee, 1978 and 1979. TVA Office of Natural Resources, Muscle Shoals, Alabama. 116 p. HIATT, F.A., S.C. HERN, J.W. HILGERT, V.W. LAMBOU,F.A. MORRIS, M.K. MORRIS, L.R. WILLIAMS, and W.D. TAYLOR. 1978. Distribution of phytoplankton in Tennessee Lakes. Working Paper No. 691. Natl. Eutrophication Surv., U.S. EPA, Environmental Monitoring and Support Laboratory, Las Vegas, Nevada, Rep. No. EPA600/3-3-78-016. 76 p. KENTUCKY NATURAL RESOURCES AND ENVIRONMENTAL PROTECTION CABINET.

1984.

Trophic state and restoration assessments of Kentucky Lake. Kentucky Division of Water, Frankfort, Kentucky. 476 p. PRESCOTT, G.W. 1978. How to know the freshwater algae. W.C. Brown Co., Dubuque, Iowa. 293 p. WILLIAMS, L.G. 1972. Plankton diatom species biomasses and the quality of American rivers and the Great Lakes. Ecology 53:1038-1050.

G.L.J.: J.M.K.: WATER MANAGEMENT SECTION DEPT. OF BIOLOGICAL SCIENCE U.S. ARMY CORPS OF ENGINEERS MURRAY STATE UNIVERSITY LOUISVILLE, KENTUCKY40201-0059 MURRAY, KENTUCKY 42071 Received January 8, 1990; Accepted September 8, 1990.

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