United States
Environmental Protection
Agency
Region 5,
Great Lakes National     EPA-905/3-88-001
Program Office        GLNPO Report No. 3
230 South Dearborn Street  February 1988
Chicago, Illinois 60604
Phytoplankton  and
Zooplankton in  Lakes
Erie, Huron,  and
Michigan:  1984
 Do not WEED. This document
 should be retained in the EPA
 Region 5 Library Collection.


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                                             EPA-905/3-88-001
                                             GLNPO Report Mb.3
                                             February 1988
          Phytoplankton and Zooplankton
In Lakes Erne, Lake Huron and Lake Michigan:  1984
         Volume 1 - Interpretive Report
                        by
               Joseph C.  Makarewicz
        Department of Biological Sciences
      State University of New York at Brockport
            Brockport,  New York  14420

                   March 1987
                Project Officer
                   Paul Bertram
       U.S. Environmental Protection Agency
       Great Lakes National Program Office
            230 South Dearborn Street
                Chicago,  Illinois 60604
        U.S. Environmental Protection Afencf
        Region 5, library (PI-12J)
        77 West Jackson Boulevard, 12th Floor
        Chicago, II  60604-3590

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                                 Abstract

     With  the acknowledgement that biological monitoring was  fundamental
to charting ecosystem health (Great  Lakes Water Quality Agreement  1978),
EPA's program was  developed for Lakes  Erie, Huron and  Michigan to:   1)
monitor seasonal patterns,  ranges of abundance and, in general,  structure
of the phytoplankton and zooptankton communities; 2) relate the biological
components  to  variations  in  the  physical,  nutrient  and   biological
environment; and 3) assess the  annual variance to allow better  long-term
assessments of trophic  structure and  state.   Several offshore  stations
(9-11) on several cruises (9-11) during  the spring, summer and autumn  of
1984 and winter of 1985 were sampled.
     By examining changes in the phytoplankton and zooplankton in relation
to water chemistry,  evidence was  found suggesting little  change in  the
trophic status of  Lakes Huron and  Michigan while an  improvement in  the
trophic status of  Lake Erie  was evident.   The offshore  region of  Lake
Michigan  is  experiencing  changes   in  phytoplankton  and   zooplankton
composition consistent with nutrient control and top-down control by fish.
Even so,  the biomass  of phytoplankton  and zooplankton  and the  trophic
status of the  lake have not  changed significantly.   The appearance  and
establishment of  Daphnia  pul icari a  in offshore  waters  of  Lake  Huron
suggest a change   in the  forage fish  base.   With the  exception of  the
resurgence of Asterjonella formosa in Lake Erie, plankton composition  has
changed little since the 60's.  However, dramatic reductions in biomass of
nuisance and eutrophic indicator species have occurred.  These changes are
consistent with expectations  of long-term nutrient  control.  However,  a
change  in  piscivory   is  evident  that   has  apparently  allowed   the
establishment of the large cladoceran Daphnia pulicaria.
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                                DISCLAIMER
This report has been reviewed by the Great Lakes National  Program  Office,
U.S.  Environmental  Protection  Agency,   and  approved  for  publication.
Approval does not signify that the contents necessarily reflect the  views
and policies of the U.S. Environmental  Protection Agency,  nor does mention
of  trade  names   or  commercial  products   constitute  endorsement   or
recommendation for use.

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                                                                      IV
                                 FOREWARD

The Great  Lakes National   Program  Office (GLNPO)   of the  United  States
Environmental Protection Agency was established  in Region V,  Chicago,   to
focus  attention  on   the  significant  and   complex  natural    resource
represented by the Great Lakes.

GLNPO implements a multi-media environmental  management program drawing on
a wide  range of  expertise represented  by universities,   private  firms,
State, Federal and  Canadian Governmental  Agencies   and the  International
Joint Commission.  The goal of  the GLNPO program is to develop  programs,
practices and technology necessary for a better understanding  of the Great
Lakes  ecosystem  and  to  eliminate  or  reduce to  the   maximum  extent
practicable the discharge of pollutants into the Great Lakes system.    The
Office also coordinates  U.S. actions  in fulfillment of  the  Great  Lakes
Water Quality Agreement of  1978 between Canada and  the United States   of
America.

This report presents results of the phytoplankton and zooplankton portions
of the  water quality  surveillance program  conducted by   GLNPO on  Lakes
Michigan, Huron and Erie in  1984 and in winter of   1985.   Results of  the
physical and chemical portions of the surveillance  program may be found in
a companion report:

          Lesht, Barry M.  and David C. Rockwell.  1987. The State
          of the Middle Great Lakes:  Results of the 1984  Water
          pual ity Survey of Lakes Erie,  Huron and Michigan.
          Publication Number ANL/ER-87-1.   Argonne  National
          Laboratory, Argonne, Illinois   60439.

GLNPO grateful Iy  acknowledges  the  contribution to  this  study  of  the
Bionetics Corporation, with  whom GLNPO contracted   for assistance in  the
collection of samples and  for the  identification  and enumeration of  the
phytoplankton and zooplankton.  In  particular, we  extend  appreciation   to
Norman A. Andresen, Mark A. Lamb,  Louis L. Lipsey,  Donna Page  and  Heather
K. TrulI I.

Funds for this report  were provided by  U.S.E.P.A., Great Lakes  National
Program Office under Grant Number R005772-01.

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                            TABLE OF CONTENTS

                                                                 Ease.
Abstract	I i
Disclaimer	..	 i i i
Foreward	iv
Tab! e of Contents	 v
L i st of Tab! es.	"	v i i I
L ist of Figures	 .xi i
Acknow I edgments	xv i I

OVERV IEW	1

SUMMARY
   Lake Mich igan	1
   Lake Huron	5
   Lake Erie	8

INTRODUCTION	11

METHODS
   Samp I ing Si tes	14
   Chem i stry	14
   Phytop I ankton	..14
   Zoop I ankton.	16
   Data Organization.	17

RESULTS AND DISCUSSION - LAKE MICHIGAN
   Phytop I ankton	19
      Annual  Abundance of Major Algal  Groups	....19

      Seasonal  Abundance and Distribution of Major Algal  Groups..19
      Regional  and Seasonal Trends In the Abundance of
         Common Taxa	21
      Vert ica I  D i str i but ion	27
      W i nter Cru i se	28
      Historical  Changes in Species Composition	29
      P i cop I ankton	33
      Geographical  Abundance and Distribution	..33
      Indicator Species...	36
      Historical  Changes in Community Abundance	37

   Zoop I ankton	39
      Annual  Abundance of Zoopl ankton Groups	..39
      Seasonal  Abundance and Distribution of Major
        Zoop I ankton Groups	39
      Geographical  Abundance and Distribution of
        Zoopl ankton Groups	40
      Common  Species.	 .40
      Changes in Species Composition	41
        Crustacea	41
        Rot i f er a	45
      Historical  Changes in Zoop I ankton Biomass	46
      Indicators of Trophic Status	..46

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                                                                  VI
                                                                 Page
      Troph ic Interactions	49
RESULTS AND DISCUSSION - LAKE HURON
   Phytop I ankton	52
      Annual  Abundance of Major Algal  Groups	52

      Seasonal  Abundance and Distribution of Major
         Algal  Groups	52
      Geographical  Abundance and Distribution of Major
         AI gal  Groups	53
      Regional  and Seasonal  Trends in the Abundance of
         Common Taxa	54
      Vertical  Distribution	...,58
      W i nter Cru i ses	59
      Historical Changes in Species Composition	60
      P i cop I ankton	61
      Indicator Species	61
      Historical Changes in Community Abundance and Biomass	63

   Zoopl ankton	65
      Annual  Abundance of Zoop I ankton Groups	65
      Seasonal  Abundance and Distribution of Major
         Zoop I ankton Groups	65
      Common Species	66
      Changes in Species Composition	66
         Crustacea	66
         Rot i f era	69
      Geographical  Abundance and Distribution of
         Zoop I ankton Groups	70
      I nd i cators of Troph i c Status	71
      Historical Trends  in Abundance	73
      Troph ic I nteract ions	74

RESULTS AND DISCUSSION - LAKE ERIE
   Phytopl ankton	76
      Annual  Abundance of Major Algal Groups	76
      Seasonal  Abundance and Distribution of Major
         AI gal  Groups	77
      Geographical  Abundance on Distribution Major
         of Major Algal Groups..	77
      P i cop I ankton	78
      Regional  and Seasonal Trends in the Abundance
         of Common Spec i es	79
      Changes in Species Composition	82
      Indicator Species	84
      Historical Changes  in Community Biomass	85

   Zoopl ankton.	87
      Annual  Abundance of Zoopl ankton Groups	87
      Seasonal  Abundance and Distribution of Major
         Zoopi ankton Groups	87
      Geographical  Abundance and Distribution of

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                                                                   vn

                                                                 Page
         Zoop I ankton Groups	.88
      Common Species	 .89
      Changes in Species Composition	89
         Crustacea.	89
         Rot i f era	92
      East-West Species Distribution	93
      Indicators of  Trophic Status	..93
      Historical Changes in Abundance	...95
      Trophic Interactions	97
L ITERATURE CITED	102

TABLES	113

F IGURES	168

APPENDICES
  Species List - Phytoplankton
     Table A.  Lake Michigan	238
     Table B.  Lake Huron	248
     Table C.  Lake Erie	257
  Species List - ZoopIankton
     Table D.  Lake Michigan..	267
     Table E.  Lake Huron	270
     Table F.  Lake Erie	273

  VOLUME 2. DATA SUMMARY REPORT.   Summary sheets of phytopiankton
     and zoopI ankton data	ATTACHED MICROFICHE

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                                                                  vm
                             TABLE LEGENDS
                                                                 Page
TABLE 1       Plankton sampling dates for Lakes Michigan,
             Huron and Erie in 1984 and 1985	113

TABLE 2      Latitude and longitude of plankton sampling
             stat ions, 1984	114

TABLE 3      Sample dates and stations for Lake Michigan,
             1984 and 1985	116

TABLE 4      Sample dates and stations for Lake Huron,
             1984 and 1985	117

TABLE 5      Comparison of calculated crustacean dry weights
             to measured dry weights in Lake Michigan	118

TABLE 6      Mean values of physical-chemical  parameters
             (ApriI-October) from a 1-m depth  for Lakes Erie,
             Michigan and Huron,  1984	119

TABLE 7      Number of species and genera observed in each
             algal division or grouping in Lake Michigan,
             1983 and 1984	120

TABLE 8      Relative abundance of major phytoplankton
             divisions in Lake Michigan, 1983  and 1984	121

TABLE  9     Abundance of Rh i zosoI en i a eriensis in Lake
             Michigan in 1983 and 1984	122

TABLE 10     Summary of common phytoplankton species occurrence
             in Lake Michigan, 1984	123

TABLE 11     Common species observed in either 1983 or 1984
             but not both years,  Lake Michigan	124

TABLE 12     Number of species in Lake Michigan with depth
             at Station 47, 15 August 1986	125

TABLE 13     Comparison of abundance of Cyclotella species
             at offshore sites in August of 1970,  1983 and
             1984, Lake Michigan	126

TABLE 14     Comparison of nutrient levels between Stations
             6, 64,  77 and all other stations  during the
             spring and fall, Lake Michigan	127

TABLE 15     Distribution of  indicator diatom species in  Lake
             Michigan	128

TABLE 16     Relative abundance of zoopIankton  In Lake Michigan..129

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                                                                   IX

                                                                 Page
TABLE 17     Summary of common zooplankton species occurrence
             in Lake Michigan during 1984	130

TABLE 18     Cladoceran abundance in 1954, 1966,  1968,  1983
             and 1984 in Lake Michigan	131

TABLE 19     Copepoda abundance in 1954, 1966,  1968,  1983
             and 1984 in Lake Michigan.....	133

TABLE 20     Average crustacean zooplankton biomass for 1976
             and 1984, Lake Michigan	135

TABLE 21     The ratio of calanoids to cyclopoids plus
             cladocerans geographically  in Lake Michigan,
             1983 and 1984	136

TABLE 22     Correlation of phytoplankton with total  phosphorus
             concentrations and zooplankton abundance
             within  individual cruises  in Lake Michigan	137

TABLE 23     Number of species and genera observed in each
             algal  division or grouping, Lake Huron,  1983
             and 1984	138

TABLE 24     Relative abundance of major phytoplankton
             divisions in Lake Huron, 1983 and 1984	139

TABLE 25     Abundance of Rhizosolenia eriensis in Lake
             Huron,  1983 and 1984	140

TABLE 26     Summary of common phytoplankton species occurrence
             in Lake Huron during 1984 and winter of 1985	141

TABLE 27     Common  species observed  in either 1983 or 1984
             but not  in both years,  Lake Huron	142

TABLE 28     Distribution of  indicator diatom species  in
             Lake Huron	143

TABLE 29     Relative abundance of zooplankton in Lake Huron	144

TABLE 30     Summary of common zooplankton species occurrence
             in Lake Huron during 1984	145

TABLE 31     Comparison of mean crustacean abundance for the
             sampling period  in 1971  (Apr!I-November),
             1974/75  (April-November),  1983 (August-October)
             and 1984 (April-December), Lake Huron	146

TABLE 32     Abundance of selected zooplankton species in
             northern and southern Lake  Huron in 1984	147

TABLE 33     Ratio of Calanoida to Cladocera plus Cyclopoida
             in Lake Huron,  1983 and  1984	148

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                                                                  X

                                                                 Page
TABLE 34     Comparison of the plankton ratio (Calanoida/
             Cyclopoida+Cladocera) between the northern
             stations of Lake Huron and Lake Michigan	.149

TABLE 35     Mean abundance of rotifers in Lake Huron in
             1974 and 1983	150

TABLE 36     Correlation of phytoplankton abundance with
             total phosphorus concentrations and zoopIankton
             abundance within individual  cruises,  Lake
             Huron	151

TABLE 37     Number of species and genera observed in each
             algal division or grouping,  Lake Erie,  1983
             and 1984	152

TABLE 38     Number of species identified and percentage of
             species belonging to various taxonomic groups,
             Lake Erie	153

TABLE 39     Annual phytoplankton biomass for the entire lake
             and the western, central  and eastern basins of
             Lake Erie, 1983 and 1984	154

TABLE 40     Summary of common phytoplankton species occurrence
             in Lake Erie, 1984 and winter of 1985	155

TABLE 41     Location of maximum abundance of selected species
             in 1983 and 1984, Lake Erie	156

TABLE 42     Common species observed in either 1983 or 1984
             but not both years,  Lake  Erie.	157

TABLE 43     Importance of Asterionel I a iooacisa during the
             spring of 1984,  Lake Erie	158

TABLE 44     Mean maximum biomass of selected common phyto-
             plankton species in 1970  and 1983,  Lake Erie	159

TABLE 45     Distribution of indicator diatom species in the
             western basin of Lake Erie.....	160

TABLE 46     Trophic status of the western,  central  and eastern
             basins of Lake Erie in 1970 and 1983/84	161

TABLE 47     Relative abundance of zooplankton in Lake Erie	162

TABLE 48     Summary of common zooplankton species occurrence
             in Lake Erie during 1984	.163

TABLE 49     Occurrence of eutrophic zooplankton indicator
             species in Lake Erie, 1984	164

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                                                                  XI


                                                                 Page
TABLE 50     Ratio of calanoids to cladocerans plus cyclopoids
             in Lake Erie,  1983 and 1984	165

TABLE 51     Turbidity levels in 1978 and 1984,  Lake Erie	166

Table 52     Correlation of phytoplankton abundance with total
             phosphorus concentration and zoopIankton abundance
             within  individual  cruises,  Lake Erie	167

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                                                                   xi i


                              FIGURE LEGENDS

                                                                  Page
FIGURE 1.  Lake Michigan plankton sampling stations,  1984-85	168

FIGURE 2.  Lake Huron plankton sampling stations,  1984-85	169

FIGURE 3.  Lake Erie plankton sampling stations, 1984-85	170

FIGURE 4.  Seasonal phytoplankton abundance and biovolume
           trends  in Lake Michigan,  1984-85	171

FIGURE 5.  Seasonal distribution of  algal  divisions in Lake
           Michigan, 1984-85	172

FIGURE 6.  Mean seasonal distribution of Cyclotella
           ocelIataf Synedra ulna v.  chaseana,
           Synedra f iI iformis and Rhizosolenia
           long! set a,  Lake Michigan	173

FIGURE 7.  Mean seasonal distribution of Nitzsch ia
           Iauenburgiana, Qpcystis submarina,
           Dictyosphaerium ehrenbergianum and
           Cryptomonas rostratiformisf Lake Michigan	174

FIGURE 8.  Mean seasonal distribution of Qsc iI  Iator i a
           minimaf Lake Michigan	175

FIGURE 9.  Seasonal and geographical distribution of
           Cyclotella pee I|ata,  Oocystis submarina,
           Dictyosphaerium ehrenberg ianum and
           Osci I I ator i a min ima,  Lake Mi ch igan	176

FIGURE 10.  Vertical distribution of phytoplankton at Station
            47, 15 August  1984,  Lake Michigan	177

FIGURE 11.  Vertical distribution of phytoplankton at Station
            18, 15 August  1984,  Lake Michigan	178

FIGURE 12.  Annual geographical  distribution of major algal
            divisions in Lake Mich igan, 1984-85	179

FIGURE 13.  Geographical distribution of phytoplankton abundance
            on all cruises, Lake Mich igan, 1984-85	180

FIGURE 14.  Historical  abundance of  phytoplankton  in Lake
            Michigan	181

FIGURE 15.  Seasonal zooplankton abundance in Lake Michigan,
            1984	182

FIGURE 16.  Seasonal fluctuation (numerical) of zooplankton
            groups in Lake Michigan, 1984.......	183

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                                                                  xm

                                                                  Page
FIGURE 17.  Seasonal  fluctuation (biomass) of zooplankton
            groups in Lake Michigan,  1984	184

FIGURE 18.  Geographical  distribution (numerical) of major
            zooplankton groups in Lake Michigan, 1984	..185

FIGURE 19.  Geographical  distribution (biomass) of major
            zooplankton groups in Lake Michigan, 1984	186

FIGURE 20.  Geographical  distribution of Diaptomus sici1 is
            in Lake Michigan, 1984	187

FIGURE 21.  Geographical  distribution of selected zooplankton
            in Lake Michigan, 1984	188

FIGURE 22.  Geographical  distributin of selected zooplankton
            in Lake Michigan, 1984	189

FIGURE 23.  Historical trends in zooplankton biomass during July
            and August, Lake Mich igan	 190

FIGURE 24.  Seasonal  phytoplankton abundance and biovolume
            trends in Lake Huron, 1984-1985	191

FIGURE 25.  Seasonal  distribution of algal (% biovolume)
            divisions  in Lake Huron,  1984-85	192

FIGURE 26.  Annual geographical  distribution of major algal
            division  in Lake Huron,  1983	193

FIGURE 27.  Annual geographical  distribution of major algal
            divisions  in Lake Huron,  1984-85	194

FIGURE 28.  Geographical  distribution of phytoplankton
            abundance on all cruises, Lake Huron, 1984-85	195

FIGURE 29.  Mean seasonal distribution of Cyclotella
            stelI  igeraf Stephanod iscus alp inus,
            Stephanod iscus minutus and Cosmarium sp., Lake
            Huron	196

FIGURE 30.  Mean seasonal distribution of Osc 1.1.1 atari a
            minima. Lake Huron	197

FIGURE 31.  Vertical  distribution of phytoplankton at
            Station 37, 15 August 1984, Lake Huron	198

FIGURE 32.  Vertical  distribution of phytoplankton at
            Station 15, 15 August 1984, Lake Huron	199

FIGURE 33.  Historical offshore biomass trends  in Lake Huron	200

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                                                                      x;v

                                                                  Page
FIGURE 34.  Seasonal  zooplankton biomass and abundance
            in Lake Huron,  1984	201

FIGURE 35.  Seasonal  fluctuation (numerical) of zooplankton
            groups in Lake Huron,  1984	202

FIGURE 36.  Seasonal  fluctuation (biomass) of zooplankton
            groups in Lake Huron,  1984	203

FIGURE 37.  Geographical distribution of Daphnl.a pul icaria
            in 1983 and 1984,  Lake Huron	204

FIGURE 38.  Geographical distribution (numerical) of  major
            zooplankton groups in Lake Huron, 1984	205

FIGURE 39.  Geographical distribution (biomass) of major
            zooplankton groups in Lake Huron, 1984	206

FIGURE 40.  Geographical distribution of major zooplankton
            groups in Lake Huron,  1983	207

FIGURE 41.  Geographical distribution of selected Rotifera
            and Copepoda in Lake Huron,  1984	208

FIGURE 42.  Water chemistry along the  north-south axis of
            Lake Huron, 1984	209

FIGURE 43.  Crustacean abundance of Lake Huron, 1970-1984	210

FIGURE 44.  Abundance of Rotifera in Lake Huron in 1974,
            1983 and 1984	211

FIGURE 45.  Seasonal  phytoplankton abundance and biovolume
            trends in Lake Erie, 1984-85	212

FIGURE 46.  Seasonal  distribution of algal divisions in Lake
            Erie, 1984-85	213

FIGURE 47.  Annual geographical distribution of major algal
            divisions  in Lake Erie,  1984-85..	214

FIGURE 48.  Geographical distribution of phytoplankton
            abundance on all cruises, Lake Erie, 1984-85	215

FIGURE 49.  Geographical distribution of selected species,
            Lake Erie, 1984-85	216

FIGURE 50.  Geographical distribution of selected species,
            Lake Erie, 1984-85	217

FIGURE 51.  Mean seasonal distribution of Asterionella
            formosa,  Me Ios i ra isl andicaf Anabaena
            sp. and Crucigenia rectangul aris, Lake Erie	218

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                                                                  XV

                                                                  Page
FIGURE 52.   Seasonal  fluctuation of weighted mean phytoplankton
            biomass in  1970,  1983 and 1984,  Lake Erie	219

FIGURE 53.   Regression of phytoplankton biomass versus time
            in western Lake Erie	220

FIGURE 54.   Time trend in annual cruise mean concentration of
            corrected chlorophyll  _a since 1970,  Lake Erie........221

FIGURE 55.   Time trend in annual cruise average of total  phosphorus
            since 1970,  Lake Erie	222

FIGURE 56.   Seasonal  zoopIankton abundance and biomass in
            Lake Erie, 1984..	223

FIGURE 57.   Seasonal  abundance distribution of zoopIankton
            groups in Lake Erie, 1984	224

FIGURE 58.   Seasonal  biomass distribution of zoopIankton
            groups in Lake Erie, 1984	225

FIGURE 59.   Geographical  distribution (numerical) of
            zoop I ankton groups in Lake Erie,  1984	226

FIGURE 60.   Geographical  distribution (biomass)  of
            zoopl ankton groups,  1984,  Lake Erie...	227

FIGURE 61.   Geographical  distribution of selected Crustacea
            in Lake Erie, 1984	228

FIGURE 62.   Geographical  distribution of selected Rotifera
            in Lake Erie, 1984	229

FIGURE 63.   Crustacean zoopIankton abundance since 1939 in
            the western basin of Lake Erie	230

FIGURE 64.   July and August abundance of Cladocera and
            Copepoda in the western basin of Lake Erie
            s i nee 1939	231

FIGURE 65.   Seasonal  fluctuation of weighted mean Crustacea
            (nauplii  excluded) abundance in 1970, 1983,  1984,
            Lake Erie	232

FIGURE 66.   Seasonal  fluctuation of Rotifera in the western
            basin of Lake Erie from 1939 - 1983	233

FIGURE 67.   Abundance of fishable walleye in western Lake Erie
            (Ohio waters)	234

FIGURE 68.   Sport angler harvest of walleye from the central
            basin of Lake Erie	235

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                                                                  XVI

                                                                  Page
FIGURE 69.   Time trend of emerald and  spottail   shiner  abundance
            in the central  basin and alewife from the western
            basin of Lake Erie	236

FIGURE 70.   Seasonal and geographical  turbidity  trends  in  Lake
            Erie	237

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                                                                     xvn
                             ACKNOWLEDGEMEOTS

   Ted  Lewis provided the  technical  expertise on  the INFO database  and
other software.   Diane Oleson inputed the raw  data into the computer.   P.
Bertram,  D.  Rockwell,  G.   Fahnenstiel   and  other  anonymous  reviewers
provided constructive comments that improved the manuscript.  I  thank  them
for their time and effort.

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                                 OVERVIEW

    With the acknowledgement that biological  monitoring was fundamental  to
charting ecosystem  health (Great  Lakes  Water Qua I Ity  Agreement  1978),
EPA's program was  developed for Lakes  Erie,  Huron and  Michigan to:    1)
monitor seasonal  patterns,  ranges  of abundance  and  structure  of   the
phytoplankton  and  zooplankton  communities;   2)  relate  the  biological
components  to  variations  in  the  physical,   nutrient  and   biological
environment; and 3) assess the  annual variance to allow better  long-term
assessments of trophic structure and state.
      The program  has proven  successful.   By examining  changes in   the
phytoplankton and zooplankton in relation to water chemistry,  evidence was
found suggesting little change  in the trophic status  of Lakes Huron   and
Michigan while  an improvement  in the  trophic status  of Lake  Erie   was
evident within the past ten years.   The offshore region of Lake  Michigan
is experiencing  changes   in  phytoplankton  and  zooplankton  composition
consistent with nutrient control and top-down  control by fish.   Even   so,
the biomass of phytoplankton and zooplankton and the trophic status of the
lake have not changed significantly.  The appearance and establishment  of
Daphnia pulIcaria In offshore waters of Lake Huron suggest a change In the
forage fish base.   With the exception of  the resurgence of  Asterlone I la
formosa in Lake Erie,  plankton  composition has changed little since   the
60fs.  However, dramatic reductions  in biomass of nuisance and  eutrophic
indicator species  have  occurred.   These  changes  are  consistent  with
expectations  of  long-term  nutrient  control.    However,  a  change  in
piscivory  is evident that has apparently allowed the establishment of   the
large cladoceran Daphnia pulicaria.
    The following summaries for Lakes Michigan, Huron and Erie outline the
major observations of the 1984  intensive sampling of the offshore  region.
As such, the 1983 (Makarewicz 1987)  and 1984 studies provide a basis   for
long-term monitoring of the structure and functioning of the Great Lakes.

                                 SUMMARY
                               Lake Michigan

1.   In 1984,  327 algal and 52 zooplankton species were observed.  Compared
to 1983, a  15? and 24%  reduction in the number of algal and  zooplankton
species were observed.   As the same  samp I ing, enumeration procedure   and
taxonomists were employed, the observed flucuatlons In species composition
are  due  to  both  natural  and  sampling  variability  of  the  plankton
population.

2.  Compared to  Lake Huron,  variability in  common algal species in  Lake
Michigan in 1983 and 1984 was high.  76? of the common species observed in
1984 were also common species In 1983.  31? of  the common species observed
in 1983 were not common in 1984.

3.   Average phytopIankton  and  zooplankton abundances  were  22,220+1400
cells/mL and 59,764+8,284 organlsms/m  for  the study  period.   Mean algal
and zooplankton biomass  were 0.55+.038  g/m  and 33.2+4.9  mg/m  for   the
study period.

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4.  As in Lakes Erfe  and Huron, diatoms possessed the greatest  diversity
of species  (166)  and  biomass  (70.0$  of  the  total)  in  1984.     The
Cryptophyta accounted for the second highest biomass in 1984.

5.  Picopiankton represented 82.9$ of the total  abundance but  only 1.4$ of
the algal biomass.

6.  Diatoms were dominant throughout  the study  period,  accounting for  as
much as 80$ but  never less than  55$ of the  phytopIankton biomass.    The
overwhelming dominance  of the  diatoms in  1984 precluded  the  prominent
seasonal  succession of aigal divisions observed  in 1983.

7.  The  large drop  in diatom bfomass observed in  August of 1983 was  not
observed in  1984,   A bloom  of Rhizosolenfa erlensis during  1984,  not
observed In  1983, was  the major  cause of  the dominance  of diatoms  in
August of 1984.  A similar situation was observed in Lake Huron in 1984.

8.  Abundance of phytopIankton decreased from the most northern station to
Station 57 and remained the same southward to the most southerly  station,
where it increased slightly.

9.  Vertical distribution studies indicated that an increase In  abundance
occurred and a 100$+ Increase In species diversity occurred with depth  at
Station 47.  The  Increase in  abundance and diversity  correlated with  the
decrease In temperature associated with the metalImnion.

10.  Winter samples  were analyzed in 1985.   Algal  biomass and  abundance
were low during the winter but  were not significantly different from  the
autumn and spring values.   Diatoms and  cryptophytes  were predominant  as
during the non-winter  period.   However,  the relative  importance of  the
Cryptophyta increasd by a factor of >2 (11.6 to  25.3$).

11.  The  phytoplankton composition  of Lake  Michigan has  changed.    The
following subdominant or dominant species have decreased In abundance from
the 60's  and  70's:    Cyciotel I a  m!ch iganlana,   Cy dote I I a   stel I igera,
MeIosIra   is!apd ?caf    Synedra   acus   and  Ank ? strodesmus     falcatus.
QscII Iatoria  Iimnetlca   has  Increased   In abundance.    Abundance   of
Rhizosolenfa eriensts increased in 1984 after a  general  decrease since the
60's and 70«s.

12.  Dominant  diatom species  included the  mesotrophic forms  TabelI aria
flocculosa  and  Frag!I aria   crotonensis  and   the  oligotrophic   forms
Cyciotella ocelI ata  and  Rhizosoienia erlensis.     Compared to  the  1983
cruises where  mesotrophic forms  were predominant,   the same   mesotrophic
forms were present in 1984 along with the oligotrophic Indicators.

13.  The ratio of mesotrophic  to eutrophic algal  species (trophic  ratio)
suggests a  eutrophic  status for  nearshore  waters In  1977,   while  the
offshore  waters   In   1970-71,  1983   and   1984 would  be   in   the
oIigotrophlc-mesotrophic range.

14.  Based  on the classification  scheme of Munawar  and Munawar  (19824,
Lake Michigan's algal  biomass in 1983  (0.42 mg/m )  and 1984 (0.55  mg/m )
suggests an oligotrophic status for the offshore waters of Lake Michigan.

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15.   Phytop Iankton  abundance of  the  offshore waters  appears  to  have
increased from 1962-63 to 1976-77  but has not significantly changed  from
1976 to 1984.   Because of  the difference In enumeration methodology  used
In the  1962-63  study compared  with  the other  surveys,   the  suggested
increase in algal abundance from 1962-63 to 1976-77 has to be  interpreted
cautiously,

16.  The  trend  in zooplankton  biomass was similar  to the  phytoplankton
trend between 1976 and 1984 in  that no significant change in  zooplankton
biomass was observed.

17.   The  Rotifera possessed  the  largest  number of  species  (29)  and
relative abundance (67.5$).   The  Rotifera contributed only  2.6% of  the
biomass, while  the  Ciadocera  accounted for  39.8$  of  the  zooplankton
biomass.

18.  Abundance  of zooplankton  generally Increased from  north to  south.
The far northern stations (64 and 77) had a significantly higher abundance
than the  rest of  the lake.   The  northern Stations  64 and  77 and  the
southern Stations 5 and 6 are best described as nearshore stations.

19.  Both the  1983 and 1984 dominant  rotifer composition was similar  to
the nearshore and to Ahlstrom's (1936) offshore composition.

20.   The species  composition  of the  predominant rotifers  suggests  an
ollgotrophic offshore assemblage.  Further support is provided by the high
relative abundance of Diaptomus ^icii Ls and L imnocalanus macrurus and  the
occurrence of  Senecella calanoides, all ol Igotrophic crustacean  indicator
species.

21.   The  plankton  ratio (Calanoida/Cladocera  +  Cyclopoida)  was  high
relative to Lake  Erie but  lower than  Lake Huron.   Except  for the  far
northern and southern extremes of the lake,  the ratio was high and similar
indicating a similar high quality of water.   At the far northern stations,
abundance of the olIgotrophic L imnocalanus macrurus and Diaptomus  siciI is
was lower,   while  E"bosmina  coregonI  and  Bosmlna  Iong1rostrisf  often
associated with eutrophic conditions, increased.  In addition, four diatom
species Indicative  of  mesotrophic  conditions were  more  abundant,  and
phytoplankton  abundance in general  was  higher at these northern  stations
suggesting a lower water quality for the northern region.  At Station  77,
si I lea and total  phosphorus were higher than in the rest of  the lake.

22.  The changing nature of the zooplankton community of Lake Michigan was
evident in 1984.    The abundance of Daphnia .-pul icaria,  first observed  in
1978,  dropped  from  376/m  in  1983 to  78/m  in  1984.    Abundance of Q.
gal eata, rare  In 1966  and 1968,  was three  times the density observed  In
1954 (1200/m ).     In  general,   the  larger  cladocerans,   calanoids  and
cyclopoid copepods,  observed to  have decreased  In the  early 60fs,  had
Increased in abundance to values similar to those In August of 1954.

23.  With a  phytoplankton and zooplankton  abundance and biomass  between
those of Lakes Erie  and Huron,  the presence  of the oligotrophic  rotifer
associaton and  the oltgotrophic  crustacean indicator  species  Diaptomus
siciI Is and Limnoqalanus macrurusf  the predominance  of mesotrophic  and

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ollgotrophic diatom species,  and the  similarity of the plankton ratio  on
the north-south axis suggest that the offshore waters are currently In the
upper ol igotrophic-lower mesotrophic range (i.e. meso-oligotrophic).

24.  A significant change in zooplankton composition has occurred with the
establishment of Daphnia pulIcaria in  the entire offshore region of   Lake
Michigan.    Decline  of  the alewlfe  population  has  apparently  reduced
predatory  pressure  from  alewife releasing  the  suppressed  large-bodied
zooplankton such as Dqphnla pulIcaria (Scavla et al. 1986).   In  addition,
abundances of Leptodora k i ndt i i f   Daphnia galeata,  DIaptomus ash I and!   and
Cyclops bicuspldatus have returned to  or exceeded abundances observed  in
1954 during a period of low alewlfe abundances.

25.  Correlation analysis suggests  that the Increases in Daphnia  galeata
mendotaef  as well  as £. pul icarta, have exerted greater grazing  pressures
on the phytoplankton community.

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                                 SUMMARY

                                Lake Huron

1.   In 1984, 315  algal and 53 zooplankton  species were observed in  Lake
Huron.  Compared to 1983, a 4.3$ and 8.6% reduction In the number of algal
and  zooplankton  species   occurred.    These   flucuations  in   species
composition are due to both natural  and seasonal  sampling variability.

2.  Compared  to Lake Erie,  variability in common  algal species in  Lake
Huron between 1983 and 1984 was low.   94$ of the common species  observed
in 1984 were also common species in 1983.  10% of the common algal  species
observed in 1983 were not common in 1984.

3.  Average biomass,of the phytoplankton and zooplankton was 0.38±.10 g/m
and  27.3+2.3  mg/m   for  the  study  period.    Mean  phytoplankton  and
zoopIankton3 abundance   were   17,200±890   cells/mL   and   55,400+7,200
organisms/m .

4.  Diatoms  possessed the greatest  number of species  (156) and  biomass
(61.9$ of  the total)  in 1984.    The Chrysophyta  were the  second  most
important division (9.5% of the total) in 1984,  which represented a change
from 1983 when the Cryptophyta were second In Importance.

5.  Picoplankton accounted for 83.9$ of the total abundance but only  1.4$
of the biomass.  This finding  is similar to that of 1983.

6.  Considering biomass,   the diatoms were  dominant throughout the  study
period accounting  for as  much as  12% but  never less  than 44?  of  the
biomass.  The large drop in  the relative importance of diatoms In  August
of 1983 was not  observed In 1984.    A bloom  of Rhizosolenia eriensis  in
August of 1984, not observed in 1983, was the major cause of the dominance
of diatoms during the summer of 1984.

7.   Average phytoplankton  abundance for  the sampling  period  generally
decreased from  the  northern stations  to  ~Station 15,  where  abundance
increased and  then  decreased  si ightly  southward.     The  mean  station
zooplankton abundance was higher in the northern half than in the southern
half of the lake due primarily to higher rotifer abundance in the north.

8.   In general, offshore  species compositon of  phytoplankton has  changed
little since the  early 70's.    StephanodIscus minutus was  not common  in
1971, 1974,  1975,  1980 and  1983.    In 1984 it  was common with an  average
density of  19.4 cells/mL  because  of the  inclusion of  winter  samples.
Abundance averaged 63 cells/mL in February.

9.    Vertical   distribution  studies   Indicated  that  an  increase   in
picoplankton, BactIlariophyta and Chrysophyta occurred to a 30-m depth  at
Station 37.     The abundance   increase  correlates with  the  decrease  In
temperature associated with the metalImnion.

10.  Both in  1983 and 1984 the  dominant diatom assemblages were  species
characterized as indicators of oligotrophic or mesotrophic conditions.

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11.  The ratio of mesotrophlc  to eutrophic algal  species (trophic  ratio)
has not changed since 1971.   This suggests that the trophic status of  the
offshore waters of Lake Huron has not changed since 1971.

12.   The  Rotifera possessed  the  largest  number of  species  (31)  and
relative abundance (56.0%).   The Calanolda (42.0$) dominated on a  blomass
basis followed by the Cladocera  (27.5$).   Rotifera contributed only  2.5$
of the zooplankton biomass.

13.  Species composition  of zooplankton was similar  In 1971,  1974,  1983
and 1984.   Diaptomus oregonensis  was more  prevalent in  1983 and  1984,
while]), ash I and I and £. sic!I Is  have Increased in abundance since  1971.
Limnocalanus macrurus  appears to  be decreasing  In abundance.    Bosmlna
IongIrostrIs and HQI oped I urn gibberum were more abundant in 1971 than 1984.

14.  Daphnla pulI car I a was first observed In offshore waters In 1983.   In
1984, lakewlde abundance decreased.  Within the Cladocera, rank  abundance
dropped from third In 1983 to fifth in 1984.

15.  A new cladoceran species,  Bythotrephes cederstromllf was observed  in
the offshore waters of Lake Huron.

16.  The rotifer  community was dominated by  an assemblage indicative  of
oligotrophic conditions  in 1983  and  1984.   In addition,  the  calanold
PJaptoma.5 sjcjJig, an oligotrophic Indicator, was fairly abundant.

17.   The  plankton  ratio  (Calanoida/Cladocera  +  Cyclopoid)  was  high
compared to Lake Erie  but similar for the  entire offshore region,  which
suggests a similar high quality of  water over the entire offshore  region
except for the far northern Station 61.  The plankton ratio at Station  61
was similar to that of the Straits of Mackinac and northern Lake Michigan.

18.  The presence of  the oligotrophic rotifer assemblage, the  domination
of the calanoids, the fairly abundant ollgotrophlc Diaptomus s?cllisf  and
the low  zooplankton  abundance  compared  to  those  of  Lakes  Erie  and
Michigan, suggest the offshore waters of Lake Huron In 1983 and 1984  were
ol igotroph ic.

19,   Phytop Iankton  biomass and  zooplankton  abundance of  the  offshore
waters of Lake Huron in 1971,  1980, 1983 and 1984 were not  significantly
different.  Similarly, offshore zooplankton biomass was not  significantly
different between 1976 and 1984.

20.  The consistency of the trophic ratio and algal biomass through  time,
the  Insignificant difference in zooplankton abundance from 1970-1984,  the
occurrence of oligotrophic  and mesotrophic algal   indicator species,  the
ollgotrophlc zooplankton assemblage,  and the similarity  of the  plankton
ratio over the entire offshore suggest  that no significant change  in  the
trophic  status of the offshore waters of Lake Huron since 1970.

21.  With a mean  algal biomass of 0.38 and  0.42 g/m3 for 1984 and  1983,
respectively,  Lake  Huron  would  be classified  as  oligotrophic   by  the
classification scheme of Munawar and Munawar (1982).

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22.  The appearance of Daphnla  pul icaria In Lake Huron suggests that  the
zooplankton community has been released from size-selective planktivory.

23.  The correlation of phytoplankton abundance with total  phosphorus  and
zooplankton abundance within individual  cruises suggests that "top   down"
and "bottom  up" control   of the  trophic web  of lake  ecosystems  exists
simultaneously and that it varies with season.

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                                 SUMMARY
                                Lake Erie

1.  In 1984,  356 species of phytoplankton  and 81  species of  zooplankton
were observed.   As compared to  1983,  a 4.3$  reduction In  phytoplankton
species,  mostly Chlorophyta, and an 18.5? increase in zooplankton species,
mostly  Rot Ifera,  were  observed.     As  the  same  sampling  enumeration
procedure and taxonomy were employed,  the observed flucuations In  species
composition are due to both natural and sampling variability.

2.  Compared  to Lakes Michigan  and Huron,  a  high variability In  common
algal  species existed  between 1983 and  1984 in Lake  Erie.    Eighty-four
percent of the common species observed  In 1984 were also common in  1983.
Thirty of the  common species observed  In 1983 were  not common In  1984.
The number  of  common zooplankton  species  between 1983  and  1984  were
simiIar.

3.    Mean  phytoplankton  and  zooplankton  abundance  were  45,100±4,200
cells/mL and 159,600+25,300  organisms/m  for the  study period.   Average
biomass of phytoplankton and zooplankton was 1.00+.16 and .053+.0062  g/m
in 1984.    Phytop Iankton biomass  varied within  Lake Erie.   The  western
basin possessed, a  greater  biomass (1.38+0.23 ,g/m )  than  the  eastern
(0.54±0.082 g/m  >  and  central  (0.76±0.09  g/m )  basins.    Zooplankton
abundance increased in a  similar fashion Into the  western basin but  not
zooplankton biomass.

4.  Diatoms possessed the greatest diversity of species (171) and  biomass
(47.8$ of the total)  in 1984.   Compared to 1970, a significant change  in
diversity of phytoplankton has occurred.   In 1970 only 21 diatom  species
were observed that  accounted for  53$ of  the biomass.   The  Chlorophyta
possessed the largest number of species (78) In 1970.

5.  Picoplankton accounted  for 89.6$ of the  total abundance.  A  similar
finding was observed  In 1983.

6.  Diatoms  were dominant   in April  and May  and were  succeeded by  the
Cryptophyta  in  July and  the  Chlorophyta in  August.   By  December  and
through the winter months, the diatoms were again dominant.

7.  The  historically highly  productive western  basin has  had a  steady
decrease in algal  biomass from 1958  to 1984.   Similarly, chlorophyll  .a
levels have decreased In all basins, but most dramatically In the  western
basin.  However, algal biomass is  still higher In the western basin  than
in the central and eastern basins.

8.  Lakewide, the mean weighted alga! biomass was 3.4, 1.5 and 0.8 g/m  In
1970,  1983  and 1984, respectively.   A 56 to 76$ reduction  in algal biomass
has occurred  in offshore waters of the  lake from 1970 to 1983/84.

9.  Although occurrences  of common and dominant  species were similar  in
1970,  1983  and 1984,  dramatic decreases in the  biomass of these  species
were evident.  For example, a 96$ reduction  in the maximum biomass of  the
nuisance species ftphanizomenon  fIos-aquae has occurred  since 1970.   The

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eutrophlc  Indicator  species  Stephanod\scus  binderanus  and  Frag 11aria
capucina have had a >90.$ reduction In maximum blomass.

10. Aster toneI la formosa has not been  prevalent in Lake Erie since  prior
to 1950.  In the 1984 spring cruises,  .A., formosa was the dominant  species
on a  blomass basis.    Meios!ra isiandjca,  a mesotrophic  indicator   not
common  in 1983,  was common in 1984.

9.  The Rotifera possessd the largest number of species (48) and  relative
abundance (80.1$) of the  zooplankton.  On a  biomass basis, the  Rotifera
represented only  13.6$ of  the zooplankton  biomass while  the  Cladocera
contributed 40.5$ of the biomass.

10.  A shift  in zooplankton composition is  occurring with a new  species
Daphnia pulIcaria being observed for the first time in 1984.  On a biomass
basis, fi. pulicaria was  the dominant Cladocera in  the lake with a major
bloom in  August.   However,  it  was  most prominent  in the  central   and
eastern basins.     The  prevalence  of  the  eutrophic  cyclopoid  Cyclops
vernal is has decreased within the lake,  especially within the central   and
eastern basins.

11.  A decrease  in summer Cladocera and Copepoda abundance in the  western
basin Is suggested from 1961  to  1984.  Rotifera abundance In the  western
basin has increased since 1934.   A number of eutrophic rotifer  Indicator
species had  abundances  restricted  to or  significantly  higher  in   the
western basin.   The plankton ratio also suggests a more productive  status
for the western basin.

13.  There is a  lack  of dominance of eutrophic rotifer Indicator  species
for the entire lake.   This suggests that  Lake Erie  in 1984 as a unit   is
not eutrophic.     The  number  of dominant  eutrophic  algal  species   has
decreased,  while the number of  dominant mesotrophic species has Increased;
that  is, the  trophic ratio  has increased, suggesting  an improvement   in
water quality.

14.  Evidence  of a shift  in trophic status  of Lake Erie  since 1970   Is
provided by   the  trophic ratio,  the  plankton ratio,   phytoplankton   and
zooplankton  indicator species,  declines in total abundance and blomass   of
total phytoplankton and zooplankton since  the mld-60's and 70's,  declines
in abundance of  nuisance species and eutrophic species,  declines in total
phosphorus and chlorophyll .a, and the current total  blomass and  abundance
of plankton.

15.  The trophic condition of Lake Erie appears to be improving.  However,
compared to   Lakes  Huron  and   Michigan In  1983  and  1984,   biomass   of
phytoplankton and zooplankton was higher,  the plankton and trophic  ratios
were  lower,   and the  phytoplankton and zooplankton species  compositions
suggest a more productive status for Lake  Erie.

16.   Based   on  the  classification  schemes of  Vollenwelder  (1968)   and
Munawar and  Munawar  (1982)  utilizing maximum  and average algal  biomass,
the western  basin would be meso-eutroph!c,  the central  basJn  mesotrophic,
and the eastern  oligo-mesotrophic.   This conclusion is supported by other
indicators of the trophic status noted above.

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                                                                      10
17.    The  decreases  in  phytoplankton  abundance,   chlorophyll,    total
phosphorus and  turbidity are  consistent with  expectations of  long-term
nutrient control.  However,  the significant changes In the composition  of
the zoopIankton community  with the  appearance and  establishment of  the
large  cladoceran  Daphnia  pulicarla  are  attributed  to  a  change   In
planktivory.    The  planktivorous  emerald  and  spottall  shiners   have
dramatically declined,  perhaps due to a resurgence of the walleye and  the
salmonfne stocking programs.

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                                                                      11





                               INTRODUCTION








      Nutrient loading  of lakes and  rivers, navigation, fish  management



policies, fishing, shoreline  alteration, contaminant  production and,   in



general, economic  development,   ultimately  affect  the  lake  ecosystem.



Effects of  perturbations are  not  always known  and  can not  always  be



monitored individually in large,  complex systems such as the Great  Lakes.



Biological monitoring is an Integrative monitoring strategy (Johannson  et



at. 1985).   Ecosystems  respond to  stress with  compensatory changes  in



community structure and function mediated at the population level  (Boesch



and Rosenberg  1981).    Therefore,  changes In  ecosytem  health  can  be



detected by monitoring changes  in the biotic  community (Nlcholls et  al.



1980, Oil Ion et al. 1978).



        Any  monitoring  program must  first  document the  state  of  the



ecosystem, namely, the species composition, biomass and production of each



community component, including  the normal range  of temporal  and  spatial



variation.    The  second  step    Is  to  examine  the  relationship   and



interactions amongst the  ecosystem components in  order to interpret  and



possibly predict future changes in community structure or function.  Thus,



the value of  such monitoring  programs goes far  beyond its  surveillance



capabilities; It can  form the backbone  for research activities,  thereby



encouraging a detailed understanding of the system.



       This  project  reported here  was initiated  by the  United  States



Environmental Protection  Agency,   Great  Lakes  National  Program  Office



(GLNPO), to analyze phytoplankton and zooplankton samples from Lakes Erie,



Huron and  Michigan  taken  in 1984  and  the  winter of  1985.    Because



phytoplankton are sensitive to water quality conditions and possess  short

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                                                                      12




carbon turnover rates,   the determination of   phytoplankton  abundance  and



species composition has become established as a method  to trace  long-term



changes  in  the  lakes  (Stoermer   1978,  Munawar  and   Munawar   1982).



Similarly,  zooplankton have value as  indicators of water quality and  the



structure of the biotic community and have proved useful  for complementing



phytoplankton to assess the apparent  effects of water  quality   conditions



(Gannon and  Stemberger 1978)   and of  fish populations  (e.g.  Brooks  and



Dodson 1965) on biota.   This report represents the second year  of  similar



samp I ing intensity and pattern of the offshore region of  Lakes  Erie,  Huron



and Michigan.



        An   in-depth planktonic  (phyto-  and zooplankton)  comparison  is



presented based  on extensive  seasonal   lake-wide surveys  including  the



winter of  1985.   This  comparison was  achieved  by the  application  of



standard and  consistent identification,  enumeration and  data-processing



techniques of plankton that were collected along north-south transects  in



Lakes Huron  and  Michigan and  east-west  transects  In  Lake  Erie.     In



addition, the  vertical distribution of phytoplankton was examined In each



lake during the year.



     The primary objectives of this report include:



(1)  To organize plankton data for use  in eutrophication  models;



(2)  To characterize the  composition and abundance  of the  phytoplankton



     and zooplankton for  comparison with  past conditions  to  the  extent




     that they are known;



(3)  To provide firm  documentation with  which future   assessment of  the



     changes In water quality of the lakes can be made;



(4)  To characterize  the  water quality  by   studying  the  abundance  and



     autecology of phytoplankton and zooplankton; and

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                                                                     13
(5)   To characterize within   and  between year  plankton variance to  allow



     better long-term  assessments of changes  in plankton structure.

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                                                                    14




                                  METHODS





Samp I ing Sites




       Phytoplankton and  zooplankton samples  from Lakes  Erie,  Huron   and



Michigan were collected by GLNPO  personnel   during several  cruises  (9-11)



during the  spring,  summer  and autumn  of  1984 and  the winter  of  1985.



Collection dates and  station locations  of  routine  plankton sampling   are



given in Tables 1  and 2 and in Figures 1  - 3.  Locations of  sampling  sites



on Lakes Michigan and  Huron were not seasonally  consistent (Tables 3   and



4).  By design, alternate east-west stations were sampled (e.g.  5 or 6,   10



or 11;  Fig. 1) on  various cruises.  This selection  of sites was based   on



previous studies which indicated that adjacent east-west sites were  within



homogeneous areas of  Lake Michigan  (Moll et  al. 1985).   For  analytical



purposes, east-west  stations are  combined, assuming  that  no  significant



difference in  species abundance  and composition  exist between  east-west



stations, to give a single north-south  transect.  All  sites are also  part



of the Great Lakes International Surveillance Program.





Chemistry



     Only selected  water quality variables collected during the study   are



presented in  this  report.    Results  of  the  complete water  chemistry



investigation are reported  elsewhere (Lesht and  Rockwell 1987).   Methods



used were standard procedures (Lesht and RockwelI 1987).






PhytopIankton



      An 8-liter PVC Niskin bottle  mounted on a General Oceanics  Rossette



sampler with a Guildline electrobathythermograph (EBT)  was used to  collect



phytopIankton.  One-liter composite phytoplankton samples were obtained   at



each station by compositing equal aliquots from water samples collected   at

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                                                                       15



depths of the surface,  5m, 10m and  20m as allowed  by total water  depth.


Vertical distribution samples were collected at selected stations from  the


surface, 5m, 10m, 15m and 20m (occasionally to 30m).


     Phytoplankton samples were immediately preserved with 10 ml of Lugol's


solution, while formaldehyde was added upon arrival in the laboratory.  The


settling chamber procedure (Utermohl 1958) was used to Identify (except for


diatoms) and enumerate phytoplankton under  phase contrast microscopy at  a


magnification of  500x.    Objects  (spheres <  lum,  rods  <  3um  length)


possessing a  bluish  cast were  identified  as picoplankton,  while  those


appearing as dull grey were not  counted.  The designation Haptophyte  spp.


used represents  a col lection  of  morphological forms  more  appropriately


titled Haptophyceae.  A second identification and enumeration of diatoms at


1250x was performed after  the organic portion was  oxidized with 30?  H202


and HNO,.  The cleaned diatom concentrate was air dried on a cover slip and

                                       TM
mounted on a slide (75x25mm) with HYRAX   mounting medium.  Identifications


and counts were done by Dr. Norman A.  Andresen,  Mr. Mark A. Lamb,  Dr. Louis


L. Lipsey, Ms.  Heather K.  TrulIi  and Dr.  Marc Tuchman  of the  Bionetics


Corporation.


       The cell  volume  of each  species was computed  by applying  average


dimensions from each  sampling station  and date to  the geometrical   shape


such as  sphere,   cylinder,  prolate  spheroid,   etc.,  that  most  closely


resembled the species  form.   At  least 10 specimens  of each species  were


measured for the  cell volume calculation.   When fewer  than 10  specimens


were present,  those present  were measured  as they  occurred.   For  most


organisms,  the measurements  were taken  from the outside  wall to  outside


wall.  The  protoplast  was  measured   with  loricated  forms,   while   the


individual  cells  of  filaments and  colonial   forms were  measured.     For

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                                                                       16





comparative purposes, biovolume  (urn /L) was converted  to biomass  (mg/m )



assuming the  specific gravity  of phytopiankton  to be  1.0  (mm /L=mg/m )



(Willen 1959,  Nauwerck 1963).
ZoopIankton



     A Wildco Mode!  30-E28  conical  style net (62-um mesh net;  D:L ratio  =



1:3) with 0.5m opening (radius=0.25m) was used to collect,  where  possible,



two vertical zooplankton samples at each station.  Vertical  tows were taken



from 2m above  the bottom to  the surface (long  tow) and from   20m to  the



surface (short tow).  The short tow was analagous to an epilimnetic tow  in



stratified waters.   Filtration volume and towing efficiency were determined



with a Kahl flow meter (Model  OOSWA200)  mounted in the center  of the  net.



Filtration efficiency  averaged 83.4,  75.9  and 85.8$,  respectively,   for



Lakes Erie, Huron and Michigan for  the entire sampling season.   Following



collection, the net contents  were quantitatively transferred to  0.5-IIter



sample bottles, narcotized with club  soda and preserved with 5%  formalin.



Identification and enumeration  of zooplankton followed  Gannon (1971)   and



Stemberger (1979) and were performed by D. Page, H. Trulli  and  L. Stokes of



the Bionetics Corporation.



     Raw counts were converted to number/m  by Bionetics, Inc.   The  volume



of each rotifer species  was computed by using  the geometrical shape  that



most closely  resembled the  species  (Downing and  Rigler  1984).   It  is



essential that the measurements  are made on  the population being  studied



since they vary In different habitats for some species up to 100$ and  more



(BottrelI et al. 1976).  For each  cruise, length of at least 20  specimens



of each rotifer species was measured.   Width and depth were also  measured



on one date for each lake  to develop length-width and length-depth  ratios



for use in the simplified formulas of  BottrelI et al. (1976).   Assuming  a
                                 T

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                                                                      17




specific gravity of one,   volume was converted to  fresh weight and to  dry



weight assuming a ratio of dry to  wet weight of 0.1  (Doohan 1973) for  all



rotifer species except Asplanchna  spp.  A dry  weight/wet weight ratio  of



0.039 was used for Asplanchna spp.  (Dumont et al. 1975).



      Because  of the considerable  variability In length  and thus  weight



encountered In the Crustacea, the dry weights of Crustacea were  calculated



using length-weight  relationships  (Downing  and RIgler  1984).     Average



length of crustaceans (maximum of 20  for each station) was determined  for



each station  of  each cruise.    A  comparison of  calculated  weights  to



measured weights of Crustacea in  Lake Michigan suggests good agreement  at



the minimum weight range (Table 5).  The  use of the mean and the high  end



of the range for comparison  Is tenuous because they are affected by  sample



size and  selective feeding  of  predators.   The  weight of  the  Copepoda



nauplil followed Hawkins and Evans (1979).





Data Organization



        Abundances and  dimensions of  each  species of  phytoplankton  and



zooplankton were entered Into  a Prime 750 computer  using the INFO  (Henco



Software, Inc., 100 Fifth Avenue,  Waltham, Mass.) data management  system.



Blomass was calculated  for phytoplankton and  zooplankton and placed  into



summaries  for  each  sampling   station  containing  density   (cells/ml),



biovolume (urn /ml) and relative abundances  of species.  In addition,  each



division was  summarized by  station.   Summary  information Is  stored  on



magnetic tape and is available for further analysis.





Definitions



      Common phytoplankton species were defined  as having an abundance  of



>0.1J8 of the total cells or >0.5$ of the total biovolume.

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                                                                       18






     Common crustacean zooplankton species were defined as having >0.1?  of



the total abundance or  >1.0$£ of the total  blomass.   Rotifer species  were



considered common If they accounted for >1.0$ of the total abundance.





     Species diversity refers simply to the number of species observed.





     Dominance refers  to a  community property reflected  in the  relative



abundance pattern of species.  A  species was considered to be dominant  if



It possessed  the highest  relative  abundance or  biomass of  a  taxonomic



grouping (i.e. division).





     Importance refers to a group of measurements by which the species in a



community can be compared (Whlttaker 1975).   Abundance or biomass was  the



Importance value used  In the discussion.

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                                                                       19




                          RESULTS AND DISCUSSION






LAKE MICHIGAN



Phytop Iankton



     Species lists (Table  A1) and summary tables of abundance (Table  A2)



and biovolume (Table  A3) are in  Volume 2 -  Data Report.   A summary  of



water chemistry parameters is presented in Table 6.






Annual Abundance of. Major Algal  Groups



      The phytop Iankton  assemblage of 1984 was  comprised of 327  species



representing 91 genera  from eight divisions.   Compared to  1983,  a  ~A5%



reduction in  the  number  of  genera and  species  was  observed.     This



difference was  mostly  attributable  to  a  decrease  in  the  number  of



Chlorophyta, Chrysophyta and Cyanophyta (Table 7).



     Similar to 1983, the Bad 11ariophyta possessed the largest number  of



species (166)  and biovolume  (70.0$ of  the total,  Table 8),   while  the



second largest number  of species  (63) was observed  for the  Chlorophyta



(Table 7).  The Cryptophyta,  as in 1983,  accounted for the second  highest



biovolume (11.6$)  (Table 8).    Highest overall  densities were attained  by



the picoplankton  (82.9$ of  the total).    Both  the Pyrrhophyta  and  the



Chlorophyta had much lower biovolumes in 1984 than in 1983 (Table 8).  The



annual  average  phytop Iankton  density  and  biomass  were   22,220+1,400



cells/mL,  (mean±S.E.)   (29,839   cells/mL,    1983)  and   0.55   g/m3±.038



(mean±S.E.)  (0.42  g/m ,  1983),  respectively.






Seasonal  Abundance and Distribution of Major Algal Groups



      Seasonally,  abundance (cells/mL) was low   during the spring and  had



increased by July.   Because sampling in the present study was designed  to



monitor the  early  pre-bloom conditions,  the spring bloom observed in  May,

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                                                                      20





June and July  of 1976  (Bartone and Schelske  1982)  was  not observed   In



1984.  A secondary abundance maxima  was observed In  August (Fig.  4a)  but



was not observed in the biovolume seasonal  distribution (Fig.  4b).   During



August, a general  downward trend  in biomass occurred.   Because   samples



were not taken in October,  the large autumn peak (48,305 cells/mL)  seen  in



1983 (Makarewicz 1987) was not observed In 1984.  Similarly,  a fall   bloom



was not  observed  in 1976  by  Bartone and  Schelske  (1982).  This was



attributed either  to  a  weak bloom  that  was  not  observed  or   to the



occurrence of the bloom at a time when samples were not taken.



     Considering  biovolume, the Baci11ariophyta were dominant  throughout



the study period accounting for as much  as 80$, but  never less than 55$,



of the phytoplankton biovolume  (Fig. 5).   The overwhelming dominance   of



the diatoms throughout the study  period precluded the prominent  seasonal



succession of algal divisions observed in 1983 (Makarewicz 1987).



      The large drop   in biovolume of BaciIIariophyta  (to -10$) noted   in



August of 1983 (Makarewicz  1987) was not  observed In 1984.   A bloom   of



Rh I zoso I en i a er Iejisls during the summer of 1984, not observed in 1983,  was



the major cause of the dominance of the diatoms  in August (Table 9).   For



example, on the  12-14 August cruise,  abundance of £.  erlens Is was only



17.5 cells/mL,  but the  biovolume per  cell  was high.   Thus,  this  one



species accounted for 26.9$ of the total biovolume during the cruise.



     The small decrease In diatoms  in August of  1984 corresponded with   an



increase  In the Cryptophyta,  while  in 1983 the  major decline  in  diatoms



corresponded with an   increase  in  the Pyrrhophyta.   A  similar shift   in



biovolume composition was observed  in 1976 with  diatoms decreasing to  17$



In August  when  greens and  blue-green  algae predominated   (Bartone  and



Schelske  1982).

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                                                                      21
Regional  and Seasonal  Trends in the Abundance of  Common Taxa



        Common  species  (Table  10) were  arbitrarily  defined  as  those



possessing a relative abundance  of >0.1$ of  the   total  cells or >Q.5%  of



the total biovolume.   Forty-three common species  were observed In  1984-85



compared to  45  In 1983.     Seventy-six  percent of   the  common  species



observed In 1984 were also common  species in 1983; thirty-one percent  of



the common species  observed in 1983  were not common  in 1984 (Table  11)



(Makarewlcz 1987).     The  cause  of these  differences  is  difficult  to



evaluate.  Natural  annual  variability in the lake has never been evaluated



and cannot be evaluated until a longer data set exists.   Seasonal  sampling



variability exists between 1983  and 1984 and is   the most probable  cause



for the  species  differences  observed.   For example,   Dlcthyosphaerlum



ehrenberglanum was a common  species for the 1984-85   survey,  but not  the



1983 survey, by  virtue of the  inclusion of  a winter sampling period  in



1984-85.   J2.  ehrenbergianum was  prevalent only   In  the  winter of  1985.



Since no winter samples were analyzed In 1983-1984, this species was not a



common species for the entire 1983 sampling period .



      Because of the similarity between  the 1984 common species list  and



the 1983 Iist, a species by species description of autecology and regional



and seasonal  trends are  not warranted  here and  can be  referred to  In



Makarewlcz (1987).   Only new common species are discussed below.






BaciIlarlophyta



  Cyc|otella oceI Iata Pant.



       Cyclotelia  oce11ata was  observed in  the southern  basin of  Lake



Michigan in low numbers in 1963 (Stoermer and Kopczynska 1967a).  In  1967



this species was most abundant at offshore localities In the northern part




of the lake  although occasional  populations were noted  in the  southern

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                                                                      22





basin (Stoermer and  Yang 1970).   In  1972 Holland  (1980) reported  this



species as  most abundant  during the  summer (maximum  abundance range  =



50-70 cells/ml).  However,  it was not a common species in 1983 (Makarewicz



1987).  Cyclotella oceliata  is generally abundant in  areas of the  Great



Lakes which have  not undergone significant  eutrophication (Stoermer  and



Krefs 1980); i.e.,  associated with oligotrophic  conditions In the  Great



Lakes (Stoermer and Yang 1970).



      In 1984  abundance increased into the  summer (mean maximum  station



abundance = 39  cells/mL),  dropped by  mid August and  stayed low In  late



autumn (Fig. 6a).  Mean abundance and biomass were 23.3 cells/mL (0.10$ of



the total cells) and 2.1 mg/m3 (0.38$ of the total biomass) (Table 10).  A



maximum abundance of  265 cells/mL  occurred on  8-9 July  at Station  17.



Mean abundance was high at the most northerly station (77) (45.2 cells/mL)



and at Stations 17 and 22 (46.5 cells/mL) (Fig. 6a, Fig.  9a).






  Synedra ulna var. chaseana Thomas



      Stoermer and Kopczynska  (1967a and  b) reported this variety,  along



withJS. 111 na  var. danica,   as reaching 100  cells/mL In  early August  of



1962.  Although several  members of  the genus occur In Lake Michigan,  the



only numerically  important  taxa were .$_.  ul na var. chaseana  and _S_.  u I na



var. danica in 1962 and 1963,    Abundance of this variety was low in 1983



(0.16 cells/mL) (Makarewicz 1987).  Stoermer and Yang (1970) characterized




£. ulna var. chaseana as an oligotrophic offshore dominant.



      In  the present  study,  a July  maximum was observed  followed by  a



population crash by  early August (Fig.  6b).  Spring,  autumn and  winter



abundances were low.  Mean density and biomass were 2.2 cells/mL and  17.2



mg/m , respectively.  This species  represented 3.1$ of the total  biomass

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                                                                      23




for the entire  sampling period.   Maximum  abundance was  23 cells/ml  at



Station 34 in early July.






  Synedra f ?Iiformis Grun.



     Earlier work had  suggested this species to be largely restricted  to



the offshore  waters  (Stoermer and  Yang  1970) and  highly  oligotrophic



regions such as Grand  Traverse Bay (Stoermer et  al.  1972).  However,  it



was fairly abundant in Green Bay in  1977 with an average density of  14.3



cells/ml.   Similarly,  density  averaged  36.9  cells/mL  (0,95%  of  the



population) in the nearshore of southern  Lake Michigan in 1977.    Average



abundance in 1983 was 2.59 cells/mL (maximum of 25.5 cells/mL).



     Abundance  was high in April,  May and  July of 1984 (Fig.  6c).   Mean



seasonal abundance  reached a  maximum  of 30.8  cells/mL  in July.    The



maximum density  observed was  118  cells/mL  at Station  34 on  7-9  July.



Average abundance and biomass for the non-winter period was 11.2  ceils/mL



and 4.2 mg/m , respectively (Table 10).






  Rh izosolenia longlseta Zach.



      During 1962 and  1963, R. eriensis was  the dominant member of  this



genus with a small population of R.  graciI is also noted in Lake  Michigan



(Stoermer and Kopczynska 1967a).  Holland (1980) observed densities of  R.



eriensis reaching  ~750 cells/mL  in 1970  but did  not report  any  other



species of Rhizosotenia.    In the  1977  study of  Green Bay (Stoermer  and



Stevenson 1979),  only R, eriensis (maximum = 90 cells/mL) and R.   graciI is



(maximum = 46.1 cells/mL) were observed.  Similar maximum abundances  were



observed for the  nearshore zone  of southern Lake  Michigan [JB.   eriensis



(maximum  =  81.7  cells/mL);  R.  gracllis  (maximum  =  46.1   cells/mL)3



(Stoermer and Tuchman 1979).   In 1983,  R.  eriensis and R. longlseta  (R.

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                                                                      24





long!seta = £• graclI is) were observed (Makarewicz 1987).   In the  present



study,  E. long I seta was the more  abundant,  but £. erTens Is contributed  a



greater biomass (Table 10).



       £.  long I seta  abundance was  highest in  the spring,   appeared to



decrease to late July  and increased in mid-August.  Late autumn and winter



abundances were  low  (Fig. 6d).    Average abundance  was  21.2  cells/mL



representing 0.1$ of the total  cells and 4.38$ of the total  biomass (Table



10).






  Nltzsch ia Iauenburgfana Hust.



      Stoermer  and Kopczynska  (1967a), Holland (1980)  and Stoermer  and



Tuchman (1979) did not report this species in Lake Michigan.   Stoermer and



Yang (1970) did not  list U. lauenburgtana as  a dominant  plankton in  the



Great Lakes.   However,  it  has been reported  as occurring  in Green  Bay



(mean = 0.41 cells/mL;  maximum = 16.1  cells/mL) (Stoermer and  Stevenson



1979).    In 1983  this species  occurred  only seven  times (mean  =  1.36



cells/mL).     In 1984  it  was a  common species  by  virtue of  Its  large



biovolume (Table 10).   Maximum mean seasonal abundance occurred during the



spring sampling (Fig.  7a).






Chlorophyta



  Oocy st i s submarina Lagerh.



     Stoermer and Kopczynska (1967a) noted that in 1962 and 1963 the three



most common species  of OocystIs were  Q. el Iiptlca, Q.  submarina and Q.



lacustris.   Abundance ranged from 2 to 10 cells/mL.  Oocystis spp. was one



of the most abundant taxa observed in August and October of 1977 in  Green



Bay  (Stoermer  and  Stevenson  1979).    Mean density  was  133.8  cells/mL



representing 2.4$ of the population.  Similarly, the abundance of OocystIs

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                                                                      25





spp. was relatively high (mean =  30.9 cells/ml;  0.57? of the  population)



in the nearshore of Lake Michigan in 1977 (Stoermer and Tuchman 1979).



     Q. submarine was observed in 1983 but was not a common species  (mean



= 0.2  cells/ml; <.01$  of total   population).   In the  present study,   a



maximum  pulse  of  254  cells/ml  was  observed,   but  the  average   was



considerably lower (mean = 26.5  cells/ml; 0.12$  of the total   population)



(Table 10).    Abundance  was  low during  the  spring  and  progressively



increased to a  peak in  late August.    Abundance  was again  low by  late



autumn (Fig. 7b).  Generally, abundance of Q.  submarine was higher at  the



northern stations (Stations 64 and 77) and the southern stations (Stations



6, 10, 18 and 22) as compared to the mid-lake  region (Fig.  9b).






  Dictyosphaerium ehrenbergianum Naeg.



        In the  1962-63 study  of  the southern  basin of  Lake  Michigan,



Dictyosphaerium  was  observed   in  the  autumn    (usually  <1    eel1/mL,



occasionally 5 cells/mL)  but was not  noted in the  spring (Stoermer  and



Kopczynska 1967a  and  b).   Stoermer  and Ladewski  (1976)  reported  the



abundance of this species as being high in 1971 (peaks over 200  cells/mL,



many occurrences over 100  cells/mL).   Average abundance  In Green Bay   in



1977 was 10.3  cells/mL with  a maximum  of 106.8  cells/mL (Stoermer  and



Stevenson 1979).



     This was not a common species in 1983 (Makarewicz 1987) and would  not



have been in 1984 without  the addition of the  winter sampling date.    In



1984, mean abundance  was 23.6  cells/mL (0.11? of  the total   population)



(Table 10) with  a maximum abundance  of 298  cells/mL at Station  6 on  7



February 1985.  Mean  cruise abundance for the  February cruise was  105.2



cells/mL.   Seasonally,   spring abundance  was  ~25 cells/mL   followed by  a



decrease into the summer and a  major buildup  into late autumn and  winter

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                                                                       26




 (Fig. 7c).  Abundance  was substantially greater  in  the southern half   of



the  lake  (Fig. 9c).






Cryptophyta



  Cryptompnas rostratiformis Skuja



      Little historical   Information  is available  on the distribution   or



occurrence of this species  in Lake Michigan.   Much of the previous  work



simply  identifies a few  major species of Cryptomonas  and then  lumps  the



other occurrences under Cryptomonas sp.   For Green Bay, £. marssonIif   C.



Pvata, £. erosa and  £. gracile  were observed  in 1977 but apparently  not



£• rostratI form is.



      In 1983 JQ. rostratiformis was not a common species (Makarewicz  1987).



Abundance was low (1.3 cells/mL, Table 10)  in 1984 but biovolume was  high



 (4.57 mg/m , 0.84$ of total biovolume).  Seasonally, abundance was  low   in



the spring  and early  summer, increased  to -"2  cells/mL In  August,  and



maintained that level of-^2 cells/mL  into February (Fig. 7d).






Cyanophyta



  Osc iI Iator!a m1n i ma GIckIh.



      Both  Ahlstrom (1936) and  Stoermer and Kopczynska  (1967a) list  Q.



mougeot i i as the only species of the genus abundant In their  collections.



JQ. J linnet I ca and Q. bornet i i were  also observed by Stoermer and  Ladewski



 (1976).  Q. agardh!if Q.  Iimnetica, Q. subbrevis, Q. tenuis and Q.  m i n i ma



were observed In 1983 (Makarewicz 1987)  and 1984.  In 1983, Q.  IImnetica



and Q. agardhi i were common, while In 1984 Q. 11mnet i ca and Q. m i n i ma were



common.



        JQ. minlitp  abundance was  high  in 1984  (mean abundance  =  175.5



cells/mL, 0.79? of the  total  population) (Table  10).  Maximum  abundance

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                                                                       27





was 4,132  cells/ml  at Station  32  on 12  August  1984.   Abundance  was



greatest during the  summer period  (F!g. 8).   Geographically,  abundance



appears greatest at the mid-lake stations (Fig. 9d).





Vertical Distribution




     Besides the routine  Integrated samples, a vertical series of  samples



were taken at  two stations (18  and 47) on  15 August 1984  and were  not



integrated.   Abundance increased  with depth  at Station  47 and  can  be



primarily attributed  to  an   Increase  In  the  ptcoplankton  (Fig.  10a).



However,  BaciIlariophyta,  Chrysophyta,   Cryptophyta,  Chlorophyta   and



Cyanophyta also  increased with  depth  (Fig.  10b).   With depth,  species



diversity  Increased.  In  particular,  a 100$+  Increase In diatom  species



was observed between the surface and  the 10-m depth (Table 12).   Species



such as Aster? one I la fprmpsg,,  Frag 11 ar I a crotonens i sf  Cyclotel I a  oce I  I ata



(Fig. lOc), Rh izosolenia  er?enslsf fi.  Iong?setaf Chroomonas  porstedtiI,



Rhodomonas ml nuta  var.   nannoplankticaf  Osc11 Iator1 a  I linnet lea  and  JQ.



minima all increased  In abundance  with depth.   One species,   Cyclotella



comensis (Fig. 10c), was observed to decrease with depth.



      A similar Increase  in non-pI cop Iankton  species was not observed  at



Station 18 (Fig.  11a).    In contrast to Station  47, the abundance of  the



Baclllariophyta and Cryptophyta dfd not vary  In the top 20m of the  water



column.    The Chrysophyta  decreased  with depth.   All  other  divisions,



except the Cryptophyta,    increased in  abundance to the  30-m depth  (Fig.



11b).  As with Station 47, species diversity of BaciIlariophyta  increased



with depth, not to  the 20-m depth as   In Station 47,  but  from the 20  to



30-m depth.




     The Increase In the abundance and species of diatoms correlates  well



with the decrease  In temperature  associated with  the metalImnion  (Fig.

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                                                                      28




11b).  The appearance of an apparent sub-surface maximum In  phytoplankton



abundance is of  interest but not  surprising.  Brooks  and Torke  (1977),



Mortonson (1977) and Bartone and Schelske (1982) have previously  reported



a sub-surface  chlorophyll  maximum  in  Lake Michigan.   Reasons  for  the



existence  of  the  layer  are  not  clear  and  are  apparently  complex,



encompassing  physical,  chemical  and  biological   factors  (Bartone  and



Schelske 1982).






Winter Cruise




         Biomass  and  abundance  were  low  during  the  winter  and  not



significantly different from the autumn and spring values (Fig.  4).   As in



the non-winter season, the Baci I lariophyta (42.85? of the biomass) and  the



Cryptophyta (25.3$ of the biomass) were the dominant divisions.    However,



the Cryptophyta accounted for twice the biomass than during the non-winter



season (11.6?).




      Stephanodlscus minutus was the dominant  winter diatom (mean =   24.2



cells/ml); however,  this species was not a common species during the   rest



of  the  year.    Other  major  winter  diatoms,  FragiI aria  crotonensis,



label I aria floculosa and Aster loneI la  formosa were common species  (Table



10) during the non-winter period.



         Dicthyosphaerturn ehrenbergianum  was  the  dominant  Chlorophyta.



Seasonally,  abundance  of this  species was  low throughout  the year  and



reached Its peak abundance  in winter (93.5 cells/ml).   By virtue of  its



high winter abundance, It became a common species for the year (Table 10).



     Common winter  species of Cryptophyta and Cyanophyta were  Rhodomonas



minuta var.  nannoplanktlca  and QsciI later I a 11mnet1ca  and m1nIma,   which



were also common non-winter species.

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                                                                      29




Historical Changes In Species Composition



  Division Trends



       Because  common  species abundance,  blomass and  distribution  are



similar between 1983 and 1984, division and species trends are essentially



the same as those In an earlier  report (Makarewlcz 1987) and do not  need



to be repeated In detail here.   In August of 1962, an analysis of samples



from southern Lake  Michigan revealed  that the  diatoms were  numerically



dominant (Stoermer and  Kopczynka 1967a).    Relative abundance of  diatoms



was never lower than  ~70? of the  total assemblage at  all  stations.   By



1969 green,  blue-green and golden brown algae were the major phytoplankton



components (Schelske  and  Stoermer 1972).    Similarly, Schelske  et  al.



(1971) observed that blue-green and green  algae constituted 56 to 85%  of



the phytoplankton during  August and September.    In a  detailed study  of



southern Lake Michigan,  Stoermer (cited in  Tarapchak and Stoermer  1976)



observed that blue-green algae contributed up to 80% of the  phytoplankton



cells In August of 1971.



      By 1977, another shift In  algae composition was evident.   Relative



abundance of blue-greens dropped to 22.9% In August.  However, flagellates



(-42?) rather  than  diatoms  (22$)  were  the  dominant  group  of  algae



(Rockwell et al.  1980).  A similar composition as  in 1977 was observed  in



August of  1984  (diatoms  =  12.2%,  blue-greens  =  16.4?,  unidentified



flagellates = 42.1?)  if pIce-plankton  are not Included  in the  analysis.



However, in  addition  to  the  cyanophytes,  both  the  cryptophytes  and



chrysophytes were still numerically more important than the diatoms (Table



8) In 1983,  while In 1984 the chrysophytes were.   The numerical decline of



the diatoms  has  been  attributed  to the  high  phosphorus  loading  and

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                                                                      30




concomitant silica depletion (Schelske and  Stoermer 1971).  On a  biomass



basis, however, diatoms were the dominant group in 1983 and 1984.






  Species Trends




     Changes In common  species between 1983 and 1984 are discussed  under



Regional and Seasonal Trends  In the Abundance of  Common Taxa (Page  20).



Dominant diatoms  in 1983  Included  the numerically  dominant  Cyclotella



comensiSf FragiI aria crotonensls and Me IosIra itallea subsp. subartlca;  on



a biomass basis,  Tabe\\ar\a f.ioccuIosa was predominant (Makarewicz  1987).



In  1984  Cyclotel |a  comensls  and  Frag 11aria  crotonensls,   along  with



Cyclotel la  oce 11 at .a,  were  numerical ly   dominant.     M.   Ital I pa  subsp.



subarctlca was common but not dominant.  On a bfomass basis,  Rhizosolenla



erlensls and TabelI aria fIoccuIosa were predominant  in 1984.



      The Haptophyceae,  Monoraphldium contortuji (Chlorophyta),  Dinobrypn.



socI aIe   var.    amer 1canum   (Chrysophyta),   Rhodomonas   minuta    var.



nannoplanktica and Chroomonas  norstedIi  (Cryptophyta),  AnacystIs  montana.



var.   minor  and  OsciIlatorla  I imnetica  (Cyanophyta)   were  numerically



dominant in both 1983 and 1984.



    Of the 1983 and 1984 dominant diatoms, only Frag 11 aria crotonensis and



perhaps TabelI aria  fIoccuIosa were  the major  components of  the  diatom



assemblage In 1962-63.   Stoermer and  Kopczynska (1967a) noted  taxonomic



difficulties with TabelI aria and noted that most populations of TabelI aria



"are probably to be referred to J.  fenestrata ...."



      The dominant species of CyclotelI a  in 1962-63 was £.  mlchIganiana.



Rockwell et al. (1980) reported that  Cyclotella spp. were common In  1977



but were  never  dominant.     A  dramatic  decrease   in  some  species  of



Cyclotella, such as £. mlchiganlana and £. stelIigera,  which were offshore



dominants In August of 1970,  was evident (Table 13).  CyclotelI a cgmensis.

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                                                                      31




believed to be tolerant of higher nutrient and lower silica concentrations



than most members of  thfs genus, was the  numerically dominant diatom  In



the offshore waters.   In  1984,  however, Cyclotella  oceI Iata, a  species



generally associated with oligotrophlc conditions,  was also dominant.



      Yearly variation  in dominance of species  of Melosira was  evident.



Melosira Island lea was  dominant in  1962-63.   In 1983  M.  islandlca  was



present (mean = 12.1 cells/ml), but  E.  itallea subsp. subarctica (mean  =



37.6 cells/ml) was  more abundant.    In 1984  .M..  Islandlca and M>  itallea



subsp. 5ubart?ca  had similar  abundances  (-10-12 cells/mL)  (Table  10).



Similarly,  Synedra acus was common  throughout the southern basin in  1977



(Rockwell et al.   1980) but  in 1983 represented  only 0.1?  of the  total




eel Is,



      Makarewicz (1987) has suggested an  apparent decline in B.  eriensls



since 1962.  In  May of 1962, relatively  high (100 cells/mL)  populations



were observed in southern Lake  Michigan (Stoermer and Kopczynska  1967a).



During May and June of 1970, mean abundances for offshore stations were 63



and 611 cells/mL, respectively (Holland and Beeton 1972).  Rockwell  et al.



(1980) reported a  mean density of  28.7 cells/mL for  R. eriensls  during



June of 1977.  Abundance in 1983 was 2.6 cells/ml for the entire lake.   A



bloom (133 cells/mL) in the northern Station 77 did occur In October.    In



1984-85, mean  lake abundance  increased  to 18.2  cells/mL.   Similar  to



species  of  Meloslraf   considerable  yearly   variation  in  abundance  of



Rhlzpsolenla from 1983 to 1984 was observed.



       Anklstrodesmus  falcatus  Increased  In abundance  to 1977  and  had



decreased by 1983.   Ahlstrom (1936)  reported this species  as rare,   but



Stoermer and Kopczynska  (1967a) noted  that it had   Increased by  1962-63



(range = 20-60 cells/mL).  Rockwell  et al. (1980) suggested that by  1977

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                                                                      32





it had increased further (range = 20-160 cells/ml).  In 1983 this  species



was observed only  once during  the study  at Station  32 (6.5  cells/ml).



This species was not observed in 1984.



        Dominant  chrysophytes in  1962-63  were Dinobryon  divergeng,  J3.



cylindricum and Q. soc i aIe (Stoermer  and Kopczynska 1967a).  Rockwell  et



al. (1980) reported them  as dominant or subdominant  offshore.  With  the



exception of £. cyl indricum In 1984,   I>. divergens, ]).  cyl indricum and  D.



soc?aIe were common species  in 1983 and 1984.   However, the  haptophytes



were numerically the dominant  group within the  chrysophytes  in 1983  and



1984.



      Dominant cryptophytes  in 1983 and  1984 included Cryptomonas  erosa



var.  refIexa,  £.  _e_co_sfl  and  Rhodomonas  minuta  var»   nannoplanktica.



Stoermer and Kopczynska (1967b) and Stoermer (1978) reported these species



as uncommon in Lake Michigan, but  Vollenweider et al.  (1974) noted  these



species as commonly found.  Similarly, Munawar and Munawar (1975), Claflin



(1975) and Rockwell et al. (1980) had reported £. .exQSa and £. minuta var.



nannopIanktica to be dominant, abundant and perhaps increasing in  number.



From the 1983 and 1984 study, It is apparent that £. erosa was numerically



uncommon but on a biomass basis was the second most Important  cryptophyte



(Table 10).  Evaluation of abundance  of £. minuta  In earlier studies  was



not possible because it was grouped into phytoflagellates, flagellates  or



simply Rhodomonas.   What  can be  reported about  Rhodomonas minuta  var.



nannoplanktica is that in 1983 and 1984 It was the dominant cryptophyte on




a  numerical basis.



        Oscillator I a Itmnetlca  has become  more  prevalent  In  the   lake.



Ahlstrom (1936) and Stoermer and Kopczynska (1967a) listed £. mougeot11 as



the only species of  this genus abundant In  their collections.   Stoermer

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                                                                      33


and Ladewski (1976) reported that Q. I fmnetTea had generally Increased  In


abundance in  Lake Michigan.   Rockwell  et al.  (1980) observed  that  Q.

11mnet1ca was  common throughout  the  basin In  April  and June  and  was


especially abundant  in  September  of  1977 at  certain  stations.    Not


considering the pI cop Iankton, which were not counted  In previous  studies,


Q. I 1mnetic§ was  the numerically  dominant offshore  blue-green algae  In


1983 (Makarewicz 1987)  and was second  In abundance  in  1984 (Table  10).


Anacystls montana var.   minor was  the dominant blue-green  algae In  1984


(Table 10).



PJco.plan.Kt.Q-n
                                                                         4
     Picoplankton abundance  In  1984 (mean = 18,409; maximum of 4.3 x  10

                                                                         5
cells/ml) was not dissimilar from 1983 (mean = 23,607; maximum of 1 x  10


cells/ml).  On a  numerical basis, the  plcoplankton represented 82.856  of


the total cells in 1984.   Their dominance of the phytop Iankton  community


in 1984 was comparable to that In  1983 (89.4? of total cells).  Prior  to


the 1983 study (Makarewicz  1987), no other  researchers on Lake  Michigan


have  routinely  reported  this  group  of  organisms.    Because  of  the


overwhelming dominance  of this  group, analysis  and discussion  of  this

group would be  facilitated with  verification of the  systematics of  the

spheres (Anacystls  marina?),  rods (Coccochloris  pentocystis?)  and  the


spherical-fI ageIlates.



Geographical Abundance and Distribution


      Average phytopIankton abundance  for the non-winter sampling  period


generally decreased from the north (Station 77) to the south at Station 57


(Fig.  12).   Overall   abundance  remained roughly  the same  southward  to


Station 18.  At the most southerly sampling station (Station 6), abundance

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                                                                      34




was higher than  In the  rest of  the lake  except for  the most  northern



stations (Station 77  and 64).   Thfs pattern is  not dissimilar from  the



geographical pattern observed in 1983 for Lake Michigan (Makarewicz 1987).



This abundance  pattern  is attributed  mainly  to the  picoplankton,   the



BaciIlariophyta and the Cyanophyta which all  have higher abundances at the



northern stations.   The  higher abundance at Station  6 was caused by  the



picoplankton.  The peak in abundance  of the Cryptophyta at Station 41   in



1983 (Makarewicz 1987) was again observed in 1984 but was not as prominent



as  in 1983.  A peak in Chlorophyta at Station 41  was not observed in  1984



as  it was  In 1983.    Cyanophyta were in  higher abundance at the  northern



stations (77 and 64) and at Stations 41-27 (Fig.  12).



      Seasonally, the spring and  autumn cruises possessed a  geographical



abundance pattern similar  to the mean  annual phytopIankton  distribution



with abundance peaks  at the northern  (Stations 64 and  77) and  southern



(Station 6) stations (Fig.  13).   Abundances of BaciIlariophyta,  Cyanophyta



and picoplankton peaked  at these stations.    Only on  the 27-29  November



cruise did a  maximum in abundance  not occur  at Station 6.  The  summer



cruises did  not  display  the distinctive  northern  and  southern  peaks



observed in the spring and autumn of 1984.  Similar geographical peaks  in



abundance were  observed at  the northern  and southern  stations In  1983



(Makarewicz 1987).




        Interestingly, many  of  the  same  species had  distinctly  higher



abundances in  1983  and  1984  at the  northern  and  southern   stations.



TabeI Iar ja  f locculosa,   Fragt I aria   crotonensls,  Cyclotel la   comensis,.



Coelasphaerturn naegellanum  and  picoplankton  were  more abundant  at  the



northern stations than  in the  rest of  the lake in  1983 and  1984.    In



addition, Oocystis submarine was abundant in 1984, while Cyciotella comta,

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                                                                      35





Chroomonas norstedt I i and OsciI later I a agardh i i  had a greater abundance at



the northern stations 64 and  77 In 1983.   Except for £. comensis,,   whose



ecological affinities  are poorly  known, the  other diatom  species  more



prevalent at Stations 64 and 77 are generally associated with  mesotrophic



conditions.



      Besides picoplankton, the abundance peak  at Station 6 in both  1983



and 1984  was  attributed to  DInobryon  soc?aIe var.  americanum  and  J).



divergens.  Species of haptophytes prevalent at Station 6 in 1983 were not



prevalent in 1984.



     The northern stations 64  (depth = 25m) and 77 (depth = 23m) and  the



southern station 5/6 (mean  depth = 50m) are  best described as  nearshore



stations Cdepths  are less  than or  equal to  50m (Bartone  and  Schelske



1982)3.  The physical   and chemical characteristics  of the nearshore  and



the Straits of Mackfnac stations  differ significantly from the open  lake



stations  (Bartone  and   Schelske  1982).     Thus  the  differences   In



phytoplankton abundances observed at the northern and southern stations in



this study should be related to known differences in water quality.   There



is some evidence  to support this  hypothesis.   A  comparison of  nutrient



data from  the  nearshore  and  offshore  stations  indicates  that   total



phosphorus was higher at  Station 64 and silica  was higher at Station  77



compared to the  rest of  the lake  (Table 14).    Station  6 had  nutrient



levels similar to the rest of the lake.



     Temperature may also be a factor in the occurrence of the  geographic



abundance peaks  observed.    For example,  on  6  and 7  May  the  higher



temperatures and abundances at Stations 6, 64 and 77 correlate well   (Fig.



13).

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                                                                      36





 Indicator Species



      Stoermer  and Yang (1970),   in a comparison  of modern and  historic



records, reported that  taxa characteristic of  disturbed situations  were



rapidly Increasing in relative abundance in Lake Michigan in the 60's.  In



the nearshore area, a shift In oligotrophlc forms to those which  dominate



under eutrophlc conditions was evident.   Occurrence of certain  eutrophlc



species was also evident In offshore waters (Stoermer and Yang 1970).



        Dominant  diatom  species In  the  offshore waters  In  1983  were



Cyclotel la  comensIs,   £•   comtar   Tabel I aria   f locculosa.    Frag 11 aria



crotonensis and Meloslpa .Itai.l.ca subsp. subartlca.  The same five  diatoms



were dominant In 1984 with the exception  of £. £Qffi±fl and the addition  of



Rhtzosolenla erlens Is and  Cyclotella ocellatg  (Table 10).    In fact,  £.



erlensls accounted for ~25% of  the total blomass of phytoplankton  during



1984.



     Rhlzosolenlg erlensls  may be an opportunistic species which is  able



to rapidly develop  fairly high abundances  when conditions are  favorable



(Stoermer and LadewskI 1976).   Stoermer and Yang (1970) listed £. erlensls



with the ollgotrophlc offshore dominants, which includes £.  QgeUataf   but



noted that B« erlensls seemed to occur In greater abundance In areas  that



have received some degree of  nutrient enrichment.  TabelI aria  fIoccuIosa



and £. crotonensis are mesotrophic forms, while the ecological  affinities



of £. comens1s are  poorly understood.   CyclotelI a comensIs was  formerly



found in primarily ollgotrophlc areas (Stoermer and Stevenson 1979)  under



some nutrient stress (Stoermer  and Tuchman 1979).   Compared to the  1983



cruises (Makarewlcz 1987), where  mesotrophic forms were predominant,   the



same mesotrophic  forms  were  present In  1984  along  with  ollgotrophlc



indicators.

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                                                                      37




      The indicator diatom species and  the distribution of them  (trophic



ratio) (Table 15) suggest a eutrophic status for nearshore waters In 1977,



while the  offshore waters  in 1970-71,   1983  and 1984  would be  in  the



oligotrophic-mesotrophic range.  With the high mesotrophic/eutrophlc ratio



in 1970-71  (M/E =  8) as  compared to  1983 and  1984 (M/E  = 4),   It  is



tempting to  suggest a  slightly more  mesotrophtc status  in more  recent



years.  At best, this observation has to be viewed with caution since only



one species difference is  required to achieve the  observed change.    The



M/E ratio  has  to  be  interpreted conservatively  as  it  is  Influenced



somewhat by the definition  of the dominant species  (e.g. 1$ of  biomass)



utilizied.  Nevertheless,   the trophic status  as determined by  Indicator



species and the M/E  ratio agrees well with  the 1976 assessment based  on



particulate phosphorus concentrations that place  the open lake waters  of



Lake Michigan in the oligotrophic-mesotrophic range (Bartone and  Schelske



1982).






Histprical Changes In Community Abundance



       A  comparison of  abundance trends  over the  entire lake  was  not



possible because of the  lack of comparable offshore  data prior to  1983.



Figure 14  plots  the  1962-63  and  the  1976-77  data  of  Stoermer  and



Kopczynska  (1967a  and  b)  and   Rockwell  et  al.  (1980),   which   are



representative of  the southern  portion of  the lake.    Only  a range  of



abundance is available  for 1962-63,  while the mean,   standard error  and



range are  plotted for  the other  data.   Because plcoplankton  were  not



counted prior  to 1983,  they are  removed  from the  1983 and  1984  data



presented In Figure 14.  Although a mean is not available, it is  apparent



that abundance increasd from  1962-63 to 1976-77.   From 1976 to 1983  and



1984,  abundance was not  significantly different (P=0.05).   Based on  the

-------
                                                                      38



classification scheme of  Munawar and Munawar  (1982),  which utilizes  the



mean phytopIankton  biomass  as  an  Indicator  of  trophic  status,   Lake



Michigan would be classified as ollgotrophtc in 1984.   This designation Is



supported by  the trophic ratio and composition of indicator species.

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                                                                       39





LAKE MICHIGAN



ZoopIankton



Annual Abundance of ZoopIankton Groups



     Species lists (Table  A4) and summary tables of abundance (Table  A5)



and biomass (Table A6)  are in Volume  2 - Data  Report.  The  zoopIankton



assemblage of 1984 comprised  52 species representing  34 genera from  the



Calanoida, Cladocera, Cyclopotda,  Harpacticoida, Mysidacea and  Rot ifera.



Compared to  1983, reductions  of 21%  and 24%   in numbers  of genera  and



species,  respectively,  were  observed.     This  difference  is   mostly



attributable to a decrease in number of Cladocera and Rot ifera.



       The  Rot ifera  possessed the  largest number  of species  (29)  and



relative abundance (67.5?)  followed by the  Cladocera (10 species)  which



accounted for 39.8$ of the zoopIankton  biomass  (Table 16).  The  Rot ifera



contributed only 2.6% of  the total biomass (Table  16).  Average  density



and biomass for the study  period was 59,764±8,284 organisms/m  (mean±S.E.)



(1983 = 69,353) and 33.2+4.9 mg/m3 (mean±S.E.) (Table 6).





Seasonal Abundance and Distribution of Major ZoopIankton Groups



     The seasonal  abundance and biomass pattern were virtually  identical



(Fig.  15) with  a maximum  in August.  The secondary  maximum observed  in



October of 1983 (Makarewicz  1987) was not observed  in 1984.  This  major



difference between 1983 and  1984 is apparent and  is probably related  to



the difference in the  seasonal  samp I ing pattern  between years.    Samples



were not  taken in  September and  October of   1984.    A sampling  pattern



including the June-July and September-October period is required to  fully



evaluate the differences in the seasonal distribution pattern.



     Seasonally,  abundance and biomass of all  groups were higher in August



as compared to the early spring and late fall  (Figs.  16 and 17).   In  1983

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                                                                      40





a peak in rotifer abundance  occurred In October (Makarewicz  1987),   which



was not observed  In 1984  due to  a lack of  October samples.    The  high



abundance of  Cyclopolda, Calanoida,  Cladocera  and Copepoda  nauplii  in



August of 1984 was not observed in 1983 (Makarewicz 1987).






Geographical Abundance and Distribution of Zooplankton Groups



      A definite trend of  increasing zooplankton abundance occurred  from



south to north In Lake Michigan  (Fig.  18).  Zooplankton abundance at  the



far northern Stations 64 and 77 was  higher than in the rest  of the  lake.



Abundances of Rotifera, Cladocera and Copepoda nauplii were all higher  at



these far northern stations.  Biomass,  however, was similar southward from



Station 77 to Station  18, after which  biomass decreases southward  (Fig.



19).  These patterns were not observed In 1983 (Makarewicz 1987).



     Abundance of  Diaptomus sic!I is was higher In southern Lake  Michigan



(Makarewicz 1987) in  1983.  However,  a similar pattern  was not  evident



(Fig. 20) in 1984.  Copepodites  of Diaptomug averaged a higher  abundance



in southern  Michigan.   Similar to  1983 was  the increase  in  Cladocera



abundance at the far  northern stations (Fig. 19).   Abundance of  Posmipa



jongirostris dramatically increased  at these  stations In  1983 and  1984



(Fig. 21).  Also, Eubosmina cgregoni, Notholca laurentiaef  &. squamula, M.



foliacea and  HoI opediurn  gibberurn a I I   had  abundance peaks  at  the  far



northern end of the lake  In 1983 and 1984 (Fig. 22).  Polyarthra  vuigaris




and £. remata had higher abundances  in 1984 only at the northern  Stations



64 and 77 (Fig. 22).






Common Species



     Common Crustacea species  (Table 17) were arbitrarily defined as those



possessing  a relative abundance of >0.1$ of the total abundance or 1.0$ of

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                                                                     41




the total  biomass,   Rotifera  species  were considered  common  if  they



accounted for >1.0/£ of  the total  zooplankton abundance  or biomass.   The



number of common species (1983 = 25 species; 1984 = 24 species)  and common



species composition  were  essentially the  same  between 1983  and  1984.



Rotifer composition  differed with  NothoIca fo1iaceaf  &. laurentiae  and



Polyarthra  remata  being  common  In  1984  only,  while  £.   major,   £.



doIIchoptera, Keratella crassa and j£. earlinea were common in  1983.






Historic Changes in Species Composition






Crustacea



     Numerous recent  studies (Williams 1966; Johnson 1972; Gannon et  al.



1982a, 1982b; Evans et al.  1980) of the nearshore region of Lake  Michigan



exist, along with  data from as  far back  as 1927 (Eddy  1927).   Several



researchers have compared the nearshore  with the offshore zooplankton  In



discussions of  eutrophication of  the entire  lake.    Comparisons of  the



inshore with the offshore stations  should be viewed with caution  because



effects are not necessarily due to eutrophfeat ion or fish predation (Evans



et al. 1980).



     Although no  intensive zooplankton studies of the offshore waters of



the entire lake  basin have  taken place,   some offshore  studies of  Lake



Michigan zooplankton do exist.  Wells (1960, 1970) sampled Crustacea  with



a number 2 (366um)   net on four  dates in June,  July  and August in  1954,



1966 and 1968 from the offshore region off Grand Haven,  Michigan.  On  six



dates (March 1969  to January 1970),   Gannon (1975) collected   crustaceans



with a 64-um mesh net from the offshore and Inshore of Lake Michigan along



a cross-lake transect from Milwaukee to Ludington.  In September of  1973,



northern Lake Michigan was sampled with a 250-um mesh net (Schelske et al.

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                                                                      42




1976).  Also,  Stemberger and  Evans (1984) provided abundance data  (76-um




net) for a few zooplankters from offshore waters of the southeastern  Lake



Michigan area.




     The data of Wells (1960,  1970) and Gannon (1975) are useful but  have



to be  used  with  caution.   A  366-um  and a  250-um  net  are  probably



quantitative for larger crustaceans but certainly would not be for smaller



crustaceans such as Chydorus  sphaericus, Bosmina longirostris,.  Eubosmina



coregon i f  Ceriodaphnia   spp.,    Tropocyclops   prasinus   and   copepods



(Makarewicz and Likens 1979).



       The  zooplankton populations  in Lake  Michigan underwent  striking



size-related changes between  1954 and  1966 (Wells 1970).   Species  that



declined sharply were the largest cladocerans (Leptodora kindti?,  Daphnia



galeata  mendotae  and  Q..  retrocurva),  the  largest  calanoid  copepods



(L imnocgl anus macrurusj. Epischura Iacustris and Diaptomus sici I is) and the



largest cyclopoid  copepod  (Mesocyclops  edax).   Medium-sized  or  small



species  (£.  longiremisr  JU.   gibberum,  Polyphemus  pediculus,   Bosmiaa



longirostriSf  Ceriodaphn ia  sp., Cyclops  bicuspidatus, Cyclops  vernal \.sr



D i aptomu s ash I an d i) increased in number, probably in response to selective



alewife predation.   After the  alewife dieback,  J^. eda?<  and D.  galeata



mendotae were still rare  in 1968  when the composition of the  zooplankton



community shifted back toward one similar of 1954 (Wells 1970).



        In northern  Lake Michigan  during September  of 1973,  predominant



species  were  Daphni a  galeata  mendotae,  Q.  retrocurvaf   L imnocalanus



macrurusf Pi aptomu s oregonensis,. Eubosmi na coregon i and Diaptomus sici I is.



Cyclopoid copepods were a minor component  of the fauna in 1973  (Schelske



et al. 1976).

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                                                                       43




     The changing nature of the zooplankton community of Lake Michigan was



evident in 1983. Daphnia gal eata mendotae,  D.  pulicaria and D.  retrocurva



were the second, third  and fourth most abundant  cladocerans in the  lake



(Makarewicz 1987).  Q. galeata mendotae  and D.  retrocurva were again  the



prominent daphnids  in 1984  along with  the dominant  cladoceran  Rosmina



longirostris.  Abundance of Daphnia  pul icaria dropped from an average  of



376/m3 in 1983  to 78/m3 in  1984.  In  August of  1983,  abundances of  JD.



galeata.  rare in 1966 and 1968, were half of those in 1954 (1,200/m3)  and



three times the 1954 abundance In 1984 (Table 18).



      The 1983 abundance of Daphnia  retrocurva was similar to the  August



1966 abundance rather  than to those  of 1954 or  1968.  However,  maximum



abundance in October of 1983 (3,161/m ) was comparable to the 1954 or 1968



observations.  Perhaps related  to the low abundance  of D. retrocurva  in



August of  1983 was  the appearance  of  the large  (-*2 mm)  (Evans  1985)



cladoceran Paphnia pul icaria,.  which reached a maximum abundance  In August.



When .Q. pulicaria dropped  in abundance in  1984,  £.  retrocurva  abundance



increased to a density comparable to those of 1954 and 1968 (Table 17).



     Evans (1985)  recently reported that Q. pulicaria was first  observed



in Lake Michigan  in 1978.    Abundance remained low  in southeastern  Lake



Michigan until 1982  and 1983   when  they dominated  the offshore  summer



Daphnia community and  at an  offshore station southeast  of Grand  Haven,



Michigan.    In  1983 this  species  was  the dominant  cladoceran  in  the



offshore waters of  Lake Michigan  from the short   and long  hauls.   Mean



station abundance  reached  1,741  organisms/m  in  early  August  with  a



maximum of 6,056/m .  In 1984, abundance of D. pulicaria dropped to a mean



of 248/m3 from 1011/m3 in 1983 (Table 18).

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                                                                      44




     The occurrence of Daphnia .d_ub_La, a new species observed in 1983,   was



not confirmed in 1984.   In a review of the  1983 material  by a  different



taxonomist, this species was not observed.



        The large  cladoceran Leptodora  k indti i   appears to  be  steadily



increasing in abundance since 1954 in Lake Michigan (Table 18).  Eubosmlna



coregon i f B.  longirostrig  and the   larger Ho I oped 1 urn  gibberum have  also



increased  in abundance since 1954 (Table 18).  The increase in U.  gibberum



was probably real.  It is  doubtful  that this large cladoceran would  pass



through a 366-um mesh net like that used in Wells (1960,  1970) studies  of



1954-68.    However,  the  net  employed by.  Wells'  would  not  have  been



quantitative for £. coregon i and £.   longirostris.



    Cyclops bicuspidatus was the dominant cyclopoid in 1983 and 1984  with



Diaptomus ash I andj being the dominant  calanoid  in 1983 (Makarewicz  1987)



and £). siciI is in 1984 (Table 17).  Abundance of  Mesocyclops edax was  low



in August  of 1983  and 1984  compared  to 1954,   but abundance  in  early



October of  1983  reached  a comparable  151  organisms/m   (mean  station



abundance).   Diaptomus minutus  appears to  have decreased  in  abundance



since 1968, while JD. oregonensis abundance remained similar to 1954 (Table



19).   D.  siciI is  has  increased steadily  since  1968.    Abundance  of



Limnocalanus macrurus was  lower during  August of 1983  than  in  1954-68.



However,  abundance  in 1984 was similar to 1954 and 1966.   The abundance of



Epischura  lacustris in August was still  low in  1983 and 1984 relative  to




1954, but  reached   111  organisms/m  (mean  station abundance)  in  late



Octoberof  1984.



      By   1983 and  1984,  the large cladocerans,   calanoids and  cyclopoid



copepods, observed  by Wells (1970) to have decreased sharply in the  early



60's, had  increased in abundance to  densities similar to those in  August

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                                                                       45





of 1954.  In some instances, abundance  was not as high in August but  was



as high at other times of the year.  In addition,  a new large  cladoceran,



Daphnia pulicaria, has become estabiished  in the offshore waters of  Lake



Michigan.



      The resurgence  of larger zooplankton in  Lake Michigan is  probably



related to the sharp decline in the abundance of the planktivorous alewife



in 1982 and 1983.   The lakewide catch of adult  alewifes was only 31%  of



that of 1982 and only 12$ of the 1981  catch.   Bloater chubs are  replacing



the alewifes and have been  experiencing a dramatic increase in  abundance



since 1970  (Wells  and  Hatch 1983).     Bloaters  above ^18  cm  in  size



primarily feed on _My_s_Ls and Pontoporeia.  Only smaller individuals feed on



zooplankton (Wells and Beeton 1963).






Rot ifera



      Rotifer studies  reported in the literature  are primarily from  the



nearshore region of  the lake.    In the  nearshore, KeratelI a  cochlearis,



Polyarthra  vulgar is,  KeI Itcott i a   longispina,  Synchaeta  sty I ata   and



gynchaeta tremuI a were  dominant in  1926-27  (Eddy 1927).    Keratella  and



Polyarthra were  the dominant  genera in  1962 (Williams  1966), while  £.



cochlearis and £. vulgar is were dominant  in  1970 (Johnson 1972).    Gannon



et al. (1982a) noted that the following rotifers were predominant in 1977:



Keratella  cochlear?sf  JK.    crassaf   Conochi1 us  unicorn is,    KeI I? cott i a



longispina,  Polyarthra vulgaris and £.  remata.



     Abundance of  rotifers in Lake Michigan  generally decreased from  the



nearshore into the  offshore (Gannon   et al.  1982a,  Stemberger and  Evans



1984) although the species composition  of the nearshore and offshore  was



relatively similar.    In 1983  the predominant offshore  rotifers were  in



descending  order:   Polyarthra   vulgar is,    Synchaeta   sp.,    KeratelI a

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                                                                      46




cochI earisf Polyarthra  major,  KeI Ii cott i a  longisp Fnaf  Keratella  crassaf




Gastropus sty lifer and Colletheca sp.  (Makarewicz 1987).  The  predominant



rotifers   in  1984  were  KeratelI a  cochI ear\st   KeI Ii cott i a  long!spinaf



Polyarthra vulgar is and Synchaeta sp.  (Table 17).  Both  the 1983 and  1984



dominant rotifer composition is similar to the nearshore and  to Ahlstrom's



(1936) offshore observations of predominant species (Keratella cochlearis,



gynchaeta sty I ata and Polyarthra vulgaris).






Historical Changes in Zooplankton Biomass




     Offshore crustacean  zooplankton biomass data is available from  1976



(Bartone and Schelske 1982) for northern Lake Michigan.   No information is



presented on  sampling intensity  or technique.   A  comparison with  1984



(Table 20) reveals  that no significant  difference in crustacean  biomass



exists between 1976 and 1984.



     Another longer sequence of data is described by Scavia et al. (1986).



Except for 1977, 1982, 1983  and 1984,  zooplankton samples were  primarily



from an  offshore station  (40-m depth)  west  of Benton  Harbor, Ml.    A



comparison of the mean  offshore 1984 lake-wide  biomass data to  Scavia's



station indicates good agreement (Fig.  23). From Figure  23,  there appears



to be no obvious trends in zooplankton biomass.





Indicators of Trophic Status



       Zooplankton have  potential  value  as assessors  of trophic  status



(Gannon and  Stemberger  1978).   Rotifers,  in particular,  respond  more



quickly to  environmental changes  than do  the crustacean  plankton  and,



therefore, are  more sensitive  indicators of  changes  in  water  quality.



Composition of the rotifer community  (Gannon and Stemberger 1978) can  be



used to evaluate trophic status.

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                                                                      47





      In 1983 the six predominant rotifers in descending order of  relative



abundance were £.  vulgar is, Synchaeta  sp., J<. cochI ear is,  £. major,  K.



longlspina and C. unicornis, while  in 1984 the predominant rotifers  were



J<. cpchlearis, J£. longisp inaf £. vulgaris and Synchaeta sp.  The 1983  and



1984  rotifer composition suggests an oligotrophic association.  A  rotifer



community  dominated   by  Polyarthra   vulgaris,  Keratella   cochI ear is,



ConochiI us unicornis and Ke|Iicottia longisp ina has been considered to  be



an association   indicative  of an  oligotrophic  community by  Gannon  and



Stemberger (1978).



      The high  relative abundance of  Diaptomus slciIis and  Limnocalanus



macrurus (Table  17)  and  the occurrence  of  Senecel I a  cal anoides,  al I



oligotrophic  indicators  (Gannon  and  Stemberger 1978,  McNaught  et  al.



1980), also  suggested oligotrophic  offshore  conditions for  the  entire



lake.



       The calanoid/cyclopoid  plus cladoceran ratio  has been  used as  a



measure of trophic status in the Great Lakes (Gannon and Stemberger  1978,



McNaught et al. 1980).   Calanoid copepods  generally  appear best  adapted



for oligotrophic  waters,  while  cladocerans and  cylopoid  copepods  are



relatively  more  abundant   in  eutrophic  waters.    On  the  north-south



transect, the plankton  ratios were high  and similar,  except  at the  far



north and the southern extreme of the  lake  (Table 21).  This pattern  was



repeated in 1984 and suggests that a lower quality of  water occurred south



of Station 18 and  north of Station 57.   The eutrophic rotifer  indicator



species  Trichocerca  pusi I  I a  was  observed  exclusively  at  Station  6,



reinforcing the  Idea that  a lower  water quality  exists at  Station  6.



Similarly,  Trichocerca multicrinis,  a  eutrophic  indicator, was  prevalent



at northern stations.

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                                                                   48
     The  low plankton ratios (0.20 - .41;   Table 21) at the far  northern



end of  Lake Michigan  (Stations 64  and 77)  were very  similar to  those




observed in 1973 at the Straits of Mackinac (Gannon and Stemberger  1978).



Gannon and Stemberger (1978) implied that more eutrophic conditions  exist



within this area  of a low  calanoid to cladoceran  plus cyclopoid  ratio.



Abundance of the ol igotropic  L imnocalanus macrurus and Diaptomus  siciI is



was significantly  lower  in these far  northern stations,  while  Eubosmina



coregoni and Bosmina long?rostris,  often associated with more  productive



conditions,  increased  at  the  far  northern  stations  (Fig.  21).     In



addition, several mesotrophic algal species  were more predominant at  the



northern stations.



      Notholea fol iacea is  often associated with ol igotrophic  conditions



(Gannon and Stemberger 1978).   In this study, several indicators  suggest



that the northern end  of Lake Michigan near  the Straits of Mackinac  has



waters associated  with  more  productive  conditions.  Yet  abundance  of



Nptholca fol iacea  increased at  the northern  stations.   The use  of  U.



fol iacea as an ol igotrophic  indicator has to be viewed with caution.



      With a zooplankton abundance between  those of Lakes Erie and  Huron



(Table 6), the presence of an oligotrophic rotifer association, a plankton



ratio between those of Huron and Erie,  the domination of the calanoids and



the  fairly  abundant  presence  of  the  ol igotrophic  indicator  species



Diaptomus sfciI  is  and L imnocalanus macrurus,  the offshore waters of  Lake



Michigan  in 1984   are best characterized  as mesotrophic/oligotrophic.   A



similar conclusion utilizing zooplankton abundance and species composition



was drawn   in 1983.   Phytoplankton  composition and  abundance and  water



chemistry  suggest a similar trophic status (This Study).

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                                                                  49






Trophic Interactions



      Between 1975 and 1984, gradual   declines In spring total  phosphorus



and summer ep I limnetic chlorophyll  .a are reported  (Scavia et al.  1986).



However, long-term changes  of phytopl ankton and  zooplankton biomass  are



not apparent in  this study.   Perhaps,   the minimal  changes observed  in



chlorophyll a are not reflected in the high variability phytoplankton  and



zooplankton estimates.  Scavia et al.  (1986) points out that the  changes



in total phosphorus and chlorophyll .a are consistent with expectations  of




nutrient load control.



        However,  the significant  lake-wide  changes in  zooplankton  and



phytoplankton composition may not  be expected from  nutrient control.   A



species new to the plankton assemblage,   Daphnia pul icaria, is at least  a



sub-dominant  organism  within  the  offshore.    In  addition,   Leptodora



kindti I,  Daphnia  galeata  mendotaef   Diaptomus  ashjajidl  and   Cyclops



             have returned  to and  exceeded abundances  observed in  1954
during a period of low alewife abundance.



      Scavia et al. (1986) suggests  that predatory pressure from  alewife



suppressed large-bodied zooplankton  until the early  1980's.  Decline  of



the alewife population  as the  major forage  fish (Jude  and Tesar  1985,



Wells and Hatch  1983) has  been linked  to the  increasing population  of



stocked salmonines in Lake Michigan (Stewart  et al. 1981, Jude and  Tesar



1985).   The  decrease In  alewife  abundance has  reduced  size-selective



predation on  larger zooplankton  allowing  larger zooplankton  to  return



(Scavfa et al. 1986,  Wells 1970, Kltchell and Carpenter 1986).



      Table 22 lists  correlation coefficients of phytoplankton  abundance



versus total  phosphorus and zooplankton for each cruise.  For each cruise,



11 stations covering  the entire length  of the lake  were sampled over  a

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                                                                   50




short period  of  time.    Interpretations of  the  correlations  were  as



follows:    A  negative  correlation  between  a  zooplankton  group   and



phytoplankton implied grazing pressure on phytoplankton, while a  positive



correlation between  total phosphorus  and phytoplankton  abundance  would



suggest an  enhancement  of  phytoplankton  abundance  due  to  phosphorus



availability.  Except for  the early winter  cruise, correlation of  total



phosphorus to phytoplankton abundance  was weak as  compared to Lake  Erie



(Table 22).  Grazing pressure appeared to be particularly heavy during the



May series of samples.



     As suggested by Scavia et al.  (1986), .Q. pulicaria appears to have  a



negative impact on phytoplankton  abundance expecially during  mid-August.



Interestingly, when Daphnia galeata mendotae  is added to the  correlation



analysis,   the  correlation  coefficient  increases  from  -.27  to   -.50



suggesting that  Q. galeata  mendotae is  also having  a major  effect  on



phytoplankton abundance during August.  This  would be an added effect  in



that D.   galeata  has  increased  since 1954  apparently  in  response  to



decreased selective  pressure by  the alewife.   The  calanoids appear  to



exert grazing pressure in  the spring and early  winter as opposed to  the



summer (Table 22).



     The causes of the changes in species composition of phytoplankton are



much more difficult to evaluate.   Changes in herbivore species composition



could affect algal  species composition.    Certain zooplankton  feed on  a



wide variety of algae of different  sizes and shapes,  and with or  without



sheaths (Gliwicz 1980,  McNaught et  al.  1980b,  Porter  and Orcutt  1980).



Other zooplankton  are  highly  selective in  the  algal  types  ingested.



Cellular forms are  ingested more  readily than  filamentous or  spinuosus



forms and  zoopIanktonic filtration  rates,   growth and  survivorship   are

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                                                                   51   ,




greater when  feeding  on  cellular  forms  (Porter  1973,  Arnold   1971).



Selective grazing and utilization can remove species or reduce  population



size  in  the algal  community.   Alternatively, grazer  utilization of  an



algal species can result   in enhancement of  primary productivity of  that



species by  increased selection for faster growing genotypes (Crumpton  and



Wetzel 1982).



        In  summary,  zooplankton  community  structure   is  important  in



determining the responses  of algal assemblages  to grazing (Bergquist  et



al. 1985). Small  algal taxa increase  in abundance when  grazed by  small



zooplankton, but decrease  in density  when grazed  by large  zooplankton.



Conversely, large phytoplankton  become less abundant  in the presence  of



small zooplankton,  but  increase in  density  in the  presence  of  large



zooplankton (Bergquist et al. 1985).  Perhaps the increase in abundance of



the large d i atom Rh i zosoI en i a spp. during the summer of 1984 is related to



increased grazing pressure of large Daphnia



    Nutrient  effects can also affect  composition of phytoplankton.   For



example,  Asterione I  I a  is a  successful  competitor  at high  Si/P  ratios,



FragiI I art a can dominate at  intermediate ratios and Stephanodiscus  grows



well when Si/P ratios are low  (KM ham and KM ham 1978j Kilham and  Til man



1979; Tilman 1978,  1980).   At high Si/P  ratios,  diatoms can  effectively



out compete blue-green  algae (Holm  and Armstrong 1981).    Similarly,   as



silica is reduced and combined nitrogen declines,  green algae can  compete



less effectively with nitrogen-fixing  blue-greens (Smith 1983).   Effects



on phytoplankton composition  from both  top-down and bottom-up routes  are



expected but are difficult to separate in this  descriptive study.

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                                                                   52





LAKE HURON



PhytopIankton



     Species lists (Table  A7)  and summary tables of abundance (Table  A8)



and biovolume (Table  A9) are in  Volume 2 -  Data Report.    A summary   of



water chemistry parameters is presented In Table 6.






Annual Abundance of Major Algal  Groups



      The phytoplankton  assemblage of 1984 was  comprised  of 315   species



representing 92 genera from eight divisions (Table 23).   Compared  to 1983,



a 4.3$ reduction  in the  number of  species and  a 4.5%  increase in  the'



number of genera were observed.



      The annual average phytoplankton  density and biovolume were  17,209



cells/ml (19,147  cells/ml;  1983)  and 0.38  mm3/L  (0.42   mm3/L;  1983),



respectively.  Similar to 1983,  the BaciIlariophyta possessed the   largest



number of species  (156) and  biovolume (61.9$  of the  total, Table  24),



while the  second largest  number of  species (64)  was observed  for  the



Chlorophyta  (Table  24).    Although   the  relative  biovolume  of   the



Cryptophyta  in  1984 (9.1$)  was similar  to 1983  (8.3$),  their  relative



importance dropped  from second  to  third (Table  24).   The  Chrysophyta



accounted for  the  second highest  biovolume  (9.45$).   Highest   overall



densities were attained by  the picoplankton (83.9$ of  the total).   Both



the Cyanophyta and the  Chlorophyta had lower  average blovolumes  in  1984




than  in 1983, while Pyrrhophyta biovolume increased (Table  24).






Seasonal Abundance and Distribution of Major Algal Groups



      Seasonally,  abundance (cells/mL) and  biovolume (mm  /ml)   increased



from April to a  maximum (33,355 cells/mL)  in early July  (Fig. 24b).    A



secondary maximum in abundance (19,663  cells/mL) was observed  in   August,

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                                                                   53




due to pIcoplankton.  A secondary  peak was not observed in the  biovolume



seasonal distribution  (Fig. 24a)  because  of the  low biovolume  of  the



picop Iankton.  Samples  were not taken  during the late  summer and  early



fall.  Abundance  was low in  November and decreased  into early  December



(11,388 cells/ml).  Abundance increased  slightly in January but  returned



to December  levels  in February.   Abundances in the early spring, fall  and



winter  were  not  significantly  different.    Also,   biovolume  was  not



significantly different between the early spring, fall,  winter and  August




(Fig. 24a).



     Considering  biovolume, the BaciIlariophyta were dominant  throughout



the study period accounting for as much as 12% but never less than 44?  of



the phytoplankton biovolume  (Fig. 25).    The large drop  in the  relative



importance of diatoms in August of  1983 (to -30? of the total   biovolume,



Makarewicz 1987) was not observed in 1984.  A drop to 47? of the biovolume



did occur  in August.  The  occurrence of a bloom of Rhizosolenia  erJens is



in August  of 1984,   not observed  in 1983,  was the  major cause  of  the



dominance of the diatoms during the summer (Table 25).   With the  decrease



in the relative biovolume of diatoms, a seasonal succession of Pyrrhophyta



peaking in July, Cryptophyta in early August,  and Chrysophyta in August is



evident.  Diatoms regained their  spring predominant position by  February



(Fig. 25).   Cryptophyta  appeared to  increase  in  importance during  the



study period accounting for 18? of the total biovolume in the late  autumn



and winter samples.





Geographical Abundance and Distribution of Major Algal  Groups



      In  1983 the mean  phytoplankton abundance for  the sampling  period



generally decreased from north  to south to  -Station 15, where  abundance



increased and  then decreased  slightly  southward (Fig.   26)  (Makarewicz

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                                                                   54   .



1987).   In 1983 Aster ione! I a.  formosa, Cycloteila comensis,  £. comta,  and



£. oceI  Iata all had a higher biomass at Station 61 than at other  stations



(Makarewicz 1987).   A  similar algal  geographical distribution  was  not



observed during  the 1984-85  sampling period  (Fig. 27).    There  was  no



obvious pattern on a cruise basis either (Fig. 28).  Not one common  algal



species had  an  abundance  maximum  at the  northern  stations  in  1984.



Although not likely,  this difference  may be attributed  to the  sampling



patterns between 1983 and 1984.  !n 1983 six of the seven cruises  sampled




the same stations, while  in 1984 only 50% of the cruises sampled the  same



stations (Table  4).   This  sampling pattern  apparently did  not  affect



zooplankton data.  Similar to 1983, zooplankton populations were higher at



Station 61  in 1984.






Regional and Seasonal Trends in the Abundance of Common Taxa



        Common  species   (Table  26) were  arbitrarily  defined  as  those



possessing a relative abundance  of >0.1$ of the  total cells or >0.5$  of



the total biovolume.  Ninety-four  percent of the common species  observed



in 1984 were  also common  species in  1983.   Ten percent  of the  common



species observed in 1983  were not common in  1984 (Table 27)  (Makarewicz



1987).



     The causes of  these differences are difficult to evaluate.    Natural



annual variability  of  plankton  populations in  the lake  has never  been



evaluated and cannot be evaluated until a more extensive data set  exists.



Seasonal sampling variability exists between 1983 and 1984 and is the most



probable cause for the  species differences observed.   For example,  both



Osc iI  Iator i a m i n i ma and Stephanodiscus minutus were common in 1984 because



of their high density in the winter  of 1984-85.  Winter samples were  not



available in 1983-84.

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                                                                   55




     Because of the similarity of the 1984 common species list to the 1983



list, a  species by  species description  of autecology  and regional  and



seasonal trends  are  not  warranted  here  and  can  be  referred  to  in



Makarewicz (1987).  Only new common species are discussed below.






BaciIlariophyta



  Cyclotella stelIigera (Cl. and Grun.) V.H.



         This  species  is a  common  offshore  dominant  in  Great  Lakes



phytoplankton assemblages  (Stoermer  and Kreis  1980).   It is  apparently



intolerant of  highly  eutrophic  conditions in  the  natural   environment



(Stoermer and Kreis 1980).  In 1971 Munawar and Munawar (1982) reported C.



ste]Iigera to  be a  common  lakewide species  (>5% of  the  phytoplankton



biomass).    In southern  Lake Huron  during 1974,   mean abundance  was  54



cells/mL with a maximum of 720 cells/mL in July (Stoermer and Kreis 1980).



At a single  offshore station  in southern  Lake Huron,  Lin and   Schelske



(1978) observed a maximum of 762 cells/mL in late July with an average for



the sampling  period  (March-December 1975)  of  111 cells/mL.    Offshore



average abundance and  maximum abundance  in 1980 were  10.9 cells/mL  and



60.7 cells/mL,  respectively (GLNPO Data Base).   Abundance in 1983 was only



6.5 cells/mL (Makarewicz 1987),  while  in 1984 average abundance  was  25.3



cells/mL with  a  maximum of  267  cells/mL.   The  lower  mean  abundance



observed in 1983 and 1984  is probably caused by  the lack of  sampling  in



the month  of  July in  both  years,  when this  species  is  historically



dominant.  Abundance was   low during the spring,  fall and winter of  1984



(Fig. 29a).    The population did  increase by the  5-7 July sampling  date



(mean station  abundance =  123  cells/mL)  but did  not reach   the  higher



abundances historically observed later in July.

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                                                                   56




  Stephanpd|scu5 alpinus Hust.  (= £.  astrea var minutul us)




      Munawar and Munawar  (1979) did not list  this species as common  in



1971.   Similarly,   abundance  was  generally low  in  1974  (mean  =   2.6



cells/ml) (Stoermer and Kreis 1980),  in  1975 (Lin and Schelske 1978),  in



1980 (mean =  0.1 cells/ml) (GLNPO  DataBase)  and in 1983  (mean =  0.25



cells/ml) (Makarewicz 1987).   Abundance was  also low in 1984 (mean =   1.5



cells/ml), but biovolume represented 0.91$ of the total  biovolume, thereby



qualifying it as a common species (Table 26).  In 1984 seasonal  abundance



peaked in early  July (Fig. 29b)  and was  low (< 2  cells/ml)  during   the



remainder of the samp I ing period.



         This  species  is a  common  minor  element  of  the  Lake  Huron



phytoplankton assemblage.   It  appears to  be favored  by  low   levels  of



eutrophication, but it is not  tolerant of extreme levels of  perturbation



(Stoermer and Kreis 1980).






  Stephanodiscus^ mJnutus Grun.  (= .S_.  miputul us)



     .S.  minutus is generally considered to  be a fall or winter  blooming



species  in mesotrophic  or eutrophic lakes   (Stoermer and Ladewski  1976).



It was not common  in 1971  (Munawar  and Munawar 1979) and possessed a  low



abundance in 1974  (mean = 7.5 cells/mL) (Stoermer and Kreis 1980),  in  1975



(mean =  4.2  cells/mL) (Lin  and  Schelske  1978),  in  1980 (mean  =   4.2



cells/mL) (GLNPO Data Base) and  in 1983 (mean = 2.56 cells/mL)  (Makarewicz




1987).



      In  1984, average density was 19.4 cells/mL with a maximum density  of



84 cells/mL.  Seasonal  abundance was low during the summer (< 6 cells/mL),



was  higher  during spring  and   autumn (-25  cells/mL)  and peaked  at  63



cells/mL  in  February  of  1985   (Fig.  29c).   This  species is   a  winter

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                                                                   55




     Because of the similarity of the 1984 common species list to the 1983



I ist, a  species by  species description  of autecology  and regional  and



seasonal trends  are  not  warranted  here  and  can  be  referred  to  in



Makarewicz (1987).  Only new common species are discussed below.






BaciIlariophyta



  Cyclotella stelIigera (Cl. and Grun.) V.H.



         This  species  is a  common  offshore  dominant  in  Great  Lakes



phytoplankton assemblages (Stoermer  and Kreis  1980).   It is  apparently



intolerant of  highly  eutrophic  conditions in  the  natural   environment



(Stoermer and Kreis 1980).  In 1971 Munawar and Munawar (1982) reported C.



stelIigera to  be a  common  lakewide species  (>5% of  the  phytoplankton



biomass).   In southern  Lake Huron  during 1974,   mean abundance  was  54



cells/mL with a maximum of 720 cells/mL in July (Stoermer and Kreis 1980).



At a single  offshore station  in southern  Lake Huron,  Lin and   Schelske



(1978) observed a maximum of 762 cells/mL in late July with an average for



the sampling  period  (March-December 1975)  of  111 cells/mL.    Offshore



average abundance and  maximum abundance  in 1980 were  10.9 cells/mL  and



60.7 cells/mL, respectively (GLNPO Data Base).   Abundance in 1983 was only



6.5 cells/mL (Makarewicz 1987),  while  in 1984 average abundance  was  25.3



cells/mL with  a  maximum of  267  cells/mL.   The  lower  mean  abundance



observed in 1983 and 1984  is probably caused by  the lack of sampling  in



the month  of  July in  both  years,  when this  species  is  historically



dominant.  Abundance was  low during the spring,   fall and winter of  1984



(Fig. 29a).    The population did  increase by the  5-7 July sampling  date



(mean station  abundance =  123 cells/mL)  but did  not reach  the  higher



abundances historically observed later In July.

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                                                                   56




  Stephanod!sous alpinus Must.  (= .S.  astrea var minutulus)



      Munawar and Munawar  (1979) did not list  this species as common  in



1971.   Similarly,   abundance  was  generally low  in  1974  (mean  =   2.6



cells/ml) (Stoermer and Kreis 1980),  in  1975 (Lin and Schelske 1978),   in



1980 (mean =  0.1 cells/ml) (GLNPO  DataBase)  and in 1983  (mean =  0.25



cells/ml) (Makarewicz 1987).   Abundance was  also low in 1984 (mean =   1.5



cells/mL), but biovolume represented 0.91$ of the total  biovolume, thereby



qualifying  it as a common species (Table 26).  In 1984 seasonal  abundance



peaked in early  July (Fig. 29b)   and was  low (< 2  cells/mL)  during   the



remainder of the sampling period.



         This  species  is a  common  minor  element  of  the  Lake  Huron



phytoplankton assemblage.   It  appears to  be favored  by  low   levels  of



eutrophication, but it is not  tolerant of extreme levels of  perturbation



(Stoermer and Kreis 1980).






  Stephanodlscus minutus Grun.  (= 2..  mjnutul us)



     £.  mi putus is generally considered to  be a fa I I or winter  blooming



species  in  mesotrophic  or eutrophic lakes  (Stoermer and Ladewski  1976).



It was not  common  in 1971  (Munawar  and Munawar 1979) and possessed a  low



abundance in 1974  (mean = 7.5 cells/mL) (Stoermer and Kreis 1980),  in 1975



(mean =  4.2  cells/mL) (Lin  and  Schelske 1978),  in  1980 (mean  =  4.2



cells/mL) (GLNPO Data Base) and  in 1983 (mean = 2.56 cells/mL)  (Makarewicz



1987).



      In  1984, average density was 19.4 cells/mL with a maximum density  of



84 cells/mL.  Seasonal abundance was low during the summer (< 6 cells/mL),



was  higher  during spring  and   autumn (-25  cells/mL)  and peaked  at  63



cells/mL  in February  of  1985   (Fig.  29c).   This  species is   a  winter

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                                                                   57



species.  The low abundances historically observed are related to the lack



of w inter samp I ing.






Chlorophyta



  Cosmarium sp.



      Abundance of Cosmarium was low  in 1971 (Munawar and Munawar  1979),



1974 (Stoermer and Kreis 1980), 1975 (Lin and Schelske 1978),  1980  (GLNPO



DataBase), 1983 (Makarewicz 1987) and even  in 1984 (this study).  Because



of its  relatively high  individual  biovolume,  it qualifies  as a  common



taxon (Table 26).   Abundance peaks were evident  in early July and  early



winter (Fig. 29d).






Cryptophyta



  Cryptomonas rostratiform is Skuja



     Mean abundance of  this species was low  (0.8 cells/mL).  However,   the



high biovolume of the individual cell causes it to be a common species  in



1984.    £.  rostratJlormJs apparently  was  not observed  by  Munawar  and



Munawar (1979),  Stoermer and  Kreis (1980) or Lin  and Schelske (1978)  in



Lake Huron.  In 1983 abundance was low (mean = 0.35 cells/mL).  A  maximum



of 8 cells/mL was observed with a mean of 0.8 cells/ml in 1984 (Table 26).



Seasonal abundance showed a great deal of variability, perhaps because  of



the low abundance.






Cyanophyta



  OsciI Iatori a m i n i ma G i ckIh.



     Many of the previous workers (Munawar and Munawar 1979, Stoermer  and



Kreis 1980, Lin  and Schelske  1978) on Lake  Huron did  not  identify  all



fopms of Osc iI Iator i a to the species level.  Average abundance in 1983 and



1984 was 2.9 (Makarewicz 1987)  and 17.3 cells/mL, respectively.    Maximum

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                                                                   58  .




abundance was  335 cells/mL  at Station  45 on  3 August  1984.   Seasonal



abundance was bimodal with a maxima in mid-summer and winter (Fig. 30).






Vertical Distribution



      On 15 August 1984, a  series of vertical phytoplankton samples  were



taken at two stations (Stations 15  and 37).  The abundance increase  with



depth at  Station  37  can  be primarily  attributed  to  an   increase  in



picopIankton (Fig. 31).     In general,   non-pi cop Iankton abundance did  not



increase with  depth with  the  exception of  Cyclotel l_g peel I ata.  and  £.



kuetzingiana var.  planetophora  (Fig.  31).   There  was  no  correlation



between the abundance increase and temperature (Fig. 31).



      At Station 15,   phytoplankton samples were taken  to a depth of  30m



compared to 20m at Station  37.  An increase  in picoplankton,  as well  as



BaciI Iariophyta and Chrysophyta,  was evident.   The abundance  increase  in



these groups correlated with the  decrease  in temperature associated  with



the  metal imn ion  (Fig.   32),     Cyclotel I a  comensis,  £.  ocel I ata   and



Tabellaria flocculosa were responsible for the diatom abundance increases,



while Dinobryon sociale and Q, divergent  were the primary causes for  the



Chrysophyta  increase  with   depth  (Fig.   32).      A  similar   vertical



distribution pattern was observed in  Lake Michigan in 1984 (This  Study).



Brooks and Torke (1977), Mortonson (1977) and Bartone and Schelske  (1982)



have reported sub-surface  maximum in the  Great Lakes.    Reasons for  the



existence  of  the  layer  are  not  clear  and  are  apparently   complex,



encompassing  physical,   chemical   and  biological  factors  (Bartone  and



Schelske 1982).

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                                                                     59





Winter Cruises



         Biomass  and  abundance  were  low  during  the  winter  and  not



significantly different from the autumn and  spring values (Fig.  24).   As



during the  non-winter season,   the BaciI Iariophyta  (58.2$ of  the  total



biovolume) was the  dominant division.   However,   the Chrysophyta,   which



were second in importance during the  entire sampling period (9.5% of  the



total biovolume),  represented only 0.46$ of the total  biovolume during the



winter.  Similar to the Lake Michigan winter assemblage,  the importance of



the Cryptophyta  increased by  a factor  of greater  than three  from  the



non-winter (5.7$ -  7.1$ for  the entire  sampling period)  to the  winter



period (17.8$).



       FragiI aria crotonensis (mean  = 65.4  cells/ml) and  Stephanodiscus



minutus (mean =54.1 cells/mL)  were the dominant diatoms during the winter



period.  Abundance of  .S_.  minutus was high   only during periods of  cooler



water temperatures (Fig. 29c).   Because  of the high winter abundance,   it



became a common species  for the year (Table  26).  Similarly,   Fragi iaria



intermedia var. fall ax, Cryptomonas pyrenoidi fera and OsciIIator ia  mini ma



became common species for  the year (Table  26)  by virtue of their  higher



abundance or  secondary maxima (Fig. 30) during the winter.



       Other   major  winter diatoms,   Cyclotella comensis  and  TabelIaria



fenestrata were common  species (Table 26)   during the non-winter  period.



Common  winter,  as  well   as  non-winter,   species  of  Cryptophyta   and



Cyanophyta were   Gomphosphaeria   Iacustris,  Rhodomonas   minuta   var.



nannoplanktica (1.19$  of the  total  cells-winter)  and Cryptomonas  erosa



(5.30$ of total biovoIume-winter).

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                                                                      60





Historical  Changes in Species Composition



     The literature pertaining to phytoplankton of the offshore waters  of



Lake Huron is  sparse.   Fenwick (1962,  1968)  published some  qualitative



data, and Parkos et al.  (1969) listed species observed.   Quantitative data



from a single offshore station in  1971  exists (Munawar and Munawar  1982,



Vollenweider et al.  1974) and  preliminary data covering  21 stations  in



1971 are  partially analyzed  (Munawar and  Munawar 1979).    Stoermer  and



Kreis (1980) reported on  an extensive sampling  program in southern  Lake



Huron  including  Saginaw  Bay  during  1974  and  provided  an  extensive



bibliography on Huron algal research.  Lin and Schelske (1978) reported on



a single offshore  station sampled   in 1975.   An intensive  study of  the



entire lake basin was performed in  1980 (Stevenson 1985),  but only a  few



offshore stations were sampled.



     Since 1971 diatoms have been the dominant division.  Dominant diatoms



in  1971  included  species   of  Asterione I I a  formesaf A-    grac?I Iimaf



CyclotelI a comtaf  £. glomerata,  £.  ocelI ata,  £. mich iganiana,  Melosira



island ica and  M* granulata.    In addition,  species such   as  FragiI aria



crotonensis and  Tabellaria fenestrata  were common,   while  cryptomonads,



such as Rhodomonas tninuta and Cryptomonas erosa, contributed very  heavily



during different seasons.



     The following similar common diatoms (>0.1$ of the total cells) wlere



observed in  1974,  1983  and  1984:     Asterionella  formosa,  Cyclotella



comensi sf  £. mich igan iana, Q. ocelIataf  Fragi i aria crotonensis, TabeI Iar i a



fenestrata,  J.  flocculosa var.  i inearis and  Rh izosolen ia sp.    Synedra



fiI?form is was present  in 1983  and 1984 but was not  as common as  in  the



1974 southern Lake Huron plus Saginaw Bay data.  The lower abundance of £.



stelIigera in 1983 (Makarewicz  1987) and 1984  compared to 1971  (Munawar

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                                                                     61


and Munawar  1979), 1974  (Stoermer  and Kreis  1980)  and 1975  (Lin  and


Schelske 1978) was caused by the lack of sampling during mid and late July


when this species  is dominant.


     Both Cryptomonas erosa and Rhodomonas minuta var. nannoplanktica were


dominant in 1971,  1974, 1983 and 1984.  Numerically dominant  chrysophytes


in 1971 were Dinobryon divergens and ChrysosphaerelI a longisp ina.  In 1983


and 1984, these two species were  common along with Q. cylindricum and  J2.


sociaIe var. americanum  (Table 26).   Haptophytes  were also  numerically


abundant.    In  general,  the  diatom  Synedra  fiIiformis  decreased  in


abundance after 1974, while J3. cylindricum and D. sociale var.   americanum


have increased in abundance.   In general,  species composition of  common


offshore algae has changed  little since 1971.



P i cop i ankton

                                                                         4
     Picoplankton abundance in  1984 (mean = 14,396;  maximum of 3.5 x  10


cells/ml) was not dissimilar  from 1983 (mean =  19.343; maximum of 6.3  x


10  cells/mL).  On a  numerical basis, the picoplankton represented  83.9$


of the total cells in 1984 but  because of their small biomass, only  1.6$


of the total biovolume.   Their relative numerical  dominance  in 1984  was


comparable to 1983 (86.6$).   Prior to the  1983 study (Makarewicz  1987),


other researchers have not routinely reported on this group of organisms.




Indicator Species


     Dominant diatoms in Lake Huron  in 1983 and 1984 were Rhizosolenia sp.


(£. erJens is  in 1984),  TabelI aria  flocculosa (biomass)  and  Cyclotella


comensis (numerically).    Four species  of  CyclotelI a (£.  comensis,  C.


cpmta,  £. kuetz i ngiana var. pignetophora and £. ocelI ata) represented 9.4$


of the total biomass  in 1983 (Makarewicz  1987).  In  1984 the same  four

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                                                                     62




species plus  Cyclotella  stelIFgerg  accounted for  6.63$  of  the  total



biomass (Table  26).   £.  eriens is  is often  grouped  with  oligotrophic



offshore dominants even though it may occur in greater abundance in  areas



receiving some degree  of nutrient  enrichment (Stoermer  and Yang  1970).



Except for £. comensisr  whose ecological affinities are poorly  understood



(Stoermer and Kreis 1980), these species are associated with  oligotrophic



or mesotrophic conditions.   Tabellaria flocculosa is commonly  associated



with mesotrophic conditions (Tarapchak and Stoermer 1976).



     Dominant chrysophytes included Dinobryojl sociale var. americanum,   J2.



divergens and  I), cy I  indr icum,.  which are  often associated  with  several



small members  of  the  genus  Cyclotella (Schelske  et  al.  1972,  1974)



included in  the classical  oligotrophic  diatom plankton  association  of



Hutchinson (1967).  Dominant cryptophytes, cyanophytes and dinofIagellates



were  Rhodomonas   minuta  var.      nannoplanktica,   Cryptomonas   erosaf



picoplankton and Ceratium hirundinelI a.



     Because of the   Iimited number of studies of the Lake Huron  offshore



phytoplankton assemblage, there  was also a  limited basis for  evaluating



the  long-term  effects of  eutrophication.   The ratio  of mesotrophic  to



eutrophic species in  Lake Huron has  not changed since  1971 (Table  28).



This suggests that the trophic status of the lake has not changed.  Because



the trophic ratio has  not been extensively  used,  interpretations of  the



trophic ratio have to  be carefully considered. For  example, the  lack  of




change in  the  ratio  in  Lake  Huron may  simply  represent  a   lack  of



sensitivity  in the ratio. However,   interpretations using the trophic ratio



in collaboration with  other  indicators  suggest interpretations  of   the



trophic ratio parallel the other indicators.

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                                                                     63 .




      Those studies available (Munawar and  Munawar 1979,  Nicholls et  at.



1977a, Schelske et al.  1972, 1974) indicate that  the waters of  northern



Lake Huron  generally  contain  phytoplankton  assemblages  indicative of



oligotrophic conditions.    The  designation of  the  offshore  waters of



southern Lake Huron as oligotrophic based on phytoplankton composition  in



1983 and 1984 is not unlike  the trophic status suggested  by Stoermer  and



Kreis (1980) for the offshore waters of  southern Lake Huron in 1974.   This



agrees  well  with  the  trophic  status  as  determined  by  the  biomass



classification scheme of Munawar and Munawar (1982).   With a mean  biomass



of 0.38 g/m   and 0.42 g/m   for 1983 and  1984,  respectively,  Lake  Huron



would be classified as oligotrophic.






Historical Chajigeg.in Community Abundance and Biomass



     Some quantitative phytopIankton data exist for the offshore waters of



Lake Huron from  at least  1971.  The  collections of  Stoermer and  Kreis



(1980) were from 44 stations in southern Lake Huron and Saginaw Bay.   Lin



and Schelske (1978) collected from one offshore station in 1975.  In   both



studies, phytoplankton were concentrated on millipore filters rather   than



by the settling chamber procedure used in the 1980 (GLNPO  Data Base),  1983



(Makarewicz 1987) and  1984 studies.   Thus,   data sets  are not  strictly



comparable.



     Munawar and Munawar (1982) collected with a 20-m integrating  sampler



from April  to December of 1971.   Because Utermb'hl's (1958) procedure  for



enumeration of algae was employed,  these data were directly comparable to



the 1980,  1983 and 1984 data  sets.  Unfortunately, biomass data for   only



one offshore station  of Lake Huron  was available for  1971 (Munawar  and



Munawar 1979).   Phytoplankton  biomass between 1971,   1980, 1983 and   1984



was not  significantly  different  (Fig.   33).    The  consistency  of the

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                                                                     64




mesotrophic-eutrophic  ratio   through   time  and   the   occurrence   of



oligotrophic and mesotrophic  indicator species suggest  little change  in



the trophic status of the offshore waters of Lake Huron.

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                                                                     65





LAKE HURON



ZoopIankton



Annual  Abundance of Zpoplankton Groups



     Species lists (Table A10) and summary tables of abundance (Table A11)



and biomass (Table A12) are  in Volume 2 -  Data Report.   The  zooplankton



assemblage of 1984 comprised  53 species representing  31  genera from  the



Amphipoda, Calanoida, Cladocera, Cyclopoida  and Rotifera.   The  diversity



of species was similar to 1983 (58 species,  33 genera).



       The  Rotifera  possessed the  largest number  of species  (31)  and



relative abundance (56.0$) followed by the Calanoida and Cyclopoida.    The



Copepoda nauplii accounted  for 18.6/5 of  the total  zooplankton  abundance



(Table 29).   The  Calanoida  (42.0$) followed  by the  Cladocera  (27.5$)



contributed the  most  biomass to  the  zooplankton community.     Rotifera



represented only 2.5$  of the  zooplankton biomass.    Average density  and



biomass were  55,369+7,176 (mean±S.E.)  organisms/m  (46,230  - 1983)  and



27.3+2.3 (mean±S.E.) mg/m  (Table 6).






Seasonal Abundance and Distribution of Major Zooplankton Groups



       Seasonally, the abundance  and biomass  were essentially  identical



(Fig.  34) with a maximum in August.   This pattern was similar to that  of



Lake Michigan in 1984 (Fig.  16).   A comparison  to 1983 was not  possible



because of the lack of samples in the spring and summer (Makarewicz 1987).



Except for the  naupl ius stage  of the  Copepoda,  abundance  of the  major



zooplankton groups was  highest in August  as compared to  the spring  and



late autumn samples  (Fig. 35).  Nauplii abundance was high throughout  the



year with a general trend of  decreasing abundance towards the winter.   A



similar pattern was observed with biomass distribution with the  exception



of the Calanoida.   Calanoid biomass did not decrease markedly in the  late

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                                                                     66




fall as compared  to other  groups (Fig.  36).    Growth  of the  individual



Calanoids, even with a decreasing abundance,  kept Calanoida biomass high.





Common Species




     Common Crustacea species (Table 30)  were arbitrarily  defined  as those



possessing  a  relative  abundance  of  >0.1$  of  the  total   zooplankton



abundance or 1.0$ of the total  biomass.   Rot ifera species  were  considered



common if they accounted for >\,0%  of the total  zooplankton abundance   or



biomass.    The  number of  common  species were identical   in  1983  (22)



(Makarewicz 1987)  and 1984  (22).   Some  differences in   common   species



composition were  evident.    Polyarthra  remataf   Notholea  squamula,   and



Leptodora k indti i were common in  1984 but not in  1983.   Even though  the



cladoceran Leptodora kindtii  was not abundant  in 1984,  it  was a  common



species because of its high biomass per organism.   In 1983 biomass was  not



evaluated in the designation  of common species.    Daphnia retrocurva,   Q.



schodleri and D. catawba were common in 1983  but not in 1984.   D.   catawba



was observed  only in  long hauls  in  1983,  while  D.  schodleri  was  not



observed at alI  in 1984.





Changes in Species Composition



Crustacea



     Crustacean studies of the offshore waters of the Lake Huron basin  are



few in number.  Patalas (1972)   sampled 51 stations including Saginaw  Bay



in August of 1968  with a 77-um mesh  net.  In 1971  eleven stations on  a



transect from the Straits of Mackinac to  the  origin of the St.  Clair River



were sampled from May to November  with a 64-um net (Watson and Carpenter



1974).  A 64-um mesh  net was used to sample   ~18 stations on eight  dates



from April to October of 1974 in southern Lake Huron including Saginaw  Bay

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                                                                     67





(McNaught et al. 1980a).   The 1980 study  of Evans (1983, 1986)  included




stations mostly from  the nearshore rather  than the offshore.   The  1983




sampling cruises included 10 stations sampled (64-um mesh net) for each of




the three sampl ing  dates between  August and  September.   In 1984  eight




stations on  five cruises  (64-um mesh  net) from  May-December 1984  were




sampled.




     In August of 1968, calanoids were dominated by Diaptomus siciI is,   D.




ash I and i and D.  minutus (Patalas 1972).   These same  three species  were




predominant in  1971, 1974/75, 1983 and 1984 with the addition of D i aptomu s




oregonensis in  1983 and 1984 (Table 31).  Abundance of Diaptomus  .ash I.and I




and Diaptomus siciI  is  appears to  have increased since  1971  (Table  31).




The 1974 D. minutus  abundance was higher than  either the 1971,  1983  and




1984 samples.    However,  the 1971,   1983  and 1984  data were  only  from




offshore sites, while 1974 data included samples from the eutrophic waters




of  Saginaw  Bay.    The  ol  igotrophic  indicator  species,    L imnocaI anus




macrurus, appears to be decreasing in abundance (Table 31).




        In  1971,  1974/75 and  1983,  the  dominant cyclopoid  was  Cyclops




bicuspidatus thomasi (Table 31).  Tropocyclops prasinus mexicanus increased




in abundance from 1971  to 1983.   However,  a notable decline occurred  from



1983 (577/m )  to 1984 (21/m3),  which may be related to the differences  in




the timing of   the fall  sampling in these  two years.   Mesocyclops  edax




appears to  have increased   in abundance  (Table 31)  from 1971  to  1983.




Abundance was    lower  in 1984  than  in  1983.    Cyclops  vernal is,,   often




associated with eutrophic conditions in Lake Erie, was higher in abundance




in the 1974 data.   This higher abundance may again  have been due to  the




inclusion of the eutrophic of Saginaw Bay stations in the 1974 data set.

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                                                                     63





         Dominant  cladoceran species  in  August  of  1968  were  Rosming



longirostris  and  Ho I oped i urn  gibberum.     Similarly,   H.   gibberinri,    fi.



longirostris and Eubosmina  coregonj were dominant  in the  August-October



period in 1974.   Comparison  of the 1971  and 1984  August data  suggests



decreases in abundance of B. longirostris and iJ.  gibberum.



     Quantitative data on species of daphnids were not available for 1971,



but Daphnia retrocurva,  Daphnia galeatg mendotag  and J).  longiremis  were



commonly found in Lake  Huron (Watson and Carpenter  1974).   The  dominant



daphnid species in 1983 and 1984 was JD.  gal eata mendotae.



       Evans (1985)  recently reported that  Daphnia pul icaria  was a  new



species dominating the Lake Michigan  zooplankton assemblage.  In 1983  in



Lake Huron,   D. pul icaria  was observed  to be  the third   most  important



cladoceran,  while  in  1984  it dropped  to fifth in  rank  abundance  (Table



30).  Mean  station abundance increased  from north to  south with a  mean



density of 431 organisms/m  for stations south of Saginaw  Bay in 1983.   In



1984, abundance never reached the levels of 1983  (Fig.  37).



      J2.   catawba was  first reported  in  waters of  Lake Huron  in  1983



(Makarewicz 1987).  This species was not considered to be  either a  common



or a less  common species of  the Great Lakes  (Balcer et  al.  1984).    It



appeared exclusively in the long hauls from Lake  Huron in  1983.   A maximum



abundance of 1,610 organisms/m  was observed in August at  Station 12.    It



was not observed in 1984.




      Bythotrephes cederstromii was observed in  Lake Huron for the  first



time in the long haul  of 1984.    This European invader was first  observed



in the Great Lakes in Lake Ontario in 1985 (Lange and Cap  1986).   It is  a



conspicuous species in the plankton of European oligotrophic lakes.

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                                                                     69
Rot i fera



     Stemberger et al.   (1979) collected rotifers with a Nisken bottle  at



5-m intervals to 20m followed by 10-m intervals to the bottom of the  lake



at a number  of offshore  and nearshore areas.    Samples  were pooled  and



filtered through a 54-um mesh net  on the vessel.  The greatest  abundance



of rotifers in Lake Huron in 1974 occurred in late spring and early summer



(Stemberger et al. 1979),  a period in which no samples were taken in  1983



and  1984.    Comparison  of  the  August-October  samples  suggested  the



following between the  1974, 1983 and  1984 data sets;   abundant  rotifer



species in both  studies were Conoch?I us  un icornisf  Polyarthrg  vulgari sf



KeratelI a cochlearis  and Ke11i cott i a  longispina; £. unicornis  was  the



dominant rotifer in 1983;   and KeratelI a cochlearis  was dominant in  1974



(Table 30).



     Evans' (1986)  study of mostly nearshore areas suggests a  difference



in dominant rotifer  species between  the offshore  and nearshore  waters.



Dominant rotifer  species   in  this study  included  in  descending  rank:



Keratel I a cochlearis,.  Ke I I i cott i a longisp inaf  Synchaeta sp.  and Conoch i 1 us



unicornis.   Polyarthra  vulgaris  and ConochiI us  unicorn is,   which  were



dominant in the offshore waters in 1974,  1983 and 1984,  were less abundant



in the nearshore waters.



       These  differences in  horizontal  distribution  of zooplankton  are



expected in Lake Huron and are  affected by the physical Iimnology of  the



lake (McNaught et al.   1980a).  For example  in the warmer inshore  areas,



cladocerans grow best,  while calanoids tend to be found in offshore waters



(McNaught et al.  1980a).    Nearshore waters  are also  influenced by  the



movement of the zoopIankton-rich eutrophic waters of  Saginaw Bay into  the



nearshore zone  south  of  the  Bay.    In  general,   inshore  zooplankton

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                                                                     70




densities are greater  than offshore  densities (McNaught  et al.   1980a).



Similarly, abundance  and species  composition  of rotifers  increase  and



differ in the shallow more productive waters of the western basin  of  Lake



Erie (Fig. 62).






Geographical  Abundance and Distribution of Zooplankton Groups



      The mean  station zooplankton abundance was  higher in the  northern



half than in the southern  half of Lake Huron  (Fig.  38)  due primarily  to



higher rotifer abundance in the north.  A similar pattern was observed  in



1983 (Makarewicz 1987).  With a biomass comparison, no obvious  difference



between the northern and southern half of the lake was evident (Fig.  39).



Cyclopoida and  Cladocera  abundance  was  relatively  similar  along  the



north-south axis.   The Calanoida and naupl ius stage of the Copepoda had  a



geographical  distribution pattern similar to the Rotifera with  increasing



abundance from Station 61 to 45,  descending abundance from Station 45  to



27 and 12, and increasing abundance southward (Fig. 39).   McNaught et  al.



(1980a) observed  abundance increases  of  the cyclopoid   copepodites,   £.



bicusp idatus and J. praslnus,  north to  south in southern Lake Huron.   In



1983 rotifers also decreased in abundance from north to south to  Stations



9 and 6 where a si  ight increase was evident.



       The  1983  and 1984  data (Figs.  37  and 39)   suggest a trend  of



increasing total  zooplankton abundance from Station 12 northward with  the



exception of Station  32 in 1983.   Station 32,   located  northeast of  the



mouth of Saginaw Bay, would appear to be too far offshore to be influenced



by the higher abundances in the  Bay.  However,  Stoermer  and Kreis  (1980)



have observed mid lake stations   in southern Lake Huron  to be affected  by



populations of  phytoplankton from  Saginaw  Bay in  1974.   Although  the



transport of  eutrophication-tolerant algal   populations  into  Lake  Huron

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                                                                     71




from Saginaw Bay has been mitigated in recent years (Stoermer and  Theriot



1985), the mechanism of transport still   exists and thus the transport  of



zooplankton could still take place from Saginaw Bay.



      A number of  zooplankton species possessed horizontal   distributions



that varied along the  north-south axis.  These  differed between 1983  and



1984.   In 1983  Diaptomus minutus  abundance was  lower in   the  northern



portion of the lake, while Daphnia retrocurva had a maximum  limited to the



far northern  stations.    Abundance  of   both  ConochiI us  unicprnis  and



Ke111cott i a longisp ina decreased from north to south.  Ho I opediurn  gibberum



had a  higher  abundance north  of  Saginaw Bay,  while   Mesocyclops  edax



abundances were higher south of Saginaw Bay. Cyclops bicuspidatus  thomasi



was more abundant at  the far northern  stations than  in  the rest of  the



lake (Makarewicz 1987).



      In 1984 Diaptomus s i c iIi st  the copepodite of Mesocyclops,   Notholca



squ annulaf  Polyarthra vulgaris and Synchaeta  sp. had abundances that  were



higher in southern Lake Huron (Fig. 41).   Mesocyclops  edgx adults did  not



have a higher  abundance in  southern Lake  Huron as  in 1983  (Makarewicz



1987).  However,  juveniles of Mesocyclops were higher  in the southern Lake



Huron (Fig.  41).   Similar to  1983,  abundances  of Ho I opediurn  gibberum,



KelIicottia longispina and  ConochiJus unicorn is were  higher in  northern



Lake Huron (Table 32), while Diaptomus minutus was lower in the  northern



half of  the lake.  A similar north-south  distribution  of algal  populations



was not observed in 1984.






Indicators of Trophic Status



     The calanoid/cyclopoid plus cladoceran ratio (the plankton ratio) has



been employed as a  measure of trophic status  in the  Great  Lakes  (Gannon



and Stemberger 1978, McNaught et al.  1980a).  Calanoid copepods  generally

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                                                                     72




appear best  adapted for  oligotrophic conditions,   while cladocerans  and



cyclopoid copepods  are  relatively  more abundant   in  eutrophic  waters.



Using this ratio, McNaught et  al.  (1980a) identified the offshore  waters



of southern Lake Huron to have  a higher quality water than the  nearshore



waters.  Because the 1983 and 1984  samples were all  from the offshore,   no



such comparison could be made.  However, the 1984 plankton ratio was  high



and variable from north to south (Table 33).   A comparison of the 1983 and



1984 mean  phytoplankton  ratio  suggests  a lower   quality  of  water  at



Stations 6 and 9 and perhaps at Station 61.   Water  chemistry data suggests



these southern stations  have higher chloride,   sulfate,  total   phosphorus



and turbidity levels  and lower silica  levels than the  rest of the  lake



(Fig. 42).




      Station 61 might  be influenced by waters  from Lake Michigan.    The



plankton ratio at Station 61  in Lake Huron is comparable more to  northern



Lake Michigan  than the  rest of Lake  Huron (Table  34).    The  physical



transport of plankton  populations  by  water currents  from Lake  Michigan



into Lake  Huron through  the Straits  of Mackinac   has been  demonstrated



(Schelske et al. 1976).



      Species considered  to be indicators of  eutrophic waters were  rare



compared to  the western  basin of   Lake Erie  and  restricted  to  extreme



southern Lake Huron (Station  6) (F iIinia long iseta [6.6/m H,   Trichocerca



mill tier in Is p2.3/m 3) or  not detected  (Brachionus spp.).    The  rotifer



community in 1983 and 1984 was dominated by  Polyarthra vulgar is,  Keratella



cochlearis,   Conoch iI us  unicornis   and  KeI Ii cott i a  long isp ina.     This



association has been considered to   be indicative of an oligotrophic  lake



(Gannon and  Stemberger  1978).  The  offshore abundances  of  Ho I opediurn



g ibberum, Conoch iI us  unicorn is and  Ke iIi cott i a long isp ina  were  greater

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                                                                      73




north of Saginaw Bay than south  of it (Table 32) suggesting better  water



quality in northern Lake Huron.  Ho I opediurn gibberum has been reported  as




an indicator of oligotrophic lakes in Sweden (Pejler 1965)  but was  widely



distributed in both  oligotrophic and eutrophic  waters in  the  Laurentian



Great Lakes region (Gannon and Stemberger 1978),



      The low zooplakton  abundance,  compared to those  of  Lakes Erie  and



Michigan (Table 6), the presence of the oligotrophic rotifer  association,



the domination of the calanoids, and  the fairly  abundant presence of  the



oligotrophic  Diaptomus   sic?I is   (McNaught  et   al.   1980a)   suggest



ol. igotrophic offshore waters for Lake Huron in 1983 and 1984.






Historical  Trends  in Abundance



     Offshore crustacean zooplankton data collected with similar mesh size



nets (64 urn) exist for Lake Huron.   The 1970 study (Watson and  Carpenter



1974; 88  collections) sampled  the  whole lake,   while the  1974/75  work



(McNaught et al. 1980a; 46 collections)  was from southern  Lake Huron.   A



comparison of the cruise averages for Crustacea (excluding  nauplii)  (Fig.



43) suggests an increase in abundance from 1970 to 1974 and 1983  followed



by an abundance drop in 1984.  However, an ANOVA indicates  that the  means



are not significantly  different.  A  similar conclusion of  no change  in



trophic status since 1970 was reached with phytoplankton abundance.



     Stemberger et al. (1979) collected Rotifera samples from 44  stations



in southern Lake Huron in 1974.   Samples were taken with a Nisken  bottle



at 5-m  intervals  to 20  m  and at  10-m  Intervals below   that.    After



collection,  samples were immediately pooled  and  filtered through a  54-um



net.   In 1983 (Makarewicz 1987) and  1984, a vertical  tow (64-um net)  was



taken from 20 m to the surface.  Both studies are not directly  comparable



in that  Stemberger's et  al.   (1979) work  represented the  entire  water

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                                                                     74




column, while the 1983  and 1984 studies were  basically samples from  the



epilimnion.   The 1974  and 1984  sampling periods  are not  significantly



different.  A comparison of mean station seasonal  abundance suggests  that



the spring and autumn abundance  in 1983 and 1984   was lower than in  1974



(Fig.44).  Also,   abundance of major  species was  lower  in 1983 and  1984



than in 1974 (Table 35)   This difference in  abundance is related to  two



things:   (1)   Stemberger et al.  (1979) used a  smaller meshed net  which



gives a  more accurate  quantitative sample  and thus  a higher  abundance



(Likens and Gilbert 1970);  and (2)  two  different segments of water  are



being sampled and  compared.   For example,  Makarewicz  and Likens  (1979)



demonstrated higher abundances  and different species  composition in  the



hypollmnion as compared to the epilimnion of Mirror Lake,  New Hampshire.






Trophic  Interactions



     Within  the offshore, there appears to  be few changes that could  be



attributed to  nutrient control.   Phytoplankton  biomass and  zooplankton



abundance of the  offshore waters of  Lake Huron  in  1971, 1980, 1983  and



1984 are  not  significantly  different.    In  general,  offshore  species



composition of  phytopiankton has  changed  little  since the  early  70's.



However,  there  has  been  a  significant   lake-wide  change   in  species



composition of  zooplankton.   Prior  to  1983,  there  are no  records  of



Daphnia  pul icaria  in Lake  Huron.   In 1983   and 1984, this species  ranked



third  and  fifth  in abundance  in Lake Huron,  respectively.  The  appearance




of the   large JD.  pul icaria   is generally   attributed  to a  release  from



size-selective predation of forage  fish  in  Lake  Michigan (Scavia et  al.



1986,  This  Study) and Lake Erie  (This Study).  Daphnia pulicaria abundance



is correlated with decreased  phytoplankton   abundance  in  1984  (Table  36),



which  suggests an  additional grazing pressure  on pytoplankton stocks   in

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                                                                     75




Lake Huron.    This  may lead  to changes  in phytoplankton   abundance and



composition  (See Discussion in Lake Michigan on  Trophic  Interactions).



     A top-down effect on zooplankton is likely  in  Lake  Huron.   A   careful



examination  of  the time  trends in  the forage   fish  base   of  Lake  Huron



similar to what was done on  Lake Erie (This Study)  would provide   further



insight on this hypothesis.

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                                                                     76




LAKE ERIE



PhytopIankton




     Species lists (Table A13) and summary tables of abundance (Table A14)



and biovolume (Table  A15) are in  Volume 2  - Data Report.   A summary  of



water chemistry paramters is presented in Table 6.





Annual  Abundance of Major Algal  Groups




      The phytop Iankton  assemblage of 1984 was  comprised of 356  species



representing 104 genera (Table 37).  Compared to 1983,  a 4.3$ reduction in



the number of species  and a 1.0$  increase in the  number of genera  were



observed.  Seventy-five  percent of  the decrease in  species number  from



1983 to 1984 was due to a  decrease in species of Chlorophyta.   The  total



number of species  in 1983 (372)  and 1984 (356)  was considerably  higher



than the 125 to 150  species observed in all   basins in 1970 (Munawar  and



Munawar 1976).




      In 1984 the diatoms possessed  the greatest number of  species  (171,



48$ of the total species) and biomass (47.8$ of the total) (Tables 37  and



38), while the second largest number of  species (96) was observed for  the



Chlorophyta  (Table  37).    A   similar  observation  occurred  in   1983



(Makarewicz 1987).   These  diversity observations  represent  significant



changes from 1970, when  the Chlorophyta possessed  the largest number  of



species (78)  and only  21  diatom species  were  observed (16.3$  of  the



species) (Table 38).   However, diatoms in 1970 still accounted for 53$  of



the biomass (Munawar and Munawar 1976).



     Highest relative densities were attained by the picoplankton  (89.6$)



in 1984.  In 1983 the Chlorophyta had the second highest biomass, while in



1984  they  were   fourth,  si ightly  lower   than  the  Pyrrhophyta   and



Cryptophyta.

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                                                                     77
Seasonal  Abundance and Disbribution of Major Algal  Group?



     The average density  and biomass for the sampling period were  45,080



cells/ml  (40,055  cells/ml;  1983)  and  1.00  g/m3  (1.36  g/m3;  1984).



Seasonally,  abundance (cells/mL)  peaked in mid-April   at 88,762  cells/mL



(mean abundance station),  decreased through May and July, and leveled  off



during August.  A fall/early winter secondary maximum at -40,000  cells/mL



was observed before a decline to 28,200 cells/mL in February of  1985 (Fig.




45a).



       A different  pattern emerged  from the  seasonal  biovolume  totals.



Similar to the seasonal abundance pattern,  peak biomass occurred In April.



However,  biovolume was low in  July and steadily increased into   September



(Fig. 45b) followed by a decrease from December into January and February.



Except for the lower biomass  in 1983 and 1984,  the timing of the  spring



and autumn biomass peaks is similar to that observed in 1970 (Munawar  and




Munawar 1976).



      Diatoms were the  dominant group throughout the  year (47.8$ of  the



total  biovolume).      However,  seasonally   their   importance   varied



considerably (Fig. 46) but in a pattern similar to 1983 (Makarewlcz 1987).



Diatoms were dominant in  April  and May (~6Q%  of the biovolume) and  were



succeeded by the  Cryptophyta in July  and the Chlorophyta  in August.    A



similar  succession  and  relative   importance  were  observed   in   1983



(Makarewicz 1987) and In 1970 (Munawar and Munawar 1976).  By December and



through the winter months,  the diatoms were again dominant accounting  for



as much as 78$ of the biovolume.





Geographical Abundance and Distribution of Major Algal Groups



       Abundance for  the sampling  period varied  geographically but  was



similar  to  1983  observations  (Makarewicz  1987).    Biomass   generally

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                                                                     78

decreased eastward.  The western basfn (Stations 60,  57 and 55)   possessed
                           3                                            3
a greater biomass (1.38 g/m ,  S.E.=.23) than the eastern basin (0.54 g/m ,

S.E.=.82) (Stations  18, 15  and  9) and  the  central  basin  (0.76  g/m3,

S.E.=.09) (Stations 42, 73, 37,  78  and 79) (Table 39).  The  considerably

greater abundance of the western basin was attributed to the  picoplankton

(Fig. 47).  However,  the higher biomass of  the western basin (Table  39)

was  due  to  greater  abundance  and  biomass  of  the   BaciIlariophyta,

Cyanophyta,  Chlorophyta, Cryptophyta and Chlorophyta in the western basfn.

The  increase in the total abundance,  but not in biomass (Table  39),   east

of Station 78 was attributed to the higher abundance of picoplankton (Fig.

47).  Picoplankton contributed little  to community biomass (~1.5$,   Table

40) because of their extremely small size (0.5 to 2.0um).

     As   in 1983, the general  pattern  of higher abundance in the  western

basin was observed on each sampling date except for perhaps the  late  fall

and early winter cruise (Fig.  48).   It appeared that with cooling of  the

lake in the  autumn, abundance  became similar throughout  the lake  (Fig.

48).  In  1983 (Makarewicz 1987),   at least 12  common species had  higher

abundances in the  western basin.    Similarly in  1984, many  of the  same

species had geographical abundance patterns with maxima in the western  or

central  basin  (Table 41)  (Figs.  48  and 49).   A  difference in  species

abundance from  the  various  basins  of Lake  Erie  has  been  documented

previously (Munawar and Munawar 1976, Davis 1969a).


Pi cop Iankton

      Picoplankton abundance in 1984 (mean  = 38,075 cells/mL; maximum  of
        c
3.8 x 10  cells/mL) was not dissimilar from 1983 (mean = 33,171   cells/ml;
                    c
maximum of 1.4 x  10  cells/mL).  On  a numerical basis, the  picoplankton

represented 88.2$  of the  total  cells and  6,1$  of the  total   biomass.

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                                                                     79 ,





Picoplankton relative  numerical  dominance  in 1984  was similar  to  1983



(84.5$ of total cells).  Prior to the 1983 study (Makarewicz 1987),   other



researchers have not routinely reported on this group of organisms.






Regional and Seasonal Trends in the Abundance of Common Species



        Common  species  (Table  40) were  arbitrarily  defined  as  those



possessing a relative abundance  of >0.1$ of the  total cells or >0.5%  of



the total biovolume.  Eighty-four  percent of the common species  observed



in 1984 were also common  species in 1983.   Thirty percent of the  common



species observed in 1983 were not common  in 1984 (Table 41).



      The causes of these differences  is difficult to evaluate.   Natural



annual variability  of plankton  populations in  the lake  has never  been



evaluated and  can  not be  evaluated  until  a longer  data  set  exists.



Considerable seasonal sampling  variability exists between  1983 and  1984



and is the most probable cause for the species differences observed.   For



example, Coelastrum  microporum was  common in  1983 because  of its  high



density  in October  (Makarewicz 1987).   October,   September and  November



samples were not taken in 1984-85.



     Because of the similarity of the 1984 common species list to the 1983



list,  a  species by  species description  of autecology  and regional  and



seasonal trends  are  not  warranted  here  and  can  be  referred  to  in



Makarewicz (1987).   Only new common species are discussed below.






BaciIlariophyta



  Asterione I I a formosa Mass.



      A common species in  Lake Michigan and Lake  Huron in 1983 and  1984



(Makarewicz 1987,  This report), .&. formosa was a dominant species in  Lake



Erie prior to  1950 (Verduin 1964).    Hohn (1969)   stated that A.  formosa

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                                                                     80




maintained constant  densities  between 1938  and  1965 but  its  relative



importance declined.   Between  1967-1975,  a decline  in A-  formosa  was



evident from nearshore data (Nichols et  al. 1977b).   Munawar and  Munawar



(1976), working with  samples from  the entire lake,   observed that  those



species, such as  .A,, formosa,  dominant  before 1950 continued  to be  less



important in 1970.  During February of 1976, .A.  formosa comprised 10.3? of



the total biomass but contributed less than 5% of the total biomass on all



sampling dates in the western basfn (Gladish and Munawar 1980).



     In 1975-76,  h. formosa  was a common species in the central basin  in



early April  (Reuter 1979).  However, it was not a common species in  1983.
Average abundance  and biomass  in 1983  were only  8.7 cells/tnL  and  2.6



mg/m ,  respectively.  It was a dominant species in 1984 (Table 40).



       Average abundance and  biomass in  1984 were 73.4  cells/mL and  48



mg/m ,  respectively.  Maximum abundance was 942 cells/mL at Station 42  on



1 May 1984.  Seasonally, abundance was  high in April  and peaked by  early



May (mean station abundance = 278  cells/mL).  However, abundance was  low



the rest of the year (Fig. 51a).






  Melosira islandica 0.  Mull.



      Historically,  M.  is Iandica has  not been a  common species  in  Lake



Erie.  Michalski  (1968) noted  it  as sub-dominant during  the vernal  and



autumnal period from nearshore  data In 1966-67.   Similarly, Nicholls  et



al. (1977b) believed it to be a spring species between 1967 and 1975.   In



1970 M. islandica  represented 27.5$  of the  total biomass  on the  21-26



October cruise  of Lake  Erie (Munawar  and Munawar  1976).   Giadish  and



Munawar (1980) did not report this species as common in the western  basin



 in 1975-76; E.  granuiata was common  in 1975-76.    Similarly In 1983,  M.

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                                                                     81




granulata was common (Makarewlcz  1987) while E.  tslandica was not  (mean



abundance = 2.9 cells/mL; mean biomass = 3.0 mg/m ).



      In 1984 this mesotrophic  indicator species (Tarapchak and  Stoermer



1976) was the  fourth most common  diatom on a  biomass basis (Table  40).



Average abundance was 31.5  cells/mL with a maximum  of 1,564 ceils/mL  at



Station 55 on  20 April  1984.  Abundance  peaked in  April (mean  station



abundance = 190 cells/mL)  and was low  the rest of  the year (Fig.  51b).



Geographically, abundance was definitely higher in the western basin (Fig.



49).  The high abundance of M, isiandica in the western basin of Lake Erie



is correlated  with the  spring bloom  of this  species in  southern  Lake



Huron.






Chlorophyta



  Crucigenia rectangularts (Brawn) Gay



       This species  is usually a  minor element  of summer  phytoplankton



assemblages of  mesotrophic  to  eutrophic lakes  (Stoermer  and  Ladewski



1976).   Historically,  this  species has  not been  common in  Lake  Erie.



Abundance  in 1983 (mean  = 1.9 cells/mL) and  1984 (mean = 5.1   cells/mL)



was low  .    Because of  its relatively  large size,  ft became  a  common



species accounting for 1.01? of the total biomass (Table 40).   Seasonally,



abundance peaked in August (mean station abundance = 41.7 cells/mL)  (Fig.



51d).






Cyanophyta



  Anabaena sp.




     Starting in 1958,  Anabaena became more prevalent  during the fall   and



summer at  least through  1963  (Davis 1969a).    In  1966-67,   short-lived



summer pulses  of  Anabaena  were  observed  at  a  nearshore  station  by

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                                                                     82




Michalski (1968).    Munawar  and Munawar  (1976)  observed  populations  of



Anabaen^ spiroides to be "weI I  developed" in both the western and  central



basins during  the summer  of  1970.    During  1975-76,  Cyanophyta  biomass



never exceeded 20  mg/m .   Aphan izomenon fIos-aquae was  the most  common



taxa encountered in 1975-76,  wh ile Anabaepa sp. occurred less commonly  in



the central  basin (Reuter  1979).  Although species  are not mentioned,   a



decrease in Cyanophyta biomass  was observed at  a nearshore site  between



1967 and 1975 (Nichols  et al.  1977b).    Abundance in  1983 was low  (mean



abundance = 15.1 cells/ml) (Makarewicz 1987).



      In  1984 mean  abundance was 47.8  cells/mL.  The  percent of  total



biomass (0.87$) for Anabaena sp. was the same as Aphanizomenon  flos-aquae



during the 1984  study (Table 40).    Seasonally,  abundance  peaked at  255



cells/mL (mean station abundance) on 7 August 1984 (Fig. 51c).  A  maximum



abundance of 867 cells/mL  was observed at Station  55 on 19 August  1984.



No obvious geographical pattern was observed.






Changes in Species Composition



     Davis (1969a) has  reviewed the extensive earlier work on Lake  Erie,



while Munawar and Munawar (1982), Gladish and Munawar (1980) and  Nicholls



(1981) discuss the  more recent  material.  Verduin  (1964) has  concluded



that before 1950 the phytoplankton of western Lake Erie had been dominated



by Asterionella  formosaf  TabelI aria  fenestrata  and  Melosira  amb igua,



whereas fn  1960-61  the  dominant forms  had  been  fragilaria  capucina,



Coscinodiscus radiatus (probably  Actinocyclus normanii  f. subsalsa)  and



Melosira b inderana (=Stephanod tscus b inderanus).



     As with Munawar and Munawar (1976), the 1983 study (Makarewicz  1987)



confirmed Verduin's (1964) observations that those species dominant before



1950 (.A.,  formosa, J.  fenestrata and  .M,. ambigua)  continued to  be  less

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                                                                     83




important in  the 1983  collections.   Act i nocycI us normanIi  f.  subsalsa



(=Coscinodi sous roth i i)  and StephanodIscus  binderanus were  dominant  Fn



1961-62 (Verduin 1964) and in 1970 (Munawar and Munawar 1976).  FragiI aria



capucina was a dominant  in 1961 but  not in 1970.   By 1983  Actinocyclus



normani i f.  subsalsa was  only the fifth most  prevalent diatom, but on  a



numerical  basis FragiI aria capucina was  the second most prevalent  diatom



in the western basin  and in the entire  lake (Makarewicz, 1987). In  1984



Actinocyclus normanli  f.  subsalsa was  not even a  common species  (Table



40).



       Dominant species  in 1983  and 1984  were Stephanodiscus  nlagarae,



Fragi I aria  crotonensis,,  Cosmar i urn  sp.,  Cryptomonas  erosa,  Rhodomonas



mJnu.ta var.   nannopl ankt ica,  Osci I latoria  subbrevis, Osci I lator ia  tenuis



and Qeratium  h irupdinel I a (Table  40).    Fragi laria capucina,,  Coel astrum



mlcroporum,  Osci I Iatoria subbrevis  and Q.  tentiIs were  dominant in  1983



only (Makarewicz 1987),  while  Anabaena sp.,  Aphanizomenon flos-aquae  and



AsterionelI a formosa were also dominant in 1984,



     Aster?one!I a formosa has not been prevalent in Lake Erie since  prior



to 1950.  Verduin (1964) stated that before 1950 AsterionelI a formosa  was



a dominant  species  in  western  Lake Erie.     Similarly,  Davis  (1969a)



reported AsterionelI a as the dominant organism in the spring pulse of  the



central basin prior to 1949.   Numerous workers (Hohn 1969, Nichols et  al.



1977b,  Munawar  and Munawar  1976,  Gladish  and Munawar  1980) reported  a



decl ine in .A,, formosa  after 1950.    The low  abundance  of A- formosa  was



apparent into 1983  (mean = 8.7 cells/mL, Makarewicz 1987).



     Average density of A.  formosa was 73.4  cells/mL in 1984  representing



5.6$ of the  biomass (Table 40).   Maximum density in March of 1938 was 96.6



cells/mL with a March  mean of 553  cells/mL  (Hohn 1969).  No samples  were

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                                                                     84




taken in March of  1984, but the April  average was 226 cells/ml  (maximum



abundance = 942 cells/ml in  May ).   In 1984  during the three cruises  in



April and May, Asterionella formosa was  the dominant spring species on  a



biomass basis and the  second most important diatom  on a numerical   basis



(Table 43).




     Although occurrences of common and dominant species in 1970, 1983 and



1984 were similar, dramatic decreases  in abundance of these species  were



evident (Table 44).  This pattern was evident in all three basins.








Indicator Species




          Munawar  and  Munawar  (1982)  concluded  that  the  species  of



phytoplankton found in 1970 usually occurred in mesotrophic and  eutrophic



conditions.    Common  species  in  1983  included  eutrophic    indicators



(FragiI aria  capucina,   Melosira  granulata,   Peridinium   aciculiferum,



Pediastrum  s i mp i exf   Scenedesmus  ecornis)   and  mesotrophic    indicators



(Stephanodiscus niagarae, FragiI aria  crotonensisf TabelI aria  flocculosa)



(Makarewicz 1987).   A similar  set of  major common  species occurred  in



1984,  including  the  mesotrophic  indicators  Stephanodiscus   n iagarae,



FragiI aria  crotonensis  and  TabelI aria  flocculosa  and  the   eutrophic



indicators Fragi I aria  capucina.   Per id in ium actcul iferum  and  Pediastrum



simplex.    The eutrophic  indicators  Melosira granulata  and  Scenedesmus



ecornisf  common in 1983, were present  in 1984 but were not common  (>0.1$



of the total   cells or >Q.5%  of the total   biovolume).   Interestingly,   a



mesotrophic indicator, Melosira island lea,  not common in 1983,  was  common



in 1984 accounting for 4.1? of the total  biomass (Table 40).  However, the



abundance of  M. islandica in western Lake Erie appears to be influenced by



the Lake Huron M,  islandica population.

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                                                                     85 .





      Evidence of a shift  in trophic status since  1970 is provided by  a



comparison of distribution of dominant  diatom indicator species in  1970,



1983 and 1984 (Table  45).  The number  of dominant eutrophic species  has



decreased, while the number of dominant mesotrophic species has increased.



The mesotrophic-eutrophic ratio suggests a shift to mesotrophic conditions



for the western basin.





Historical Changes in Community Biomass




     Between 1927 and  1964,  a large and consistent increase in the  total



abundance of phytoplankton of the central  basin had occurred (Davis  1964,



1969a).   Nichols et  al. (1977b)  observed that  a decline  in  nearshore



phytoplankton of  the  western  basin  occurred  between  1967  and  1975.



However, Gladish and Munawar (1980) discounted this finding and  suggested



that no realistic conclusion could be  drawn from a comparison of  biomass



between 1970 and 1975.



     The mean basin weighted  biomass was 3.4, 1.49 and 0.8 g/m  in  1970,



1983 and 1984,  respectively.   A 56  to 76$ reduction in algal  biomass  has



occurred in  offshore waters  of Lake  Erie from  1970 to  1983/84.   This



reduction in biomass  is evident for  all  seasons of  the year (Fig.  52).



The historically highly productive western basin (Munawar and Burns  1976)



has had a steady decrease in biomass  from 1958 to 1984 (Fig.  53).    Since



1975 chlorophyll concentrations  have decreased in  all basins (Fig.  54).



Phosphorus levels  have  also  decreased  in  all   sub-basins  (Fig.  55),



Between 1970  and 1983-1984,   dramatic reductions  In maximum  biomass  of



common species have  occurred (Table 44),    For example,   in the  nuisance



species Aphapizomenpn f los-aquae,. a 96%  reduction in the maximum  biomass



observed has occurred since 1970.  Stephanodiscus binderanusf  a  eutrophic



indicator species,  has decreased in biomass  by 90$ in the western  basin.

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                                                                    86

Similary, Frag!I aria capucina,  another eutrophic indicator,  has  decreased

(99% reduction) dramatically within the phytopiankton community.

     Based on maximum biomass concentrations (Vollenweider 1968),   Munawar

and Munawar (1976) classified the  western basin as highly eutrophic,   the

eastern basin as mesotrophic and the central basin between the mesotrophic

and eutrophic  conditions.     Using  the  same  classification  system  of

Vollenweider (1968):

               Ultraol igotrophic    <1 g/m     ,
               Mesotrophic           3 to 5,g/m
               Highly eutrophic     >10 g/m

the western basin (maximum biomass = 6.6 g/m ,  Station 55, April)  in  1984

would be between  mesotrophic and  eutrophic,  the  central basin  (maximum

biomass =  3.0 g/m ,  Station 37,  August) would  be mesotrophic  and   the

eastern basin (maximum  biomass = 2.0  g/m , Station 15,   April)  would  be

between oligotrophic  and  mesotrophic.    Similarly,  the  classification

scheme of Munawar and Munawar (1982), based on mean phytopiankton  biomass,

suggests an improvement in water  quality between 1970 and 1983/84  (Table

46) in all basins of Lake Erie.

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LAKE ERIE



ZoopIankton



Annual Abundance of Zooplankton Groups



     Species lists (Table A16) and summary tables of abundance (Table A17)



and biomass (Table A18) are  in Volume 2 -  Data Report.  The  zoopIankton



assemblage of 1984 comprised  81 species representing  39 genera from  the



Amphipoda,  Calanoida,  Cladocera,   Cyclopoida,  Harpacticolda  and   the



Rot ifera.  Compared to 1983 (37 genera, 66 species), an 18.5$ increase  in



number of species was observed.   This difference was mostly  attributable



to an increase in number of rotifers (34 to 48).



       The  Rot ifera  possessed the  largest number  of species  (48)  and



relative abundance (80.1?) followed by the Cyclopoida and Calanoida.    The



nauplius  stage  of  the  Copepoda  accounted  for  10.4?  of  the   total



zooplankton abundance (Table 47).  On  a biomass basis, the importance  of



the Rot ifera dropped to 13.6$ of the zooplankton biomass because of  their



small size, while the  Cladocera contributed 40.5?  of the biomass  (Table



47).   Average density and biomass for the study period were 159,615+34,000



organisms /m3 (mean±S.E.)  (288,100/m3 - 1983)  and 53.6±6.2 mg/m3  (Table



6).





Seasonal  Abundance gnd Distribution of Major Zooplankton Groups



     Seasonally,  biomass distribution (Fig. 56a) was unimodal,  peaking  in



August.   The seasonal abundance pattern suggested two peaks: one in spring



and a second  in late summer  (Fig. 56b),   which were caused  by peaks  in



rotifer abundance.    A sampling  pattern that includes  the June-July  and



September-October  period  is  needed  to  fully  evaluate  the   seasonal



distribution patterns.

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                                                                     88




        The 1984  seasonal   abundance pattern  (Fig.   57) of  the  various



zooplankton groups  was  similar  to 1983  (Makarewlcz  1987).     Rotifera



abundance peaked in  May and  a secondary peak  was noted  in late  August



(Fig. 57).  Cladocera and Calanolda abundance was low in spring,  peaked in



early August and  decreased the  rest of  the year.    Cyclopoida  achieved



their highest abundance in late August  (Fig. 57a).   The biomass  seasonal



distribution pattern of  the major zooplankton  groups generally  mimicked



the abundance pattern (Fig. 58).






Geographical Abundance and Distribution of Zooplankton Groups



     Geographically, zooplankton abundance was similar to 1983 (Makarewicz



1987), with abundance  being higher  in the western  basin and  decreasing



easterly to Station 79 (Fig.   59).  Abundance increased slightly  eastward



through the eastern basin (Stations 18, 15 and 9).   The Rotifera were  the



cause of the high zooplankton abundance in the western basin,  although the



Copepoda nauplii also had a slightly higher abundance in the western basin



(Stations 60,  57, 55) (Fig. 59b).



      Interestingly, biomass was  similar in all  three basins of Lake  Erie



(Fig. 60a) even though Rotifera biomass was highest in the western  basin,



particularly et the most western Station 60.   The high rotifer biomass was



countered by a low  Cladocera biomass at Station  60 (Fig. 60b),  while  at



the next easterly  station (57),  Rotifera biomass was  low and  Cladocera



biomass was high.  A low Cladocera abundance was observed at Station 60 in



1983  (Makarewicz 1987).   Perhaps  there is  an influence  of the  Detroit



River at  this station  that affects  Cladocera abundance  negatively  and



Rotifera positively.



       Except  for  Station 60,  Cladocera abundance  generally  decreased



eastward  into  and through  the  central basin.     In the  eastern  basin,

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                                                                     89




Cladocera biomass  (Fig. 60b),  but not  abundance (Fig.  59b),   increased



easterly.  Cyclopoida and  Calanoida abundance was  higher in the  central



and eastern basin as compared to the western basin.



Common Species




     Common Crustacea species (Table 48) were arbitrarily defined as those



possessing a relative abundance  >0.1$ of the total  abundance or 1.0$  of



the total  biomass.   Rot ifera  species  were considered  common  if  they



accounted for >1.0$ of  the total zooplankton abundance  or biomass.   The



number of common species in 1983 (25) and 1984 (27) was similar, but there



were changes in  composition of  the common species.    Daphnia  pulicaria,



common in 1984, was not observed in 1983 in Lake Erie.  Leptodora kindti if



KeratelI a earlinae and Notho lea squamulaf common in 1984, were present  in



1983 but not common.    Common species observed in  1983, but not  in  1984,



included Diaptomus sic!loldes, Diaphanosomg Ieuchtenbergianumf  Colletheca



sp. and Ke11?cott i a longispina.






Changes in Species Composition



Crustacea



      Brooks  (1969)  suggested that  a shift in  the Lake Erie  cladoceran



assemblage was  evident  by  1948-49 with  smaller  cladocerans,  such  as



Daphn ja galeata mendotae.  D.  retrocurva  and Diaphanosoma sp., being  more



abundant than in 1938-39.   In 1970 the most commonly  found Daphnia species



were D.  retrocurva,  Q.  galeata mendotae  and D.  longiremis (Watson  and



Carpenter 1974)j  Bosmina longirostris  and Eubosmina  coregoni were  more



abundant (Watson and  Carpenter 1974).    Predominant cladoceran species  in



1983 were small   forms similar to  those observed  in 1970.    In 1983  the



predominant Cladocera in descending order were Eubosmina coregon i f Daphnia



gaIeatq mendotae,  Bosmina longirostris, Diaphanosoma  Ieuchtenbergfanum and

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                                                                     90




Chydorus sphaericus (Makarewicz 1987).  In 1984,  on a numerical  basis,  the



predominant Cladocera were Daphnia  galaeta men dot ae f Eubosmina  coregoni,.



Bosmina longirostris,  Daphnia pul icari a,   Daphnia retrocurva and  Chydorgs



sphaericus (Table 48).  Between 1983 and 1984,  essentially the same common



species, with the  exception of  D. pul icaria,  were  present with  minimal



change  in rank abundance.  These  changes in rank order may be  attributed



to the difference in the seasonal  sampling pattern between 1983 and 1984.



     On a biomass basis, Daphnia pul icaria was the dominant Cladocera  for



the lake,  with  a major bloom  in  August.   It was most  prominent in  the



central and eastern basins (Fig.  61a).  A reexamination of the August 1983



samples revealed that JD. pul icaria was present.  Apparently the taxonomist



included this species under Daphnia spp.  in the 1983 counts (N.   Andresen,



Personal Communication).   The existence of  the Iarge J).  pul icaria is  a



major finding.  This species was  first observed in Lake Michigan in  1978



(Evans  1985) and was a dominant species  in 1983 in Lake Michigan and  the



third most important cladoceran in  Lake Huron in 1983 (Makarewicz  1987).



The occurrence of this  species in  large  numbers in Lake  Erie may be  an



important factor, along with decreasing phosphorus loading, in  explaining



the decreasing phytoplankton abundance observed in Lake Erie  (This study).



Large populations of Daphnia pul icaria have been correlated with low algal



biomass (Osgood 1983,  Vanni 1983).



     A  rare species in the offshore waters of the western basin  in 1929-30



(Tidd  1955), Chydorus sphaericus was a prominent constituent  in the 1950's



(Davis  1962) and  in 1970  with a higher  abundance  in  the western  basin



(Watson and Carpenter 1974).   In  1983 and 1984,  this species  contributed



0.2$ and  0.1$, respectively,  of the  total abundance   (Makarewicz  1987)

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                                                                     911




 (Table 48).    Chydoriis sphaerlcus  has  established  Itself  as  a  common



 species  in Lake Erie.



       The prevalence of  Cyclops vernal Is  has changed over  the past  50



 years.   In the 1930's, £. vernal is  was  found only  in the extreme  western



 end of Lake  Erie at  the mouth  of the  Detroit and  Maumee Rivers  (Tidd



 1955).  By 1967 it had spread throughout the lake (Davis 1969b).   Fatal as



 (1972) and Watson (1976) reported it  as numerous in the western basin  of



 Lake Erie during the  late 60's and 70's. This species was not observed  in



 1983 (Makarewicz 1987), while in 1984 it was not common (Table 48) but did



 average 25.9  organisms/m  for  the entire  lake.   However, It  was  more



 prevalent  in the  western basin  (83/m ) as  compared to  the eastern  and



 central basins (3.3/m  ).



       The dominant  cyclopoid copepod   in 1970  was Cyclops  bicuspIdatus



 thomasi with tyesocycjpps edax common  In the summer (Watson and  Carpenter



 1974).   Cap (1980)  documented a  shift in  predominant copepods  in  the



 eastern basin from calanoids in 1928 to cyclopoid copepods, mainly Cyclops



Mcuspidatus t homas i,  in 1974.   Trppocyclops pr 35 ings was present in  low



 numbers  (Watson and  Carpenter 1974).    In 1983 and  1984,  the same  three



 species  (£. JjLcuspidatgs  thomasi,  N|. edax  and J. praglnus)  predominated



 (Makarewicz 1987) (Table 48).



     Abundance of Diaptomus siciIoides has increased in Lake Erie  (Gannon



 1981).   It was most prevalent in the western basin and western portion  of



the central basin in the late  60's and 70's (Patalas 1972, Watson  1976).



Abundant diaptomids  in  the  eastern  and central  basins  in  1970  were



Diaptomus oregonensis and J2. siciloides,   which were also the  predominant



calanoids in Lake Erie in 1983 and 1984  (Makarewicz 1987) (Table 48).    D.

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                                                                     92






sic!loides was not a  common species (1.0$ of  total  zooplankton)  in  1984



but was the second most abundant calanofd.



Rot i fera



     Davis' studies (1968,  1969a) of the zooplankton  of Lake Erie  included



rotifers.  Certain soft-bodied rotifers  were not identified nor were  the



samples quantitative  for  rotifers  as  a number  20  net  was employed.



However, it is apparently  the only lake-wide study  of the offshore  that



included the rotifers.



     Species observed to be abundant in 1967 were Brach ionus angular is, E.



calyciflorus,  Conoch iI us  unicornis,  KeratelI a  coch|ear is,  J< quadrata,



KelIicottia Ipngisp inaf Synchaeta sty I ata  and Polyarthra vulgaris  (Davis



1968, 1969a).    In  1983  a similar group  of abundant  rotifers was  found



(Makarewicz 1987).  In decreasing order of relative abundance ($ of  total



abundance), the  abundant  species  in 1983  were:    Polyarthra  vulgaris



(18.4$), Synchaeta  sp. (9.5$),  Keratella cochI ear is  (7.3$),  ConochiI us



unicornis  (5.3$),  Keratella  hiemalis   (3.5$),   Rrachionus  sp.    (3.0$)



(Makarewicz 1987).  Polyarthra vulgaris (22.49$)  and Synchaeta sp. (9.46$)



were still dominant in 1984  along with Notholca sguamula (11.06$),  which



was not  a  common  species  in  1983.   Other  abundant rotifers  in   1984



included  Polyarthra  major  (4.94$),  Keratella  cochI ear is  (4.91$)  and



Notholca laurentiae (3.21$) (Table  48).  Except for  the addition of  the



species of Notholca   in 1984,  the   1983 and  1984  rotifer composition  was



similar to  1967.   Although   it was  only  the fourteenth  most  abundant



rotifer, KelIicottia  longispina was still prevalent  in 1983, but not 1984,



representing  1.3$  of  the  total  abundance  (Makarewicz  1987).     Only



Keratella quadrata  was apparently not as  prominent in  1983 and 1984 as  it



was  in  1967.

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                                                                     93
East-West Species Distribution



      Numerous researchers (e.g. Davis  1969b, Watson 1974,  Fatal as  1972,



Gannon 1981) have  documented the differences  in species composition  and



abundance from the central, western and  eastern basins of Lake Erie.   As



in 1983 (Makarewicz 1987), a number of species, all  rotifers in 1984,  had



higher abundances in  the western  basin (Figs.  59a&b).   Geographically,



Cyclops bicusp idatus thomast, Mesocyclops  edax and Diaptomus  oregonensis




had geographical abundance patterns  with maxima in  the central  basin  !n



1983 (Makarewicz 1987) and  1984 (Fig. 61a).   Ho I opediurn gibberum  (1983)



and Tropocyclops prasinus mexicanus (1983 and 1984) were more prevalent in



the eastern basin (Fig. 61b).





Indicators of Trophic Status



       Zooplankton have  potential  value  as assessors  of trophic  status



(Gannon and  Stemberger  1978).   Rotifers,   in particular,   respond  more



quickly to  environmental  changes than  do  the crustacean  plankton  and



appear to be sensitive Indicators of changes in water quality (Gannon  and



Stemberger  1978).    Brach ionus   angularisf  E«  calyciflorus,    FiIinia



long iseta and Trichocerca multicrinis are four rotifer species  indicative



of eutrophy.  Also,  species in the genus Brach ionus are particularly  good



indicators of eutrophy   in the Great  Lakes (Gannon 1981).   Of the  three



dominant rotifer species in Lake Erie, £.  vulgar is Is a eurytopic species;



Notholea  squamuls   is   a   cold  stenotherm   often   associated   with



oligo-mesotrophic lakes  (Gannon and  Stemberger 1978)  during the  summer



that is also  often encountered in  eutrophic lakes during  the winter  or



early spring (as In Lake Erie  in 1984); and some species of  Synchaeta  are



eutrophic indicators (Gannon and Stemberger 1978).   The lack of  dominance



of eutrophic indicator species for the entire lake suggests that Lake Erie

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                                                                     94




in 1984, as  a unit, is  not eutrophic.   This would agree  well  with  the



conclusion from the  phytoplankton indicator  species and  from the  algal



biomass classification of trophic status of Lake Erie..



          However,  the  eutrophic  indicators  Brach ionus  caudatus,   E.



calyciflorus, fi.. angular is,  F i Ii n i a longiseta, Trichocerca muI tier in is and



Trichocerca cylindrica  had  abundances  restricted  to   or  significantly



higher  in the western basin (Table  49).  Total  zooplankton abundance  was



also higher  in the  western basin.    As  with phytoplankton  biomass  and



species  composition,  both  rotifer  abundance  and  species  composition



indicated a greater degree  of  eutrophy in the  western  basin than in  the



central or eastern basins.




       Another measure  of trophic  status is  the caIanoid/cylopoid  plus



cladoceran ratio (plankton ratio) (Gannon and Stemberger 1978,  McNaught et



al. 1980a, Krieger 1981).   Calanoid copepods generally appear best adapted



for oligotrophic conditions, while cladocerans and cyclopoid copepods  are



relatively more abundant in eutrophic waters (Gannon and Stemberger 1978).



In Lake Erie,  this ratio increased  from west  to east  in  1983 and  1984



(Table 50) indicating a  more productive status for  the western basin  as



compared to the rest of the lake.



     The higher algal biomass (Table 39) of the western  basin as  compared



to the central  and eastern basins was reflected in the higher abundance of



zooplankton,   eutrophic  zooplankton  species  composition  and  the   low



plankton ratio.  Compared  to Lakes Huron and  Michigan  in 1983 and  1984,



abundance of zooplankton was greatest and the plankton ratio



was lower in Lake Erie (Table  6),  indicating the higher trophic status of



Lake Erie compared to Lakes Huron and Michigan.

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                                                                     95




Historical Changes fn Abundances



     Zooplankton data exists for the western basin of Lake Erie from  1939



to 1984.  The 1939 (Chandler  1940; 49 collections),  1949 (Bradshaw  1964;



30 collections)  and 1959  (Hubschmann 1960;  daily collections  July  and



August) collections were taken with a 10-1 iter Juday  trap equipped with  a



64-um mesh net in the western basin.  A 1970 study by Nalepa (1972) is not



included  in the analysis  because it is  from the far  western end of  the



basin and may not be representative of the entire western basin.   The 1961



study of  Brltt et  al.  (1973) sampled  twice  monthly from  mid-June  to



mid-September, while Davis (1968) used a  76-um mesh  net in July  of  1967.



Because of the comparable net sizes, all these studies, with the  exception



of Nalepa's (1972), are comparable to the 1983 (Makarewicz 1987)  and  1984




surveys.





      A comparison  of the ApriI-December Crustacea  means of 1939,  1949,



1983 and  1984 suggests an increase  in zooplankton abundance from 1939  to



1949 (Fig. 63).   Similarly,  the mean abundance  for  July and August  from



1939 to 1961 suggests a similar  increase in zooplankton (Fig. 64).   Both



Bradshaw  (1964) and Gannon (1981)  concluded similarly.  Average   ice-free



abundances from  1949  to  1983 suggest  a  decreasing  but  insignificant



downward trend (Fig.  63).   A major decrease  in zooplankton abundance  is



suggested from 1983 to 1984 (Fig. 63).   It is difficult to evaluate  this



drop in biomass because of  the large gap in data  from 1950 to 1983.   It



could simply  be  annual  natural   variability.   However,  the  Huron  and



Michigan zooplankton abundance  did not display  such a great  variability



from 1983 to  1984.   Focusing on  July and  August,  where  more  data  are



available,  an  abundance  decrease  in  Cladocera,  Copepoda  and   total



Crustacea from the 1961 maximum (Fig,  64) is evident.

-------
     A data point in the  early 70's would be of interest.   Data do  exist



for the 70's.   However,  Nalepa's (1972) study  is from  the far  western



portion of the  western basin.    Watson and Carpenter  (1974) sampled  the



western basin, as welI  as the central  and eastern basins in 1970.    Their



data is  reported as  a weighted  lake  average and  is not  available  to



compare with other  years in the  western basin.   As the sampling   method



(1970; vertical hauls,  64-um mesh) is comparable to those used in 1983 and



1984, these data  are also directly  comparable on a  lake-wide basis.   A



seasonal comparison of  weighted lake-wide means suggests little change  in



zooplankton abundance from  1970 and  1983 during the  spring and  autumn.



However, 1984 values are  generally lower than 1983  and 1970 data   points



(Fig. 65).   The importance of   a sampling point  between mid-May  through



July in 1983 and 1984,  the generally recognized period of peak  abundance,



is apparent from this figure.




     The  1939 and 1961 rotifer samples  were collected with a 64-um  mesh



net, as in the 1983 and 1984 works.  An increase in Rot ifera abundance  in



the western basin is suggested  since 1939 (Fig. 66).





Trophic Interactions



     Long-term  changes of  phytoplankton and  zooplankton abundance  were



apparent.   A 56 to 76$   reduction in lake-wide offshore algal biomass  has



occurred from 1970 to  1983 to  1984.   Total phosphorus and chlorophyll  .a



levels  in  each  basin   decreased  (Figs. 53  and  54).  Similarly,  where



comparable data are available,  zooplankton abundance and biomass decreased



in the western  basin,  while  a decrease in  lakewide zooplankton  biomass



during the summer  period from  1970 to 1984   is suggested.  With the  N/P



ratio  currently exceeding 30 to 1, apparently due to P-control,   nuisance



blue-green algae  species,  such as  Aphanizomenon  flos-aquaf  decreased.

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                                                                     97





These changes  are  consistent  with expectations  of  long-term  nutrient



control .



     There are,  however, significant  changes in the  composition of  the



zooplankton community  that  can  not be  attributed  solely  to  nutrient



control.  The appearance of the large cladoceran Daphnig pulicar Ia in Lake



Erie was evident in 1983  and 1984.  Its dominance  with a major bloom  in



August of 1984 was surprising for it suggested  changes in planktivory  in



Lake Erie (Wells  1970, Brooks  and Dodson  1965, Carpenter  el al.   1985,



Scavia et al. 1986).



     A recovery  in the  walleye fishery of  Lake Erie is  evident by  the



increasing harvest  and abundance  (Fig.  67).    Annual   walleye  harvest



rapidly increased from 112,000 fish in 1975 to 2.2 million fish in 1977 in



the Ohio Lake Erie waters (western and central basins) (Ohio Department of



Natural Resources 1985).   Annual harvests since 1978 have stayed high  but



ranged from 1.7 million to the record 4.1 million in 1984 (Ohio Department



of Natural  Resources  1985).    Central  basin  harvests  have  increased



dramatically over the past  two years (Fig. 68).   The initial  recovery  of



the walleye fishery is attributed to the closing of the walleye fishery in



1970 due  to mercury  contamination  and to  the exclusion  of  commercial



fishing for walleyes in U.S. waters since 1972 (Kutkahn et al. 1976).



        In  addition,   salmonid  stocking  programs  exist  In  New   York,



Pennsylvania, Ohio and Ontario.  New  York,  which has the largest  stocking



program, has a target  stocking of -1  million  fish in 1987  (F.Cornelius,



Personal  Communication). Lake trout,  Chinook and Coho salmon  and  various



strains of rainbow/steel head trout are  stocked in New York waters.   These



fish are primarily feeding on smelt (NYSDEC 1987).

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                                                                     98




     Seasonal   diets of walleye  closely followed  changes in  forage-fish



availability (Knight et al.  1984).    Between 1979 and 1981 in the  western



basin of Lake Erie,   walleye ate (100$ by  volume) age-1  shiners  Notropis



atherinoldes (emerald shiner) and U.  hudsonius (spottail  shiner) in spring



but switched to age-0 clupeids (60-90$) Dorpsoma cepedianum (gizzard shad)



and Alosa pseudoharengus  (alewife) in  late July.   Clupeids and  shiners



composed 25-70$ and 10-40$,  respectively,  of  the diets of age-1 or  older



walleyes in  autumn (Knight  et al.  1984).   There does  appear to  be  a



difference  in walleye foraging from west to east.  Recent stomach analyses



of walleye  from New  York  and Pennsylvania  waters indicate  that  smelt



represent  90$  of  their  diet  (NYSDEC  1987  and  R.  Kenyon,  Personal



Communication,).   Smelt  are not  abundant  in the  western  and  central



basins.



    Dramatic changes have occurred  in the forage species of Erie.   It  is



apparent that alewife, spottail shiner and emerald shiner have declined in



the western and central  basins (Fig. 69) and  in Pennsylvania waters  (R.



Kenyon, Personal  Communication).   The decline  of spottail  and  emerald



shiners between 1982-1984 is impressive in view of the massive  increase in



walleye harvest   in the  central  basin since  1982  (Fig. 68).    Fishery



biologists  have no  specific reason for  this decline. Besides  predation,



other possible causes of the  decline  include climatic factors,  turbidity



changes, toxic chemicals  and the commercial  bait industry. Whatever  the



cause, a decrease In planktivorous shiners has occurred.



     Emerald and  spottail shiners feed  heavily on microcrustacean,  some



midge  larvae and  algae   (Scott and Grossman 1973,  Smith  and Kramer  1964,



McCann 1959).   Evidence  gathered  by Gray  (1942)  in Lake  Erie  during



December  indicated that Diaptomusf Daphnia,  Cyclops and Bosmi na were  a I I

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                                                                     99




 important  in the diet of the emerald shiner but at different times of  the



 day.  Dymond  (1926) noted that  in the spottail  shiner of Lake  Nipigon,



 Daphnia formed 40$ of the diet  although Bosmfna, Sidg and Leptodorg  were



 also eaten.  A study  on current  shiner diets   would  be useful  to  the



 ongoing discussion on trophic interaction in Lake Erie.



     There  is good evidence that pianktivorous fish abundance has  changed



 as a result of the walleye  resurgence but perhaps also from the  salmonid



 stocking program in Lake  Erie.   Release  from planktivore pressure  could



 have led to the establ ishment of the large Daphnia pulicaria in Lake Erie.



 Other top-down effects are difficult to evaluate.  For instance, a clearer



 water column, as  observed in  Lake Michigan and  attributed to  cascading



 effects (Scavia et al. 1986), is  difficult to evaluate in Lake Erie.  For



 example, the decrease  in Aphanizomenon  flos-aquge in Lake  Erie is  more



 readily  attributed  to   decreased  phosphorus   concentration  and   the



 increasing N/P ratio (Smith 1983) than the influence of large  zooplankton



 such as Daphnia  pulicaria on  the phytoplankton  assemblage (Lynch  1980,



Bergquist et  a I   1985).     However,  the  reappearance  and  dominance  of



Aster tone I  I a formosp  in  1984  may  be related  to  the  presence  of  fi.



 pulicarjg  (e.g.  Bergquist  et al. 1985).    In an  ecosystem  dominated  by



 large and more efficient herbivores,  such as Daphnia pul icaria,. a  grazing



effect on phytoplankton would be expected.



    The  index of dispersion (Elliot  1971)  indicates a highly  contagious



 distribution of  phytoplankton  and zooplankton  In Lake Erie.   Could  the



patchy distribution of phytoplankton   be related to zooplankton  herb ivory



on phytoplankton;  that is,   was  there top down  control (i.e. grazing)  on



phytoplankton  on  a  short-term  basis?     Table  51   lists   correlation



coefficients  of   phytoplankton   abundance  versus  total   phosphorus  and

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                                                                    100




zooplankton abundance for each  cruise on Lake Erie.   For each cruise,   11



stations were sampled covering the entire length of the lake over a  short



period of time.   Interpretation of  the correlations were  as follows:  A



negative correlation between a zooplankton group and phytoplankton implied



grazing pressure on  phytoplankton, while a  positive correlation  between



total phosphorus and phytoplankton abundance would suggest an  enhancement



of phytoplankton abundance due to phosphorus.



     All correlations were positive  in April, suggesting that  phosphorus



was  influencing the food  web. A different situation  was evident by  May.



Phytoplankton were blooming (Fig. 45) and all zooplankton groups increased



in  abundance  (Fig.  56).     High  negative  correlations  existed   for



zooplankton suggesting a  top-down influence  on phytoplankton  abundance.



Interestingly, a negative correlation existed for TP versus  phytoplankton



implying    that   phosphorus  was  not   the  major  factor   controlling



phytopl ankton abundance  on  this  spring  date.   As  expected,  when  B..



pulfearia became dominant   in August, a  negative fairly high  correlation



existed betwen J3. pulicgria and  phytoplankton.  During this same  period,



Daphnia spp.,  in general,  and Rot ifera  were not  negatively  correlated



spatially with changes   in phytoplankton  abundance.   By December,  other



species of Daphnia and Calanoida  exerted some  influence on  phytoplankton



abundance.



      Calanoids were  negatively correlated  with phytoplankton  abundance



throughout the  year,  except  in  April,  suggesting a  constant  baseline




effect on phytoplankton  abundance.   In a  lake such as Erie, where a  large



efficient Daphnia sp. is  added to the food  web, the new species   induces



grazing pressures previously not present  during the summer.  Thus  during



the  summer, a greater  grazing pressure  leads to  a decrease  in algae,  an

-------
                                                                    101




increase in  transparency  and a  decrease  In turbidity.   A  decrease  in



turbidity during the  August bloom of  U.  pulicaria was  observed in  Lake



Erie in  1984 (Fig.  70).   Turbidity levels  in the  central  and  western



basins have decreased since 1978 (Table 52).  Similarly,  a large  increase



in transparency was attributed to grazing  of D,  pulicaria in Lake Michigan



(Scavia et al. 1986).



    Except for the May bloom,  total phosphorus positively  correlated  well



with phytoplankton abundance spatially on  Lake Erie.   At  least two factors



were controlling  the  phytoplankton abundance.     Because of   the  higher



correlation,   it is  tempting to  suggest that phosphorus   was  the  primary



control on phytoplankton abundance.  This   was not true during the  spring



phytoplankton  bloom  where  zooplankton  obviously  affected   the  bloom.



Although P-control was evident during  the summer,  there  were  also  fairly



high negative correlations between phytoplankton and Daphnia pulicaria and



calanoids.    This exercise    suggests that  "top  down"  and  "bottom   up"



control  of the trophic web  of lake ecosystems exists simultaneously   and



that it varies with season.

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                                                                      102
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                                                                     1Q6


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                                                                      109
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                                                                     no
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                                                                     w
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                                                                     112
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                                                                     113
TABLE  1.  Plankton sampling dates for Lakes Michigan,  Huron and  Erie  in
1984 and 1985.  Only phytoplankton samples were taken during the  winter
helicopter cruises of 1985.
          1984
          1985
Cruise
1
2
3
4
5
6
7
8
9
10
11
Lake
Michigan
4/9-12
5/6-7
-
7/8-9
8/1-3
8/12-14
8/15-16
11/27-29
12/13-18
-
2/7-9
Lake
Huron
4/12-15
5/4-5
-
7/5-7
8/3-4
8/10-12
8/17-18
11/30-12/2
12/10-12
1/15-16
2/9-10
Lake
Erie
4/18-19
4/20-21
5/1-2
7/2-3
8/5-6
8/7-9
8/19-20
12/4-5
12/5-8
1/14-14
2/17-18

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                                                                      114
TABLE
1984.
2.  Latitude and longitude of plankton sampling stations,
Station
Number
LAKE ERIE
LE60
LE57
LE55
LE42
LE73
LE37
LE78
LE79
LE18
LE15
LE09
Latitude

41°53'30"
41 49 54
41 44 18
41 57 54
41 58 40
42 06 36
42 07 00
42 15 00
42 25 18
42 31 00
42 32 18
Longitude

83°11'48"
83 01 06
82 44 00
82 02 30
81 45 25
81 34 30
81 15 00
80 48 00
80 04 48
79 53 36
79 37 00
        LAKE HURON

          LH93
          LH92
          LH91
          LH90
          LH61
          LH57
          LH54
          LH53
          LH48
          LH45
          LH43
          LH38
          LH37
          LH34
          LH32
          LH29
          LH27
          LH15
          LH12
          LH09
          LH06
44 06 00
43 48 30
43 42 00
43 24 00
45 45 00
45 40 00
45 31 00
45 27 00
45 16 42
45 08 12
45 00 48
44 44 24
44 45 42
44 38 24
44 27 12
44 22 00
44 11 54
44 00 00
43 53 24
43 38 00
43 28 00
82 07 00
82 22 00
82 01 00
82 18 00
83 55 00
83 43 36
83 25 00
82 54 54
82 27 06
82 59 00
82 00 30
82 03 36
82 47 00
83 13 54
82 20 30
81 50 00
82 30 12
82 21 00
82 03 24
82 13 00
82 00 00

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                                                                     115
Table 2 (continued).

      LAKE MICHIGAN
        LM05                   42 00 00           87  25  00
        LM06                   42 00 00           87  00  00
        LM10                   42 23 00           87  25  00
        LM11                   42 23 00           87  00  00
        LM17                   42 44 00           87  25  00
        LM18                   42 44 00           87  00  00
        LM22                   43 08 00           87  25  00
        LM23                   43 08 00           87  00  00
        LM26                   43 36 00           87  22  00
        LM27                   43 36 00           86  55  00
        LM32                   44 08 24           87  14  00
        LM34                   44 05 24           86  46  00
        LM40                   44 45 36           86  58  00
        LM41                   44 44 12           86  43  18
        LM46                   45 13 24           86  36  48
        LM47                   45 10 42           86  22  30
        LM56                   45 37 30           86  18  00
        LM57                   45 38 12           86  03  30
        LM64                   45 57 00           85  35  12
        LM77                   45 47 24           84  49  24

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                                                                       116

TABLE  3.  Sample dates and stations for Lake Michigan, 1984 and 1985.


    Station    4/9   5/6   7/8   8/1   8/12  8/15 11/27 12/13  2/7
    Number
5
6
10
11
17
18
22
23
26
27
32
34
40
41
46
47
56
57
64
77a
XXX X
X X X X X
XXX X
X X X X X
XXX X
X X X X X
X XXX
XX X XX
X XXX
XX X XX
X XXX
XX X XX
X XXX
XX X XX
XXX X
X X X X X
XXX X
X X X X X
xxxxxxxx
xxxx xxxx

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                                                                     117
Table 4.  Sample dates and stations for Lake Huron, 1984 and 1985.


Station   4/     5/    7/   8/    8/     8/    ll/   12/     I/    2/
Number  12-15   4-5   5-7  3-4  10-12  17-18  27-2  10-13  15-16  9-10

  6       x            xx            xx
  90             x                x                   xxx

  9x            xx            xx            xx
  91             x                x

  12      x            xx            xx
  92             x                x                   x

  15      x            xx            xx            xx

  27      x            xx            xx
  93             x                x                   x

  29             x                x                   x
  32      x            xx            xx

  34                                                         xx

  37      x            xx            xx            xx
  38             x                x                   x
43
i 45
]
I 48
1
53
1 54
57
61
XX X
X XX XX XX

XX X
XX X
X XX XX X
XX XX
X XX XX X

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                                                                      118
TABLE 5.  Comparison of calculated crustacean dry weights (ug)  to
measured dry weights in Lake Michigan. Measured weights from Hawkins and
Evans (1979).
                                  Calculated
Measured
      Species
Cyclops bicuspidatus
   thomasi

Cyclops vernalis
Diaptomus ashlandi


Diaptomus minutus


Diaptomus oregonensis


Diaptomus sicilis


Limnocalanus macrurus


Tropocyclops prasinus
   mexicanus

Cyclopoid copepodite

Bosmina  longirostris

Chydorus sphaericus

Daphnia  galeata mendotae

Daphnia  retrocurva

Eubosmina coregoni

Holopedium gibberum

Polyphemus pediculus

Epischura lacustris

Eurytemera affinis
mean
weight
4.2
8.6
2.6
2.4
5.1
7.1
29.3
1.3
.7
1.4
1.9
10.0
5.2
2.5
12.0
13.6
9.4
4.9
mean
length
.97
.97
.96
.93
1.26
1.44
2.57
.59
.54
.39
.34
1.32
1.04
.49
.95
.87
1.61
1.25
range
3.0-5.3
5.9-12.2
2.2-3.2
1.9-2.9
3.2-6.7
5.3-10.2
20.5-35.0
0.9-1.8
0.2-1.4
0.6-3.2
1.7-2.1
2.6-20.1
1.6-11.2
0.9-6.1
6.3-20.2
4.3-36.9
5.4-13.0
_
(n)
63
8
65
63
45
61
59
59
65
39
4
42
26
36
1>3
6
22
1
mean
we ight
3.2
1.9
5.6
2.5
4.8
3.9
3.2
2.8
6.9
5.6
17.3
11.4
45.3
33.8
.9
1.2
1.0
1.0
4.0
2.9
1.7
5.0
2.9
10.8
4.8
range
1.9-5.6
1.2-2.9
4,8-6.4
2.4-2.6
2.1-7.1
1.9-5.6
1.7-4.9
1.8-4.4
3.8-10.9
3.3-10.1
13.4-23.6
8.6-11.4
13.2-88.2
16.7-55.8
0.7-1.2
0.6-2.2
0.6-1.8
0.8-1.2
2.5-8.9
1.2-6.5
1.2-2.5
1.9-10.9
2.2-4.1
5.9-13.9
3.9-5.3

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                                                                     119
  Table 6.  Mean values (±S.E.) of physical-chemical parameters
(April-October) from a 1-m depth for Lakes Erie,  Michigan and Huron*  1984.
Values are in mg/L unless noted otherwise. Values in parentheses  represent
number of samples analyzed.  Phytoplankton samples are from April to  February.
Zooplankton samples are from April to November.   The trophic ratio and
zooplankton ratio are discussed in the text.
                               Erie
                     Michigan
                      Huron
pH

Alkalinity

Conductivity (umhos/cm)

Turbidity (NTU)

Soluble Reactive
Silica (mg/L)

Chloride

Sulfate

Nitrite + Nitrate

Total phosphorus (ug/L)

Soluble Reactive
Phosphorus (ug/L)

Sodium

Potassium

Chi a

Phytoplankton
(1000x*/mL)
  (g/m3)

Zooplankton
(1000x*/m )
 (mg/m )

Trophic ratio

Zooplankton ratio
 8.25±.03(101)

 92.5+.43U06)

 272.4+1.54(106)

 4.5±.85(88)


 178.9+21.7(106)

 14.61+.25(105)

 22.70+.20U05)

 0.29±.02(106)

 16.55+1.41(105)


 2.0+.33UOO)

 7.18±.16(32)

 1.40±.02(32)

 3.45+.3K105)
45.1+4.2  (117)
1.0±.08(117)
159.6+25.3(65)
 53.6±6.2(65)

    1.8

    0.35
8.23+.03(83)

106.9+.56(85)

273±1.38(85)

.39+.03(75)


360.1±22(85)

8.79+.08(85)

21.22+.14(84)

0.22+.006(85)

4.63±.24(84)


0.92+.KK71)

4.75±.03(30)

1.30+.OK30)

0.86+.05(84)
22.2+1.4(97)
0.55+.038(97)
59.8±8.3(65)
33.2+4.9(65)

  4

  0.64
 8.02+.03(101)

 77.4±.31(106)

 202.9+.83(106)

 0.32+.02(88)


644.6±19.9(106)

 5.66±.05(105)

 16.09+.1K105)

 0.30+.004(106)

 3.70+.25(105)


 0.80+.10UOO)

 3.17±.05(32)

 0.94±.01(32)

 0.64+.04(105)
17.2+.89(95)
0.38±.10(95)
55.4+7.2(49)
27.3±2.3(49)

 3.8*

 1.50
   * Average of 1983 and 1984

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                                                                    120
TABLE  7.  Number of species and genera observed in each algal
division or grouping in Lake Michigan. 1983 and 1984.  Results are
for the non-winter period.
Division                         Species               Genera
                             1983	1984        1983	1984

Bacillariophyta               168       166          33        29

Chlorophyta                    86        63          36        26

Chrysophyta                    49        33          13        11

Cryptophyta                    23        20           44

Cyanophyta                     21        13          10         8

Picoplankton                   (2)1       3          (2)1       3

Colorless flagellates          16        15           65

Pyrrhophyta                     97           43

Euglenophyta                    11           11

Unidentified                    55           -

Chlorotnanophyta                 10           10
                  Total       379       327         108        91
      Included in Cyanophyta in 1983

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                                                                121
TABLE   8.  Relative abundance of major phytoplankton divisions in
Lake Michigan, 1983 and 1984.  Bac=Bacillariophyta,
Cat=Chloromanophyta, Chl=Chlorophyta, Ohr=Chrysophyta,
Col=Color less flagellates, Cry=Cryptophyta, Cya=Cyanophyta»
Pic=Picoplankton, Eug=Euglenophyta, Pyr=Pyrrhophyta,
Uni=Unidentified.
                           Biovolume/mL            Cells/mL
            Division      1983      1984        1983      1984
Bac
Cat
Chi
Chr
Col
Cry
Cya
Pic
Eug
Pyr
Dni
56.41
0.02
5.25
6.53
0.75
13.43
5.56*
-
0.04
7.32
4.68
69.97
0.00
1.99
5.01
0.41
11.61
1.65
1.39
0.07
2.36
5.53
1.07
0.01
0.65
1.49
0.13
1.24
92.21
-
0.01
0.01
3.20
2.04
0.00
0.67
2.18
0.30
1.50
3.54
82.85
<0.01
0.02
6.89
                   *Picoplankton are included with the Cyanophyta
            in 1983.

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                                                                      122
Table 9.  Abundance of Rhizosolenia eriensis in Lake Michigan in 1983
and 1984. Values in parentheses represent &. eriensis+R. longiseta.
     4/17
     4/26
     5/4
     7/4
     8/3
     8/17
    10/12
    10/26
                  1983
    Date    cells/mL  % biovolume
 0.0
 0.0
 0.2
 0.0
 0.0
 0.0
10.9
 7.1
0.0
0.0
0.1
0.0
0.0(.05)
0.0
9.1(9.2)
2.1(10.7)
                                    1984
                          Date   cells/mL  % biovolume
 4/9
 5/6
 7/8
 8/1
 8/12
 8/15
11/27
12/13
 2/7
10.3
 9.3
52.4
22.6
17,
21,
 3.2
 8.3
 4.8
.5
.9
17.5(30.5)
 8.6(17.4)
33.4(36.1)
23.2(25.0)
26.9(30.1)
39.2(44.6)
 7.9 (8.3)
16.4(17.0)
 4.6 (5.2)

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                                                                                                             123
       TABLE 10.  Summary of common phytoplankton species  occurrence  in  Lake Michigan  daring  1984  and  winter
      of 1985.  Summary is based on all  samples analyzed.  Summary includes the maximum population density
      encountered, the average population   density and  biovolume. and  the relative   abundance  (Z of  total
      cells and Z of total biovolune).   Common species  were arbitrarily  defined  as having an abundance  of
      X3.1X of tbe total cells or >0.5I of tbe total biovolnme.

                 Taxon
BACILLARIOPHVTA
   Asterionella formosa
   Cyclotella comensis v. 1
   Cyclotella comta
   Cyclotella ocellata
   Fragilaria capucina
   Fragilaria crotonensis
   Meloaira islandica
   Helosira italica snbsp. subarctic*
   Nitzschia lauenburgiana
   Khizosolenia eriensis
   Rhizosolenia longiseta
   Stepbanodiscus alpinus
   Stephanodiscus alpinus?
   Stephanodiscua niagarae
   Stepbanodiscus transilvanicus
   Synedra filifoncis
   Synedra ulna v. cbaseana
   Tabellaria flocculoaa

CHLOROPHTTA
   Honoraphidium contortun
   Oocystis submarina
   Dictyosphaerium ehrenbergianum

CHKSOPHYTA
   Cbrysophycean coccoids
   Diaobryon divergens
   Dinobryon sociale v. americanum
   Haptophyte sp.

COLORLESS FLAGELLATES
   Colorless flagellates
   Honosiga ovata

CRYPTOPHITA
   Chroomonas norstedtii
   Cryptomonas erosa
   Cryptomonas marsaonii
   Cryptomonas rostratifornis
   Khodomonas minuta v. nannoplanktica

CYASOPHYTA
   Anacystis montana v. minor
   Coelosphaerium naegelianum
   Oscillatoria limnetica
   Oscillatoria minima

PICOPLAWCTON
   rods
   spheres
   spherical - flagellates

FYKRBOPKfTA
   Gymnodinium sp.
   Peridinium sp.

UNIDENTIFIED
   Unidentified flagellate - ovoid
   Unidentified flagellate - spherical
Maximum
Cells/mL
184
2. 568
96
265
161
376
96
74
10
110
162
18
11
14
7
118
23
82
344
254
278
630
303
1,743
1.456
311
352
270
65
25
25
965
2,790
982
2,070
4.132
4.287
43.541
2.847
16
16
4.287
1.350
Average
Cells/ml
22.4
115.6
4.4
23.3
11.9
74.3
12.6
10.8
0.7
18.2
21.2
2.2
0.7
1.1
0.8
11.2
2.2
13.9
36.8
25.8
23.6
83.1
26.5
111.7
182.3
26.5
24.5
48.8
11.2
3.7
1.3
232.5
292.6
31.9
209.8
175.5
886.6
16.716.3
805.6
0.5
1.5
1.0026.3
503.2
Z of Total
Cells
0.10
0.52
0.02
0.10
0.05
0.33
0.06
0.05
0.00
0.08
0.10
0.01
0.00
0.01
0.00
0.05
0.01
0.06
0.17
0.12
0.11
0.37
0.12
0.50
0.82
0.12
0.11
0.22
0.05
0.02
0.01
1.05
1.32
0.14
0.94
0.79
3.99
75.23
3.63
0.00
0.01
4.62
2.26
Mean
Biovolume
urn /mL
6.130
3.539
11.561
2.079
3.940
48.175
13.538
2.784
4.506
129.063
23.928
8.318
4.267
17.571
16,294
4.225
17.151
41.459
385
417
196
320
5.443
11.052
1.633
424
310
1.480
25.171
4.948
4,572
17.683
1,276
153
1.023
3.737
2.415
4.481
714
4.111
4.275
23.103
6.771
Z of Total
Biovolume
1.12
0.65
2.12
0.38
0.72
8.83
2.48
0.51
0.83
23.64
4.38
1.52
0.78
3.22
2.99
0.77
3.14
7.60
0.07
0.08
0.03
0.06
1.00
2.02
0.30
0.08
0.06
0.27
4.61
0.91
0.84
3.24
0.23
0.03
0.19
0.68
0.44
0.82
0.13
0.75
0.78
4.23
1.24

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                                                                    124
TABLE 11.  Common species observed in either 1983 or 1984»  but  not
both years, Lake Michigan.  Common species were arbitrarily defined
as having an abundance of >0.1% of the total cells or >0.5% of  the
total biovolume.
            1983
      1984/1985
Bacillariophyta

     Cyclotella michiganiana
     Cymatopleura solea
     Entomoneis ornata
     Fragilaria vaucheriae
     Tabellaria fenestrata
Cyclotella ocellata
Rhizosolenia longiseta
Nitzschia lauenburgiana
Synedra filiformis
Synedra ulna v. chaseana
Chlorophyta
     Cosmarium sp.
     Stichococcus sp,
Oocystis submarina
Dictyosphaerium ehrenbergianum
Chrysophyta
     Dinobryon cylindricum
     Stylotheca aurea
Cryptophyta
     Cryptomonas erosa v. reflexa
     Cryptomonas pyrenoidifera
Cryptomonas rostratiformis
Cyanophyta
     Gomphosphaeria naegelianum
     Oscillatoria agardhii
Oscillatoria minima
Pyrrhophyta

     Ceratium hirundinella

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                                                                      125
Table  12.  Number of species in Lake Michigan with depth at Station 47,
15 August 1986.

                                    Division
Depth
(m)
1
5
10
15
20
Bac
13
11
30
23
27
Chi
1
3
4
5
10
Chr
9
9
7
7
8
Cry
5
4
5
3
6
Cya
1
2
2
3
5
Pic
3
3
3
3
3

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                                                                      125
  TABLE 13.  Comparison of abundance of Cyclotella species  at  offshore
sites in August of 1970, 1983 and 1984, Lake Michigan.   Data from
Holland and Beeton (1972), Makarewicz (1987) and this  study.  Stations
22 and 27 are geographically comparable to Holland and  Beeton's  offshore
sites.  Values are in cells/mL.
                     11 August 70        17 August 83     15 August 84
                  (offshore stations)  (Stations 22&27)   (Stations22&27)
Cyclotella
  michiganiana        71 - 182            0.44 - 6.8        0.38 - 4.5

Cyclotella
  stelligera         300 - 613            0.17 - 2.2        1.7 - 2.8

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                                                                      127
TABLE 14.  Comparison of nutrient levels between Stations 6, 64, 77 and
all other stations during the spring and fall, Lake Michigan. Mean+j>.E.


Station 77
Station 64
Lake Mean
( exc lud ing
Station 77)

Silica
(ug/L)
632.7+23.2
364.5+22.8
501.0+14.4
Total
Phosphorus
(ug/L)
4.67+1.08
6.35+2.16
5. 14+. 35
Nitrate +
Nitrite
(mg/L)
.27+.01
.20+. 01
.26+.01
      Station 6
502.8+38.9
4.05+.41
.27±.01

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                                                                      128
  TABLE  15.  Distribution of indicator diatom species in Lake Michigan.
The classification scheme followed Tarapchak and Stoermer (1976).
M,=mesotrophic but intolerant of nutrient enrichment•  M =mesotrophic and
tolerant of moderate nutrient enrichment. E=eutrophic.  1970-71, 1977
and 1983 data are from  Holland and Beeton (1972),  Stoermer and Tuchman
(1979) and Makarewicz (1987).
Ml
6
5
6
5
M2
5
3
2
3
E
7
1
2
2
M1+M2'
1.6
8.0
4.0
4.0
      19772(Nearshore)

      1970-713

      19831

      19841
            Only diatoms contributing >.5% of the biomass for a cruise
      are classified.

         2  Only diatoms contributing >1% (1977) or >0.1% (1984) of the
      abundance are classified.
         3
            Only "predominant" species are classified.

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                                                                      129
  Table 16.  Relative abundance of zooplankton in Lake Michigan.
                             Percent                     Percent
                                                        Abundance
Rot ifera

Cladocera

Copepoda nauplii

Cyclopoida

Calanoida

Mysidacea

Harpacticoida
                         1983       1984            1983         1984
N
0
T










C
A
L
C
U
L
A
T
E
D

2

39

11

15

30

0
<
.6

.8

.2

.8

.4

.2
.1
59

3

21

5

10

<
<
.7

.2

.3

.7

.1

.1
.1
67.5

4.1

15.6

6.2

6.6

<.01
<.01

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                                                                      130
  TABLE  17.  Summary of common zooplankton species occurrence in Lake
Michigan during 1984.  Values are from the short zooplankton hauls only.
Species were arbitrarily classified as common if they accounted for
>0.1% of the total abundance or 1.0% of the total biomass,  with the
exception of rotifers.  Rotifer species were considered common if they
accounted for >1.0% of the total abundance.
Taxon
COPEPODA
 Copepoda - nauplii
 Cyclopoida
   Cyclopoid - copepodite
   Cyclops bicuspidatus
      thomasi
   Tropocyclops prasinus
      mexicanus
 Calanoida
   Diaptomus - copepodite
   Diaptomus ashlandi
   Diaptomus minutus
   Diaptomus sicilis
   Limnocalanus macrurus
CLADOCERA
   Bosmina longirostris
   Daphnia galeata mendotae
   Daphnia pulicaria
   Daphnia retrocurva
   Eubosmina coregoni
   Holopedium gibberum
   Leptodora kindtii
ROTIFERA
   Collotheca sp.
   Conochilus unicornis
   Gastropus stylifer
   Kellicottia longispina
   Keratella cochlearis
   Notholca foliacea
   Notholca laurentiae
   Notholca squamula
   Polyarthra remata
   Polyarthra vulgaris
   Synchaeta sp.
Maximum
Dens ity
#/m3
62127
14358
5475
439
30508
5098
695
1062
469

29566
9110
690
5286
1465
4333
255

6814
8850
18843
43489
124128
21396
52609
50381
20550
47790
27545


Average
Density
#/m3
9183
2767
749
60
2518
848
132
2157
56
TOTAL
942
846
78
238
125
136
27
TOTAL
1134
942
1241
5649
11764
798
2325
2200
1105
5785
4223
TOTAL

% of
Total
Density
15.60
4.70
1.27
.10
4.28
1.44
.22
.37
.09
22.07
1.60
1.44
.13
.40
.21
.23
.05
4.06
1.93
1.60
2.11
9.60
19.99
1.36
3.95
3.74
1.88
9.83
7.18
63.15
95.29
Mean
Biomass
ug/m
3673
1797
3057
73
3676
2162
321
1478
1637

876
6825
1638
1389
271
1132
779

8
17
18
49
65
20
77
37
30
82
98


% of
Total
Biomass
11.23
5.50
9.35
.22
11.24
6.61
.98
4.52
5.01
54.67
2.68
20.88
5.01
4.25
.83
3.46
2.38
39.49
.02
.05
.05
.15
.20
.06
.24
.11
.09
.25
.30
1.53
95.69

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                                                                      131
  TABLE  18.  Cladoceran abundance in 1954, 1966, 1968, 1983 and 1984 in
Lake Michigan.  Data from Well^ (1970), Makarewicz (1987) and this
study. Values are in number /m
Species and Year
Leptodora kindtii
1954
1966
1968
1983
1984
Daphnia galeata
1954
1966
1968
1983
1984
Daphnia retrocurva
1954
1966
1968
1983
1984
Diaphanosoma brachyurum
1954
1966
1968
1983
1984
Daphnia longiremis
1954
1966
1968
1983
1984
Daphnia pulicaria
1954
1966
1968
1983
1984
Holopedium gibberum
1954
1966
1968
1983
1984
Polyphemus pediculus
1954
1966
1968
Early
August
29
4
16
34
98
1200
0
0.4
514
3508
1400
79
2100
82
1061
2
0
0
1
0
0
16
0
0
14
0
0
0
1011
248
0
2
5
456
536
2
15
10

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                                                          132
TABLE
18. (continued)
1983
1984
Bosmina longirostris
1954
1966
1968
1983
1984
Eubosmina coregoni
1954
1966
1968
1983
1984
Ceriodaphnia quadrangula
1954
1966
1968
1983
1984

13
7

26
98
16
342
5231(141)*

0
1
16
159
208

0
4
1
0
0
* Bloom at Station 77 and 64. Mean for the offshore waters
minus Station 77 and 64 is in parentheses.

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                                                                    133
  TA.BLE  19.  Copepod abundance in 1954, 1966, 1968, 1983 and
1984 in Lake Michigan.  Data frgm Wells (1970), Makarewicz (1987) and
this study. Values are number/m.

                                                    Early
                     Species and Year               August

                     Limnocalarms macrurus
                         1954                          91
                         1966                          34
                         1968                         270
                         1983                          18
                         1984                          64
                     Epischura lacustris
                         1954                          41
                         1966                           7
                         1968                          21
                         1983                          19
                         1984                          14
                     Diaptomus sicilis
                         1954                           3
                         1966                           1
                         1968                           3
                         1983                          79
                         1984                         155
                     Mesocyclops edax
                         1954                         200
                         1966                           0
                         1968                           0
                         1983                          13
                         1984                          31
                     Senecella calanoides
                         1954                         0.2
                         1966                         0.2
                         1968                         0.1
                         1983                         1.4
                         1984                           0
                     Cyclops bicuspidatus
                         1954                         310
                         1966                        1000
                         1968                         860
                         1983                        1457
                         1984                        2807
                     Diaptomus ashlandi
                         1954                         140
                         1966                         220
                         1968                          13
                         1983                        1256
                         1984                        1733
                     Cyclops vernalis
                         1954                           0
                         1966                           0
                         1968                           0
                         1983                           0

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                                                          134
TABLE  19.  (continued).
            1984                          16
        Eurytemora affinis
            1954                           0
            1966                          33
            1968                           3
            1983                           0
            1984                           0
        Diaptomus oregonensis
            1954                          63
            1966                          58
            1968                         100
            1983                         138
            1984                          58
        Diaptomus tninutus
            1954                          39
            1966                          25
            1968                        1500
            1983                         151
            1984                         183

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                                                                     135
Table 20.  Average crustacean zooplankton biomass (dry weight)  for 1976
and 1984, Lake Michigan.  The 1976 data (Bartone and Schelske 1982) were
converted to dry weight assuming carbon content was 50% of dry  weight.
                       1976           50.0+14.8 mg/m3
                       1984           33.6+14.7 mg/m3

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TABLE  21.  The ratio of calanoids to cyclopoids  plus  cladocerans
geographically in Lake Michigan,  1983 and 1984.
                                                                     136
                                              Calanolda
at ion

77 (North)
64
57
47
41
34
27
23
18
11
6 (South)
Cyclopoida + (
1983
0.37
0.41
1.74
1.52
1.10
1.03
1.53
1.15
3.01
1.71
0.87
Jladoce:
1984
0.23
0.20
0.69
0.57
0.57
0.80
0.84
1.32
1.93
1.09
0.75

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                                                                      137
Table 22.  Correlation of phytoplankton with total  phosphorus
concentrations and zooplankton abundance within individual  cruises  (  11
stations) in Lake Michigan,  1984.  NO = not observed.

4/9-12
5/6-7
8/1-3
8/15-16
11/27-29
12/18
Daphnia
pulicaria
NO
-.132
-.021
-.272
-.171
-.095
Daphnia
spp.
.794
-.327
.137
-.496
-.016
.594
Rot if era
.395
.715
.768
-.031
.680
.763
Calanoida
-.707
-.738
-.059
.243
.455
-.164
Tota
Phosphi
-.385
-.113
.330
.191
-.156
.653

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                                                                      138
TABLE 23.  Number of species and genera observed in each algal  division
or groupingt Lake Huron, 1983 and 1984.
      Division         	Species          	Genera

BAG
CHL
CHR
CRY
CYA
PIC
COL
PYR
BUG
UNI
CAT
1983
158
73
36
22
13
(2)*
13
10
4
3
1
1984
156
64
35
17
13
3
13
9
1
4
0
1983
29
28
10
3
6
(2)*
4
4
3
-
1
1984
28
28
12
4
7
3
5
4
1
-
0
              Total     329     315           88        92
          *  Included in Cyanophyta in 1983

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                                                                      139
TABLE 24.  Relative abundance of major phytoplankton divisions in Lake
Huron, 1983 and 1984.  In 1983 picoplankton are included with the
Cyanophyta.  BAC=Bacillariophyta, CAT=Chloromanophyta» CHL=Chlorophyta,
CHR=Chrysophyta, COL=Colorless Flagellates, CRY=Cryptophyta,
CYA=Cyanophyta, PIC=Picoplankton, EUG=Euglenophyta, PYR=Pyrrhophyta»
UNI=Unidenti£ied.
   Division             Biovo ]uiPft/UlL           	Cells/mL

BAG
CAT
CHL
CHR
COL
CRY
CYA
PIC
EUG
PYR
UNI
1983
68.20
.02
3.45
7.11
.14
8.29
4.31*
-
.11
3.25
5.11
1984
61.90
0.00
2.72
9.45
.19
9.10
1.41
1.60
.06
7.15
6.41
1983
1.16
.01
.42
1.60
.06
1.13
89.53*
-
.01
.01
6.09
1984
2.78
0.00
.58
2.08
.14
1.24
4.15
83.85
.01
.02
5.14
              * Picoplankton included in Cyanophyta in 1983.

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                                                                     14.0
Table 25. Abundance of Rhizosolenia eriensis in Lake Huront 1983 and
1984. Values in parentheses in 1983 represent Rhizosolenia sp. and in
1984 R.. longiseta.
                 1983
                                    1984
     Date  cells/mL  % biovolume
     4/21
     5/6
     7/2
     8/4
     8/19

    10/16
    10/24
0.1
0.2
0.0
0.0
0.0

0.4
0.0
0.01
0.01(38.3)
0.0 (59.2)
0.0 (11.3)
0.0 (12.8)
                        Date  cells/mL  % biovolume
1.0
0.0
(6.1)
(8.7)
 4/12
 5/4
 7/5
 8/3
 8/10
 8/17
11/27
12/10
 1/15
 2/9
 6.3
 5.4
51.0
26.7
33.1
 9.9
 5.8
 2.9
 2.4
10.7
 9.0(0.43)
 6.3(0.46)
18.1(0.81)
30.4(0.92)
   1(0.15)
   1(0.51)
   1(0.39)
10.3(0.44)
 4.4(0.0)
12.4(0.17)
35
29
16

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                                                                                                 141


 TABLE  26,  Summary of common phytoplankton species  occurrence  in Lake  Huron during  1984  and winter of  1965*
Summary is based on all samples analyzed.   Summary includes  the  maximum  population density encountered*
 the average population density and biovolume,  and the relative  abundance  (!  of  total cells and  I of total
biovolume). Common species were arbitrarily defined as having  an abundance of >0.1Z of the total cells or
>0.5Z of the total biovolume.
                                                                                       Mean
      Taxon                                       Maximum     Average    X  of  Total    Biovolume  Z  of Total
                                                 cells/ml   cells/ml/         Cells      um /mL     Biovolume
BACIIXARIOPHYTA
    Asterionella formosa                              168        27.5
    Cyclotella comencis                              1386       122.2
    Cyclotella comta                                   35         2.3
    Cyclotella kuetzingiana v. planetophora?          135        13.2
    Cyclotella ocellata                              1000       113.2
    Cyclotella stelligera                             267        25.3
    Fragilaria crotonensis                            375        44.7
    Fragilaria intermedia v. fallax                    25         2.6
    Melosira islandica                                 43         6.5
    Rhizosolenia eriensis                             131        17.2
    Rhizosolenia longiseta                             33         2.9
    Stephanodiscus alpinus                             19         1.5
    Stephanodiscus minutus                             85        19.4
    Stephanodiscus niagarae                             2         0.2
    Tabellaria flocculosa                             181        25.0
.16
.71
.01
.08
.66
.15
.26
.02
.04
.10
.02
.01
.11
.00
,15
9.225
5.781
8.178
3.902
9.784
614
39,333
2.233
8.752
81.644
2.355
3.950
851
3.562
69.337
2.13
1.49
1.89
.90
2.26
.14
9.09
.52
2.02
18.87
.54
.91
.20
.82
16.02
CHL080PHYTA
    Cosmarium sp.
                                                       16
                                                                   .7
                                                                               .00
                                                                                          2,173
                                                                                                         .5
CHRYSOPHYTA
    Chrysopbycean coccoids
    Chrysosphaerella longispina
    Dinobryon cylindricum
    Dinobryon divergens
    Dinobryon sociale
    Dinobryon sociale v. americanum
    Haptophyte sp.
CRYPTOPHYTA
    Chroomonas norstedtii
    Cryptomonas erosa
    Cryptomonas pyrenoidifera
    Cryptomonas rostratiformis
    Rhodomonas minute v. nannoplanktica
CtANOPHYTA
    Anacystis montana v. minor
    Coelosphaerium naegelianum
    Gomphosphaeria lacustris
    Oscillatoria limnetica
    Oscillatoria minima
PICOPLANKTON
    rods
    spheres
    spherical - flagellates
PYRRHOPHYTA
    Ceratium hirundinella
    Gymnodinium helveticum f. achroum
    Gymnodinium sp.
    Gymnodinium sp.  2
UNIDENTIFIED
    Unidentified flagellate - ovoid
    Unidentified flagellate - spherical
160
1325
196
254
589
540
589
115
31
33
8
360
4606
1047
851
942
335
2741
29690
2160
36.2
31.3
13.3
32.0
65.6
27.8
110.1
22.8
4.5
4.2
.8
155.1
445.4
77.6
79.0
45.9
17.3
811.6
13021.0
563.2
                                                     1481
                                                     2193
                                                                   .1
                                                                   .2
                                                                   .5
                                                                   .3
615.9
264.7
.21
.18
.08
.19
.38
.16
.64
.13
.03
.02
.00
.90
2.59
.45
.46
.27
.10
4.73
75.84
3.28
.00
.00
.00
.00
3.59
1.54
189
8.313
4,298
6.544
10,771
4,716
1.460
724
10,333
2.450
3.290
13,772
2,205
335
380
219
453
2,568
3,768
592
14,991
3,566
3,312
5.816
17.740
9.767
                                                                                                        .04
                                                                                                       1.92
                                                                                                        .99
                                                                                                       1.51
                                                                                                       2.49
                                                                                                       1.09
                                                                                                        .34
                                                                                                        .17
                                                                                                       2.39
                                                                                                        .57
                                                                                                        .76
                                                                                                       3.18
                                                                                                        .51
                                                                                                        .08
                                                                                                        .09
                                                                                                        .05
                                                                                                        .10
                                                                                                       .59
                                                                                                       .87
                                                                                                       .14
                                       3.46
                                        .82
                                        .77
                                       1.34
4.10
2.26

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                                                                       142
TABLE 27.  Common species observed in either 1983 or 1984 but  not  in
both years. Lake Huron.
             1983
Bacillariophyta

   Stephanodiscus transiIvanicus
     1984/85
Cyclotella stelligera
Stephanodiscus alpinus
Stephanodiscus minutus
Chlorophyta
Cryptophyta
                                         Cosmarium sp.
                                         Cryptomonas rostratiformis
Cyanophyta

   Anacystis thermalis
   Coccochloris elabans
Oscillatoria minima

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                                                                     143
   Table 28.  Distribution of indicator diatom species in Lake Huron.
The classification scheme of Tarapchak and Stoermer (1976) was utilized,
M^meso trophic but intolerant of nutrient enrichment,  M =me so trophic and
tolerant of moderate nutrient enrichment, E=eutrophic.  1971, 1975-76
and 1983 data are from Munawar and Munawar (1979),  Lin and Schelske
(1978) and Makarewicz (1987).
Ml
6
2
7
6
M2
3
4
2
3
E
3
2
2
3
M1+M2
3.0
3.0
4.5
3.0
      19711

      1975-762

      19833

      19843
            Only diatoms cantributing >5% of the seasonal biomass are
      classified.
         2
            Only "abundant" diatom species are classified.
         3
            Only diatoms contributing >0.5% of the biomass for the study
      period are classified.

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                                                                 144
  Table 29.  Relative abundance of  zooplankton in  Lake Huron.
Rotifera

Cladocera

Copepoda nauplii

Cyclopoida

Calanoida

Amphipoda

Mysidacea
                             Percent                     Percent
                         	Biomass               	Abundance

                         1983       1984           1983         1984
N
0
T












C
A
L
C
U
L
A
T
E
D

2

27

14

13

42

<

0
.5

.5

.7

.3

.0

.1

.0
41

4

23

11

19

0

<
.1

.8

.1

.2

.8

.0

.1
56

2

18

7

15

<

0
.0

.9

.6

.3

.3

.1

.0

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                                                                      145
  TABLE 30.  Summary of common zooplankton species occurrence in Lake
Huron during 1984.  Values are from the short zooplankton hauls  only.
Species were arbitrarily classified as common if they accounted  for
>0.1% of the total abundance or 1.0% of the total biomass, with  the
exception of rotifers.  Rotifer species were considered common if they
accounted for >1.0% of the total abundance.
Taxon
Maximum  Average    % of     Mean     % of
Density  Density   Total   Biomass   Total
           ///m    Density   ug/m    Biomass
COPEPODA
 Copepoda - nauplii
 Cyclopoida
   Cyclopoid - copepodite
   Cyclops bicuspidatus
      thomasi
   Mesocyclops
      copepodite
   Mesocyclops edax
 Calanoida
   Diaptomus - copepodite
   Diaptomus ashlandi
   Diaptomus minutus
   Diaptomus oregonensis
   Diaptomus sicilis
   Limnocalanus macrurus
CLADOCERA
   Bosmina longirostris
   Daphnia galaeta
      mendotae
   Daphnia pulicaria
   Eubosmina coregoni
   Holopedium gibberum
   Leptodora kindtii
ROTIFERA
   Collotheca sp.
   Conochilus unicornis
   Gastropus stylifer
   Kellicottia longispina
   Keratella cochlearis
   Notholca squamula
   Polyarthra remata
   Polyarthra vulgaris
   Synchaeta sp.
24749
12791
1487
3262
270
22584
2960
1306
256
2044
266

3304
4127
935
3441
2124
133

3584
66009
9855
19274
51995
6804
5916
18086
12963


10071
3254
316
300
40
6174
1071
369
93
502
20
TOTAL
338
586
71
326
158
16
TOTAL
672
10878
1094
3784
6652
570
650
2917
1489
TOTAL

18.59
6.01
.58
.55
.07
11.40
1.98
.68
.17
.93
.04
41.00
.62
1.08
.13
.60
.29
.03
2.76
1.24
20.08
2.02
6.99
12.28
1.05
1.20
5.38
2.75
52.99
96.75
4028
1750
1356
205
283
5020
2189
720
363
2377
525

303
3136
1017
709
1658
416

4
239
26
45
24
11
17
117
42


14.73
6.40
4.96
.75
1.03
18.36
8.01
2.63
1.33
8.69
1.92
68.81
1.11
11.47
3.72
2.59
6.06
1.52
26.48
.01
.87
.09
.16
.09
.04
.06
.43
.16
1.92
97.21

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                                                                     146
  Table 31.  Comparison of mean crustacean abundance for the sampling
period in 1971 (April-November), 1974/75 (April-November),  1983
(August-October) and 1984 (April-December), Lake Huron.  1971 data
modified from Watson and Carpenter (1974), 1974/75 data from McNaught et
al. (1980) and J983 data from Makarewicz (1987).  NF = not  found. Values
are in number/m
j
Cladocera
  Bosmina longirostris
  Eubosmina coregoni
  Daphnia retrocurva
  Daphnia galeata mendotae
  Daphnia longiremis
  Daphnia pulicaria
  Chydorus sphaericus
  Holopedium gibberum

Cyclopoida
  Cyclops bicuspidatus
       thomasi
  Cyclops vernalis
  Tropocyclops prasinus
     mexicanus
  Mesocyclops edax

Calanoida
  Diaptomus ashlandi
  Diaptomus minutus
  Diaptomus sicilis
  Diaptomus oregonensis
  Limnocalanus macrurus
                                1971
              553 (1047)*
              330 (765)*

              339 (852)*

                0 (0)
               18
              229 (580)*
             3764 (3274)*
              7.5 (5)*

               63 (61)*
                5 (6.7)*
              246 (37)*
              462 (322)*
              117 (77)*
              109 (92)*
               64 (44)*
                            1974/75**    1983***
4109
2084
 361
 692

   0
 391
 576
1271
 117

 310
  91
 745
 966
 496
 192
  34
 518
 229
  74
1029

 363
  NF
  58
2346
  .5

 577
 115
 206
 465
 145
 140
 9.3
                     1984
 338
 326
  36
 586

  71
  NF
 158
 316
 1.5

  21
  40
1071
 369
 502
  93
  20
       *   August, September and October average
       **  Includes Saginaw Bay
       *** August and October average

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                                                                      147
  Table 32.  Abundance of selected zooplankton species  in  northern  and
 mthern Lake Huron in 1984.   Valu<
is defined as south of Station 27.
southern Lake Huron in 1984.   Values  are number/m .   Southern Lake  Huron
                 Conochilus     Kellicottia    Diaptomus     Ho lopedium
                 unicornis     longispina     minutus       gibberum
  Northern        12,526          3,897          298            239

  Southern         4,729          2,449          383             29

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                                                               148
TABLE 33. Ratio of Calanoida to Cladocera plus  Cyclopoida  in Lake
Huron, 1983 and 1984.

                       	Calanoida	
Station

   61   (North)

   54

   45

   37

   32

   27

   15

   12

   09

   06   (South)
                       Cyclopoida + Cladocera
1983

0.67

1.11

1.19

1.57

2.13

1.37

1.60

1.98

1.31

1.23
1984

0.90

1.36

1.84

1.33

1.46

1.16



1.83

2.00

1.89
Mean

0.74

1.24

1.52

1.45

1.80

1.27



1.91

1.66

1.56

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                                                                     149
Table 34.   Comparison of the plankton ratio
(Calanoida/Cyclopoida+Cladocera)  between the  northern  stations of Lake
Huron and Lake Michigan.
                                1983    1984     mean
          Lake Michigan
            Station 77          0.37    0.23     0.32

          Lake Huron
            Station 61          0.67    0.90     0.78
            Lake Mean           1.49    1.61     1.55

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                                                                     150
  TABLE 35.  Mean abundance of rotifers in Lake Huron in 1974 and  1983.   Data
from Stemberger et al. (1979), Makarewicz (1987) and this study.   NF  = not
found in short tow.
Colletheca sp.

Conochilus unicornis

Filinia longiseta

Gastropus stylifer

Kellicottia longispina

Keratella cochlearis

Keratella earlinae

Notholca squamula

Polyarthra dolichoptera

Polyarthra remata

Polyarthra vulgaris

Synchaeta kitina

Synchaeta stylata

Synchaeta sp.
1974
April-Nov.
#/L
0.8
15.0
3.4
5.2
6.8
41.9
10.9
7.4
3.0
6.8
17.6
8.1
7.1
2.4
1980
April-July
#/L
0.0
0.79
<.01
0.27
1.15
1.86
<.01
1.8
0.12
0.12
0.05
NF
NF
1.03
1983
Aug. -Oct.
#/L
0.90
7.10
0.004
1.10
2.10
2.00
0.08
NF
0.07
0.01
3.00
NF
NF
0.10
1984
April-Dec.
0.67
10.87
0.007
1.09
3.78
6.65
0.10
0.57
0.43
0.65
2.92
NF
NF
1.5

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                                                                     151
  Table 36.  Correlation (r) of phytoplankton abundance with total
phosphorus concentrations and zooplankton abundance within individual
cruises (10 stations) in Lake Huron»  1984.  NO = observed.

5/4-5
8/3-4
8/17-18
11/30-12/2
12/10-13
Daphnia
pu lie aria
-.110
-.258
-.286
-.218
.380
Daphnia
spp.
-.110
-.698
-.060
-.460
.415
Rotifera
.393
.595
-.662
.420
.049
Calanoida
-.370
.010
-.549
.101
-.192
Total
Phospho:
-.032
.144
-.314
-.168
.378

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                                                                      152
Table  37.  Number of species and genera observed in each algal division
or grouping, Lake Erie, 1983 and 1984.  Bac=Bacillariophyta»
Cat=Chloromanophyta» Chl=Chlorophyta, Chr=Chrysophyta»  Col=Colorless
flagellates, Cry=Cryptophyta» Cya=Cyanophyta, Pic=Picoplankton,
Eug=Euglenophyta, Pyr=Pyrrhophyta, Uni=Unidentified.
                      	Species	          	Genera
    Division
    BAG

    CHL

    CHR

    CRY

    CYA

    PIC

    COL

    PYR

    BUG

    UNI

    CAT
1983
176
108
29
14
16
-
15
8
2
3
1
1984
171
96
28
15
18
3*
11
9
0
4
1
1983
30
38
11
3
9
-
6
4
2
0
0
1984
30
38
14
4
10
0
4
4
0
0
0
               TOTAL    372        356          103       104
             * Included in Cyanophyta in 1983.

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                                                                      153
  TABLE  38.  Number of species identified and percentage of species
belonging to various taxonomic groups* Lake Erie.   1970 data represent
the mean for the central, western and eastern basins [modified from
Munawar and Munawar (1976)].
Number of Species
Division
BAG
CHL
CHR
CYA
CRY
EUG
PYR
PIC
UNI
COL
1970
134.3

16.3
58.0
6.3
11.2
3.3
0.7
4.0
-
-
_
1983
372
Percent Composition
47.3
29.0
7.8
4.3
3.8
0.5
2.2
-
0.8
4.0
1984
356

48.0
27.0
7.9
5.1
4.2
0.0
0.0
0.8
1.1
3.1

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                                                                     15.4
Table  39.  Phytoplankton and zooplankton biomass*  total phosphorus and
chlorophyll a. concentrations in the western, central and eastern basins
of Lake Erie, 1983 and 1984. Values are in g/m  unless noted otherwise.
estern
1.49
1.38
1.44
Central
1.59
0.76
1.18
Eastern
0.84
0.54
0.69
Entire
Lake
(mean±S.E.
1.36+.12
1.00+.16
1.18
    Phytoplankton

         1983

         1984

         mean

    Zooplankton

         1984         0.055      0.052      0.054    0.053+.0062

         1984 (#/L)   295.6      94.3       130.4    159.6±25

    Total
     Phosphorus

         1983(ug/L)   26.77      16.82      12.79

         1984(ug/L)   23.91      19.37      12.41

    Chlorophyll a

         1983(ug/L)    5.68       4.05       2.22

         1984(ug/L)    5.10       3.27       2.11

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                                                                                               155
 TABLE  40.  Summary of common phytop lankton species  occurrence  in Lake Erie during  1984 and winter of 1985.
Summary is based on all samples analyzed.  Summary includes  the  maximum population density encountered,
 the average population density and biovolume,  and the  relative  abundance  (% of  total cells and I of total
biovolume). Common species were arbitrarily defined as  having  an abundance of  >0.1X  of the total cells or
>0.5S! of the total biovolume.
             Taxon
BACILLARIOPHY1A
      Asterionella formosa
      Fragilaria capucina
      Fragilaria crotonensis
      Melosira islandica
      Stephanodiscus alpinus
      Stephanodiecus binderanus
      Stephanodiscus niagarse
      Stephanodiscus sp.
      labellaria flocculosa

 CHLOROPHYTA
      Cosmarium sp.
      Crucigenia rectangularis
      Oocystis borgei
      Fedlastrun simplex v. duodenarium

CHRYSOPHYTA
      Haptophyte sp.

COLORLESS FLAGELLATES
      Colorless flagellates
      Stelexmonas dichotoma

CRYPTOPHYTA
      Chrooraonas norstedtii
      Cryptomonae eroea
      Cryptomonas rostratiformie
      Rhodomonas minuta v* nannoplanktica

CYANOPHYTA
      Anabaena sp.
      Anacystis montana v. minor
      Aphanizomenon flos-aquae
      Coelosphaerium naegelianum
      Herismopedia tenuissima
      Oscillatoria limnetica

PICOPLANKTON
      rods
      spheres
      spherical - flagellates

PYRRHOPHYTA
      Ceratium  hirundinella
      Gymnodinium sp.  2
      Peridinium aciculiferum
      Peridinium sp.

UHIDEHTIFIED
      Unidentified flagellate - ovoid
      Unidentified flagellate - spherical
Maximum
Cells/mL
942
407
826
1564
198
2506
120
781
207
25
295
180
393
1317
2119
1186
425
295
33
2348
1162
22253
2643
3436
6218
5179
10987
379.888
1726
82
33
41
82
4303
2479
Average X of Total
Cells/mL Cells
73.4
38.2
77.9
31.5
8.7
59.2
4.9
78.3
15.8
1.0
5.1
8.0
7.8
151.9
65.2
87.8
50.9
24.3
1.6
499.1
47.8
1052.1
103.6
78.1
85.6
112.7
1.128.5
38,075.3
544.8
2.8
2.2
1.3
5.5
1177.5
558.1
0.16
0.08
0.17
0.07
0.02
0.13
0.01
0.17
0.04
0.00
0.01
0.02
0.02
0.34
0.14
0.19
0.11
0.05
0.00
1.11
0.11
2.33
0.23
0.17
0.19
0.25
2.50
84.46
1.21
0.01
0.00
0.00
0.01
2.61
1.24
Mean Z of Total
Bio volume Biovolume
urn /mL
48,802
10.764
66.983
35,812
17,522
21.539
135.855
6,991
33.732
39.142
10.087
9.357
10.685
2,670
1.619
3.500
1.219
45.760
5,132
39.038
7,603
4.892
7.598
333
103
421
3.154
10.207
644
37.283
31,523
12.634
25.308
51.864
16.609
5.57
1.23
7.53
4.09
2.00
2.46
15.51
0.80
3.85
4.47
1.15
1.07
1.22
0.30
0.18
0.40
0.14
5.23
0.59
4.46
0.87
0.56
0.87
0.04
0.01
0.05
0.36
1.J7
0.07
4.26
3.60
1.44
2.89
5.92
1.90

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                                                                    156
  Table  41.  Location of maximum abundance of selected species  in 1983
and 1984, Lake Erie.
                                            1983
Fragilaria crotonensis
Fragilaria capucina
Melosira granulata
Melosira islandica
Stephanodiscus sp.
Stephanodiscus binderanus
Tabellaria flocculosa
Oscillatoria tenuis
Oscillatoria limnetica
Oscillatoria subbrevis
Anacystis montana var. minor
Aphanizomenon flos-aquae
spheres
Cryptomonas erosa
Chroomonas norstedtii
Merismopedia tenuissima
Pediastrum simplex var. duodenarium
Coelosphaerium naegelianum
Scenedesmus ecornis
Peridinium aciculiferum
Stephanodiscus
Asterionella formosa
Gymnodinium sp.#2
Haptophyte
Western
Western
Western
not common
not common
Western
Western
Western
Western
Western
not common
not common
Western
Western
not common
not common
Central
Central
Central
Central
Central
not common
                    1984
Western
Western
not common
Western
Western
Western
Western
not common
Western
not common
Western
Western
Western
Western
Western
Western
Western
Western
Western
Central
Western
Central
Central
Central

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                                                                      157
  TABLE  42.  Common species observed in either 1983 or 1984 but  not
both years. Lake Erie.  1983 data are from Makarewicz (1987).
            1983
Bacillariophyta

   Actinocyclus normanii f. subsalsa
   Melosira granulata
   Rhizosolenia sp.
           1984
Asterionella formosa
Melosira islandica
Stephanodiscus sp.
Chlorophyta

   Coelastrum microporum
   Monoraphidium contortum
   Mougeotia sp.
   Scenedesmus ecornis
   Staurastrum paradoxum
Crucigenia rectangularis
Cyanophyta

   Agemenellum quadruplicatum
   Oscillatoria subbrevis
   Oscillatoria tenuis
Anacystis sp.

-------
                                                                      158
  TABLE 43.  Importance of Asterionella formosa during
the spring of 1984,  Lake Erie.   Sampling dates:  4/18, 4/20,  5/1/84.
   Rank             Species



    1        Asterionella formosa

    2        Fragilaria crotonensis

    3        Melosira islandica

    4        Gymnodinium sp. #2
B iovolume(g An )
  All Species
   0.162

   0.160

   0.123

   0.109
   Rank             Species


    1        Stephanodiscus sp.

    2        Asterionella formosa

    3        Fragilaria crotonensis

    4        Stephanodiscus parvus
Abundance (#/mL)
  Diatoms

   238

   224

   170

   117

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                                                                      159

  TABLE  44.  Mean maximum bio mass  of selected  common phytoplankton species  in
1970 and 1983, Lake Erie.   Data from Munawar and Munawar  (1976)  and this
study.  1970 data - graphical accuracy.   Percent reduction  is  from 1970 to
1984.

Actinocyclus
normanii
Stephanodiscus
niagarae
Stephanodiscus
tenuis
Stephanodiscus
binderanus
Fragilaria
crotonensis
Fragilaria
capucina
Peridinium
aciculiferum
Ceratium
hirundinella
Rhodomonas
minuta
Cryptomonas
erosa
Pediastrum
simplex
Staurastrum
paradoxum
Aphanizomenon
BASIN
Western
Eastern
Central
Western
Western
Western
Eastern
Central
Western
Central
Eastern
Central
Eastern
Central
Eastern
Eastern
Central
Western
Central
Central
Western
197§
g/m
4.7
1.4
2.3
0.6
1.8
0.5
1.0
3.4
7.9
2.4
0.4
0.2
1.0
1.8
2.0
1.6
0.4
2.0
0.4
0.4
2.0
1983*
g/m
0.30
1.05
2.19
0.12
0.001
0.11
0.15
0.11
0.18
0.02
0.04
0.06
0.05
0.35
0.31
0.04
0.10
0.63
0.06
0.07
0.10
1984.
g/m
0.05
0.22
0.23
0.17
0.002
0.04
0.45
0.16
0.29
0.03
0.01
0.18
0.03
0.13
0.35
0.05
0.14
0.40
0.00
0.00
0.09
Percent
Reduction
99
84
90
72
99
92
54
95
96
99
99
10
95
93
83
97
65
37
100
100
96
   flos-aquae

-------
                                                                      150
  Table  45.  Distribution of indicator diatom species in the western
basin of Lake Erie.  The classification scheme of Tarapchak and Stoermer
(1976) was utilized.  Only diatoms contributing 5% or more of the
biomass for a cruise are classified.  M. = mesotrophic but intolerant of
nutrient enrichment* M_ = mesotrophic and tolerant of moderate nutrient
enrichment, E = eutropnic.  1970 data are from Munawar and Munawar
(1976).  1978 data are from Devault and Rockwell (1986).
                                   M2       E
           1970           0        1        5        0.2

           1978           0        3        3        1.0

           1983           1        2        3        1.0

           1984           3        2        2        2.5

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                                                                      161
  Table  46.  Trophic status of the western, central and eastern basins
of Lake Erie in 1970 and 1983/84.  The classification scheme of Munawar
and Munawar (1982) is used.  1970 data is from Munawar and Munawar
(1982). Based on average biomass of basins in 1983 and 1984.
                                     1970             1983 + 84


Eastern Basin                    mesoeutrophic       oligotrophic

Central Basin                    mesoeutrophic       mesotrophic

Western Basin                    eutrophic           mesotrophic

-------
                                                                    162
  Table 47.  Relative abundance of zooplankton  in Lake Erie.
Rotifera

Cladocera

Copepoda nauplii

Cyclopoida

Calanoida

Harpacticoida

Amphipoda
                          Percent                         Percent
                     	Bjomass	              	Abundance

                      1983       1984              1983         1984
N
0
T










C
A
L
C
U
L
A
T
E
D

13

40

12

17

16

<
<
.6

.5

.3

.1

.5

.1
.1
69

6

15

5

3

<
0
.2

.0

.8

.4

.7

.1
.0
80

3

10

3

2

<
<
.1

.2

.4

.9

.5

.1
.1

-------
                                                                    163
  TABLE 48.  Summary of common zooplankton species occurrence in Lake
Erie during 1984.  Values are from the short zooplankton hauls only.
Species were arbitrarily classified as common if they accounted for
>0.1% of the total abundance or 1.0% of the total biomass,  with the
exception of rotifers.  Rotifer species were considered common if they
accounted for >1.0% of the total abundance.
Taxon
COPEPODA
 Copepoda - nauplii
 Cyclopoida
   Cyclopoid - copepodite
   Cyclops bicuspidatus
      thomasi
   Mesocyclops - copepodite
   Mesocyclops edax
   Tropocyclops prasinus
      mexicanus
 Calanoida
   Diaptomus - copepodite
   Diaptomus oregonensis
CLADOCERA
   Bosmina longirostris
   Chydorus sphaericus
   Daphnia galaeta mendotae
   Daphnia pulicaria
   Daphnia retrocurva
   Eubosmina coregoni
   Holopedium gibberum
   Leptodora kindtii
ROTIFERA
   Ascomorpha ovalis
   Asplanchna priodonta
   Brachionus sp.
   Conochilus unicornis
   Keratella cochlearis
   Keratella crassa
   Keratella earlinae
   Notholca foliacea
   Notholca laurentiae
   Notholca squamula
   Polyarthra dolichoptera
   Polyarthra major
   Polyarthra remata
   Polyarthra vulgaris
   Synchaeta sp.
Maximum  Average    % of
Density  Density   Total
  #/m      #/m    Density
  Mean    % of
Biomass   Total
 ug/m   Biomass
79012
13367
4519
6311
3095
1407
20178
7731

4772
6675
21410
3752
6903
11215
807
623

57498
52038
157414
57762
40170
37236
42931
56316
93031
348455
61171
102788
18399
340262
340262


16275
3625
790
954
413
234
2652
890
TOTAL
710
157
1932
492
287
1209
63
35
TOTAL
6159
1806
3418
3404
7726
1575
1831
2825
5125
17392
4430
7768
2537
35357
14864
TOTAL

10.35
2.31
.50
.61
.26
.15
1.69
.57
16.44
.45
.10
1.23
.31
.18
.77
.04
.02
3.11
3.92
1.15
2.17
2.17
4.91
1.00
1.16
1.80
3.26
11.06
2.82
4.94
1.61
22.49
9.46
73.93
93.48
6510
2614
3637
758
1608
255
4249
3631

832
126
7506
7784
982
2417
754
627

77
1582
203
37
29
77
65
74
363
347
208
711
44
1597
1115


12.34
4.95
6.89
1.44
3.05
.48
8.05
6.88
44.08
1.58
.24
14.22
14.75
1.86
4.58
1.43
1.19
39.85
.15
3.00
.38
.07
.06
.15
.12
.14
.69
.66
.39
1.35
.08
3.03
2.11
12.37
96.30

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                                                                      164
  TABLE  49.  Occurrence of eutrophic zoqplankton indicator species in
Lake Erie. 1984.  Values are in number/tn .
                                              BASIN
                                   Western    Central    Eastern

Brachionus angularis                 177         0         0

B. budapestinen*                      92         0         0

B. calyciflorus                       97         0         0

B. caudatus                           81         0         0

Filinia longiseta                    459        2.8        0

Keratella cochlearis f. tecta       2062        9.2        0

Trichocerca cylindrica               397         0         0

T. elongata*                         907         0         0

T. multicrinis                       477        42         0

T. pusilla                            36         0         0

    *Not listed as eutrophic species by Gannon and  Stemberger (1978).

-------
                                                                     165
TABLE 50.  Ratio of calanoids to cladocerans  plus  cyclopoids  in Lake
Erie, 1983 and 1984.
                   WESTERN     CENTRAL    EASTERN    MEAN
                    BASIN       BASIN      BASIN

         1983        0.19        0.31       0.45     0.32

         1984        0.27        0.42       0.36     0.35

-------
                                                                        166
Table 51.    Correlation (r)   of phytoplankton abundance  with  total  phosphorus
concentration and zooplankton abundance within individual cruises  (11   stations)
in Lake Erie, 1984.  N.0.=not observed.

4/18-19
5/1-2
8/5-6
8/19-20
12/4-5
Daphnia
pulicaria
N.O.
N.O.
-.509
-.548
N.O.
Daphnia
spp.
.535
-.941
-.079
.061
-.448
Rot if era
.714
-.771
.021
.929
.097
Calanoida
.343
-.922
-.534
-.383
-.345
Tota
Phospho
.801
-.811
.756
.910
.505

-------
                                                            167
Table 52.   Turbidity levels in 1978 and 1984,
Lake Erie. 1978 values represent graphical
accuracy.


                      1978             1984


                    mean±S.E.        mean±S.E.

     Western        4.2+1.5          2.66±.43

     Central        0.7              0.40+.04

     Eastern        0.5              0.52+.09

-------
                                          168
          Station Locations
    Lake Michigan - Main Lake
               Manistique
                         Traverse City
                  34   A   Michigan
                      Ludington
                                  Petosky
 Milwaukee
       <
    Racine
 Waukegon[
  Chicago
  Muskegon
Benton Harbor
FIGURE 1. Lake Michigan plankton sampling stations, 1984-85

-------
                       Lake Huron
                       Main Lake
                       Sampling Locations
                                                    CTv
FIGURE 2.  Lake Huron plankton sampling stations, 1984-85. >

-------
                  Lake Erie
       Main Lake Sampling Station
                                             Lake Ontario
 A
 N

Michigan
                                       Erie   New York
                                  Pennsylvania
                     Cleveland
            Ohio
United States
     FIGURE 3. Lake Erie plankton sampling stations, 1984-85.

-------
                                                          2T
o
a
o
LU
u
40-




30-




20-
     10--
                   LAKE  MICHIGAN
                            o  Total
Total  minus

 picoplankton
         AMJJASONOJF
E
X


E
                                                          0--
                                                                       LAKE MICHIGAN
                                                                                                B
                                                                                —-er*"-	o
                                                         AMJJASONDJF
                      FIGURE 4.  Seasonal phytoplankton abundance (Aa) and biovolume (4b)

                      trends in  Lake Michigan* 1984-8^.

-------
                                                         172
.8-
01
5 .Bn
r— 1
0
o
•rl
CD
4-
C« *T
O
••H
i.
o
a.
o p-
c • *
Q.
0-
\J
_^_ LAKE MICHIGAN
"^"A __^*
\r~~~~~~~
•— — • BAC »
* 	 « CRY
B— —c PIC

0 	 e CYA
0 	 — 0 UNI
« 	 « COL
•— — • CHR

B— — a PYR
•—- -— » CHL x
. 	 —-^-^^ ~-~~"~~~~~
p;;^:."^i^]^=;^^fe=s=s=s==:..s5Jg|
	 1 	 1 t 	 1 	 i — 	 	 1 	 1 	 1 i
M
                     J
0
N     D
FIGURE 5.  Seasonal distribution of algal divisions in Lake Michigan,
Bac=Bacillariophyta, Chl=Chlorophyta, Chr=Chrysophyta» Col=colorless
flagellates, Cry=Cryptophyta, Cya=Cyanophyta,  Pic=picoplankton,
Pyr=Pyrrhophyta, Uni=unidentified flagellates, 1984-85.

-------
                   NUMBER/ML
NUMBER/ML
                   NUMBER/ML
NUMBER/ML
 14
 O
 CO
 O
O

H-
00
03
                                                                 o

-------
                   NUMBER/ML
                                    NUMBER/ML
O t-i
n o
  a
CO 50
  PI
  ?
  05
  05
  n>
* s
  *
O O
p n-
ito H
O H-
3" o-
  C
T> rt
-t I—
— O
  o
D Mi

1 »
•0 •— '
D>
3

i b
n
D


3 cr1
Si v-x
NUMBER/ML
NUMBER/ML
O IV) *•
t * * '

X

t-t

«-.


3^


0
2
D


(_i
•n





^v.
^v.
J*
m
*

t
\
\
jt
/
I
f
j
cn oo o IV)
i i i i
X

n
X
TJ
0
3
O
3
O
~fD
i
o
Q
^
O
T
3
n
a

-------
 •*)

 I
 90
 M
NUMBER/ML
 oo
 to
 3

 
-------
    Cyclotella ocellata
N -•»  S
                                       Dictyosphaerium
                                       N
       Qocystis submarina
N
I
         B
Oscillator!a  minima
                                        N -
          FIGURE  9.  Seasonal and  geographical distribution of a) Cyclotella
          ocellata. b) Oocystis SMbTMJJLat  °) Dictyosphaerium ehrenbergiamim and d)
                      . ^inj^na , Lake Michigan

-------
                       CELLS XI000 / ML

                                    40
                                     11
                                                         177
60
                                       +  Total

                                       o  Non-picoplanKton

                                       x  Picoplanhton


                                           Temperature
                    19   20   21   22   23   24   25


                         TEMPERATURE   <°C  )
                        CELLS    XI000 / ML

                   0123
           a.
              10-
              15-
              20 A
                            CELLS / ML
                                      120     160
                                   comensl* - all vor.
                             o  A.  formoBa
                             x  F.  crotoneneiB
FIGURE 10.   Vertical  distribution of phytoplankton at Station A7.  15  August
1984, Lake Michigan

-------
    X
    Q.
    a.
    LU
    o
        10-
       20- c
       30 -L
        10-
    X   20- i
        30-
        40-
                                                178
                  CELLS XI000 / ML
                     20
         40
         —H-
                            60
                                  o  non -
                                      picoplankton

                                  x  Total  cells
                  CELLS XI000 / ML
                                              2
                                             •H
       o  Baci1lariophyta
       x  Chrysophyta
       *  Chlorophyta
       ^  Cryptophyta
       D  Cyanophyta
       •  Temperature
                                            B
21
22
                               23
24
25
                    TEMPERATURE   C C )
FIGURE tl.  Vertical distribution of phytoplankton at Station 18. 15 August
1984 , Lake Michigan

-------
                                                              179
o
o
o
r—I
X

CO
u
LJ
u
    40-r
    30-
    20-
    10-
      0
    1.6
*   1.2
\

o
o
o
1—I

x    .8--
     . 4--
       1   \
                        LAKE    MICHIGAN



                               o   Total

                               y.   Picoplankton

                               •+•   Non-pi cop lankton
            _,	1	>	 i
                    NORTH
A
                                               -i	,	,_
                                              SOUTH
                     -i	t    i
                                 A
                                      I
CYA

CHL


BAG


CRY

CHR
                                        \
            •i  •   t
           77  64  57  47  41  34   27  23   18  11  06


                             STATION
     FIGURE 12. Annual geographical distribution of major algal divisions

     in Lake Michigan.  Bac=Bacillariophyta. Chl=Chlorophytai

     Chr=Chrysophyta, Cry=Cryptophyta,  Cya=Cyanophyta» Pic=picoplankton,

     Pytspyrrhophyta, 1984-85.

-------
                                                                  180
Z


N


O
D
O


X


(/>



LU
U
      _j
      o
      o
      o

      X

      1/1
      111
      (J
         60-r
         40--
         20--
         40
         30--
    20--
         10--
         40"
^  30--
o
o
a

x  20-•
S  10
                LAKE MICHIGAN
                                                    H
                                       April 0-12

                                       May 6-7

                                       Ma*. 27-29

                                       tec. 13 - IB

                                           r
              NORTH
                                       I
                                           ""«
                                        SOUTH
                             o  Abundance

                             x  Temperature
                                                  --4
                                                  --3
                                                  --2
                                                 if)

                                                 •-*
                                                 10
                                                 _j
                                                 LU
                                                 U
              77 64 57 47  41 34 27 23 18 11 06


                          STATION
FIGURE 13.  Geographical distribution of phytoplankton abundance on

all cruises,  Lake Michigan,  1984-85.

-------
       10-r
o
a
o
^-t
x

tn
6--
4--
LU
u
        0
              1960    1965    1970    1975     1980    1985
           FIGURE 14.  Historical abundance of phytoplankton in Lake Michigan.

           Horizontal bars are the mean.  Wide vertical lines are the standard error.

           Thin vertical  lines are the range.  Data are from Stoermer and Kopczynska

           (1967a and b). Rockwell et al. (1980), Makarewicz (1987) and this study.
                                                                                       CO

-------
                                                               182

0>
E
X
a
o
o
en
D


n
O
O
0
•—I
X
111
u
z
Q
z
ID
m

lOO-i
80-



60-
40-
20-
175:
140-

105-


70-

35-

0-
i_nr\c. muniuniN
/\
1 \

1 \
j \
1 ®L
// X
/ X.s^


- A
t \
/ \
/ \
/ \
/ \
/ \
/ \
°^ ^0

                AMJJASONDJF
FIGURE  15.  Seasonal zooplankton abundance in Lake Michigan, 1984,

-------
                                                               183




m
Z
0
O
o
\BUNDANCE XI






25-
20-
15-

10-
5-
120:
100-
80-
60-

40-

20-
0-
i r \ i \ i_ ii * ^^ i • * •«• • • • ••
L A o Calonoldo
. v x Clodocara
' \ * Copapoda
/ 4i a Cyelapotda
// \
/ A ^\
r's/'\Z^

' . Rot If or a
/ \
/ \
/ \
/ ^
/ \
*-** \
	 < 	 . 	 . 	 , 	 , 	 . — : — . 	 . 	 . 	 . 	 .
                     MJJASONDJ
FIGURE  16.  Seasonal fluctuation (numerical)  of  zooplankton groups  in Lake
Michigan, 1984.  Copepoda refers  to the  nauplius  stage  of  the  Copepoda.

-------
                                                             184






PI
2
o
o
o
X
07
3




20-
16-

12-

8-

4-
60:
45-
30-

15-
0-
I_/\r\L_ n x u*i i x uf \m
e Copopoda
f\ x Cyclopolda
/ \ * Rotlfwra
/ \
/ \
/ /^^^^
fs^ "^ ^
'&-*< ^ ^^*\


/ . o Colanoida
/ * x Cladocara
- / V
/ Q \
/ /\ \
/ s' V- ^
&-*f V_
•»-? 	 	 , , , ,
              AMJJASONDJF
FIGURE  17.  Seasonal fluctuation (biomass)  of  zooplankton groups  in Lake
Michigan.  1984.  Copepoda refers  to  the nauplius  stage  of the Copepoda.

-------
                                                     185
     16--
      12--

-------
                                                   186 ,
                      LAKE   MICHIGAN






tn
^C*
2
O
o
o
f- 1
X
en







50-
40-

30-
20-


10-


30:


24-
18-

12-
6-

VM. r * i « i,^. 1 1 ab ^^ 1 1 A ^^ f * 1 V
Total
/r~ -K A\
^^v X / v/A
V \
\
\
fe





o Colanoida
x C 1 adocero
* Copepoda
O Cyclopoido . \ O
O Rotifsro A ' \ /
/ ' \
^~^^ \ / V \
/^v-^r'"^ VN
- ^-/ ^X^"^ ^^ ^
N^ ^ \fZ- •*— >= *= ^ »
$ — -v — -_^_ ._ o _. ^^___ 	 	 y
it i i i , ... i 	 + 	 i t i i I
           77   64  57  47  41   34  27  23   18  11  06
                             STATION
FIGURE  19.  Geographical distribution (biomass) of major  zooplankton
groups in Lake Michigan, 1984.  Copepoda = Copepoda nauplius.

-------
              LAKE   MICHIGAN


CE
LU
L_
l
1— 1
_J
a:
Ld
CD
2:
Z)
~z.




61

5-


4-

3-
2-
1-

0-
H»
o Diaptomus
sici 1 if
x Diaptomus -



" A
. ^ / V
. ^w
0
o 	 e — •"
77 64 57 47

K
5 / \
copcpodite / \
/ \
/ i
/ \
i \
/ \
f \
X / \
v \
X
^^ 	 _ —CXa^fVi—. .. _ JJI^Up"^^** " ^^^ T»y^^*^ _ ^_*^
	 Q — --"^r '^•"•'J i***^— *^J
l 1 1 1 1 1 1 1
41 34 27 23 18 11 06
STATION
FIGURE 20. Geographical distribution of Diaptomus sicilis in Lake
Michigan, 1984.
                                                        CO

-------


o:
UJ
HH
_J
\
(T
UJ
m
Z)


6-1

5-

4-

3-


2-

1-
o-
§•
^ LAKE MICHIGAN
\
'F o Eubosmina corQgoni
\ x Bosmina longirostris
\ + HolopQdium gibborum
\ a Conochilus unicornis
&-!._„
' \ ^\
' S \ \
\ \
- \ \ \ __
^=--*=:4t=:^-a^84^=:S^:*=:S^.^
    77 64  57  47 41 34  27  23 18  11  06


                      STATION
FIGURE  21. Geographical distribution of selected zooplankton in Lake

Michigan, 1984.
oo
oo

-------
              LAKE   MICHIGAN

o:
UJ
i—
i— *
_j
\
NUMBER ,

141
12-
10-
8-
6-
4-
2-
0-
r o Notholca foliacea
x N. laurentiaQ
R * N. squamula
\ f\ O Polyarthra vulgar is
/ \ Q P. romata
' / \\\ ^*-- 	
/ A \^ \ »
rt \ \ ^\ ^DtL \X' L/' /'
^> — — -\or ^*^\**&~^ ^6^ -£* -
&—- -s^^'!*^3*^'5^^^^^-^
*j~— --^^- 	 Q 	 O — Q 	 =BI — :--€3
	 . 	 1 	 • 	 • 	 1 	 1 	 1 	 1 	 1 	 i i 	 A
    77 64  57  47  41  34 27 23 18 11  06

                     STATION
FIGURE 22.  Geographical distributin of selected zooplankton in Lake
Michigan, 1984.
oo

-------


1
— 1
O")
D
to
0
E
O
•fH
GQ

1DU-
120-

90-

60-


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••
H
/ \
/ \
7 \
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x 40m Stati
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\
/ \
. \
\ f *^^^^
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-4 V A
; \
on / \
on / \
© Lakawtda offshore avaragQ / Daphnia
& — e — ®— --e---'s*-^Q^ spp.
I 	 1 	 1 	 1 	 i i 	 —4 	 1 	 1 	 1 	 « 	 1
  1975
1980
1985
Figure 23.   Historical trends  in zooplankton biomass during July and
Augusts Lake Michigan. The 1984 data point represents the mean of all
offshore stations. Modified from Scavia  et al. (1986).
                                                      VD
                                                      O

-------
                   LAKE HURON
E


X
t!
        AMJJASONDJF
                                                          40 T
30--
-I

\
o
o
o
                                                     x   20--
                                                     ui
                                                     u
10--
               LAKE  HURON
                                         B
                                                                                 o   Total
                           Total  minus

                            pi cop lank ton
    AMJJASONDJF
                           FIGURE  24.   Seasonal phytoplankton biovolume  (4a) and abundance (4b)

                           trends  in Lake Huron, 1984-1985.

-------
           Proportion Biovoluma
M o 35 ij
*O M C M
00 "•< H Q
*• II O C ->,.
1 0 3 » •>*
• t3 fO
ll9" ^
ft OS (&
co n co
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^s CD Co
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000.

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n sr o.
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\ X /
\ /
\ /
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k / ^
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\ / /
1 / /
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X \ ^
40 x «


R-
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-------
                                                           193
         10  -
                61   5*4  45  37  32  27  15
                                STATION
FIGURE 26.   Annual  geographical distribution of major algal division in
Lake Huron.   Bac=Bacillariophyta. Chl=ChlorophytaB  Chr=Chrysophyta,
Cry=Cryptophyta»  Cya=Cyanophyta, Pyr=Pyrrhophyta,  1983.

-------
o
V)
u
\

o
o
o
«—I
X

(ft
LU
CJ
     30-r
     20--
      10-
     1.2-
                                                          194
                           LAKE  HURON
                     NORTH

              \
\,
A
/
                          \
                                           o TOTAL
                                           x PIC
                                    ->   SOUTH
                                            -»-
                                             BAC

                                             CRY

                                             CHL

                                             CYA


                                             CHR
                     \	/
                                                /
             61  54   45  37  32   27   12  09   06

                            STATION
FIGURE 27.  Annual geographical distribution of major algal divisions in

Lake Huron.  Bac=Bacillariophyta.  Chl=Chlorophyta, Chr=Chrysophyta,

Cry=Cryptophyta, Cya=Cyanophyta, Pic=picoplankton, Pyr=Pyrrhophyta.

1984-85.

-------
                                                   195
/u-


_l
z- 50-
o
o
o
x 30-
Ul
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10-
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* Aug 3-4
* Aug 10 - 12
• Aug 17 - 16
D Nov 27 - Dec 2
B
«^^ ^^V^ *"*""' - * 	
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1 1 1 1 1 1 1 1 1 1
o Apr 12 - 15
x Hoy 4-5
* Dec 10 - 13 NUKIH " bUU 1 H
« Jan 15 - 16
a Fab Q - 10
A >
/ \ /^i ^«
/*^\J*^'^ /S*^*^/
	 t t 	 1 	 : 	 1 	 1| 1 	 < 1 1 	 1
         61  54   45  37  32   27   12  09   06
                        STATION
FIGURE 28.  Geographical distribution of phytoplankton abundance on all
cruises. Lake Huron, 1984-85.

-------
                        NUMBER/ML
                                       NUMBER/ML
so *n
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  M



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-------
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-------
                                                        198
                         CELLS  XI000 /  ML
           I

           a.
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              25
            o  Total


         >   x  Picoplankton


            •*•  non-pi cop 1 ankt on
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              20




              25
                             CELLS/ML
                            40
                            —i—
                             80
                             —i—
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              x   C.  kuotzingiana v.
                                 pianetophora
  120
                               i

                               10
                              15
 i

20
                                             °C
FIGURE  31.  Vertical distribution of phytoplankton at Station 37, 15

August 1984, Lake Huron
 i

25

-------
                                                                    199
        CELLS XI000 /  ML

        10       20      30
                                                               40
                      ~  10-
                      fc  20
                      D
                          30-•
                               \

                                                   \
                                          CELLS / ML


                                      200      400      BOD
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 f

H.H
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                               j\

                          30-•  *  V
                                    400    BOO    1200    1600
                       I

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                          10--
                          SO--
                          SO-
                                     \   o  Chryoophytq

                                     /   x  Dinebryon oociol*

                                         *  0. dlvwganc
FIGURE   32.  Vertical distribution of phytoplankton at  Station 15,  15

August  1984* Lake Huron

-------
m
  LD
           2-r
       1.  5-
0-
19
•
70

1974

1978

1982

1986
         FIGURE  33.  Historical  offshore  biomass trends in Lake Huron.  Values are
         the mean±S.E. and the range.   Data are from Munawar and Munawar (1979),
         Makarewicz (1987) and this  study.  1980 offshore data are modified from a
         GLNPO data base.
                                                                                           ro
                                                                                           o
                                                                                           o

-------
                                                               201

o
X
O
o
D
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PI
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150-
125-
100-
75-

50-
25-
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LAKE HURON A
A

' ®v
/ ^x
f \^
/ \
/ \
/ N*
' d
B
/ \
/ \
/ ^
/ x.
- /
d ^&**
~. 	 III! 	 < « 	 1 	 1 1 1 	
                  AMJJASONDJF
FIGURE 34.  Seasonal zooplankton biomass  (a)  and  abundance  (b)  in Lake
Huron, 1984.

-------
                                                             202




CD

X
O
o
o
><

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L)

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z
m






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16-
12-
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40

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0
LAKE HUKUN A
Q o Calonolda
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B

*
/ \
/ • Rot if era
/ \
/ ^x

/ X\
/ X
' , ,,4 	 1 	 ..II 	 • 	 ' ' _ «
MJ
                                    SONDJF
FIGURE 35.  Seasonal fluctuation (numerical)  of  zooplankton groups  in Lake
Huron, 1984.  Copepoda  refer  to  the  nauplius  stage of the Copepoda.

-------
                                                             203






m
X
| xlQQQ
O7




30 -]

24-

18-
12-
6-
»:
12-

9-
6-
3-
0-

LAKE HURON A

o Caionolda
H C 1 OQOC QT*Q
\
y^^V-^^
• -/ \
/ w
B
o Copapoda
x Cyclopolda
* Rot 1 fora
/ \
G— - 	 ^f_ *
7^ 	 Q KV.

              AMJJASONDJF
FIGURE 36 .   Seasonal fluctuation (biomass)  of  zooplankton groups  in Lake
Huron     1984.  Copepoda refers  to  the  nauplius  stage of the Copepoda.

-------
i -
Q£ p
LU ' °
• 	
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Daphnia pulicaria ' \
/ \
- o 1984 / \
x 1983 / x
1
1
l
I
NORTH ^* l SOUTH
^ N/ A
/ \

61 54 45 37 32 27 12 09 06
STATION
FIGURE 37 . Geographical distribution of Daphnia pulicaria in 1983 and
                                                                                             ro
                                                                                             o
1984, Lake Huron.

-------
                                                   205



pi
o
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-------
                                                      206



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. i 	 1 	 1 	 1 i i 	 1 	 1 	 *
             61  54  45  37  32  27  12  09   06

                             STATION
FIGURE 39.  Geographical distribution (biomass) of major zooplankton
groups in Lake Huron,  1984.  Copepoda refers to the nauplius stage of the
Copepoda.

-------
                                                        207
  10
  .£
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                         LAKE  HURON
                                   CALANOIDA •-
            NORTH
                           CLADOCERA »-
CYC LOP DID A, •-

  NAUPLII
          I   J   i   i   I   J   I   I
                                      TOTAL •-
                                     SOUTH
             I    i    i    r   r   i    i    ITT
            61  54 45 37 32  27  15   12  9   6
                           STATION
FIGURE 40.   Geographical distribution of major zooplankton groups in
Lake Huron,  1983.

-------
                                                  208


CK
LU
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          61   54   45   37  32  27  12  09  06
                          STATION
FIGURE 41 .  Geographical distribution of selected Rotifera (a) and
Copepoda  (b) in Lake Huron, 1984.

-------
                                                               209
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     61 54 45 37 33 27 12  09  06
                                        61 54 45 37 32 27 12  09  06
Figure 42.   Water  chemistry  along the north-south axis of Lake Huron,
1984. Values are the  station  mean ± S.E.

-------
cc
LU
Of
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GQ
20--
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                     LAKE     HURON
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                          1975
1980
1985
        Figure 43.  Crustacean abundance (excluding nauplii) of Lake Huron,
                  1970-1984. 1970:Watson and Carpenter (1974). 1974:McNaught
                  (1980), 1983:Makarewicz (1987). 1974 data represent
                  offshore sites only (Areas 9 and 10). Values are the mean ± S.E.
                                                                     ro
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-------




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1 i
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           M    J     J
S    0    N     D    J
Figure 44.  Abundance of Rotifera in Lake  Huron  in  1974,  1983 and 1984.  1974
data are from offshore stations  only (Stemberger et a]. 1979).
                                                                                           ro

-------
                                                       212
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 20
                           LAKE  ERIE
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                           x  Total  minus

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               AMJJASONDJF
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               AMJJASONDJF
FIGURE 45.  Seasonal phytoplankton abundance (a)  and biovolume (b) trends

in Lake Erie, 1984-85.

-------
                      PERCENT  BIOVOLUME
O W Ml
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 11  ii <=3
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to  n ..
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                                                     1

-------
                                                          214
90-
70-
50-
30-
N.
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0
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\j r~ c T 	 	 	 	 , 	 - 	 r~ A c T
WLo 1 tAo 1
              60 57 55  42 73  37 78  79 18  15 09

                           STATIONS

FIGURE 47.   Annual geographical distribution  of major algal divisions in
Lake Erie.   Bac=Bacillariophyta,  Chl=Chlorophyta» Chr=Chrysophyta»
Cry=Cryptophyta» Cya=Cyanophyta,  Pic=Picoplankton, 1984-85.

-------
        400
*  300

O
O


5  20°
     UJ
     u
            ..    \
                         e April 18-18

                         x April 20-21
1- 2

2- 3
                                  LAKE  ERIE
              60 57 55  42 73 37 78 79  18 15 09
         75T
                            o  Aug.  5-6


                            K  Aug.  7-9
                                                g
1 55-
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\
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&=*• 4fZ

mf
* Aug. 19-20
j/£—^f**\^
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         75T
    55--
     o
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     tn
      UJ
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         60 57  55 42 73 37  78 79 IB  15  09

                            e  0«c.  4-5     C


                            K  D«c.  5-9
                    A
                       \
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                                 »  Jon.  13-14  19BS
                         V
                                  V

                                               N
              60 57 55 42 73 37 78 79 18  15 09

                           STATION
                                                   215
FIGURE 48'.   Geographical distribution of phytoplankton abundance on all

cruises, Lake Erie,  1984-85.

-------
                                                                    216
300 T
                °  A»t«rlon«lla formoBa
                x  Frog 11 or la crotoncnsii
                +  M«lo«lra ialandica
                     A
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   tn
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                      duodanar i urn
              »  Chroomonas normtadti i
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" 50-

0-
600-1
V^ ;
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t i^^^ «^*«^^ •^•••••B ^U«^B «^^^_ ^M>^^ ^M««m .^ri_^ -"
** ™ -» — ^^p~^— ™^r^^~ ^w^^ ^W*«"» "^F^^ •
-------
                                                                  217
3500-
2800-
X2100-
if)
j 1400-
ai
u
700-
0-
80 n
' *
^ Anocyctic Montana v. Minor
\
\
\
\
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A
     ^   60 +


     o
     a

     2   40 +
     x

     to

     d   20 +
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                \
                 \
    \
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        900--
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     _
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          0--
o RhodoMonas •loota v. narmoplanktica

x Oscillatoria limnatlca

* Marismopadia tanuiealaa
               60 57 55 42  73 37  78 79 18  15  09
FIGURE 50.  Geographical distribution of selected species,  Lake Erie,

1984-85.

-------
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                                                                    219
                 1983
             f-01970
                 >I984
to
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                M
A   '   S      0

    MONTHS
N
       FIGURE 52.   Seasonal fluctuation of weighted mean phytoplankton biomass in
       1970, 1983  and  1984, Lake Erie.   1970 data modified from Munawar and
       Munawar (1976).  1983 data from  Makarewicz (1987).  Values are corrected by
       using the weighting factors of 15.6%, 59.6% and 24.6%  for the western,
       central and eastern basins (after Munawar and Munawar  1976) .

-------
                                                               220
i<;-
9-
m
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LAKE ERIE




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^^**" * *""*J








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-







-
— i
1955 1965 1975 1985
FIGURE 53.  Regression (r =79.2%)  of phytoplankton biomass  versus  time in
western Lake Erie.  Modifed from Gladish and Munawar (1980).   1956-58  data
are from the Bass Island region.  1970 data from Point  Pelee  and near  the
mouth of Detroit River.  1975-76 data are from northern portions of  the
western basin.  1978 data are from similar geographic areas as 1970
(Devault and Rockwell 1986). 1979 data are not included because of a
reduced sampling regime and other technical difficulties (Devault  and
Rockwell 1986). 1983-84 data are from Stations 60, 57 and 55.  Except  for
the 1956 and the 1957-58 data sets, all enumeration was by  the Utermohl
technique.  In 1956 and 1957-58, a settling technique was used, but  counts
were not made on an inverted microscope. Thin vertical  lines are the range.
Wide vertical lines are the  standard error.

-------
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    12-
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CL
o
L
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            LAKE    ERIE
        1970
                       o   V/QstQrn
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                       •*•   Eastern
                                     \
                                 ^

                                      v
i   i   <  <   i   (   <   i   t  t   (   i—i  i >•
            1975
1980
1985
      Figure 54.  Time trend in annual cruise mean concentration of

      corrected chlorophyll a since 1970. Data from Rathke (1984) and this

      study.
                                                           ro
                                                           ro

-------
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-------
                                                                223


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250-

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100-


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r LAKE ERIE
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e

	 1 	 1 	 1 i < 	 1 	 1 . • • i
                   AMJJASONDJF
FIGURE  56.  Seasonal zooplankton abundance  and biomass in Lake Erie,
1984.  Short hauls are plotted.

-------
                                                          224



^
1)
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8-

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240-
200-
160-
120-
80-
40-
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LAKE ERIE ° J*1""01*'
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/ ^v. ' \
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/ \
- / \
./
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- 	 ^ 	 V

                AMJJASONDJ

                              MONTHS
FIGURE  57.   Seasonal abundance distribution of zooplankton groups in Lake
Erie* 1984.   Copepoda refer to the nauplius stage of  the  Copepoda.

-------
                                                             225



q
2
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20-

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LAKE LKit
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Col one Ida
Cladocara
Copepodo
Cyclopolda
Rot If era


\^
*
                 AMJJASONDJF
FIGURE  58.  Seasonal biomass  distribution of  zooplankton groups in Lake
Eriet 1984.  Copepoda refer  to the  nauplius  stage of the Copepoda.

-------
                                                         226
O
O
O
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                      LAKE    ERIE
                          o  Total

                          x  Rot i fora

                             Total  crustacQa
o  Co1anaIda

x  Clodocera

*  Copapoda

   Cyclopolda
            60 57 55 42  73 37  78 79 18  15  09


                          STATION
 FIGURE  59.  Geographical distribution (numerical) of zooplankton groups in

 Lake Erie, 1984.  Copepoda refer to the nauplius  stage of the Copepoda.

-------
                                                  227




ro
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: 	 . 	 . 	 • 	 • 	 . — — . 	 • 	 • 	 1 	 • 	 1
            60  57  55  42 73  37  78  79  18  15  09

                            STATION
FIGURE  60.  Geographical distribution (biomass) of zooplankton groups,
1984.  Copepoda refer to the  nauplius stage of the Copepoda , Lake Erie

-------
                                               228



a:
UJ
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. 4-

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o Cyclops bicuspidatus
thomasa
LAKE ERIE « x Diaptomus oregonensis
/' \ * Daphn i a pulicaria
\n Mesocy clops edax
/ \
/\
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u
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^^*^v x jt \ w / *S / ^v_.
L^ H ^» * ^VL \ « i *m / TQ
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o Ho 1 oped i urn gibberum
x Tropocyclops prasinus
mexicanus

/ \
/
^ -A /.A --
,«/' ^N^' Nt---*' y'/ ^^

          60  57 55 42 73  37  78  79  18  15 09
                          STATION
FIGURE 61. Geographical distribution of selected Crustacea in Lake Erie,
1984.

-------
                                             229


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LAKE LKlt 0 Notholca foliacea
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i_ 	 . 	 • 	 1 	 1 	 1 — * — — • 	 • 	 ' 	 ' — — • 	 1
         60 57  55  42 73 37  78  79  18  15  09
                         STATION
FIGURE 62. Geographical distribution of selected Rotifera in Lake Erie,
1984.

-------
CC
LJ
LL!
CD
       100T
        80-
        60--
         40-
        20-
          0
LAKE     ERIE
   o  Copepoda
   x  Cladocera
   •*•  Total  Crustacea

	?	.	_	
              —'—

                1940
                     1960
                  1980
             FIGURE 63. Crustacean zooplankton abundance since 1939  in the western
             basin of Lake Erie.  1939 data are from Chandler (1940; April-October).
             1949 data are from Bradshaw (1964; April-October).  1983  data are from
             Makarewicz (1987; April-November) and 1984 (This study; April-December).
             Values are the mean ± Standard Error.
                                                                      ro
                                                                      oo
                                                                      o

-------
UJ
h-
I—»
_J
\
a:
LU
m
150-
120-
90-
60-
30-
n
LAKE bKIt
o CladocQra • \
' x Adult Copepoda / «
-«- Total Copopoda * / \
/ \ 1 \
/ V -
/ K \ '
• / i \ 	 \_t
s ^f\ \\
^'- ^JEf v^ 	 KO
°^1 	 , 	 1 — — . 	 1 	 . 	 1 	 ' 	 1 	 •
                1940
1950      1960      1970
1980
          FIGURE 64.  July and August  abundance  of Cladocera and Copepoda in the
          western basin of Lake Erie since 1939.  Total Copepoda refers to adults
          plus the nauplius stage.  Data are from Chandler  (1940), Bradshaw (1964),
          Hubschman (1960), Britt et al. (1973),  Davis (1969b) Makarewicz (1987) and
          this study.  The number of adults and total copepods in 1939 and 1959
          follow Bradshaw's (1964) calculations.
                                                                                         ro
                                                                                         CO

-------


QL
Ul
•
r-
t— i
-J

^

o:
LU
m

«•_
2

240-
200-

160-


120-



80-


40-
P>
•
j\ + 1970
/ \ x 1983
• i
- i *
i %
/ \ o 1984
v
/ ^
/ V
' V
/ A
/ .
/ *3**~96^
| ^^0 \ ^^^^
*"fU0^" ... ^ 	 .
A'MJJASONDJF
FIGURE  65.  Seasonal fluctuation of weighted mean Crustacea (nauplii
excluded) abundance in 1970,  1983, 1984,  Lake Erie.  1970 data follow
Watson and Carpenter (1974).   1983 data from Makarewicz (1987).  1983
and 1984 values are corrected by using the weighting factors of 15.6%,
59.6% and 24.6% for the western, central  and eastern basins (after Munawar
and Munawar 1976).
tXJ
CO

-------




o:
UJ
i 	
i —
\
o:
LU
CD
Z)

720j A


600-


480-
360-


240-
120-
0-
J\

i v
I \
1 x
" i y
\\f
/
i /
• J/

i — i — i
LAKE ERIE
A
/ \
/ \
ff \
v/ \ o 1939
» \ x 1961
\ A D 1983
\ /X * 1984
\ • V x
\ / \\
\ .' \ \
\ \ \
\ 1 V \
V \ TX-
* XT \
/ \i \
/* u * *•
	 , 	 , 	 , 	 1 1 	 1 1 1 	 1
   AMJJASONDJ
                          MONTHS
FIGURE  66.  Seasonal fluctuation of Rotifera in  the western basin of Lake
Erie from  1939 - 1983.  Sources:  1939 - Chandler  (1940); 1961 - Britt et
al. (1973); 1983 - Makarewicz  (1987).  The 1970 samples of Nalepa (1972)
are not  included because they  are from the far western end of the basin and
may not  be represented of the  entire basin.
ro
oo
co

-------
01
X
01
35 j

30-

25-

20-

15-

10-

  j" •

  0--
               New  York  waters
                  Western  basin
                i
ji
                      1975
                                     1980
                     1985
        Figure 67.  Abundance (millions of fish) of fishable walleye (age 2+)
        in western Lake Erie (Ohio waters).  For New York waters, values
        represent catch per 22 net meters in variable mesh nets. Data from
        Ohio Department of Natural Resources (1985) and New York State
        Department of Environmental Conservation (1985).
                                                                                oo

-------
a
o
a
 L
 O

X
auu-
600-
400-
200-
0-
»
Wai leye
m

1975 1980
1



\

— L»— 	 1
1985
Figure 68. Sport angler harvest of walleye from the central basin of
Lake Erie. Modified from the Ohio Department of Natural Resources.
                                                                                                                                                    ro
                                                                                                                                                    oo
                                                                                                                                                    en

-------
                                                       236










a
o
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3
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3000-
2500-
2000-

1500-

1000-
500-

240-

200-
160-
120-
80-
40-
1500:
1200-

900-


600-

300-

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1 • M ew i f e
/\
I •
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-------
25-r
20-
15-
10-
 0--
LAKE   ERIE
/A
\   \
 Y\
o
X
       18  April
       19  August
        4  DecQmbQr
               V
                           -*
                   --x
                         -I	I	1	1	I	1
     60 57 55  42 73 37 78 79  18  15 09
                 STATION
  Figure 70.  Seasonal and geographical turbidity trends in Lake Erie,
  1984.
                                                      ro
                                                      oo

-------
                                             238
        Table A

      Phytoplankton
Species List:  Lake Michigan

-------
                                                             239
          SPECIES LIST - LAKE MICHIGAN PHYTOPLANKTON  (1983)

DIV  TAXON                                     AUTHORITY
bAC
Achnanthes af f ini s
Achnanthes bi aso 1 et t iana
Achnanthes clevei
Achnanthes clevei v. rostrata
Achnanthes conspicua
Achnanthes deflexa
Achnanthes ex igua
Achnanthes exigua v. constricta
Achnanthes f 1 exe 1 la
Achnanthes hatckiana
Achnanthes
Achnanthes
Achnanthes
Achnant hes
Achnanthes
Achnanthes
anceo la ta
anceo la ta v
apponica v.
apponica v.
i near i s
i near i s to.
Achnanthes rainutissima
Achnanthes oestrupii v.
Achnanthes sp.
Achnanthes suchlandtii


. dub ia
n i ncke i
n i ncke i ?

cur ta

1 anceo lata


     Actinocyclus noriranii f. subsalsa
     AmphipIeura pellucida
     Amphora  ova I i s
     Amphora  ovalis v. aff ins
     Amphora  ovalis v, pediculius
     Amphora  perpusilla
     Amphora  sp.
     Amphora  thumensus
     Anomoeoneis vitrea
     Asterionella forrrosa
     CaI one is sp.
     Cocconeis diminuta
               d i scuILS
               placentula v. euglypta
               placertula v. lineata
               thutr.ensis
                ant i qua
                ant i qua?
                at orrus
                comensi s
                comensis - auxospore
                correns i s v . 1
                c o rii e n s i s v . Z
                con ta
                coirta - auxospore
                corrta v. o I i gact is
                crypt i ca
                kuetz i ng iana
                meneghi n iana
                m i ch igan iana
                michiganiana - auxospore
                oce I I ata
Coccone i s
Cocconei s
Coccone i s
Coccone i
Cyclotel
Cyclotel
CycIote
Cyclote
Cyclote
CycIote
Cyclote
CycIote
Cyclote
Cyclote
Cyclote
CycIote
CycIote
CycIote
CycIotella
CycIoteI la
s
 a
 a
 a
 a
 a
 a
 a
 a
 a
 a
la
la
la
la
Gr un.
(Kutz.) Grun.
Gr un .
Hust.
A. Mayer
Re in.  in Patr . £ Re i m.
Grun.
(Grun.) Hust.
(Kutz.) Brun.
Grun.
(Breb.) Greg.
Grun.
(Guerm. t Hang.) Reicn.
(Guerm. £ Mang.) Rein.
(M.  Sin.) Grun.
H.L. Sm.
Kutz.
Hust .

Hust.
(Juh I ,-Oannf.) Hust.
(Kutz.) Kutz.
(Kutz.) Kutz.
(Kutz.) V.H. ex DeT.
(Kutz.) V.H. ex DeT.
(Grun.) Grun.

(Mayer) A. Cl.
(Grun.) Patr. £ Re im.
Hass.

Pant.
(Schum.) Cl.
(Ehr .) Cl .
(Ehr .) Cl.
A. Mayer
W. Sm.
W. Sir.
Pant.
Gr un.
(Ehr.) Kutz.

(Ehr.) Grun.
Re i m. et a I.
Thw.
Kutz.
Skv.

Pant.

-------
                                                                240
          SPECIES LIST - LAKE MICHIGAN PHYTOPLANKTON (1983)

DIV  7AXON                                     AUTHORITY
6AC
Cyc I otel la
Cyc I otel la
CycIote I I a
Cyclotel la
CycIoteI la
CycIoteI la
CycIoteI la
CymatopIeu ra
Cymatop I eura
                opercuI a ta
                operculata unipunctata
                pseudosteI I j gera
                sp.
                sp. SI
                sp . - au xo spor e
                ste I I i ge ra
                  e I I i pt i ca
                  so I ea
Cymbe I la cesat i i
Cymbella cistula  v. gibbosa
Cymbe I la de I i catu la
Cymbella micrccephala
Cymbe I la tn i nuta
Cymbella minute  v. silesiaca
Cymbella norvecjca
Cymbella prostrata v. auerswaldii
Cymbe I I a si nuata
Cymbe I la sp.
Cymbella trianculum
Denticula tenuis  v. crassula
D latoma tenue
Diatoma tenue  v.  elongatum
Dip) one is e I I i pt i ca
D i p I one is ocu I a ta
Dip! one i s parnr.a
Dip! one is sp .
Entomoneis ornata
Eunot i a i nc i sa
Fragilaria brevistriata
           brevistriata
           brevistriata
           capucina
           capucina v. mesolepta
           construens
           construens
           construens
           construens
           construens
           crotonensis
           leptostauron
           pi nnata
           pi nnata v.
           pi nnata v.
           sp.
           vaucheriae
           vaucheriae
           affine
           dichotomum
           gracile
           parvulum
           sp .
     Fragilaria
     Fragilaria
     Fragilaria
     Fragilaria
     Fragilaria
     Fragilaria
     Fragilaria
     Fragilaria
     Fragilaria
     Fragilaria
     Fragilaria
     Fragilaria
     Fragilaria
     Fragilaria
     Fragilaria
     Fragilaria
     Fragilaria
     Gomphonema
     Gomphoneira
     Gomphonema
     Gomphonema
     Gomphonema
  v.
  v.
  i nfIata
  subcapi tata
v.
v.
v.
v.
b i nodi s
m i nuta
subsaIi na
venter
intercedens
lancettu I a
v. cap i teI Iata
                    (Ag.) Kutz.
                    Hust .
                    Hust.
(Cl. £ Grun.) V.H.
(Breb. )  W.Sm.
(Breb. £ Godey) W. Sir.
(Rabh.)  Grun. ex A.S.
Br un.
Kutz.
Grun.
Hi Ise
(Blei sch ) Re im.
Gr un.
(Rabh.)  Re i m.
Greg.

(Ehr .) C I .
(Nag.) W. £ G.S. West.
Ag.
Lyngb.
(Kutz.)  Cl.
(Breb.)  Cl.
Cl.

( J.W. Ba i I .) ke im.
W. Sm.
Gr un .
(Pant.)  Hust.
Gr un .
Desm.
(Rabh.)  Grun.
(Ehr .) Grun.
(Ehr . ) Grun.
Temp . £  Per.
Hust.
(Ehr . ) G run.
Kitton
(Ehr.) Hust.
Ehr .
(Grun.)  Hust.
(Schum.) Hust.

(Kutz.)  Peters.
(Grun.)  Patr.
Kutz.
Kutz.
Ehr. em. V.H.
Kutz.

-------
                                                             241
          SPECIES LIST - LAKE MICHIGAN PHYTOPLANKTON (1983)

DIV  TAXON                                     AUTHORITY
                        v« angustissima
                             subarcti ca
                         s i gnata
                         subsa I sa

                        •  hur gar i ca
6AC  Gyrosigma scictense
     Me Ios i ra ambi qua
     Me I os i ra a i starts
     Me I os i ra granu lata
     Melosira granulata
     Me I os i ra i s I and i ca
     Melosira i taI ica
     Melosira italica suDsp.
     Melosira sp.
     Meridion circulare
     Nav i cu la angI ica v.
     Navicula angIica v.
     Nav i cuI a capitate
     Nav i cula cap i tata v
     Navicula cincta
     Navicula cryptocephaI a
     Navicula cryptocepha I a v. veneta
     Navicula exigua v. capitata
     Nav i cula grac i I o ides
     Nav i cuI a gregar i a
     Nav icula irtegra
     Navicula jaerneteIdtii
     Nav icula I a custr i s
     Navicula lanceolata
     Navicula meniscu!us v. upsaliensis
     Nav icula m in i ira
     Navicula pseucoreinhardtii?
     Nav i cula pupu la
     Nav i cuI a rad i osa
     Navicula radiosa v. tenella
     Nav i cuI a re inharoti i
     Navicula seminu I oides
     Navicula seminulum
     Nav icula sp.
     Navicula tripurctata
     Navicula tripunctata v.  senjzonemoi
     Nav icula tuscu I a
     Navicula v i r i duI a
     Neiduim sp.  #1
                                        des
     N i tzsch i a
     N i t z s c h i a
     Nitzsch ia
     Ni tzsch ia
     N i tzsch ia
     Ni tzschia
     Ni tzschia
     Ni tzschia
     Ki tzsch i a
     Nitzsch ia
     N i tzsch ia
     N i tzsch i a
     Ni tzsch ia
               ac i cular i oides
               ac i cu lar i s
               acu I a
               acute
               arnph i b i a
               angustata
               angustata  v. acuta
               bacata
               cap i teI I a ta
               conf i n i s
               c o n f i n i s ?
               d i ss ipata
               f ont i cola
(Sul I i v. t. Worm I ey )  Cl
(Grun.)  0. Mull.
(Ehr.)  Kutz.
(Ehr.)  Ralfs
0. MulI.
0. Mul I .
(Ehr.)  Kutz.
0. Mul I .

(Greg.)  Ag.
Hust.
(Grun.)  CI.
Ehr .
(Grun.)  Ross
(Ehr.)  Ralfs
Kutz.
(Kutz.)  Rabh.
Patr .
A. Mayer
Donk .
(k. Sm.) RaIfs
Hust.
Greg.
(Ag.)  Kutz.
(Grun.)  Grun.
Gr un.
Patr .
Kutz.
Kutz.
(breb. )  Cl. £ Mo I  I.
(Grun.)  Grun.
Hust.
Grun .

(O.F.MulI.) Bory
(Breb.  ex Grun.)  V.H.
Ehr .
(Kutz.)  Ehr.

Arch,  non Hust.
(Kutz.)  W. Sm.
Hantz.  ex Cl . I Gr un.
Hantz.
Grun.
(W. Sm.) Grun.
Gr un .
Hust.
Hust.
Hust.
Hust.
(Kutz.)  Gr un.
Grun .

-------
                                                              242
          SPECIES LIST - LAKE MICHIGAN PHYTOPLANKTON  (1983)

DIV  TAXON                                     AUTHORITY
8AC -sNitzschia frustulum
   ~ Nitzschia frustulum v. minutula
   " Nitzschia gancersheim i ensis
    -Nitzschia gracilis
    -N i tzsch ia i mp ressa
    ^Nitzschia kuetzingiana
    -Nitzschia lauerburgiana
     Nitzschia linear is
     Nitzschia pa Iea
     Nitzschia palea v. debilis
     Nitzschia paleacea
     N~i tzsch ia pura?
     ffitzschia recta
     N'itzschia romana
     N-J tzsch ia sociabi I i s
     Nitzschia sp.
     N'itzschia spicUum
    '•Nitzschia subacicularis
    "Nitzschia sublinearis
     'Nitzschia sublinearis?
     tiitzschia subrostrata
    -N i tzsch ia tenu is
     "Nitzschia valdestrita
     Opephora  mar ty i
     Rh i zosoI en i a er i ens is
     Rhizosolenia Icngiseta
     Rh i zosoI en ia sp.
     Rhoiocosphenia curvata
     Skeletonema  pctairos
     Stauroneis smfthii  v. rcinuta
     Stephanodiscus alpinus
     Stephanodiscus
     Stephanod i scu s
     Stephanod i scus
     S tephanod i scus
     S tephanod i scus
     Stephanodiscu s
     Stephanod i scus
     S tephanod i sc us
     S tephanod i sc us
     S tephanod i scus
     Stephanod i scus
     Stephanodiscus
     Stephanod i scus
     S tephanod i scus
a Ip inus?
binder anus
b i nder anus?
hantzsch i  i
m i nu tus
n iagarae
sp.
sp. S03
sp. -auxospore
subt tlis
tenu is
tenuis v.  #01
tenu is v.  #02
tenu i s?
tran si I van i cus
     Stephanod i scus
     Sur i re I I a  angusta
     Synedra  amphicephaI a  v.
     Synedra  eye Iopum
     Synedra  delicatissima  v
     Synedra  fame i I i ca
     Synedra  f i I if ormi s
         austr i ca
          angust iss ima
 (Kutz. )  Grun .

 Kr asske
 Hantz.
 Hust .
 Hi Ise
 Hust.
 W. Sir.
 (Kutz.)  W.  Sm.
 (Kutz. 1  Grun .
 Gr un .
 Hust .
 Hantz.
 Grun .
 Hust.

 Hust .
 Hust.
 Hust.
 Hust.
 Hust .
 W. Sir.
 Aleem £  Hust.
 Her i b.
 H.L. Sm.
 Zach .

 (Kutz. )  Grun.
 (Ueber)  Hasle
 Haw.
 Hust.
 Hust .
 (Kutz.)  Krieg.
 (Kutz. )  Kr ieg.
 Grun.
 Gr un .
 Ehr.
                                         £  Evens,
(Van Goor )
Hust.
           A. C I
Hust.
Pant.
Kutz.
(Grun. ) Hust.
Br ut schy
Gr un.
Kutz.
Grun.

-------
                                                             243
          SPECIES LIST - LAKE MICHIGAN PHYTOPLANKTON  (1983)

DIV  TAXON                                     AUTHORITY
BAG
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
T a b e I 1 a r i a
Tabe I I ar i a
f i I i f oriri s
mini scula
paras i t i ca
rad i ans
sp.
u I na
u I na
u I na
v.  e x i I i s
                  v. chaseana
                  v . dan i ca
             ulna v . I on gi ss ima
                f erestrata
                ferestrata v. geniculata
     Tabe I lar i a
     Tabe I (aria
           fIocculosa
           fIocculosa
                 Ii near i s
CAT  VacuoIar i a sp •

CHL  Ankistrodesmus
     Ankistrodesmus
     Ank i strodesmus
     Ank i strodesmus
     Arthrodesmus bi
               fa Ic at us
               geIi factum
               sp.  #01
               sp.?
               f idus
     Botryoc occus
     Carte r i a sp.
     Chlamydocapsa
     ChIamydocapsa
     ChIamydomonas
     Ch tamydoironas
     Chlamydoironas
     C I osteri ops i s
     C I ost er i urn
     C I oster i urn
     CoeI astr urn
     CoeI astr urn
     Coelastrum
     Coenocyst i s
             Braun i i

              plank toni ca
              sp.
              sp.
              sp. -  ovo i d
                  -  sphere
              sp
              sp.
           ac i cular e
           grac i le
           carrbr i cum
           m i cropor urn
           sp.
            sp.
     Co smar i um sp .
     Crucigenia irregularis
     Crucigenia quacrata
     Crucigenia rectangu laris
     D i ctyosphaer
     D i ctyosphaer
     EIakatothr i x
     Elakatothr i x
     EIakatothr i x
     G I oedact i n i urn
     Go I enk i n t ps i s
     Green cocco i d
     Gr een cocco i d
     Green cocco i d
     Green coccoid
     Green cocco i d
     Green coccoid
             um ehrenbergianum
             urr pu I chel I um
             geI at i no sa
             v i r idi s
             viridi s?
              I irrne t i cum
              sp.

              #04
              - aci cular
              - bac iI I  iform
              - b i c e I I s
              - fus iform
A. C I.
Gr un.
W. Sm.
Kutz.

(Nitz.) Ehr.
Thomas
(Kutz.) V.H.
(W. Sm.)  Brun.
Kutz.
A. Cl .
(Roth) Kutz.
Koppen
                                  (Corda) Ralfs
                                  (Chod.) Bourr.
                                  Br eb.
                                  Kutz.

                                  (w. £ G.S. West) Fott
                       T.  West
                       Br eb .
                       Arch .
                       Nag. in A
                       Wi II e
                       Mor r en
                       A.  Braun
                       Nag.
                       Wood.
                       Wi I le
                       (Snow)
                       (Snow )
                                             Braun
                                         Printz
                                         Printz
                                  G . M . Sm .

-------
                                                              244
          SPECIES LIST - LAKE MICHIGAN PHYTOPLANKTON  (1983)

DIV  TAXON                                     AUTHORITY
CHL  Green coccoid
     Gr een coccoid
     Gr een coccoid
     Green coccoid
     Gr een cocco i d
     Gr een coccoid
     Kirchneriella
     Monoraph i d i urn
     Monoraph i d i urn
     Monoraph i d i urn
     Monoraph id i urn
     Monoraph id i urn
     Monoraph id i urn
     Nephrocyt i urn
     Nephrocyt i urn
     Gedogonium sp
  - fusiform bicells
  - oocyst i s-l ike bi ceI I
  - ova I
  - ren i f orm
  - sphere
  - sphere ( large)
  conto r ta
  contor turn
  i r reguI are
  ir i nut urn
  saxat i I e
  set i f orraae
  torti le
 Agarch ianum
 I linnet i cum
     Oocyst i s  sp.
     Oocystis  sp. #1
     Oocyst is borge i
     Oocyst is crassa
     Oocyst i s I acustr i s
     Oocys t i s mar s en i i
     Oocyst is parva
     Oocyst is pusi I I a
     Oocyst i s so I i tar ia
     Oocyst i s submar i na
     Ped iastrurn  sp.?
     Phacotus minuscula
     Phacotus sp.
     Planktonema lauterbornii
     PIank tonema sp .
     Pter omona s  sp .
     Pyramidomonas sp.
     Scenedesmus act-ir, i na tus
     Sceneoesirus
     S c e n e d e s nr u s
     Scenedesirus
     Scenedesmus
     Scenedesmus
     Scenedesmus
     Scenedesmus
     Schro eder i a
     Sphae reI Iocys t i s
     Sphae re I Iocyst i s
eccr nis
quaar i cauda
quadricau da v.
s ec ur if or mi s
ser ratu s
sp.
sp inosus
set i ge r a
     I a cu str i s
     lateral is
I ongsp i na
     Sphaerocystis  schroeteri
     St i chococcus  sp.
     Tetraedron  caudatum
     Tetraedron  minimum
     Tetraspora  lacustris
     Tetrastrum  glabrum
     Treubaria  planktonica
     Treubar i a  set i gera
               (Schirid.) Bohlir,
               (Thur.) Kom.-Legn.
               (G.M. Sm.) Kom.-Legn.
               (Nag.) Kom.-Legn.
               Kom,-Legn.
               (Nyg.) Kom.-Legn.
               (H. £ h.) Kom.-Legn.
               Nag.
               (G.M. Sm.) G.M. Sm.
                              Snow
                              Mi ttr .  in Wi ttr. £  Nord,
                              Chod.
                              Lemm.
                              West  £  West
                              Hansg.
                              Wi ttr .  in Mi ttr . £  Nor d,
                              Lagerh .

                              Bour r.

                              SchmidIe
(Lagerh.) Choa.
(RaIfs) Chod.
( Tur p. ) Breb.
(Chod.) G.M. Stn.
Playf.
(Corda) Bohlm

Chod .
(Schroed.)  Lemm.
Skuja
Fott  £ Novak.
Chod.

(Cor da) Hansg.
(A. Braun)  Hansg.
Lemm.

(G.M. Sm.)  Korch.
(Arch.) G.M. Sm.

-------
                                                              245
          SPECIES LIST - LAKE MICHIGAN PHYTOPLANKTDN  (1983)

UIV  TAXON                                     AUTHORITY
CHR  Bi tr i ch ia chodat i i
     B i tr i ch j a ohri di ana
     Chromul ina sp.
     Chrysococcus sp.?
     Chrysolykos angulatus
     Chrysolykos planktonicus
     Chrysolykos skujae
     ChrysoIykos sp.
     Chrysosphaere I I a longispina
     D i nobr yon - cyst
     Dinobryon acuirinatum
     Dinobryon bavaricum
     D i nobr yon bor ge i
     Dinobryon cylinoricum
     Oinobryon divergens
     Oinobryon eurystcma?
     Dinobryor sertularia
     D i nobr yon soc i a Ie
     Dinobryon sociale v.
     Dinobryon sociale v.
     D i nobr yon sp•
     Dinobryon stokesii  v.
     Dinobryon tubaeformae
     Dinobryon utriculus v
     Halobryon sp . ?
     Haptophyte sp.
           airer i canum
           sti ptatum

            ep i p lank ton i cum

           . tabeI Iar ia e
               sp,
               sp.
               sp.
               sp.
Kephyr i on
Kephyrion
Kephyr i on
Kephyr i on
Kephyr i on
Kephyr ion
Kephyr i on
Kephyr i on
Ma I Iomonas
Mai Iomonas
Ma I Iomonas
Qchromonas
Ochromonas
Ochromonas
Ochromonas
Paraphysomonas
Paraphysomonas
Pseudokephyr i on
Pseudokephyr i on
Pseudokephyr i on
Pseudokephyr i on
Pseudokephyr i on
Un ident i f ied
Un ident i f i ed
               cupu I i for mae
               do I  i oI urn
               rub i-caIu st r i
    #1 -Pseudokephyrion entzi
    #2
    #3
sp i rale
 majcrens is
 sp .
 sp.  #3
 sp.
 sp.  - ova I
 sp.  - ov oi d
 sp.  - sphere
     sp.
     sp.?
      con i cuir
      I at urn
      ir i I lerense
      sp. #1
      undu lat i ss imum
   cocco i a - ovo i 0
   coccoi a - sphere
                                           (Rev .) Chod.
                                           (Fott) Mich.
                                           (Wi I len) Niauw.
                                           Mack .
                                           (Nauw.) Bourr.

                                           Laut. em. N i ch.

                                           Rutt.
                                           Imhof
                                           Lemm.
                                           Imhof
                                           Imhof
                                           (Stokes) Lemm.
                                           Ehr .
                                           Ehr .
                                           (Br unnth.)  Bac hm,
                                           (S te in) Lemrr,.

                                           Sku ja
                                           Nyg.
                                           Lemm.
                                Conr .
                                Conr .
                                Conr .
                                                (Lack.) Conr.
                                                Skuja
                                                (Sen ill.) Schuir.
                                                (Sen i I I . 1 Schutn.
                                                Mich.

                                                Scherff.
     Un identif i ed
     Un i dent i f i ed
             coccoi ds
             I or ica te -
                             sphere

-------
                                                                246
          SPECIES LIST - LAKE MICHIGAN PHYTOPLANKTON (1983)

UIV  TAXON                                     AUTHORITY
CHR  Unidentified I oricate-fI age I I ate sphere
COL
CRY
Bicoeca campanulata
Bicoeca lacustris?
B i coeca m i t ra v.?
Bicoeca sp.
Bicoeca sp. *C4
B i coeca tub if ormi s
Codonosiga sp.
Co I or less f lage Mate
Co I or I ess f I age I late
Colorless flagellates
Hast i geI I a sp.
Monos iga ovata
Salpingoeca arrphcrae
SaIp ingoeca grac iIi s
SaIp ingoeca sp.
Stylotheca aurea
                          - ovoid
                          - sphere
Chroomonas
Chroomonas
Chroomonas
Chroomonas
Cryptomonas
Cr yptomonas
Cryptomonas
Cr yptomonas
Cryptomonas
Cryptomonas
Cr yptomonas
Cr yptorconas
Cr yptomonas
Cr yptomonas
Cr yptomonas
Cr yptomonas
Cr yptomonas
Cr yptomonas
Cr yptomonas
Cr yptomonas
Cr yptomonas
Cryptomonas
Cr yptomonas
Rhodomonas
Rhodomonas
                acita
                caudata
                norstedt i i
                pochmann i
                 - cyst
                 brev i s
                 brev is?
                 caudata
                 er csa
                 ercsa  v. reflexa
                 I obata
                 mar sson i i
                 rrar sson i i  v. ?
                 o vat a
                 parapyrenoidifera
                 phaseolus
                 pus iI la
                 pyreno i oi fer a
                 reflexa v. er osa
                 rostrat if ormi s
                 sp.
                 tenu i s
                 tetrapyreniodiosa
                lacustr i s
                lens
CYA
Rhodomonas mi nuta
Rhodomonas roinuta v.
Senn i a par vuI a
Senn ia parvuI a?

Anabaena flos-aquae
Anabaena sp.
Anacyst is mar i na
                          nannoplankt i ca
                                               (Lack.) Bourr. em. Skuja
                                               J. Clark
                                               Skuja
Kent
Kent
Clark

(Bachm.) Bo Ioch.

Uterm.
Ge i t.
Hansg.
Huber-Pest.

Schi I I .
Schi I I .
Schi I I .
Ehr.
Marss.
Korsch.
Skuja
Skuja
Ehr .
Skuja
Skuja
Bachm.
Geit I .

Skuja

Pasch.
Skuja
Pasch. t kutt.
Pasch. £ Rutt.
Skuja
Skuja
Skuja
Skuja

(Lyngb.) Breb.

(Hansg.) Dr.  t Dai ly

-------
          SPECIES LIST - LAKE MICHIGAN PHYTQPLANKTON  (1983)

DIV  TAXON                                     AUTHORITY
                                                              247
CYA  Anacystis montana
     Anacystis montana v. minor
     Anacyst i s therrra I is
     Aphanothece gelatinosa
     Coccochloris elabans
     Coccochloris peniocystis
     Coe I osphaer i uir, naegelianum
     DactyIococcopsis Smithii
     DactyIococcopsis sp.
     Gioeothece ruprestris
     Gomphosphaeria lacustris
     Lyngbya limneticuro
                  acardh i i
                  Iimnet i ca
                  I imnetica?
                  ir in ima
                  sp.
                  subbre v i s
                  tenu i s
                  tenu i s v . natans
                  tenuis v. tergistina
                  bIue-g reens
PYR
UNI
Osci
Osc i
Osc i
Osc i
Osci
Osc i
Osci
Osc i
Osc i









ato r i a
ato r i a
ato r i a
ato r i a
ato r ia
ato r i a
ator i a
a to r i a
ator i a
     Unident if fed
EUG  Euglena sp.
     Amph i d in i urn sp.
     Ceratium hirundinella
     D i nof I age I late cyst
     Gymnod i n i urn sp•
     Gymnodiniurn sp. #1
     Gymnodiniurr sp. #2
     Gymnod i n i urn sp. #3
     P er i d i n i urn c i nc turn
     Peridinium inconspicuum
     Per i din i urn sp .
     Unident if i ed
     Un ident if i ed
     Unident i f i ed
     Un ident i f i ed
     Un i dent i f i ed
ccccoi d fI age Ilates
f lagel late  #01
f lagel late  #03
f I age I late  - ovo i d
flagellate  - spherical
                                               Dr. £ Da iIy
                                               Dr. £ Dai ly
                                               (Menegh. )  Or.  £ Dai I y
                                               (Henn.) Lemm.
                                               Or . £ Da iIy
                                               (Kutz. ) Dr .  £  Daily
                                               Unger
                                               Chod. £ Chod.

                                               (Lyngb.)  born.
                                               Chod.
                                               Lemm.
                                               Gom.
                                               Lemm.
                                               Lemm.
                                               GickIn.

                                               S c h m i d .
                                               C.A. Ag.
                                               Gom.
                                               (Kutz.) Rabh.
                                               (O.F .hulI.) Schrank
                                               (MulI.) Ehr.
                                               Lemm.

-------
                                           248
        Table B,

      Phytoplankton
Species List:  Lake Huron

-------
                                                             249
            SPECIES LIST - LAKE HURON PHYTOPLANKTON 11983)

OIV  TAXON                                     AUTHORITY
BAC
Achnanthes
Achnanthes
Achnanthes
Achnanthes
Ac hnanthes
Achnanthes
Achnanthes
Achnanthes
Achnanthes
Achnanthes
Achnanthes
Achnanthes
Ac hnanthes
Achnanthes
Achnanthes
Achnanthes
Achnanthes
Achnanthes
Achnanthes
Achnanthes
Achnanthes
Amphipleura
aff inis
bi asolett i ana
brevipes v. intermedia
cI eve i
c I eve i  v . cost rata
consp i cua?
aetha
ex igua
exigua  v. heterovalva
f I exe I la
ha uckj ana
 anceolata
 anceolata  v. dubia
 apponica v. ninckei
 aterostrata
  i near i s
  i near is to. cur ta
marg i nuIata
mi crocepha la
m i nut i ss ima
sp.
 pe I Iuc i aa
                          v. grac iI Iima
Amphora cotfeiformis
Amphora inariensis
Ampho ra ova I i  s
Amphora oval is v. pediculius
Amphora perpusilla
Amphora sp•
Anomoeoneis vitrea
Asterionella  forirosa
Asterionella  forirosa
CaI one i s bac iI I urn
Cocconeis diminuta
Cocconeis disculis
Cocconeis placentula v. euglypta
Cocconeis placertula v. lineata
CycIostephanos duoius
CycIotelI a ant i qua?
           catenata
           comens i s
           cofrensis - auxospore
           coirens i s v . 1
           correns i s v . 2
           c o  m t a
           comta - auxospore
           coirta v. #2
           corrta v. o I igact is
           crypt ica
           kuetz ingiana
           kuetz ing iana v.
           kuetz i ng iana v.
           kuetz i ng iana v.
Cyclotel la
Cyclotella
Cyc I otella
CycIoteI 1
Cyclotel I
Cyclotel I
CycIotel 1
CycIotelI
Cyclotel
Cyc I oteI
CycIoteI
CycIotel
CycIotel
Cyclotel
                                pIane tophora
                                plane tophora?
                                rad i osa
Gr un .
(Kutz.)  Grun.
(Kutz.)  Cl.
Grun.
Hust.
A. Mayer
Hohn £ HelIerm.
Grun .
Kr as ske
(Kutz.)  Brun.
Grun.
(Breb.)  Greg.
Grun.
(Guerm.  £ Mang.) Rein.
Hust.
(M. Sm.) Grun.
H.L. Sm.
Grun.
(Kutz.)  Grun.
Kutz.

(Kutz.)  Kutz.
(Ag.) Kutz.
Kr am.
(Kutz.)  Kutz.
(Kutz.)  V.H. ex DeT.
(Grun.)  Grun.

(Grun.)  Patr. £ Reim.
Mass.
(Hantz.) Grun
(Grun.)  CI.
Pant.
(Schum.) Cl.
(Ehr.) Cl.
(Ehr.) Cl.
(Fr i eke  ) Round
M. Sm.
Br un .
Grun.
                                                (Ehr.) Kutz.
                               (Ehr.) Grun.
                               Re i m. et al.
                               Thw.
                               F r i c k e
                               Fr i eke
                               Fr icke

-------
                                                             250
            SPECIES LIST - LAKE HURON PHYTOPLANKTON (1983)

DIV  TAXON                                     AUTHORITY
BAG  Cyclotella meneghintana
     Cyclotella michiganiana
     Cyclotella ocellata
     Cyclotel la operculata
     Cyclotella pseudoste I I igera
     Cyclotella sp.
     Cyc lotel la sp. #1
     Cyclotel la sp. #2
     Cyclotella sp. - auxospore
     CycIoteI la ste I I  igera
     Cymatopleura solea v. apiculata
     Cymbella  angustata
     CymbeI I a  laev i s
     Cymbella  micrccepha la
     CymbeI I a  m i nuta
     Cymbella  minuta v. silesiaca
     Cymbella  navicu I  iformjs
     CymbeI la  sp.
     Cymbella  trianculum
     Dent i cu la sp.
     Denticula tenuis v. crassula
     D iatoma tenue
     Diatoma tenue v. elongatum
     Di pI one is eI I i pt i ca
     D i pI one is ooIongeI I a
     D i pI one i s ocu I sta
     Entomoneis ornata
     Eunot ia praerupta
     Fragilaria brevistriata
     Fragilaria brevistriata v. subcapitata
     Fragilaria capucina
     Fragilaria capucina v.  mesolepta
     Fragilaria construens
     Fragilaria construens
     Fragilaria construens
     Fragilaria construens
     Fragilaria construens
     Fragilaria crotcnensis
     Fragilaria intermedia v
     Fragilaria leptostauron
     Fragilaria leptostauron
     Fragilaria pi nnata
     Fragilaria pinnata v.
     Fragilaria pi nnata v.
     Fragilaria sp.
     Fragilaria vaucheriae
     Gomphonema angustatum
     Gomphoneira dichotomum
     Gomphoneir.a grac i le
     Gomphonema olivaceum
     Gomphonema parvulum
     Gomphonema sp.
v.
v.
v.
v.
minuta
pumiI a
subsa I i na
venter

fat lax

.  dub ia
intercedens
lancettuI a
                    Kutz.
                    Skv.
                    Pant.
                    (Ag.)  Kutz.
                    Hust.
(Cl . t Grun.) V.H.
(W.  Sm.) Ralfs
(W.  Sm.) Cl.
Naeg. ex Kutz.
Gr un .
Hi Ise
(B le isch )  Re im.
Auer sw.

(Ehr .) CI.

(Nag.) W.  £ G.S. West.
Ag.
Lyngb.
(Kutz.) Cl.
(Naeg.ex Kutz.) Ross
(Breb.) Cl.
( J.W. Ba i I.) Re im.
Ehr.
Grun.
Grun.
Desm.
(Rabh.) Grun.
(Ehr .) Grun.
Temp . £ Per •
Grun.
Hust.
(Ehr .) Grun.
Ki tton
(Grun.) Stoerm. £ Yang
(Ehr.) Hust.
(Grun.I Hust.
Ehr.
(Grun.) Hust.
(Schunu ) hust.

(Kutz.) Peters.
(Kutz.) Rabh.
Kutz.
Ehr. em. V.H.
(Lyngb.) Kutz.
Kutz .

-------
                                                              251
            SPECIES LIST - LAKE HURON PHYTDPLANKTON  (1983)




OIV  TAXON                                     AUTHORITY







BAC
Hantzschia amphioxys
Me 1 os i ra d i stans
Me 1 os i ra d i stans?
Melosira granulata
Melosira granulata v. angustissima
Me 1 os i ra i s 1 and i ca
Melosira italica subsp. subarctica
Melosira sp.
Navicula acceptata
Navicula atornus
Navicula capitata v. 1 unebur gens i s
Na v icu la c i ncta
Navicula confervacea
Navicula ctfnterta v. biceps
Navicula cr yptocepha 1 a v. veneta
Navicula gottlandica
Nav i cu la med i ccr i s
Nav i cu 1 a m i n i ma
Nav i cu la mura 1 i s
Nav i cu 1 a mura 1 i s?
Navicula mutica
Navicul a perpusi 1 la
Nav i cu la rad i osa
Navicula radiosa v. parva
Navicula raaiosa v. tenella
Navicula seminulum
Nav i cu 1 a s im i 1 i s?
Nav i cu la sp.
Nav i cu la sp . 816
Navicula sp. #18
Navicula submtralis
Navicula subt i 1 i ss i ma
Nav i cu la tant u la
Navicula viridula v. avenacea
Navicula viridula v. rostellata?
Nitzschia ac i cu 1 ar i o i cies
Nitzschia acicularis
Nitzschia acu la
Ni tzsch ia amph ib ia
Nitzschia angustata
Nitzschia angustata v. acuta
Nitzschia confinis
Nitzschia diss ipata
Nitzschia font! co la
Nitzschia frustulum
Nitzschia frustulum v. perpusilla
N i tzschia grac i 1 is
Nitzschia kuetzmgiana
Nitzschia lauerburg ia na
Nitzschia pa 1 ea
Nitzschia paleacea
N i tzsch ia pura
( Ehr . Grun .
(Ehr. Kutz.
(Ehr. Kutz.
(Ehr. Ralfs
0. Mu 1.
0. Mu 1.
0. Mu 1.

Hust.
(Kutz.) Grun.
(Grun. ) Patr .
(Ehr.) Ralfs
Kutz.
(Am.) V.H.
(Kutz. ) Rabh.
Grun .
Krasske
Gr un .
Gr un .
Gr un •
Kutz.
(Kutz.) Grun.
Kutz.
Wa 1 lace
(Breb. ) Cl . 1 Mo I 1 .
Grun.
Krasske



Hust.
Cl.
Hust.
(Breb.) V.H.
(Kutz.) Cl.
Arch, non Hust.
(Kutz.) W. Sm.
Hantz. ex C 1 . £. Gr un .
Grun.
(W. Sm.) Grun.
Grun.
Hust.
(Kutz. ) Grun.
Grun .
(Kutz. 1 Grun.
(Rabh. ) Gr un .
Hantz.
Hi Ise
Hust.
(Kutz.) W. Sm.
Gr un .
Hust.

-------
                                                              252
            SPECIES LIST - LAKE HURON PHYTOPLANKTON (1983)

DIV  TAXON                                     AUTHORITY
BAG
N i tzsch i a
N i tzsch i a
N i tzsch i a
N i tzsch i a
N i tzsch la
Ni tzschia
Ni tzschia
Nitzschia
Opephora
               pus iI la
               recta
               romana
               rosteI lata
               sp.
               sub I i near i s
               subrostrata
               tenu i s
              mar ty i
     Pinnularia tn i cr cstaur on
     Rhizosolenia eriensis
     Rh i zosoI en ja sp.
     Stephanodiscus alpinus
     S tephanod i sc us
     S tephanod i scus
     Stephanod i scus
     Stephanod i scu s
     Ste phanod i sc us
     S tephanod i scus
     S tephanoc i scus
     Stephanod i scus
     Stephanod i scus
     Stephanod i scus
     S tephanod i sc us
     S tephanod i scus
     Stephanod i scus
     Stephanod i scus
     Stephanod i scus
     Stephanod i scus
     Stephanod i scus
     Sur i re I I a ovata
     Surirella ovata v.  salina
     Synedra amphicephaI a v. austrica
             eye Iocum
             de I i cat i ss i ma
             deli cat i ss i ma
             fame i Ii ca?
             f iI if crmi s
             f i I if crmis  v.
             m i ni scuI a
             nana
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
Synedra
Tabe Maria
Tabe I lar i a
Tabe I I ar i a
               alpinus - auxospore
               aIp i nus?
               b inderanus
               b i nder anus?
               hantzsch ii
               minu tus
               n iagarae
               niagarae - auxospore
               sp.
               sp. #03
               sp. #05
               sp. -auxospore
               tenu i s
               tenu is v. #01
               tenu is v. #02
               tenu i s?
               t ran s i I van i cus
           v. angust iss iroa
           ex i I is
             paras i t
             rad i ans
             rumpens
             rumpens
             sp.
             uIna  v .
             uIna  v.
             u I na  v .
     ca
     v.  f ragiIar i o i des
     chaseana
     dan ica
     Iong i ss ima
f erestrata
ferestrata  v. geniculata
fIoccuIo sa
                                (Kutz.) Grun. em. L.-B.
                                Hantz.
                                Grun.
                                Hust.

                                Must.
                                Hust.
                                W. Sir.
                                Her i b.
                                (Ehr .) C I.
                                H.L. Sin.

                                Hust.

                                Hust.
                                (Kutz.) Kr i eg.
                                (Kutz.) Krieg.
                                Grun.
                                Gr un.
                                Ehr .
                               Hust.
Hust.
Pant.
Kutz.
(M. Sm. )  Hust.
(Grun.) Hust.
Brut schy
W. Sm.
Gr un.
Kutz.
Grun .
A. C I .
Gr un.
Heister
W. Sm.
Kutz.
Kutz.
Grun.

Thomas
(Kutz.) V.H.
(H. Sm.1  Br un.
Kutz.
A. Cl.
(Roth) Kutz.

-------
                                                              253
            SPECIES LIST - LAKE HURON PHYTOPLANKTON (1983)

OIV  TAXON                                     AUTHORITY
BAG  Tabellaria flocculosa
     Tabe Maria sp.
     ThaI assjrosira sp.
                         I  ineari s
                                          Koppen
CAT
CHL
Vacua I ar i a
Vacuo I ar ia
                sp .
                sp.?
                    falcatus
                    falcatus v.
                    gelifactum
                    sp. #01
                    sp. #02
                    spiral is
                    stipitatus?
                   plank ton i ca
                   sp.
                   sp.?
                   sp.
                   sp. - ovo i a
                   sp. - sphere
Ank i s trodesmus
Ank i strodesmus
Ank i s trodesmus
Ank i strodesmus
Ank i strodesmus
Ank i strodesmus
Ank i strodesmus
Botryococcus Brauni i
Ch I amydocapsa bacillus
Ch lamydocapsa
Ch I amydocapsa
Ch lamydocapsa
Ch lamydomonas
Ch lamydomonas
Ch I amydomonas
Coelastrum microporum
Cosmar i urn sp .
Cosmar ium sp. »1
Crucigenia irregularis
Crucigenia quadrata
Crucigenia r e ctangu lar i s
D i c ty osphaer i urr. pulchellum
Ech i nosphaere I I a limnetica
Elakatothrix gelatinosa
E I aka to th r i x v i r i di s
Eudor i na el egars
France i a ova I i s
G I oeocyst is  sp.  #3
Go I enk i nia r aa i ata
Green coccoid #02
      coccoid
      coccoid
      coccoid
      cocco i d
      cocco i d
      coccoid
      cocco i d
      cocco i d
K i r chner i e I I a
Lager he i in i a c i I iata
Micractinium pus ill urn
Monoraph i d i urn contortum
Monoraph i d i urn convolution
Monoraph i d ium (rinutum
Monoraph id i urn saxat i I e
                                mirabiI is
     Green
     Green
     Green
     Green
     Green
     Green
     Green
     Green
                   #03
                   - ac i cu lar
                   - bac i I  I if orrn
                   - bic el  Is
                   - fus i form
                   - ova I
                   - sph er e
                   conto r ta
(Corda)  Ralfs
(West £  West) G.S.  West
(Chod.)  Bour r.
                                               (Turner) Lemm.
                                               (Chod.) Kom.-Legn.
                                               Kutz.
                                               (Tei I.) Fott
                                               (M.  I G.S. West) Fott
                                               Nag. in A. Braun
                                               Wi I I e
                                               Hor r en
                                               A. Braun
                                               Mood.
                                               G • M •  S m •
                                               Mi I I e
                                               (Snow) Pr intz
                                               Ehr .
                                               (France) Lemm.

                                               (Chod.) UiI Ie
                                               (Schnrid.) Bohiti,
                                               (Lagerh.) Choa.
                                               Freseni us
                                               (Thur.) Kom.-Legn.
                                               (Corda) Kom.-Legn.
                                               (Nag.) Kom.-Legn.
                                               Kom.-Legn.

-------
                                                             254
            SPECIES LIST - LAKE HURON PHYTOPLANKTON (1983)

DIV  TAXON                                     AUTHORITY
CHL
CHR
Monoraphidium setiformae
Mougeotia sp.
Oocyst is  sp.
Oocystis  sp. »1
Oocystis Borgei
Oocystis crassa
Oocystis lacustris
Oocyst is mar s en i i
Oocystis parva
Oocystis pus i I la
Oocyst is so Ii tar ia
Pyramidomonas sp.
Scenedesmus abcndans
Scenedesmus
Scenedesirus
Scenedesmus
Scenedesmus
Scenedesmus
Scenedesmus
Scenedesrrus
Scenedesmus
            dent i culatus
            eccrn i s
            secu r if or mi s
            secur if oriri s ?
            serratu s
            sp.
            subsp i cat us
            ve I itar i s
SphaereI Iocystis lateralis
Sphaerocystis  schroeteri
St i chococcus sp.
Synechococcus  sp.
TetrachI ore I I a alternans
Tetraedron minimum
Treubaria planktonica
Treubaria planktonica?
Treubaria setigera

B i tr i ch i a chodati i
Chrysolykos planktonicus
Chrysolykos sktjae
Chrysolykos sp.
ChrysosphaereI I a longispina
Dinobryon - statospore
Dinobryon acuiri i na turn
Dinobryon bavaricum
D i nobryon bor ge i
Dinobryon cylindricum
Dinobryon cylindricum
Dinobryon divergens
Dinobryon divergens -
Dinobryon eurystoma
Dinobryon sertularia
Dinobryor sertularia
D inobryon soc iaIe
D i nobryon soc i a Ie v .
D inobryon stokes i i  v
Dinobryor utriculus
Haptophyte sp.
                           v . aIp i num

                           statospor es


                          v. pro tuberans

                          arrer i canum
                         • ep i pIanktoni cum
                         v . tabeI Iar iae
                                                (Nyg.) Kom.-Legn.
                                               Snow
                                               Mi ttr .  in W i ttr . i Nord,
                                               Chod.
                                               Lemm.
                                               West £ West
                                               Hansg.
                                               Wittr.  in Wi ttr. £ Nord.

                                               (Kirch.) Chod.
                                               Lagerh.
                                               (Ralfs) Chod.
                                               Playf.
                                               Playf.
                                               (Cor da) Boh I m

                                               Chod.
                                               Kom.
                                               Fott t Novak.
                                               Choa.
                                          (G.M. Smith) Kors.
                                          (A. Braun) Hansg.
                                          (G.M. Sen.) Korch.
                                          (G.M. Sro.) Korch.
                                          (Arch.) G.M. Sm.

                                          (Rev.) Chod.
                                          Mack.
                                          (Nauw. ) Bour r.

                                          Laut. em. Nich.

                                          Rutt.
                                          Imhof
                                          Lemm.
                                          Imhof
                                          (Imhof) Bachm.
                                          Imhof

                                          (Stokes) Lemm.
                                          Ehr.
                                          (Lemm. ) Kr i eg.
                                          Ehr .
                                          (Brunnth.) Bachm.
                                          Sku ja
                                          Lemm.

-------
                                                             255
            SPECIES LIST - LAKE HURON PHYTCjPLANKTON (1983)

DIV  TAXON                                     AUTHORITY
CHR
COL
CRY
Kephyr i on
Kephyr i on
Kephyr i on
Kephyr i on
Kephyr i on
Ma I Iomonas
Ma I Iomonas sp•
Mai Iomonas sp.
Ochromonas sp.
Ochromonas sp.
Ochromonas sp.
Paraphysomona s
Paraphysoroonas
Pseudokephry i on
Pseudokephyr i on
Pseudokephyr i on
Pseudokephyr i on
Pseudokephyr i on
Uni dent i f i ed
Un i dent if i ed
cupu I i for mae
sp. #1  -Pseudokephyrion
sp. #2
sp. #3
sp i ra I e
 sp.
     #1
     #3
                                       entzi i
                    - ovoi d
                    - sphere
                    sp.
                    sp.?
                     entzi i
                     con icutr
                     I at urn
                     IT i I lerense
                     sp. #1
                  ccccoias
                  I or i ca te - ovo i d
Unidentified  loricate - sphere

Bicoeca campanulata
6 i coeca crystal Iina
Bicoeca mitra  v.  suecica
B i coeca socia I i s
B i coeca sp.
Bicoeca sp. 804
Bicoeca tub if or mis
Colorless flagellates
Monos i ga ovata
Monos i gna ova I i s
Salpingoeca amphorae
Salpingoeca gracilis
Stylotheca aurea
Chroomonas
Chroomonas
Chroomonas
Cryptomonas
Cr yptomonas
Cr yptomonas
Cryptomonas
Cryptomonas
Cr yp toironas
Cryptomonas
Cryptomonas
Cr yptomonas
Cr yptomonas
Cryptomonas
Cryptomonas
Cryptomonas
 a c f t a
 cabdata
 norstedti i
  - cyst
  brev i s
  caudata
  er csa
  ercsa v.  reflexa
  trar sson i i
  obovata ?
  ovata
  parapyrenoidifera
  phaseoI us
  phaseolus ?
  pus i I la
  pyreno i ci fer a
                                               Conr ,
                                               (Lack.) Conr.
                                Conr .
                                (SchilI.)
                                (Sch ill.)
                                Ni ch.
          Schum.
          Schum.
                                (Lack.) Bourr. em. Skuja
                                Sku ja
                                Skuja
                                Lauter b.
Skuja

Kent
Kent
Kent
Clark
(Bachm.) Boloch.

Uterm.
Ge it.
Hansg.

Schi II .
SchiI I.
Ehr .
Marss.
Skuja
Skuja
Ehr.
Skuja
Skuja
Skuja
Bachm.
Geit I.

-------
                                                             256
            SPECIES LIST - LAKE HURON PHYTOPLANKTON (1983)

OIV  TAXON                                     AUTHORITY
CRY
CYA
BUG
Cr yptomonas
Cryptomonas
Cr yptomona s
Cryptomonas
Cr yptomonas
Rhodomonas
Rhodomonas
                 refIexa
                 rest rat if orm i s
                 sp.
                 tenu i s
                 tetrapyrenoidiosa?
                Iacustr i s
                I ens
     Rhodomonas minuta
     Rhodomonas minuta v.
     Unidentified coccoid
                     nannop lankti ca
     Anabaena circinalis
     Anabaena sp.
     Anacyst is mar i na
     Anacystis montana v. minor
     Anacyst is therira I is
     Coccochloris elabans
     Coccochloris peniocystis
     CoeIosphaeri im  Naegel ianum
     Gomphosphaeria   lacustris
     Osc iI Iator ia
     Osc iI Iato r i a
     Osc iI Iator ia
     Osc iI I ator i a
             Iimnet i ca
             rr i n ima
             subbrev i s
             tenu i s
Euglena sp.
Phacus sp.
TracheIomonas hispiaa
TracheIomonas sp.
PYR  Amphidinium sp.
     Ceratium hirundinella
     Gymnod i n i urn sp.
     Gymnod i n i urn
     Gymnod i n i urn
     Gymnod i n i urn
     Gyirnod in i urn
     Peridinium inconspi
     Per i d i n i urn sp.
     Peridinium sp. #02
            sp.
            sp.
            sp.
            sp.
#1
#2
#3
#5
UNI  Un identi f i ed
     Un i dent i fi ed
     Un identi f ied
             flagel late #01
             f I age Mate - ovo i d
             flagellate - spherical
                          Sku ja
                          Sku ja

                          Pasch.
                          Skuja
                          Pasch.
                          Pasch.
                          Skuja
                          Skuja
£ Rutt.
£ Rutt.
                          Rabenhorst

                          (Hansg .) Or . £ Da i ly
                          Or. £ Da i I y
                          (Menegh . )  Dr . £ Da iIy
                          Dr. £ Da i ly
                          (Kutz.) Dr. £ Da ily
                          Unger
                          Chod.
                          Lemm.
                          GickIn.
                          Schm id.
                          C.A. Ag.
                                               (Perty ) Ste in em. DefI
                                          (O.F.MuI  I.) Schrank
                                          Lemm,

-------
                                          257'
        Table C

      Phytoplankton
Species List:  Lake Erie

-------
                                                              258 .
            SPECIES LIST - LAKE ERIE PHYTOPLANKTON (1983)

UIV  TAXON                                     AUTHORITY
SAC
     Achnanthes
     Achnanthes
     Ac hnanthes
     Achnanthes
     Achnanthes
     Achnanthes
     Achnanthes
     Achnanthes
     Achnanthes
     Achnanthes
     Achnanthes
     Achnanthes
     Achnanthes
     Achnanthes
     Achnanthes
     Achnanthes
     Act i nocycI us
     Act i nocycI us
b i asoIet t iana
b i oret i
cI eve i
c I eve i  v. rostrata
consp i cua
ex igua
haucki ana
lanceolata v. dubia
I emirerirann t
I inear is
I inear is f o.
m i cr ocep ha la
mi nut i ss ima
sp.
sp.?
sub laev i s
  rorman i i  f ,
  sp.
cur ta
 subsa I sa
            roinuta
             euglypta
             lineata
     Amphora oval is v. affins
     Amphora ovalis v. pediculius
     Amphora perpus ilia
     Amphora sp.
     Amphora tenuistriata
     Anomoeoneis  vitrea
     Asterionella forir
-------
                                                              259
            SPECIES LIST - LAKE ERIE PNYTOPLANKTON <1983)

DIV  TAXON                                     AUTHORITY
bAC  CycIotel I a
     Cyclotel la
     CycIotel la
     CycIotel la
     Cyc lotel la
     CyclotelI a
     Cymatopleura solea
     CyroatopIeura solea
     CymbeI  la at f i ni s
     CymbeI  I a
     CymbeI  la
     CymbeI  la
     CymbeI  la
     Cymbe I  I a
           operculata
           pseuaosteI I i gera
           sp.
           sp. SI
           stelIige ra
           wo Itereck i
                   v. apiculata
         m i crccepha la
         minuta
         mi nuts v. si lesiaca
         prostrata v. auerswaldii
         pusi I la
CymbeI la sp .
Denticula tenuis v. crassula
0 iatoma anceps
Oiatoma tenue v. el on gat urn
Diatoma vulgare
0ip I one is ocu I ata
Entomoneis ornata
Fragilaria brevistriata
Fragilaria brevistriata v. inflata
Fragilaria capucina
Fragilaria construens
Fragilaria construens v. minuta
Fragilaria construens v. pumila
Fragilaria construens v. venter
Fragilaria crotonensi s
Fragilaria inter/red ia v. fallax
Fragilaria leptostauron
Fragilaria leptostauron v. dubia
Fragilaria nitzschicides
Frag i far i a pi nnata
Fragilaria pinnata v. lancettula
Fragilaria pinnata v. pinnata
Fragilaria sp.
Fragilaria vaucheriae
Gomphoneira clevei
Gomphonema dichotomum
Gorophonema parvulum
Gomphonema sp.
Gomphonema tergestinum
Gyrosigma attenuatuiri
Gyrosigma scictense
Melosira agassizii v. malayensis
he I os i ra di stans
Melosira distans v.  limnetica
Melosira granulata
Melosira granulata v. angustissima
Melosira granulata?
Melosira i sI and i ca
(Ag.) Kutz.
Must.
(CI. £. Grun.) V.H.
Hust.
IBreb. i. Godey) W. Sm.
(W. Sm.) kalfs
Kutz.
Grun.
Hi Ise
(Sleisch) Reim.
(Rabh,)  Reim.
Grun.

(Nag.) k. t G.S. West.
(Ehr.) Kirchn.
Lyngb.
Bory
(Breb.)  CI.
(J.W. Bai I .) Re i m.
Grun.
(Pant.)  Hust.
Oesm.
(Ehr.) Grun.
Temp. £  Per.
Grun .
(Ehr.) Grun.
Ki tton
(Grun.)  Stoerm. £ Yang
(Ehr.) Hust.
(Grun.)  Hust.
Grun •
Ehr.
(Schum.) Hust.
                                               (Kutz.) Peters.
                                               Fr icKe
                                               Kutz.
                                               Kutz .

                                               (Grun. ) Fr i eke
                                               (Kutz.) Rabh.
                                               (SulIiv. L Wormley) CI .
                                               Ostenf.
                                               (Ehr.) Kutz.
                                               0. MulI.
                                               (Ehr.) kalfs
                                               0. Mul I.
                                               (Ehr.) Ralfs
                                               0. MulI.

-------
                                                           260
            SPECIES LIST - LAKE ERIE PHYTOPLANKTON 11983)

DIV  TAXON                                     AUTHORITY
                          hurgari ca
                          Iuneburgens i s
                               veneta
BAG  Melosira italica subsp. subarctica
     Melosira sp.
     Navicula acceptata
     Nav i cuI a angI i ca
     Nav i cuI a capitate
     Navicula capitata v,
     Nav i cula cap i tata v <
     Navicula c i nc ta
     Navicula cocconeiformis
     Navicula cryptocephaI a
     Navicula cryptocephala v,
     Navicula ex i gua
     Navicula exigua v. capitata
     Navicula lanceolata
     Nav i cuI a meni scuI us
     Navicula menisculus v. upsaliensis
     Nav i cuI a minima
     Navicula pseudoscutiformis
     Nav i cuI a pupu la
     Navicula radiosa v. tenella
     Navicula salinarurr v.  intermedia
     Navicula seminulcides
     Navicula seminulum
     Nav i cuI a sp.
     Nav icula stroen i i
     Navicula terminate
     Navicula tripunctata
     Navicula viridula v. rostellata
     Navicula vitabunda
     Nav i cula zanon i
     Ne id i um af f i ne
     Ni tzsch13
     Ni tzsch ta
     N i tzsch ia
     N i tzsch i a
     Ni tzsch ia
     Ni tzsch ia
     N i tzsch ia
     N 11 z s c h i a
     N i tzsch ia
     N i tzsch i a
     Ni tzsch ia
     N i tzsch ia
     Ni tzschia
     N i tzsch ia
     N i tzsch ia
     Ni tzschia
     N i tzsch ia
     N i tzsch ia
     N i tzsch ia
     Nitzschia
     N i tzsch i a
               ac i cuIar i o ides
               ac i cular i s
               acict'lari s?
               acu I a
               araphi D ia
               angustata
               angustata  v. acuta
               ap i cuIata
               ar chba I di i
               closter ium
               conf in i s
               d i ssipata
               d i ss i pata  v. media
               f ont i co la
               fr ustuI urn
               gancershe im i ens i s
               grac i I i s
               hantzschiana
               i nconsp i c ua
               i ntermed i a
               Kuetz i ng i ana
0. MulI.

Hust.
Rat f s
Ehr .
(Grun.)  Ross
(Grun.)  Pat r.
(Ehr.) Ralfs
Greg .
Kutz .
(Kutz.)  Rabh.
Greg . ex Grun•
Patr .
(Ag.) Kutz.
Schum.
(Grun.)  Grun.
Grun .
Hust.
Kutz.
(Breb.)  Cl. £ Mo I I.
(Grun.)  CI.
Hust.
Gr un.

Hust.
Hust.
(O.F.MulI.) Bory
(Kutz.)  CI.
Hust .
Hust.
Pf i tz.
Arch, non Hust.
(Kutz.)  W. Sni.
(Kutz.)  W. Snu
Hantz. ex Cl. 6 Grun,
Grun .
(W. Sm.) Grun.
Gr un .
(Greb.)  Grun.
L.-B.
(Ehr.) W. Sm.
Hust.
(Kutz.)  Grun.
(Hantz.) Grun.
Grun.
(Kutz.)  Grun.
KrassKe
Hantz.
Rabh.
Grun .
Hantz.
Hi Ise

-------
                                                                261
            SPECIES LIST - LAKE ERIE PHYTOPLANKTON (1983)

DIV  TAXON                                     AUTHORITY
BAG
Ni tzsch ia
Ni tzschia
Ni tzsch ia
N i tzsch i a
Ni tzsch ia
N i tzsch ia
Ni tzschia
Nitzschia
Ni tzschia
Ni tzsch ia
Nitzsch ia
N i tzsch ia
N i tzsch ia
Nitzsch ia
N i tzsch ia
N i tzsch ia
N i tzsch ia
N i tzsch i a
Nitzschia
Ni tzsch ia
Ni tzschia
N i tzsch ia
Ni tzschia
Nitzsch ia
Rh izosoI en ia
Rh i zoso ten ia
                        debi I is
                        tenu i rostr i
kuetz irtgi oi des?
lauerburgiana
I  inear i s
pa I ea
pa Iea  v.
pa Iea  v.
pa I eacea
puni la
pur a
pus i I la
r  ecta
romana
r  osteIlata
soc iab iIi s
sp.
sp i c LIc i des
subac icularis
sub I inear is
tenu is
tr op i ca
trybIi one I  I a
trybli one I  I a
trybIi one I  I a
trybli one I  I a
   er iens is
   Icngiseta
                            v.
                            v.
                            v.
debiI is
v i ctor iae
v i ctor i ae?
     Rhi zosoI en ia sp.
     Skeletonema pctairos
     Stauroneis kriegeri
     Stephanodiscus alpinus
     Stephanodi scus
     Stephanod i scus
     Stephanoci scus
     Stephanod i scus
     Stephanod i scus
     Stephanod i scus
     Stephanod i scus
     Stephanod i scus
     Stephanodi scus
     Stephanod i scus
     Stephanod i scus
     Stephanod i scus
     Stephanod i scus
     Stephanodi scus
     Stephanod i scus
     Stephanod i scus
     Stephanod i scus
     S tephanod i scus
               alpinus - auxospore
               a Ip i nus?
               binder anus
               hantzsch i i
               m inu tu s
               minutus - auxospore
               n iagar ae
                        - auxospore
                        v. magnif ica
               sp.
               sp.
               sp.
               sp.
     Sur i r eI I a
     Surirella
     Surirella
     Sur i r el I a
     n iagar ae
     n iagar ae
     sp.
         #03
         #04
         #07
         "auxospore
     tenu i s
     tenu i s v. #01
     tenu is v• #02
     tenu i s?
b i rostrata
o vata
ovata v. pinnata
ovata v. sa I ina
Hust.
W. Sir.
(KutZ.) W. Sm.
(Kutz.) Grun.
Gr un.
Grun.
Hust.
Hust.
(Kutz.) Grun. em. L.-6.
Hantz.
Grun.
Hust.
Hust.

Hust.
Hust.
Hust.
W. Sir.
Hust.

(Arnott) A. Mayer
Grun.
Grun .
H.L. Sm.
Zach.

(Weber) Hasle £ Evens.
Patr.
Hust.

Hust.
(Kutz.) Krieg.
Grun.
Gr un.

Ehr.

Fr icke
                                          Hust.
                                          Hust.
                                          Hust.
                                          Kutz.
                                          (W. Sm.) Hus.t.
                                          (W. Sm.) Hust.

-------
                                                                262
            SPECIES LIST - LAKE ERIE PHYTOPLANKTON  (1983)

LlIV  TAXON                                      AUTHORITY
                   austr i ca
                    angust iss ima
BAG  Sur i re I I a sp.
     Sur i r e I I a turgida
     Synedra  acus?
             amph i cephala v.
             deli  cat issi ma
             deIi  cat i ss i ma v.
             f i I i  f o r ITI i s
             filiforiris v. exilis
             rr i ni  scu I a
             paras 11 i ca
             u I na v. Iong i ss i roa
                f  enestr ata
                ferestrata v. genicutata
                fIoccuIo sa
                flocculosa v. linearis
                sp.
     Synedra
     Synedra
     Synedra
     Synedra
     Synedra
     Synedra
     Synedra
     Synedra
     Tabe
     Tabe
     Tabe
     Tabe
     Tabe
I ar i a
lar i a
I ar i a
I ar i a
I ar ia
     ThaI assiosira fluviatilts

CAT  Vacuolar i a sp .

CHL  Actinastrum gracilimum
     Anh i strodesmus sp. feO*!
     AnKyra juday i
     Car ter ia  sp.
     Car ter ia  sp. -cvo i d
     Carteria  sp. -sphere
     ChIamydocapsa planktonica
     ChIamydocapsa
     ChIamydomonas
     Chlamydomonas
     Ch tamydomonas
     ChI orogon ium
         sp.
         sp.
         sp.  - ovoid
         sp .  - spher e
        tr in imu m
     ChIorogoniurn sp.
     Closterium aciculare
     Closter ium
     Clo ster ium
     Coelastrurn
     CoeIastrum
     CoeIastrum
     Co smar i urn
      par vu I urn
      sp.
      canbri cure.
      m i cr opor urn
      sp.
     sp.
     Crucigenia irregularis
     Crucigenia quadrata
     Crucigenia rectangu laris
     Crucigenia tetrapedia
     D i ct yosphae r i urr ehr enber g ianum
     D i ct yosphaer i uir, pulchellum
     Elakatothrix  gelatinosa
     Elakatothrix  viridis
     Eudor i na eIegars
     France ia ova Ii s
     Golenkinia radiata
     Green F i lament
W. Sm.
Kutz.
(Grun.) Hust.
W. Sm.
Gr un.
Grun.
A. Cl.
Grun.
W. Sm.
(W. Sm.)  Br un.
Kutz.
A. Cl .
(Roth) Kutz.
Koppen

Hust.
G.M. Smith

(G.M. Sm.) Fott



(W. £ G.S. West) Fott




Playf.

T. West
Nag.

Arch.
Nag. in A. Braun
                                     Mi I I e
                                     Mor ren
                                     A.  Braun
                                     (Kirch.) W. £ G.S. West
                                     Nag.
                                     Wood.
                                     Wi I le
                                     (Snow ) Pr i ntz
                                     Ehr .
                                     (France) Lemm.
                                     (Chod.) WiIle

-------
                                                             263
            SPECIES LIST - LAKE ERIE PHYTOPLANKTDN (1983)

DIV  TAXON                                     AUTHORITY
                   #04
                   - aci cu I ar
                   - bac i I  I iform
                   - bicel  Is
                   - fusiform bicetls
                   - ova I
                   - ovo i d
                   - sph er  e
CHL  Gr een cocco i d
     Green coccoid
     Gr een coccoid
     Gr een coccoid
     Gr een coccoid
     Gr een cocco i d
     Green cocco i d
     Green coccoid
     Green flagellate - sphere
     KirchnerieI I a contorta
     K i rchner i eI la obesa
     Lagerheimia balatonica
                 c iIiata
                 geneven si s
                 Iong i se ta v. major
                 quadr i seta
                 sp.
                 subsaIsa
     Lager he im i a
     Lager he i m ia
     Lager he i m i a
     Lagerhe i ir i a
     Lagerhe i ir ia
     Liger heimia
     Lobomonas sp . ?
     Micractiniuro p u s i I I urn
     Monoraphidiurn contortum
     Monorapfl i d i urn griffitrsii
     Monoraphidiurn irregulare
     Monoraphidium nirutum
     Mougeotia sp.
     Nephrocytium Agardhianum
     Nephrocytiurn limneticum
     Nephrocytium liirneticura?
     Oedogonium  sp.
     Oocyst i s
     Oocyst is
     Oocyst is
     Oocyst is
     Oocyst i s
     Oocyst i s
     Oocyst i s
     Oocyst is
     Oocyst i s
     Oocyst I s
     Oocyst i s
     Oocyst i s
     Pandor ina
                   *1
 sp,
 sp,
 sp.?
borge i
crassa
el Ii pt i ca
lacustri s
ma r sen ii
par va
pus i I I a
so Ii tar ia
submar i na
 mor unr ?
                        v. minor
     Paradoxia multiseta
     Pediastrum boryanum
     Ped i astrum
     Ped i astrutn
     Pediastrum
     Ped i as trurn
     Pediastrum
     S c e n e d e s m u s
                dup lex v .
                dup lex v .
                simplex
                simplex v.
                sp.
                 abtndan s
            c lathratum
            ret i cuIatum

             duodenari um
     Scenedesmus
     Scenedesmus
                 actir, inatus
                 ar cuatus
                                 (Schm id.) Bohlm
                                 (W. fcest) Schmidle
                                 (Scherf f. in KoI) Hind.
                                 (Lagerh.) Choo.
                                 (Chod.) Chod.
                                 G.M, Sm •
                                 (Lerom.) G.M. Sm.

                                 Lemm.

                                 Fr esen i us
                                 (Thur.) Kom.-Legn.
                                 (Berkel.) Kom.-Legn.
                                 (G.M.  Sm.) Kom.-Legn.
                                 (Nag.) Kom.-Legn.

                                 Nag.
                                 (G.M.  Sm.) G.M. Sm.
                                 (G.M.  Sm.) G.M. Sm.
Snow
Wittr . in Mi ttr. £ Nord.
M. West
Chod.
Lemm.
West £ West
Hansg.
Wi ttr. in Wi ttr. £ Nord.
Lagerh.
(Muell.)  Bory
Swir.
(Turp.) Menegh.
(A. Braun) Lagerh.
Lagerh.
(Meyen) Lemm.
(Bail.) Rabh.

(Kirch.)  Chod.
(Lagerh.) Chod.
Lemm.

-------
                                                              264 ,
            SPECIES LIST - LAKE ERIE PHYTOPLANKTON (1983)

UIV  TAXON                                     AUTHORITY
CHL  Scenedesmus arrrattis
     Scenedesmus bicaudatus
     Scenedesnr.us carinatus
     Scenedesmus denticulatus
     Scenedesmus eccrnis
     Scenedesmus intermeaius
     Scenedesmus interrreaius v. bicaudatus
     Sceneaesrrus quadricauda
     Scenedestrus s ecu r if or mi s
     Scenedesmus sp.
     Scenedesmus spinosus
     Scenedesmus spinosus?
     Schroederia setigera
     SphaereI Iocystis lateralis
     SphaereI I ops i s sp.
     Sphaerocystis  schroeteri
     Staurastrum paradoxum
     Staurastrurn sp•
     St i chococcus sp.
     Tetraedron  caudatum
     Tetraedr on m i n imum
     Tetraedron mut i cum
     Tetraedron  regulare v. incus
     Tetraspora  lacustris
     Tetrastrum heteracanthum
     Tetrastrum staurogeniaeforme
     Treubaria planktonica
CHR
     Treubar ia
     Treubaria
          set
          sp.
gera
B i tr i chia chodati i
Chrysolykos planktonicus
Chrysolykos skcjae
ChrysosphaereI I a longispina
Dinobryon acuirinatum
Dinobryon bavaricum
Dinobryon cylinaricum
Dinobryon divergens
Dinobryon sertularia
Dinobryon sociale v. arrericanum
0 i nobryon sp.
Dinobryon stokesii v. epipIanktonicum
Dinobryor utriculus v. tabellariae
Haptophyte sp.
Kephyrion cupuliformae
Kephyrion sp. #1 -Pseudokephyrion entzi
Kephyr i on sp. #2
Kephyrion sp. #3
Ma I I omonas sp.
Ochromonas sp.
Qchromonas sp. - ovoid
Paraphysomonas sp.?
                                          (Chod.) G.M. Sm.
                                          (Hansg.) Chod.
                                          (Lernm. ) Chod.
                                          Lagerh.
                                          (RaIfs ) Choa.
                                          Chod.
                                          Hor tob.
                                          (Turp. ) Breb.
                                          PIayf.

                                          Chod.
                                          Chod.
                                          (Schroed.) Lemm.
                                          Fott  t. Novak.

                                          Chod.
                                          Meyen
                                          (Corda) Hansg.
                                          (A. Br aun) Hansg.
                                          ( A . Br aun ) Hansg.
                                          Te i1ung
                                          Lemm.
                                          (Nor dst.) Chod.
                                          ( Schroed.) Lemm.
                                          (G.M. Srr.) Korch.
                                          (Arch.) G.M. Sm.
                            (Rev.) Chod.
                            Mack .
                            ( NiauM . )  Bour r .
                            Laut. em. Mich.
                            Rutt.
                            Imhof
                            Imhof
                            Imhof
                            Ehr.
                            (Brunnth.) Bachm.

                            Skuja
                            Lemm.

                            Conr .

-------
                                                               265 .
            SPECIES LIST - LAKE ERIE PHYTOPLANKTDN (1983)

DIV  TAXON                                     AUTHORITY
CHR  Pseudokephyr ion millerense
     Pseudokephyrion sp. #1
     Pseudotetraedron neglectum
     Unidentified coccoids
     Unidentified flagellate
     Unidentified loricate - ovoid
     Unidentified loricate - sphere

COL  B coeca campanulata
     B coeca crystaI Iina
     B coeca sp.
     B coeca sp. *01
     B coeca sp. #G4
     B coeca sp. #05
     B coeca tubifcrir.is
     Codonosiga sp.
     Colorless flagellates
     Colorless flagellates - colonial
     Monosi ga ovata
     Salpingoeca amphorae
     Salplngoeca gracilis
     Stelexmonas dichotoma
     Stylotheca aurea

CRY  Chroomonas acuta
     Chroomonas norstedtii
     Cryptomonas -  cyst
     Cryptomonas catoata
     Cryptomonas curvata
     Cryptomonas curvata?
     Cryptomonas ercsa
     Cryptomonas ercsa v. reflexa
     Cryptomonas rrarssonii
     Cryptomonas marssonii v.?
     Cryptomonas ovata
     Cryptomonas phaseolus
     Cryptomonas pyrenoioifera
     Cryptomonas reflexa
     Cryptomonas rostrat iformis
     Cryptomonas rostrat iformis?
     Cryptomonas sp.
     Rhodomonas lens
     Rhodomonas minuta
     Rhodomonas minuta v. nannoplanktica

CYA  Agmenellum quadrupl icatum
     Anabaena sp.
     Anabaena spircides
     Anacyst i s mar ina
     Anacystis montana v. minor
     Anacystis therrralis
     Anacystis theriralis f. major
                                               Nich.

                                               Pasch.
                                               (Lack.) Bourr. em. Skuja
                                               Skuja
                                               Skuja
                                               Kent
                                               Kent
                                               Clark
                                               Lack .
                                               (6achm.> Bo I och.

                                               Uterm.
                                               Hansg.

                                               Schi I I .
                                               Ehr .
                                               Ehr .
                                               Ehr .
                                               Marss.
                                               Skuja
                                               Skuja
                                               Ehr .
                                               Skuja
                                               Geit I .
                                               Skuja
                                               Skuja
                                               Skuja

                                               Pasch. fc Rutt.
                                               Skuja
                                               Skuja

                                               (rtenegh. ) Breb.

                                               Kleb.
                                               (Hansg.) Dr.  i. Dai ly
                                               Dr. £ Da i ly
                                               (Menegh. ) Or. i Da i  I y
                                               (Lagerh. I Dr. £ Da iIy

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                                                             266
            SPECIES LIST - LAKE ERIE PHYTOPLANKTON  11983)

DIV  TAXON                                     AUTHORITY
CYA
EUG
PYR
Aphanizomenon flos-aquae
Coccochlorts elabans
Coccochloris peniocystis
CoeIosphaeriurn dubiurn
CoeIosphaeriurn naegelianum
Gomphosphaeria lacustris
Merismopedia tenuissima
     Osc iI lator ia
     Osc iI I atoria
     OsciI Iator ia
     Osc iI I ator i a
             I irnnet ica
             subbrev i s
             tenu is
             tenu i s ?
Euglena sp.
TracheIomonas sp.

Amphidinium sp.
Ceratium hirundinella
Ceratium hirundinella - cyst
Gymnod in i urn sp.
Gymnodinium sp. H2
Gymnodiniurn sp. S3
Peridinium acicu I iferurn
Peridinium acicu I iferurn?
Peridinium inconspicuum
Per i d i n i urn sp.
UNI  Unidentified
     Unidentif ied
     Unidentif i ed
             flagel late #01
             fI age I late - ovoid
             flagellate - spherical
(L.) Ralfs
Dr. £ Da i ly
(Kutz. )  Dr. £ Da i ly
Grun. in Rabh.
Unge r
Chod.
Lemm.
Lemm.
Schm id.
C.A. Ag.
C.A. Ag.
                                               (O.F .Mul I .) Schrank
                                               (O.F.MulI.) Schrank
                                               Lemm.
                                               Lemm.
                                               Lemm .

-------
                                                    267
                Table  D




Zooplankton Species List:  Lake Michigan

-------
                                                           268
 DIVISION
GREAT LAKES ZODPLANKTON SPECIES LIST
           LAKE MICHIGAN
               (1983)

          TAXON
 Ca lano i da
C I adocera
Copepoda

CycIopo i da
Harpact i co i da

Mys i aacea

Rot i fera
                       Ca lano i d  -
                       Di aptomus
                       D i ap tomus
                       Di aptomus
                       0i aptomus
                       D j aptomus
                       Ep i schura
                       Eu ry temora
                     copepodi te
                    ash land i
                    mi nutus
                    oregonensi s
                    sici I is
                    si c iIo i des
                    lacustr i s
                     aff ini s
          Limnocalanus  macrurus
          Senecella  calanoides

          A Iona af f i ni s
          Bosmina  longirostris
          Camptocercus  rectirostris
          Ceriodaphnia  lacustris
          Chydori dae
          Chydorus  sphaericus
          Daphnia  catawba
          Caphnia  dubia
          Daphnia  galaeta  mendota
          Daphnia  immatures
          Daphnia  longiremis
          Daphnia  middendorffiana
          Daphnia  puIi car i a
          Daphnia  retrorurva
          Daphnia  schod leti
          Daphnia  sp.
          Diaphanosoma  I euchtenbergianum
          Eubosmina coregoni
          Eurycercus  lamellatus
          Holopedium  gtbberum
          Ilyocryptus  spinifer
          Leptodora k i ndt i i
          Polyphemus  pediculus

          Copepoda NaupIi i

          Cyclopoid - copepodite
          Cyclops bicuspidatus  thomasi
          Eucyclops prionophorus
          Mesocyclops edax
          Tropocyclops  prasinus mexicanus

          Harpact i co i da

          My si s r el i eta

          Ascomorpha sp.
          Asplanchna priodonta
          Brachionus quadridentatus

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                                                         269
            GREAT LAKES ZOOPLANKTON SPECIES LIST
                       LAKE MICHIGAN
                           (19B3)
DIVISION              TAXON
Rotifera              Cephaloflella sp.
                      Co I Iotheca sp.
                      Conoch i Ic i des sp•
                      Conochi IDS unicornis
                      Encentrum sp.
                      Euchlanis sp.
                      F i Ii na Iong i seta
                      Gastropus sty Ii fer
                      Kellicottia longispina
                      Keratella cochlearis
                      Keratella crassa
                      Kerate1 la ear Ii nae
                      Ke rate I I a hi emaI i s
                      Keratella quadrata
                      Lecane tenuiseta
                      Monostyla sp.
                      Notholca acuminata
                      No tholea fo I iacea
                      Notholca laurentiae
                      Notholca squamula
                      Notholca str iata
                      PIoesoma sp.
                      Polyarthra dolichoptera
                      Po lyar thra major
                      Po lyar thra remata
                      Polyarthra vulgaris
                      Synchaeta sp.
                      Trichocerca cylindrica
                      Trichocerca multicrinis
                      Tr ichocerca sp.

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                                                  270.
              Table E




Zooplankton Species List:  Lake Huron

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                                                         271
DIVISION


Ca lanoi da
            GREAT LAKES ZOOPLANKTON SPECIES LIST
                         LAKE HURON
                           (1983)

                      TAXON
                      Catanoid - copepodite
                      Diaptomus ashlandi
                      Diaptomus minutus
                      Diaptomus oregonensis
                      Di aptomus s i c i I i s
                      Diaptomus siciloides
                      Ep ischura lacustr i s
                      Limnocalanus  macrurus
                      Senecella calanoides
Cladocera
Copepoda

Cyclopoida
Mysidacea

kotifera
                      Bosmina
                      Daphnia
                      Daphnia
                      Daphnia
                      Daphnia
                      Daphnia
                      Daphnia
                      Da phn ia
                      Di aphanosoma
                      Diaphanosoma
                              longirostris
                              catawba
                              dubia
                              galaeta mendota
                              pu I i car i a
                              retrocurva
                              schod ler i
                              sp •
                                   I euc htenber g ianum
                                   sp .
                      Eubosmina coregoni
                      Holopedium giboerum
                      Leptodora K i ndt i i
                      Polyphemus pediculus
                      Si 03. crystal I ina

                      Copepoda Nauplii

                      Cyclopoid - copepodite
                      Cyclops bicuspidatus thomasf
                      Cy c I ops vernal is
                      Mesocyclops edax
                      Tropocyclops  prasinus mexicanus

                      Mysis relicta

                      Ascomorpha sp.
                      Asplanchna priodonta
                      Cephal ode I la  sp.
                      Co 1 1 otheca sp.
                      Conochilus unicornis
                      Eu ch Ian i s sp.
                      F i I i na  I ong i seta
                      Gastropus sp.
                      Cast ropus sty I i f er
                      Kellicottia longispina
                      Keratella cochlearis
                      Keratella cochlearis hispida
                      Ke rate I la crassa
                      Ke rate I la ear I inae

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                                                         272
            GREAT LAKES ZOOPLANKTON SPECIES LIST
                         LAKE HURON
                           (1983)
DIVISION              TAXON
Rotifera              Keratella hiemalis
                      Keratella quadrata
                      MonostyI a Iunar i s
                      No tho I ca foil acea
                      Notholca laurentiae
                      Notholca squamula
                      PIoesoma sp•
                      Polyarthra dolichoptera
                      Po lyar thra major
                      Polyarthra remata
                      PoIyar thra vuI gar i s
                      Rotifer - soft  body
                      Synchaeta sp.
                      Trichocerca  cylindrica
                      Tr ichocerca  multicrinis
                      Tr ichocerca  sp.
                      Trichotria pocilium

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                                                   273
              Table F




Zooplankton Species List:  Lake Erie

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                                                           274
            GREAT LAKES
DIVISION
  ZOOPLANKTON
   LAKE ERIE
     (1983)
                                    SPECIES LIST
                      TAXON
Ca lanoIda
CIadocera
Copepoda

CycIopoida
Harpact i co i da

Rotifera
Ca lano i d
0 i ap tomus
D j ap tomus
D i aptomus
D i aptomus
0 i aptomus
Ep i schura
Euryteroora
                                 copepodi te
                                ash land i
                                ID inutus
                                or egonensi s
                                sic i I  is
                                s i c iIoi des
                                lacustri s
                                 atfinis
Limnoca lanus macrurus
Senecella calanoides

Bosmina longirostris
Ceriodaphnia lacustris
Ceriodaphnia reticulata
Ceriodaphnia sp.
Chydorus sphaericus
Daphnia catawba
        ga laeta mendota
        retrocurva
        schod ler i
        sp.
             ecaudi s
             leuchtenbergianum
                      Daphnia
                      Daphnia
                      Daphnia
                      Daphn ia
                      D i aphanosoma
                      Di aphanosoma
Eubosmina coregoni
Eurycercus lamellatus
Holopedium gibberum
11yocryptus sp i n i fer
Leptodora k indt ii
S i oa cr ystalIi na

Copepoda NaupI i i

Cyclopoio - copepodite
Cyclops bicuspidatus thomasi
Eucyclops edax
Eucyclops prionophorus
Me socyclops edax
Tropocyclops  prasinus mexicanus

Ha rpact i co i da

Alona quadranquIaris
Ascomorpha ecaudis
As comorpha sp.
Asplanchna priodonta
Bdel loi d Roti fera
Brachionus bidentata
Brachionus caudatus
Br ac h i onus sp.
Co I  Iotheca sp .

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                                                         275
            GREAT LAKES ZOOPLANKTDN SPECIES LIST
                         LAKE ERIE
                           (1983)
DIVISION              TAXON
Kotifera              Co nochi Ioides sp.
                      Co nochi I us unicornis
                      Eu ch Ian i s sp.
                      F i I i na  Iong i seta
                      Gastropus sp.
                      Gastropus sty I i fer
                      Kellicottia longispina
                      Keratella cochlearis
                      Ke rate I la crassa
                      KerateI la ear Iinae
                      Keratella hi emal is
                      Keratella quadrats
                      Lepadel la sp.
                      Notholca  foliacea
                      Notholca  laurentiae
                      Notholca  squamula
                      PIoesoma  sp.
                      Polyarthra do I ichoptera
                      Po lyar thra major
                      Polyarthra remata
                      Polyarthra vulgaris
                      Synchaeta sp.
                      Trichocerca cylindrica
                      Trichocerca multicrinis
                      Trichocerca simills
                      Tr ichocerca sp.

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                                    TECHNICAL REPORT DATA
                            (flcaif rtadlmlrucDom on thr rrirrx btforr complctinf
        NO
   EPA- 905/3-88-001
                                                            3 RECIPIENT'S ACCESSIO*NO
 A TIT LE AND SUBT IT Li
   Phytoplankton and Zooplankton in Lakes  Erie,  Huron
   and Michigan: 1984
                                                            f> REPORT DATE
                                                              February 1988
e PERFORMING ORGANIZATION CODE
  5GL
 7 AUTHOR(S)

   Joseph C.  Makarewicz
e PERFORMING ORGANIZATION REPORT NO

  GLNPO Report No. 3
 9 PERFORMING ORGANIZATION NAME AND ADDRESS
   Department of Biological Sciences
   State University of New York
   College at Brockport
   Brockport, New York 14420
10 PROGRAM ELEMENT NO.
11. CONTRACT/GRANT NO.

  R005772-02
 12. SPONSORING AGENCY NAME AND ADDRESS
   U.S.  Environmental Protection Agency
   Great Lakes National Program Office
   230 South Dearborn Street
   Chicago, Illinois 60604
13. TYPE OF REPORT AND PERIOD COVERED
  Final  1984-1985
14. SPONSORING AGENCY CODE
  Great Lakes National Program
  Office, U.S. EPA, Region V
 15. SUPPLEMENTARY NOTES
   Paul  Bertram,  Project Officer
 16. ABSTRACT                                                                           	

   During the spring, summer and autmn of 1984 and winter  of 1985,  the structure
   of the plankton community in the offshore waters of  Lakes Erie,  Huron and
   Michigan  was monitored.  By examining changes  in the phytoplankton and zooplankton
   in relation to water chemistry, evidence was found suggesting  little change in
   the trophic status of Lake Huron and Michigan  while  an  improvement in the trophic
   status of Lake Erie was evident over the past  several years.   The offshore region
   of Lake Michigan is experiencing changes in phytoplankton and  zooplankton composition
   consistent with nutrient control and top-down  control by fish.   Even so, the biomass
   of phytoplankton and zooplankton and the trophic status  of the  lake have not changed
   significantly.  The appearance and establishment of  Daphm'a pulicaria in offshore
   waters of Lake Huron suggest a change in the forage  fish base.   With the exception
   of the resurgence of Asterione!1 a formosa in Lake Erie,  plankton composition has
   changed little since the 1960's.  However, dramatic  reductions  in biomass of nuisance
   and eutrophic indicator species have occurred.  These changes are consistent with
   expectations of long-term nutrient control.  However, a  change  in piscivory is
   evident that has apparently allowed the establishment of the large caldoceran
   Daphm'a pulicaria.

17. KEY WORDS AND DOCUMENT ANALYSIS
•d DESCRIPTORS
Lake Michigan Eutrophication
Lake Huron Great Lakes
Lake Erie Community structure
Plankton
Phytoplankton
Zooplankton
Limnology
18 DISTRIBUTION STATEMENT
Document is available to the Public
through the National Technical Information
Service(NTIS), Springfield, VA 22161
b. IDENTIFIERS/OPEN ENDED TERMS

19. SECURITY CLASS tTMs Report i
20. SECURITY CLASS (This page)
c. COSATl Field/Group

21. NO. OF PAGES
296
22. PRICE
EPA Form 2220-1 (9-73)
* U.S GOVERNMENT PRINTING OFFICE. 1988 543-858

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