eccmgv AND SEASGNAL' VARIATIGNQF THE; ,
PSAMMONFLQRA OF A SANDYBEACHJTRANSECT‘;g[7._':;11:'_}_4.fk-‘Zi-ngf»._

Thesis for fi’ha Dagéeeof Ph D 7: >

‘ MECHEGAN 5mm UNEVER-SITY ’ '

Wiiiiam Davies I ‘ . .   
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Michigan State
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This is to certifg that the

thesis entitled
ECULUGY AND SEASONAL VARIATION
OF THE PSAMMUNFLURA OF A SANDY
BEACH TRANSECT

presented bg

William Davies

has been accepted towards fulfillment
of the requirements for

M degree in M

   

Major professor

Date W

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ABSTRACT

ECOLOGY AND SEASONAL VARIATION
OF THE PSAMMDNFLDRA OF A SANDY
BEACH TRANSECT

by Nilliam Davies

   

The investigation involved a 12-month study of the

euflogy of the algae of the psammolittoral zone of Duck
Lake, a hardwater lake in Oakland County, Michigan. The
study was undertaken to ascertain the practicability of‘
adaptation of‘ techniques of. soil science and limnology to
psmmmlittoral zone investigation, to establish evidence of
zonation of‘ psammon algal communities and evidence of‘ sea-
smmlcmantitative and qualitative variation in these algal
commmities within a sandy beach, to attempt correlation of

I beach algal zonation and seasonal variation with physico-
chmfital conditions prevailing in the habitat, and to com-
pare the algal Flora of‘ the psammolittoral zone with that
of the adjacent lake.

The transect method of‘ community investigation was
utilized in study of the hygro~ and eupsammon portions of
the psammolittoral zone selected. Each collection date,
qualitative and quantitative phytoplankton samples and sur-
faceumter samples were taken offshore From the transect
and interstitial water withdrawn From the beach transect

hygropsammon zone. Determinations made upon lake and

 

 

 

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William Davies

  

orthophosphate, total iron, silica, and sulfates. Each
collection, at each 25 cm. interval along the transect,
determinations were made of’ temperature, pH, alkalinity,
substrate percentage water content, "black layer" position,
and total organic content.

Qualitative and quantitative sand samples were also
takm1monthly at each 25 cm. interval. Only algae present
h1samples From the 25, 100, 200, and 300 cm. intervals oF
theimansect were included in the study. Also investigated
unefluctuation of substrate phreatic levels and extent 0F
11mm penetration into the substrate. At each 25 cm. in-
temml, determinations were made oF substrate harmonic mean
grafl1size diameter, grain size grade distribution, eFFec-
tive grain size, grain size uniFormity coefficient,
porosity, and unit weight.

Values For substrate water content, harmonic mean
mahwsiza, eFFective grain size, grain size uniformity
coeflfitient, porosity, temperature, pH, and alkalinity
umrefpund Functions 0F distance From waterline.

In interstitial water, mean values For total and
bhmrmrmte alkalinity, calcium and carbonate hardness,
fifialtmrdness, dissolved £02, nitrate and nitrite nitrogen,
irmn sulfates, silicates, and total and volatile solids

wemaappreciably higher than those in the adjacent lake.

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William Davies

Fates, orthophosphates, iron, and nitrate and nitrite

pmliggen were observed in interstitial and lake water.

ield experimentation suggested chemical analyses of‘ inter-

‘ stitial water were not reFlective oF the environment con-

ditions experienced by psammon algae.

Qualitative phytoplankton collections were dominated
by members oF the Chrysophyta in all months but January and
June. Least phytoplankton species diversity was exhibited
during the December-March period 0F lake ice cover. Quanti-
tatively the phytoplankton was dominated by chrysophytes
and cyanophytes in all months but June. A clear seasonal
progression in quantitative dominance oF the phytoplankton
appeared during the study. The phytoplankton population
Fluctuation pattern was bimodal during the 12-month study,
mhflma occurring in winter and mid—summer, and maxima
occurring in spring and early Fall. Relations between
physicochemical Factors and qualitative and quantitative
Phytoplankton variation were discussed.

Qualitatively, the psammon algal Flora was essen-
Halhrone comprised 0F members oF the Hormogonales and
Females. Each oF the Four transect intervals were quali-
taflweh/dominated throughout the 12-month period by File.
mentous blue-green algae and diatoms. The psammon Flora
Wasqumfifltatively dominated throughout the year by File.
mentous blue-green algae and diatoms. Fluctuation in

total number of‘ algal individuals per cc. 0F beach Sand

   

 

 

 

 

William Davies

   

(lump the 12-month study was essentially bimodal in char-
".guer at all beach intervals. Total number of algal in-
dividuals at all beach interVals reached a primary maximum
during the December-March period of Frozen substrate and a
lesser maximum during the May—July period. Algal population
minima in the psammolittoral zone occurred in the March—
April period of thaw and fluctuating lake levels and again
in the late summer. A temporal inverse relationship was
evident in quantitative algal abundance in the lake and at
each beach transect interval. Total number of algal in-
dividuals per cc. of sand ranged from 114,523 to 18.3 times
the density of phytoplankton per cc. of lake water on the
same dates.

Indices of diversity were calculated for the algal
community at each beach interval each month of the 12—month
sampling period in order to identify evidence of physico-
chemical factor relationships to the algal community.

Direct relationship was identified between algal community
indices of diversity at each transect interval and water

_ content of the substrate. Inverse relationships were ap-
parent between algal community indices of diversity and
both harmonic mean grain size diameters and grain siZe
uniformity coefficients. Decreasing magnitudes of indices
Of diversity with increasing distance from shoreline in-

dicated an increasingly severe environment shoreward from

 

 

 

Waterline in the psammolittoral zone.

 

 

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William Davies

   

Community coefficients were calculated for the
communities within the lake and each beach transect
Val each month. A fundamental difference in community
u structure was indicated between lake and beach. Many of the
moflzqualitatively and quantitatively abundant algal species
inthebeadihave been characterized in the literature as
soil algae and a relatively large number were char-
acteristic of arid soil environments.

No ecotone could be identified in the beach as a
Mmlecm at any interval. It was concluded there was no
trmmitfln1in evidence from an aquatic to a terrestrial en—
virmmmnt along the beach transect. Rather, it was con-
chumd that the beach algal community exhibited a qualita-
thm amiquantitative attenuation shoreward from waterline

hIresponse to an increasingly severe environment.

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ECOLOGY AND SEASONAL VARIATION
OF THE PSAMMONFLORA OF A SANDY
BEACH TRANSECT

By

Hmrdd william Davies In

A THESIS

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY
Department of Botany and Plant Pathology

1966

 

 

 

 

CDCopyright by
Harold William Davies Jr.

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ACKNOWLEDGMENTS

  

The author wishes to thank Dr. G.M. Prescott of
the Michigan State University Botany Department for his
assistance, counsel, and patience. Acknowledgment and
gratitude are extended to Drs. J.E. Cantlon and M.B. Drew
of the Department of Botany and Dr. 8.8. Ellis of the
Department of Soil Science for their helpful advice and
provision of research facilities as well as to the National
Science Foundation for its financial support. Greatly
appreciated also is the assistance rendered by Drs. Ruth
Patrick and c.w. Reimer of the Department of Limnology,
Philadelphia Academy of Natural Science, in diatom identi-
fication and in the provision of research facilities and
financial support during the period of study at the Academy.
The author wishes also to express gratitude to
the personnel of the phycology laboratory at Michigan State,
Particularly Rosa Bicudo, R. Kullberg, and D.C. Jackson, for
their assistance. And special acknowledgment is due to
Norma Davies for her encouragement and her help in thesis

Preparation .

ii'

 

 

 

TABLE OF CONTENTS

ACKNOWLEDGMENTS . . . . . . . . . . . . .

LIST OF TABLES C . . O O l O O O C D C 0

LIST OF FIGURES I C O O O O O O I O C O 0

Chapter

IV.

C
n

INTRODUCTION . . . . . . . . . .

Results of Preliminary Studies
Statement of Problem . . . . .

LITERATURE RE\IIEUJ O O I U U I I 0
DESCRIPTION OF THE STUDY AREA . .

General Description of the Area
Description of Beach Transact .

METHODS AND MATERIALS . . . . . .

Granulometric Description of the Beach

Phreatic Levels . . . . . .
Physical Data . . .
Chemical Data . . .
Biological Data . .

RESULTS 0 o o o o o v o o o e o o

General Information . .

O

Granulometric Characteristics of the

Transact Substrate . . . . .
Beach and Lake Physical Data .
Beach and Lake Chemical Data .

Biological Data, Qualitative Lake Samples

Biological Data, Quantitative Lake

Samples . . . . .

0

Biological Data, Qualitative Psammon
Samples, 300 cm. Interval . . . . . .

Biological Data, Quantitative Psammon

Samples, 300 cm. Interval . . . . . .

iii

 

O

Page

a . mjwinl. a:

 

   
  

Biological Data, Qualitative Psammon
Samples, 200 cm. Interval . . . . . .
Biological Data, Quantitative Psammon
Samples, 200 cm. Interval . . . . . .
Biological Data, Qualitative Psammon
Samples, 100 cm. Interval . . . . . .
Biological Data, Quantitative Psammon
Samples, 100 cm. Interval . . . . . .
Biological Data, Qualitative Psammon
Samples, 25 cm. Interval . . . . . .
Biological Data, Quantitative Psammon
Samples, 25 cm. Interval . . . .
Biological Data, Recapitulation of
Results of Monthly Qualitative and
Quantitative Sampling for 12-month
Study periOd . e o o o o o o I o o 0
Biological Data, Indices of Diversity
of Lake and Beach Transact Intervals
Biological Data, Community Coefficients

VIQDIscussroN...............

Terminology Relative to the
Psammolittoral .
Porosity and Grade
Temperature . . . .
Light 0 O O O I O 0
Black Layer . . . .
Organic Matter . .
Mater Content . .
Chemistry of Interstitia n a
water: pH . . . .
Chemistry of Lake and Interstitial
Mater: Dissolved Gases and Dissolved
SOlidS O O O O O O O O O O O 0
Lake phytoplankton o o o o o
Psammon Algae: Qualitative Aspects . .
Psammon Algae: Quantitative Aspects .
Indices of Diversity of Lake and Beach
Transact Intervals . . . . . . . . .
Community Coefficients . . . . . . . .

C
O
I
U
I
O

I O O U I D O
I I O O O I I
I O O O Q 0 O

7:00.000.

F‘s-coco.
(l)

0-. O O O O I I

i;

 

VII! SUMMARY 0 o u o o o e e o e o e I o o o 0

LITERATURE CITED 0 n e o o o o o o a o o e e a o

PLATES

I O Q 0 O I I 0 o O o O U I o 0 e I o 0 0

iv

 

Page

171
184
199
212
229
244

315
325
345
345
357
364
370
374
376
381
406
412
428
440

452
454

459
471

489

 

 

LIST OF TABLES

   

Table

1. Major Morphometric Characters of Duck Lake
and REaCheS A and A1 0 o I o o o e o o o I

2. Daily Precipitation Data in Inches for
Months October, 1963, Through October,
1964, Highland, Michigan, Station of the
Detroit Metropolitan Area Raingage Network

3. Daily Precipitation Data in Inches for
Months October, 1963, Through October,
1964, White Lake, Michigan, Station of the
Detroit Metropolitan Area Raingage Network

4. Percentage Composition 150 gm. Incinerated
Beach Transact Sand Samples by Dimension
Based on U.S. Sieve Series for Intervals
‘100 to +300 CIT]. e e e o o o o e o I o o o

5. Granulometric Data for Beach Transect
Intervals ‘100 to +300 Cr”. 0 o o o v o o .
6. Monghly Lake and Transect Temperatures in
C

7. Percentage Water Cemposition of Monthly Sand
Samples at Each 25 cm. Interval on Wet
Gross Weight Basis . . . . . . . . . . . .

3. Percentage Water Content of Monthly Sand
Samples at Each 25 cm. Interval on Net
Weight Basis Expressed as Per Cent of
Saturated Capacity . . . . . . . . . . . .

9. Percentage Volatile Matter on Dry Gross
Weight Basis at Each Transect Interval
monthly 0 o o O O I o o o o O O o o o o 0

10. Monthly Determinations of pH of Lake Water,
Interstitial Water, and of the Substrate
at Each 25 cm. Transect Interval . . . . .

Page

19

27

28

SO

51

65

58

72

 

 

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1-3.4

 

 

 

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20.

21.

22.

23.

24.

25.

Bicarbonate Alkalinity as ppm. CaCDg per
Month Each 25 cm. Interval Along Transect

Lake Water Chemical Data for Samples
November 7 to October 11 . . . . . . . . .

Hygropsammon Zone Interstitial Water
Chemistry . . . . . . . . . . . . . . . .

Percentage Calcium and Carbonate Hardness of
Total Hardness for Lake and Interstitial'
water I O C O I O O O O O O O O O I O O I

Lake Phytoplankton Qualitative and
Quantitative Data for 12—month Sampling
period 0 O I O O O O O C O O O O C O C O 0

Summary of Lake Phytoplankton Qualitative
Sampling for 12-month Sampling Period . .

Quantitative Lake Phytoplankton Samples
Species Percentage Composition . . . . . .

Order and Division Percentage Composition of
Monthly Lake Quantitative Samples . . . .

Qualitative and Quantitative Data for
Psammonflora Algae Present at +300 cm.
Interval Monthly for 12-month Sampling
Period 0 Q 0 O 0 O O O 0 O O O I O O O O I

Summary of 300 cm. Interval Qualitative
Sampling for 12-month Sampling Period . .

Quantitative Sample Percentage Composition
Monthly, 300 cm. Interval . . . . . . . .
Order and Division Percentage Composition of
Monthly Quantitative Samples, 300 cm.
Interval . . . . . . . . . . . . . . . . .
Qualitative and Quantitative Data for
Psammonflora Algae Present at +200 cm.
Interval Monthly for 12-month Sampling
Pariod.........-o--o.-..
Summary of 200 cm. Interval Qualitative
Sampling for 12-month Sampling Period . .
Quantitative Sample Percentage Composition
Monthly, ZOO cm. Interval . . . . . . .

vi

Page

7?

79

81

93

111

123

135

144

166

172

1B?

16-—

 

 

Order and Division Percentage Composition
of Monthly Quantitative Samples, 200 cm.
IntervaleoeI-eoeooeeoeoo

Qualitative and Quantitative Data for
Psammonflora Algae Present at +100 cm.
Interval Monthly for 12-month Sampling
period 0 O O O C C O O O O O O O O O I 0

Summary of 100 cm. Interval Qualitative
Sampling for 12-month Sampling Period .

Quantitative Sample Percentage Composition
Monthly, 100 cm. Interval . . . . . . .

Order and Division Percentage Composition
of Monthly Quantitative Samples, 100 cm.
Interval O C I O O O O O O O O O I O O O

Qualitative and Quantitative Data for
Psammonflora Algae Present at +25 cm.
Interval Monthly for 12-month Sampling
period . O O O C D D O O I O O O O O O 0

Summary of +25 cm. Interval Qualitative
Sampling for 12-month Sampling Period .

Quantitative Sample Percentage Composition
Monthly, 25 cm. Interval . . . . . . . .

Order and Division Percentage Composition
of Monthly Quantitative Samples, 25 cm.
Interval O O O O I O O O O O O O O I 6 C

 

35. Summation of Qualitative and Quantitative
Contribution of Lake and Psammon Zone
Algae for 12-month Study Period . . . . .

36. Division and Order Summation of Species
Reported in Qualitative Samples Taken from
Lake and Beach Transect Intervals During
12-m0nth Study a o c o o a o o o o o o o .

‘ 37. Summary of Total Species Number in Lake and
Beach Transect Qualitative Samples Monthly
‘ During 12-m0nth Study a e o o a o o o e l

1 38. Summary of Division and Order Contributions

‘ to QUalitative Beach and Lake Qualitative
1 Samples, November to October . . . . . . .

1 vii

 

Page

193

200

209

216

224

247

255

291

293

295

Page

Summary of Quantitative Data, Total Number
of Individuals per cc. of Sand in
Quantitative Samples, November Through
October, Lake and Beach Transect Intervals 300

Relative Monthly Quantitative Abundance of
Algae in Beach Transect Intervals,
November Through October, With Respect
to Lake Phytoplankton Density . . . . . . 307

Index of Diversity for Lake and Each Beach
Transect Interval Monthly, November to
October . . . . . . . . . . . . . . . . . 320

Coefficients of Community and Species in
Common, Lake and Beach Transect Intervals,
Monthly November to October . . . . . . . 328

Summary of 12-month Means for Values of C

and Number of Species Shared in Common by

Each Collection Site in Relationship to

the Other Four . . . . . . . . . . . . . . 338

 

viii

i

 

2.
3.

4.
5L

5.

7.
8.
9.

10.

11.

12.

13.

 

    
 

I Figure
1.

LIST OF FIGURES

Map of Clyde, Michigan, and Vicinity Showing
Location of Duck Lake

Outline Map of Reach A of Duck Lake Showing
Location of Transect (X)

Beach Profile at Transect Site Showing Late
Summer Phreatic Levels .

Massed Samples Grain Size Cradation Curve .

Diagrammatic Representation of "Black Layer"
Distribution During Months Observed . . .

Graphic Comparison of Transect Intervals
Twelve Month Means Volatile Matter
Percentage on Dry Gross Weight Basis and
Percentage Water Content on Wet Gross
Weight Basis .

. O O I D I O O O 0 O O I 0

Monthly Variation in pH of Lake and
Interstitial Water .

0 O 0 O C C O Q C I O

Dissolved Carbon Dioxide in ppm. for Monthly
Lake and Interstitial Water Samples . . .

Dissolved Oxygen in ppm. for Monthly Lake
and Interstitial Water Samples . . . . . .

Dissolved Oxygen as Per Cent Saturation for
Lake and Interstitial Water Monthly
Samples . . .

Total and Volatile Dissolved Solids in
mg./l. for Monthly Lake Collections . . .
Total and Volatile Dissolved Solids in

mg./l. for Monthly Interstitial Water
Samples . . .

Volatile Dissolved Solids Expressed as
Percentage of Total Dissolved Solids for
Monthly Lake and Interstitial Water
Samples . . . .

O O O O 0 O O O O 0 O O 0

ix

Page

18

2O

31

53

61

7O

74

83

85

B7

B9

89

90

2. Lake
a

‘ “hrs
II“

‘- Eta}

‘ ‘0", 1,,”

  
     

Monthly for Total, Calcium, and Carbonatg
Hardness . . .

I . . U 0 O C 0 O O O C . O

Interstitial Water Hardness Expressed as

ppm. CaCO3 Monthly for Total, Calcium, and
Carbonate Hardness .

O C O O O O i O O O C

Total, Bicarbonate, and Carbonate Alkalinity
of Lake Water at Monthly Collections . . .

17. Total and Bicarbonate Alkalinity of
Interstitial Water at Monthly Collections

18. Total Ferrous and Ferric Iron in ppm. for
Monthly Lake and Interstitial Water
Samples 0 O C C O Q 0 O O O O O O I O O O

19. Sulfates in ppm. for Monthly Lake Water and
Interstitial Water Samples . . . . . . . .

20. Silicates in ppm. for Monthly Lake Water and
Interstitial Water Samples . . . . . . . .

21.

Orthophosphate in ppm. for Monthly Lake
Water and Interstitial Water Samples .

Q 0

22. Inorganic Nitrate and Nitrite Nitrogen in

ppm. for Monthly Lake Water Samples

23. Inorganic Nitrate and Nitrite Nitrogen in
ppm. for Monthly Interstitial Water
Samples . . .

O O O O O D U I O O O O O O

24. Diagram of Bore Field Utilized in Tracing
‘1 Hygropsammon Zone Interstitial Water
Movement . . .

O O O O O C O O C D 0 C O O

25. Total Number of Individuals per Liter,
Quantitative Phytoplankton Samples
November Through October .

O C O C . O I

26. Division Percentage Composition in

Quantitative Lake Samples, November
Through October . .

Q 0 O C I U I O O O O

27. Total Number of Individual Algal Units per
cc. of Sand, Quantitative Monthly Samples,
300 cm. Interval . . .

o o o o o e e e e e

 

Page

91

91

97

9B

99

100

101

103

104

105

108

127

145

160

L
r
7
f.

 

Page

   

Summary of Percentage Composition by
Divisions for 300 cm. Interval
Quantitative Samples, November Through
notch-rooeoesooo-ooooooo167

29. Total Number of Individual Algal Units per

cc. of Sand, Quantitative Monthly Samples,

200 cm. Interval O O O I O O D O O I O O O 185

30. Summary of Percentage Composition by
Divisions for 200 cm. Interval
Quantitative Samples, November Through
October 0 O C O O O O O O O O O O O O O O 194

31. Total Number of Individual Algal Units per
cc. of Sand, Quantitative Monthly Samples,
100 cm. Interval . . . . . . . . . . . . . 213

32. Summary of Percentage Composition by
Divisions for 100 cm. Interval
Quantitative Samples, November Through
DCtOber O O O O O O I O D O I O O O O O O 225

33. Total Numbers of Individual Algal Units per
cc. of Sand, Quantitative Monthly Samples,
25 cm. Interval C O l O O I C I I 0 O O O 245

34. Summary of Percentage Composition by
Divisions for 25 cm. Interval Quantitative
Samples, November Through October . . . . 256

1
1
1 35. Summary of Quantitative Date, Total Number
1 of Individuals per cc. of Quantitative

Samples, November Through October, for
Lake and Beach Transect Intervals . . . . 301
1
1
i
1

36. Relative Monthly Quantitative Abundance of
Algae in Beach Transect Intervals,
November Through October, with Respect to
Lake Phytoplankton Density . . . . . . . . 308

37. Index of Diversity for Lake and Each Beach
Transect Interval Monthly, November to
DCtOber O O I O C I O I C C O C O O O O O 321

(J:
(23
0

Community Coefficients for Reach A1 and Four
Beach Transect Intervals Monthly, November
to OCtObEr O O O I O O O O O O O I O I O D 329

 

Page

   

, Community Coefficients for the 25 cm.
‘ Interval in Relationship to Reach A1 and
Other Beach Transect Intervals Monthly,
November to October . . . . . . . . . . . 331
Community Coefficients for the 100 cm.

Interval in Relationship to Reach A1 and

Other Beach Transect Intervals Monthly,

November to October . . . . . . . . . . . 333

Community Coefficients for the ZOO cm.
Interval in Relationship to Reach A1 and
Other Beach Transect Intervals Monthly,
November to October . . . . . . . . . . .

Community Coefficients for the 300 cm.
Interval in Relationship to Reach A1 and
the Other Beach Transect Intervals
Monthly, November to October . . . . . . . 336

Comparison of Index of Diversity with Mater
Content on Net Gross Weight Basis for Each
Beach Interval Monthly, November to

OCtOber I O O C O O O I O O O O C C O I O 342

 

 

 

 

 

 

 

 

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CHAPTER I
INTRODUCTION

Since the recognition of Sassuchin, Kabanov, and
Neisuestnova in 1927 that interstices within a sandy beach
contain a well-developed microflora and fauna, considerable
research has been done upOn this environment (see Chapter II,
Literature Review). Most published reports upon investiga—
tions of the sandy beach, however, have emphasized the
taxonomic approach, particularly that of the rotifer com-
ponent of the microfauna. In comparison with the extensive
literature relating to the ecology of the algae of aquatic
and terrestrial environments, that of the microflora of the
sandy beach transition zone between the two has been largely
ignored.

Controversy still exists considering the terminology
and ecological concept of the interstitial water ecotone
within a beach. For that matter, definitions applied to
the zones of the beach itself are often vague and contra-
dictory (welsh, 1952, pp. 315-319). Sassuchin gt _a_1. (1927)
considered the environment transitional between aquatic
and soil habitats, referring to it as the "psammon", but
Nisznismski (1934) considered it rather a unique type of

aquatic littoral habitat, the "psammolittoral". Subsequent

    

 

jg“ body of water.

Generalizations concerning the physical-chemical

aspects of the psammon environment must be qualified as to

3 the relative proximity of the normal shoreline in that the

literature of research upon the ecology of the psammon
indicates that any given physical-chemical factor tends to
be a function of the distance from the shoreline. It is this
horizontal Variation of physical-chemical factors with dis-
tance from shoreline that has been used to justify
Wiszniewski's concept of the sandy beach as a unique type
01’ aquatic environment, the psammolittoral, more or less
transitional between the aquatic and terrestrial environments.
Most investigators have utilized miszniewski's
terminology to delimit zones of the psammon as based upon
moisture content and exposure to wave action. These are
as follows: (1) the hydropsammon of permanently submerged
sand below the water's edge, (2) the hygropsammon zone
adjacent to the water's edge and completely saturated by
capillarity and wave action, and (3) the eupsammon zone,
landward from the hygropsammon zone, which is partially
saturated and exhibits frequently a thin, dry surface layer.
Pannak (1940), Neel (1948), and mum (1952) compare the
hygropsammon zone to the inner beach and eupsammon to the

middle beach. Review of the literature indicates confusion

 

.'. .n'b
-n_' lat-II

“rh- .5
In“ .
"u

u
u

an,
.il"
-- .

'-

“Wan—3‘» N'—

3

  

ng delimitation between the hygro- and eupsammon
and it is evident differentiation between them is
y arbitrary.

Pennak (1939, 1940, 1950) has stated that perhaps
other environment supports such a dense and diversified
group of microorganisms as the sandy beach. Subsequent
investigations have tended to verify this and his charac-
terization of the psammonflora as basically dominated by
diatoms associated with blue-green algae and, to a lesser
extent, green algael Most investigations have indicated
the psammonfauna to be composed primarily of testaceous
rhizopods, nematodes, rotifers, oligochaetes, tardigrades,
and copepods, many species of these being unique to the
psammon or exhibiting a definite distributional perference
to the psammon habitat. The biota of the psammon apparently
tends to be restricted to the surface few centimeters of
the sand, the producers within the photosynthetic 20ne at
1 the surface, and the consumers barred from depths of more
than a few centimeters by chemical conditions developing
From organic decomposition at lower levels. Apparently,
too, the abundance of organisms within the hydropsammon is
far less than that of emergent portions of the beach. The
l high biological productivity of the psammon zone micro-
ecosystem has been attributed to the close vertical rela-
tion of the photosynthetic and decomposition zone (Neel,
1948).

Strangenberg (1934) has described the psammo-

littoral as an extremely eutrophic water medium, and

 

 

u re sub

"‘ {rely

Trent T

. ....

4

  
    
  

to) gliding on an average 54 per cent more dissolved in-
mé‘gnic material and 42 per cent more dissolved salts than
Uia‘adjacent lake. Physical-chemical conditions in this
medium are subject to extreme seasonal and diurnal varia-
tions largely produced by interaction of factors influenc-
ing movement of water through the voids between adjacent
sand grains, the fluctuations becoming more pronounced with
distance landward from the shore. Too, the milling action
of the sand matrix induced by wave action, diurnal expan-
sion and contraction produced by thermal variatiOn, and
major shearing and plowing by ice movements on lee shores
results in an unstable physical environment. That such
an environment supports such numerous inhabitants assuredly
warrants far more investigation than it has received.
wiszniewski's concept of the psammolittoral as
contrasted to Sassuchin's psammon has been adhered to in
most ecologically oriented psammon studies, limnological
techniques having been utilized to the almost complete
exclusion of techniques of soil study. A search of the
literature indicates that no study has been made incorporat-
ing or comparing to any extent the standard methods used
by the limnologist and soil scientist in environmental
research. Ruttner-Kolisko (1956), however, has pointed
Out the necessity of some standardization of approach to
study of this unique environment.

Most ecologically oriented studies of the psammon

have been extensive rather than intensive, and, although

 

a ram
ta'nmti
H.615" l

E' 2112:!

 
 
   

5

‘ all investigators have reported upon evidence of sea-

study conducted at regular intervals throughout the year
on a freshwater beach. Neither does there appear to be

any study of algal communities as such in the freshwater
beach comparable to those of Wieser (1959) or of King (1960,
1962) upon invertebrates in marine beaches.

In 1952 Welch stated that little, if any, informa-
tion was available with regard to the fate of psammon
populations in winter when beaches are frozen, when falling
water level causes a gradual migration of the water-
saturated beach zone, .and in the presence of shore erosion
by ice. He noted at that time (1952) that the general
reletiee of the psammon to the main water mass of adjacent

lake or stream was still uncertain.

Results of greliminary studies

1 During 1959 and 1960, preliminary psammon studies
were conducted upon several beaches in south central

lower Michigan to investigate the feasibility of conduc-
tion of year around field research upon the psammon
environment at that latitude. These preliminary studies
indicated that, contrary to generalizations in the litera-
ture, the psammonflora is not necessarily dominated by
diatoms, at least not throughout the year. Definite
ssasonal qualitative and quantitative periodicity was

evident in both algal and invertebrate populations during

 

 

 

it"s: Id

1'5 «)1

I'll
22:1 I!
Yeats:
Till!!! I
112':

t: E
3353! I}

E :32?)

6

   

l invertebrates and autotrophic and heterotrophic protistans
immediately upon thawing. Initial winter freeze in Lake
Lansing and Big Portage lake produced a frozen psammon
layer 6 to 8 inches thick which was sheared from the
substrate by expansion and movement of lake ice. It was
these psammon layers then buried under one to three feet
of snow and fractured lake ice that yielded numerous
living algae and invertebrates upon thawing.

Attempts to use tool steel cores and chemical
thawing for quantitative winter psammon samples were re-
Peatedly unsuccessful. In addition, the time involved in
thawing gross sand samples nullified attempts at chemical
determinations of interstitial water. It was seen,
however, that south-facing beach slopes frequently thawed
to an appreciable depth suitable for quantitative algal
samples on sunny winter afternoons, but still such beaches
did not yield sufficient interstitial water for chemical
determinations in the winter months.

In all, preliminary observations were made on a
total of 89 lakes within a 60 mile radius of East Lansing
with respect to the presence or absence of a south-facing

beach Possessing the selection criteria listed in

 

 

2'21 m
{12:55.

7

  
   

~‘4rIII, Description of Beach Transect. Observations

This investigation was concerned with the fol-
lowing problems:
1. to attempt standardization in psammon study
of physical-chemical determinations by amalgama-
tion of soil science techniques and those of
limnology,
2. to establish evidence of zonation of algal
communities in a sandy beach,

3 to establish evidence of seasonal qualitative

and quantitative variation in algal populations

in a sandy beach,

J:
o

to attempt correlation of zonation and seasonal
variation with physical-chemical conditions
prevailing in the habitat,

5. to compare the algal flora of the psammon with

that of the adjacent lake.

 

 

 

 

 

 

 

 

 

 

 

 

CHAPTER I I

LITERATURE REVIEW

  
 

Probably the first published record of investiga-
tion of the psammobiota is that of Sassuchin (1926) who
investigated the microbiota, particularly the Protozoa,
of the sandy shores of the Dka River, Russia. Olson-
Seffer (1909) had previously studied hydrodynamic factors
influencing plant life on sandy sea shores, and Oliver
(1912) had described the shingle beach as a plant habitat.
Hill and Henley (1914) investigated the source of inter-
stitial water in marine shingle beaches, and warren and
Mudd (1924) reported on the active migration of bacteria

1 through sand substrates. Herdman (1921) described dis-
taloration of the sand at Port Erin produced by dino-
flagellates and other algae.

It is believed that Sassuchin, Kabanov, and
Neiswestnova (1927) were the first, however, to recognize
the psammolittoral as a distinct environment, regarding it
1 as a transition zone between the aquatic and soil habitat
1 and coining the term "psammon" to designate this environ-
ment. Their report, presenting the results of study of
1‘ the microbiota inhabiting the beach of the Uka River, was

concerned primarily with the microfauna of the environment,

 

 

mustta‘

'2.

Wméxi

   
  
 
 
  

9
”fiflgh some phy91ca1-chemical data were also given.

#ushin elaborated upon ecological conditions of life in

and Shadina described faunal differences occurring in the
v sandy substrate from midstream to the banks of the Oka
River.

In 1921 Klie described the copepod component of
the sand-dwelling fauna of north German beaches, and in
the same year Bracher investigated the ecology of the
banks of the brackish mouth of the Avon River, noting
distinct communities of algae -- primarily Euglena and
diatoms -- that showed spatial distribution on the basis
of substrate moisture content, seasonal qualitative and
quantitative variation, and diurnal vertical migratiOn
attributed to light regimes.

In 1932 Pirre, Bruce, and Moore reported the
results of a quantitative study of the metazoan population
0? a sandy marine beach at Port Erin in which they in-

‘ eluded pH, mechanical analysis of grain size, grade,
salinity, and organic content data in reference to the
substrate. Distinct distribution patterns were attributed
to pore siza and grade. Bruce (1928a, 1928b) had pre-
viously published accounts of investigations of physical-
chemical factors occurring in marine beaches at Plymouth,
England. Also in 1932, wilson reported the occurrence

Of several species of copepods from marine beaches in the

Vicinity of woods Hole, and in 1935 he described the sandy

 

 

10

Rob as a new and important copepod habitat, referring to
copepods of such a habitat as terraqueous and attributing
discovery of the habitat to N.A. Cobb.

miszniewski in 1934(a) published the first of an
extensive series of reports upon the freshwater sandy
beach as a special type of environment for microscopic
life. Strangenberg in the same issue of the publication
described the environment, the psammolittoral, as an
extremely eutrophic water medium as based upon data col-
lected pertaining to the chemistry of interstitial water
from two beaches in Poland. Probably no other investigator
of the psammolittoral biota has been so prolific in pub-
lication (1934a, 1934b, 1935a, 1935b, 1936a, 1936b, 1936c,
1937a, 1937b, 1947). Most of these publications deal with
1 the taxonomy and biology of rotifers inhabiting sandy
beaches of lakes and rivers at various locations in Poland,
although some (1934b, 1947) contain considerable references
to ecology of the psammolittoral zone.

Aside from the investigation of the environmental
factors acting in the sandy beaches of New Brunswick and
Preliminary designations of their intertidal communities
reported in 1935 by Newcombe, published accounts of re-
search upon the psammolittoral in the 30's and early 40‘s
were concerned almost entirely with the taxonomy of its
metazoan inhabitants. Nicholle in Scotland (1935, 1939)
and later (1945a, 1945b) in western Australia and Schulz

(1937) and Kunz (1938) in Germany described new copepod

 

L'Eilhab
dad a]

:1 mm!

11

  
     

fies inhabiting marine beaches. Also contributing to

a; are of almost exclusively metazoan taxonomic approach
{in the psammolittoral zone were Varga (1938) with a report
n the micropsammofauna of Hungarian lakes and Gieysztor
(1938) reporting upon turbellariana from the freshwater
psammon of Germany. Choppuis (1942, 1944) described the
ieopods of the sandy margin of a Hungarian stream, and
Stentscheff (1944) reported upon the nematodes of the
psammolittoral of a Holstein lake. In this country Myers
(1936) listed the species of rotifers found in the psammo-
littoral of several acid lakes in New Jersey, and Tressler
(1940) described a number of ostracods from a marine sandy
beach near Beaufort, North Carolina. Pennak (1939, 1939b)
described a new rotifer and new copepod from the psammo-
littoral zone of some Wisconsin lakes.

Pennak subsequent to his taxonomic papers of 1939
later (1939c, 1940) emphasized the ecology of microscopic
metazoans inhabiting the psammolittoral of misconsin lakes.
In 1950 he published a summary of what was then known of
the comparative ecology of interstitial fauna of marine
and freshwater beaches. Papers describing copepods from
intertidal beaches near woods Hole (1942, 1942a), a new
order of crustacean from an intertidal beach (1943) and a
new species of isopod found in marine beaches of the Gulf
of Mexico (1958) have also been published by Pennak.

In 1942 Pearse, Humm, and wharton reported upOn

the ecology of marine sandy beaches of North Carolina.

 

 

 

12

   

wwvhwgh emphasis in the report was upon the rnetazoa, some
emeralizations were made concerning the relative abundance
sf various algal classes in the psammobiota and their

, morphological adaptations to the environment. In the same
year Lund reported upon seasonal variation and phototactic
response of the psammon algae of English ponds. Previously
(1940) Beanland investigated sand and mud communities of

the Dover estuary utilizing quantitative techniques. She
related faunal distribution to grade size of the substrate,
exposure time, and salinity gradation.

Papers closely related to psammolittoral zone
research were published by Moore (1939) who described a
limnological investigation of the microscopic benthic
fauna of Douglas Lake, and Noffett (1943) who described
the limnology and seasonal variation of the macroscopic
hydropsammon zone fauna of the same Michigan lake. Neel
(1948), investigating the same lake, reported upon the
limnological relationships of the psammolittoral zone.
Considerable attention was given to the ecology of the
algal component of the psammobiota.

Aleem (1950) described diatom communities present
in the surface two mm. of the substrate of English inter-
tidal mudflats. Quantitative determinations of diatom
communities were correlated with temperature, light penetra-
tion, organic content, and gradation of pore size. Definite
seasonal variation was observed as was vertical migration

Df’ diatoms in response to variations in light intensity.

 

 

‘11:: :n

“i“- I
--.... 4

:0 or; -‘

‘Iu-gu

:- A.‘

~- :1

‘g n .__
"- u .

13

     

. 1961). Canapati, Rao, and Rao (1959) reported on the tidal
rhythm of diatoms and dinoflagellates of intertidal sands
of Indian beaches, attributing alternation of low tide
photophilic phase and high tide photophobic phase to in-
herent periodicity determined by tidal rhythms. Janika
(1954) published a brief summary concerning generalizations
about the composition of psammon flora and its relation-
ships to physical-chemical factors of the psammolittoral
zone;

Levi (1950) described the isopods of French beaches,
Remacne and Siewing (1953) psammon rotifers from marine
beaches in Brazil; Among a series of papers describing
the interstitial fauna of African beaches were those of
1 Gnanamuthu (1954), Chappuis (1954, 1954a, 1954b, 1956b)

‘ and Delamare and Chappuis (1956a, 1956b, 1957). Chappuis

i (1956a) also described the interstitial fauna of the
Bahamas. Koepscke and Koepscke (1952) discussed the origin
of organic matter in marine beaches of the coast of Peru.

Investigations concerned primarily with the micro-
Fauna of European marine beaches include those of Vacelet
(1961) who described seasonal variation of the infusorian
microfauna of a medium sand beach near marseille, Janseon
(1960) who reported upon the taxonomic study of inter-
stitial fauna of a sandy beach with some reference to

Organic matter and water relationships, Prenant (1960) who

 

 

‘lll‘rcnsta

into: 1
turned
'e‘. .1352)

:1! 11:1

  
    

reuseed granulometric methods of ecological investigation
a)? intercoastal sands, and Scott (1960) who described the
fp'erse fauna of a shifting beach at St. Kilda. Although
not concerned with the microfauna of marine beaches,
Selvat (1962) used quantitative“ methods to demonstrate
seasonal variation of macrofauna during year-round regular
sampling of French beaches. And Boaden (1963a) has re—
ported upon the taxonomy of marine gastrotrichs from the
interstitial fauna of some north wales beaches. Additional
reports by Boaden (1963b, 19630) were concerned with the
distribution of nematodes on many marine beaches, over 100
new species being reported with distribution patterns
apparently determined by grain siZe of the substrate.
Recent publications referring to investigations
1 01“ European freshwater psammolittoral habitats include
those of Milosevic (1960) who presented generalizations
concerning biological activity of different levels of
river sands, Sakharova (1963) who described the micro-
bsnthos of sandy beaches of the Uchinsk Reservoir in
Russia, and Altherr (1963) who presented a list of nematodes
Occurring in the interstitial water of sandy banks of the
Moselle River. Ruttner-Kolisko (1953, 1954, 1955a, 1955b,
1956, 1961, 1962) has published a series of reports of
ecologically oriented studies on lentic and lotic psammo-
littoral environments in Scandanavia and Italy. Although
stressing the rotifer component of the microfauna, she

has included some generalizations pertinent to the algae

 

 

 

 

 

::
a...
5;. :

 

I?“ u
n
\L

 

 

1S

  
   

h psammolittoral Zone. Evans (1958, 1959, 1960) has

substrate. He, too, has noted vertical migration of
the algae corresponding to moisture content changes.

Round (1960, 1961a, 1961b) in investigations related to
the periphyton of the hydropsammon zone has reported
seasonal cycles in abundance of the Cyanophyta and diatoms
in English lakes correlating with variatiOns in light,
temperature, silicates, and nitrates, the cycles of the
periphytes differing in considerable degree to those of
the phytoplankton.

Recent investigations concerned with the psammo-
littoral zone in the United States have been primarily
related to studies of the microfauna of marine beaches.
Wieser (1959) has reported upon the effect of grain size
upon distribution of small invertebrates inhabiting the
beaches of Puget Sound, distribution and barriers to even
distribution apparently being determined by the grain
size of the substrate. Crites (1961) has listed marine
nematodes living in the upper three inches of sandy North
Carolina beaches. King (1960, 1962) has published results
01’ investigations dealing with the ecology of psammolittoral ‘
POPUIations and communities of nematodes of the north-
eastern Gulf of Mexico and has found evidence of communities

related to average grain size of the substrate and organic

 

16

   

contained therein. Weber (1963 unpublished VHS.
711-) has made a study of seasonal variation of the
fauna of the psammolittoral zone of a Michigan
some reference also being made to seasonal variation

the psammoflora.

 

 

 

 

 

 

l
:‘ --.
I l

 

 

CHAPTER III

  
 

DESCRIPTION OF THE STUDY AREA

General description of the area
Duck Lake is located at an altitude of 1016.3 feet

in sections 11, 12, and 14 of Highland Township, TEN, RTE,
Oakland County, michigan. Humphrys and Green (1962)

assign lake number 569 to Duck Lake and describe it as a
natural lake of 253 acre area and 51 foot maximum depth,
the area being estimated by reference to the 1909 U.S.G.S.
Milford quadrangle topographic survey map. No more

recent topographic map of this quadrangle exists to data.

Figure 1 diagrams the lake and vicinity, the lake
outline as of spring, 1964, being based upon 1957 aerial
‘ photographs of the U.S.D.A. Commodity Stabilization
Service, Oakland County Drain Commission maps, and aerial
Photographs supplied by Mr. John Namish, owner and devel-
Oper of the lake.

In the winter of 1958-1959, a channel was dredged
approximately 0.05 miles toward the WNW to connect Duck
Lake to a former segment isolated and separated by
maintainance of the main lake at its then current level.
Although assigned lake number 573 in Michigan Lake
Inventory Bulletin No. 63, since 1959 the body of water

17

J— ,_,,_,,,__,,,,. ,,_ ,

 

 

 

 

 
   
   
       

Duck Lake

Milford
Road

Harvey Lek: Road

 

White Lake Road

 

Ghesapede and Ohio
Milford Road

Peninsula

 

L.__

Fig. 1.--Map of Clyde, Michigan, and vicinity
“mmhm location of Duck Lake.

Drain

1(2

g4

 

 

 

 

 

' L
2'54“
Hands
In
I '1 ‘

//

     

‘l 19
{-

nias been directly connected to Duck Lake. Henceforth in
study, this body of water will be referred to as

‘ Reach AL‘. Figure 2 shows the present form of Reach A

based on information provided by the sources previously
cited.

Subsequent to the dredging of a channel to Reach A,
its previous morphology was greatly modified by use of a
drag line to remove deposits which had partially filled its
basin. Reaches A1, A2, and A3 (Fig. 2), formerly separated
from Reach A, were thereby connected to Reach A and thus
returned to Duck Lake.

From Reach A1, the body of water upon which the
psammon transect studied is located, muck three to twelve
feet deep was remOVed until the quartz sand substrate was
exposed. Several large subsurface springs were uncovered
during the dredging at the WSW end of Reach A1 as material
was removed from the basin, piled upon the shore, and
leveled. Table 1 presents the major physical characters
of Duck Lake and Reaches A and A1 at the time of this
study as based upon Figures 1 and 2 and morphometric
techniques outlined by welch (1948).

TABLE 1.--Major morphometric characters of Duck Lake and
Reaches A and A1

 

Duck Lake Reach A Reach A1

—__._
k
M.

Area in sq. miles .456 » .083 .007
Area in Acres 291.8 53.1 4.5
Shore line length 7.9 1.6 .18
Shore line development 3.3 1.6 *

 

 

 

 

L , a ,, ,

 

 

 

 

 

 

20
Reach A
£535... 3“”-
1;: ._-.'-“
A1 ”-74. E:
‘ .lA2 5‘
A3
‘ L
9 0.1 0.2 0.3 0.4 0,5
Fig. 2. --0utline map of Reach A of Duck Lake
wahm location of transect (X).

 

1 2::

.4
. -._.
" '- 3.

.....

 

  
  
   

21
Western Oakland County lies within the Hillsdale-

ciassification is characterized by altitudes of 800 to
:12300 feet and topography of undulating rolling plains,
15in part of arcs of morainic highlands, the substrate being
mainly sandy drift in origin (Veatch, 1953).

Underlying bedrock of Duck Lake and vicinity is
Mississippian, Iowan series, Kinderhook group, Goldwater
ehale formation. Overlaid upon this bedrock is approxi-
mately 300 to 400 feet of glacial drift carried to the
area from the Canadian highlands lying to the northeast
during the Wisconsin stage of Pleistocene glaciation,
deposits of Wisconsin age being superimposed upon debris
0f earlier Illinoian glaciation (Leverett and Taylor, 1915;
Leverett, 1917). The topography of this region is dominated
by glaciation rather than by the overlaid rocks, which are
so deeply buried that they contribute no character to
present day topography although their contours influenced
the course and flow of the ice sheet and deposition of
drift (Bingham, 1945).

} Determination of the origin of the deposits about
Duck Lake is made difficult because of the interlocking
marainic systems formed with advance and retreat of the
Saginaw and Huron-Erie lobes of the wisconsin ice sheet
Which met and combined in this immediate area. The Saginaw

lobe of“ Misconsin flow was southwestward through and beyond

 

 

i
e-m int
I!" am:

..:.,

 

 

 

 

 

   
    

, 22
mggiizua‘w Bay, and the Huron-Erie westward through the Lake
[5-3135 basin into Indiana. At Duck Lake, interlocking flat
Utweeh aprons are associated with moraines of differing
origins.

Reference to surface formation maps (Leverett,
1924; Martin, 1955) indicates that Duck Lake is situated
in an outwesh apron, lying about one mile south of the
southern margin of an extensive glacial till plain or
ground moraine. No streams flow into or out of the lake,
drainage into the lake being that of surface and subsurface
flow from the glacial outwash aprons derived from the
moraines which surround the lake on all sides but the east.
Aridge of glacial till at the margin of this ground
moraine forms the northern limit of the Duck Lake watershed.
Drainage to the north of the ridge is to Saginaw Bay via
the Shiawassee River. Overflow from Duck Lake is via a
‘ II control sluice which drains to the south and west into
Pettibone Creek and eventually southeast into Lake Erie
via the Huron River.

The ground moraine lying north of Duck Lake is
assigned to part of the Ionia moraine system of the
Saginaw lobe (Bergquist, 1932). A mile to the south and
west of Duck Lake, extending from Clyde to Highland roughly
along the route of the Chesapeake and Ohio railroad, is the
northern terminus of a morainic mound which forms the
westernmost limit of the Duck Lake watershed. This moraine

and its outwash apron component are attributed to the

 

 

9": Eu

,f- n

.
...I
'e. n
u
a. :
a.
q.» 1
: .._.
e .‘
..__
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M
I.
'-.
.
- .
. ‘1
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f
.
.g‘.

23

  
  

ation in southern Michigan. The barrow ditches lin-

fixtreme western end of Reach A1.

A morainic mound arcs to the northeast from
Highland to the cemmunity of White Lake, the northern
terminus forming essentially the eastern margin of white
Lake. The southern terminus lies about 1.5 miles south
of Duck Lake and apparently delimits the southern margin
of the Duck Lake watershed. Duck Lake receives water
draining from White Lake, the latter's level being main-
tained at 1018.6 feet above sea level by a control sluice.
This morainic mound southeast and east of the basins of
Duck and White Lakes is assigned to the Defiance System of
the Huron-Erie lobe, the ice apparently having advanced
from the south and east rather than north and west as
was true for the ice movements which deposited the debris
to the west and north of Duck Lake (Leverett and Taylor,
1915). According to Leverett and Taylor, the drift in
this area is largely gravel and sand with only scattered
deposits of clayey till; the outwash aprons being composed,
too, of gravel and sand.

No comprehensive soil surveys are on record as
ever having been made in western Oakland County. Descrip-
tion of the soils in the Duck Lake area is therefore based
"PO" generalizations relative to soil characteristics of the

general area as gathered frOm diverse sources.

 

 

: .1 3m
with
-: Ian! 0

I “'L
rt“:

 

 

 

5’
l

we

24

   

smooth sand and gravel plains, only slightly pitted

or undulating with few lakes or muck basins. Soils of
this land type they characterize as sandy loams with a
clayf-sand hardpan, six inches to several feet in thick-
nass, lying one to two feet below the soil surface. Below
the hardpan is send and gravel. Specifically they assign
the soils of the Duck Lake vicinity to the Fox Soil type.
Veatch (1953) describes the Fox soil type as characteristic
0? glacial outwash flats and glacial drainage valleys.
These soilslhave light-colored surface horizons of friable
sandy loams, a subsurface horizon of sand and gravel

bound into a slightly to strongly coherent mass by reddish
Clay and colloids, a pervious substratum of variably

1 calcareous gravel and sand, and an underlying mass of
interetratified sandy clay, silt, sands, and gravel.

The climate of western Oakland county is subject
to the moderating and stabilizing influence of Lake
Michigan and to a lesser extent that of Lake Huron.

Winters are milder and summers cooler than at like latitude
in Wisconsin and Minnesota. Seeley (1917) listed a mean

‘ annual temperature of 47°F. for western Oakland county

with a January mean of 22°F. and one of 70°F. for July.
Weather Bureau records at Pontiac, the nearest first class

U.S. Weather Bureau station to Duck Lake, give a 32-year

 

 

  

 

25

ghost usually occurring in the first week of October and

the last the second week in May.

Mean annual precipitation in the vicinity of Duck
Lake is approximately 28.5 inches with the precipitation
relatively evenly distributed throughout the year. For
May, the wettest month, the 32-year mean precipitation is
approximately 3.5 inches and that of January, the driest
month, approximately 1.7 inches. From April through
September, 59 per cent of the total annual precipitation
normally occurs.

Prevailing winds in western Oakland county are
from the southwest, the average velocity approximating 8
to 10 mph. In the winter, cold continental air passing
eastward over Lake Michigan, and occasionally westward
over Lake Huron, becomes warmed and laden with moisture,
thus producing unstable cloudy conditions in southeast
lower Michigan. In December and January, the Duck Lake
region receives on the average only 22 per cent of p05-
sibla sunshine. On an average there are only four clear
days per month in December and January, as contrasted to
an average of 11 clear days per month for the period July
through October. According to the U.S. Weather Bureau,
sautheastern Michigan sunshine averages only about 50

Far cent of possible incidence annually.

 

r: tis ii

men III
a! 23:11
3‘. art
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11' he

26

   

It should be noted that abnormal climatic condi-
Vt‘is prevailed in lower Michigan during the period within
{hich this investigation was made. During 1963 the Duck
M.eke area received only 60 to 70 per cent of the normal
annual precipitation, the second consecutive year ‘of‘ below
normal precipitation. In 1964 as well rainfall in the
vicinity of Duck Lake was approximately six inches below
the annual mean of‘ 31 inches. Levels in many Oakland
county lakes reached record lows, some becoming completely
dry during the period October, 1963, to November, 1964.
During this period, Lake Michigan and Lake Huron reached
the lowest levels since record keeping began in the 1860‘s.
Although no meteorological data were recorded at
the transect site during the period of“ study, Tables 2
and 3 record the precipitation occurring daily at the
communities of Highland and Duck Lake, the Former location
3.1 miles SSH! and the latter 5.3 miles ESE of‘ the beach

(Detroit Metropolitan Network Annual Summary, 1963 - 1964),

Description of‘ beach transect

The beach transect investigated From November,
1963, to October, 1964, was located on the northern margin
OF Reach A1 at the Far western end of‘ Duck Lake (Fig. 2),
At the initiation of‘ the study a well-defined white quartz
sand beach extended For 0.12 miles along the northern
margin of" Reach A1. In width the landward margin of‘ this
beach varied from two to six meters distance From the

Shoreline. Using descriptive criteria developed by Veatch

 

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29

   

she Humphrys (1955), the beach is of the low shore, hard

fifh‘ore, sand beach type. Page 19 outlines the manner in
which the morphometry of‘ the beach was formed about the
. I margin of Reach A1 in the winter of 1958-1959.

The southern margin of Reach A1 was also a sandy
beach during the study, similar in width to that of the
northern margin (Fig. 2). The western limit of Reach A1
during the growing season was a marshy area, characterized
by a thick, massed grOwth of emergent, rooted hydrophytes.
A shallow sill extended across the channel connecting
Reaches A1 and A2 at the point of narrowest constriction.
During low water periods in July, August, and September
this sill was exposed, separating Reaches A1 and A2 from
the main body of Duck Lake.

The site of the transect was selected with refer—
ence to homogeneity of slope, Freedom of area from
macrophytes and surface organic deposits, relative homo-
geneity of sand grain size, nature of slope, and surface
0? the hydropsammon zone sand surface in the event of
water level regression. Upon selection of the site,
one piece of one—inch copper tubing was driven two feet
into the sand at the shoreline. Another rod was placed
300 cm. landward and one BOO cm. lakeward from the zero
Delhi: to form a straight line, perpendicular to the shore-
line, between the three index points used as references
for all subsequent linear measurements made during the

study. All linear measurements made along the index line

 

 

 

3E!

    

_ dward from point Zero position and negative numbers
to stations lakeward from the original point zero.
“ The general topography of the area making up the
northern margin of Reach A1 suggests that the slope
investigated lies on the extreme flank of a morainic
mound originating approximately 100 meters ENE of the
transect and terminating just short of the western limit
of Reach A1.

Inasmuch as the linear dimensions of the transect
did not warrant use of beach slope measurement techniques
such as those described by Emery (1961), a temporary
wooden framework was constructed with reference to the
1 } three index points, the horizontal member being positioned
by means of a bubble level. A plumb line was used in
‘ measuring distances from sand surface to Framework
‘ horizontal member every 12.5 cm. along the transect index
line. The process was repeated for a parallel line 10 cm.
Either side of the index line and the three readings
averaged for each 12.5 cm. location. Figure 3 presents
the beach profile thus determined.

A number of ‘12‘ x 2 x 2 inch pine wood blocks
were placed on the sand surface above shoreline at in-
tervals of 25 cm. along the index line, beginning at
+12.5 cm., in order to determine the extent of wave action

between collection dates. As the lake level retreated,

 

—t\r-lI-I. .I....:e .-..~

 
  
 

 

 

 

 

 

31

 

 

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mm mm D mNI omI me earl
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mm.

 

 

 

.. Erin;

.‘1I1'. the }
-:.': it II

‘3: -“.2

32

  

:Tbitional blocks were added in the former hydropsammon
'w‘ne. During the study, the blocks were found dislodged
only on the April 6 and May 6 collection dates. On

April 6 it was found that all blocks from +12.S cm.
through +112.5 cm. were missing, and there was visual
evidence of churning and plowing of the sand up to the
+125 cm. mark. This activity was apparently related to
the break-up of the winter ice cover. It would appear
that at no time other than in March-April did wave action
influence the beach at any point 12.5 cm. or more above
the then current shoreline.

From November through March the level in Reach A1
remained unchanged. At the April 6 collection, however,
the shoreline was found to have advanced to the +30 cm.
mark and to the +40 cm. mark on May 6. In June the lake
had receded to the -30 cm. mark, in July to ~50 cm. and
in August to -225 cm. mark. Heavy August rains returned
the level to the -175 cm. mark in September. By October,
the lake level had retreated to -250 cm. mark.

In November of 1963 Reach A1 offshore frOm the
transect was characterized by an almost pure population
Offl'lfl‘a vulgaris L., beginning at the -125 cm. mark and
extending in an arch around the western end of Reach A1
t0 appear in similar extent on the southern margin of
the body of water. This growth flourished throughout
the winter under the ice, but was almost completely

eradicated in the March-April ice breakup, after which

 

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:w. 31

My. M1
'2: ball!
air; I! 9
rain! t!
M E 3::
5?! :r‘t
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33

   

In. July, Auguet, September, and October, however, the

 

Chara thalli became covered with a thick mucilaginous
coating of Phormidium angustissimum N. at (3.5. west to

the point that almost the entire surface of Reach A1

became a brown, decaying entangled mass of' Chara fragments
at the center of heavy sheaths of B. angustissimum and

numerous cyanophytes entrapped in the Phormidium mucilage.

 

From November through May the beach in the vicinity
of the transect was practically devoid of macrophytes,
but by June a relatively thick growth of Juncus acuminatus
Michx. and Juncus bufonius L. had developed in a band
roughly 100 cm. wide between 100 and 200 cm. inward from
the November shoreline. Scattered individuals of
Ambrosia elatior L. and Erigeron ramosus Watt.) 8.5.9.
also appeared. This development of macrophytes, however,
did not intrude into the transect collection zone.

An ice cover ranging from 10 to 25 cm. in thick-
noes covered Reach A1 from December through March. The
sand of the beach itself was frozan into an icy concretion
from December 8 through March 6. In December the lake

was covered by a layer of ice 10 cm. thick, and the sand

 

Of the entire transect from point zero to +300 cm. was
frozen except for the surface three cm. Ice 18 cm. thick
covered the lake on January 4, 1964. On that date ten cm.

0f snow covered the beach, but the top three to five Cm.

 

 

   

. . 34

:599end beneath were friable, quickly solidifying when
in snow cover was removed. The sand had a distinct

‘ blue-green tinge from the +150 to the +300 cm. mark. On
February 5 ice cover on the lake was 25‘ cm. thick, and
thesamiwas solidly frozen except for the top two cm.
Again-the beach sands between the +150 and the +250 cm.
mark were distinctly blue-green. A layer of frozen sand
one cm. thick lay one cm. below the surface of the beach
unmard16, this layer being two to three cm. above the
comfletely frozen layer of sand parallel to the surface.
Tm1cm. of ice covered the lake on the March 6 collection
(hte. On each of the December, February, and March
mfllections the sand was completely frozen until the
mmface few cm. of the south-facing slope thawed around
mid-day. By late afternoon the sand was again firmly

\
l
‘ frozen.
\
\

 

 

 

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.
.._
.‘-t-
. ..

 

    

 

CHAPTER IV
METHODS AND MATERIALS

Granulometric fiscription of‘ the beach

m

 

Mechanical analyses of‘ approximately 150 grams of‘
incinerated sand collected From the surface three cm.
were made For each station along the transect index line
at 25 cm. intervals beginning at the -‘IUU cm. and con-
tinuing through the +300 cm. mark. (See page 29 For
description of‘ transect site). Mechanical analysis For
sand grade distribution was made Following standard pro-
cedures as outlined by Kilmer and Alexander (1949), Passing
the gross sample through a 2.5 inch U.S.D.A. mechanical
analysis sieve set having standard U.S. sieve series
components of” 18, 35, 60, 140, and 300 mesh (U.S.D.A.
Bull. #170). Each sample was subjected to 15 minutes of‘
agitation on a lateral shaker operating at 1140 reversals
par minute. Percentage of‘ gross sample retained by each
screen was calculated and data plotted as a grain-size
accumulation curve in order to determine harmonic mean
grain diameter, grain size distribution, ef‘f‘ective grain
size, and uniformity coefficient at each 25 cm.- interval

following methods of Heugh (1957).

35

 

itinhy

1* a 53‘
3'3. in
it‘s-g an:
if. 131
E1 can
I: 32:93

'n: 4‘

   

36
Porosity was determined by standard volumetric

procedures (Hough, ibid.). Standard soil porosity deter-

 

minations by means of volumetric procedures were modified
in that a soil sampling ring 7.6 cm. in diameter and

2.54 cm. in depth was driven into the sand surface, and
the ring and 113.4 cc. of sand contained therein removed
without disturbing the sample by insertion of a sheet
meta13quare along the lower surface of the ring. The
upper surface of each ring and its contents were then
covered with a No. 50 Filter paper disc and cheesecloth
and then inverted in distilled water for 24 hours to
saturate the sample. The sample was next weighed quickly,
any water draining onto the pan during weighing being
included in gross wet weight. Finally the sample was
placed in a 103°C. oven for 24 hours, after which the
sample was again weighed. Wet gross weight less dry
gross weight gave void volume. Weight of water per unit
sample at saturation was also given by this procedure.
Volume of the ring was considered total soil volume in
calculations. Six samples were taken at each 25 cm.
interval, three on each side of the transect limits, and
the porosity values averaged.

Unit weight or weight per unit volume of the sand
at each 25 cm. interval was computed as the mean of six
samples following standard methods (Russell, 1949). This
method essentially involved determination of even dry net

Night of sand sample divided by the sample VOIUIHB, 113.4 cc.

 

J— ,,\,7_r ii ‘

 

51. Baal?

3.1. east

i:. h at

so,

5 ' :5
Ti? 3

 

37

     

1964. Parallel to the index line of the transect and

100 cm. east of it a series of holes approximately 30 to
40 cm. in diameter were dug vertically into the sand at
marks -100, ~50, zero, +50, +125, +180, and +300 cm.

These holes ranged in depth from 75 cm. at the -100 cm.
mark to 180 cm. at the +300 cm. mark. The symbol "0" on
Figure 3 indicates the approximate depth in each hole

at which a stratum of coarse gravel and heavily calcareous
coarse sand was encountered. Above this substrate, size
and nature of the sand particles at each hole were more

or less those exposed on the beach surface. Reference to
Figure 3 indicates the upper surface of the coarse
calcareous deposits to be quite irregular. In the +300 cm.
hole at 15 cm. a thick deposit of very dense clay was

found superimposed upon the calcareous substrate.

The depth of the phreatic surface below beach
surface was recorded for each hole at regular intervals
beginning in early July and continuing through September.
Each hole was covered with plywood and sand to reduce
eVaDDI‘ation and increment of windblown sand and debris
within the holes. It was found that stabilization of the
Phreatic surface with varying lake levels occurred in
approximately 48 hours, fluctuation after 48 hours being

five per cent or less.

38

   

. Figure 3 shows the phreatic surface levels beneath
‘Ve beach surface for three series of measurements made

' I over 52-hour periods during which the lake level was at
-75, -150, and -275 cm. respectively. Interestingly, it
is seen that the phreatic surface in each series lies
considerably below the lake level prevailing during each
series of measurements and that the phreatic surface
distance to the beach surface is essentially the same frOm
the +25 to the +300 cm. mark over a relatively wide range
of lake levels.

Unfortunately phreatic surface depth measurements
were not made during the period November - May when the
lake level stood at or above the zero mark. Thus it is
not known whether the phreatic surface profiles constructed
on Figure 3 are illustrative of conditions prevailing

throughout the study.

Physical data
T Depth of light penetration into the sand was

determined by utilizing a 4x 5" press camera magazine

‘ with a detachable glass face. The film magazine was in-
serted'horizontally into the sand at depths below the

‘ surface of 0.5, 1.0, 2.0, and 3.0 cm., frameworks of these
1 thicknesses being temporarily attached to the magazine

to maintain accurate and even distribution of send over
the glass face. Fine grain positive film was exposed at
each depth for two minutes, two series of exposures being

made at the zero, +150, and the +300 om. marks.

 

 

T
m! sw’

Eaters!
e and

11'. :33

‘53::-
‘II ‘

 

39

   

Temperature readings were determined at 25 cm.
'ervals along the beach transect index line by thrusting
1;; the bulb of a corrected laboratory thermometer just below
the sand surface, any readings thus being illustrative of
temperatures in the top 1.5 cm. of sand. Air readings
were recorded at the +50 cm. mark at breast height and
one cm. above the sand surface. Lake bottom temperatures
were taken just below the sand surface at the most landward
point along the transect index line at which a horizontal
linear differential of 20 cm. occurred between lake bottom
and lake surface. Lake surface temperature was also
recorded at this point.

Visual observations of the "black layer" were
made each month that the sand was not frozen, and its
vertical and horizontal extent plotted on graph paper.
The "black layer" has been described by Neel (1948) as
that portion of the sandy beach in which anaerobic de-
composition of organic material has resulted in reduction
01’ iron oxides by sulfides to iron sulfide. A trench
20 cm. deep was dug from waterline to the +350 cm. mark
Parallel to and 50 cm. distant from the base line. The
trench was located on alternate sides of base line each
month to reduce beach disturbance.

Methods utilized by Pennak (1940) were modified
to determine percentage of water content and total
Organic content of sand at each 25 cm. interval along

the beach transect index line at each collection date.

 

exist:
:3. if a

E's“:- 11‘

'Illhc c]

Inn—l I

':' tie

a sigh

l'itatl

40

   

Approximately 20-gm. samples were removed from the top

two cm. of exposed sand at each interval and immediately
sealed in labelled plastic vials. Following transfer to
porcelain crucibles and determination of gross wet weight,
each sample was evaporated to oven dryness to obtain dry
'gross weight. Each sample was then incinerated over a
burner at low red heat for one hour to determine loss of
volatile matter (i.e. an approximation of contained

1., 194:5).

organic matter according to Theroux e_t.
Dissolved volatile matter (i.e. organic matter)
was determined for lake water and interstitial water by

modification of methods recommended by Theroux _e_l:._ _a_1.

(flJ. 0ne hundred—ml. samples of lake and of inter-
stitial water filtered through a 0.45 micron Millipore
filter were evaporated to oven dryness in porcelain
crucibles. Following gravimetric determination of total
dissolved solids, the samples were ignited over a burner
at low red heat until the ash became white. The crucibles
were then cooled, weighed, and loss of volatile matter

COmpUth.

Chemical data
Interstitial water was drawn from the hygropsammon
zone by means of an aspirator. The tip of the glass
SUPply tube was thrust vertically into the sand to a
depth of four cm. and slurry drawn into a 150 cc. suction

flask, the supernantant fluid being decanted and retained

i until 500 to 600 cc. of water had been obtained. To

 

 

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.h‘ I.
:. .- _
.Ie' :-
In“...
.~..: _
n'.
“anal
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r.
"1. -‘El
."4
':- .
'I... '

41

   

several points along a line 50 cm. landward and parallel
to the shoreline.

Interstitial water sampling began in April and
continued to the October collection, the sampling site
migrating along the transect index line as the lake
level advanced or retreated. In monthly sequence, the
sampling points were as follows: +80, +50, +20, zero,
-75, -100 and at -100 in October, the shoreline in each
case being 50 cm. lakeward except in August, September,
and October when it was 200 cm., 75 cm., and 150 cm.
distant respectively. In August, September, and October
no water could be obtained directly from the sand with
the aspirator. A sump 10 to 20 cm. deep was dug at the
sample point and the water collecting therein withdrawn
with the aspirator.

Lake water samples were taken by surface dipping
0f one liter of water along the transect index line
offshore approximately five meters.

Dissolved oxygen in lake and interstitial water

‘ was determined using the Alsterberg modification of the

1 Winkler method as described in W Md; f_o_1_~

r Examination o_f Ina—ta; _a_n_d_ w_aiLe ‘Jd_at_e_r (1960). Titration

‘ with .025N phenylarsene oxide was conducted in the labora-

tory, preliminary steps being performed in the field.

 

is

re use 1'

r:- :utli

r 211
as 1252:
in“.

- em":

21:: 5-1!

42

Dissolved carbon dioxide determinations for lake

  

water were made in the field according to the titrimetric
method outlined in Water Analysis Procedures, Cat.’ #8
(Hach Chemical 00.). A 200 ml. sample was utilized for
lake water determinations. Inasmuch as high turbidity of
interstitial water made colorimetric and titrimetric
methods subject to considerable error because of difficulty
in detection of end points, the method presented in
Theroux it Q. (1943) was utilized. CompariSOn of monthly
data with those obtained by use of the carbon dioxide
nomographic method as presented in Standard Methods
(22. fl') gave results with less than 10 per cent
difference. It is data obtained by use of the Theroux
method that is presented in Table 7.

For all chemical determinations other than those
for dissolved gases, interstitial water samples were
passed through a Millipore filter apparatus utilizing a
filter of 0.45 micron apertures. A Bauch and Lomb

Spectronic 20 spectrophotometer calibrated for reagents

used in Hach procedures was used in all colorimetric
determinations. Calibration curves were constructed using
standard solutions recommended by Standard Methods (1960).
‘ In the chemical determinations listed below,

1 methods presented in water Analysis Procedures, Cat. #8
(Hack Chemical 80., 1963), were employed using a 8. 8: L.

w Spectronic 20 as a colorimeter.

 

 

5
7

43

1. Alkalinity

2. Calcium hardness -- Calver II modification of
standard EDTA

3. Total hardness -- Monover modification of
standard EDTA titration

4. Total ferrous and ferric iron -- 1—10

phenanthroline colorimetry

 

5. Nitrate nitrogen -- Brucine calorimetry as
recommended in Vol. 7 Water Analysis Procedures
(Hach Chemical Co.)

6. Nitrite nitrogen -- NitriVer modification of
sulfanilic-l-napthylamine colorimetry

7. Orthophosphate -- stannous chloride calorimetry

8. Silica -- sodium sulfite reduction calorimetry

9. Sulfate -- barium sulfate turbidimetric method

The pH of lake water and beach interstitial water

wascbtermined through utilization of a Beckman 180

portable pH meter. From November through to May the pH

Ofthe sand at 25 cm. intervals along the transect index

lineums obtained by inserting the tip of the electrode

Prmm into the sand to a depth of one cm. From May to

Bummer this method was supplimented by additional field

dauaobtained following standard soil pH determination

tedmiques. This method involved the mixing in a beaker

DfZU cc. of sand from each 25 cm. interval with 20 cc. of

distilled water adjusted to pH 7.0 and subsequent deter-

mhmtim1of the pH of this slurry by insertion of the pH

meter probe.

 

 

 

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5‘3‘

44

For correlation with pH determinations along the

  
  
  
  
  
  
 

transect, alkalinity determinations were also made at

each 25 cm; interval. On each collection date, a plastic
vial was filled with sand from each station and tightly
capped. In the laboratory 5 gm. aliquots were added to
distilled water previously adjusted to zero alkalinity and
alkalinity of the sample determined following procedures
used in determination of interstitial and lake water
alkalinity. The remainder of each sample was oven dried

and alkalinity determinations conducted as above.

Biological data

3 Qualitative phytoplankton lake samples were taken
with a No. 20 bolting silk plankton net from a point
approximately 10 meters offshore from the beach transect.
Twelve liters of water for quantitative plankton study
were dipped from the same point prior to qualitative
sampling. For subsequent quantitative study, an aspirator
tube provided with a four-ply filter of No. 20 bolting
silk at the inlet was used to concentrate two liter
aliquots to 500 cc. Each 500 cc. concentrate was then
reduced to 10 cc. volume by means of filtration through
a 0.45u Millipore filter.

One 20 cc. qualitative sand sample was taken each
CUllection date from the upper three cm. of beach sand
at each 25 cm. interval along the transect index line.

R8 was the case with quantitative sand samples, sampling

 

';

45

    

samples being taken 10 cm. east of the index line, January
samples 10 cm. west of the line, February samples on the
line, etc., so that three months elapsed between successiVe
samples at the same site.

A copper tube of 1.25 cm. internal diameter was
inserted two cm. into the sand to obtain a 2;4 cc. quan-
titative psammon sample. By inserting a sheet metal
strip at the lower end of the tube, the tube and undis-
turbed sand core within could be withdrawn from the beach.
The core was then transferred directly to a six dram vial
by flooding the tube interior with distilled water over
a glass funnel directed into the vial. Three quantitative
samples were taken each collection date at each 25 cm.
interval.

All lake and beach quantitative samples were
preserved in the field with FAA solution as were all but
one lake qualitative plankton sample. A 10 cc. aliquot
0? each 20 cc. qualitative sand sample was preserved in
FAA within 24 hours after collection, the remaining
Portion of each being placed in a 250 cc. Erlenmeyer
Flask which had been previously autoclaved with its contents
0? 100 cc. of 50 per cent Knop‘s solution. These cultures
were incubated at approximately 20°C. under fluorescent
illumination of 200 to 400 f.c. for a maximum of six
munths, after which time the culture was preserved in FAA

salution. Twenty cc. of lake qualitative plankton samples

 

 

 

 

 

 

I. “
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. y.
..‘

."I.
I“.

 

46

    

cultures were examined periodically during taxonomic
‘ investigations.‘

Separation of the algae from the preserved quali-
tative and quantitative sand samples was accomplished
following the method recommended by Standard Methods (1960)
in describing the Sedgwich-Rafter sand filtration procedure.
Thorough mixing of the sand included three minutes of
stirring with a fine stream of compressed air delivered
through a one ml. pipette. The three minutes of stirring
were divided into six 30 second periods and six decanta-
tions. The approximately 250 cc. of water and suspended
material obtained by repeated stirring and decanting of
the diluted 2.4 cc. sand sample were filtered through a
0;45u Millipore filter, and the residue brought up to a
standard 5 cc. volume in 10 per cent FAA solution. This
standard 5 cc. volume was diluted when above average
quantities of detritus and silt were encountered.
Quantitative counts of plankton algae were made

using the Drop-Sedimentation method as given in Standard

 

Mods (i_t1i_d_.). Quantitative counts of lake plankton
were made at 675x, all algae present in 1/7 cc. aliquots
0f 10 cc. concentrates of the standard two liter samples
being counted under 25 cm2. cover slips. Three mounts
were made for each sampling date, the results averaged,
and the data for the average 1/7 cc. aliquot extrapolated

to number per liter.

1\ ,— ,

 

 

47

For each quantitative psammon sample, 1/7 cc.

  

aliquots of the 5 cc. concentrates were mounted under
25 cm2. coverslips and counts made at 675x. Each count
involved counts of all organisms present in the 1/7 cc.
sliquot.‘ The resultant counts for each of the three
quantitative samples taken monthly at each interval were
then averaged and the mean number per 1/7 cc. aliquot
extrapolated to number per cc. of sand sample.
Construction of quantitative and qualitative
lists for lake plankton and psammon intervals on each
collection date involved preparation of permanent slides

of acid-cleaned diatom cemponents of the microflora.

 

  
 

 

 

CHAPTER \I

RESULTS

  

General information

Prior to the Duck Lake beach transect investiga—
tion, a previous transect study of the psammonflora of a
beach on Big Portage Lake, Jackson County, Michigan, was
prematurely terminated after six months because of disap-
pearance of the beach transect as a result of ice mOVe-
ments. In the Duck Lake study, twelve series of col-
lections were made at approximately monthly intervals
during the 12-month period of investigation initiated on
November 7, 1963, and terminated on October 11, 1964.
See Diggiption of 921°" transect for description of
meteorological and climatological conditions occurring
during the investigation period. See also Chapter IV for
methods utilized in obtaining physical-chemical and
biological data for the beach transect and adjacent lake
at each monthly collection. Although core collections
were made at each 25 cm. of exposed beach from shoreline
to the +350 cm. sampling station, in this paper biological
data is presented for only the lake and the +300, +200,

+100: and +25 cm. beach transect intervals.

48

 

 

 

 

49

   

Granulometric characteristics
of the transect substrate

Table 4 presents a summation of determinations
of grain dimension percentage composition in incinerated
sand samples as based upon retention by U.S. sieve series.
With the exception of samples collected at the +25 cm.
mark, samples collected at each of the 25 cm. intervals
along the transect were composed primarily of sand
granules lying in the range of .25 to .10 mm. maximum
dimension. Such a distribution as evidenced in Table 4
would categorize the beach studied as fine sand with
reference to criteria selected by the American Bureau
of Soils and the International Soils Association.

Harmonic mean grain size diameter (Mavis and
Tsung-Pei, 1939) ranged from .148 mm. at the +300 cm.
Mark to .085 mm. at the -75 cm. mark with a harmonic
mean of .109 mm. for grouped samples run independently
(Table 5). The arithmetic mean of the individual harmonic
means was .115 mm. Thus on the basis of criteria selected
by the American Bureau of Soils (Russell, 1949), the
harmonic mean grain diameter determinations would indicate
that the substrate of the transect as a whole COUld be
characterized as fine sand, stations +25, -50 and -75 cm.
as Very fine sand.

Ef‘l’ective grain siZe, defined by Hazen as the
grain dimension of a soil sample less than that of 90 per

cent of the sample and more than 10 per cent of it

 

 

 

r.
7.51:“. ,‘

  

50

.‘~-.ABLE 4.--Percentage composition 150 gm. incinerated beach
-i‘ansect sand samples by dimension based on U.S. sieve series
For intervals -100 to +300 cm.

 

 

Per cent of total sample

 

larger larger larger larger larger larger smaller
Station than than than than than than than
2.36mm. .991mm. .495mm. .246mm. .104mm. .048mm. .048mm.

 

+388 1.57 4.98 15.88 29.27 43.89 3.78 8.81
+275 .58 9.88 13.15 22.84 45.79 7.89 8.88
+258 2.29 5.45 12.32 28.58 45.85 7.85 8.48
+225 .48 1.89 9.45 35.88 48.83 8.48 8.83
+288 1.89 1.27 13.18 21.93 51.48 18.38 8.22
+175 1.78 3.18 8.88 23.17 52.51 9.95 8.84
+158 2.22 3.33 8.85 18.45 51.82 14.28 1.25
+125 7.59 4.51 18.38 19.81 44.88 12.72 1.12
+188 2.59 4.17 7.58 19.17 58.88 14.79 1.11
+75 4.39 4.49 8.79 28.81 49.98 11.11 8.88
+50 4.88 4.79 18.58 18.78 47.11 12.83 1.31
+25 1.87 2.22 5.88 14.48 32.85 41.38 2.38
0 1.72 2.48 15.93 13.88 52.23 14.48 8.21
-25 8.77 1.28 12.77 18.77 52.75 13.57 8.18
'50 2.81 1.74 5.87 15.55 53.98 18.99 1.88
~75 8.48 8.89 18.54 7.82 37.87 38.78 8.71

”on 0-04 0.34 18.37 8.15 59.88 14.87 0.38
Arith- M

mgfic 2.13 3.33 11.39 19.49 48.12 14.80 0-74
Ban

 

 

 

 

E 5.--Granulometric

 

51

u

data for beach transect intervals -100

 

 

 

 

+300 cm.
harmonic tJniForm. wt. HOH volumet. unit
mean D60 D10 coeffic. at sat. porosity weight
grain D/60/D10 as 7. as o in
dia.inmm. _ gm/cc.
+388 .148 .315 .114 2.75 21.2 41.3 1.58-
+275 .137 .298 .118 2.84 21.8 42.5 1.52
+258 .134 .283 .118 2.57 22.9 44.1 1.45
+225 .132 .283 .123 2.13 21.4 43.8 1.88
+288 .119 .243 .184 2.33 22.3 43.7 1.43
+175 .119 .235 .182 2.38 23.3 44.3 1.55
+158 .118 .221 .888 2.51 23.5 47.2 1.45
+125 .123 .288 .895 2.74 24.4 48.4 1.35
+100 .110 .215 .898 2.39 24.3 48.3 1.28
+75 +120 .237 .188 2.37 28.8 48.5 1.31
+58 .119 .243 .891 2.87 25.8 45.9 1.23
+25 .879 .158 .878 2.14 23.9 43.5 1.38
0 .111 .‘289 .891 2.38 27.2 58.8 1.35
‘25 .118 .218 .898 2.33 23.8 45.8 1.51
'5” ~096 .175 .874 2.35 23.8 58.3 1.69
I ~75 .885 .184 .881 2.89 24.8 48.7 1.48
‘100 .184 .158 .884 1.98 35.8 82.8 1.18
NM
Total
.894 2.43 24.4 45.8 1.43

 

mean .115 .223
\

 

 

 

52

(Hough, 1957), as determined From individual grain size
accumulation curves For incinerated sand From each 25 cm.
interval along the transect ranged From .123 mm. at
+225 cm. to .061 mm. at the -75 cm. mark. The mean comput-
ed FrOm the individual ef‘f‘ective grain size at each 25 cm.
interval was .094 mm. Such Figures are comparable to
those reported by Hough For clean Fine river sand. An
overall tendency For eFFective grain size to increase
with distance From shoreline was apparent.

UniFormity coeFFicients, def‘ined by Hazen (in
Hough, Egg.) as the ratio between the grain size Finer
than 40 per cent 0F the material and the grain siza
Finer than 90 per cent 01“ the material, 01" 150 gm.
incinerated samples 0F sand collected at each 25 cm.
interval along the transect ranged Frorn 2.76 at the +300 cm.
mark to 1.90 at the -100 cm. mark. The mean cemputed
From individual uniformity coef‘Ficients was 2.43. A
grain size accumulation curve (Figure 4) constructed on
the basis of arithmetic means oF individual stations
percentage composition Figures gave an overall uniformity
coeFFicient oF 2.70 For the transect substrate. Such
Figures are similar to those reported by Hough (En—d.) f‘Or
Fine or Ottawa send (a commercially processed silica
sand with a uniformity coeFFicient of‘ 1.1). Hough re-
Ported uniformity coeFFicients For Ottawa sand, clean Fine
sand, and silty sand 01" 1.1, 4 to 6, and 15 to 300 respec-

tively. Ordinary beach sands he reports as having a

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tiniformity coefficient ranging From 2.0 to 6.0. A
uniformity coefficient of 1.0 indicates homogeneity of
grain size in the sample. A uniformity coefficient of

more than 1.0 indicates relative heterogereity of grain
size. Reference to Table 5 indicates no significant
difference in homogeneity of the substrate at each 25 cm.
interval except For station -100. At the -100 interval
grain dimension exhibited a conspicuous tendency toward
uniformity as compared to that exhibited by grain dimen-
sion at the +300 interval. Thus it would appear that

in general, for the beach studied in this investigation,
dimension of interstitial spaces between sand grains would
be more uniform the closer samples are taken toward the
normal shoreline, becoming more irregular the Further
landward.

1 I Factors responsible For grain dimension sorting
upon the beach apparently produce a relatively homogeneous
distribution within samples collected at each regular
interval along the transect, although the harmonic mean
Grain size diameter and effective diameter increase
gradually with distance from normal shoreline. Thus a
relatively invariable uniFormity coefficient accompanied
by increasing harmonic mean grain size diameter and
effective grain size landwards indicated an overall
tendency for all grain diameters to increase with distance
i from the beach.

1‘ The amount of water present in non-incinerated

saMPlos at saturation and expressed as percentage of wet

 

‘l— 27 ,7 2

55
gross weight ranged from 21.2 per cent at the +300 cm.

  

mark to 35.0 per cent at the ~100 cm. mark. Mean value
for determinations made at 25 cm. intervals along the
transect was 24.4 per cent, a value approached most
frequently at the mid-point of the transect. Sand samples
taken at those int-Wale closest to the normal shoreline
showed a greater water retention capacity than those
located further inland. Maximum water holding capacity

for quartz sand has been reported by Hough (ibid.) as

 

28.3 per cent, for 4/5 sand and 1/5 peat as 47.8 per cent,
‘ I and for 1/2 sand and 1/2 peat as 89.1 per cent.
i I. In general water retention capacity of samples
taken along the transect paralleled closely the porosity
0f the sand samples as determined volumetrically. Lowest
volumetric porosity was indicated at the +300 cm. mark,
highest at the -100 cm. mark, the values being 41.3 and
52.6 per cent respectively (Table 5). A tendency for
Porosity to increase from most landward point to most
lakeward point along the transect was apparent. Mean
Porosity along the transect was 46.8 per cent, roughly

that of silty sand as reported by Hough (ibid.). Accord-

 

ing to Savor (1959), porosities in the ranges reported
above are characteristic of silt and clay and are far in
excess of values characteristic of fine sand, which Bauer
indicates as 35 per cent.

Density or volume weight for sand samples col-

lected at 25 cm. intervals along the transect ranged

 

 

   

 

‘ 56

1.10 g/cc. at the -100 cm. mark to 1.69 g/cc. at
-50 cm. mark. In general there was a tendency For
'.;B'lume weight to increase the Further the sampling point
was located away From the shoreline. Mean volume weight

for massed samples was 1.43 g/cc.

Beach physical data

Following procedures outlined in Chapter IV,
two series of‘ positive Film exposures were made in full
sunlight at horizontal depths beneath the beach surface
of 0.5, 1.0, 2.0, and 3.0 cm. The two series of exposures
were conducted at the zero, +150, and +300 cm. intervals
along the transect. Not one of’ the six plates exposed
at 3.0 cm. depths exhibited any evidence of" fogging, nor
was any evidence of‘ light penetration visible on any of
the six plates exposed at 2.0 cm. depth. Una each of’
the two exposures made at 1.0 cm. depth at the zero,
+150, and +300 cm. intervals showed some evidence of
faint fogging, apparently in each case around a pebble
01' small stone embedded in the one cm. of‘ sand above
the plate. All six plates exposed at depths of 0.5 cm.
below the beach surface showed evidence of‘ slight Fogging
distributed in irregular random patches over the plate
surfaces. 0n the basis of‘ this rather limited investiga-
tion it would appear that effective light penetration
into the beach sand was semewhat less than 0.5 cm. With
some random penetration in heterogeneous deposits to

maximum depths of‘ one centimeter. Thus it might be

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57

  
   
 
   
   
  
   
   
 

r ofs-tu1ated that photolithotrophs would be concentrated
“in the top 0.5 cm. of sand with perhaps a Few stragglers
active at depths of up to 1.0 cm.

Table 6 presents the record of‘ temperature
determinations made at the transect site during the twelve-
month period of sampling. Air temperatures encountered
during the study ranged From a high of 27.0%. at the
September collection to a low of 0.508. in January.
Temperature differentials between air temperature at
ground level and breast height level at the +50 cm.
interVal did not exceed 4°C., the maximum differential
occurring at the June collection. Generally air tempera-
tures at ground level were somewhat lower than those at
breast height except For the months of‘ July and August.

l The relatively f‘ew temperature data negate attempts to
identify evidence of orderly relationship between air
temperature at breast height and ground level, surface
water temperature, bottom temperature, and hygropsammon
zone temperature at the sampling positiOn.

Temperatures recorded at sampling stations along
the beach transect ranged From U.0°C. recorded at all
stations in January and February to 35.00C. at the +275 cm.
mark in September. Here too a paucity of‘ data precludes
detailed observations concerning relationships of‘ tempera-
tures along the transect with those of‘ the overlying
air and the neighboring lake. However, reference to

Table 6 reveals a strong tendency For temperatures of

 

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59

  
  
  
  
  
  
  
  
  
  
  
  

the meet lakeward stations to approximate within three
at degrees C. that of the lake surface, with those most
landward approximating within three degrees [2. breast
height air temperature. Only in July, August, and
September did temperature at the most landward point
markedly differ from that of air temperature, being
respectively 6.5, 7.5, and 7.5 CO. higher on those monthly
collections. Temperature variation between the most
lakeward and the most landward stations on the transect
ranged from zero in January, February, and March to
maxima of 6.000. and 5.505. in April and September respec-
tivaly. Temperature variation along the transect tended
to be lowest during the winter months and greatest in
spring and fall.

Observations of the "black layer" could not be
made in December, January, February, and March because
0? the frozen condition of‘ the beach. There was no
evidence of’ a black layer beneath the beach surface to
a depth of 20 cm.’ in November, April, May, or June. By
July, however, a lens-shaped black layer about 2 cm. in
maximum thickness was found seven to eight centimeters
below the surface and extending From the -30 to the
-30 cm. interval along the transect. Another ill-defined
lens-shaped black layer about one cm. in maximum thicanSS
was also observed in July extending From the +30 to the
+125 cm. interval about two centimeters below the surface.

In August, on the opposite side of the transect, no black

 

 

 

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60
layer was observed except For small patches about two cm.
thick lying at depths of three centimeters below the
surface between the +20 and the +70 cm. interval and
again at the -20 to the ~75 cm. interval. In September
there was again no continuous black layer. Isolated
lenses three to five cm. long and one to two cm. thick
lay at depths from one to eight cm. below the surface
from the +30 to the +70 cm. marks. Beginning at the
-50 cm. interval and six to eight cm. below the surface,
a narrow black layer two cm. thick ran parallel to the
surface out to ~17S cm., the shoreline on that date.

In October no black layer was evident except
from the -50 cm. interval to the -250 cm. shoreline. A
layer two to three cm. thick extended from -50 to ~15!) cm.
parallel to the surface and about six cm. below it. At
the ~150 cm. interval the band abruptly widened to the
bottom of the trench. An uninterrupted layer thus
extended from the ~150 cm. to the -250 cm. mark with an
upper margin parallel to and about six cm. below the
surface and extending downward to a depth of‘ at least
20 cm. Figure 5 presents diagrammatically the extent of’
the black layer during those months it was observed.
I Table 7 presents results of‘ determinations of‘
percentage water content on a gross wet weight basis for
each of the 2'5 cm. intervals along the transect from
November through October. Percentage water content

ranged f‘rOm a low of‘ 0.01 per cent on August 6 at the

 

 

       

 

 

 

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63

+300 interval to 49.18 per cent at the +25 cm. interval
on December 8. Twelve-month mean percentage water content
ranged from 3.00 per cent at the +300 cm. interval to
24.57 per cent at the +25 cm. interval. With the excep-
tion of the +250 cm. interval, a direct relationship was
apparent between the twelve-month mean percentage water
content and distance from shoreline. with few exceptions
this relationship occurred as well at each monthly col-
lection period. No abrupt differential was obvious
between adjacent intervals which would suggest the most
landward transect point substrate was not beyond the
limit of interstitial water movement by capillarity.
The mean percentage water content of‘ interval +250 higher
than that of both adjacent intervals was not obvious each
collection period. In December percentage water content
was lower at interval +250 than both adjacent intervals,
and in November, January, April, and August it lay
between that of both adjacent water levels.

Following relatively heavy precipitation immediate-
ly prior to the collection date in march, April, June,
and July it was seen that the maximum percentage water
content along the transect was not only 25 cm. from the
Shoreline but also 100 to 125 cm. landward from the shore-
line. Too, on those collection dates it was seen that
Percentage water-content at the +250 cm. interval was
generally appreciably higher than adjacent intervals.

Reference to Figure 3 suggests these maxima in percentage

 

64

 

water-content were not related to depressions on the
transect since alight rises occur at both the +150 cm.
and the +250 cm. area of the transect.

Maximum differentials in percentage water-content
between shoreline and the +300 cm. interval, i.e. range,
occurred at those collection periods during which the
substrate was Frozen. The lowest range, 12.78 per cent,
occurred on April 6, immediately after a heavy rain in
the collection area. The twelve-month mean range was
2157 per cent between the +300 cm. interval and the
+25 cm. interval.

0n the basis of' twelve month means, the +300 cm.
interval contained only 12.21 per cent as much water as
did the +25 cm. interval. 0n collection dates in December,
May, June, July, August, September, and October the
driest interval (usually +300 cm.) contained less than
10 per cent of’ the amount of“ water present in the sand
0' the wettest interval (usually +25 cm.). In December,
July, August, September, and October the driest interval
contained less than 5 per cent of the amount of water
Present at the wettest interval.

Table 8 presents the results 01’ determinations
of the amount of water present in monthly samples from
each interval expressed as a percentage of the amount
Of water held in saturated samples of‘ sand collected at
each interval in September (see Chapter IV for methods

Utilized). With reference to twelve month means for

iP , h

 

 

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65

 

 

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66
determinations at each interval, it is apparent that the
+300 interval contained only 14.15 per cent of the theoreti-
cal maximum water retention capacity whereas the +25 cm.
interval contained 102.8 per cent. It is postulated that
all Figures in excess of” 100 per cent theoretical maximum
capacity are influenced by the fact that organic content
within the samples was considerably higher at the initia-
tion of the study than at its near termination when
water retention determinations were made. As in Table 7,
a direct relationship exists between the amount of‘ water
actually held by samples From each interval and distance
from water line, again with the exception of‘ the +250 cm.
interval.

Except For the exceptions noted below, results
within each monthly collection parallel the twelve month
mean tendencies. Potential water retention capacity was
most nearly realized on those collection dates at which
the beach was Frozen and on the April collection date
following a 1.25 inch rain. Greatest departure From
Potential water retention capacity occurred during summer
months when the most landward intervals Frequently con-
tained less than 10 per cent of‘ their theoretical capacity.
Considerable irregularity in realiZation of theoretical
capacity at intervals along the transect were particulary
Obvious in August, September, and October. Inasmuch as
such irregularities are not apparent in granulometric

conditions at each interval nor in transect profile, their

 

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67

  
  
  
  
  
   
 
  
  
 

f;,.p1anetion is apparently partially related to substrate
organic content. It should be noted that in August Four
of sixteen intervals contained less than 1.0 per cent of“
their potential water retention capacity, much of that
water apparently being hygrosc0pic. Also to be noted is
the fact that in the winter months those intervals 75 cm.
or less from the shoreline characteristically contained
more than 100 per cent of‘ theoretical water retention
capacity.

Table 9 presents the results of‘ monthly determina-
tions of’ percentage volatile matter on a dry gross weight
basis for each 25 cm. interval along the transect. Values
ranged frOm a maximum of‘ 10.98 per cent recorded December
flat the +25 cm. interval to a minimum of‘ 0.39 per cent
recorded on May 6 at the same interval. The greatest
range along the transect, 10.49 per cent, occurred in the
December collection. The smallest range, 0.94 per cent,
Occurred in the May collections. A mean range For the
twelve-month collection period was recorded as 3.22 per
cent. In general the greatest occurred during those
months in which the transect substrate was in a Frozen
condition, the smallest ranges occurring during the

Spring and early summer months. when the minimum volatile
percentage within a monthly collection is compared with
the respective maximum, it appears that the minimum most
closely approximates the maximum in the spring and departs

t0 the greatest degree during those months in which the

 

 

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69

substrate was frozen. Considered on the basis of the
twelve-month mean, the minimum percentage volatile content
of the substrate approximated 20.7 per cent of‘ the maximum
percentage volatile content of’ the transect substrate.

Reference to Figure 6 reveals considerable simi-
larity in twelve month means of‘ per cent volatile material
and per cent water at each interval along the transect.
It should be noted that means of intervals zero to ~75
are for only four series of‘ collections. Readings from
the +300 interval to the +175 cm. interval show distinct
correlation between means of’ volatile material content
and that of“ water. From the +175 cm. interval lakeward,
the marked variation between the two Factors considered
is probably attributable to the increasing proximity of
the transect intervals to the phreatic level and the
lake itself (see Fig. 3). Reference to Table 7 indicates
an abrupt marked decrease in percentage water composition
From the +150 to +200 cm. intervals from November through
July, those months during which the shoreline lay approxi-
mately at the zero index point. Reference to Table 5
shows no abrupt change in volumetric porosity, unit
weight, or uniformity coef‘f‘icient at any point along the
transect except at the ~75 to -100 cm. intervals. It
should be noted that the harmonic mean grain diameter
and effective grain 3129 do change rapidly in the +175
to +225 cm. portion of the transect. However, the data

For the critical Factor of‘ volumetric porosity do not

 
 

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71

   

teal the explanation on a granulometric basis For the
_ regularities in water retention by the substrate at

[ those intervals landward From the +175 cm. interVal. A
hypothesis might be made that percentage water content

at each interval along the transect is directly related

to porosity and distance From the shoreline and/or phreatic
surface up to a certain critical distance, in this case

about 175 cm., and beyond the critical distance it becomes

secondarily related to amount oF organic matter present.

Beach chemical data

Table 10 presents the results or” monthly pH
determinations of‘ lake and interstitial water as well as
in'situ determinations of‘ pH at each 25 cm. interval
along the transect. From Nevember through May all pH
readings were made by inserting the probe of” a Beckman
portable pH meter directly into the water sample or the
substrate at each 25 cm. interval. From June through
October, the method was supplemented at each interval
along the transect by standard soil pH determinations
(see Chapter IV, Methods and Materials).

The abrupt and marked drying of‘ the transect
substrate landward From the +150 cm. interval in May
necessitated modification of‘ pH determination techniques
in that readings made with the probe obviously were
inaccurate. Laboratory tests suggested the use of‘ probe

alone was not an accurate index of pH when percentage

 

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F 73
titer-content in the substrate fell below 10.0 per cent.
zaomparieon of data in Table 10 with those of Table 7 shows
Etlosely similar results between the two pH determination
techniques utilized until percentage water-content of

the sample falls below 10.0 per cent. Therefore, con-
siderations of pH readings made from June through October
“‘0 based upon the lower segment of dual numbers reported
on Table 10.

Lake surface water pH immediately offshore From
the transect ranged From 7.2 recorded under seven inches
of ice on January 4 to 9.5 recorded on June 6, 1964.
Median pH during the twelve-month period was 8.4 with a
twelve-month range of" 2.3 pH units. In general lake
water pH most nearly approached neutrality during the
winter months and became most alkaline in spring and
early summer (Fig. 7). Reference to Table 10 reveals
that interstitial water withdrawn from the beach with an
napirator or from a collection sump ranged in pH from
6.8 recorded on July 6 to 8.3 recorded on April 6. Refer-
ence to Figure 7 reveals that pH of‘ the interstitial water
tended to be considerably more acidic than that of the
lake offshore. A tendency is also indicated For an
inverse relationship between lake and interstitial water
PH. The two most alkaline readings f‘or lake water, For
instance, occurred on dates when pH of‘ interstitial water

was most acidic.

 

 

 

 

 

 

      

—-——- lake water

--- - — —- interstitial water

 

7.0.

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\
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608 J I A l I A I ‘1 L 1

Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct,

. Fig. 7.--.‘Ylonthly variation in pH of‘ lake and
interstitial water.

 

 

 

 

 

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75

   

£3? The pH of intervals along the transect ranged from
,; 8.6 recorded at the Zero interval in May and June and
the +25 cm. in May to 7.0 recorded in October at- the
+125 cm. interval and in March at the +25 cm. interval.

From November through April pH range along the transect

 

ordinarily did not exceed a value of 0.4 units. Ranges
in other months were considerably greater, becoming 1.0
or more pH units in June, July, September, and October.

- In all months but February and March the pH shifted
toward the acidic side a10ng the transect with distance
landward from the shoreline. In February and March the
pH became slightly more alkaline at the most landward
portion of the transect with reference to those intervals

closest to the lake. It should be noted that it was in

 

January, February, and March that lake pH was closest to
neutrality.

Comparison of Figure 7 with Table 10 reveals
that monthly shifts in pH of lake water tended to be
mirrored by similar shifts in any given interval along
the transect. Relatively acidic conditions of lake water
developing January through March also occurred at all
intervals, winter minimum pH being last realized, as a
generalization, by those intervals most distant from the
shore. The November through March shift toward neutrality
in lake pH was paralleled most closely by that of the
index location interval. All intervals exhibited a rapid

Shift toward alkaline conditions following warming

 

 

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76

 

u

'tunstitial water was not mirrored by any interval. The

early fall increase in pH and subsequent decrease in

October was exhibited by all intervals as well as by

 

lake and interstitial water.
Results of both methods of alkalinity determina-
tions (see Chapter IV, Methods and Materials) used on

samples of‘ sand taken monthly From each 25 cm. interval

 

along the transect showed no significant nor consistent
variation. Figures given in Table 11 are thus means of“
Values of‘ both methods rounded off‘ to the nearest whole

number. Although only bicarbonate alkalinity was indi-

 

cated as present in samples of‘ transect substrate through-

 

out the study on the basis of‘ alkalinity determination
techniques used, on several occasions pH exceeded 8.3.
Obviously, therefore, alkalinity determinations made

Upon the substrate are suspect. Figures presented on
Table 11 are For comparison purposes only even though
they are reported as bicarbonate alkalinity as ppm. (law?)
for Five grams of‘ sand.

Reference to Table 11 indicates that on any given
collection date alkalinity tended to decrease markedly
with distance f‘rom shoreline, often by a f‘actor as great
as 14 but usually on the order of Five times, the greatest
differences occurring in the summer months. Since

alkalinity is ordinarily produced by the presence of the

 

 

 

 

 

77

  

 

 

 

 

 

 

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78

salts of weak acids, it is not known whether the determina-
tions recorded on Table 11 are reflections of‘ the amount
of bicarbonates or rather of‘ silicates, humic salts, etc.
It is surmized the high organic content of‘ the intervals
nearest the shoreline contributed considerable amounts of‘
organic acids.

Because of the large number of unmeasured variables
which produce alkalinity, perhaps the only conclusion
which can be drawn validly From Table 11 is that organisms
present in the interstitial water of" more landward por-

Pertions of the transect were more subject to pH varia-

 

tiOns than those in the more shoreward portion, the higher
alkalinity acting as a buffer to pH Variation.

Table 12 presents the results of chemical deter-

 

minations made upon water samples withdrawn just below
the lake surface approximately f‘ive meters of‘f‘shore of
the shoreline each collection period. Table 13 presents
the results of‘ chemical determinations made upon samples
01’ interstitial water withdrawn monthly April thI‘OUQh
UCtober from the hygropsammon zone of the transect.
Dissolved carbon dioxide content in lake water
ranged t"I‘Om a high of‘ 18.0 ppm. on January 4 to a low of‘
0'0 ppm. recorded in June, July, September, and October.
In general dissolved carbon dioxide tended to be hiQhBSt
during the winter months and lowest in summer (”9- 8)'
DissolVed carbon dioxide content in interstitial water

ranged From a high of‘ 100.0 Ppm. recorded on July 5 to a

 

 

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Nae om NNe No as em as eN eN Dee mNe ewe .Eaaemwowxea
Ne Ne o mN we sN o o a a me o .Eaa ca .xes moo
am mN NNe en o as as «N sN use Nee ewe .Eaa cs .xem moo:
e.m e.m m.e m.m m.m m.m n.m s.N m.N N.N n.m s.m In mumcoaw
.wwo .mmm .N:< xman omen xms .mo< .mwa .MoL .Non .mmo .noz

 

 

   

ee emanate on

N. HmnEm>DZ mmHQEwm HOL NHNU HmowEmto henna mxmulloNv Mdmvfi.

 

 

 

 

 

 

 

 

 

 

 

 

I’ll.

 

 

 

 

 

ON.e Om.N Oe.m Om. OO.e Om. .eaa as sueeem
O.em O.NN O.NN O.me O.Om O.me .Eaa ce sOm
Dec. Dec. moo. So. go. 0ND. .53 CH Hmpop cogs.”
Ne.O Oe.O m0.0 Oe.O OO.O Ne.O .EOQ cs .moca-oceeo
eOO. NOO. mOO. eOO. OOO. sOO. .Eaa as .Ooasec NOz
OmO. mmO. NOO. sOe. ONO. esO. .Eaa ca .Ooesec mO2
m.nNee m.OOm O.Nmm N.eNN O.emN s.eOe H\EOE cs museum .men eases
ee .eOO m .amm a .On< O sees a mass O ewe

m ON.e mm. ON. mm.e mm. Om. .Eaa ce mueeem
O.nN O.Oe O.Oe O.NO O.mN O.me .EOQ Ce eOm
ONO. eOO. OeO. OmO. Oee. Oee. .Eaa ce sweep cogs
so.O mO.O OO.e mm.e mm.e NN.e .EQO ca .moca-oceeo
OeO. aOO. eOO. meO. meO. meO. .EQO cs .Ooasec NOz
mmO. snO. nOe. wee. mee. mee. .sOa cs .Ooasec mO2
O.mmN m.mee m.OOe s.ONN N.eeN m.ONn H\EOE cs museum .meu Hmong

m ..Hn< m .eme m .33 e .52. m .03 N :62 \

 

 

 

DmDCHHCOUIloNr uqm<h

 

 

 

 

 

 

81

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

s.NOO O.NON N.NeN O.eeO N.O e . . empeH\EOE ce assume
N m eme o QNe maeumao>ucoc um>Hommwu
O.eOO O.eON N.NOO O.OOO . . . ameee\memquHHeE OH
O OOe O eNe O OeN posses meepmHo> Omseommeu
O.O O.N O.O O.OOe O.O O.N O.s .EOO Ce
NOO Omseommeu
O.Ne O.O O.O O.NN O.O O.e O.eO suspecnemm e as
NO Om>eommeu
O.e N.O O.O e.N O.O O.O s.O .EOO as
No um>aowmwu
mm m a o a a me .eoe cw mmmcuumc
memconumoncoc
one we .eau cw mmmcuum;
mNN «em Dem man mm m e .eua CH mwmcnemc
a owe Hence
oar r .Eua cw
mm are new mm mm mm wmmcnumc mu
One OON NON Oes O e .EOO as eeeceemme
o Nee owe a Hopes
.Ea ce Neeceemme
a o o o a o m mumconumo
one .Eoo cw >uwcefimxam
. HON Now one moe Nee use mumcooemowa
e.e O.N O.O O.O O.N O.O n.O IO
er .000 m .umm m .m:< m xaon m moon m >mE m .uo<
>HumHEmLU Hmams Hmwuwvmhmucw mcoN CDEEmmoonm>Ill.nH UAmQF

 

 

 

 

 

 

 

82

 

 

O.O O.ON Oqu O.NN O.ON O.NN O.Ne 0O as messmamaems
0.0 O.e s.O Oe.Oe OO.N OO.N Os.N .EOO cs OOHHHO
0.0e 0.0 0.0s O.O O.ON O.ON O.eN .EOO as OOOOOHOO
OO0.0 OOO.O eOO.O OeO.O OOe.O OON.O OOO.O ce cwfimaemeoe
NN.O O0.0 ON.O Oe.O OO.O OO.O OO.e mpOOmmmzmwcseo
OeO. eeO. sNO. NOO. OOO. ONO. OOO. .OOHWWMOOMWaOec
eOO. Oee. ONN. OOO. NOO. OOO. Ose. .Ooawwmammwesec
O.<Oee O.Ome «.mmO O.OeO O.ONO e.ONO O.OOO Wmmwmmmmummwwmm
N.<N O.Oe e.NO O.eO OOOO O.NO O.eO OOeHOMOOM>MMMMMwomOsos
ee upon O .amO O .O:< O seas O mans O ems O .uaq

 

 

 

 

 

noDCHpcoUIl.ne Udm<e

 

 

 

 

__

 

 

 

83

 

lake water "
95- "
————-- interstitial water "

 

 

—+—

45-

J}
C]
‘

bl
U1
A

carbon dioxide in ppm;
N
u:

I
l
I
I
I
I
l
I
I
I
I
I
I
I
I
I
I
20. '
I
I
I
I
I
I
I
I

O
I $\
’/ “

(A
O
.
.—-—
———— ——-———————-———
——————._——

 

 

 

w A l l ‘ ‘
NOV. Dec. Jan. Feb. mar. Apr. May June July Aue- 599‘ 0°t°

8.--Dissolved carbon dioxide in ppm. For

Fig.
mo
”thly lake and interstitial water samples.

 

 

L O a

 

 

I :1 lama

' mason ‘

I
it». I

filial II

 

84

  
  
  
  
   
 
  
  
  
  
   
   

am of 2.0 ppm. reported on May 6. As expected, dissolved
£02 maxima corresponded closely with pH lows. £02 minima
corresponded with pH highs for both lake and interstitial
water. No relationships appeared between lake and inter-
stitial water with reference to relative amounts of dis-

solved carbon dioxide reported monthly.

 

Dissolved oxygen content in lake water reported
as ppm. ranged from a high of 13.6 on December 8 under
four inches of ice to a low of 4.3 ppm. recorded on
November 7. In general, in lake water, dissolved oxygen
reported as ppm. was highest during the winter months and
lowest during the summer, peaks in dissolved oxygen con-
tent occurring in December and March followed by a summer
decline and fall recovery (Fig. 9). Such a pattern is
typical of a body of water such as Reach A1 (Burner and
Leist, 1953; Schultz, 1952; Dineen, 1953; Tiemeier and
Moorman, 1957). With reference to interstitial water,
dissolved oxygen content in ppm. ranged from a high of
3.4 ppm. on April 6 to a low oF 0.0 ppm. reported on
August 6. Reference to Figure 9 reveals an apparent
close relationship between dissolved oxygen content in
lake and interstitial water from April through October,
the amount of oxygen in interstitial water being con-
Siderably less, however, than that present in the lake
on any collection date.

Although no relationship was seen between relative

amounts of dissolved oxygen and carbon dioxide, reported

 

OL—

 

 

 

 

85

 

13. _—

  

dissolved oxygen in ppm.
01 ON N1 CD \D
I l l a l

J:-
I

 

0
Nov. Dec. Jan. Feb. Mar. Apr. May

andinterstitial water samples.

 

lake water
interstitial water

 

O \"——‘/

June July Aug. Sep. Oct;

 

Fig. 9.--Dissolved oxygen in ppm. for monthly lake

 

 

31‘-

. u

   

 

86

ppm., in lake water, comparison of Figures 8 and 9

3;; show direct relationship between the amount of the gases

   
   
   
  
   
  
  
   

dissolved in interstitial water in that both gases display
essentially parallel concentration curves from April
through October.

For lake water, dissolved oxygen content reported
as per cent saturation ranged from a high of 103.0 record-
ed on July 6 and October 11 to a low of 39.0 recorded on
November 7. No seasonal trends were apparent, maxima and
minima fluctuating irregularly throughout the 12 month
study (Fig. 10). Neither was there significant correla-
tion between dissolved oxygen in lake water as per cent
saturation and any other physical-chemical factor measured
with the exception of dissolved oxygen in ppm. Per cent
dissolved oxygen saturation in interstitial water ranged
from an April 6 high of 31.0 to an August low of 0.0.
Again, no seasonal trends were apparent. An apparent
direct relationship did occur, however, between the
degree of oxygen saturation in lake and interstitial
water (Fig. 10).

Total dissolved solids recorded as milligrams
per liter in lake water during the 12-month period of
the study ranged from a high of 1173.6 mg/l. on October
11 to a low of 108.8 mg/l. on February 6. Seasonal
variation was indicated, total dissolved solids decreas-
ing markedly following formation of ice cover in

December to the late winter minima and increasing rapidly

 

 

 

87

 

110

100'

90-

03
D
I

 

.q
C
l

per cent saturation
U1 O"
O O
I A

J:-
O
n

M
D
a

20-

10‘

 

0

 

 

—— lake water

—————— interstitial water
0‘ .
\ /\
\ I \
\ ,’ ‘
\ I \
\ I \ ,
\ I \ /
'\ ‘ I \ /
‘ l /

 

}‘_,—u

 

I r I I I ‘

Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct

19. 10.--Dissolved oxygen as per cent saturation

F'
forlake and interstitial water monthly samples.

IMII I
Ian In I
I arm I‘
:- 22: mm
. III of I
I iwlved
his I
3:12.; II

"n
. .E'I‘,

 

 

88
. to the autumn level following the disappearance of ice
cover in the spring (Fig. 11). Volatile dissolved solids
reported as milligrams per liter at the same collection
dates ranged from a high of 942.0 mg./l. on October 11 to
a low of 61.6 mg./l. on March 6. A seasonal variation of
dissolved volatiles was also apparent, their relative
amounts closely paralleling total dissolved solids.
During the 12-month study no general trends in volatile
content expressed as a percentage of total dissolved
solids were apparent (Fig. 13). Only in January, September,
and October did volatile percentage content depart mark-
edly from a mean value of about 55 per cent. In January,
percentage volatile-content dropped to (10.8 and in
September and October it jumped abruptly to 68.0 and 80.2
respectively.

In interstitial water, total dissolved solids
ranged from a low of 326.4 mg./l. recorded on May 6 to
a high of 1194.0 mg./l. recorded on October 11. A peak
in July and August was followed by a September decline
and abrupt October increase. Total volatile solids
showed a similar pattern of abundance. A May low of
171.6 mg./l. was followed by a July peak, a September
decline, and an abrupt October increase to a maximum of
1194.0 mg./l. (Fig. 12). Values for percentage volatile
content of total dissolved solids in interstitial water
fluctuated irregularly around an approximate 60 per cent

mean with no obvious trends (Fig. 13). There were no clear

2 O____O h ,

89

 

1000 ———————— Total dissolved solids
800 - ----- Total volatile dissolved solids I

 

100
90
80
7o . \. l
60 ' l [~‘j l 1 41 n .. .
Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. 59p, Oct

Fig. 11.--Total and volatile dissolved solids in mg./l.
for monthly lake collections.

1200

1000

800
700
600

500 -

 

 

 

 

 

   
   

Total dissolved solids

_—————-

n m e e
__—-— Total \910 atile issolved
solids in mg. l.

I“‘I

400 -

"190/1.

30 ’
U I I \

200 .. \ ,

 

 

 

100 I n I t J . l A l I
Nov. Dec. Jan. Feb. Mar. Apr. May June JUIY AUQ- Sep. Oct.

Fig. 12.--Tota1 and volatile dissolved solids in mg./1.
Fornwnthly interstitial water samples.

 

P l a, ,, l a

 

 

 

 

90
100

 

90 p

80 r

‘1
D
6

0"
O

 

Percentage
U1
0
l

J)-
C)
I

Lake water

 

30 ' ——————— Interstitial water

20

 

 

l I 4 J I I I L

ONOV. Dec. Jan. Feb. mar. Apr. May June July Aug. Sep. Oct.
Fig. 13.—-Volatile dissolved solids expressed as

percentage of total dissolved solids For monthly lake and

interstitial water samples.

correlations of total dissolved solids between lake and

hmerstitial waters and none For volatile solids.

Total hardness in lake water ranged from a high
09158.0 ppm. recorded on October 11 to a low of 68.0 ppm.
recorded on February 6. Evidence of seasonal variation
was indicated in the early spring increase which Followed
the winter low and preceded that of early summer (Fig. 14).

Maximum total hardness For interstitial water was recorded

 

‘ 15'0—

1U!)-
enJ
80: ;_
_ . 70*
V
‘I E 60W
l 50—

 

AUJ .0} ’a 3: ‘ \ V //’I
A.” -' \’tota1 hardness
304 1. f —————— calcium hardness

- - - - - - .. - “carbonate hardness

 

‘ o
Ian-0.0....

 

 

 

20 A L A J. 1 l— l 1 J 1

Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.

Fig. 14.--Lake water hardness expressed as ppm.
Cams monthly For total, calcium, and carbonate hardness.

400

350
300 -——total hardness
ZSDL -~-—~—Ca hardness
........ €03 hardness

 

Vij

200 '

5150
Q.

 

\

60 , o—-"‘

 

50 I . L .1 l I l A

Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep, Oct.

 

 

PPm. CaC03 monthly f‘or total, calcium, and carbonate

Fig. 1S.-—Interstitial water hardness expressed as
hardness.

 

 

imd thi

faith a1
w: in m
it, rig
1‘ assigr

eiatial

 

   
  
   
  
   
 
  
  
  

92

as 388.0 ppm. on July 6, a June 6 minimum content of

1‘ 98.0 ppm. immediately preceding the maximum. It is be-

lieved the abundance of‘ precipitated calcium carbonate

visible at the site of' withdrawal depth of interstitial

water in July contributed to the maximum recorded on that

data, migration of‘ the collecting site making it dif‘f’icult

to assign fluctuations exhibited on Figure 15 to seasonal

variation.

With reference to the cation Factors of" total

hardness, calcium hardness of‘ lake water showed essen-

tially parallel Fluctuations through the twelve months

to those of‘ total hardness (Fig. 14). Maximum calcium
hardness ranged From 90.0 ppm. in October to a 10w of

I 34.0 ppm. reported on February 6. Reference to Figure 14
reveals similar seasonal variation in calcium hardness
to that of total hardness. Only in December and June
did the calcium percentage of‘ cation hardness components
comprise less than 50 per cent of‘ the total hardness.
Percentage calcium content of‘ total hardness during the
twelve month period of“ sampling beginning in November
and terminating in October is presented in Table 14.
Since Ferrous iron hardness component during this period
was negligible, it would appear magnesium ions were
Significant contributors to total hardness of‘ the waters
0? Reach A1.

In interstitial waters, calcium hardness ranged

From a high of 240.0 ppm. on July 6 to lows of‘ 60.0

1%

 

 

 

 

D.Dor c.00r macaw

n.mm e.me e.ms n.sm

<.Ne o.es .N.na

93

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w.mm

0.0m m.Nm 0.0m

o.oor m.mm

q.vm N.m<

0.009

o.om

umpme Hmwuwpmuopca
co mmmcuum: ampop
Lo pcmcoaeoo mum
Iconnmo mmmpcmomma

Hopes mme
Lo mwmcunmc Hmuop
Lo ucmcoaeoo mpm
nconumo mmmpcmouma
Hopes Hmwpflpm
upwaca Lo mmmcomm;
amuou so pcmcoa
1500 mu mmmaceoemo
mxma Lo mmmcnnm;
ampop mo ucecoa
1200 mu mmmpcmouma

 

«new .m:< >Hon econ

.unq .ums .nec

0C0“,

00mg

.>oz

 

 

 

ucm exwa AOL mwecunmz Hmvov Lo

 

 

momma Hmwuwpmnmucw

awesome; eumconpmo com eawoamo

moppcmoneaun.er mumqh

 

 

i

end 62.0

I 11!!!ch

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94

  

“or! 62.0 ppm. reported on May 6 and June 6 respectively.
Reference to Figure 15 reveals closely parallel fluctua-
tions in seasonal calcium content to that of‘ total and
carbonate hardness but relatively little parallel to com-
parable hardness fluctuation in lake water (Fig. 14).

For interstitial water, percentage calcium content of‘
total hardness during the seven month period of sampling
beginning in April and continuing through October is pre-
sented in Table 14. As in lake water, the neglibible
amount of Ferrous ions present during this period would
indicate magnesium ions were of‘ considerable importance in
composition of cation components of total hardness of‘
interstitial water.

Carbonate hardness in the waters of Reach A1 OFF-
shore of‘ the transect ranged From a high of‘ 136.0 ppm. on
January 4 to a low of 24.0 ppm. recorded on both February
6 and March 6. Figure 14 indicates somewhat parallel
trends in seasonal variation of carbonate hardness to
those of‘ total and calcium hardness. For the twelve-month
period of‘ the study, November through October, carbonate
recorded as ppm. CaC03 with reference to percentage of‘
total hardness is given in Table 14. Inasmuch as the
major anions producing nan-carbonate hardness include
sulfates, nitrates, and silicates, it would seem likely
that these ions must have been relatively most abundant
during those months of‘ February and March when carbonate

hardness was lowest. Reference to Figures 19, 22, and 20

2
2

.

2J—

 

95
however, reveals these ions also were at relative minimum
abundance in February and March.

In interstitial water, carbonate hardness during
the April through October sampling period ranged From a
high of‘ 388.8 ppm. on July 6 to a low of‘ 98.0 ppm. re-
corded on the sixth of‘ the previous month.‘ For that
period, carbonate hardness as percentage of‘ monthly total
hardness amounts is presented in Table 14. No obvious
explanation For the relatively low October reading is
apparent. No correlation of‘ interstitial water carbonate
hardness was apparent with that of‘ the water of‘ Reach A1.

Negative non-carbonate hardness in the water of

Reach A1 was recorded on November 7 and January 4, being

 

20.0 and 4.0 ppm. respectively. Such a condition occurs
when a surplus of‘ bicarbonate ions is available to

Satisfy the divalent metallic ions present, thus indicat-
ing the association of bicarbonate ions with sodium and
PDtassium. Negative non-carbonate hardness also occurred
in interstitial water on the sixth of‘ May, June, July,
and August, the amount of‘ ppm. being 4.0, 10.0, 82.0,

and 22.0 respectively.

Alkalinity in both interstitial water and that

Of‘ Reach A1 throughout the study was that of" carbonates

 

and bicarbonates, no hydroxide alkalinity being present

at any collection date. In water of‘ Reach A1 maximum

total alkalinity as ppm. Ca003 was encountered on January

4. Minimum readings were recorded as 24.0 ppm. on

 

 

1
I

'_ he lintI

, muss?

Eat 1|

' hilly

I! 9359
1'11 in
‘15 :cr

225M

 

, 96

February 6 and March 6. Figure 16 indicates a general

u late winter alkalinity minimum, an early spring pulse
followed by a mid—summer secondary minimum, and late
summer increase in total alkalinity to approximately that
of early winter. In the waters of Reach A1 alkalinity
was essentially that contributed by bicarbonate ions.
Only in the May, June, and July collections did carbonate
ions-contribute appreciably to total alkalinity. Maximum
carbonate alkalinity coincided with the lowest pH value
0f 9.5 recorded on June 6.

Total alkalinity of interstitial water ranged
from a maximum of 470.0 ppm. recorded on July 6 con-
comitant with the most acidic pH recorded, 6.8, to a
minimum total alkalinity of 108.0 ppm. recorded on June 6
(Fig. 17). Only on April 6 was carbonate alkalinity
Observed. Aside from the 3.0 ppm. carbonate alkalinity
recorded on April 6, all alkalinity in interstitial
water was that contributed by bicarbonate ions.

Total ferrous and ferric iron in water collected
from Reach A1 at each monthly collection date ranged
from a high of 0.140 mg./l. on November ‘7 to a low of
0.001 mg./l. occurring on March 6 (Fig. 18). Figure 18
indicates a precipitous decrease in the amount of dis-
solved or suspended iron throughout the winter until the
disappearance of the ice cover prior to the April 6 col...
lection. A spring pulse of total iron content declined

in the summer to a secondary low in August followed by

 

 

 

97

 

' '150
—— total alkalinity

—————— H003 alkalinity
e e e o I e e e . ...CDS alkalinity

140-

   

130-
12m
110.

100

3

CaCU
as \D
J? E

q
C)
l

 

alkalinity in ppm.
Ch
1;?

0'!
P

am
3%

20.

 

 

e
c
O . o. ---n-- .. .4... ..ed 4 . . z: -

th Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.

 

F . 16.--Total, bicarbonate, and carbonate
alkalinity of lake water at monthly collections.

 

 
    

..u
n:-
\A|- unu- null-n.
/

 

 

 

98

 

 

  

500 total alkalinity

_ .. - --.— bicarbonate alkalinity

Ca003

alkalinity as ppm.
:3
Q

 

4n I L n A

A;

 

 

 

"Nov. Dec. Jan. Feb. Mar. Apr. NJay Jdne July Aug. Sep.
Fig. 17.--Total and bicarbonate alkalinity of
interstitial water at monthly collections.
partial fall recovery. A somewhat similar spring pulse,
summer decline, and fall recovery of total iron content
occurred in interstitial water (Fig. 18). Maximum iron
content of 0.260 ppm. was recorded on May 6 and minimum
content of 0.001 ppm. recorded on August 6. No con-

spicuous relationship between total iron and dissolved

Oct.

oxygen was apparent in either lake or interstitial water.

No evidence was visible that the anaerobic condition of
the interstitial water on May 6, June 6, August 6, or
September 5 promoted increased reduction of ferric iron

to the more soluble ferrous form.

 

 

 

 

 

99

 

1.0

.6 ‘ -———-——-—-1ake water

————— interstitial water

 

.06 "

a

D

J—\
L

o
C)
N

4L

.01%
.008

.

.006<

total Ferrous and ferric iron in p rn

.004]

.002&

 

 

 

'0n1L\I¥ l I 1 g | I ' 'g‘

ROV- DEC. Jan. Feb. mar. Apr. Nay June JUlY AUG. 395- fitt'

f0 Fig. lB.--Total ferrous and ferric iron in pom.
r menthly lake and interstitial water samples.

 

m a hi
r: 1m
‘ “A111

 

item

'! emu
I. 1‘ July

_ m1,

S‘JIE'J 1

100

Dissolved sulfate as ppm. in lake water ranged

   

—: from a high of 32.0 ppm. recorded on January 4 to a low
of 15.0 ppm. on May 6. No seasonal trends were apparent,
monthly figures being sporadic in amount (Fig. 19). In
interstitial water, dissolved sulfate as ppm. ranged from
a high of 45.0 ppm. on October 11 to a low of 3.0 ppm.

on July 6. As was the situation in lake water, the
monthly amount of dissolved sulfate was sporadic and

showed no seasonal trends. Reference to Figure 19 reveals,

 

100

—————— lake water

30 . ——-——-—- interstitial water

70

60.

01
D
l

sulfates in ppm.
#-
C)
I
\

 

 

 

 

U A I A L 41 I n A A ‘

Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct

Fig. 19.--Su1fatss in ppm. for monthly lake water
and interstitial water samples.

 

 

 

fifths
mun

-. Agra?

 

 

101

 

‘fl however, some evidence of somewhat parallel fluctuation of
sulfate content in lake and interstitial water in June
through October.

In the water of Reach A1 during the 12-month period
of the study dissolved silica ranged from a high of 3.10
ppm. on August 6 to a low of 0.25 ppm. on February 6.

Figure 20 reveals the abundance of dissolved silica was

 

 

20.0
A
lake water I \\
100% _— ————— interstitial water / \\
8.0

0‘
£‘l

as"
‘? ¢

M

o

O
A

N
P

Silicates in ppm.

.Frfle

06‘
.5

.¢
03‘

 

 

 

I I J I I ‘ l A I
.1 ‘

 

Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.

Fig. 20.--Silicates in ppm. for monthly lake water
and interstitial water samples.

1
I
.

 

 

I. Lbrsti‘

I parent

cl.

.
I an

i“- turn

102

V regular through the year, three major peaks apparently

  
  
  
  
   
 
  
  
   
  
 

. occurring in January, April, and August. Each peak was
immediately followed by a rather precipitous decline. In
interstitial water only one peak in dissolved silica was
apparent. The July maximum of 13.10 ppm. was preceded by
a rapid increase from the May 6 minimum of 2.00 ppm. and
in turn preceded a rapid decline in the fall.
Orthophosphate as ppm. in lake water ranged from
a December 8 maximum of 1.86 ppm. to a May 6 minimum of
0.42 ppm. Figure 21 indicates the December-Nay decline
to be rapid and regular. Following the June increase to
0.30 ppm., dissolved orthophosphate in the water of
Reach A1 fluctuated in the vicinity of 0.80 ppm. until
termination of the study. Such amounts are far in excess
of the 0.003 ppm. mean value of soluble phosphorous
reported in 479 lakes in northeastern L'Jisconsin (Juday

and Birge, 1931).

In interstitial water, orthophosphate ranged
from an initial April 6 maximum of 1.00 ppm. to a minimum
of 0.18 ppm. recorded on July 6 (Fig. 21). A late

‘ Summer recovery to the September 6 value of 0.36 ppm.
preceded another decline at the termination of the study.
In the water of Reach A1 nitrate nitrogen ranged
‘ from the initial maximum of 0.116 ppm. on November 7,

‘ December 8, and January 4 to a minimum of 0.020 ppm.

\

recorded on June 6. The June minimum was preceded by a

secondary march decline and April recovery which was also

 

l g

 

103

 

 

0
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- —————— interstitial water

 

 

 

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Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.

Fig. 21.--0rthophosphate in ppm. for monthly lake
water and interstitial water samples-

 

 

 

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to the
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104

  
  
  
  
  
  
  
  

apparent with nitrite nitrogen quantities during the 12-
month study (Fig. 22). Nitrite nitrogen quantity ranged
from the November through January maxima of .016 ppm. to
the minimum of .000 ppm. recorded on June 6. In inter-

stitial water, the nitrate nitrogen maximum was recorded
on August 6 at 0.226 ppm. Minimum nitrite nitrogen con-

tent was recorded as 0.007 ppm. on July 6. Fluctuations

 

.15

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Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.

 

 

 

Fig. 22.-~Inorganic nitrate and nitrite nitrogen in
PM. for monthly lake water samples.

 

!
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1 in mm

1 malls!

 

   
 
   

The literature reporting results of chemical

105

lparalleled essentially those in interstitial water

(Fig. 23).

determinations upon samples of interstitial water From

gfin nitrite and nitrate nitrogen quantities in lake water

 

 
 
  

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Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct

 

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106

e psammolittoral zone indicates that, apparently, the

  
  
  
  
  
  
  
  
  
  

Ci assumption has been made by most investigators that the

r mater withdrawn is that which surrounds the psammobiota.
There appears to be no published report with reFerence

to investigations 0F the source 01’ water withdrawn From
the sand by means oF the standard aspirator sampling
device. IF it is assumed that the hydrological phenonema
associated with the cone oF depression developing around
a well bore during water withdrawal (NeinZer, 1942)

occur as well about an aspirator thrust into the hygro-
psammon zone 01“ a sandy beach, then it seems unlikely
that the volume of‘ water normally withdrawn From the beach
For chemical determinations can all have come From the
three or Four centimeters of surFace sand which lies
above the lowermost point oF the aspirator probe.

Mavis and Tsung-Pei (1939) report that horizontal
water movement below the peizometric water level in
artiFicial sand-Filled Flumes is substantially constant,
but velocity in the capillary Fringe varies with a well-
deFined regularity From zero at the top oF the Fringe to
the velocity in the piezometric zone at the bottom oF
the Fringe. Luthin and Day (1955) Found considerable
Capillary Flow oF water laterally above the water table,
but their data are not applicable to the withdrawal oF
water From the hygropsammon zone. ReFerence to investi-
gations oF capillary Flow and water conduction in soils

by Moore (1939), Richards, (1931, 1933), Lambe (1951),

 

 

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107

‘Kirkham and Feng (1949), and Remson and Fox (1955) reveals

no generalizatiOns which can be made with regard to the
source oF the interstitial water withdrawn From the
hygropsammon zone sand with an aspirator.

In an attempt to determine the origin oF the
interstitial water withdrawn From the hygropsammon zone,
a number oF experiments were conducted adjacent to the
beach transect. ReFerral to publications by Dole (1906),
Ambrose (1921), Kaufman and Todd (1955), and KauFman and
0rlob (1956) suggested the use 01“ uranine and/or acridine
Orange as tracers in plotting movements oF water in the
hygropsammon zone.

In the First series 0F tests, conducted in July,
the aspirator probe was inserted into the sand to a depth
0F Four centimeters approximately 60 centimeters FrOm
the water's edge. Four bores one cm. in diameter and
Five cm. deep were then made at radii oF 13.0 cm. From
the probe and an additional Four at 25 cm. radii (see
Fig. 24). Uranine solution was then poured into all
bores until it stabilized at ground level. An aspirator
next was utilized to withdraw 250 cc. oF interstitial
water over a one-hour period, uranine solution being
added when necessary to maintain the solution level in
the bores. Finally, a straight edge was drawn across
two parallel wooden guides to shave oFF 2.5 cm. layers

across the bore Fields until uranine stains were no longer

 

 

108

 

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0 A1
50 cm. to water's
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.34 .A41 13 C01. - 0A2 . 82
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0 A3

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Fig. 24.--Diagram oF bore Field utilized in
tracing hygropsammon zone interstitial water movement.
visible in the sand. This procedure was repeated at
three diFFerent sites along the beach.

Horizontal sectioning oF the three bore Fields
revealed that the mean maximum depth 0F uranine penetra-
tion in the bores at 25 cm. radii was 11.5 cm. with a
range From 7.5 to 13.0 cm. There was no evidence oF
signiFicant departure oF the uranine tracer From a
Cylindrical pattern about the bores at any depth achieved,

Mean maximum depth oF uranine tracer penetration in the

 

 

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109
bores at 13.0 cm. radii was 8.1 cm. with a range of 5.0
to 10.0 cm. All bores at 13 cm. radii showed a slight but
distinct eccentricity oF uranine solution pattern toward
the position oF the aspirator probe at 5.0 cm. depth but
none was apparent at the surFace or other depths.

In a subsequent series oF tests, uranine solution
was added to bores positioned as in the First series and
250 cc. oF water withdrawn From the center oF each oF
three bore Fields over one-hour periods. Sections oF the
bore Fields were not made; rather the 250 cc. inter-
stitial samples were examined under ultraviolet light
For evidence oF Fluorescence. None was detected.

Both series oF tests would seemingly indicate
that the interstitial water withdrawn From the hygropsammOn
zone with an aspirator is not From the surFace layers oF
the send but rather apparently is derived From depths
below the terminus 0F the aspirator probe. Consequently
the chemical data resulting From determinations upon the
interstitial water withdrawn From the beach are most likely
not those oF the upper photic layers but instead those 0F
the aphotic and essentially abiotic zone beneath. There-
FOre, the interstitial water chemistry reported in the
preceding section oF the Results might best be considered
the'Taw material" to be modiFied by the organisms oF the
photic zone rather than indicative oF the chemical environ-

ment within the photic zone itselF.

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110

31010 ical data ualitative
lake samples

Table 15 presents the results oF qualitative and
quantitative phytoplankton sampling From the water oF
Reach A1 oFFshore oF the beach transect during the period
November 9, 1963, to October 10, 1964. (See pp. 44 - 46
For methods utilized in data collection.)

From qualitative samples, 187 species were
identiFied during the 12-month sampling period. As
indicated on Table 16, minimum number oF species were
reported present December through March, and maximum
species numbers were present in April through October.
The minimum number oF species, 30, was reported For the
January collection, 16 0F these being members oF the
Cyanophyta. Thirty-six oF the 90 species recorded pre-
sent For qualitative samples taken July 6, the collection
in which the maximum number oF species was present,
were members oF the Chrysophyta. Qualitative collections
were dominated by members oF the Chrysophyta in all months
but January and June. This qualitative dominance 0F the
Phytoplankton by diatoms was lessened in January when
members oF the Cyanophyta became qualitatively more
abundant. Members oF the Chlorophyta dominated the
qualitative plankton Flora collections only in June.

At all times oF the year except July through
September members oF the Chlorococcales qualitatively

dominated the green algal component oF the phytoplankton.

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|Members of the orders Volvocales, fledogoniales, and
Tetrasporales appeared Only in samples taken March through
August.

In qualitative samples throughout the 12-month
sampling peried, as a generalization, members of the
Chlorophyta were in number of species less abundant than
either the Cyanophyta or Chrysophyta. An exception to
the generalization occurred in May, August, and September
when green algal species outnumbered those of the
Cyanophyta but trailed Far behind the Chrysophyta.

No clear seasonal qualitative Fluctuation in
dominance of the blue-green companent e? the lake phyto-
Plankton by members of the Hormogonales as contrasted

to members of the Chroococcales was apparent. Throughout
the study, each month the number 0? species within the
two groups was essentially equal.

Each month, in the Chrysophyta, members of the
Pennales were qualitatively Far more abundant than those
of the Chrysomonadales or Centrales; Members of the
Chrysomonadales were entirely absent From qualitative
Samples during the period June through August. Members
of the Centrales were inconspicuous in qualitative
samples, being rare or absent except February through
May.

Qualitative distribution of members of the
pYrrhophyta during the 12-month sampling period showed

definite seasonal variation, the maximum number of species

 

 

 

125
occurring from March through October, members of the
group being qualitatively absent or negligible from
November through February.
Only 10 species of algae were present in qualita-
tive samples throughout the-year. A list of these species
includes Anacystis mar inata, Aghanocagsa elachista,

Lyngbya hieron musii, Phormidium luridum, Tetraedron

 

minimum, Anomoeoneis vitrea, Cymbella microce hala,

 

Navicula heuFleri Var heufleri, Navicula heufleri var
legtoceghala, and Navicula radiosa var tenella. In
addition to these were 11 species which were absent only

in one or two months, usually in the January-March period.
These include Agmenellum guadruplicatum (absent March-
April), Phormidium angustissimum (March), Spirulina
subsalsa (January), Tetraedron muticum (February),

Achnanthes linearis F. curta (January), Achnanthes

 

linearis var. linearis (December-January), Cymbella )

affinis (January), Navicula minima (January), Nitzachia

H

 

linearis (December-January), and geratium hirundinella

-——--——_.~

 

(February and April). Thus 10 of the 21 species of

algae which might be considered most representative of

the year around phytoplankton of Reach A1 were members

0? the Pennales. Seven of the year-around species were

Cyanophytes. Significantly, 45 per cent of the 187

species reported in phytoplankton samples From Reach A1

during the study were diatoms, and 23 per cent were 1

Cyanophytes. From November through April, chrysophytes

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126

and cyanophytes collectively contributed 78 per cent or
more of the total species present monthly. In February,
98 per cent of phytoplankton species in Reach A1 were
chrysophytes or cyanophytes. From May through October,
chrysophytes and cyanophytes collectively contributed

at most 69 per cent of total phytoplankton species pre-
sent in monthly samples. In August, the minimum cyano-
phyte-chrysophyte contribution of 58 per cent occurred.

Biological data, Quantitative
lake samples

Figure 25 presents in graph Form the monthly
variation in total number of individual phytoplankters
per liter in quantitative plankton samples during the
12—menth study. One entry was made For each individual
cell or For each aggregation of cells bearing the char-
acteristics of a species. That is, one Chlamydomonas
cell was recorded as one unit, one Agmenellum colony as
one unit, one Lyngbya trichome as one unit, one Dinobryon
colony or any isolated individual thereof as one unit,
etc. Thus interpretations of tables and graphs of
Quantitative lake sample data should include considera-
tion that cells of Filamentous and colonial forms are
Far more numerous than total counts indicate.

Extreme Fluctuation in total number of individuals
per liter occurred From a low of only 345 individuals
per liter on March 5 to a peak of 119,010 individuals

per liter on September 5. Minimum counts correlate

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127
Nov. Dec. Jan. Feb. Mar. Apr. may June July Aug. Sep. Oct.

  
 

 

100000
00000

60000

40000

20000

2000

..s
C3
C3
0

Total number of individuals per liter
(1)
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O

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is
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Fig. 25.--Total number of individuals per liter,
Quantitative phytoplankton samples November through
October.

 

 

128
closely with those manthly collections at which the sur-
face of Reach A1 had a thick ice cover, that in March at
the end of four months of apparently uninterrupted ice
cover and stable lake levels. Increased counts of 1770
and 6264 individuals per liter respectively on April 6
and May 6 followed extensive churning of shoreline and
shallow water areas during the break up between March 5
and April 6. As noted previously, this churning was
apparently sufficiently vigorous to eradicate the luxuriant

growth of Chara vulgaris that had flourished throughout

 

the winter under the ice.

The relative decline in total number of phyto-
plankters per liter during June through August coincided
with a progressive decline in the water level in Reach A1
during these months. Heavy August rains and partial ‘
return of lake level to that of November-March preceded
the 10-fold September increase in total number of phyto-
plankters per liter, and the October 10 decline in phyto-
Plenkton abundance was concomitant with a rapid drop of
water level.

Comparison of Figure 25 with Figure 11 (total and
dissolved volatile solids in mg./1. for monthly lake
collections) shows a relatively close correlation between
the total number of phytoplankters per liter and dissolved
volatile solids in lake water. However, the volatile
solids content (i.e. organic matter) of the lake water

at any given collection date cannot be considered an

——_r~.-

 

 

129

accurate indication of phytoplankton abundance at that
time, although the literature of limnological research
contains frequent references to dry weight measurements
of plankton and/or seston determination as measures of
phytoplankton productivity. Notes taken on examination

of quantitative plankton samples in December, for example,
refer to abundant ghaga fragments and debris from decay-
ing vascular plants which line Reach A1. January 4 notes
indicate a bloom of copepod nauplii, abundant detritus,

and numerous aquatic fungi. In February, Chara Fragments

 

were again very common and detritus abundant. March
notes refer to a bloom of copepod nauplii, very abundant
detritus, and large numbers of Actinomycetes and yeasts.
In August detritus content in quantitative samples was
very heavy as it was also in October. Zooplankters were
recorded as being very abundant in October. Thus Figure 11
is as illustrative of a heavy organic load as it is of
the phytoplankton bloom which developed in late summer in
Reach A1.

It should be again noted that from July through
October a dense floc of decaying algae, consisting pri-
marily of clumps of Phormidium spp. ensheathing floating
Fragments of ghgga, covered almost the entire surface of
Reach A1 making phytoplankton collection quite difficult.

Disparity between quantitative plankton counts
and obvious dense algal growths at the surface of Reach A1

during the summer months suggested possible error in

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130
data collection techniques. Consequently, a number oF
tests were made to determine the accuracy of the con—
centrating method used in collection oF quantitative
phytoplankton data For Reach A1 (see pp. 44-46). Tests
were conducted upon two each oF the November and May
qualitative and quantitative plankton samples.

The 10 cc. concentrates oF two liter quantitative
samples were diluted to one liter and again concentrated
to 10 cc. volume using the standard procedure utilized
throughout the study. The Filtrate From each concentrate
was next passed through a 0.45 u Millipore Filter and the
Filter either cleared in cedar oil and scanned under
675x For plankters or the residue scraped From the Filter
and brought up to 10 cc. with distilled water. No identi-
fiable plankters were observed by direct scanning oF the
Filter oF either the November or Nay quantitative sample,
nor in mounts oF 1/7 cc. aliquots oF the 10 cc. residue
suspensions. The bolting silk Filter pads used in recon-
centration oF the November and May quantitative samples
were then each placed in 10 cc. oF distilled water and
agitated vigorously with a stream oF compressed air.

Three 1/? cc. aliquots oF each 0F these suspensions were
then mounted and examined For plankters Following standard
counting procedures. None was Found in the November
sample and but two Fragments oF Phormidium angustissimum
Found in the May sample.

mm?
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131

The above procedure was then repeated substituting
qualitative samples For quantitative. It is estimated
each qualitative sample represented the plankters Found
in approximately 1200 liters oF water. In both the
November and May collection the amount oF detritus pre-
sent on the transparent Millipore Filter made direct
counts oF the number oF organisms per liter oF qualitative
sample Filtrate impossible.

When residue oF the Filtrate was scraped From
the Millipore Filter and brought up to 10 cc., three
counts oF 1/7 cc. aliquots gave an average oF 1218
organiSms per cc. 0F 10 cc. concentrate oF one liter oF
Filtrate. Thus one liter oF bolting silk Filtrate con-
tained approximately 12,180 organisms. IF it is assumed
that 12,180 organisms are lost in the Filtrate when
1200 liters are concentrated to 10 cc., then it might
be Further assumed that approximately 20 individuals
might be lost to the Filtrate when two liters oF the May
sample were concentrated to 10 cc. Une liter oF bolting
silk Filtrate From the November qualitative collection
contained approximately 7140 organisms, suggesting that
about 12 organisms were lost to the Filtrate when the
two liter November quantitative collections were con-
centrated to 10 cc. In both November and May it would
appear that loss oF plankters through the bolting silk

Filters was negligible.

‘
i
w

of ml

untrai

 

cunt

 

 

 

  
 

 

132

Counts oF 1/7 cc. aliquots oF 10 cc. suspensions
of washings From the bolting silk Filters used in recon-
centrating the May qualitative samples gave an approximate
count of 280 organisms in each of the 10 cc. oF washwater
From the bolting silk Filters. November qualitative
samples gave an approximate count of 210 organisms per
cc. of washwater. In both instances, apparent loss oF such
a relatively small number oF organisms From the qualitative
samples suggesb less than one per cent 0F plankters were
lost in the standard concentration procedure utilized.
Consequently, on the basis oF the tests described, it
would appear that the number oF organisms lost in the
concentration procedure utilized was negligible.

Inasmuch as the Drop-sedimentation technique 0F
quantitative count determination involved the count oF
all phytoplankters in the three 1/7 cc. aliquots rather
than extrapolation From an arbitrary number oF Fields,
the standard procedure as outlined in Standard Methods
(1960), it is concluded that the quantitative phyto-
plankton data reported in Table 15 and Figure 25 is indica-
tive oF a quantitatively sparse phytoplankton Flora
which prevailed December through April in the waters oF
Reach A1.

Comparison oF Figure 25 with those illustrating
monthly variation in pH, dissolved carbon dioxide, and
dissolved oxygen in both ppm. and per cent saturation

shows little relationship. Neither is any correlation

Minus
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133
obvious between the total quantitative variation in
phytoplankton abundance as shown in Figure 25 and total,
calcium, or carbonate hardness (Fig. 14) nor For the
three Forms oF alkalinity illustrated in Figure 16. In
addition, no relationship is evident between Fluctuations
in total phytoplankton abundance (Fig. 25) and sulFates
in ppm. (Fig. 19).

The general decline oF total Ferrous and Ferric
iron in ppm. (Fig. 18) From November through March and
rapid increase in April Following ice cover disappearance
Parallels that oF total phytoplankton abundance, but this
relationship is not clearly evident From May through
October.

Little correlation appears between relative
total phytoplankton abundance and that oF silicates in
Ppm. (Fig. 20) throughout the year, although there is
some indication that the low phytoplankton counts oF
February and March were accompanied by minimal amounts
BF dissolved silicates, and the April and September phyto-
plankton pulses were concomitant with silicates maxima.
Any relationship between total phytoplankton abundance
and that oF orthophosphates in ppm. (Fig. 21) is inverse,
maximum amounts 0F orthophosphate tending to occur during
periods oF lowest total phytoplankton abundance, and
lowest amounts occurring during periods oF maximum phyto—
plankton populations. Although no deFinite correlation

is apparent between nitrate or nitrite nitrogen in ppm.

(fig. 2

Mali
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3
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134
(Fig. 22) and total phytoplankton, a general tendency For
an inverse relationship between nitrites and nitrates
abundance and that oF total phytoplankton abundance is
indicated.

Table 17 presents data 0F quantitative lake phyto-
plankton samples on the basis 0F species percentage
composition oF total phytoplankton For each month of the
12-month sampling period. Table 18 presents a summary oF
data From Table 17, the species data 0F Table 17 being
grouped to Division and Order. It should be noted that
all percentages were rounded oFF to the nearest 0.1 per
cent. Consequently, totals For all data in Tables 17
and 18 do not sum to 100.0 per cent.

A clear seasonal progression in quantitative
dominance oF lake phytoplankton appeared during the
study. Data From Table 18 indicate that members oF the
Cyanophyta dominated quantitative collections in November,
December and January, comprising respectively 76.1, 68.0,
and 81.8 per cent oF the total phytoplankters, and during
July and August when they comprised 46.6 and 47.5 per cent
0F the total phytoplankton respectively (see also Fig. 26),
During the months oF February, March, April, and May,
members oF the Chrysophyta in turn dominated quantitative
samples From Reach A1, contributing to the total phyto-
plankton oF those months percentages oF 62.6, 63.6,

52.2 and 56.6 respectively. Again in September and

October the Chrysophyta became dominant in quantitative

 

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Per cent of total phytoplankton

 

145

Nov. Dec. Jan. Feb. Ear. Apr. Hay June July Aug. Sep. Oct.

 

100 ' I fii '
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--—-- Chrysophyta
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Fig. 26.--Division percentage composition in
quantitative lake samples, November through October.
samples, making up 84.1 per cent of the total population
in September and 73.7 per cent in October.

Members of the Chlorophyta dominated quantitative
lake samples only in June, when they comprised 47.6 per
cent of the phytoplankton count. At no time during the
study did the fourth Division represented, the Pyrrhophyta'
contribute more than 16.5 per cent oF the total monthly

phytoplankton population.

 

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146
The dominance of quantitative lake samples From
November through January by cyanophytes was primarily
attributable to relatively large populations of members

of the Hormogonales, particularly Phormidium angustissimum,

 

E. africanum, and E. minnesotense. In November, Phormidium
africanum was at 19.5 per cent the most abundant phyto-
plankter, accompanied in relatively significant numbers

by E. angustissimum (16.2 per cent), E. minnesotense

(12.0 per cent), flflQELEELE marginata (7.1 per cent),

Aghanocagsg glachista (4.9 per cent), and Qeratigl

 

hirundinena (4.8 per cent). The December collaetijn was

 

characterized by relatively large numbers of BHQEEEQLEE
angustiggimum (28.2 per cent), Anacystis marginata

(8.9 per cent), A. montana (7.7 per cent), Aghanocapsg
elachista (3.5 per cent), Lyngbya aerugiqgg—caerulea
(5.3 per cent), Tetraedron minimum (5.4 per cent) and
Uroglenogsis americana (7.1 per cent). Again most

abundant (23.3 per cent) in January, Phormidium angustissimum
was in that month accompanied in relatively significant
numbers by Anacystis marginata (11.3 per cent), A. montana
(6.9 per cent), Gomghospheria lacustris (8.7 per cent),
Lyngbya hieronymusii (4.4 per cent), L. aerugineo-caerulea

(5.9 per cent), Aulosira laxa (4.4 per cent), and

 

Uroglenopsis americana (8.7 per cent).
The quantitative succession of cyanophytes by
Chrysophytes obvious in Figure 26 From February through

May was largely attributable to the appearance of relatively

 

largo
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147
large numbers of members of the genus Dinobryon.
Dinobryon divergens dominated the February collections,
contributing 48.8 per cent of the total plankton count
(Table 17). In March, Q. sociale var. americanum assumed
dominance by contributing 30.1 per cent of the total
phytoplankton count, to be replaced in turn in April and
May by Q. sociale, which made up 27.0 and 40.4 per cent
of the total phytoplankton count respectively in those
two months. Other signiFicant contributors to quanti-
tative phytoplankton counts in February were Nostoc

galudosum (8.1 per cent) and Peloglea bacillifera (4.1

 

per cent). Accompanying Q. sociale var. americanum in
significant percentages in March were Anacystis marginata
(5.2 per cent), Chroococcus turgidus (5.2 per cent),

Cosmarium angulosum (5.2 per cent), and Navicula radiosa

var. tenella (3.4 per cent).

In addition to Q. sociale in April, other signifi-
cant contributors to the total phytoplankton counts were
Phormidium angustissimum (4.6 per cent), Dinobryon

sociale var. americanum (17.6 per cent), and Peridinium

cinctium (8.8 per cent). In May, D. sociale was accompanied

 

in significant numbers by Tetraedron muticum (7.7 per cent),

Oocystis elliptica (5.7 per cent) and Q. sociale var

americanum (5.9 per cent).

June quantitative samples were dominated by the
Chloroohyta, but the most abundant green alga species,

Uocystis elliptica, contributed only 13.8 per cent oF the

m1 pl}
ll thl .‘
(10.6 91

em
" (5.9 pl

 

 

148

total phytoplankton count. Other significant contributors
to the June phytoplankton were Phormidigm angustissimum
(10.6 per cent), Plectonema notatum (6.4 per cent),
Oedogonium rufescens (5.3 per cent), Cosmarium angulosum
(6.9 per cent), and Ceratium hirundiqgQa (16.5 per cent).

Members of the Cyanophyta re-established dominance
in July and August, although the single most abundant
species in July was Peridinium inconspicugm (13.4 per
cent). Other important members in the July phytoplankton
were Anacystis marginata (6.5 per cent), Phormidium
angustissimum (11.3 per cent), Lyngbya hieronymusii
(7.0 per cent) and Tetraedron minimum (4.7 per cent).

The most abundant member of August samples was
Phormidium angustissimum (26.2 per cent), accompanied in
relatively significant abundance by Lyngbya hieronymusii

(4.3 per cent), Tetraedron minimum (9.0 per cent),

Navicula radiosa var. tenella (6.6 per cent) and Nitzschia

acicularioides (2.9 per cent).

The pronounced September pulse of Chrysophytes
exhibited in Figures 25 and 26 is attributed to Dinobryon
divergens, which contributed 74.9 per cent of the phyto-
plankters in September to be followed by one of Q. social»
which, absent from quantitative and qualitative samples
since June, abruptly appeared to contribute 52.5 per cent
0? the October phytoplankton count. On the basis of

percentage composition, the only other significant members

of the September phytoplankton were Oocystis elliptica

 

 

u: [up

 

 

149

var minor (1.6 per cent), Phormidium angustissimum (2.8

 

per cent), 3. luridum (1.2 per cent), Tetraedron minimum

(1.4 per cent), 1. muticum (1.2 per cent), Navicula radiosa

var. tenella (2.1 per cent), Nitzschia linearis vars. (1.4
per cent), and Achnanthes linearis F. 2233; (1.1 per cent).
Reference to Table 15 however, reveals that many phyto-
plankters were in numbers per liter more abundant in
September than at any other time in the study.

Dinobryon sociale was accompanied in significant
relative abundance in October collections by Anacystis

montana (2.9 per cent), Phormidigm anqustissimum (2.9 per

 

cent), E. luridum (3.5 per cent), Oocystis elliptiqg

var. minor (2.5 per cent) and Dinobryon divergqu (16.1

 

 

per cent).

Biological data, Qualitative
psammon samp es, 300 cm. interval

Table 19 presents the results of qualitative and
quantitative sampling From the beach transect at the
300 cm. interval during the period November 9, 1963, to
October 10, 1964. (See pp. 44 - 47 For methods utilized
in data collectiod} Table 20 summarizes data From Table
19 with respect to Order and Division totals.

From qualitative samples taken at the 300 cm.
interval, 62 species were identified during the 12~month
Sampling period. The maximum number of species, 40, was
recorded on February 5. Minimum species number of 12

was reported on September 5. As indicated on Table 20,

 

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157
minimum numbers oF species were present July through
October when total species number approximated halF or
less that oF the November - June period. Maximum species
numbers were reported From November through June, during
-which months chrysophytes comprised at least halF oF the
total number 0F species present in qualitative samples.
The qualitative dominance 0F total species number during
the November ~ June period by members oF the Chrysophyta
was primarily attributable to members 0F the Pennales.
During the same period, the Cyanophyta closely approached
the Chrysophyta in number of species monthly. Only in
September did the Hormogonales species comprise less than
halF oF the total number oF cyanophyte species present.
Although the Hormogonales reached maximum qualitative
abundance in February with 10 species present, the total
number oF Hormogonales species in each monthly collection
From November through June never was less than eight.
The Chroococcales component oF the 300 cm. psammonFlora
showed a similar seasonal distribution, maximum number
of species occurring From November through June with an
abrupt July to October decline.

Throughout the 12-month study, members oF the
Chrysophyta approached or exceeded 50 per cent 0F the
total species present. Members oF the Pennales con-
tributed From BO to 100 per cent 0F the total chrysophyte
species present in qualitativa samples throughout the

12 months. The three species OF the Heterococcales

 

...__— __ _._—-.._.

 

Ehry:

abun
and
non
pre
gm

thl

 

158
identiFied during the 12-month study were Found only From
December through August, all three appearing in February
and May. Phaeothamnion, the only member oF the
Chrysotrichales identiFied, was present only in February.

Species oF the Chlorophyta were relatively most
abundant From November through August, but only in July
and August oF the 12-month period did they contribute
more than 18 per cent oF the total number oF species
present. In September and October, only one species oF
green alga was identiFied in qualitative samples From
the BOD cm. interval. The only member oF the Tetrasporales
identiFied during the 12-month period, Oloeocystis,
appeared only on February 5. Members oF the Volvocales
and Chlorococcales were sporadic in appearance, being most
common From November through August and practically
disappearing in September and October.

Qualitatively, members 0F the Chlorococcales
dominated the green algal cemponent 0F the psammon Flora
throughout the 12 months.

Only seven species oF algae were present at the
300 cm. interval throughout the entire 12-month sampling
period. These included Syflechococcus aeurqinosus,

Chlorococcum humicola, Cymbella microce hala, Navicula

M

 

 

decussis, N. heuFleri var. leptocephala, N. radiosa var

-—-————_.—_._

tenella, and Rhopalodia qibba. Plectonema notatum was

present in all qualitative samples except those oF

September 5, and Cymbella aFFinis was absent only in

 

 

w

vari
No
the
d9:

 

159
October. Navicula hungarica var linearis was absent From
qualitative samples only in September and October.
Biological data, Quantitative
psammon samples. SOO gm. interval

Figure 27 presents in graph Form the monthly
variation in total number oF algal members oF the psammon-
Flora per co. in quantitative 2.4 cc. samples taken during
the 12~month study. (Techniques oF data collection are
described on pp. 44 - 47.)

Total number oF individuals per cc. oF sand From
the 300 cm. interval ranged From a minimum oF 1,369 on
August 6 to a maximum oF 13,296 individuals per cc. re-
corded on January 6.

Comparison 0F Figure 27 with Figure 25 (total
number oF individuals per liter in quantitative lake
plankton samples November through October) shows an
almost absolute reversal oF seasonal variation in quanti-
tative abundance oF algae. Naximum algal population
densities in the sand at the 3GB cm. interval occurred
in months in which lake phytoplankton was least abundant,
i.e. during the December-February period, and were at
minimum during those months in which lake phytoplankton
was most abundant, i.e. November and Nay through October.
An exception to the generalization occurred, however, in
March when lake population density was at minimum and
beach +3OO cm. interVal population exhibited a late winter

depression. It should be noted here that maximum

 

 

7* ..g»fir=: z. -- l —- #: Fag- 7 ,

 

 

 

 

J
C)

CO.

 

Thousands oF individuals per

 

 

 

 

Fig. 27.--Total number oF individual algal units
1 per cc. oF sand, quantitative monthly samples, 300 cm.
i interval.

PW"
sun:
10:!

flat:

 

 

161

population densities occurred either when the beach was
solidly Frozen December through February or in April Fol-
lowing the March thaw. Minimum densities occurred in
March as thawing began and again From May through October
as the sand became progressively drier. Even at minimum
beach population density in August, iF density per cc. oF
beach sand at the 300 cm. interval be multiplied by 1000
to convert to density per liter and compared with maximum
lake phytoplankton abundance in September, density oF
beach population at minimum exceeds that 0F lake phyto-
plankton at maximum by a Factor oF approximately 10. A
comparison oF maximum beach population density at the
300 cm. interval in January with that oF maximum lake
phytoplankton abundance in September indicates the maximum
beach population density to be approximately 112 times
that oF the lake.

Table 21 presents data oF quantitative psammon-
Flora abundance at the 300 cm. interval on the basis oF
species percentage composition oF total individuals counted
each month oF the 12-month sampling period. Table 22
presents a summary oF data From Table 21, the species
data From Table 21 being grouped to Order and Division.
All percentages are rounded oFF to the nearest 0.01 per cent.

Data From Tables 21 and 22 indicate that in
quantitative samples the Cyanophyta numerically dominated
the ZOO cm. interval psammonFlora From November through

June and again in September and October. This dominance

162

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167
was particularly pronounced in November through January
when members of the Cyanophyta comprised 85 per cent or
more of the total number of algae present in sand From
the 300 cc. interval. During those three months, members
of the Homogonales contributed 80 per cent or more of the
total number of algal individuals present.
Figure 28 presents in graph Form a summary of

percentage composition of total algal population during

 

Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.

—— Cyanophyta

_u___n——-Chrysophyta
____._._ Chlorophyta

......... Euglenophyta

Ch
(3
4*

Percentage of total count
(,1 b 'n
O D J

N
O
L

.1
O
:_

 

 

 

 

 

 

Fig. 28.-—Summary of percentage composition by
Divisions For 300 cm. interval quantitative samples,
November through October.

 

 

..__.____._—_._. .

the
div
Fig
abt

 

168
the twelve-month study with reference to the four
divisions represented in the collections. Reference to
Figures 27 and 28 shows that the general decline in total
abundance of algae present in the sand at the 300 cm.
interval is relatively closely paralleled by a similar
decline in the percentage contribution of the Cyanophyta
to the total algal population. The relatively minor
"bloom" of Phormidium microtgflgm in September and October
produced a disproportionately large upturn in the
Eyanophyta component of Figure 28 because of the rela-
tively small total number of algal individuals present
in the sand at that time. The apparent increasing im- T
portance of members of the Chrysophyta and Chlorophyta

From March through August indicated on Figure 28 is not

 

illustrative so much of significant population "blooms"
within those two Divisions as it is of the relative
decreasa in abundance of members of the Cyanophyta.

Only in July and August did the Chrysophyta
numerically dominate the quantitative samples of the 300 cm.
interval. The numerical importance 0? the Chrysophyta in
July was attributable primarily to relatively large
numbers of members of the Heterococcales, particularly
Botrydiopsis arhiza, which alone contributed 30.08 per
cent of the total individuals present that month.
Bgtrydiopsis arhiza was also the most common species in
August, outnumbering all other Chrysophytes combined

with 26.58 per cent of the total individuals present.

The

Ehri

hav

uh!

981

.I. ..Illll‘llc |

 

169
The large (40.70 per cent) September contribution of the
Chrysophyta to the total algal population was attributable
to the Pennales, particularly Navicula heufleri vars.,
Navicula decussis, and Navicula radiosa var. tenella,
which respectively contributed 11.07, 16.74, and 5.53
per cent of the total algal population. An important
contributor in the June 6 collection was Navicula heufleri
vars., which accounted for 10.80 per cent of the total
number of individuals that month.

Individual species of the Chlorophyta rarely
'contributed more than five per cent to the total algal
population during the 12—month study, and only in May
through September did the chlorophytes comprise more than
10 per cent of total individuals present. 0f the
Chlorophyta, only the species Chlorococcum humicola was
a significant contributor to the algal flora of quanti-
tative samples at the 300 cm. interval throughout the
study. In June, July, August, and September Q. humicola
comprised respectively 15.45, 18.32, 24.83, and 16.65
per cent of the total individuals present in quantitative
samples.

0f the Chroococcales present in the 300 cm.
interval quantitative samples, gyflgchococcus aeruoingsgg
was the only species present throughout the 12-month
period and present in significant amounts in any one
collection. In May, July, and August samples §.

aeruginosus contributed respectively 13.58, 15.68, and

 

 

 

trih

tote

 

170

21.25 per cent of the total algal population. Chroococcus
minutus, appearing only in September and October con-
tributed 26.68 and 10.70 per cent respectively of the
total count.

In the Hormogonales component of quantitative
psammon samples frOm the 300 cm. interval Schizothrix
friesii. Phormidium microtgmgm, Nostoc paludosum,

Microcoleus acutissimus, fl. vaginatus, Plectonema notatum,

 

and Cylindrospermum minimum were significant. In
November, December, January, and February 5. friesii
contributed 58.22, 57.47, 61.96, and 24.88 per cent of
the total count respectively. Although Phormidium

microtomum appeared only in September and October, in

 

October 49.53 per cent of the algal individuals iden-
tified were of this species. Cylindrospermum minimum,
although present only sporadically through the year, in
April and May became a significant contributor to the
total count, supplying 28.82 and 15.22 per cent respec-
tively in those months. Plectonema notatum, also present
only sporadically through the year, contributed 19.10

per cent of the total count in November and 27.88 per

cent of the count in June.

In summary, for quantitative samples at the 300 cm.
interval, Schizothrix friesii dominated November through
February and was joined in dominance in March by
1icrocoleus acutissimus and Botrydiopsis arhiza. In

N

April and Nay Anabaena anomala and 8. arhiza characteriZed

 

 

 

 

Elk-‘3'

 

171
quantitative samples. In June Plectonema notatum,

Chlorococcum humicola, g. arhiza, and Navicula heuFleri

 

vars. deminated. July and August samples were characterized
by relatively large numbers of Syflgchococcus aeruoinoggs,
Chlorococcum humicola, 8. arhiZa. Navicula ggcussis. fl.
radiosa var. tenella, and fl. heufleri vars. By September
and October quantitative samples were dominated by

Chroococcus minutus, Phormidium microtomum, Chlorococcum

__——

 

humicola, Navicula decussis, and Navicula heufleri vars.

 

Biological data, Qualitative
psammon samples, 200 cm. interval

Table 23 presents the results of qualitative and
quantitative sampling from the beach transect at the
200 cm. interval during the period November 9, 1963,
to October 10, 1964. From qualitative samples taken at
the 200 cm. interval, 104 species were identified during
the 12-month sampling period. Total number of species
present in monthly qualitative samples ranged from a
maximum of 81 on February 5 to a minimum of 23 recorded
on October 10. As indicated on Table 24, the greatest
diversity of species occurred from January through may.
FrOm June through October there was a progressive decline
in total number of species.

Throughout the 12-month sampling period, cyanophytes
and chrysophytes dominated qualitative samples from the
+200 cm. interval. Only in October did the Cyanophyta

contribute less than 30 per cent of the monthly species

 

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132
list. Cyanophyte contribution to monthly lists ranged
from a maximum of 48.1 per cent in February to a minimum
of 21.7 per cent in October with diversity of species
approximating or exceeding 45 per cent of the total species
count from January through July. This important quali-
tative contribution of the Cyanophyta was attributable
primarily to members of the Hormogonales.

With reference to total number of species present
in monthly qualitative samples, the Hormogonales exhibited
maximum numbers of species January through July with
greatest number-- 29 --occurring in February when the
beach had been Frozen for three months. The least number
of species were present in August, September, and October.
Members of the Chroococcalee showed a similar seasonal
distribution, the largest monthly number of species being
present in February and the lowest in August through
October.

Species of Chlorophyta were relatively most
abundant in February when 13 species were recorded, but
members of the Chlorophyta comprised only 17.3 per cent
or the total species count in that month. In relative
abundance, members of the Chlorophyta were most abundant
February through June, comprising during those months
From 17.0 to 13.4 per cent of the total species present
monthly, and in October and November when they contributed
17.4 and 16.4 per cent respectively. Species of green

algae reached lowest numbers in July through September.

 

The

sent

memt

DB?

mun

 

183
The Four species present in the October collection repre-
sented 17.4 per cent of species present. At no time did
members of the Chlorococcales contribute less than 66.6
per cent of the total green algal species present in
monthly quantitative samples.

The chrysophyte component of monthly qualitative
samples From the ZOO cm. interval was dominated by members
of the Bacilliarophyceae, which contributed 90 per cent
or more of the total chrysophyte species monthly. No
obvious seasonal distribution patterns For the
Bacilliarophyceae as a group were apparent, although
there was an obvious decline in number of species during

the study From the initial high of 25 to 27 species in

 

November and December. At no time during the 12-month
period did the Chrysophyta contribute less than 34.6
per cent of the monthly total species list. Minimum
percentage contribution to the total species list took
place in February and May when chrysophyte species com-
prised respectively 34.6 and 39.0 per cent of the total
number of species present. Beginning in August,
chrysophyte species became increasingly important in
qualitative samples, reaching a 12—month peak oF 61 per
cent in October.

Thirteen species of algae were present in quali-
tative collections at the ZOO cm. interval throughout the
entire 12-month sampling period. These included

Synechococcus aeruginosus, Phormidiufl_angustissimu

.11

 

184

Chlorococcum humicola, Achnanthes linearis f. curta,

 

 

 

Achnanthes minutissima, Cymbella microce hala, Navicula
cryptoceghala, Navicula decussis, Navicula pugula var.

mutica, Navicula radiosa var. tenella, Navicula heuFleri

 

var. le toce hala, Navicula hungarica var. linearis,

and Rhogalodia gibba.

Biolo ical data uantitative
MM eral

Figure 29 presents in graph Form the monthly
variation in total number of algal members of the psammon—
Flora per cc. in quantitative 2.4 cc. samples taken during
the 12-month study. ‘

Total numbers of individuals per cc. of sand

 

from the 200 cm. interval ranged From a minimum of 1,602

 

on October 10 to a maximum of 65,559 individuals per cc.
recorded on February 5. Comparison of Figure 29 with
Figure 27, page 160, shows general agreement of seasonal
variation in total individual counts between the SOD and
200 cm. intervals. Maximum numbers of psammon algae were
reported at both intervals in monthly collections made

in those months in which the beach was solidly Frozen
with another peak in early Spring. The Summer and Fall
decline in algae abundance at the 380 cm. interval was
essentially paralleled at the ZOO cm. interval, although
throughout the study total numbers of algae in quanti-
tative monthly collections at the ZOO cm. interval were

much more irregular.

 

Utah t- ID”. “I: 'F‘vlr

185
Ne]. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.
T l I I F f I I | I

60 —

40 '

 

thousands of individuals per cc. of sand
A
O

 

1.5

 

 

Fig. 29.~-Total number of individual algal units
per cc. of sand, quantitative monthly samples, ZOO cm.
interval.

 

 

 

 

186

Comparison of Figure 29 with Figure 25 shows
little relationship between seasonal variations in algae
abundance at the ZOO cm. interval and that of quantitative
lake plankton samples. The relationships, iF any, as
were those of the SOD cm. interval and lake plankton,
are inverse. At minimum algal population density at the
ZOO cm. interval in October density of algae at the ZOO cm.
interval exceeded the maximum lake plankton density of
September by a Factor oF approximately 18. Maximum
algal population density at the ZOO cm. interval in
February exceeded that of September maximum lake phyto-
plankton density by a Factor of approximately 730. When
maxima of algae population densities at the 200 cm.
interval in February and that of the 300 cm. interval in
January are compared, that of the ZOO cm. interval ex-
ceeded that of the BOO cm. interval by a Factor of
approximately Five.

Table 25 presents data of quantitative psammon

alQae abundance at the ZOO cm. interval on the basis oF
Species percentage composition of total individuals
counted each month of the 12-month sampling period.
Table 26 presents a summary of data From Table 25, the
Species data From Table 26 being grouped to Order and
Division. All percentages are rounded oFF to the nearest
0.01 per cent.

Data From Tables 25 and 26 indicate that in

quantitative samples the Cyanophyta numerically dominated

 

 

 

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194

the 200 cm. interval psammonflora throughout the 12-month
sampling period. During this period cyanophyte individuals
comprised approximately 60 per cent or more of the total
algal individuals present in each monthly quantitative
sample. From March 5 through July 6, cyanophytes con-
tributed more than 75 per cent of the total quantitative
counts. On May 6 more than 95 per cent oF the total count
were members of the Cyanophyta.

Figure 30 presents in graph Form a summary of

percentage cemposition of the total algal population

 

Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.
I I I I | l l I I

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Fig. 30.-—Summary of percentage composition by
DiViSions For 200 cm. interval quantitative samples,
November through October.

 

195

monthly during the 12-month study with reference to the
three divisions present in the collections. From November
through August, the dominance of quantitative samples
by cyanophytes was attributable primarily to members oF
the Hormogonales, which during this period contributed
52 per cent or more of the total individual counts monthly.
The algae density curve exhibited on Figure 29 is there-
Fore by and large an expression of relative abundance of
the Hormogonales, particularly the species Phormidium
angustissimum and Schizothrix Freisii. Relatively large
numbers of the species giggghococcus aeruqingggg in
September and October accounted For the cyanophyte
dominance during those two months 0? minimum algae numbers.

Reference to Table 23 indicates that total numbers
of members of the Chlorophyta and Chrysophyta as a
generalization did not Fluctuate to any great extent dur—
ing the 12-month study. Members of the Chlorophyta in
total never contributed more than 14.2 per cent of the
total monthly individual counts during the study. Numbers
of the species Chlorococcum humicola, the primary con-
tributor to the total count of green algae at each monthly
collection showed little significant Fluctuation during
the 12-month study. Trochiscia granulata and 1. species 2,
the only other significant contributors to green algae
numbers during the study, together never exceeded 10
per cent oF the total count at any given monthly col-

lection. The apparently large contribution oF 9.97 per

 

 

 

196
cent of the total September count by Trochiscia species 2
was based on a contribution of only 218 individuals per
cc. of sand.

The apparent pulses of Chrysophyta in November
through January and in August were attributable essentially
to reduction in relative numbers of cyanophytes during
those months. During the apparent pulses of Chrysophyta
in November-January and in August, Botrydiogsis arhiza
was a significant species, contributing 14.4 per cent of
the total algal individual count in December and 9.0 per
cent in August.

Several members of the Order Pennales made signifi-
cant contributions to the psammonflora at the ZOO cm.
interval during the 12-month study. Navicula heuFleri
varieties comprised 7.10 per cent of the total algal
individuals on November 9, 24.98 per cent on January 4,
8.43 per cent on March 5, 11.82 per cent on June 6, and
6.43 per cent on August 6. Navicula radiosa var. tenella
contributed 5.82 per cent of the algae present in the
December 8 samples, and Navicula decussis contributed
4.71 per cent On November 9 and 5.42 per cent on January 4.

From November through January, the algal Flora
of the 238 cm. interval quantitative samples was dominated

by Fhormidium angustissimum, Microcoleus vaoinatus,

 

Botrydiogsis arhiza, and Navicula heuFleri vars. In
November these 4 species contributed respectively 14.8,

15.6, 4.3, and 7.1 per cent of the total individual

 

 

 

 

 

197
species count. Other important contributors to the
November quantitative counts and their respective per-
centages were Navicula decussis, 4.7, and Porphyrosiphon

Fuscens, 7.2. The December contributions by Phormidium

 

angustissimum, Botrydiopsis arhiza, and Navicula heuFleri

 

vars. were in per cent respectively 34.6, 14.4, and 3.0.
Other species present in significant numbers and their
December percentage contributions were Lyngbya aestuari,
7.2; Chlorococcum humicola, 6.3; and Navicula radiosa
var. tenella, 5.82.

In January Phormidium angustissimum contributed
14.1 per cent of the total count and Navicula heuFleri
Vars. 25.0 per cent. Navicula decussis contributed 5.4
per cent of the total count in January and Microcoleus
chthonoplastes 5.9 per cent.

In February 3. gfltflgflgplastgg individuals com-

 

prised 9.0 per cent of the total count and E. anoustissimum
33.6 per cent. In February 81 species were involved in

the total quantitative count of 65,559 individuals per

cc. of sand.

March 200 cm. interval quantitative counts were
dominated by Lyngbya aesturari, 5.1 per cent; Nostoc
oaludosum, 10.2 per cent; Navicula heuFleri vars., 8.4
per cent; and Phormidigm anqustissimgm, 33.4 per cent.

Phormidium anqustissimgm dominated the psammOn-

Flora algae in April and May quantitative samples, con-

tributing 40.5 and 81.9 per cent of the total counts in

 

198
those two months respectively. The May dominance of B.
angustissimum was such that no one other of the 59 species
present that month contributed more than 2.3 per cent to
the total count. In April dominance of E. angustissimum
was shared by Anabaena sp. 2, 10.5 per cent; Nostoc
galudosum, 10.5 per cent; and Oscillatoria tormosa,
6.3 per cent.

In June 3. angustissimum continued to dominate
quantitative samples with 17.0 per cent of the total
individuals present. Also numerically important in June
were the species Anabaena sp. 2, 7.6 per cent; microcoleus
chthenoplastes, 5.1 per cent; Phormidium aFricanum,

12.8 per cent; 3. luridum, 6.5 per cent; Schizothrix
Freisii, 5.3 per cent; Chlorococcum humicola, 5.9 per cent;
and Navicula heuFleri vars., 11.8 per cent.

Although in July E. angustissimum contributed
15.2 per cent of the individuals present in quantitative
samples, the species gynechococcus aeruqinosug was most
abundant with a 17.2 per cent contribution. Anabaena
sp. 2 contributed 10.5 per cent, microcolegg chthoflgglgstes
and Chlorococcum humicola each contributed 5.0 per cent,
and Schizothrix Freisii 13.6 per cent.

August quantitative samples were cominated by
Schizothrix Freisii, which contributed 22.7 per cent of
the total count, and to a lesser extent Microcoleus

M

chthonoglastes, 14.3 per cent; Synechococcus aeruginosus,

10.5 per cent; Botrydiopsis arhiza. 9.0 per cent;

 

 

 

 

199
Anabaena sp. 2 and E. angustissimum both contributing 5.3

per cent; Navicula hungarica var. linearis, 7.5 per cent;

 

and Navicula heuFleri vars., 6.4 per cent.

The rapidly diminishing September algal Flora in
200 cm. quantitative samples was dominated by g.
aeru inosus, which contributed 35.? per cent oF the count,
Microcoleus chthonoplastes and E. angustissimum both
contributing 8.4 per cent, Trochiscia sp. 2, 10.0 per cent,
Schizothrix Freisii contributing 6.7 per cent, and Navicula
hungarica var. linearis, 6.6 per cent. 0F the 1602
individuals present per cc. of sand in the October 200 cm.
quantitative samples, g. aeruginosus contributed 27.3
per cent, 3. angustissimum, 19.6 per cent, Schizothrix

Freisii, 24.2 per cent, and Chlorococcum humicola con-

 

tributed 5.7 per cent.

Biological data, gualitative
psammon samples, 100 cm; interval

Table 27 presents the results of qualitative and
quantitative sampling From the beach transect at the
100 cm. interval during the period November 9, 1963, to
October 10, 1964. Table 26 summarizes data From Table
27 with respect to Order and Division qualitative totals.
(See pp. 44-47 For methods utilized in data collection.)

From qualitative samples taken at the 100 cm.
interval, 118 species were identiFied during the 12—month
Sampling period. The maximum number 0F species, 72, was

present in January 4 samples, and the minimum species

 

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210
number oF 45 was reported From September 5 and October 10
samples. As indicated on Table 28, during the 12-month
sampling period the most diversity 0F species composition,
61 to 72, occurred during the December through March
period when the collecting site was solidly Frozen. An-
other pulse in number 0F species, primarily attributable
to the Pennales, occurred in the July 6 samples when 64
species were reported. Following the July pulse, a marked
decline in number oF species was observed to the end of
the sampling period.

The contribution oF cyanophyte species to the
qualitative monthly species list at the 100 cm. interval
ranged From a low oF 33.3 per cent 0F the total species
list to a maximum 0F 53.3 per cent on September 5. The
mean contribution oF cyanophytes to the total species
list each month was 40.2 per cent. This signiFicant con-
tribution was largely attributable to members oF the
Hormogonales, which with 20 species contributed From a
maximum oF 44.4 per cent oF the total species list in
the September collection to a minimum oF 23.5 per cent
with 12 species in the June 6 species list. Species of
the Hormogonales were qualitatively most abundant in
early winter and mid-summer and least abundant in late
spring and early summer.

Species cF Chlorophyta were relatively qualita-
tively most abundant in those months in which the beach

was solidly Frozen and least abundant in late summer and

 

 

 

 

 

211
early Fall. At no time during the 12-month study did
chlorophytes comprise more than 24.6 per cent 0F the
qualitative species list (February 5) or less than 6.7
per cent (September 5 and October 10). The monthly mean
centribution oF green algal species during the 12-month
period was 17.7 per cent oF the total species present.
This contribution was attributable essentially to members
of the Chlorococcales and Zygnematales, particularly the
Former, which in February and March contributed 17.4 and
19.0 per cent oF the total qualitative species list
respectively.

Except in January and February when they contri-
buted 4.2 and 2.9 per cent respectively of the total
species list, members oF the Euglenophyta were insigniFi-
cant contributors to qualitative samples.

members 0F the Chrysophyta contributed From 31.9
to 46.9 per cent 0F the total monthly qualitative sample
species lists with a mean monthly contribution oF 41.2
per cent. This contribution to the total species list
was essentially ascribable to members oF the Pennales,
members oF neither the Heterococcales nor Centrales cen—
tributing more than 2.0 or 3.9 per cent respectively to
the monthly species lists. No clearly apparent seasonal
variation in abundance oF diatom species could be dis-
cerned, although qualitatively diatoms were least abundant
in September and October and most abundant January through

July.

 

 

 

 

212

Twenty-Four species oF algae were present in
qualitative collections at the 100 cm. interval through-
out the 12-month sampling period. These included
§yflg§hococcus aeruginosus, leindrosgermum minimum,

Lynobya aerugineo—caerulea, Nostoc paludosqm, Phormidiqm
angustissimum, Phormidiqm luridqm, Schizothrix Friesii,

 

Chlorococcum humicola, Tetraedron mininqm, Euastrum

insulare, Achnanthes linearis F. curta, A. minutissima,

“—..— —-

 

 

C mbella aFFinis C. microce hala Navicula cryptocephala
_1_______.______9 _ v __ 9

 

fl. EgsEidata, fl. heuFleri var. heuFleri, fl. heuFleri
var. leptocephala, fl. hunoariqa var. linearis, fl. minima,

N. radiosa var. tenella, Hitzschia palea, and Rhogalodig

 

gibba. Absent only in October was Lyngby§.gieronymusii,
and Phormidigg aflggglg was absent From qualitative samples
only in June.

Biological data, quantitative

psammon samples, 100 cm. interval

Figure 31 presents in graph Form the monthly

 

Variation in total number oF algal members 0F the psammon
Flora per co. in quantitative 2.4 cc. samples taken dur-
ing the 12-month study. In the quantitative samples
collected at the 100 cm. interval oF the beach transect,
total number oF individuals per cc. of sand ranged From

a minimum oF 4,887 on April 6 to a maximum oF 31,926
recorded in December 8 collections. Comparison oF

Figure 31 with Figure 25 indicates relatively little

relationship between seasonal variation in algal abundance

 

 

  

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213
N v. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.
40%. Hr. - NH. . . .

M
D

N
D
l

A 4.4.344

thousands oF individuals per cc. oF sand
Ch Kl mtOD—tNmDUT
I

 

 

 

 

Fig. 31.—-Total number oF individual algal units
per cc. oF sand, quantitative monthly samples, 100 cm.
interval.
at the 100 cm. interval and that in quantitative lake
PhytOplankton samples as Far as month by month cemparison
is concerned. It should be noted that both Figures,
however, do show limited relationship in that both in~
dicate an abrupt decrease in algal abundance in the march-

April period oF thaw Followed by early summer recovery to

 

. 1 at}!

214

an abundance level approximately that oF the initial
collections in November. The June-July depression Fol-
lowed by the September—Uctober ”bloom" exhibited by the
lake algal community, however, was not exhibited at the
100 cm. interval where, instead, a June-July pulse was
Followed by a gradual decline in algal abundance From
July onward to the last collection in October.

Comparison oF Figure 31 with Figure 27 indicates
a somewhat closer agreement oF seasonal variation in
quantitative algal abundance between the 100 cm. interval
and that oF the SOD cm. interval as does comparison oF
Figure 31 with Figure 29, which presents in graph Form
seasonal variation in algal abundance at the 200 cm.
interval. A more detailed analysis oF seasonal variation
in algal abundance in the lake and the 300, 200, 190,

and 25 cm. intervals is given later in this chapter.

 

It should be noted at this point, however, that the 300,

 

200, and 100 cm. intervals all exhibited a marked maximum
in quantitative algal abundance during the period in which
the beach was solidly Frozen, Followed by a precipitous
decline in the March-April period oF thaw. All three
intervals exhibited an April-May recovery oF algal
abundance to or exceeding that oF the initial November
collection prior to the congealing OF the environment,
Followed by a June slump, partial recovery in July-
August, and more or less rapid decline in algal abundance

in the September-October collections.

 

 

 

 

215

At minimum algal population density oF the 180 cm.
interval in April, density oF algae at the 100 cm. interval
exceeded that oF the maximum lake phytoplankton density
inSeptember by a Factor oF approximately 48. flaximum
algal population density at the 100 cm. interval in
December exceeded that oF the September lake quantitative
samples by a Factor oF approximately 268.

Table 29 presents data 0F quantitative abundance
oF psammon algae at the 100 cm. interval on the basis
0F species percentage composition oF total individuals
counted each month oF the 12-month sampling period.
Table 30 presents a summary oF data From Table 29 with
species data being grouped to Order and Division. All
percentages are rounded oFF to the nearest 0.01 per cent.

Data From Tables 29 and 30 indicate that, in
quantitative samples collected at the 100 cm. interval,
members oF the Cyanophyta and Chrysophyta dominated the
100 cm. interval psammon algae community throughout the
12-month period oF the study. Members oF the Cyanophyta
contributed From a maximum oF 59.44 per cent oF the
total number oF algae enumerated in December 8 samples
to a minimum 0F 23.34 per cent in April 6 samples.
From December 8 to February 5 cyanophytes contributed
54.47 to 59.44 per cent oF the total count. Cyanophytes
apparently diminished in quantitative importance at the
100 cm. interval during the March—April period. In

May, cyanophytes began a gradual return to their position

 

 

 

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225
of dominance until they reached 50.22 per cent abundance
in August 5 samples. In September and October, cyanophytes
abruptly diminished in relative importance (see Fig. 32).
At no time during the study did cyanophytes contribute
less than the combined monthly totals contributed by
members of the Chlorophyta and Euglenophyta. The sig-

nificant contribution of the Cyanophyta to the 100 cm.

 

 

Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.

 

 

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rcentage composition by
quantitative samples,

Fig. 32.--Summary of p8
Divisions For 100 cm. interval
November through October.

 

 

 

 

226
interval algal community was attributable primarily to
members of the Hormogonales. At no collection date did
members of the Hormogonales contribute less than 19.39
per cent of the total individual algae present nor less
than 66.37 per cent of the total number of blue-green
algae present. Apparent seasonal variation in quantita-
tive abundance of blue-green algae at the 100 cm. interval

was largely produced by Fluctuation in numbers of Lyngbya

 

aerugineo-gaerulea. L. hieronymusii, Microcoleus
chthonoplastes, E. vs inatus, Oscillatoria sglendng,

Phormidium angustissimum, Phormidigm luridum, Schizothrix

 

Friesii, and Symgloca kieneri.

Reference to Tables 29 and 30 reveals that members
0? the Chlorococcales were primarily responsible For the
somewhat limited numbers of the Chlorophyta of the algal
Flora of the 100 cm. interval. Members of the Chlorophyta
were relatively most abundant in the months December
through March with maximum contribution of 16.29 per cent
of the total count occurring on March 5. Fluctuations
in quantitative abundance of the green algae during the
study were essentially expressive of abundance oF

Chlorococcgfl humicola, Oocystis elliptica var. minor, and

 

Tetraedron minimum.

Members of the Chrysophyta, particularly the
Pennales, dominated the 100 cm. interval algal Flora
from march S to October 10 (see Fig. 31) except For the

August 6 collection in which large numbers of Schizothrix

 

 

 

 

 

16”“? a .

227

friesii and Microcoleus vaginatus together with a sudden

dimunition of diatom abundance resulted in a temporary
dominance by blue—green algae. At no time during the
12-month study did the Pennales render less than the 25.89
per cent contribution of the December 8 collection nor
more than the 67.93 per cent contribution of the April 5
samples. Fluctuations in monthly abundance of diatoms

at the 100 cm. interval were largely attributable to
abundance of Cymbella microcgphala. flavicula decussis,

Navicula heuFleri varieties, fl. hungarica var. linearis

 

and fl. radiosa var. tenella.

Quantitative samples at the 100 cm. interval
were characterized throughout the study by large numbers
of Navicula heuFleri varieties heuFleri and legtocephala
which together contributed From 12.46 (February 5) to
47.13 per cent (September 5) 0? the monthly total counts.
Together with Phormidium angustissimgm, Schizothrix
Friesii, and flagicula decussis, flavicula heuFleri domin-
ated quantitative samples From November through January,
the Four collectively contributing 52.38, 35.66, and 52.92
per cent respectively of the total counts in those months.
Other important species and their per cent contributions
in November samples were Lyngbya aerugiqgg—caerulea,
6.06; L. hieron musii, 6.44; and Navicula radiosa var.

tenella, 6.55. In December 8.15 per cent of the 31,926

 

species present per cc. of sand were classified as

Anacystis montana. And in January Micr0coleus vaginatus

 

 

 

 

 

228
and Qggillatoria tenuis each contributed 6.93 per cent of
the total count.
In February the species Lyngbya hieron musii,

Microcoleus va inatus, Phormidium an ustissimum, and

 

Navicula radiosa var. tenella together with y. heuFleri
varieties comprised 40.70 per cent of the total species
count of 12,552 individuals. Together with Phormidium
luridum, Navicula decussis, and fl. trigunctata, fl. hgyjlggi
vars. contributed 31.54 per cent of the algal species
present in March. And in April the 30.85 per cent con-
tributed by Navicula heuFleri varieties with Phormidium

anoustissimum, Cymbella microcephala, and Navicula

 

decussis comprised 48.53 per cent of the 4,887 individual
algae present per cc. of sand.

Phormidium apgustissimum, E. luridgm and the two

 

varieties of Navicula heuFleri accounted For 50.00 per
cent of the total count in May. These three species
together with the 6.80 per cent contributed by Rhogalodia
gibgfl included 53.76 per cent of the individuals present

in June. Oscillatoria splendida and Navicula heuFleri

 

along contributed 48.22 per cent of the 23,451 individuals
per cc. of sand collected on July 6. Three members of
the Hormogonales, Micr0coleus chthonoplastes. Schizothrix

Friesii, and Symgloca kisneri, with fiavicula heuFleri

 

comprised 53.86 per cent of the individuals present in
August. And in September, Navicula heuFleri vars. and

Oscillatoria agardhii contributed 53.81 per cent of the

M—

 

 

 

 

g . ell

 

 

 

229

total count. In October, 61.31 per cent of the 11,277
individuals present in each cc. of sand were assigned
to Qgcillatoria tenuis. Symploca kieneri, flavicula decussis,
or Navicula heuFleri varieties.
Biological data, qualitative
psammon samples, 25 cm. interval

Table 31 presents the results of qualitative and
quantitative sampling From the beach transect at the +25
cm. interval during the period November 9, 1963, to
October 10, 1964. Table 32 summarizes data From Table 31
with respect to Order and Division qualitative totals.
(See pp. 44 - 47 For methods utilized in data collection.)

Qualitative samples taken at the 25 cm. interval
monthly during the 12-month sampling period yielded a
total of 131 identifiable species. Total number of
species present in monthly qualitative samples ranged
From a maximum oF 71 species reported in November,
February, and March to a minimum oF 18 present in April 6
samples. Table 32 indicates greatest species diversity
occurred From November through March. During this period
the +25 cm. interval was 25 cm. distant From the shoreline,
and in December through March the beach was solidly Frozen.
Subsequent to the March 5 collection and prior to April 6,
the break-up of lake ice cover resulted in clearly evident
disruption of the transect substrate up to and including
the +112.5 om. position. On April 6 and Way 6, the +25 cm.

interval was under approximately 5 cm. of water (see pg. 32).

 

 

 

 

230

 

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241

By June 6, the + 25 cm. interval was again exposed and
about 50 cm. distant From the shoreline. Recovery of
species diversity, essentially a reflection of increase
of Pennales species, produced a second peak in species
diversity of 69 species in July. By October 10, a pro—
gressive decline in species abundance resulted in only 50
species being present.

As was true of qualitative samples From the
+300, +200, and +100 cm. transect intervals throughout
the 12-month sampling period, cyanophytes and chrysophytes
dominated the psammonflora algae. Fluctuation in total
number of species present each month was essentially
attributable to variation in the number of species which
were assigned to the Hormogonales and Pennales.

The contribution of cyanophytes to monthly quali-
tative species lists ranged From a maximum of 38.9 per cent
in the April 6 collection to a minimUm of 23.9 per cent
of the total species list in the November 9 collection.
The mean contribution of members of the Cyanophyta to
monthly qualitative species lists was 32.4 per cent, with
contributions above the mean occurring in February, April,
and June through October. At no time did members of the
Chroococcales contribute more than 11.6 per cent of the
total species list nor more than 41.2 per cent of the
total monthly cyanophyte species contribution, both
occurrences taking place in the January 4 collection. on

a 12-month mean basis, species assigned to the Chroococcales

 

 

 

 

242

 

contributed no more.than 25.2 per cent of the total
cyanophyte species present monthly.

Consequently, the qualitative importance of
cyanophytes to monthly species counts was attributable
largely to members of the Hormogonales. Species assigned
to the Hormogonales were most abundant February through
March and again June through September, although they were
relatively most important (33.3 per cent) From the basis
of a percentage contribution to the total species list
in the April 6 collection.

The two species assigned to the Euglenophyta
were of little importance in qualitative species lists.

Species of the Division Chlorophyta were most
abundant in November and December when 15 green algal
species were recorded, but in those months they contributed
only 21.1 and 21.4 per cent respectively of the total
species count. In the April 6 collection, no green algae
were reported. The monthly mean contribution of green
algae species during the 12—month sampling period was
14.1 per cent of the total species present. Only in
September were members of the Chlorococcales less abundant
than species assigned to the Zygnematales, but even in
that month members of the Zygnematales comprised only 9.7
per cent of the total species count.

Species assigned to the Division Chrysophyta con~
tributed FrOm a maximum of 64.0 per cent of the total

Species list in the May 6 collection to a minimum of 43.7

 

 

. LCWNIMLF , « a.

243

per cent in February with a mean monthly contribution of
52.6 per cent. This dominance of qualitative samples
was ascribable to members of the Pennales, members of the
Centrales contributing at maximum only 5.6 per cent of
the total species list in the May 6 collection and on a
12-month mean basis contributing only 6.3 per cent of the
total monthly chrysophyte species present. Diatom species
were most abundant in November through March and May
through September.

Only 13 species of algae were reported present
in qualitative collections From the 25 cm. interval through-
out the 12-month sampling period. These included
Qscillatoria tenuis, Phormidiufl angustissimum, Cyclotella
compta, Achnanthes liggagig F. gurtg, Cymbella affinis,
E. microce hala, Navicula or toes hala, fl. decussis,
fl. heuFleri var. heuFleri, fl. heuFleri var. leptocephala,
fl. minima, fl. radiosa var. tenella and Rhopalodig gibgg.
In addition, eight species were absent only in April.

These included Spirulina subsalsa, Tetraedron minimum,

 

Euastrum insulare, Cymbella sp. 1, Navicula cus idata,
fl. tripunctata, Nitzachia acicularflides, and flitgsghifl
SD. 4. Phormidiqm luridqm was absent From qualitative
CDllections only in January. Thus 16 of the 22 species
which were present at the 25 cm. interval throughout the

collection period were diatoms.

 

 

 

. 1 2 [It]:

fit. . - ... .

 

244

Biological data, Quantitative
psammon samples. 25 cm. interval

Figure 33 presents in graph form results relative

 

to monthly variation in total number of algal members of
the psammonflora per co. in 2.4 cc. quantitative samples
taken at the +25 cm. interval during the 12-month study.
Total number of individuals per cc. of sand ranged from
a minimum of 1186 on April 6 to a maximum of 75,955
reported in December 8 samples. It should be noted that
the precipitous decline to the population minima recorded
in April and May was concomitant with late Harch ice
movements which disrupted the beach transect up to and
including the +112.5 cm. position and with inundation
with approximately 5 cm. of lake water on the April 6 and
May 6 collection dates.

In general, the algal companent of the psammon
flora at the 25 cm. interval showed an initial density

increase in December as the transect substrate became

 

congealed Followed by a subsequent regular precipitous

decline during the January-march period when the beach 1
was apparently solidly frozen. The violent disruption of .
the substrate by late March ice break-up and subsequent é
inundation of the 25 cm. collection site of April and T
day 6 collecting dates precludes any other explanation

Of the April-May population minima. In June and July

collections a gradual recovery of the algae of the psammon-

flora is evident. A marked decline in the August 6

 

 

 

-__..._

—...__.‘_.__....__

Nov.

DeCo

Jan.

245

 

90
BO ‘
7O -

SO'
40'

30‘

204

_\

thousands of individuals per cc. of sand
.J—\ 01 m \‘ICDKDD

 

Feb. Mar. Apr. May June July Aug. Sep. Oct.
I I I 1 I I '

 

 

Fig. 33.--Total numbers of individual algal units

per cc. of sand, quantitative monthly samples,

interval.

25 cm.

 

 

 

 

fi

 

 

246
samples was Followed by September 5 recovery beyond that
of July 6 but still short of the December 8 peak. In
October 10 collections the general decline of psammon
algae abundance evidenced at the other three transect
sites was again apparent at the 25 cm. interval.

Detailed comparison of seasonal variation of lake
and psammon algae population densities with that of the
25 cm. interval is made in the biological data recapitu-
lation section of this chapter, but at this point it may
be noted that comparison of Figures 25 (lake), 2? (300 cm.
interval), 29 (200 cm. interval), and 31 (100 cm. interval)
indicates that no direct and obvious correlation was seem-
ingly evident between month by month variations in algae
population density of the lake or any of the transect
collection sites.

Minimum algae population density at the 25 cm.
interval on April 5 exceeded that of the maximum lake
PhytOplankton density per comparable sampling volume in
SeDtember by a Factor oF approximately 1D. Laximum algae
PODUlation density oF the 25 cm. interval in December
exceeded that of September lake quantitative samples by
a Factor of approximately 6 9.

Table 33 presents data of quantitative abundance
of psammon algae at the 25 cm. interval on the basis of
species percentage composition of total individuals
counted each month of the 12-month sampling period. Table

34 presents a summary of data From Taole J3, the speCies

 

 

 

 

 

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Q53; ‘ 4‘

256
data from Table 33 being grouped to Order and Division.
All percentages are rounded off to the nearest 0.01 per
cent. Figure 34 in graph form presents a summary of
total algae population percentages monthly during the
12-month study with reference to the four divisions

present in the collections.

 

f

Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.
| I ' j v I F |
Cyanophyta
90- —-——— Chrysophyta
————— Chlorophyta
......... . Euglenophyta

 

80'

70'

60'

50"

 

40-

30‘

20

10‘

 

 

Fig. 34.--Summary of percentage composition by
Divisions For 25 cm. interval quantitative samples, November
through October.

 

257

Data from Tables 33 and 34 indicate that in quan—
titative samples the Chrysophyta numerically dominated
the 25 cm. interval psammonflora algae From November
through March, in June, and again in October. From
November through March the chrysophytes contributed from
51.4 per cent (November) to 72.8 per cent (March) of the
total individual algae present in each monthly quanti-
tative sample. Ice action and subsequent inundation of
the 25 cm. interval in April and May reduced the numbers
of Chrysophyta individuals to approximately 3.2 per cent
that of the March 5 collection. A slow recovery of
chrysophyte abundance, particularly by individuals as-
signed to Nitzschia spp. and Navicula spp., resulted in
temporary resumption of dominance in the still depressed
algal flora of the June 6 collection. A decline in
diatom abundance in July and August coupled with a
population pulse by Oscillateria splendida and species of
Phermidium resulted in loss of chrysophyte abundance in
July and August. Chrysophytes again resumed dominance in
September and October when pulses in numbers of indi-
viduals of Navicula spp. and Nitzschia spp. were not
equalled by concomitant pulses of members of the Cyanophyta,

Members of the Cyanophyta, as indicated on
Figure 34, dominated quantitative collections from the
25 cm. interval only in the disruption period of April-
May and later in July 6 and August 6 collections. The

April-Nay dominance was essentially a reflection of

 

 

 

 

258

reduced numbers of members of the Pennales, complete
absence of members of Euglenophyta and Chlorophyta, and
the survival of populations of Microcoleug chthonoplastes,
Lyngbya spp. and Phormidium spp. during the disruption
induced by lake ice break-up. The July-August dominance
was attributable to reduced diatom abundance and pulses
in abundance of Oscillatoria splendida and Phormidium
angustissimum. -

Members of the Division Chlorophyta attained
maximum percentage contribution of 14.7 per cent in
December 8 samples when 13.12 per cent of the individuals
present were assigned to the Chlorococcales. The
fluctuations in Chlorophyta abundance presented on
Figure 34 express the relative abundance of Chlorococcum
hgmicola. Tetraedron minimum, and I. muticum, other
members of the Division rarely exceeding 0.5 per cent
of the total individuals present in any monthly quanti-
tative sample taken at the 25 cm. interval. Together with
members of the Euglenophyta, green algae never exceeded
14.63 per cent of the individuals present in any month.

The algal camponent of the psammonflora of the
25 cm. transect interval, in short, was throughout the
year dominated by members of the Hormogonales and Pennalee.
Particularly common were the cyanophytes Lyngbya
aerugineo-caerulea, E. hieronymusii, Microcoleus
chthonoglastes, fl. va inatus, Oscillatoria tenuis,

Phormicium anqustissimum, E. luridum and the diatoms

 

 

 

259

Navicula heuFleri var. le toce hale, fl. cryptoceghala,

 

 

and fl. radiosa var. tenella.

 

From November 9 through March 5, Navicula heuFleri
var. legtocephala was the most abundant organism in 25 cm.
interval quantitative samples. In November collections,
19.6 per cent of the individuals present were assigned
to this taxon. Also common among the 51,171 individuals
per cc. of sand were Lyngbxa hieronymusii (5.62 per cent),
Phormidium angustissimum (10.1 per cent), and Spirulina
subsalsa (4.9 per cent). The 30.7 per cent contributed
by E. heuFleri varieties in December together with the
17.3 per cent contributed by Microcolggg chthonoplastes
and 7.3 per cent by Chlorococcum humicola comprised
55.2 per cent of the 75,955 individuals present per cc.
of sand in the December 8 collection. The declining
numbers 0F algae present in the January 4 collections
were dominated by the 43.3 per cent contributed by E.
heuFleri. Also common in the 24,868 individuals per cc.
of sand reported in January were Phormidium angustissimum
(9.1 per cent), Chlorococcum humicola (6.6 per cent),
and Navicula radiosa var. tenella (5.9 per cent).

February quantitative collections From the 25 cm,

interval were characterized by N. heuFleri, Phormidium

 

angustissimum, Navicula decussis, and fl. radiosa var.

 

tenella, which respectively contributed 29.5, 8.4, 10.6,
and 6.1 per cent of the 9,933 individuals present per cc.

of sand. And in March important contributors to the still

 

 

 

water‘a. ‘ .‘

268
declining population of algae, now only 8,726 individuals
per cc. of sand, were N. heuFleri (31.9 per cent), N.

~—-—-.—-— —

radiosa var. tenella (9.4 per cent), fl. decussis (6.3 per

 

cent), and Phormidium angustissimum (5.4 per cent).

As noted previously, the apparent dominance of
the algae by cyanophytes in April 6 and May 6 quantitative
collections made at the 25 cm. interval was attributable
not to large numbers of these taxa but rather to the
disappearance of other species resulting From severe
disruption of the environment as a result of ice action
and rising lake levels. The 37.8 per cent contribution
of E. angustissimum to the 1186 individuals per cc. of
sand on April 6 was through a count of only 389 individ-
uals of this taxon per cc. of sand, the lowest density of
E. angustissimum throughout the 12-month study. Only
eight species of algae were recorded in April 6 quanti-
tative samples. Microcoleus chthonoplastes contributed
27.9 per cent of the total count and fl. heuFleri 9.8
per cent. In each cc. of the submerged sand of the May 6
collection site 1206 individuals belonging to 11 taxa
were present. 0? these, 46.8 per cent were E.
angustissimum, 9.7 per cent were assigned to Lyngbya
hieronymusii, and fl. heuFleri varieties and fl. radiosa

var. tenella each contributed 12.9 per cent.

 

Following the emergence of the 25 cm. collecting
site, 4914 individuals assigned to 33 taxa were reported

per cc. of sand in the June 6 quantitative collection.

 

 

 

 

he?» .®_ ~

261

Phormidium angustissimum contributed 21.3 per cent of these,

3. luridum contributed 7.9 per cent, and Navicula heuFleri

 

contributed 11.9 per cent. Other important contributors
were Nitzschia subcapitellata and Nitzschia acicularkfdes,
which contributed respectively 7.9 and 12.6 per cent of
the total count.

OF the 26,194 individuals present per cc. of sand
in the July 6 collection, 34.0 per cent were assigned

to Oscillatoria s lendida, 19.1 per cent were assigned

 

to Navicula heuFleri, 14.5 per cent to Microcoleus

 

chthono lastes, and 7.6 per cent to Oscillatoria tenuis.

Phormidium angustissimum contributed only 4.1 per cent.

 

But in August 6 collections, E. angustissimum regained
dominance at 33.8 per cent, partially because of greatly
reduced abundance of other taxe. Other important con-

tributors were fl. heuFleri (18.2 per cent) and Microcoleus

 

vaginatus (7.0 per cent).

In September collections, algal density per cc.
of sand at the 25 cm. interval reached the highest level
since the December 8 collection. OF the 35,711 individuals
present per cc. of sand, 39.8 per cent were assigned to
E. heuFleri, 11.5 per cent to E. ngustissimum, 10.4 per

cent to Microcoleus vaginatus and 4.3 per cent to Navicula
cryptocephala. The October collection was dominated by
E. heuFleri and E. ngustissimum which respectively con-
tributing 33.3 and 14.7 per cent 0F the total count com-

prised 48.0 per cent of the 24,395 individuals present per

 

 

 

 

262
cc. of sand. Also important on a percentage basis were
microcoleus va inatus, (10.2 per cent), Nitzschia

subcagitellata (8.9 per cent) and Navicula cryptocephala
(8.8 per cent).

Biolo ical data reca itulation
of results 0? monthly Qualitative
and Quantitative sampling For
12~menth study period

Table 35 presents a summation of qualitative and
quantitative contributions of lake and psammon zone algae
during the 12-month sampling period. In the table, the
number of months in which a taxon was present in qualita-
tive samples taken From the lake or the 25, 100, 200,
or 300 cm. intervals of the beach transect is given as
a whole number in the upper portion of the line allotted
each taxon. In the lower portion of the allotted space,
the whole number represents the number of months of the
12 in which the taxon was present in quantitative col-
lections taken at that site. The decimal indicates the
mean percentage contribution of that taxon to total
quantitative counts of those months only in which the
taxon was present at that given site. Although neither
whole number is illustrative of seasonal variation, the
table does give a rough approximation of the relative

importance oF each taxon during the 12—month study.

During examination of qualitative and quantitative

samples From the lake and adjacent beach transect, 261

Species assigned to 14 Orders of five Divisions were

 

 

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268

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272

 

 

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285
tabulated. Of these, 25 species were present in quali-
tative and/or quantitative samples at least once tram
the lake and each of the beach intervals, although not
necessarily simultaneously. Three species -- Cymbella

microceghala, Navicula heuFleri var. leptocephala, and

 

Navicula radiosa var. tenella -- were, however, truly
ubiquitous in that they were present at each of the Five
sites sampled on each of the 12 sampling dates. Present
in both lake and beach samples but not recorded at least
once at each of the Four psammon zone sites along the
beach transect were 80 species. As a generalization,
most of these were absent From the 200 and/or 300 cm.
sites.

It should be noted that no taxon was counted
present unless it apparently was living at or shortly
before the time of collection. Diatoms and desmids,

For example, were not recorded if there was no organic
material present within the cell walls. A Far longer
qualitative species list -- particularly For diatoms and
desmids -- would have resulted if empty cells were included.
Numerous empty cell walls of species of both groups were
Found in the sand which were never found present in the
living state at any time during the study. Inasmuch as
positive identification 0F diatoms and many desmids requires
observation of empty frustules or semicells, an element

of error is obviously inherent in construction of species

lists For the psammon algae community. The sand of the

 

 

 

 

286
beach transect acts as a natural Filter to trap dead cells
oF many algae brought to the beach by wave action or
drainage From Further up the beach. Error in construc-
tion oF species lists and counts oF living Forms in
quantitative samples can be reduced somewhat by the rather
laborious and time consuming process oF matching camera
lucida drawings oF living Forms with cleaned material
taken as aliquots From the same sample. Ergo, very
similar Forms such as flaviggla heuFleri var. heuFleri and
var. lsptocephala were oF necessity "lumped” in raw data
because oF the diFFiculty in differentiating between them
in the living state.

Seventy-Four species were Found only in samples
taken From the beach transect intervals. At no time were
these species Found in an apparently living state within
lake samples. Eleven species —— eight oF these diatoms --
0F the 74 were Found only at the 25 cm. interval, Five
were Found only at the 100 cm. interval, six were Found
only at the 200 cm. interval, and Four species were re-
ported only at the 300 cm. interval.

Eighty—two species oF algae were reported present
only in lake samples.

DF the 26 species oF Chroococcales recorded dUr_
ing the study, nine were Found only in the lake, six were

Found only in the beach, and Five -- Anacystis marginata,

A. montana, Chroococcus turqidus, Gloeothece rupestris.

and gynechococcus aeruginosis —- were more or less

 

 

 

 

287
ubiquitous in that they were present at least once at
each beach interval and also in the lake. Ten of the
54 species assigned to the Hormogonales were found only
in the lake, 32 species were found only in the beach, 10
were Found in both beach and lake but not all beach

intervals, and the species Aulosira laxa and Nostoc

 

Qaludosum were Found at least once in the lake and at each
beach interval.

Five of the five species assigned to the Order
Euglenales were found only in the beach, only Trachelomogas
PlayFairii being present at more than two beach intervals.
It is believed that numerous species of euglenoids escaped
identiFication during the study. Although encysted forms
which apparently were euglenoids were Frequently en-
countered in beach samples, prolonged culturing did not
induce excystment, and hence identification was precluded.
All seven of the seven species assigned to the Peridiniales
were found only in lake samples.

In the Division Chlorophyta, Order Volvocales,
one of the Four species identified during the study was
Found only in the lake, two were present only in the beach
at the 200 and/or 300 cm. interval, and the organisms
identified as Chlamydomonas Snowii were Found in the lake
and at the 100 and 300 cm. beach intervals. Infrequently,
palmelloid colonies of what apparently were members oF
the Volvocales were encountered in beach samples. These

Forms resisted attempts to induce through culturing

 

 

 

 

 

 

288
resumption of the motile stage needed For identification.
Such Forms were not reported in either qualitative or
quantitative samples, but it is believed that at best they
contributed little to total number of species or indi-
viduals in either lake or beach samples. That members
of the Volvocales are apparently not of major qualitative
importance is indicated in the literature of investiga-
tion of Freshwater psammon flora (Sassuchin gt 9;, 1927;
Neel, 1948). Noel reported only three species of the
Order Uolvocales in a list oF 173 species of algae identi-
Fied as a result oF extensive sampling from the psammon
of Douglas Lake.

Of the two species of Tetrasporales reported in
the 12-month study, Gloeocystis vesiculosa was Found
only one time each at the 100 and 300 cm. intervals, and
Q. gigs; was found only two months in the lake and once at
the 100 cm. interval.

Nine of the 33 species assigned to Order
Chlorococcales were reported only From the lake, and
seven oF the 33 were found only in beach samples, three
of these being unique to only one interval. Pediastrum
Boryanum, Tetraedron minimum, and I. muticum were present
in the lake and at least once at each beach interval.
Fourteen species at some time during the study were pre-
sent both in the lake and in beach samples but were not

present at least once at all Four beach intervals.

 

 

...?‘WM' ‘ <

289

Of the 21 species assigned to the Zygnematales,

10 were found only in the lake, two were present only in
beach samples, and nine were found both in lake and beach
samples but not at all beach intervals. None were reported
from the +300 cm. interval.

Assigned to the Division Chrysophyta were 104
species belonging to a total of five orders. 0f the
four species of Chrysomonadales, all were found only in
the lake. Phaeothamnion Borzianum, the single species
assigned to the Chrysotricales, was found only at the
300 cm. interval, and of the three members of the
Heterococcales, none were found closer to the shoreline
than the +100 cm. interval.

Three of the seven species identified during the
study as members of the Centrales were found only in the
lake, Cyclotella meneqhiniaga was found in three months
only at the 25 cm. interval, and four species were found
in both lake and beach samples but not at all beach
intervals.

0f 89 species assigned to the Order Pennales, 26
were found only in the lake and 12 found only in beach
samples, seven of the 12 being present only at the 25 cm.
interval. Fifteen diatom species were present at some
time during the study in lake samples and at least once
at each beach interval. Found in both lake and beach
samples but absent throughout the 12—month study from at

least one beach interval were 36 species of diatoms.

 

 

290

Table 36 presents a summary of data from Table 35.
The total number of species appearing in qualitative
samples from the lake and beach transect intervals at
some time during the 12-month sampling period has been
grouped at Division and Order levels to show species dis-
tribution at each of the sampling sites. Reference to
Table 36 indicates that greatest species diversity during
the 12-month study occurred in lake qualitative samples.
The total number of species present in qualitative samples
from the beach transect sites shows a gradual diminution
with increasing distance from the +25 cm. interval
landward. This diminution is essentially a reflection
of decreasing species diversity in the Hormogonales,
Chlorococcales, Zygnematales, Centrales, and Pennales with
increasing distance from the lake. By and large, the
decreasing number of species landward along the transect
was a result of disappearance of species that were pre-
sent at intervals closer to the lake. Only four of the
62 species present in samples from the 300 cm. interval
were unique to that interval, and only six of the 104
species found at the 200 cm. interval were unique to that
collecting site.

Reference to Table 35 indicates that in con—
sideration of qualitative data only 15 species were found
exclusively at the 200 and/or 300 cm. intervals. Con-
sequently, as a generalization, it would appear that on

the basis of qualitative data the algae most landward

 

 

 

291

TABLE 36.--Division and Order summation of species reported
in qualitative samples taken from lake and beach transect
intervals during 12-month study

 

 

 

 

 

 

 

 

 

 

25 100 200 300 total at
lake cm. cm. cm. cm. all sites
Chroococcales 20 11 11 10 10 26
Hormogonales 22 33 33 33 15 54
CYANDPHYTA 42 44 44 '44 25 80
Euglenales 0 2 4 2 1 5
EUGLENOPHYTA o 2 4 2 1 '5
Volvocales 2 0 1 1 3 4
Tetrasporales 1 D 2 0 1 2
Dedogoniales ' 3 o o 2 o 5
Chlorococcales 26 17 18 14 5 33
Zygnematales 19 9 11 2 0 21
CHLDROPHYTA 51 26 32 19 9 65
Peridiniales 7 0 0 0 0 7
PYRRHDPHYTA 7 D D U D 7
Heterococcales 0 0 1 3 3 3
Chrysomonadales 4 0 0 0 0 4
Chrysotrichales 0 0 0 0 1 1
Centrales 6 4 3 2 0 7
Pennales 77 55 34 34 23 89
CHRYSOPHYTA 87 59 38 39 27 104
Total species 187 131 118 104 t 62 l 261

 

 

Read table thusly:

During the study, a total of 26 species assigned
to the Chroococcales was reported from lake and beach samples.
In the lake, 20 species of the Chroococcales were reported,
and in the lake a total of 187 species of algae was found
in qualitative samples.

 

 

 

292
along the beach transect were those able to survive the
severe environment of that portion of the transect rather
than a unique group established there in "preference“ to
sites further toward the lake. It should not be over-
looked, however, that 28 species at no time were found
less than 100 cm. from the shoreline, and 17 species were
never found less than 200 cm. distant from the shoreline.
These species —— including all those assigned to the
Heterococcales and Chrysotrichales —- might be considered
"preferentially" adapted to the exceedingly severe environ-
ment of the upper portion of the beach transect. Those
found only at the 200 and/or 300 cm. interval included
Agmenellum thermale, Dactylococcopsis raphidioides
Microcystis elabens, Oscillatoria limnosa, Phormidigm

jadinianum, E. microtomum, Trachelomonas ranulosa,

 

Chlamydomonas sp., E. Frankii, 0ed0qonium sp. 1, 0edogonigm

sp. 2, Scenedesmus incrassatulus var. mononae, ghlorallanthus

 

oblonqus, Chlorobotrys simplex, Phaeothamnion Borzianum,

 

Navicula gapitata, and Radium gubium, To this list might
be added those which also showed qualitative evidence
of "preference“ for the upper portion of the transect,
including Calothrix membranacea, Microcoleus acutissimus,

Nostoc linkii var. arvense, Phormidium apomala, Schizothrix

 

friesii, Achnanthes exi us, and Botrydiopsis arhiza.

 

Table 37 presents a summary of total species

diversity in lake and each beach transect interval monthly

 

 

 

 

293

TABLE 37.-—Summary of total species number in lake and beach
transect qualitative samples monthly during 12-month study

 

 

Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.

 

lake 70 46 30 60 39 72* 74* 75* 90* 88* 76* 57
25 71* 70* 60 71 71* 18 50 55 69 65 62 50
100 55 61 72* 69 63 51 59 51 64 49 45 45
200 55 49 61 81* 51 4s 59 44 46 33 .27 25

300 31 38 39 4D 33 33 30 27 16 15 12 13

 

Read table thusly:

The symbol * indicates the collection site with
greatest number of species reported each month. On November
a total of 71 species was reported from 25 cm. interval
qualitative samples.
during the 12-month study. It is constructed on the basis
of total species number rather than by Orders and Divisions.

Seasonal variation in total number of species
and the relative qualitative importance of the Order and
Division to which each belongs has been discussed at some
length in the sections of this chapter devoted to detailed
results of qualitative sampling at each collection site.

It should be noted again, however, that inspection of

Table 37 reveals that species diversity in lake samples

as evidenced by total species number was least during the
December-March period concomitant with continuous ice
cover and minima in quantitative counts. And, as noted
previously in this chapter, species diversity in the beach

transect intervals was greatest during the December-March

 

.. Q,

1 . .1 ,5,th

 

294
period when the sandy substrate was solidly frozen. From
November through march the greatest species diversity of
all five collection sites occurred in the beach rather
than in the lake. From April through October the lake
became the site of greatest species diversity.

Table 38 presents a summary of data given in
tables 16, 20, 24, 28, and 32 with reference to results
of qualitative sampling from the lake and beach transect
intervals. The table indicates the monthly contribution
of the 14 Orders to the total number of species found in
qualitative samples from each collection site.

Inspection of Table 38 reveals that for a number
of Orders in each month, as a generalization, there was
evidenced a gradual and conspicuous decline in number of
species landward along the beach transect. This diminution
along the transect was particularly pronounced in the
Centrales, Pennales, and Zygnematales if it is noted that
the irregularites in numbers exhibited in the 25 and 100
cm. beach intervals occurred during and subsequent to
those months in which ice break—up and advancing water
levels disrupted the environment. The Heterococcales
and Euglenales exhibited, however, a reversal of this
tendency. And the Chlorococcales from November through
July displayed larger numbers of species at the 100 and/or
200 cm. intervals of the transect than at the extremes.
In August through October, the Chlorococcales exhibited

a gradual decrease in species number shoreward from the

 

 

 

 

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299
25 cm. interval. with respect to the Hormogonales and
Chroococcales, and with the exception of July and August,
the greatest numbers of species were reported at the 100
or 200 cm. interval rather than at 25 or 300 cm.

Consideration of Table 38 shows that only in
January, February, April, and May did exception occur to
the gradual decline in species abundance landward From
the shoreline. The April-May exception can be explained
by the disruption of the substrate during those months.
By June, the qualitative population of the 25 and 100 cm.
intervals essentially was back to ”normal" March levels.
The January-February exception may be partially explained
by increases in the number of species of Hormogonales at
the 100, 200, and 300 cm. intervals to levels not reached
at any other time of year.

Although qualitative seasonal variation in the
algae of lake and beach transect was evident at each col-
lection site and has been discussed previously For each
site, seasonal variation at the Order level -- except For
the Hormogonales -- For the entire transect was not so
evident. Further reference to seasonal variation will be
Found later in this chapter.

Table 39 includes a summary of quantitative data
From tables 15, 19, 23, 2?, and 31 with respect to total
number of individual algae per cc. monthly From lake and
beach transect intervals during the 12-month sampling

period. Figure 35 presents the same data in histogram Form

 

 

 

300

 

 

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302

For better visualization 0F relative monthly changes in
quantitative abundance. To Facilitate construction oF
Figure 35, For lake plankton organisms per liter has been
converted to organisms per cc. and multiplied by a Factor
0F x1000. Figure 35 should be compared with Figures 25,
27, 29, 31, and 33 which present in graph Form total
number oF individual algae per liter (lake) or cc. 0F sand
(beach transect intervals) For lake, 300, 200, 100, and
25 cm. beach intervals respectively.

Reference to Figure 35 and to Figures 25, 27, 29,
31, and 33 reveals the marked dissimilitude in quantita-
tive abundance exhibited monthly by the algae at the Five
collection sites. Seasonal Fluctuation in quantitative
abundance at each collection site during the 12—month
collection period has been discussed in some length in
previous sections oF this chapter. Figures 25 and 35
indicate that the density 0F phytoplankton within the
lake underwent a decline From 5801 individuals/liter in
November to the 12—month 10w OF 345 individuals/liter in
March after Four months 0F lake surface ice cover. Sub-
sequent to ice break-up, density of phytoplankton began
gradual increase to a maximum 0F 119,010 individuals/liter
in the September 5 collection, Followed by an abrupt
decline to the October 10 density oF 10,048 individuals/
liter.

None oF the beach intervals showed patterns in

Quantitative abundance seaSOnal variation similar to that

 

 

 

303

of the lake. All beach intervals exhibited maxima in
algae density during the Four~month period when the beach
substrate was Frozen. At the 25 cm. interval, maximum
12-month density oF 75,955/cc. was recorded in December 8
collections, the same collection date that the 100 cm.
interval maximum algae density oF 31,926 individUals/cc.
0F sand was reported. The 12-month density maximum oF
13,296 individuals/cc. was reported at the 300 cm. interval
in January 4 samples, but the 200 cm. interval maximum
0F 65,559 was not reached until the February 5 collection.

Subsequent to the December 8 collection, density
oF the algal component 0F the psammonFlora at the 25 cm.
interval began a gradual and unbroken decline to the
April 6 minimum oF 1186 individuals/cc 0F sand. At the
100 cm. interval, however, the pattern oF December density
increase From lower November levels Followed by decline
to an April 6 12-month minimum was modiFied. Following a
January 4 density level oF 10,369 individuals per cc.,
a February and march partial recovery to a march 6 density
oF 14,229 individuals/cc. occurred before the density
decline resumed to the April 6 minimum 0F 40”? individuals/
cc. At the 200 cm. interval, the decline in numbers oF
algae From the February maximum to the April 6 level of
9555 individuals/cc. of sand was perhaps more pronounced

nterval with respect to magnitude

‘4.

than at any other beach
L‘

in number decrease (see :igure 29). As occurred at the

200 cm. interval the decline in number oF al as From
7

 

 

 

 

 

304
winter maxima to flarch—April minima was abrupt and pre-
cipitous at the 300 cm. interval.

At each beach interval, an algal density maximum
occurring during the December-March period was Followed
by a more or less rapid decline to a markedly reduced level
in April (25 and 100 cm. intervals) or March 6 (200 and
300 cm. intervals) quantitative samples. At each beach
interval, this decline was Followed by an increase in
quantitative abundance oF algae to a level approaching or
exceeding that oF the November collections but Far short
oF the December—March maxima. For the 25 cm. interval
the secondary maximum of 26, 194 individuals/cc. was not
reached until the July 6 collection, the same date upon
which the 100 cm. interval secondary maximum 0F 23,451
individuals/cc. oF sand was reported. Recovery at the
200 and 300 cm. intervals was more rapid, the secondary
maximum 0F 33,038 individuals/cc. occurring in the may 6
collection at the 200 cm. interval and that of 9422
individuals/cc. occurring on April 6 For the 300 cm.
interval.

Subsequent to the secondary peak in algal quanti—
tative density, a general decline in density took place at
the 300, 200, and 100 cm. intervals until the October
termination 0F the 12-month study. At the 300 cm. interval,
the October algae density was approximately one—eighth
that of the November level and about one—third that oF

the March depression. The October algae density at the

 

 

 

 

 

305

200 cm. interval was approximately one-tenth that of
November and one-ninth that of the April depression. At
the 100 cm. interval, however, the October density was
approximately 1% times that of November and three times
that of the April depression. But at the 25 cm. interval,
the October density was about half that of November and

20 times that of the April-May period of disrupted sub-
strate. During the July-October period, algae abundance at
the 25 cm. interval fluctuated irregularly From 9677 to
35,711 individuals per cc. of sand.

Thus, in summary of Figure 35 and Table 39, it is
seen that a more or less inverse relationship in quantita-
tive algae abundance was evidenced between the lake and
each beach transect interval. The Further From the lake,
the more pronounced this inverse relationship became.
maxima in quantitative algae abundance For all beach
interVals occUrred during the December-March period when
the beach substrate was congealed by low temperature. At
each interval, the winter maximum was Followed by a decline
in algae abundance to a March or April depression,

Followed by a more or less gradual recovery in May (300
and ZOO cm.) or July (25 and 100 cm. intervals) at levels
approaching or exceeding those present in November at the
initiation of the study. After this recovery, a gradUal
decline occurred in algae abundance to October record low
levels (ZOO and 300 cm.) or levels appreciably lower than

those of the early summer recovery period (25 and 100 cm.

intervals).

 

 

 

 

 

306

Table 40 presents in tabular Form a comparison oF
relative monthly quantitative abundance oF algal indi—
viduals in the beach transect intervals, November through
October, with respect to lake phytoplankton density as
individuals per cc. oF sample. Each month the number oF
phytoplankters present per cc. oF quantitative samples
has been divided into the total number oF individual
algae per cc. at each beach interval in order to illus-
trate the relative density at each interval with reFerence
to the density oF phytoplankton. Figure 36 presents the
same information in graph Form For better visualization
oF the data. Data From Table 40 should be compared with
those presented on Table 39 For better realization oF the
significance oF relative monthly quantitative abundance.

IF relative nUmber oF organisms per cc. oF sample
is considered an index oF productivity, then reFerence
to Table 40 and Figure 36 suggests that the psammon zone,
particularly 100 cm. or less From the shoreline, was a
remarkably productive environment when compared with the
water oF Reach A1 oFFshore From the beach transect. In
December, For instance, each cc. oF sand at the 25 cm.
interval had an algal density 114,523 times that oF each
cc. oF water collected From the lake the same date. Field
notes taken during the December-march period when the
beach substrate was Frozen make reFerence to the con-
spicuous bluengreen color oF the beach, particularly in

January and February. In February and March, it was noted

 

 

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307

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300

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1Dgngg~°V Déc J3” F?b mar {Pr §Y Juqe July Aug Sep Oct

  

n_n__

80000
950000: -—._ —— 25 cm. interval
‘- - / \' *---- 100 cm. interval
140000 4 \ /' \, -—-—--200 cm. interval
3 \ , \ '°"'°°'°3OO cm. interval
30000

10000
8000

6000

4000
3000

2000

1000

mm
DD
OD

400
300

200

OD

 

 

 

 

 

10

Fig. 36.--Relative monthly quantitative abundance oF
algae in beach transect intervals, November through October,
with respect to lake phytoplankton density.

 

 

 

309
that thawed samples had a suance woFt oF algal Filaments
that resisted disintegration, and it required more than
the usual amount oF agitation to homogenize the sample
For counting. ReFerence to Figures 28, 30, 32, and 34
reveals that the Cyanophyta dominated beach transect
samples during this December-March period at the 300, 200,
and 100 cm. intervals and were important contributors to
the Flora at the 25 cm. interval. Algae species present
in signiFicant numbers at this time at the 300 cm. interval
were Botrydiopsis arhiza, Cylindrospermum minimum,
Microcoleus acutissimus, fl. vaginatus, and Schizothrix

Friesii. At the 200 cm. interval, samples during the

 

December-March period were dominated by Lyngbya aestuari,
Microcoleus vaginatus, fl. chthonoplastes, Nostoc paludosum,

-———-——.—.—.

and in particular, Phormidium angustissimum. And at the

 

100 cm. interval during this period, samples were dominated
by large numbers oF Lynqbya hieronymusii, Nicrocoleus

vaginatus, Phormidium angustissimum, Schizothrix Friesii,

 

and the diatoms Navicula decussis. fl. heuFleri var.
leptocephala, and N. radiosa var. tenella. In 25 cm.
samples, microcoleus chthonoplastes and E. angustissimur
were important contributors in the December-March period,

but Navicula heuFleri var. leptecephala, N. decussis, and

 

 

N. radiosa var. tenella were the dominant species. In
the same period, lake samples were dominated by Anacystis
marginata, A. montana, Phormidium angustissimgm, Lyngbya

aerugineo-Eaerulea,and particularly by members oF the

 

 

310
Chrysomonadales, the latter group not present at any time
in the beach.

ReFerence to Table 40 and Figure 36 reveals that
during the First Five months 0F the study, the relative
algae density in the beach ranged From 114,563 to 1327
times that in the lake. From November 9 to January 4,
the 25 cm. interval was the most productive oF the beach
intervals, members oF the Pennales comprising From 49.2
to 66.5 per cent oF the total numbers oF algae present
at this interval during those months. In February 5 and
March 5 samples, the 200 cm. interval became the most
productive collection site with relative densities 0F
60,591 and 59,217 respectively during those months. In
February and March, the Hormogonales comprised respectively
78.9 and 73.9 per cent oF the algae present at the 200 cm.
interval. Nostoc paludosum and Phormidium angustissimgm
contributed heavily to these percentages.

The conspicuous decrease in relative densities at
beach intervals evident on Figure 36 during the April-June
period was concomitant with increasing density oF phyto-
plankters in the lake and decrease oF algae numbers in the
beach. During this period, as a broad generalization,
members oF the Chrysophyta and Cyanophyta dominated the
Flora in the beach transect intervals. In the same time
period, dominance oF lake phytoplankton by the
Chrysomonadales ended, and members oF the Peridiniales,
Hormogonales, and Chlorococcales became conspicuous in

quantitative samples.

 

 

 

 

 

311
At the 300 cm. interval, quantitatively important
species in the April-June period included Synechococcus

aeru inosus, Cylindrosgermum minimum, Plectonema notatum,

 

Chlorococcum humicola, and Botrydiopsis arhiza. In April

 

and May collections the greatest relative density in the
beach transect occurred at the 200 cm. interval. Important
contributors to this large number 0F algae were Anabaena
sp. 2, Nostoc paludosgm, Phormidigm anqustissimum, E.
aFricanum, Schizothrix Freisii, and Navicula heuFleri var.
legtocephala. By June 6, the site oF greatest relative
algal density had shiFted lakeward to the 100 cm. interval.
At the 100 cm. interval, important contributors to the
relative density index oF 4911.9 included Eflprmidium
angustissimum, E. luridum, Navicula heuFleri var.
legtoceghaga, N. decussis, and N. ragigsa var. tgflglla.
During this same time period, the 25 cm. interval had been
subjected to late March disruption by ice action and
inundation by lake water on April 6 and May 6 collection
dates. Consequently, relative algae densities at the 25 cm.
interval in April and May oF 670.0 and 192.5 respectively
were the lowest along the transect. By June 6, the 25 cm.
interval, now emergent, had a relative density oF 1657.3,
which was larger than that oF either the 200 or 300 cm.
intervals. Important contributors to total algal numbers
at the 25 cm. interval during the April-June period in-
cluded tyngbya hieronymusii, Oscillatoria tenuis, Phormidium

angustissimum, B. luridum, Nicrocoleus chthonoplastes,

 

 

 

 

 

312

Navicula heuFleri var. le toce hale, and fl. radiosa var.

 

tenella.

As a generalization, the July-August collection
period was one of slowly increasing lake plankton density ~-
particularly by Hormogonales, Chlorococcales, and Pennales ~-
and Fluctuating algal abundance in the beach. In July 6
quantitative collections, a count of 26,194 individual
algae/cc. of sand gave the 25 cm. interval a relative
algal density of 5412.1 when compared with the numerical
density of phytoplankton within the waters of Reach A1.

By the August 6 collection, the Fact that the numbers of
algae had decreased more rapidly at the 25 cm. interval
than at the 100 cm. interval had resulted in the 100 cm.
interval having a greater algal relative density (see
Table 39). A list of algae of large numerical importance
in the 25 cm. interval during this period includes
Hicrocolegs chthonoplastes, fl. vaginatgg, Eggillatoria
splendigg, Phormidigm anqustissimum, and Navicula DEEfiEEE
var. leptoceohala. Important at the 360 cm. interval
during this period were Syflechococcus aeruginosus,

microcoleus acutissimus, Chlorococcum humicola, Botrydiopsis

 

arhiza. Navicula decussis, fl. heuFleri var. leEtEEEEflfilfl,

 

and fl. radiosa var. tenella. During the same period,
important contributors to the Flora at the 200 cm. interval
included éyflgphococcus aeruginosus, Anabaena sp. 2,
Microcoleus chthonoplastes, Nostoc paludosum, Phormigigm

angustissimum, Schizothrix freisii, Chlorococcum humicola,

 

 

 

Botrydiopsis arhiza. and Iavicula heuFleri var. legtoceahaln.
At the 100 cm. interVal, a list of important contributors
includes Microcoleus chthonoplastes, Uscillatoria splendiga,
Schizothrix Freisii. Symploca kieneri, and Navicula
heuFleri var. leptocephala.

In September and October, the lake underwent a
"bloom" of Dinobryon spp. that overshadowed simultaneous
increases in abundance by most other phytoplankton taxa
present in the lake at that time. In the September 5
collections, the count oF 119,010 individuals per liter
was the largest reported From lake quantitative samples
during the 12-month study. Concomitant with the phyto—
plankton bloom, algae abundance in the sand at the 300,
200, and 100 cm. intervals continued the decline begun in
August. At the 25 cm. interval, however, a pulse in algae
abundance negated the August decline.

Abundance oF algae at the 25 cm. interval in
September and October reached the greatest magnitudes
since January. At the same time, abundance of algae at
the 300 and 206 cm. intervals declined to or approximated
the lowest quantities reported For those intervals during
the 12-month study. Nevertheless, the relative densities
of both the 200 and 300 cm. intervals in September were
18.3 times that of the lake.

Quantitative samples at the 300 cm. interval in
September and October exhibited a resumption of cyanophyte

dominance which was absent in July and August. Important

 

 

 

314
contributors to the psammon algae at this interval in
September-October were Chroococcus minutqs, Phormidiufl

microtomum, Plectonema notatum, Chlorococcum humicola,

 

 

Navicula decussis, fl. heuFleri var. leptocephala, and fl.
radiosa var. tenella. In 200 cm. interval samples,

important species were §yfl§chococcus aeruginosus, Microcoleus
chthonoplastes, Phormidiqm anqustissimqm, Schizothrix

Freisii. Chlorococcgm humicola, Trochiscia sp. 2, and

 

flgvicula decussis. Oscillatoria aggrdhii, Q. tenuis,
Symgloca kieneri, and Navicula heuFleri var. leptocephala
dominated 100 cm. interval quantitative samples during
September and October. Primarily responsible For the
September-October pulse in algae abundance at the 25 cm.

interval were Microcoleus va inatus, Dscillatoria Formosa,

 

Phormidigm angustissimum, Navicula cryptocephala and fl.

heuFleri var. leptocephala. The September-October pulse

 

in abundance at the 25 cm. interval was attributable to

a doubling in the number oF blue-green algae and a seven-
to eightFold increase in the number oF diatoms, particu-
larly Navicula heuFleri var. leptocephala and Eavicula
cryptocephala.

In very brieF summary with respect to quantitative
seasonal variation, the lake phytoplankton From November
to January was dominated by the Hormogonales, From
February to May by members 0F the Chrysomonadales, and in
June and July by Chlorococcales and Hormogonales. By

August, dominance was shared by Hormogonales, Chlorococcales,

   

 

 

315
and Pennales. In September and October, the Chrysomonadales
were again daminant (see Fig. 26). At the 300 cm. interval,
the Hormogonales exhibited a clear dominance throughout
the year, except in May through &mtember when the
Heterococcales and Pennales established a transitory
dominance (see Fig. 28). At the 200 cm. interval, members
0F the Hormogonales maintained a clear quantitative
dominance throughout the year (see Fig. SO). In the sand
at the 100 cm. interval, however, members oF the Hormogonales
and Pennales shared dominance throughout the year, the
Filamentous blue-green algae being somewhat more abundant
From November to February and in August, the diatoms being
the most abundant March to July and September to October
(see Fig. 32). The situation at the 25 cm. interval was
almost the reverse oF that at the 100 cm. interval (see
Fig. 34). Although members 0F the Hormogonales and
Pennales clearly dominated 25 cm. interval quantitative
samples throughout the year, members oF the Pennales were
most abundant November to March, in June, and again in
September to October. Cyanophytes were most abundant in
April and May and in July and August.

Biological data, indices oF

diversity oF lake and beach
transect intervals

 

Odum (1959) states that in any natural community
a relatively small number oF species are usually repre-

sented by large numbers oF individuals, and the largest

 

 

316
number of species are represented by a small number of
individuals. The ratio between the number oF species and
the number of individuals is termed the diversity index.
He Further states that strong physicochemical limiting
Factors or interspeciFic competition reduces the diversity
within a community. Severe physical limiting Factors
consequently result in a low diversity index as does a
period oF high productivity Followed by competition For
space or nutrients.

Gleason in 1922 reported that a deFinite rela-
tionship apparently existed between the number oF species
oF arboraceous vascular plants present in a sample and
the logarithm of the area represented by the sample.
Preston (1960) reviewed at length the rationales For
various Formulas and techniques developed since 1922 For
community description based on species-area curves. The
same author in 1948 had reviewed various Formulae For
community description based upOn the phenomenon that in
apparently any population the relationship oF the number
OF species represented by one individual in a sample to
the number oF species represented by two in a sample, etc.,
when plotted reveals a logarithmic sequential series.

He concluded that this relationship is a constant For the
sample population whatever the sample size. Fisher,

Corbet, and Williams had reported in 1943 upon the rela-
tionship between the nUmber oF species and the number 0F

individuals in random samples oF insect populations.

 

 

 
 

 

317
They concluded that in random samples oF animal populations
the number oF species in successive abundance categories
show a logarithmic series as demonstrated in Fisher's
equation

N
3: 1n 1+—
CI

where S=number oF species and
N=number oF individuals in a sample.

They Found that“, termed index oF diversity oF
species, was more or less constant over a wide range oF
samples sizes and could be considered characteristic oF
a given community. The higher the value oFCI, the greater
the species diversity, i.e. relatively Fewer individuals
per species. The lower the valuetfi’Ofi the lower diversity,
i.e. more individuals per species.

A number oF other Formulae For calculation oF
species diversity have since been devised (Kohn, 1959;
MacArthur, 196D; fiacArthur and MacArthur, 1961; fiargeleF,
1959). The equations devised by MacArthur (1960) do not
seem applicable to a study oF the psammon environment
since they assume non-overlapping communities in equilibrium,
i.e. the total number oF individuals together are more
or less constant in that iF number oF individuals oF one
species increases, then the other must decrease. The
algae oF the psammon environment, however, seem to be
opportunistic, i.e. the abundance oF a given species
apparently is determined independently oF that oF any

other.

 

 

 

318
MargaleF (1957) considered possibilities oF
analyzing natural communities oF phytoplankton by use oF
the inFormation theory and reported satisFactory results
in identiFication oF species diversity through utiliza-

tion oF the equation

_ S-l
d-rrm

where d=index 0F diversity
S=number oF species and
N=number oF individuals in a sample.

Hairston (1959) in a review 0F techniques 0F
community identiFication applicable to studies oF soil
arthropods, although critical oF application 0F inForma-
tion theory to biological problems, conceeded MargaleF's
equation is adequate For crude but completely objective
separation oF similar communities such as in plankton
studies.

Patrick and Reimer (1966) reviewing the litera—
ture oF statistical studies 0F the relationship oF diatom
Flores to environment indicate that natural river com-
munities are characterized by a large number oF species
most oF which are represented by small numbers 0F
specimens. Any environmental condition which reduces the
number of species that can live in an area but does not
reduce the nutrient level results in a shiFt in composi-
tion oF the community. Organic pollution, they state,
tends to cause a reduction in the number 0F species and
induces very large numbers 0F a relatively Few species.

Thus, it seems, the value OFCZ may diminish. They Further

 

 

 

319
note that iF the environment remains nearly constant there
is a reduction in the number oF species which may occupy
an area. They conclude that a variable but somewhat
stable environment is necessary iF a high is to be main-
tained. MacArthur and MacArthur (1961) state that there
is no adequate theory of "species—area" or "diversity-
area" curve phenonema and no real evidence 0F the causes
cF high species diversity.

In view oF the apparent unresolved controversy
with respect to usage oF index oF diversity For sole
identiFication of communities, in this study index 0F
diversity has been considered only in an attempt to deter-
mine whether there is evidence oF physiCOChemical Factors
inFluencing the cemposition 0F the algae component oF the
psammonflora along the beach transect. The Following
equation was utilized in determination oF index 0F di—

versity For the lake and each beach transect interval:

__§_.
logeN

where S=number oF species and
N=number oF individuals in each sample.

The Value 5 rather than 3—1 was used because oF the rela-
tively large number 0F species present in each sample.

Table 41 presents the index 0F diversity For the
water oF Reach A1 and each beach transect interval menthly
during the 12—month study. On the basis oF the magnitudes
0F N and “For the lake and beach each interval each

month, standard error OFCI expressed as per cent 01’ the a

 

 
 

 

320

TABLE 41.--Index of diversity for lake and each beach

transect interval monthly, November to October

 

 

 

lake +25cm. +100cm. +200cm. +300cm.
Nov. 6.07790 6.54625 6.14604 5.92466 3.45609
Dec. 7.13126 6.22696 5.66160 5.70090 4.01656
Jan. 4.52505 5.92602 7.76666 6.70456 4.10733
Feb. 6.56791 7.71456 7.31117 7.50321 4.22255
mar. 6.67405 7.62445 6.59472 5.15666 4.03796
Apr. 9.62729 2.54297 6.00400 5.21405 5.60624
May 6.46465 7.04716 6.06699 5.67013 5.61520
June 9.36153 6.47070 5.95901 5.55061 3.55346
July 10.54579 6.76251 6.36020 5.06642 2.12606
Aug. 10.16519 7.06246 5.12654 4.06625 2.07703
Sep. 6.50502 5.91421 4.77760 5.51154 1.56060
Oct. 6.16555 4.94945 4.62269 3.11695 1.75271

 

 

value determined at maximum was less than 0.86 per cent

and in most cases was considerably less than 0.28 per cent.

The standard error was estimated From standard error

tables forCI determination presented by Fisher, Corbet,

and williams (1943).

contained in Table 41.

Figure 37 presents in graph form the information

In Figure 37 the index of diversity

menthly For the lake and each beach transect interval has

been plotted against time.

Reference to Table 41 and

Figure 37 indicates that -- except For the December~May

 

 

 

 

321
gov. Dec. Jan. Feb. mar.‘ Apr. May June Jxly Aug. Sep. Oct.
10. 1 I I I I I I I l I I

 

 
 
 
  
 
 
   

10.0-
lake

9.5- --—-—--25 cm.
—--«——--‘100 cm.

9.U_rOooooo~-OU.200 Cm.
x x x x xxxSDO cm.

index of diversity

*‘h'xxx v
0* *7. *x‘h ‘ I .c
4 , 0 .. ++X x x XXX +4," .

\v* +4' I "

1.

 

 

 

Fig. 37.--Index of diversity For lake and each beach
transect interval monthly, November to October.

 

 

 

 

 

 

322

period of collection —- the indices of diversity for the
five collection sites monthly exhibit a distinctly decreas-
ing gradation from the waters of Reach A1 to the 300 cm.
beach transect interval. In collections from May 6 to
and including October 10, each collection site maintained
a clearly defined characteristic index of diversity that
when plotted against time exhibited patterns of Fluctua-
tion essentially parallel to those of the other transect
intervals. The disruption by ice movements of the environ-
ment at the 25 cm. interval in April apparently induced
the pronounced displacement of the 25 cm. interval index
of diversity From its "expected" April position between
that of the algae of Reach A1 and the +100 cm. beach
transect interval. with that exception, only in January
through March was the pattern of progressive diminution
of index of diversity with distance landward not evidenced.

At no collection date did the index of diversity
at the +300 cm. interval exceed that of the +200 cm.
interval on the same date nor that of the +200 cm. interval
exceed that of the +100 cm. interval on the same date.
The index of diversity of the +100 cm. interval exceeded
that of the +25 cm. interval on the same collection date
only on January 4 and April 6.

Reference to Figure 37 suggests that the Further
the beach transect interval was distant From the water of
Reach A1, the lower the index of diversity and the less

seasonal variation of the index of diversity resembled

 

 

 

”a _. .

323
that in the lake. Figure 37 and the data presented in
Table 41 indicate also that the Further the transect
interval was distant From the lake, the less extreme
were the monthly Fluctuations in the diversity index.

On the basis oF the previously cited statements
by Odum, it would seem apparent that in the decreasing
index 0F diversity landward along the transect in any
given month there is evidenced an increasingly severe
environment and/or increasingly limiting complex 0F
physicochemical Factors.

The concept oF the ecotone as a transition area
where increased density and variety oF species results
From contribution oF organisms From both adjacent com-
munities as well as versatile species indigenous to this
environment has been reviewed at some length by Smith
(1966), Boating (1956) and Odum (1959). IF it is assumed
that the psammon zone is, indeed, a transition zone
between the juxtaposed aquatic and terrestrial environ-
ments as Sassuchin at al. (1927) proposed, then it would
seem logical to assume that somewhere along the beach
transect such a condition oF increased density and species
diversity would be encountered.

ReFerence to Table 39 indicates that in six
months (November through January, July, and September~
October) out 0F 12 the greatest abundance oF individual
algae per unit sample volume was at the 25 cm. interval.

In February through May the site oF most algae/unit

 

 

 

 

Mm”...

324

sample volume moved landward to the 200 cm. interval, and
in June and August it migrated to the 100 cm. interval.
Table 37, on the other hand, reveals that From April
through October, the greatest species diversity in the
Five collecting sites took place in the water oF Reach A1.
Only in November, December, and March did greatest number
oF species occur at the 25 cm. interval. In January it
took place at the 100 cm. interval and in February at the
200 cm. interval. Thus, only in November, December, and
February did greatest species number and largest number
of individuals occur simultaneously at the same collect-
ing site. In November and December this occurred at the
25 cm. interval and in February at the 200 cm. interval.
ReFerence to Figure 37, however, indicates that in
November, December, and February the greatest index oF
diversity oF the Five collection sites was in the water 0F
Reach A1. It should be recalled that only 11 OF 261
species oF algae reported during the 12-month study were
unique to the 25 cm. interval, and only Five were unique
to the 100 cm. interval.

It would seem that data presented in tables 37,
39; and 41 do not reveal the presence oF an acetone at
any collecting site along the beach transect nor do they
suggest that the entire beach From shoreline to the 300 cm.
interval may be considered transitional between two dis-
tinct communities, aquatic and terrestrial. The same data,
however, do indicate that the beach is by no means a

homogeneous environment as is more or less implied in the

 

 

 

325

concept oF the "psammolittoral" as a unique type oF aquatic
littoral habitat.
Egmflunity coeFFicients

Although index 0F diversity may be used to
characterize a given algal community, the index does not
lend itselF to comparison and identiFication oF communities.
Extensive reviews oF the literature with respect to quan-
titative techniques utilized in attempts at community
classiFication have been published by Bray and Curtis
(1957), Boating (1956), Graig-Smith (1964), Culberson
(1955), weiser (1960), King (1962), Curtis and McIntosh
(1951), Saunders (1960), Kohn (1959), and MacArthur (1964).
OF the various methods described, it would appear that
those based upon Kulczynski's coeFFicient oF community
are among the most Frequently used in studies oF plant
and/or animal communities. Such methods involve the
weighing oF the contribution 0F each species considered
by a measure cF abundance, i.e. by total numbers or by
percentage contribution to total numbers.

In essence Kulczynski’s coeFFicient is obtained

through use oF the equation

 

C= 2w x 100
a+b
where a=contribution oF species considered in
community 1,
b=contribution oF species considered in
community 2,
and w=contribution 0F species shared in
common in 1 and 2, w being the lesser
oF each paired value.

In this Form the index has a range 0F 0 to 100.

 

 

 

326

Bray and Curtis (1957), reFerring to (C) as
Gleason's coeFFicient oF community, state that it is the
best approximation yet available to a linear measure oF
community relationship. weiser (1960) inFers that the
coeFFicient is the best available technique that expresses
the degree oF uniFormity 0F populations From which samples
are derived. The value (C) has also been termed "index
0F aFFinity" (Saunders, 1960) and ”common abundance value"
(King, 1962; weiser, 1960).

Culberson (1955), Bray and Curtis (1957) and
Kidd (1964) have outlined the procedures involved in
utilization oF Kulczynski's coeFFicient oF community in
comparison 0F plant communities. Bray and Curtis have
discussed at some length the diFFiculties encountered in
using the standard correlation coeFFicient (r) in com-
parison oF plant communities. They indicate that since r
has in its solution a square transFormation, a Few very
abundant species cause two communities to show a high
correlation coeFFicient on the basis oF these Few species
which have a high count in both communities. Similarity
or dissimilarity oF lesser species is thus masked in
computation 0F r.

IF data involved in determination 0F Kulczynski's
coeFFicient are expressed as percentages, the coeFFicient
becomes simply the value oF w, i.e. the sum oF the lesser
value 0F percentage centribution For each species present

in both communities (Bray and Curtis, 1957). Since the

 

 

 

327

 

basic equation is stated 0: a Ewb x 100, when a + b:

200 and the value w is the sum oF the lesser percentage
value For each species shared in common in the two popu-
lations samples, then C=w.

Table 42 presents a summary 0F determinations oF
coeFFicient 0F community (C) For each site compared monthly
with the other Four sites along the transect. Figure 38
presents in graph Form the monthly coeFFicient oF com-
munity relationships oF Reach A1 and the Four beach tran-
sect intervals. On the basis 0F 12-month means oF coeF-
Ficient oF community and number 0F species in common in
Reach A1 and beach transect intervals as entered on
Table 42, a deFinite decrease in similarity oF community
composition oF the psammon algae is evident with increas-
ing distance FrOm the shoreline (see Table 42). This
decrease in similarity is evident in both number oF
species and relative abundance oF species shared in common.
The Fact that the 100 cm. interval means show greater
similarity to Reach A1 than do those oF the 25 cm.
interval is attributable largely to the disruption oF
the 25 cm. interval by ice action in March and the re-
sulting reduction in algae abundance at that interval
until July.

Figure 38 and Table 42 indicate, however, that the
decreasing similarity 0F the Reach A1 community to the
algal component oF the psammonFlora with increasing dis-

tanct Fr0m the shoreline was not as clearly evident on a

 

 

 

 

----IIIIIIIIIIlllllIIlllJIIIIIIfiHHIIIIIIIIIIIIIIIIIIT

 

 

 

 

 

 

 

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@345.» eves”. (A

329

Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.
I l l I 1 l

45 I ) l l

' lake vs. 25
40 ‘ /\ -—-——--1ake vs. 100
/ \ -—-'---lake vs. 200
35 _ . - "“‘-“'-lake vs. 300

coefficient of community
0 N m
o m o

..x
U1

.3
D

 

a.‘ O

 

 

 

Fig. 38.-—Community coeFFicients for Reach A1 and
Four beach transect intervals monthly, November to October.
month to month basis as it was in 12-month means. When
only the number of species shared monthly by Reach A1
and the beach transect intervals is considered, a pro-
gressive decrease in number of species shared From the
100 cm. to the 300 cm. interval is apparent all 12 months,
although in three months two intervals had the same
number of species in common. With respect to community
coefficient, the value (C) exhibited a progressive de-
crease From the 100 to 380 cm. interval in only seven

months. From November to January and in July, the

 

 

 

330
composition 0F the algal community 0F Reach A1 showed
a closer similarity to that oF the 208 cm. interval than
to other beach intervals. But in March the value For C
was lowest oF all beach intervals. These anomalies were
largely attributable to wide Fluctuations in abundance oF
Phormidium angustissimum throughout the year and along
the transect.

Figure 39 presents in graph Form the C relation-
shipscf the 25 cm. interval with Reach A1 and the other
beach transect intervals monthly November to October.
ReFerence to Table 42 reveals that on the basis 0F 12-
month means 0F both number oF species and relative
abundance oF species shared in common the algae community
at the 25 cm. interval was most similar to that 0F the
100 cm. interval and least similar to that at the 300 cm.
interval. when the 25 cm. interval algae community was
compared with that oF Reach A1, and the 108, 200, and
300 cm. intervals, the monthly mean number 0F species
shared in common was respectively 12.8, 22.0, 12.3, and
7.6. The monthly mean C values For the 25 cm. interval
in relationship to Reach A1 and the 180, 200, and 300 cm.
intervals were respectively 16.52, 48.85, 32.76, and
11.54.

Figure 39 indicates that the month by month C
relationships oF the 25 cm. interval to Reach A1 and the
other beach transect intervals showed considerable varia-

tion. with only an August exception, however, the C

 

 

 

 

331

60Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.
I | I | | I I 5 I I
lake vs. 25
-- ———1OO vs. 25
\——-——-——200 vs. 25,}
\ -------- o-SOO vs. 25/\ /
\ I

 

 

N (A LA 35 J-\ 01 U1
01 0 U1 C3 (II D U1
1
\
/
\
/
z

N
C3

coeFFicient of community (C)

 

_a
U1

.3
C3

 

 

 

O

 

 

. Fig. 39.--Community coeFFicients For the 25 cm.
interval in relationship to Reach A1 and other beach
transect intervals monthly, November to October.
relationship 0F the 25 cm. interVal to the 100 and ZOO cm.
intervals was clearly greater than that OF Reach A1 and
the BOO cm. interVal. For eight oF the 12 months the
algal Flora at the 25 cm. interval was most similar to

that oF the 100 cm. interval, and in nine oF the 12 months

 

 

 

332
it was least similar to that oF the 300 cm. interval. In
February, March, and August the lowest C relationships
oF the 25 cm. interval to other collection sites were
those oF the 25 cm. interval and Reach A1.

The month by month C relationships oF the 100 cm.
interval to Reach A1 and other beach transect intervals
are presented in Figure 40. Table 42 indicates that when
the 100 cm. interval algae community is compared with
those oF Reach A1 and the 25, 200, and 300 cm. intervals,
the monthly mean number oF species shared in common was
respectively 14.0, 22.0, 17.7, and 11.6. In 100 cm.
interval relationships to Reach A1 and the 25, 200, and
300 cm. intervals, the monthly mean values For C were
respectively 17.21, 46.85, 33.34, and 20.89.

ReFerence to Figure 40 reveals that -— as was
true with Reach A1 and with the 25 cm. interval ~-
month by month C relationships between the 100 cm. inter-
val and Reach A1 and the other beach transect intervals
were highly variable. It is evident, however, that C
relationships between the 100 cm. interval and the 200
and 25 cm. intervals were consistently higher than these
between the 100 cm. interval and Reach A1 and the 300 cm.
interval. In nine 0F 12 months the C value For the algae
at the 100 cm. interval indicated closest similarity to
the algae community at the 25 cm. interval. In December,
January, and August only was the 100 cm. interval C

value greater at the 200 cm. interval than at the 25 cm.

 

 

 

 

333

Nov. Dec. Jan. Feb. Mar. Apr.

 

may June July Aug. Sep. Oct.
A
55 .\ A /\ I \
\ /\ / \ / \
\ I 1 // \ I \
50 - \ \ I, \ ./ \ I \
\ \, / ‘ I I \\ A, \
\ / ,\.\/ ‘ .’ \l" \
€345 — \ . : \ I f - Y
V \// \ ‘ I '\ 7\’ ‘
> \'/ - \ / / . -!\
v 40 - , \ ‘ / ' \ I
é, \. i , ' \ /
g 35 - \ ‘ I / '
U . . ‘ - - /
. \ l / \ .
% 30 - '3' a \\' \ / ' /
1: ' \‘ / \' '
I)
3 25 '
...4
Q.
0..
320 ‘°
0 .
\. ‘
15 \\
10 - 100 vs.
————— 100 vs. 25
_____ 1.3!] vs. 200
5 _ .......... 100 V3. 300
0

 

    
 

 

 

Fig. 40.--Community coeFFicients For the 100 cm.

interval in relationship to Reach A and other beach
transect intervals monthly, November to October.

interval.

similarity between the algal community at the 100 cm.

interval was greater at the 300 cm. interval than in

Reach A1.

Both monthly means and Figure 40 suggest that

 

 

 

334
Figure 41 Presents in graph Form the monthly
coeFFicient oF community relationships between the ZOO cm.
interval and Reach A1 and the other beach transect in-
tervals. When the 200 cm. interval algal community was
compared with that oF Reach A1 and the 25, 100, and 300

cm. intervals, the monthly mean number oF species shared

Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.
I l I I I I |

 

50 -

45 -

20 '

..A
01
I

A
O

coeFFicient oF community (C)

 

ao-aacoIZOO V8. 300

 

 

 

Fig. 41.-—Community coeFFicients For the 200 cm.
interVal in relationship to Reach A and other beach
transect intervals monthly, November to October.

 

 

 

 

335

in common in quantitative samples was respectively 7.0,
12.3, 17.7, and 10.8. The monthly mean values For C For
the 200 cm. interval in relationship to Reach A1 and the
25, 100, and 300 cm. intervals were respectively 14.86,
32.76, 33.34, and 22.17. On the basis oF 1Z-month means,
thereFore, the algae community at the 200 cm. interval
apparently was most similar to that at the 100 cm. in-
terval and least similar to that in Reach A1.

ReFerence to Figure 41 indicates that month by
month C values For the algal cemmunity at the 200 cm.
interval in comparison to the communities in Reach A1
and the other beach intervals along the transect were
very variable. In only six 0F 12 months was the greatest
C value oF the 200 cm. interval between it and 100 cm.
interval. In Five months the 200 cm. interval C value
was greatest when compared with the 25 cm. interval. In
December, the coeFFicient 0F community was highest between
the 200 cm. interval and Reach A1. The C value in
December, however, was based on only seven species shared
in common between Reach A1 and the 200 cm. interval in
quantitative samples, and 28.2 oF the 42.6 C value was
contributed by the species Phormidium anqustissimgm.
Large numbers oF this taxon were responsible as well, it
would appear, For the anomalies in the April and may C
values where closest similarity was indicated DBtWBBH
the 200 and 25 cm. intervals rather than 200 and 100 cm.

intervals.

 

 

 

 

336

Month by month C relationships of the 300 cm.
interval algal community to communities in Reach A1 and
the 25, 100, and 200 cm. beach transect intervals are
presented in Figure 42. When the 300 cm. community was
compared with those 0F Reach A1 and the 25, 100, and 200
cm. intervals, the monthly mean numbers of species shared
in common were respectively 5.0, 7.6, 11.6, and 10.8.
In 300 cm. interval relationships to Reach A1 and the
25, 100, and 200 cm. intervals, however, the monthly

mean values For C were respectively 6.60, 11.54, 20.89,

Nov. Dec. Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct.

 

40
300 vs. lake
35 . ————— 300 vs. 25
. ——.——-— 300 vs. 100
j /\_ ......... 300 vs. 200
30 ‘

N
0'!

..3
U1

_\
CD

coeFFicient oF community (C)
N
O

U1

 

 

 

Fig. 4Z.--Community coeFFicients For the 300 cm.
interval in relationship to Reach A1 and the other beach
transect intervals monthly, November to October.

 

 

 

 

337
and 22.17. Thus, it would appear that on the basis oF
12-month means a clear dimunition oF similarity in com-
munity structure to that at the 300 cm. interval was
evident with closer approximation to the shoreline.
Figure 42 and Table 42 suggest, therefore, that the algal
community at the 300 cm. interval was most similar to
that of the 200 cm. interval and least like that in
Reach A1.

Although month by month C values For the 300 cm.
interval in relationship to the other collection sites
showed considerable variation, Figure 42 indicates that
throughout the year the algal community at the 300 cm.
interval was least similar to those in Reach A1 and at
the 25 cm. interval and most similar to those at the 200
and 100 cm. intervals. There was, however, considerable
monthly Fluctuation in which oF the two closest sites
had algae communities most like that oF the 300 cm.
interval. In only Four 0F 12 months was the C value be—
tween the 300 cm. and 200 cm. intervals higher than that
0F the 300 and 100 cm. intervals.

Table 43 presents a summary oF 12-month means
For value C and number oF species shared in common by
each collection site in relationship to the other Four.
As has been previously stated, the C value and number oF
species shared by the algal community oF Reach A1 and
those 0? the 25, 100, 200, and 300 cm. beach transect

intervals shows a gradual dimunition with increasing

 

 

 

 

338

TABLE 43.--Summary oF 12-month means For values oF C and
number oF species shared in common by each collection site
in relationship to the other Four

 

 

 

Reach A1 25 100 200 300
Reach A1 (12.8) (14.0) (7.0) (5.0)
16.52 17.21 14.88 8.80
25 (12.8) (22.0) (12.3) (7.8)
18.52 46.85 32.78 11.54
100 (14.0) (22.0) (17.7) (11.6)
17.21 46.85 33.34 20.89
200 (7.0) (12.3) (17.7) (10.8)
14.88 32.76 33.34 22.17
300 (5.0) (7.8) (11.6) (10.8)
8.80 11.54 20.89 22.17

 

 

Read table thusly:

The 12—month mean community coeFFicient between
Reach A1 and the 25 cm. beach transect interval was 16.52.
The mean number oF species shared in common by these two
collection sites was 12.8.
distance landward along the transect. The possible reason
For the low C and shared species value between the lake
and the 25 cm. interval has been discussed. flith respect
to values oF C and shared species oF the 25 cm. interval
and the other collection sites, it has been noted that the
25 cm. interval algal community exhibited the greatest
similarity to that oF the 100 cm. interval and the least
to the 300 cm. interval. Apparently, the algal com—
munity at the 25 cm. interval was more similar to that oF
Reach A1 than to that oF the 300 cm. interval. It has

}_ also been noted that the algal community at the 100 cm.

 

 

339
interval was most similar to that of the 25 cm. interval
and least like that of Reach A1. The 100, 200, and 300
cm. interval communities were most similar to the adjacent
interval lakeward and least similar to the community of
Reach A1.

Reference to Table 43 suggests, therefore, that
a lake algal community did exist which differed to some
extent from that in the beach. The previous discussion
pertinent to Table 35 has indicated that qualitative and
quantitative differences in the algae present in Reach A1
and the beach were marked and distinct, the differences
becoming increasingly pronounced with the distance separat-
ing a given beach transect interval and Reach A1. There
is evidence that the algal community at the 25 cm. in-
terval was somewhat transitional between the beach and
lake algal communities, but there is no indication that
the 25 cm. interval could be considered an ecotone. The
Fact that the community at each intermediate beach transect
interval -- with the exception oF the 25 cm. collecting
site -- resembled that at the 300 cm. interval more than
that in Reach A1 is Further evidence of a distinct beach
algal community.

The fact that the postulated beach algal community
was not homogenays throughout the limits of the transect
is evident From examination of Table 35 and the results of
indices of diversity determinations previously discussed.

A consideration of the Fact that each intermediate beach

 

 

 

340
transect interval showed in C and shared species Values
greater similarity to the interval lying lakeward rather
than to that lying shoreward further suggests that the
algal community at 300 cm. interval -- although obviously
qualitatively and quantitatively different from those
closer to the lake -- was primarily an attenuated expres-
sion of those lakeward rather than a distinct terrestrial
community. This apparent "survivors" community at the
300 cm. interval has been previously discussed in the
Results section relative to determination of indices of
diversity. Thus, data in tables 41, 42, and 43 can be
interpreted to conclude that an increasingly severe en-
vironment developing in a shoreward direction along the
beach transect induced a gradual attenuation of a distinct
beach algal community.

In attempts to identify the physicochemical
factors or factor responsible for the decrease in index
of diversity with increased landward distance along the
transect and responsible for the variation in community
coefficients, the amount of water present monthly ex-
pressed as percentage wet gross weight was plotted against
index of diversity at each interval samples each col—
lection data. Although a definite relationship between
these two variables was readily apparent, a better cor-
relatiOn of data was achieved when the index of diversity

for each interval each month was plotted against the log

 

 

 

 

341
of the monthly percentage water content on wet gross
weight basis at the same interval.

Figure 43 presents the results of these compari—
sons. Calculation for standard correlation coefficient
(r) for the two variables gave an r value of 0.83258,
well within 95% confidence limits. The slope of the re-
gression line of X on Y on Figure 43 indicating the rela-
tionship of monthly index of diversity with concomitant
percentage water content of the beach substrate on wet
gross weight basis was computed by use of the equation

1=bny + axy, using log values for Y.

X

According to variance interpretation tables for
correlation coefficients when r=O.83258, approximately
64 per cent of the variance in index of diversity during
this study could be attributed to variance in percentage
water content within the beach sand substrate. Unfor-
tunately, data from coefficient of community determina—
tions did not lend themselves to statistical treatment
insofar as their relationship to water content was con-
cerned.

when index of diversity for each interval each
month was plotted against the log of per cent saturation
of the sand at the same intervals on the same dates, a
definite correction quite similar to that expressed in
Figure 43 was evident. Determination of r values for
index of diversity in relationship to per cent saturation

was not made, however, inasmuch as per cent saturation

 

 

 

 

342

 

 

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a
. . was; 1 529?;
,

 

 

 

 

 

 

343
data were subject to determinations of water retention
capacity of sand from each interval made only once dur-
ing the 12-month study.

Values for r with reference to 12-month means of
indices of diversity for each beach transect interval
were computed for porosity, for 12-month means of per-
centage volatile matter, for sand grain size uniformity

coefficients, and for harmonic mean of grain size dia-

meters at each interval. For none of these variables did
I fall within 95 per cent confidence limits. The r
value for porosity in relationship to 12-month means of
diversity indices was computed to be 0.48375. with

reference to correlation between 12-month means of di-
versity indices and percentage volatile matter at each

interval, the computed r value was 0.57584.
Cumputed r value for harmonic mean sand grain
size diameters at sampled intervals in relationship to

12‘month means for indices of diversity at the same

intervals was -0.92330. For sand grain siZe uniformity

coefficients in relationship to diversity indices at the
same intervals, the computed r value was -0.90612. The
r values for uniformity coefficients and harmonic mean
grain size diameters in relationship to indices of
diversity both closely approached, but did not reach,
95 per cent confidence limits.

Consequently, it would appear that the percentage

water on wet gross weight basis at each interval was the

 

 

 

 

344-
primary factor in determination of qualitative and quanti-
tative algal abundance at the beach transect intervals
sampled. The relationship was direct. Of considerably
lesser importance was the direct relationship between

porosity and algal abundance.

 

An inverse relationship apparently existed to
considerable degree between algal abundance and sand
grain size and uniformity. Seemingly, the finer the sand,
the more productive the environment. It would also
appear that the more homogeralm the sand grains in dia-
meter and the more uniform the general dimension of inter-
stitial spaces between sand grains, the more productive

was the beach environment.

 

 

 

 

CHAPTER VI
DISCUSSION
Terminology relative to the

psammolittoral

Reference has been made in Chapter I to the extant

 

controversy relative to the terminology applicable to the
sandy beach biotope. Sassuchin gt al. (1927) apparently
coined the term "psammon” for the microbiota of the sandy
beach, considering it analagous to the term "edaphon"
which had been applied by France (1913) to the microbiota

of the soil. Their use of the terms "nannopsammon" and

 

”micropsammcn" suggests their concept of the term psammon
as referring to biota of the sandy beach rather than to
the environment itself. References to the 1927 paper by
Sassuchin et al. in reviews of the literature concerned
with studies of the beach environment allude that the
paper employs the term "psammon” for both the biota and
the environment. In the 1927 paper, however, the authors
did not employ the term for other than biota.

Although Sassuchin gt a}. noted horizontal zona-
tion in beaches with reference to water content and
characteristic microflora and microfauna and wrote of
“zones”, they did not clearly delineate or define these

zones. Rather, they dwelt at some length upon vertical

 

 

 

346
"horizons", characterizing these horizons on the bases of
color, depth, and biota and noted the manner in which the
horizons were expressed in two zones. The two zones were
briefly described as one moist and adjacent to the lake
and the second drier and further removed from the lake.
wiszniewski (1934a) in reviewing studies of the

beach environment attributed initial use of the term

 

psammon to Sassuchin. Niszniewski (1934b) had previously
proposed the term "psammolittoral” in a 1932 paper to
represent the concept of the beach as a unique type of
littoral environment. In his Recherces ecoloqiques so;
is psammon (1934a) fiiszniewski restated his 1932 clas-
sification of the micropsammon —- in the sense of psammon

being an assemblage of aquatic microorganisms rather

 

than an environment -— as hydropsammon, hygropsammon,

and eupsammon. The first group includes those organisms
living in constantly submerged sand. The second group

he defined as those organisms found at the surface of the
exposed portion of the beach within one meter of open
water. The last group —- the eupsammon -- he defined as
those organisms living in the extreme portion of the beach
more than one meter from open water.

In the initial portion of the summary of Recherghgs
ecologigues egg lg psammon miszniewski again stated that
the terms hydro-, hygro-, and eupsammon are subdivisions
of the psammon biocenosis rather than of the psammolittoral

biotope. In a subsequent portion of the summary, however,

 

 

   

347
he refers to the "zone of the hydropsammon", "zone of the
hygropsammon", and "zone of the eupsammon". It would
appear that subsequent investigators of the psammolittoral
environment have incorrectly equated "hygropsammon" with
"zone of the hygropsammon", etc., thus leading to un-
necessary confusion in the terminology involved in study
of this unique environment.

It would seem, therefore, that confusion could be
reduced through utilization of the term psammon to
identify the biota living in the psammolittoral zone of
a beach. It would appear, also, that the terms hydro-
psammon, hygropsammon, and eupsammon should refer to the
organisms living in the hydropsammon zone, the hygro—
psammon zone, and the eupsammon zone respectively. In the
connotation apparently intended by wiszniewski, a psammo-
littoral zone exists wherever a body of water having a
sandy substrate possesses a more or less extensive sandy
exposed shore..

A review of the literature indicates that apparent—
ly there has been no definition of the word ”sandy" as
used in relationship to the psammolittoral. Spangler
(1960) has noted that terms used in the United States to
indicate texture of soil are highly arbitrary in origin,
and their connotations vary from place to place and from
One discipline to another. Beanland (1940) states that

variety of methods of soil analysis and lack of uniformity

H
H
T
T

 

 

 
 

 

348
in describing soil grades makes present data upon estuarine
soils of little value for comparative purposes.

One approach to the problem of classifying soils
is use of triangular classification charts such as those
presented by Spengler (1960) and Hough (1957). It would
appear that soil classification charts developed by

organizations such as the U.S. Corps of Engineers, U.S.

- Bureau of Public Roads, and the U.S. Department of

Agriculture all share the characteristic that any "soil"
having more than 80 per cent dry weight contributed by
particles within the size limits of 2.0 to 0.05 mm. is
classified as sand. Reference to grade distribution of
sands of psammolittoral environments investigated by Neel
(1949), Pennak (1950, 1940), Ruttner-Kolisko (1956, 1953,
1954), Niszniewski (1934a, 1934b), and that of the Reach
A1 beach transect indicates that all have had grade dis-
tributions with more than 80 per cent contribution by
particles lying in the range of 2.0 to 0.05 mm. diameters.

The hydropsammon zone of the psammolittoral
environment has been defined by wiszniewski (1934a) as
that portion of the beach continually submerged. He de—
fined the hygropsammon zone as that portion of the exposed
beach in which the interstitial water is in a capillary
state, i.e. the interstitial pores are completely filled
with water, and the beach surface is continually moist.
The eupsammon zone Uiszniewski characterized as being

inundated only by the highest waves. The surface is said

 

 

 

 

jig-91am...“ ,

349
to be in a funicular state, i.e. with air spaces within
the interstices between sand grains, and the surface
appears dry. when the surface film of sand reaches a
condition in which only a thin film of water covers the
sand grains, according to wiszniewski that portion of the

eupsammon zone is said to be in a pendular state. In

 

essence, therefore, the original definitions of the three
zones of the psammolittoral environment are based on
water content of the substrate.

Pennak (1940), Neel (1948) and welsh (1952)
equate the hygropsammon and eupsammon zones respectively
with inner and middle beach. A search of the literature
does not reveal any definition pertinent to delimitation
of the upper limit of the eupsammon zone. Unfortunately,
few investigators of the psammolittoral zone have in-
dicated in their reports the distances from the water
line at which samples have been taken. Sassuchin g3 gl.
(1927) included as psammon organisms those collected 50.0
meters from the shore, but Wiszniewski (1934a) presents
no data derived from samples taken more than eight meters
from the water line. Neel (1948) seemingly considered
the upper limit of the psammolittoral as two to six
meters from the water line. Pennak (1950, 1939, 1940)
apparently considered a distance of three to four meters
the upper limit of the psammolittoral zone. Ruttner-
Kolisko (1954, 1953, 1956) reports no eupsammon zone ‘

data more than 80 cm. distant from the water line.

 

 

 

 

 

350
Consequently, it can be assumed that the term eupsammon
may include the "outer beach" (Welch, 1952) as well as
the middle beach.

It was noted in Chapter I that review of the
literature indicates confusion regarding delimitation
between the hygro- and eupsammon zones, and it is evident
differentiation between the two is highly arbitrary. If
wiSZniewski's criteria for identification of the three
zones of the psammolittoral are utilized, reference to
Table 8, Chapter V, indicates that the +25 cm. interval
of the Reach A1 beach transect was from November through
March located in the hygropsammon zone. In April and
May the +25 cm. interval was located in the hydropsammon
zone. From the June 6 collection date to the October
termination of the study, the +25 cm. interval was located
in the eupsammon zone. with the exception of the +100
cm. interval in April and May, all other transect intervals
shoreward from the 25 cm. interval were located in the
eupsammon zone. In April and May the advancing lake water-
line positioned the +100 cm. interval in a transitory
hygropsammon zone.

Reference to tables 7 and 8 does not reveal,
either in monthly data or 12—month means, evidence of
obvious demarcation of zenes along the beach transect.
Neither data of percentage water content on wet weight
basis nor data of percentage saturation show a sharp

break in water content along the transect that would

 

 

 

 

 

351
suggest an abrupt shift from the hygropsammon to eupsammon
zone. Neither is there evidence of a line marking the
limit of capillary rise. Rather, in each month, there
is evidenced a more or less gradual ~- albeit pronounced ——
diminution of water content with distance from water line.
Unfortunately, there apparently is in the literature no
data of water content taken at relatively close intervals
during a prolonged period which can be compared with the
results of the Reach A1 transect study.

It is evident that investigators of the psammo—
littoral have made the assumption that the interstitial
water of the beach sand is derived from wave action,
tides, capillarity, and -— to a much lesser extent -—
precipitation. Bruce (1929), Newcombe (1935), Pennak
(1939, 1940, 1950), Neal (1948), Hill and Hanley (1914),
Ruttner-Kolisko (1956, 1953, 1954, 1962) and uiszniewski
(1934a), among others, have discussed at some length the
rise of water in beach sand by capillarity. In essence,
they have found that the finer the sand substrate, the
greater the capillary rise. The literature presents
instances of observed capillary rise in beach sand of
magnitudes of as much as 00 cm. (Pennak, 1940). Ruttner—
Kolisko (1956) has stated that the width of the hygro—
psammon zone is determined by sand grain size and beach
slope. The finer the grain size and the smaller the de—
gree of slope, the wider is the zone of complete satura-

tion. The data relative to field and laboratory

 

 

 

 

 

 

 

investigations of sand grain size and capillary rise
presented in reports of studies of the psammolittoral do
not indicate, however, that the distance water will rise
by capillarity can be predicted in any exactitude by
reference to grain size of the sand substrate.

Although there are in the literature numerous
descriptions of experiments and observations regarding
height of capillary rise in beach sands, it would appear
that few (Sassuchin gt. gl., 1927; Hill and Henley, 1914)
have investigated the contour of the phreatic level below
the beach surface. The phreatic surface is defined by
Todd (1959) as the upper surface of the zone of satura-
tion, i.e. that area of the substrate in which the inter-
stices are completely filled with water. The level of the
phreatic surface is revealed by the level at which water
stands in a bore penetrating the substrate. Actually,
according to Todd, the zone of saturation extends slight-
ly above the phreatic surface because of capillary at-
traction; however, water is retained there at less than
atmoapheric pressure. Above the phreatic surface, in the
zone of aeration, interstitial water may be found as
hygroscopic water, or as capillary water in a more or
less continuous film held by surface tension around sur-
face particles and moved by capillary action, or as
gravitational water which, derived from precipitation,
drains through soil to the zone of saturation. Hater in

the zone of aeration is termed vadose water. Forces of

 

 

 

 

 

353
surface tension and molecular adsorption do not permit
complete removal of interstitial water by drainage or
pumping in either the zones of saturation or aeration.

It appears that previous investigators of the
psammolittoral have assumed vadose water is derived
primarily by capillarity from the zone of saturation
below the beach. In the report by Hill and Henley (1914)
and in the Duck Lake Reach A1 beach transect study, it
has been noted that the phreatic surface declines with
shoreward distance from the water line. The question
thus arises whether vadose water at any given point on
the beach surface is that rising by capillarity from the
phreatic surface below.

Reference to Figure 3 indicates that during the
period November 9 to and including October 10 a vertical
distance of as much as 48 cm. may have separated the
phreatic surface and the surface sand at the +25 cm.
interval. This distance lies within observed values of
capillary rise reported in the literature for sands hav~
ing D10 of .070 cm. and/or mean grain diameters of .079
mm. In July, August, and September, however, the vertical
distances between the phreatic surface and the surface
sands of the +100, +200, and +300 cm. beach transect
intervals were, respectively, approximately 70, 80, and
105 cm. From November through June, it is assumed these
vertical distances were somewhat less. Table 5 indicates

that the 010 and mean grain diameter for surface sand at

 

 

 

 

 

 

354
the +100 cm. interval were respectively .090 and .110 mm.
The D13 and mean grain diameters at the +200 cm. respec-
tively were .104 and .119 mm. and at the +300 cm. were
respectively .114 and .148 mm. Even assuming the sands
at each interval were homogenele in texture from surface
to phreatic surface, which they obviously were not, the
vertical distances between surface sand and July —
September phreatic surface levels were considerably
greater than maximum capillary rise reported in the
literature for sands of the above mentioned D10 and
average grain diameters. Yet in the July — September
collections, percent saturation values at the 100 cm.
interval ranged from 58.5 to 23.6 per cent. At the

200 cm. interval, July - September per cent saturation

values ranged from 21.3 to 4.3 per cent. In those months,

the zone of saturation lying lakeward was at least 125 cm.

distant from the 100 cm. interval.

An extensive literature exists with reference to
movement of water through porous soils, including among
otheesreports by Mavis and Tsui (1939), Richards (1931),
Smith gt al. (1931), Moore (1939), Luthin and Day (1955),
Remson and Fox (1955), Lambe (1951), and Kirkham and
Feng (1949). Kirkham and Feng in summation of a review
of the literature of studies of capillary water in soil
state that, in brief, all is not known with reference to
capillary movement in soil. Lambe (1951) has stated that

numerous incorrect concepts appear in the literature

 

 

 

 

355
because of incomplete fundamental understanding of the
role of capillarity in movement and retention of soil
water and concludes that there is a great need for much
more research upon capillarity. He has found that soils
in which 90 per cent of the particles are larger than
0.08 mm. in diameter (thus being sand as defined in the
triangular classification charts previously cited) have
a maximum capillary saturation value of only 80 per cent
of void space rather than the 100 per cent previously
presumed. If this is generally true, it would appear
all data pertaining to determination of soil porosity
by standard water saturation techniques are highly sus-
pect.

Luthin and Day (1955) and mavis and Tsui (1939)
report that substantial lateral water flow takes place
in soil above a sloping water table. Mavis and Tsui
report the horizontal movement of water in the capillary
fringe above the phreatic surface varies with a well-
defined regularity from zero at the top of the fringe to
a maximum at the phreatic surface. Luthin and Day note
that no existing theories consider variation in capil-
lary conductivity in the zones of saturation and aeration,
and they further note that no quantitative measurements
had been made —— in 1955 -- which included the region
above the phreatic surface. 0reshkina (1960) states
that capillary water discontinuity occurs when substrate

moisture content becomes less than four per cent. with

 

 
 

 

356
lower moisture content, free moisture exists only in
"wedges" regardless of the state of compaction or size
of the sand particles.

Another factor which apparently has not been con-
sidered by investigators of the psammolittoral zone in
their generalizations concerning capillarity in beach
sands is the relationship of temperature to capillary
rise. mavis and Tsui (1939) state that the magnitude
of capillary rise in beach sand depends not only on the
size of the grains but also on the ”porosity factor” of
the substrate and surface tension of the water (see
Mavis and Tsui for a discussion of porosity factor).
Surface tension, they state, is a function of water
temperature. Moore (1939) has noted that in unsaturated
soils, rapid changes in soil temperature above a water
table are accompanied by rising phreatic levels with
temperature increase and falling levels with falling
temperatures.

Lambe (1951) has noted that the relative humidity
of soil air is 100 per cent. Consequently, any drop in
temperature results in condensation of water vapor upon
sand grains. Inasmuch as evaporation in unsaturated soils
takes place directly into interstitial spaces (Greshkina,
1960), it seems possible there may be appreciable water
deposition in superficial layers of the beach following
temperature decreases. Hill and Hanley (1914) found

appreciable increases in water content of beach sands

 

 

 

357
during night hours. This they attributed to dew forming
on the sand surface when moist air over the beach surface
cooled.

The present incomplete knowledge of fundamental
processes of capillary movement in beach sands and the
apparent lack of information regarding the importance of
condensation of water within interstices between sand
grains suggest that efforts (Ruttner-Kolisko, 1956) to
predict the extent of the hygropsammon and eupsammon
zones 0n the basis of beach slope and grain size distribu-
tions alone are probably premature. In short, it would
seem that at the present time it is not known whether
the water found at any given point in the psammolittoral
zone is capillary water from the underlying phreatic
surface, capillary water transported by lateral capil-
larity from the adjacent basin of water, water of conden-
sation from the atmosphere or interstitial air voids, or
gravitational water from recent precipitation. Thus, it
seems unwise to refer to the water within the interstices
of a sandy beach as capillary water. Rather, it might be
suggested that water found in the beach transect be re-

ferred to only as "interstitial” water.

Porosity and grade
Bruce (1928) has noted that the interstitial
spaces within a sandy beach, whether occupied by water

or air, are not only important as the habitat within

 

  

 

 

358
which the psammon organisms must live but also their
relative volumes determine the specific heat of the beach
and the availability of capillary water. He reports that
.the specific heat of damp sand varies from one-tenth to
one—third that of pure water, depending upon the relative
volumes occupied by sand, air, and water.

The diameter of sand grains in the beach substrate
determines the size of the interstitial voids in which
water may be found. The smaller the grain size, the less
the available space. According to mavis and Tsui (1939),
the absolute size of pores is small in fine sand and
large in coarse sand, but both materials have the same
porosity if they are geometrically similar. The absolute
volume of pore space between adjacent sand grains varies
with the cube of grain siZe, and the cross sectional
area of this interstitial space varies with the square of
the grain size.

Pennak (1939) states that the proportion of pore
space to total sand volume is approximately 40 per cent
regardless of the size of the sand grains and irrespec—
tive of the degree of heterogeneity of the particles. In
spite of varied composition of beach sands, Pennak (1940,
1950) further states, the interstitial volume of beach
sands is seldom less than 36 per cent or more than 43 per
cent. Hough (1957) in summarization of numerous sand
porosity determinations reported in the literature con~

eludes porosities in sands may range from 44 to 17 per

 

 

 

 

 

3S9
cent, and Baver (1940) states that soil porosities average
50 per cent, sand being usually less and clays usually
more. Hough indicates that sands having uniformity coef-
ficients approximating 1.0 and effective grain size (010)
near 1.0 mm. have porosities of 44 to 33 per cent. Those
with D10 approximating 0.10 and having uniformity coef-
ficients of four to six, on the other hand, may have
porosities ranging from 49 to 17 per cent. Todd (1959)
gives porosity values for mixed sands ranging from 30 to
40 per cent, coarse sands approximating 40 per cent and
fine sands 30 per cent.

Bruce (1928) found porosities of mixed beach sands
approximating 20 per cent, but mavis and Tsui (1939) found
that porosity determinations made upon sands of the Iowa
River ranged from 43 to 34 per cent. Sassuchin gt al.
(1927) reported average porosities of about 37 per cent
in beach sands. diszniewski (1934a) found porosity values
in beach sands ranging from 32 to 42 per cent. Heel
(1948) discovered porosity values for ungraded sands
varied from 37 to 41 per cent. Thus, the literature in-
dicates porosities in beach sands range from 49 to 17
per cent, most determinations having ranges from 44 to 30
per cent.

Porosity values determined for each 25 cm. inter-

val along the Reach A1 beach transect tended to be higher

than most values reported in the literature for beach

sands. Values along the beach transect above the November-

 

 

 

 

360
March water line ranged from 50.8 per cent at the water
line to 41.3 per cent at the +300 cm. interval. The mean
value for porosity determinations at the 17 intervals
along the transect was 46.8 per cent. Reference to Table
5 (see also pages 49 through 56) indicates that, as a
generalization, the larger the harmonic mean grain dia-
meter in mm. and the larger the uniformity coefficient,
the lesser was the porosity.

Unfortunately, the absence of standard methods of
porosity determinations reported in the literature of
psammolittoral studies and the fact that grade data re-
ported from this environment have not included mean dia-
meters, uniformity coefficients, or effective grain sizes
precludes any meaningful comparisons of the porosity
results of this study with those of previous investigators
of the psammolittoral zone. Ruttner-Kolisko (1956) has
previously pointed out the necessity for an improvement in
the techniques of study of this environment. It can only
be hoped that future investigators of the psammolittoral
will employ sound standard procedures for physical analysis
of the substrate.

Pennak (1950) has stated that size of sand grains,
pH, dissolved free and bound 002, and dissolved organic
and inorganic matter have no known constant relationship
to either numbers or distribution of psammon organisms.
Ruttner-Kolisko (1956), however, reports that the appreci~

able development of a psammon microfauna occurs only in

 

 

 

 

 

2
g.

361
pore systems based on grain sizes 2.0 to 0.2 mm. in dia—
meter. She argues that above this limit capillary forces
are insufficient to produce capillary rise, and below
this limit the pores are too small for movement of the
organisms. She states that a "good" habitat is one with
about 40 per cent of the sand grains having diameters of
about 0.5 mm., and a ”poor” habitat is one in which ap-
proximately 50 per cent of the sample has diameters below
0.2 mm. She concluded that for the "biota” of the psammo—
littoral the grain size distribution most favorable is
one in which 50 per cent of the sand grains have dia-
meters of about 0.4 mm. and the uniformity is such that
few grains have diameters of more than 1.0 or less than
0.2 mm.

Although few, if any, references to relationships
between algal communities and porosity of the psammo—
littoral environment appear in the literature, there are
numerous published reports of apparent relationships of
porosity and distribution of invertebrates in or upon
beach sands (Kohn, 1959; Beanland, 1940; Boaden, 1963a,

1963b; Nicholle, 1935; Pirrie at al., 1932; Vacelet, 1961;

Teiser, 1960, 1959; Ruttner-Kolisko, 1961). Boaden (loc.

git.) reported that the diameter of sand grains markedly
influenced interstitial nematode fauna. Certain species,
he states, showed obvious preference to substrates of
given sand diameter ranges, some being typical of fine

send, some of coarse sand. (ing (1962) found evidence

 

 

 

 

 

«~a_e

362

that nematode communities in marine beaches are delimited
in part by differences in average grain sizes. weiser
(1959, 1960) discovered that the distributions of some
species of nematodes and kinorhynchs apparently were deter-
mined by composition of the substrate, i.e. some prefer-
ring coarse sand, some silt, etc. Nematodes, he reported,
are very sensitive to small changes in composition of the
substrate, the size of the nematode in respect to the
size of the substrate particles apparently being a con-
tributary factor. In Puget Sound, Heiser found distribu~
tion of some invertebrates is not determined as much by
tide levels as by the pattern of distribution of certain
grades of the substrate sand. Certain mixtures of sands
constitute barriers separating major faunal components
from each other.

It has been previously noted that results of the
Reach A1 beach transect study indicate an r value of
-0.92330 between harmonic mean sand grain size diameters
at sampled intervals and 12—month means for indices of
diversity at the same intervals. This would suggest that —-
within the limits of the grain sizes present in the Reach
A1 study -- there is an inverse relationship between
grain size diameter and index of diversity, i.e. the smal-
ler the grain size, the larger the index of diversity and
thus, the more favorable the environment. It should be
noted that all 17 intervals along the Reach A1 beach

tranSect analyzed for grain size distribution had grain

 

 

 

363

 

size percentage compositions such that they fell into the
size category characterized by Ruttner-Kolisko (1956) as
"poor" habitats (see tables 4 and 5 and Figure 3). It
would seem that Ruttner-Kolisko's generalizations regard-
ing grain size and productivity are perhaps not applicable
to the algal community. It should also be noted, however,
that not only did harmonic mean grain size become smaller
the closer the sample interval to the water line (Table 5),
but in addition the amount of interstitial water also
increased the closer the sample interval to the water line
(Table 7).

It has also been previously noted that the com—
puted r value was -0.90612 for sand grain size uniformity
coefficients at each sample interval in relationship to
12-month mean diversity indices at the same intervals.
This indicates an inverse relationship between uniformity
coefficients of sand grains in the Reach A1 beach transect
and indices of diversity, i.e. the more homogeneous the
sand grains in diameter, the more favorable the environ-
ment. Again, however, it should be noted that reference
to Table 5 and Table 7 reveals that the closer to water
line the sample interval, the more homogeneous the sand
grain diameters and the greater the average water content.
It becomes obvious that laboratory experimentation with
controlled water content is needed to determine whether
there is, indeed, a relationship between sand grain size

and/or uniformity on one hand and productivity of the

E:.............._____________________ ii

 

 

 

364
psammolittoral with reference to the algal biota on the

other.

Temperature

Pennak (1950) in summarizing the ecology of the
psammolittoral zone has noted that temperature within the
interstitial voids is influenced by temperatures of the
adjacent water basin, air temperatures, amount of "capil-
lary" water and rate of its evaporation, relative humidity
above the beach, and insolation. He states that in fresh-
water beaches the surface temperatures of the sand between
50 and 250 cm. from the water's edge are relatively
homogeneous because of the predominating influence of
evaporation of the interstitial water. In this zone, he

continues, even on the warmest of days the surface tempera-

 

ture of the sand seldom exceeds 32°C. Nearer to the water
line, however, the sand temperature reflects that of the
adjacent body of water, and at the shoremost, driest
extremes of the psammolittoral zone insolation may raise
sand temperature to more than 40°C.

Pennak (1939) has found that during a 24—hour
period temperatures at the moist surface of sandy beaches
may vary as much as 100C. or more, the surface layers
exhibiting greater temperature fludwations than deeper
layers. Newcombe (1935) reports that temperatures 8 cm.
or more below the sand surface are relatively stable.

Johnson (1961) indicates that daily temperature variation

figa’iif; Mfr-T

in intertidal marine sand flats resembles that observed
in terrestrial soils. Animals living in the upper 1 cm.
of sand, he reports are exposed to daily temperature
ranges three times that of the subtidal environment. But
even under maximum natural insolation, small amounts of
water may be sufficient to keep sand temperatures low
enough to be tolerated by most psammon organisms (see
Pennak, 1940, and diszniewski, 1934a for some of the most
extensive temperature data reported from studies of the
psammolittoral).

Although wiszniewski (122. git.) with respect to
temperature characterizes the psammolittoral as having
high temperatures derived from insolation tempered by
evaporation as well as having large amplitudes of daily
temperature oscillation, it is apparently in comparative
reference to the aquatic environment that he writes. As
is true in almost all studies of the psammolittoral,
Uiszniewski's temperature data are derived from investi-
gations conducted in the summer months. In essence, his
tables and graphs show for any given transect a rather
rapid temperature increase in the first 25 cm. from water
line followed by more or less stable temperatures over
the remainder of the transect. Rarely do his tables
exhibit temperature data in excess of 330". He found,
however, no clear generalizations with regard to tempera~

ture changes with distance from lakes except that such

 

 

 

 

 

   

54ft" , ‘

366

fluctuations tend to be less rapid and less marked the

nearer the sample site to the lake.

Bruce (1928) has also noted that evaporation from

the moist beach surface usually prevents any excessive

rise in temperature.

He reported the highest tem erature

recorded in English intertidal beach sands during summer

months do not exceed 210C. in full sunshine, although

readings in dry, wind-blown sand were much higher. Pearse

23 al. (1942) have found summer air temperatures immedi~

ately above sandy marine beaches to be lower than water

temperatures in the adjacent sea.

diszniewski (1934a)

reports that the relative humidity one cm. above the

freshwater beach reaches a

maximum in the first 50 co.

from the water line, then drops, and remains relatively

stable shoreward from the

50 cm. interval.

Bruce (loc.

Cit.) reports that experimentation upon the rate of evapora-

tion from sands of various
alone has little influence
tion.
surface, he states, is not
of an equal area of water.
sand apparently contribute

loss, the actual escape of

by rate of diffusion through narrow interstices.

(1955), however, has found
from fine sand exceeds the

sand.

 

grades indicates that .rade

L0

in determining rate of evapora-

Initial rate of water loss from a saturated sandy

appreciably different from that
The internal surfaces of damp
little to total evaporation
water molecules being limited
Newcombe
that the rate of evaporation

rate of evaporation from coars

 

 

 

 

 

 

Table 6 presents the record of temperature deter—
minations made during the 12-month investigation of Reach
A1 and the psammolittoral zone transect (see also pages
57 through 59 for additional temperature information).
Reference to Table 6 reveals the strong tendency for tem—
peratures at the most lakeward stations to approximate
within three degrees C. those of the lake surface on the
same dates and for those most landward to approximate
within three degrees C. breast height air temperature.

At no time during the study did sand temperature exceed
35.00C. This reading was, however, eight degrees higher
than the breast height air temperature of 27.0 degrees
recorded on the same date, September 5. The spring
through fall temperature ranges along the transect did
not differ markedly from those reported in the literature
during comparable monthly periods.

Sand temperature differential between the most
lakeward and the most landward stations on the transect
ranged from zero in January, February, and March to maxima
Of 6.0 and 5.50C. in April and September respectively.
Temperature differentials between extremes of the transect
tended to be lowest during winter months and greatest in
spring and fall. Only in July through September did
temperature at the most landward point of the transect
differ markedly from air temperature, it being in those
months about 70C. higher than breast height air tempera~

ture. During the December—March collection period when

 

 

 

 

 

368

the substrate was more or less continually frozen, tempera-
ture differentials between extremes of the transect did
not exceed 0.50C. On January 4 and February 5 collection
dates, all temperatures recorded along the transect were
0.005. It is not known what temperature minima occurred
in the beach sand during the December-March period, but it
is obvious that daily sand temperature fluctuations during
this period were greater than temperature fluctuations in
the adjacent ice-covered water of Reach A1.

Lund (1942) and Ruttner-Kolisko (1956) have re—
ported that temperature alone apparently exerts no impor-
tant influence upon growth and reproduction of the biota

of the sand. Aleem (1950) found temperature apparently

 

has no effect upon diatom populations in mudflats, and
Pearsall (1923) has concluded that it is improbable that
temperature alone is of any great importance in determin-
ing periodicity of diatom populations. Round (1968, 1961),
however, has concluded that gross productivity of both
diatom and blue-green algal populations is correlated

with temperature in epilithic, benthic environments.
Sakharova (1963) reports that regular patterns of distri—
bution and seasonal dynamics have been established for
microfauna of the psammolittoral of Russian lakes, assumed-
ly with respect to temperature. Shields and Durrell
(1964), however, with reference to soil algae state that
because of tolerance of soil algae to extremes of heat

and cold, temperature loses significance as a factor in

 

 

 

 

 

369
determining growth and distribution of soil algae. Cameron
(1964) has found that soil algae may grow and reproduce
readily in cultures maintained at 90 to 100°F.

Table 39 and Figure 35 present summaries of quanti-
tative data relative to fluctuation in algal abundance in
the lake and in beach transect intervals during the 12-month
study of the Reach A1 beach (see also pages 301-315 for
summarization of data relative to seasonal fluctuations
in algal numbers). Figures 35 and 25 indicate that phyto-
plankton density within the waters of Reach A1 underwent
a gradual and pronounced decline from November to March.
Subsequent to the March-April ice break-up, density of
phytoplankton began a gradual increase to a 12—month peak

in September, followed by an abrupt decline to the October

 

termination of the study (see also pages 127 through 149
for further reference to quantitative variation in phyto-
plankton abundance during the 12-month study). Similar
patterns of seasonal phytoplankton quantitative variations
have been reported by Chandler (1940, 1942, 1944) Spencer
(1950), Davis (19543, 1954b), Griffith (1955), and Reed
and Olive (1956).

It has been noted (page 303) that all beach inter-
vals exhibited maxima in algal density during the four—
month period when the beach substrate was frozen. An
algal density maximum occurring in this period of con-
gealed substrate was followed by a more or less rapid

decline to a markedly reduced spring level at all intervals.

 

 

 

370
At each interval along the beach transect, this decline
was followed by a mid-summer increase to density levels
approximating those of November. Subsequent to this
secondary peak in algal abundance in the beach, a general
decline in density took place until the October termina-
tion of the study. Inasmuch as algal population peaks
occurred in periods when the beach Was solidly frozen as
well as in the mid-summer period when beach temperatures
approached or reached maxima, it would seem that tempera—
ture alone was not a primary determinant in fluctuations
in quantitative abundance of the algae in the psammo-

littoral zone during the 12-month study.

earl

Pennak (1950) has described the psammolittoral as
an essentially lightless environment, basing this premise
upon reports in the literature indicating that measurable
light does not penetrate more than 10 to 15 mm. below the
surface of the sand. Sassuchin at al. (1927) exposed
photographic film at various depths in river beach sand
and reported no light reaction upon filn buried at depths
of two or more cm. below the surface, bu Heel (1948)
found the greatest depth at which a light meter recorded
the presence of light in beach sands was approximately
0.5 cm. Aleem (1950) discovered that incident light is
reduced 95 per cent in the first two mm. of mudflat sub-

strate. Tchan (1953) reported that in dry sand, buried

 

 

371
film exposed to full sunlight indicated maximum light
penetration to be 1.5 em. but was unable to state whether
light at such depth was sufficient for algal growth. He
found little, if any, difference in the extent of light
penetration in wet and dry sand.

Investigations relative to the extent of light
penetration into the sand of the Reach A1 beach transect
indicated that full sunlight penetrated at most less than
two cm. into the sand of either the hygro— or eupsammon
zones (see pages 56-57). Film buried at depths of 1.0 cm.
beneath the sand surface exhibited some evidence of faint
fogging, but only apparently where a pebble or small stone
was embedded in the one cm. of sand above the plate. All
film plates exposed at depths of 0.5 cm. below the beach
surface showed evidence of slight fogging distributed in
irregular, random patches over the plate surfaces. It was
concluded that effective light penetration into the beach
sand of the Reach A1 transect was somewhat less than 0.5
cm. with some random penetration in heterogeneous deposits
to as much as one cm. It was postulated that photo-
lithotrophs, therefore, would be concentrated in the top
0.5 cm. of sand. There was no evidence that the extent
Of light penetration differed in wet sand in comparison-
to that in dry sand.

Tchan (1953) in laboratory sand culture experi-
ments found algal populations limited to the surface five

mm. of sand. Cameron (1964) reported that algae of desert

 

 

 

372

soils are found only in the top few millimeters of sandy
soil within, essentially, the limits of light penetra-
tion. In spite of the fact that desert soil temperatures
may be as much as 50°F. higher than breast height tempera-
tures, Cameron states that light and moisture appear to
be the most important limiting factors in growth of soil
algae. Tchan and whitehouse (1953) reported that there
is no significant growth exhibited by soil algal cultures
kept in the dark, even in carbohydrate enriched media.
They conclude that the presence of organic matter does
not support growth of algae in sandy soils in natural con-
ditions. Tchan and whitehouse (£22. git.) also found that
algae of sandy soil will not grow even in light if the
soil environment is anaerobic. If sandy soil is water-
saturated, the algal community is found only in the sur—
face few mm.

Although it may be postulated that the algal
communities of the psammolittoral zone are concentrated
in the surface five mm. of sand, there is a distinct pos-
sibility that there may be considerable vertical migration
of certain phototactic elements of the community. Aleem

(1950) reports that there is evidence of diurnal migra-

found in the interstices of brachish-water mudflats. He
disc0vered few, if any, algae on the surfaces of exposed
mudflats at night, but numerous diatoms and dinoflagellates

were present at the surface in daylight. Canapati gt Qi-

 

 

 

 

er. ' i
M I

 

373
(1959) also found that diatoms and dinoflagellates showed
definite diurnal migration patterns in marine beaches.
Apparently absent from the moist sand surface at night,
forms such as fiantzschia amphioxys, Pleurosigfla aestuari,
Nitzschia closteriufl Eympodinium splcndens, and Amphidium
gellucidum were observed to migrate to the surface in
daylight. In laboratory experiments, Canapati at al.
report, these algae migrated to the surface at times
corresponding to low tides and migrated beneath the sur—
face at times corresponding to higt tides. They were
observed to react positively to light at time of low
tide and to react negatively to light at time of high
tide.

Lund (19é2) has found many motile algae in de-
posits about pond margins to be positively phototactic,
and Round and Happey (1955) discovered a vertical migra—
tion of the diatom flora of the eoipelic association of
freshwater streams in response to light periodicity.
movement of the diatom flora up to the surface of the
sediment in streams and down beneath the surface occurred
rhythmetically once every 24 hours, reaching a peak of
cell numbers at the surface at approximately the same
time each day. Round and Happey reported similar light—
induced rhythms in epipelic diatom associations in fresh-
water ponds. Included in their list of migratory diatoms

were the species Navicula cryptocephala and Navicula

 

 

 

 

 

 

374

radiosa, which were also common in the sand of the Reach
A1 beach transect.

River bank algal communities composed primarily
of Euglena and diatoms have been reported by Bracher
(1929) to exhibit diurnal fluctuations in surface abun-
dance. Bracher observed that euglenoids "burrowed" into
the mud in darkness and on overcast days, appearing again
on the surface within 20 minutes after restoration of full
illumination. Diatoms, according to Bracher, exhibited
no phototactic responses. It would appear the possibility
exists that qualitative and quantitative data gathered
from studies of the algae of the psammolittoral could be
influenced by conditions of light intensity at the time

of collections.

Black layer

Pennak (1950) has reported that in undisturbed
beaches it is a common occurrence to find distinct black
strata at depths of 5 to 15 cm. It has been well estab-
lished, he continues, that these strata are evidence of
anaerobic conditions and are produced by reduction of
iron oxides by sulfides in the presence of anaerobic de-
composition of organic matter. Sassuchin _t._l. (1927)
referred to the iron sulfide layer as the lowermost of
the four horizontal strata characteristic of the sandy

beach and characterized the layer as being 8.5 to 15 cm.

thick, very black, and containing no living "micropsammon."

 

 

 

 

 

375
This black layer has also been observed in some detail by
Aleem (1950), Ellis (1925), Bruce (1928b), Neel (194a),
and Weber (1962), among others.

Neel (I22. 223.) reported that laboratory and
field investigations of the black layer indicate that
the upper limit of the black layer marks the division
between the anaerobic and aerobic zones in the sand.

Neel found that oxygen penetration into hydropsammon zone
sands is apparently a process that includes only the top
1.5 cm. of sand. Weber (I22. 222.) found that very few
organisms occurred at depths more than two to three cm.
below the sand. He postulated that the conditions in-
ducing formation of the black layer may be major limiting
factors in restricting vertical distribution of the
psammobiota. He reported also that in transects where
the black layer was poorly developed, organisms did not
occur to depths significantly greater than in transects
with a well-developed black layer, and he concluded that
the exact importance of the black layer could not be
evaluated in his study in comparison to other factors
restricting the depth to which psammon organisms occur.
Although there are in the literature of psammolittoral
investigations considerable data with reference to the
fact that the density of psammon organisms decreases
rapidly with depth, there apparently has been no con-

clusive proof that chemical and/or physical conditions

 

 

376

inducing black layer formation act as barriers restricting
vertical migration in the interstices between sand grains.

Figure 5 presents a diagrammatic representation of
black layer distribution in months a black layer was ob-
served present during the Reach A1 beach transect study
(see also pages 59-62 for additional observations of the
black layer made during the 12-month study). There is
no evidence that a black layer was found within 29 cm. of
the beach surface from November to June. From July to
October, a black layer was observed present under the
transect within 20 cm. of the surface, but it was relative-
ly poorly defined and sporadic in appearance and extent
during those months it was present. At no time during the
study did the black layer reach the depth penetrated by
the sampling cores. Consequently, no attempt was made to
correlate algal distribution patterns in the transect with

the black layer.

__q_0 r a 0.12. ladies.

It has been stated by Pearse 22 a}. (1942) that
the sandy beach is an incubator and digestive system that
continually supplies nutrients to the adjacent sea. Neel
(1948) has noted that processes that result in temporary
l inclusion of organic matter into beach sands of lakes af—
ford as well a supply of products of decay to the lake

following decomposition of this organic matter. It is

assumed water derived frOm wave action and precipitation

 

 

 

 

 

   

37?
when passing downward through the beach dilutes the inter—
stitial water and transports its dissolved materials to
the adjacent body of water. Neel has also noted that the
high productivity of the psammolittoral may be attributed
to the close proximity of the surface producer zone and
the zone of sub-surface reducers and consumers.

Both particulate and dissolved organic matter
within the beach interstices can be considered present
and future sources of nutrient to photolithotrophs and
consumers living in the psammolittoral and its adjacent
water basin. Organic matter, too, may retain moisture
and thus retard rate of evaporation from the interstitial
voids.

Koepcke and Koepcke (1952), Heel, (1948) and
Pennak (1939, 1940, 195 ), among others, have discuss d
the origin of particulate and dissolved organic matter
found within the interstices of the sandy beach. Apparent-
ly there have been no detailed quantitative studies rele-
vant to the relative amounts of auto- and allochthonous
organic matter within the beach. Unfortunately, too, as,
pointed out by Ruttner-Kolisko (1956), data pertinent to
organic matter content in beaches are thus far very un-
satisfactory. She indicates that the explanation is in
the serious error potentials in present methods of deter-
mination and in the difficulty in differentiating be—
tween suspended particulate and dissolved organic matter

in determination of organic content within the beach.

L

 

 

 

 

 

ivy-gm; _A ,

37B

Ruttner—Kolisko (123. git.), Schwoerbel (1961),
and other European investigators have, by and large, re-
lied upon the KmnOA-uptake method of determining organic
content. The English-language literature of psammolittoral
investigations and those of related benthic littoral
studies involving organic matter determinations indicates
that most have utilized the weight loss on ignition
technique in determination of organic content (see Pennak,
1940; Neel, 1948; Pearse at al., 1942; Aleem, 1950; Pirrie
gt al., 1932; Bracher, 1929; Lund, 1942). In far too
many instances, the methods are not given at all. Some
data have been reported as mg./1Dcc. of sample, some as
per cent dry weight, and some as per cent wet weight.
Pirrie _t _l. (lag. gig.) have noted that data based on
weight loss on ignition techniques are suspect if con-
siderable amounts of Ca503 are present in the sand.

Data resulting from monthly determinations of
percentage volatile matter on a dry gross weight basis
for each 25 cm. interval along the Reach A1 beach tran-
sect are presented in Table 9 and Figure 6 (see also
pages 68 to 72 for detailed results of monthly determina—
tions of percentage volatile matter, i.e. organic matter,
in the beach substrate). Percentage volatile matter in
gross beach samples ranged from 10.9? per cent recorded
on December 8 at the 25 cm. interval to a minimum of 0.39
per cent recorded on May 6 at the same interval. In each

month there was a general tendency for percentage volatilq

 

 

 

 

379
content to diminish with distance from shoreline. This
tendency was also exhibited in 12-month means. The great-
est range between extremes of the transect occurred when
a differential of 10.49 per cent was recorded in the

December collections. The smallest range, 0.94 per cent,

occurred in the may collections. The 12-month mean range
between extremes of the transect was 3.22 per cent. As

reported on page 68, the greatest differentials between
volatile content at the most lakeward interval and the
+300 cm. interval occurred during those months in which
the transect substrate was in a frozen condition. The
least differentials occurred in the spring and early
summer months. Although it might be postulated that the
retardation of organic decomposition in winter months
might account for the larger winter differentials, the
fact should be considered that algal populations tended
to be largest when the beach was frozen and smallest
during the spring and early summer. Data on Table 9 do
not indicate a gradual build—up of organic content at

any interval in the beach during the winter months.

The weight loss on ignition technique of organic
content determination does not differentiate between
organic material within living cells and that as detritus
or in solution. It should also be noted that some weight
loss in this technique may be from thermal decomposition

of carbonates and bicarbonates. Although the method

employed in carbonate and bicarbonate determination along

 

 

 

 

330

the transect was highly subjective (see page 44 for
methods), reference to Table 11 indicates an appreciable
differential in bicarbonates between extremes of the
transect, the amount of bicarbonates each month increas-
ing with distance toward the water line. It is, therefore,
possible that an appreciable degree of the differential
between extremes of the beach transect might be attrib-
utable to thermal decomposition of carbOnates and bicar-
bonates.

Noted on page 39 is the considerable similarity
in 12-month means of per cent volatile material and per
cent water on wet gross wet basis at each interval along
the transect (see also Figure 6). Readings from the +390

cm. interval to the +175 cm. intervnl showed distinct

graphic correlation between means of volatile material
content and that of water. From the +175 cw. interval
lakeward, the marked variation between the two factors
considered perhaps was attributable to the increasing

proximity of the transect intervals on the phreatic sur-

/

far:o end the lake L.olf. It n. h"‘h llf“ vlso th L
,Here apparently was we axplar +177 on o granulanetrlr
basis for the irregularities in rater retention by the

substrate at those intervals lrnjwari from the +173 c;
interval. The hypothesis might be made that percentage
water content at each interval along the transect was
directly related to porosity and distance from the shcr3~

line and or phreatic surface un to a certain crit'
. . 1C8

 

 

 

 

 

 

distance —- about the +175 cm. mt rval For most of the
12-month study -— and beyond the critical distance it
became secondarily related to the amount of organic matter
present.

Reference to data presented on Table 12 and Table
13 reveals that, during those months in which the amounts
of dissolved volatile matter were determined for both
beach interstitial water and the water of Reach A1, the
amount of volatile nateri al in interstitial water was
ordinarily considerably higher than in lake water (see
also figures 11 and 12 and pages 87 to 90 for detailed
observations upon results of monthly de eter ruinations of
percentage volatile matter in water of Reach A1 and in
interstitial water from the beach transect). The mean
amount of volatile material in lake water during the
April—October period when it was possible to withdraw
interstitial water from the beach was 266.7 mg./l. For
the sane period, the mean amount of volati is material in

interstitial water was 390.3 dg./l.

X

_:tor cont-a_nt

Neel (1948) has stated that it is the wgter con—
tained in the voids between sand grains that forms the
habitat of ”psamno—organisms” and that this habitat te—
comes increasingly restricted when the amount of water

in the interstices lessens. Pennak (1940, 1950) also

indicates that the complex microscopic flora and fauna

 

 

 

 

 

382

oF the psammolittoral inhabit the capillary water held

between beach sand grains. Although the argument may be

somewhat trivial, the term psammon as deFined by Sassuchin
gt 3}. (1927) apparently reFers to organisms oF the sand
rather than to those living in water within the sand.

And wiszniewski (1934a) in deFining hydropsammon, hygro-
psammon, and eupsammon writes oF these organisms living

in their respective beach zones rather than in the water

0F their respective beach zones. The Fact, however, that

wiszniewski reFerred to the psammolittoral as a unique

type oF aquatic environment and used limnological tech—

niques in its investigation may have led to the tacit
assumption by subsequent investigators that the ”micro-
psammon," which diszniewski reFerred to as the aquatic
microorganisms living in the sandy beach substrate,

obviously live in the interstitial water. Too, Strangenberg's
(1934) use oF limnological techniques in the investigation
0F the psammolittoral and his reFerence to the environ-

ment as an ”extremely eutrophic water medium” abetted this

concept 0F the psammon as aquatic organisms. Pennak (1939)

also has stated that the psammon are primarily the genera
and species encountered in the true aquatic environment.
when the sizes 0F larger members oF the Fauna
living in the eupsammon zone are compared with data 0F
porosity, grain size, and water content oF the substrate
From which these organisms have been reported, an element

oF doubt arises with respect to whether these Forms should

 

 

383
be considered aquatic or interstitial Forms. Lambe (1951)
has stated that interstitial air has a relative humidity
oF 100 per cent. It has also been noted that Dreshkina
(1960) reported that capillary discontinuity occurs in
sands oF 0.10 to 0.08 mm. mean grain diameter at approxi-
mately Four per cent moisture content. Moisture content
in the eupsammon zone Frequently is less than Four per
cent at the surFace, but the literature indicates move-
ment or migration oF certain elements 0? the psammon
within the eupsammon zone. It would seem possible, there-
Fore, that some members oF the psammon move about in and
across interstitial water discontinuities. Thus, the
"aquatic” organism 0F the psammolittoral perhaps survives
in a humid but not necessarily aquatic environment. The
Fact, as well, that in the Reach A1 study many 0F the
algae Found are characterized as "terrestrial" in phyco-
logical literature suggests that the concept oF psammon
organisms as aquatic Forms may need some revision or raw
Finement.

It is a well-established principle in the litera-
ture 0F psammolittoral studies that the amount oF water
per unit volume oF sand diminishes with distance From
shoreline (Sassuchin gt al., 1927; wiszniewski, 1934a,
1937a; Ruttner—Kolisko, 1953, 1954, 1956; Neel, 1948;
Pennak, 1939, 1940, 1950). Consequently, the amount oF
air per unit volume oF sand increases with distance From

shoreline until near the landward limit 0F the psammon

 

 

 

384

any water present is hygroscopic, held in a thin Film
upon sand and colloidal particles by Forces 0F adsorption.
Instances in the literature where exceptions occur to the
rule 0F decreasing water content with distance From shore—
line have been attributed to random concentrations oF
water-retentive prganic matter, irregularities in the
beach proFile, variations in porosity and grain size,
shade patterns, or drainage patterns within the beach
(see Ruttner-Kolisko, 1953, 1954, For discussion 0F
Factors producing variation in horizontal and vertical
water retention patterns in the sandy beach).

UnFortunately, as also pointed out by Ruttner-
Kolisko (1956), the lack oF standardization oF methods oF
presenting water content data makes generalizations con-
cerning water content in the sandy beach subject to con-
siderable error. Some investigators have reported water
content as percentage wet gross weight, some as percentage
dry weight, some as per cent saturation, some as per-
centage oF void volume, and others as percentage oF gross
sample volume. Evans (1958) encourages the presentation
0F moisture content data on the basis oF percentage wet
weight. Hough (1957), however, states the reporting oF
percentage water content on a dry weight basis is becom-
ing increasingly the accepted standard in soils engineer-
ing. But he also states that it is in some instances
desirable to indicate water content as a percentage oF

the total or wet weight 0F a soil sample and notes this

 

 

 

 

 

385
method was the prevailing practice "some years ago." It
should be noted that iF water content is reported on a
dry weight basis the water content values may exceed 100
per cent. It would seem advisable, thereFore, in ecolo-
gically oriented psammolittoral studies to report water
content on a wet gross weight basis in Future studies to
Facilitate statistical treatment 0F the data, particular—
ly in calculation oF r relationships. Too, it should be
noted that air-dried soil, even desert sands, normally
contains slight but measurable hygroscopic water (Hough,
£22. git.). ThereFore, it is also encouraged that water
content data be based on oven-dried rather than air—dried
samples.

Hough (£92. git.) has stated that For any given
soil the possible range 0F water content values is so
wide that any general statements regarding water content
are oF little value. In addition, the only other water.
content data relative to psammolittoral studies reported
in the literature are based upon single collections
isolated in time and/or space or upon other than wet or
dry weight percentages oF oven-dried soil or sand. Probab-
ly the most extensive data oF water content in beach sand
are those reported by Pennak (1940, 1950), but these are
reported as percentage saturation (see also Ruttner-
Kolisko, 1953, 1954).

Table 7 presents the results 0F determinations

0F percentage water content on gross wet weight basis For

 

 

 

 

 

386

each oF the 25 cm” intervals along the Reach A1 beach
transect From November 9 to October 10. Water content
data For the same intervals during the same time period
are presented also in Table 8 as the percentage water con-
tent oF monthly sand samples at each 25 cm. interval on
wet weight basis expressed as per cent saturated capacity
(see also pages 62 through 68 For detailed observations
upon the results 0F beach sand water content determina-
tions made during the 12—month transect study).

Twelve-month mean percentage water content ranged
From 3.00 per cent at the +300 cm. interval to 24.57 per
cent at the +25 cm. interval. An inverse relationship
was apparent between the 12—month mean percentage water
content 0F any interval and its distance From the shore-
line. With Few exceptions, this relationship occurred
as well at each monthly collection period. Percentage
water content during the 12-month study ranged From a
low oF 0.01 per cent recorded on August 6 at the +300 cm.
interval to a high oF 49.18 per cent at the +25 cm. in~
terval on December 8. No abrupt diFFerential was apparent
in water content at adjacent intervals which would sug—
gest the obvious division between the hygro- and eupsammon
zones (see also Figure 6). As has been noted previously
(pages 71, 380, and 381), however, there is some indica-
tion that the limit 0F capillarity may have been reached

at approximately the +175 cm. interval as evidenced by

 

 

 

387
the rapidly decreasing water content From that point up-
shore during most months oF the 12-month study.

As reported on page 64, maximum diFFerentials in
percentage water content between shoreline and the +300
cm. interval occurred at those collection periods during
which the substrate was Frozen. The minimum range, 12.78
per cent, occurred on April 6 within 24 hours aFter a
heavy rain in the collection area. The 12-month mean
range was 21.57 per cent between the +300 and +25 cm.
intervals. The anomaly oF maximum diFFerentials during
periods when the beach was Frozen was attributable to
the very high moisture content oF the +25 cm. interval
in the December-March period rather than to low readings
at the upshore points during the same period. It was
during the winter months that water-content maxima were
recorded in the upshore intervals along the transect.

0n the basis oF 12~month means, the +300 cm.
interval contained only 12.21 per cent as much water as
did the +25 cm. interval. 0n collection dates in
December and May through October, the driest interval
(usually +300 cm.) contained less than 10 per cent 0F
the amount oF water present in the sand 0F the wettest
interval (usually +25 cm.).

For discussion oF Table 8 and results 0F deter-
minations oF the amount oF water present in monthly
samples From each interval expressed as per cent satura-

tion, the reader is reFerred to pages 64 through 67. It

 

 

 

388
may be noted here, however, that on the basis oF 12-month
means the +300 cm. interval contained only 14.2 per cent
oF theoretical maximum water retention capacity and the
+25 cm. interval contained 102.8 per cent. The 12-month
means oF certain intervals in excess oF 100 per cent
theoretical maximum capacity was attributed to Fluctua-
tion in organic content in the sand 0F shoreline inter-
vals during the 12-month study. An inverse relationship
between the amount oF water actually held by samples From
each interval and distance From water line was apparent.
Potential water retention capacity was most nearly real-
ized those collection dates on which the beach was Frozen
or aFter a heavy rainFall. Greatest departure From po-
tential water retention capacity occurred during summer
months when the most landward intervals Frequently con-
tained less than 10 per cent 0F their theoretical water
retention capacity.

Pennak (1940) has presented mean percentage satura-
tion values For 18 series oF determinations made on the
amount oF water held in 12 wisconsin Freshwater beaches
with reFerence to vertical variation in the amount oF
water held at one cm. interVals From surFace to eight cm.
below the surFace. Pennak Found no vertical water reten-
tion gradient at the zero interval. At the 50 cm. inter-
val, however, he Found the lowest Five cm. practically
saturated and the top three cm. showing evidence oF dry-

ing with 86 per cent saturation in the uppermost cm. All

 

 

 

 

 

 

upshore intervals exhibited
350 cm. intervals, the mean
For the top cm. 0F sand was
the bottom cm. reached only

reported very Few organisms

389

vertical gradients. At the
percentage water saturation
less than one per cent, and
23 per cent saturation. He

in sand samples which had a

saturation oF less than 10 per cent.

Apparently, Pennak (1939, 1940, 1950) believed
the amount oF water within the sand to be primarily re—
sponsible For vertical distribution oF psammon organisms.
He reported (1950) that more than 95 per cent 0F the
psammon are located in the top six cm. oF sand and more
than 50 per cent are Found in the top Four cm. In drier
zones at distances oF 250 to 350 cm. From water line,
the psammon tend to be concentrated at depths oF Four to
eight cm., he stated, chieFly because oF lack oF inter—
stitial water toward the surFace. He has also noted
(1939) that there is a tendency For copepod concentra—
tion to shiFt downward where the surFace oF the sand be—
comes dry.

Ruttner-Kolisko (1956), however, attributes her
observation that all animal groups in the psammolittoral
exhibit a tendency For maximum population density to
move downward with increasing distance From the water
line to be due to increased air circulation at greater
depths in the absence oF interstitial water. It has been
noted previously in this chapter that Neel (1948) con—

cluded the psammon to be restricted essentially to the

 

 

 

   

390

uppermost two cm. oF sand because oF the Failure oF light I

to penetrate more deeply than 1.5 cm. into the sand and

the close proximity oF the anaerobic black layer. Neel

reported that the concentration oF psammon near the sur-

Face indicates the dependence oF heterotrophic Forms upon

photosynthesis For nutrition and oxygen. Those organisms

carried to lower levels by waves and gravitational water,

he believed, must move back to the surFace or perish.

Sakharova (1963) has also reported that 90 per cent oF

the psammon Fauna are concentrated in the upper two_cm.

oF substrate but makes no generalized statement as to the l

cause. '
As Far as can be determined From a more or less

exhaustive search oF the literature oF psammolittoral

studies, only Heel (1948) and Baklanovskaya (1963) have

 

made more than superFicial reFerence to the algal com-
ponent oF the psammon. Neither presents a critical eval-
uation oF the vertical distribution oF algae in the sandy
beach, although Neel does present in tabular Form evi-
dence that relatively Few algae are Found at depths great-
er than two cm. Consequently, it must be assumed that on
the basis oF inFormation presented in reports oF studies
oF terrestrial algae From substrates having textures
similar to those oF the psammolittoral that psammon algae
are Found essentially in the top cm. 0F the sand and
apparently are most heavily concentrated in the tOp Few

mm. (Tchan, 1953; Tchan and whitehouse, 1953; Cameron,

 

 

   

391
1964; Allem, 1950; Bracher, 1929; Shields and Durrell,
1954; Evans, 1953, 1959, 1960; John, 1942).

There is, however, considerable evidence 0F ver-
tical migration oF soil or psammon algae in response to
moisture variation. Peterson (1935) has reported that
algae migrate about in the top Few mm. oF soil in response
to changes in moisture. Aleem (£22. git.) Found that
Free water, at least 30 percent by wet weight, is neces-
sary For vertical movements oF diatoms and dinoFlagellates
in mudFlats. Bracher (lgg. git.) concluded that the
vertical distribution oF euglenoids and diatoms in river
shores was related to water content inasmuch as labora-
tory and Field observations indicated that euglenoids
occupied dry ridges and the diatoms moist Furrows in the
exposed shore. Evans (1958, 1959, 1960) has Found evi-
dence that motile algae descend as much as Five mm. into
pond margin litter and mud with decreasing water content
and rise to the surFace when surFace substrate water con-
tent increases. Diatoms, he reported, are Found on the
surFace when relative humidity is 70 per cent or more
and migrate downward when the humidity Falls below 79
per cent. Among the list oF migratory species presented
by Evans were Pinnularia viridis, E. microstauron, E.
subcapitata, E. braunii, Chlamydomonas sp., Euglena sp.,

Trachelomonas volvocina, Eunotia sp., Nitzschia galea,

and Gomphonema garvulum.

 

 

 

 

392

Although no attempt was made to determine vertical
distribution patterns For the algal component oF the Reach
A1 beach transect psammon, it is believed that core samples
taken to depths oF two cm. negated any irregularities in
counts produced by vertical migration, iF it occurred.

There is abundant evidence in the literature oF
psammolittoral investigations For the generalization
that horizontal distribution oF the psammon varies From
hydropsammon zone to the eupsammon zone and within each
zone as well. Ruttner—Kolisko (1956) in review oF the
literature oF psammolittoral studies notes that the diF-
Ferent taxa oF the psammon Fauna exhibit diFFerent pat-
terns oF horizontal distribution. She attributes these
distribution patterns to variations in the amount oF
oxygen, organic material, and water content From the
lakeward to upshore extremes oF any psammolittoral tran-
sect.

In a review oF numerous psammolittoral studies,
Pennak (1950) concluded that the Protozoa, Turbellaria,
Nematode, Castrotricha, and Oligochaeta display no par-
ticular horizontal distribution patterns in the sandy
Freshwater beach. These groups, he states, are scattered
about the psammolittoral at random or are conFined to
the hygropsammon where suFFicient interstitial water is
present. There apparently is in the literature OF in—
vestigations oF the marine or Freshwater psammolittoral

no conspicuous exception to the principle that the

 

 

 

 

 

 

393
hygropsammon or eupsammon zone. It should be noted,
however, that Schwoerbel (1961) has Found the hyporheic
portion oF the lotic psammolittoral to be quite produc—
tive, but it is not clear how productive this environment
may be with respect to the exposed river shore.

Pennak (lgg. git.) states, on the other hand,
that the Rotatoria, Tardigrada, and Copepoda oFten show
speciFic distribution patterns For wet or dry zones at
speciFic distances From the water line. In dry periods,
he continues, when the lake level Falls, there is a cor-
responding activo horzontal migration oF the microFauna
that may be easily traced (see Pennak, 1939, 1940, 1950,
For listings oF speciFic distribution patterns oF re-
presentatives oF the taxa listed above). Although
horizontal distribution patterns are not discussed at
length in their publications, Jiszniowski (1937a, 1934a,
1934b, 1934:), Sassuchin at al. (1927), and Ruttner-
(olisko (1953, 1954) present tables that do indicate
horizontal distribution patterns For certain components
oF the psammon Fauna.

Neel (1948) has concluded that position with
reFerence to the water line appeared to have little eFFect
upon numbers oF microorganisms except near the two ex—
tremities oF the six lake psammolittoral transects he
studied. He Found that low populations at the lakeward
end oF the transects were due to deposits oF marl and

organic debris in interstices. Relatively small amounts

 

 

 

 

 

394

oF interstitial water near the landward limits oF the
transects he Felt were responsible For low densities
there. He Found no signiFicant decrease in the number oF
organisms above water line on the transects until the
surFace oF the sand became obviously dry.

There are Few published studies oF the psammo—
littoral which include data relative to horizontal dis-

tribution oF the algal component oF the psammon.

diszniewski (1934a) makes only passing reFerence to the

algae in his review oF psammolittoral investigations up

to 1934. Although he noted that rarely do ”blooms" oF
algae appear in the hydropsammon, noticeable growths which
he postulated were diatoms appeared on the submerged sub—
strates near the water line 0F a number oF sampling areas.
Round (1965) has termed this association the "epipsamm:n“
and describes it as an epipelic association OF motile
diatoms Found in Freshwater sandy sediments. Although
Round classiFies the association as epipelic as contrasted

He notes that the diatoms are Firmly at-

to epilithic,
tached to the sand grains. fliSZniewski (lgg. gig.) stated
that in the hygropsammon zone algae appear in small
numbers and seldOm Form blooms. A characteristic oF the
eupsammon zone, he continued, ‘5 the large numbers oF
algae which typically Form blooms at times to give the
beach an intense green color, the ”green horizon" des-

cribed by Sassuchin gt a1. (1927). There the gree

...——

 

 

 

 

   

 

 

horizon, wiszniewski stated, may extend several cm. below
the sand surFace.

Although Sassuchin _e_t‘ _a_1_. (_l_o_c_. 9_i_t.) do not dis—
cuss horizontal distribution oF the psammon algae in
length, their tables do indicate algal distribution pat—
terns. It should be noted that Sassuchin gg gl. based
their generalizations about psammon upon data derived
From samples taken almost exclusively in what must have
been the eupsammon zone. Thus, reFerence to their some-
what limited and subjective data relative to horizontal
distribution oF psammon algae may lead to erroneous con-
clusions. Sassuchin g3 gl. ascribed the limits oF hori—
zontal distribution oF the green horizon to the amount
oF moisture in the sand, and they noted that the green
horizon contained almost all the microscopic biota oF the
sand. They Further noted that algae, particularly diatoms,
comprised the principle portion oF the micropsammon. Host
abundant in the green horizon, they stated, were Phormidium

sp., Oscillatorio tenuis, Navicula radiogg, fl.

cryptocelphala, Nitzschia palea, Chlamydgmonas sp., and

Scenedesmug quadricauda.

 

Quantitative data reported by Ruttner-Kolisko
(1953, 1954) relative to number oF algae per cc. oF sand
samples in hydro-, hygro—, and eupsammon zones present
no clear evidence oF “preFerence” oF various groups For
particular zones, and she makes no generalizations with

regard to horizontal distribution oF psammon algae.

 

 

 

 

 

396
Neel (1948) although presenting probably the most
voluminous quantitative and qualitative data relative to
psammon algae reported in the literature oF psammolittoral

studies, in quantitative tables lists only the genera

Herismopedia, Pediastrum, Scenedesmus, and Cosmarium,

lumping all others as diatoms, Chroococcales exclusive

oF Merismo edia, or green algae exclusive oF Pediastrum,

Scenedesmus, and Cosmarium. He concluded that diatoms,

blue—green algae, and "others" exhibited no general
horizontal distribution patterns along six transects
extending From as Far as 50 meters into the hydropsammon
to the eupsammon Five meters upshore From the water line.

Baklanovskaya (1963) in summary oF a 12—month
study oF the algal Flora oF a Freshwater sandy beach,
reported 50 species present in the hydropsammon, 58
species in the hygropsammon, and 71 species in what ap—
parently was the eupsammon zone. He noted that in number
oF individuals the blue-green algae dominated the algal
community at all beach stations, particularly those at
the water's edge. Blue—green and green algae predominated
in the hygropsammon zone. Green algae and diatoms were
cemmon in the upshore stations. He reported that in the
majority oF instances the diatOms were Found in a dying
state.

Shields and Durrell (1964) in a review oF 184
reFerences related to systematic and physiological studies

OF soil algae and their interrelations with their

 

 

 

 

39?
environment have concluded that the soil algal community
structure is inFluenced less by the chemistry oF the sub-
strate than by the amount oF available moisture. They
note that where water is not a limiting Factor, soil
nutrients tend to govern luxuriance oF the soil algal
Flora and the number oF species represented. The abun-
dance oF algae in soils, they state, is closely related
to the amount oF necessary mineral salts, Favorable pH,
and moisture conditions. Shtina and Bolyshev (1963) in
studies oF algal communities oF arid and desert steppes
Found the composition oF ephemeral crust-Forming algal
communities to be essentially similar in all sample sites
with the expression oF two basic community types depend—
ing upon the amount oF water present.

Tchan and Jhitehouse (1953) in laboratory experi-
mentation relative to variation 0F algal populations in
sandy soils discovered that optimum growth oF soil algae
occurs at approximately 63 per cent water-holding capacity.
No Further increase in growth rate was noted when soil
algae were cultured in soils retaining more than 35 per
cent oF water—holding capacity. to growth was observed
by algae in soil cultures containing less than 12 per
cent oF water-holding capacity, approximately three per
cent water by weight. Tchan and Hhitohouse concluded
that the minimum amount oF water For algal growth is
between 12 and 24 per cent oF water—holding capacity.

They Found algal populations in air dry soil seemed to

 

 

 

 

, «if-v -£§y_»:§"%> .\

398

stabilize at about 65 per cent their number in optimum
conditions. Thus, Tchan and whitehouse concluded, soil
algal populations tend to stabilize in soils below 12
per cent and above 65 per cent water-holding capacity.
They state that between these limits water content oF
soil apparently plays a major part in controlling rate
oF algal growth.

According to Shields and Durrell (log. git.),
the optimum moisture range For soil algae is about 40 to
60 per cent oF soil moisture-retention capacity, but they
review reFerences in the literature where blue—green
algae and diatoms have survived For 26 to 73 years in
sealed containers retaining soil having From only three
to ten per cent water content. Reported particularly
resistant to prolonged drought were Chlorococcum humicola,
Trochiscia aspera, Phormidium tenue, Nitzschia palea,
and Nodualaria harveyana -- all 0F which were present in
samples taken during the 12-month Reach A1 beach transect
study. Peterson (1935) has reported that HEDEEEEDEE
amphioxys and Navicula mutica are Fully active in soils
containing only 5.2 per cent water. It might be noted
here that during the Reach A1 transect study counts oF
algae ranging From 1,369 to 12,846 individuals/cc. 0F
sample were recorded From samples containing less than
one per cent water on wet gross weight basis. Shields
and Durrell note that Chlorococcum humicola, a species

conspicuous in quantitative and qualitative samples From

 

 

 

 

 

399
all Four intervals sampled during the Reach A1 study, is
considered to be the universal soil alga.

Shields and Durrell ($22. git.) state that the
physiology oF drought resistance exhibited by certain
algae is not known. The presence or absence 0F a sheath
may not be a Factor, since many 0F the most resistant
Forms lack sheaths. Fritsch and Haines (1923) have re-
ported in some length relative to the remarkable resis-
tance oF some terrestrial algae to drying. Fritsch
(1922) has noted that in England the most rapid growth
oF soil algae occurs in the winter, and he has observed
that air dry algae may display a steady increase in weight
when exposed to a humid atmosphere, showing the marked
absorption oF atmospheric moisture when humidity is high.
Durrell (1962) has reported that air-dried Nostoc can
absorb twelve times its own volume and weight oF water
in six minutes. Durrell (lgg. git.) has listed 23 species
oF algae living in Death Valley environments where the
rate 0F evaporation exceeds the amount oF rainFall by a
Factor oF 100, where the surFace temperatures may reach
1900F., and where the only available water Frequently
contains at least Five to six per cent sodium chloride.
Eight oF the 23 species listed by Durrell —— including
Chlorococcum humicola, Phormidiufl tenue, Hicrocoleus
vaoinatus, Anacystis montana, Uscillatoria agardhii,
Phormidium ambiquum, E. angustissimum, and Egflphgsph§3££fl

aponina —— were also common in the Reach A1 beach

 

 

 

 

 

 

400
transect, although not necessarily showing "preFerence"
For the driest intervals along the transect.

Microcoleus va inatus, one oF the most common
algae present in the Reach A1 beach substrate, is report-
ed by Durrell and Shields (1961) to be one oF the most
important algae involved in the Formation oF algal crusts
on the moist surFace oF soils oF the arid southwestern

United States. Nostoc, Phormidium, Nodularia, and

fiicrocoleus va inatus, according to Durrell and Shields

are the most common crust-Farmers on desert soil. They
note that crust samples air dried For more than one year
become green on the second day aFter exposure to moisture.
The colloidal sheath oF fl. vaginatus has great water-
holding capacity. The development 0F this alga and others
oF the crust complex is apparently Favored by alkaline
conditions.

Cameron (1964) in a review oF the literature oF
the soil algae oF arid areas, particularly the south-
western United States, lists 67 algal t'xa characteristic
oF arid soils. FiFty oF the ED algal species listed by
Cameron are cyanophytes, and only eight are green.
Cameron states that diatoms are not common in dry areas.
The most cemmon arid Forms, according to Cameron, are

Microcoleus vaginatus, fl. chthonoplastes, Scytonena

 

hoFmanii, Schizothrix macbridei, Nostoc muscorum, and
Plectonema nostocorum. All six have been reportcd to

resist dessication For as much as Five years and to

 

 

 

 

401

thrive at 90 to 100°F. in aqueous cultures. Microcoleus

 

vaginatus and fl. chthono lastes, reported Cameron, were
Found in 75 per cent oF soil samples examined and were
the most rapid oF the soil Flora in reporduction and
development. It is oF interest that oF the 60 algal
species listed by Cameron as characteristic oF arid soils,
almost one—third were also present in samples taken From
the Reach A1 beach transect. These included Anacystis
montana, Coccochloris peniocystis, Nodularia harveyaga,
Aulosira lgxg, Lyngbya aestuarii, flicrocoleus acutissimus,
fl. chthonoplastes, fl. va inatus, Qspillatoria Formosa,

Q. limosa, Q. sancta, Q. tenuis, Phormidium ambiguum,

and Porphyrosiphon Fuscus.

Evans (1958, 1959, 1963) in a series oF studies
concerned with the survival oF algae during unFavorable
environmental conditions discovered that the algal Flora
in marginal deposits and upon exposed banks 0F Fresh-
water pools decreasos rapidly in Frequency and number oF
species when soil moisture content Falls to less than 50
per cent On wet weight basis. He noted also succession
OF algal populations as moisture content oF soil changes
and evidence oF surFace horizontal migration oF diatoms
and euglenoids, perhaps in response to localized accumula-
tion oF nutrients or moisture. Evans also reported that
although desmids in general could not us ally withstand

exposure and drying in the vegetative state, Cosmarium

 

imDTBSSulum, Q. botrytig, and Closterium sp. were drought

 

 

 

 

 

402

resistant. It might be noted here that Q. impressulum
was the most temporally and spatially widespread oF the
desmids in samples taken during the Reach A1 beach
transect study, but Q. botrytis was present only in
Reach A1. Diatoms, Evans noted, were well able to stand

exposure, Nitzschia palea populations in Fact increasing

 

upon drying. Also noted by Evans to be particularly

drought resistant were the species Scenedesmus guadricauda,

 

Stauroneis ohoenicenteron, Nitzschia linearis an

,.

 

Oscilla toria tenuis —— all also common in e~eples col-
lected during the Reach A1 beach transect study.

dith reFerence to relationships between water
content oF the Reach A1 beach transect and distribution
oF the algal components oF the psammon in space and time,
attempts to reconcile Fluctuations in qualitative and
quantitative abundance oF any particular algal tax0n From
Division to species level with the monthly or mean water
content at a given inter'al can only be termed Fatuuus.
The biological data an ered in ta Ml s 13 through 40,
illustrated in Figures 27 through 33, and discussed in
pages 143 to 315 Frequently suggest that distribution
patterns For any given taxon mid j'ut possibly be related
to the monthly or mean water center t at certain tra.sect
intervals. For instance, when qualitative horizontal
distribution oF algae is considered at Division level
over the entire 12-month period oF the Reach A1 study

(see Table 33), it is seen that in the Cyanophyta,

 

 

 

 

 

 

403

Chlorophyta, and Chrysophyta there is a pronounced
diminution in number oF species shoreward From the +25
cm. interval to the +300 cm. interval.

At the Order level, the same tendency is evident
in the Hormogonales, Chlorococcales, Zygnematales,
Centrales, and Pennales. No general pattern oF diminu-
tion oF species with distance From shoreline was clearly
evident For the Chroococcales, Euglenales, Tetrasporales,
or Dedogoniales. The number oF species assigned to the
Volvocales, Heterococcales, and Chrysotrichales on the
other hand increased with distance From shoreline.

ReFerence to Table 3? reveals that there was
exhibited in each monthly series oF qualitative samples
a pronounced decrease in total number oF species shore-
ward along the transect (see also pages 262 through 301
For summary recapitulation oF qualitative data relative
to the temporal and spatial distribution oF the algal
component oF the psammon oF the Reach A1 beach transect
during the 12—month study).

Tables 39 and 43 and Figures 35 and 35 also in-
dicate a pronounced decrease in most months in total
number oF algal individuals per cc. 0F sand From the +25
cm. interval to the +300 cm. interval oF the beach study
(See also pages 301 through 315 For a summary recapitu—
lation oF quantitative data relative to spatial and
temporal distribution oF algae oF the Reach A1 beach tran—

sect during the 12—month study).

 

 

 

 

404

 

That species such as Dactylococcopsis rhaohidioides,
fiicrocolegs acutissimgs, Dscillatoria agardhii. Schizothrix
lacustris, Botrydiopsis arhiza. figvicula capitata, and
Heidium dubium -- to list just a Few —- apparently exhibit-
ed "preFerence" For certain intervals along the transect
is clearly evident in Table 35.

In spite oF the Fact that both qualitative and
quantitative algal abundance diminished with landward
distance From the Reach A1 waterline, and despite the
evidence that certain algal species showed deFinite dis-
tribution patterns with respect to water content within
the substrate, there is no justiFication For cencluding
that it is water content that is responsible For these
phenomena. Obviously, water content was involved in the
distribution oF the psammon algae in the Reach A1 beach
transect, and water content is probably the primary deter-
minant cF temporal and spatial distribution oF the algae
oF the psammolittoral environment. But it is also obvious
that controlled laboratory experimentation will be neces—
sary to determine to what degree water content does in—
Fluence the distribution oF algae in the psammolittoral
zone.

In an attempt to determine statistically whether
there was evidence oF water content inFluencing the com-
position oF the algal component oF the psammcn Flora
along the Reach A1 beach transect, the amount oF water

present monthly at each interval expressed as percentage

 

 

 

 

 

 

435

wet gross weight was plotted against algal index of
diversity at each interval sampled each collection date
(see pages 315 to 325 for methods of determination of
algal index of diversity and relevant discussion there—
of). As indicated on pages 340 through 344 and illus-
trated on Figure 43, calculation for standard correla-
tion coefficient (r) between algal index of diversity
and water content gave a r value of 0.83258, well within
95 per cent confidence limits. According to variance
interpretation tables for correlation coefficients when
r=G.83258, approximately 64 per cent of the variance in
index of diversity during this study could be attributed
to variance in percentage water content within the beach
sand substrate.

Apparently the percentage water on wet gross
weight basis at each interval was the primary factor in
determination of qualitative and quantitative aljal
abundance at the beach transect intervals sampled. The
relationship was direct. Although there is, therefore,
statistical evidence that water content did determine
in part the number of species of algae and the number of
algal individuals per cc. of sand at any interval sampled
on the Reach A1 beach transect, there is no evidence
that water content determined the appearance of any given

algal taxon at any given beach interval.

 

 

 

 

 

 

496

Chemistry of interstitial and
lake water: pH

A review of pH data reported in the literature of

 

psammolittoral studies indicates the principle exists
that pH values, almost without exception, increase with
distance shoreward from waterline, i.e. the relative
acidity of interstitial water increases the further the
sampling site is located from the adjacent water basin
(Neel, 1948; Pennak, 1939, 1949, 1959; Pirrie gt al.,
1932; Stangenberg, 1934; weber, 1963; Ruttner-Kolisko,
1956; fiiszniewski, 1934a, 1936a, 1936b, 1936c, 1935;
Schwoerbel, 1951). It should be emphasized that in almost
every report in the psammolittoral zone literature in
which pH is given, the pH has been determined upon inter-
stitial water drawn up from the beach with an aspirator.
Further, in most instances the determinations have been
made upon interstitial water collected only 100 cm. or
less from the waterline.

Tables 12 and 13 include pH data from the water
of Reach A1 and interstitial water respectively (see also
Table 10, Figure 7, and pages 71 to 77 for additional
information relative to pH data recorded during the 12-
month Reach A1 beach transect study). Lake surface water
pH immediately offshore from the transect ranged from 7.2
to 9.5. Median pH during the 12-month period of collec—
tions from the lake was 8.4 with a range of 2.3 pH units.

Interstitial water pH ranged from 6.9 to 8.3 with a median

 

 

 

 

407
of 7.8 and a range of 1.5 pH units (see page 41 for des-
cription of site of interstitial water collections).

In the few instances where the psammolittoral
zone investigators have ventured a hypothesis for this
increase in pH with upshore distance from the water basin,
it has been attributed to high levels of dissolved carbon
dioxide produced by decay of organic matter (Ruttner-
Kolisko, 1956; Neel, 1948). But in most instances where
dissolved carbon dioxide levels are reported in the litera—
ture of psammolittoral investigations, the amount of €02
decreases with distance upshore rather than increases.

A possibility for the increase in pH with distance
shoreward is the fact that the amount of water generally
decreases with increased distance from shoreline. Russell
(1950) has pointed out that for any given soil there is
a tendency for the pH to increase as the soil becomes
drier (see Russell pp. 98-103 for an explanation of the
diffuse double layer of cations about soil particles and
its influence upon pH of soil solution with decrease in
soil water volume). Thus, the drier the soil, the more
acidic the soil solution.

Another point to be considered is the balance be-
tween oxidation and reduction of soil solutes with respect
to dissolved oxygen. It has been previously noted that
the psammolittoral zone is frequently characterized by a
"black layer" of sulfides beneath the surface. As the

psammolittoral substrate becomes drier, the oxygen content

 

 

 

325M: ». .
E‘ ‘

 

408
apparently rises (see later). According to Russell
(lgg. git.), as soil sulfides are oxidized to sulfates
in the presence of oxygen, the pH may increase to as much
as 4.0.

Pennak (1939) in noting the difficulty involved
in procuring interstitial water for pH determinations at
distances more than 100 cm. from the waterline, pointed
out the need for a method of determining pH at points
beyond 100 cm. from the waterline. So far as can be
determined in a more or less exhaustive review of the
literature of psammolittoral investigations, there have
been no published reports of pH determinations upon
quantities of the substrate itself rather than inter-
stitial water drawn up from the substrate. Table 10
presents data resulting frow monthly determinations of
pH in the substrate at each 25 cm. interval along the
Reach A1 transect during the 12-month sampling period
(see pages 43, 71, and 73 for methods involved in sub-
strate pH determinations).

The pH of intervals along the transect ranged
from 8.6 to 7.0. In all months but February and march
the pH increased, i.e. became more acidic, along the
transect with distance landward from shoreline. From
November through April, pH range along the transect ordin—
arily did not exceed a value of 0.4 pH units. Ranges in
other months were considerably greater, becoming one or

more pH units in June, July, September, and October. In

 

 

 

 

 

409

February and March, the pH became slightly more alkaline
at the most landward portion of the transect than at the
intervals closer to the lake. It was in January through
March that lake pH was closest to neutrality.

As stated on page 75, comparison of Figure 7 with
Table 10 reveals that monthly shifts in pH of lake water
tended to be reflected by similar shifts at any given
interval along the transect. The closer the transect
interval to the lake, the more nearly did the shift of
pH of that interval compare in magnitude to that of the
lake. All intervals exhibited a rapid shift toward
alkaline conditions following warming conditions that
induced complete substrate thaw in late March.

In comparing results of pH determinations at
each interval alang the Reach A1 beach transect with
data reported in the voluminous literature of pH - algae
relationships, those references concerned with pH and the
algae of freshwater and marine environments have been
arbitrarily excluded. Inasmuch as water could be drawn
up from the sand within 100 cm. proximity to any psammon
sampling point only on the April through June collection
dates, there seems to be little point in relating psammon
algae to pH relationships reported for aquatic environ-
ments.

Lund (1962) in reviewing the literature of studies
of the ecology of soil algae with respect to pH has con-

cluded that most soil algae are found over a fairly wide

 

 

 

41D
pH range. Nearly neutral or slightly alkaline soils have
the most species but not necessarily the largest crops.
Lund states that as a generalization most Cyanophyta pre-
fer neutral or alkaline soils; most Chlorophyta exhibit
maximum growth at neutral pH; and blue-green algae and
diatoms are usually rare or absent in acid soils. He
concludes that numerous errors in taxonomic identifica-
tion and incorrect culture techniques make it difficult
to generalize in reference to pH limits tolerated in
nature.

Cameron (1964) reported that arid soils with a
pH of less than 6.8 have restricted blue-green algal
floras. Shields and Durrell (1964) also found that most
soil algae tolerate a wide range in pH, but that pH does
have a decided effect upon the nature of the dominant
species in any soil association. Although pH in itself
may not be a directly critical factor in soil algal
ecology, chemical processes in the substrate dependent
upon pH may be affected. Cyanophytes and diatoms, they
reported, are favored by alkaline soils. Lune (1945),
John (1942), and peterson (1935) all have reported that
diatoms are poorly represented in acid soils.

There apparently have been no published reports
in the literature of psammolittoral investigations in
which efforts have been made to relate algal spatial and
temporal relationships to substrate pH. No obvious rela-

tionships were apparent when pH monthly fluctuations or

 

 

 

we‘re-*3?! “T , ,

411
variations along the Reach A1 beach transect were com-
pared with algal distribution patterns observed during
the 12—month sampling period. True, in each month there
was a tendency for substrate pH to become more alkaline
with diminishing distance between sampling interval and
waterline. And reference to tables 36 and 38 indicates
that in general more species of blue-green algae and
diatoms were found at the more ”alkaline" intervals than
at those with relatively greater acidity. Comparison of
figures 28, 30, 32, and 34, however, does not indicate
any quantitative preference by blue-green algae and diatoms
as groups for any particular interval sampled. The range
of pH along the transect in any given month, too, seems
inconsequential in view of the statements in the litera-
ture that most soil algae -- or aquatic algae for that
matter —~ tolerate relatively wide ranges in pH.

In the absence of any data to the contrary, it
may be assumed that pH had little if any effect upon
quantitative or qualitative expression of the algal flora
of the Reach A1 beach transect during the 12—month study.

Alkalinity determinations made upon samples taken
at 25 cm. intervals each month along the Reach A1 beach
transect indicated that on any given collection date
alkalinity tended to decrease markedly with distance from
shoreline, the greatest differences occurring in the
summer months (see pg. 44 for methods utilized in deter-

mination of substrate alkalinity, Table 11 for results

 

412
of determinations given as bicarbonate alkalinity ex-
pressed as ppm. CaCOs, and pages 76 and 78 for detailed
observations relative to alkalinity data). Because of
the large number of unmeasured variables which produce
alkalinity, perhaps the only valid conclusion that can
be drawn from the data of substrate alkalinity determina-
tions is that organisms present in the interstitial water
of the upshore portions of the transect were more subject
to pH variations than those in the more lakeward portions.
Since alkalinity is ordinarily produced by solutions of
salts of weak acids, the higher alkalinity at the inter-
vals nearest the waterline in all likelihood acted as a
buffer to rapid and pronounced pH changes.
Eflgflistry of lake and interstitial

water: dissolved gases and
dissolved solids

 

Stangenberg (1934) characterized the psammolittoral
as an extremely eutrophic aquatic habitat as a result of
chemical analyses of interstitial water drawn up from the
hygropsammon zone of several Polish freshwater beaches.
Comparing results of determinations of pH, dissolved
oxygen, carbon dioxide, inorganic and total phosphorous,
ammonia and nitrate nitrogen, iron, chlorides, silicates,
sulfates, oxygen demand, and total organic and inorganic
dissolved solids in interstitial water and water from the
adjacent lake, Stangenberg found values for dissolved

solids to be far greater in interstitial water than in

 

 

 

413
lake water. Values for total dissolved solids in inter-
stitial water he found to be as much as 194 per cent as
great as values determined in the lake.

Pennak (1939, 1940, 1950) in reviewing the pub—
lished reports of psammolittoral investigations noted
the chemistry of interstitial water to differ greatly
from that of the adjacent water basin. Although in marine
beaches, he reported, the concentration of dissolved
solids in the beach does not differ appreciably from the
amount in the adjacent sea, in freshwater beaches inter-
stitial water contains on an average 50 per cent more
dissolved materials than are present in the adjacent lake
or stream.

It should be noted that generalizations in the
literature concerning the chemistry of interstitial water
are in almost all instances based upon determinations
made upon water drawn up from the substrate with an
aspirator (Pennak, 1939, 194a; Heel, 1948; "fiszniowski,
1934a, 1935, 1936c; ieber, 1963; Ruttner-Kolisko, 1956;
Schwoerbel, 1961). In almost all instances the water
could not be drawn up from any location more than 100 cm.
from waterline. Consequently, little chemical informa-
tion is available from any zone of the psammolittoral
but that of the hygropsammon. Pennak (1939) and Ruttner-
Kolisko (lgg. git.) have both commented on the need for

better methods of sampling interstitial water, particularly

 

 

 

 

Tessa , A

414
the small amount of water present between one and three
meters upshore from the waterline.

The assumption apparently has been made by many
investigators of the psammolittoral zone that the inter-
stitial water withdrawn from the beach has come from the
level penetrated by the tip of the aspirator probe.

Kohn (1928) has noted that it is often assumed when a
pipette is used to remove suspensions or solutions from
a given depth below the surface of a liquid that the
pipette removes a thin layer of liquid at the required
depth. It appears this assumption has been made in most
studies of interstitial water of the psammolittoral zone.
But Kohn reports that actually a sphere of liquid is
removed the volume of which equals that of the pipette,
the center of the sphere being at the end of the pipette.
Pennak (1940), who inserted the collecting probe four

cm. into the beach, stated that data resulting from
chemical determinations upon the interstitial water col—
lected represented average chemical conditions between
surface and a depth of eight centimeters. Neel (1948)
stated that chemical features of water at depth of four
cm., which was the depth of probe tip insertion, undoubted—
ly differed from those of water contained in the upper—
most layors of sand where most organisms occur. 5991
concluded that although chemical analysis nf the inter“
stitial water so collected did not furnish data relative

to the surface stratum of most abundant life, it did offer

 

 

 

 

 

 

 

 

415
information concerning processes present in the beach
and relationships with lake water.

In an effort to determine the source of inter—
stitial water drawn up from the hygropsammoo zone of the
Reach A1 beach transect, a number of investigations were
made utilizing uranine as a chemical tracer (see pages
105-109 for a description of methods employed and detail-
ed observation of the results). It was concluded from
these investigations that the interstitial water with-
drawn from the hygropsammon zone of the Roach A1 beach
transect was not from the surface layers of sand, but
rather it apparently is derived from depths below the
terminus of the aspirator probe. Consequently, the
chemical data resulting from determinations upon the

interstitial water withdrawn from the beach are most

 

likely not those of the upper photic layer but instead

 

are illuatrativo of conditions in the abiotic zone be-
neath. It was further concluded that interstitial hater ,
chemistry reported in the literature reflects conditions ‘
in the ”raw material” water of the abiotic zone which is
drawn upon by organisms in the photic zone. As has been
noted previously, there is no conclusive evidence that
interstitial water is derived from the adjacent water
basin or that chemical conditiOns within the interstitial
water may influence those within the adjacent water basin.

A review of reports upon chemical analyses of

psammolittoral zone interstitial water reveals a number

 

 

 

of apparent principles with respect to quantities of dis-
solved gases and solids. Almost without exception, it
has been reported that the amount oF dissolved oxygen in
interstitial water is less than in the water 0? the ad-
jacent basin (Heel, 1948; Stangenberg, 1934; fliszniewski,
1934a; Ruttner-Kolisko, 1956; Pennak, 1939, 1040, 1950;
fieber, 1963). Host investigators have attributed this

to rapid utilization of oxygen in decomposition of inter-
stitial matter. Pennak (1930) has noted that rarely can

a water sample be taken From a Freshwater beach at 100 cm.
From waterline that shows as much as one ppn. xygen.
Pennak believed many members of the psammon Fauna are
Faculativc anaerobes inasmuch as they can be Frequently
t:

.4 v

collected From oxygen rreo otr

Several investigators have reported pronounce”

U)
U
}..1
C
(I)
1

horizontal gradients in the amount of oxygen dis~
in interstitial water. Pennak (1Qéfl) reported that
average dissolved oxygen determinations in 13 beache
gave values of 5.? oer. at waterline, 1.5 pom. at 39 or
distance From waterline, and 0.4 mph. at 1WD cm. with
mean lake oxygen content of 9.4 ape
Ruttner—Kolisko (1??‘) has noted the principle
that dissolved carbon dioxide values in interstitial mote:
increase with distance upshore From waterline. Penna?
(1939) reported that mean dissolved carbon ”iexide value:

in interstitial water from 13 Freshwater beaches rnngnd

 

 

 

 

 

 

417
From 2.0 to 9.5 ppm. at waterline but From 4.5 to 38.0
ppm. at collection sites 100 cm. From waterline.

Although Heel (1943) Found values For bound carbon
dioxide to be variable in Freshwater beaches, he reported
that during his investigations interstitial water always
contained more bicarbonate than did lake water. He
attributed the relatively high bicarbonate values in
interstitial water to action 0F Free carbon dioxide in
solution upon deposits oF CaCUz in the sand. Weber (1953)
has also reported increasing amounts oF dissolved bicar-
bonates in interstitial water with distance landward From
waterline. Pennak (193;) has reported the interstitial
water oFten contains EU per cent more bound carbon dioxide
than does water oF the adjacent lake.

Tables 12 and 13 present results 0F chemical
analyses conducted upon interstitial water and lake water
during the 12-nonth Roach A1 peach transect study (so:
also Figures 8 through 2a and Dogs: 7o throunh 105 For
detailed observations relative to determinations oF hi—
carbonate alkalinity, carbonate alkalinity, calcium
hardness, carbonate and non—carbonate hardness, total
hardness, dissolved oxygen reported as ppm. and as psr
cent saturation, dissolved carbon dioxide, dissolved
volatile and non—volatile solids, nitrat: and nitrite
nitrogen, erthophosphates, total iron, sulFates, and

silicates).

 

 

‘1, Ma‘s-n; s

419

Values For dissolved carbon dioxide in lakewater
ranged From 18.0 to 0.0 ppm. (see Figure 8). In general
dissolved carbon dioxide values tended to be highest in
winter months and lowest in summer. Dissolved carbon
dioxide content in interstitial water ranged From 133.0
(?) to 2.0 ppm. As indicated on Figure 3, dissolved car-
bon dioxide content was appreciably higher in interstitial
than lake water on all collection dates but August 5. Ho
relationships appeared between lake and interstitial water
with reference to relative amounts oF dissolved carbon
dioxide reported monthly. During the seven—month period
when interstitial water could be withdrawn From the beach,
mean dissolved carbon dioxide value in the beach was 18.2
and in the lake 1.9 ppm.

Eigration oF the site For collection of inter—
stitial water makes it difficult to determine whether
monthly variations exhibited in interstitial water data
were nttributablc to seasonal variation or change in col—
lection site (see page 41).

Values For dissolved oxygen as ppm. in lakewetar
ranged from 13.6 to é.3 ppm. In general dissolved oxygen
in lakewater reported as ppm. was highest during winter
months and lowest during the summer. in interstitial
water, dissolved oxygen content in ppm. ranged From 7.2
to 3.0 ppm. The amount of oxygen in interstit'
was considerably less than that in the lake on any given

collection date (see Figure 9). There was evidence 0?

 

 

 

 

direct relationship between the amounts of dissolved oxygen
in lake and interstitial water from April through October.
Kean value of dissolved oxygen in ppm. during these months
when interstitial water could be withdrawn From the eeach

in interstitial water was 1.2 and in lake water 8.1.

In lake water dissolved oxygen content repertec
as per cent saturation ranged From 103.0 to 39.0. No
seasonal trends were apparent, maxima and minima Fluctuat-
ing irregularly throughout the 12-aonth study. Per cent

dissolved oxygen saturation in interstitial water ranged

From 31.0 to 0.0. Again no seasonal trends were apparcn..

LJ
H
C
U
I

An apparent direct relationship did occur, howevv

tween relative oxygen saturation in lake and interstitial

water (see Figure 10). Noon oxygen content as per cent
saturation during those months interstitial Petr“ coulfi
be withdrawn From the beach was 88.2 in 3: ch 31 and 11.?

in interstitial water.
Total dissolved solids reported as wg./l. in 1.bn

water during the 12—nonth study ranged From 117 .

133.0. Seasonal variation was evident, total dissrlvxf
solids decreasing markedly following Formation of ice
cover in December and increasing rapidly Following dis-
appearance of ice cover in the spring (Fig. 11). Values
For non-volatile solids in lake water during the same
12-month period ranged From 231.6 to 43.5 mg./l. A
narked seasonal periodicity was apparent. fiaximum values

:3.

For non—volatile dissolved solids occurred in the Fall and

 

 

 

 

420
early winter and minimum values occurred in late winter
and early spring (Fig. 11). Only in January, September,
and October did non-volatile percentage oF total dissolved
solids depart markedly From a mean value oF about 45 per

cent. In January, values reached 59.2 per cent and in

9
l

[.4

September and October non-volatile percentage 0 tota
dissolved solids dropped to 32.0 and 19.8 per cent respec-
tively (Fig. 13).

In interstitial water, total dissolved solids
ranged From 324.4 to 1194.0 ng./l. (Fig. 12). Only in
September and October did the amount oF total dissolved
solids in lake water exceed the amount in interstitial
water. Values For non-volatile dissolved solids in inter—
stitial mater ranged From 302.4 to 120.3 mg./l. Percent—
age values oF non-volatile dissolved solids with respect
to total dissolved solids were relatively somewhat less
then those in the lake during the same collection periods,
the value Fluctuating semewhat irregularly around a mean
0F 40 per cent. There were no clear monthly correlations
oF total dissolved solids between lake and interstitial
waters and none For nonovolatile solids (Fig. 13). Durino
the seven-month period when interstitial mater could he
withdrawn From the beach, the mean value For non—volatile
dissolved solids in lake water was only 57.3 per cent thnt
in interstitial meter.

Total hardness in lake water ranged From a high

OF 158.0 to 2 low oF 60.r ppm. Evidence oF seasonal

 

 

 

 

 

 

421

variation was indicated in the early spring increase oF
total hardness which Followed the winter low and preceded
that oF early summer (Fig. 14). with reFerence to cation
contributors to total hardness, calcium hardness ranged
From 90.0 to 34.0 ppm. Figure 14 shows similar seasonal
variation in calcium content to that oF total hardness.
Only in December and June did the calcium percentage oF
cation hardness components comprise less than 50 per cent
oF total hardness (Table 14). Since Ferrous iron content
in lake water samples throughout the study was negligible
in amount, it would appear magnesium ions were signiFicant
contributors to total hardness oF the waters oF leach A1.

to

Carbonate hardness in lake water ranged From 136.0
24.0 ppm. Figure 14 indicates somewhat parallel trends

in seasonal variations For carbonate hardness and those

 

oF calcium and total hardness.

In interstitial water, monthly values For total
hardness ranged From 333.0 to 92.0 ppm. (see Figure 10).
Calcium hardness during the same period ranged From 24C.u
to 03.0 ppm. ReFerence to Figure 15 reveals closely
parallel Fluctuations in calcium content and those oF
total and calcium hardness. There was relatively little
similarity to comparable hardness Fluctuations in lake
water (see also Table 14). Magnesium ions were apparent—
ly oF considerable importance in the cation component oF
total hardness oF interstitial water. Values For carbon-

ate hardness in interstitial water ranged From 338.0 to

 

 

 

 

422

98.0 ppm. No correlation oF interstitial water variations
in carbonate hardness was apparent with those oF the water
0F Reach A1.

During those months in which interstitial water
could be drawn up From the beach, mean total hardness in
the water oF Reach A1 was 111.4 ppm. In interstitial
water mean total hardness was 209.1. In the same period,
mean calcium hardness in the lake was 62.9 ppm., but in
interstitial water it was 122.0 ppm. Mean values For
carbonate and non-carbonate hardness during the same
period were 86.3 and 32.9 ppm. respectively in lake water.
In interstitial, comparable values were 193.1 and 16.0
ppm. respectively.

Alkalinity in both interstitial water and that oF
Reach A1 throughout the study was that contributed by
carbonates and bicerbonates, no hydroxide alkalinity being
present at any collection date. Total alkalinity in the
water oF Reach A1 ranged From 136.0 ppm. to 24.0 ppm.
During the 12—month collection period, alkalinity in water
oF Reach A1 was essentially that contributed by bicar-
bonate ions (Fig. 16). Only in Way to July collections
did carbonate ions contribute appreciably to total alka-
linity. During the seven—month period when interstitial
water could be drawn up From the beach transect, mean
total alkalinity, bicarbonate alkalinity, and bicarbonate
alkalinity in the water oF Reach A1 were in ppm. respec-

tively 86.0, 68.6, and 17.7.

 

 

 

 

£23

Total alkalinity oF interstitial water ranged
From 470.0 to 108.0 ppm. with a seven-month mean oF 210.0
ppm. Carbonate alkalinity was observed only in the April
6 interstitial water collection and then only 3.0 ppm. in
magnitude. During the same collection period, mean value
For bicarbonate determinations in ppm. was 209.6. No
correlations in Fluctuations in alkalinity data were ap-
parent between waters oF Reach A1 and beach interstitial
water (see Figures 16 and 17).

Total Ferrous and Ferric iron in water collected
From Reach A1 during the 12-month study ranged From 0.140
to 0.001 ppm. mean content in lake water during the seven-
month period when samples were also being collected From
the beach was 0.014 ppm. Figure 18 indicates a pre—
cipitous decrease in the amount 0F total dissolved or
suspended iron throughout the winter until the disap-
pearance oF ice cover prior to the April collection. A
spring pulse oF total iron content was Followed by a
secondary summer low and partial Fall recovery. A some-
what similar pattern oF monthly Fluctuations in iron con-
tent was observed in interstitial water (Fig. 18). Values
0F total iron content in interstitial water ranged From
0.260 to 0.001 ppm. Mean total iron content in inter-
stitial water was 0.070 ppm.

Dissolved sulFates reported as ppm. in lake water
ranged From 32.0 to 15.0. In interstitial water, dis~

solved sulFates reported as ppm. ranged From 45.0 to 3.0.

 

 

 

 

 

424
No seasonal trends in sulFate content were observed in
either interstitial or lake water (Fig. 19). Mean sul-
Fate value in interstitial water was 25.3 ppm. Lake water
samples taken during the seven-month period oF interstitial
water sampling yielded a mean sulFate content oF 24.4 ppm.
The abundance oF dissolved silica was irregular
in lake water during the 12-month sampling period. Values
ranged From 3.10 to 0.25 ppm. (Fig. 20). In April to
October collections, silica values in interstitial water
ranged From 13.10 to 2.00 ppm. with a seven-month mean oF
5.63 ppm. In the same seven-month period, the mean silica
value oF lake water was 1.60 ppm. Figure 20 indicates
a somewhat related pattern oF monthly Fluctuations in
silica content in interstitial and lake water.
Orthophosphate content reported as ppm. ranged
From 1.05 to 9.42 during the 12 months oF analysis oF
water From Reach A1. A range oF 1.00 to 0.13 ppm. was
reported in the April-October interstitial water collec-
tions with a seven—month mean nF 9.46 ppm. The mean lake
orthophosphate content during the same seven—month period
was 0.67 ppm., and as noted previously (pg. 100) this
mean was Far in excess oF the 0.903 ppm. mean value For
soluble phosphorous reported in a study oF 479 Uisconsin
lakes by Juday and Birge (1931). A somewhat similar pat-
tern oF monthly Fluctuations in phosphorous content was
exhibited in interstitial water and that 0F Reach A1 (see

Figure 21).

 

 

   

 

425

 

Values For nitrate nitrOQen in interstitial water
ranged From 9.226 to 0.032 ppm. with a seven-month mean oF

0.109 ppm. A seven-month mean 0F 0.025 ppm. was reported

For nitrite nitrogen in interstitial water. The range in
nitrite nitrogen during this period was From 0.000 to 0.007

ppm. Figure 23 indicates relative monthly Fluctuations

0F nitrate and nitrite content in interstitial water were

quite similar. To a somewhat lesser extent, relative

monthly Fluctuations 0F nitrate and nitrite nitrogen con—

tent in water 0F Reach A1 were also similar (Fig. 22).

Comparison oF Figures 22 and 23 indicates similar patterns

0F monthly nitrogen content Fluctuations in lake and

interstitial water. The 12~month range oF nitrate nitrogen

n Reach A was 0.115 to 0.020 o3m. and that oF nitrite
1 l l

1.1.

nitrogen was 0.015 to 0.000 ppm. The mean values For

nitrate and nitrite nitrogen in Rea h A1 during the

tions were also being

seven-month period when determine

tade oF interstitial water chemistry were respectively

9.062 and 0.004 ppm.

mean values For total and bicarbonate alkalinity,
calcium hardness, carbonate hardness, total hardness,
dissolved carbon dioxide, nitrate and nitrite nitrogen,
total iron, sulFates, and silicates were appreciably high—

er than those reported From Reach A1. 0nly mean values

For carbonate alkalinity, non—carbonate hardness, dissolved

and percent saturation, and orthophosphate

\ .
oxygen in pom.

were higher in the waters oF Reach A1 than in interstitial

 

426
water. Similar patterns in monthly Fluctuations in abun-
dance oF silicates, orthophosphates, total iron, and
nitrate and nitrite nitrogen were observed in interstitial
water and water 0F Reach A1.

Davis (1954) has stated that the vast accumula~
tion 0F data collected From pond and lake studies has sug-
gested the abandonment oF attempts to explain seasonal
plankton Fluctuations on the basis oF any one controlling
Factor. Nelch (1952), too, has concluded that it is pro-
bably impossible to use any single chemical or physical
Factor to explain plankton seasonal periodicity, the
Fluctuations probably being the result oF interaction oF
the Factors involved in various combinations and in-
tensities. In this, Tyron and Jackson (1952) concur.
Pennak (1950) stated in a review oF numerous investiga-
tions oF the psammolittoral zone that sand population
variations with respect to physical or chemical Factors
appear to be as insoluble as sporadic plankton varia—
tions in small ponds.

Inasmuch as the physicochemical data presented
in this study are the result 0F, at most, 12 monthly
collections, it seems Fruitless to attempt to make ex—
tensive correlations between these physicochemical Factors
and qualitative and/or quantitative Fluctuations in the
phytoplankton oF the waters oF Reach A1 (see, however,

pages 128 through 134 For comparisons oF Fluctuations in

phytoplankton abundance and chemical Factors determined

 

 

 

 

 

42?
upon water of Reach A1 during the 12-month study period).
There were few apparent correlations observable in phyto-
plankton qualitative and/or quantitative Fluctuations
with respect to chemical conditions in the water of
Reach A1. There were, however, apparent correlations in
phytoplankton quantitative abundance and relative amounts
of silicates, orthophosphates, and nitrite and nitrate
nitrogen. Inverse relationships were observable between
phytoplankton abundance and both orthophosphate and ni-
trogen abundance. In both instances, quantities of
orthophosphate and nitrogen tended to be lowest during
those periods when phytoplankton abundance was greatest.
Prescott (1962) has noted that phytoplankton maxima are.
typically concomitant with minima of phosphorous and
nitrogen content. It has been concluded that this rela-
tionship indicates the rapid utilization of these nutrients
as plant mass increases.

Silicates, on the other hand, in quantity tended
to be at minimum at periods of low phytoplankton abun-
dance and at maxima during phytoplankton pulses. Rela-
tionships between silica content and phytoplankton abun-
dance are not as clearly defined as those For phosphorous
and nitrogen. The literature of algal-silicon relation-
ships reviewed by Round (1965), however, suggests that
chrysophyte maxima are usually associated with peaks in
Silica content within the Freshwater environment. Silica

depletion, 0.5 mg./l. or less, usually results in rapid

 

 

 

 

, bill.

 

 

 

428
decrease in abundance of chrysophytes, particularly
diatoms.

It has been demonstrated that the chemistry of
beach interstitial water collected with an aspirator in
the hygropsammon zone is not necessarily that of the
photic zone inhabited by the psammon flora. Consequent-
ly, no efforts were made to correlate fluctuations in
qualitative and/or quantitative abundance of the algae
of the Reach A1 beach transect with concomitant fluctua-
tions in the chemistry of the interstitial water. It
can only be hoped that data presented in this disserta-
tion can in time be included in the mass of information
needed to make meaningful generalizations with regard to

the ecology of the psammon algae.

Lake phytoplankton

Inasmuch as the study of the phytoplankton of
Reach A1 was undertaken as an adjunct investigation pri-
marily to identify its relationship to the algae of the
adjacent beach transect, it is not the intent here to
discuss in length the fluctuations in qualitative and
quantitative phytoplankton abundance (for detailed data
and observations relative to qualitative and quantita-
tive sampling of the phytoplankton, see tables 15 through
18, figures 25 and 26, and pages 110 to 156). The litera—
ture contains reports of far more extensive and intensive

studies of the ecology of lake and/or pond phytoplankton

 

 

 

429
than the 12-month study upon Reach A1 (for example,
Chandler, 1940a, 1940b, 1942; Birge and Juday, 1922;
Davis, 1954; Griffith, 1955; Prescott, 1962; Spencer,
1950; Round, 1965). Aside from the rather high phosphorus
content in the water of Reach A1 during the 12-month
study, the somewhat lesser contribution of diatoms to
total quantitative counts, and the quantitative domin-
ance of the plankton flora of Dinobryon spp. in the
February—May and September-October periods, the overall
character of Reach A1 did not differ markedly from des-
criptions of similar lakes reported upon in the litera-
ture.

During the 12-months of qualitative and quanti-.
tative sampling of the phytoplankton of Reach A1, 187
species of algae were identified. Of these, 42 were
assigned to the Cyanophyta, 51 to the Chlorophyta, 7 to
the Pyrrhophyta, and 87 to the Chrysophyta (see Table
36). Qualitative collections were dominated by members
of the Chrysophyta in all months but January and June.

In January, members of the Cyanophyta were qualitatively
dominant, and in June members of the Chlorophyta dominat-
ed the qualitative plankton flora.

Comparison of Table 10 (lake water chemical data)
with the lake chemical classification criteria presented
by Prescott (1962) suggests that Reach A1 might best be
described as a hard water drainage lake. Such lakes are

described by Prescott as stream fed and provided with an

 

 

43D

outlet at least part of the year. In general they are
characterized by pH ranges of 7.2 to 9.4 and typically
contain more than 14 ppm. carbonates. These lakes,
according to Prescott, are further characterized by a
predominantly cyanophyte-diatom flora and frequently
harbor luxuriant growths of Ehgra. Prescott reports
that if green algae do become abundant in the "typical"
hard water lake, they are most abundant in the summer
months.

Although the qualitative structure of the Reach
A1 phytoplankton has been previously discussed (pages
110, 124-127, 293, 302, 314—315), it may here be briefly
noted that from November through April, chrysophytes
(particularly the Pennales) and cyanophytes collectively
contributed 78 per cent or more of the total species
monthly. In February, 98 per cent of the phytoplankton
species in Reach A1 were chrysophytes or cyanophytes.
For the remainder of the 12-month collection period,
chrysophytes and cyanophytes collectively contributed at
most 69 per cent and at the least 58 per cent of the
total phytoplankton species present in monthly samples.

In general, greatest species diversity in the
phytoplankton was exhibited in the fall, spring, and
summer, and least species diversity was exhibited in
those few months when the lake was subjected to uninter~

rupted ice cover.

 

 

   

431

Quantitatively, Reach A1 was dominated by
cyanophytes and chrysophytes in all months but June. A
clear seasonal progression in quantitative dominance of
lake phytoplankton appeared during the study. Members
of the Cyanophyta dominated quantitative samples from
November through January and in July and August. Members
of the Chrysophyta numerically dominated quantitative
samples in February through May and again in September
and October. The February-May and September-October
dominance was attributable essentially to large numbers
of Dinobryon spp., Q. divergens in September, for in-
stance, contributing 74.9 per cent of the total phyto-
plankton count.

The phytoplankton of Reach A1 was essentially
quantitatively dominated by members of the Chroococcales,
Hormogonales, and particularly by the Chrysomonadales.
Although not appearing in the data, the gigantic biomass

of Chara established for most of the year upon the sandy

 

bottom of Reach A1 assuredly influenced the qualitative
and quantitative expression of the phytoplankton.

welch (1951) has noted that in temperate lakes
the total annual production often takes the form of a
bimodal curve presenting two maxima, one in the spring
and one in the autumn, and two minima, one during the
summer and the other during the winter. He further states
that in the northern United States the spring maxima

usually occur in April to May followed by summer minima

 

432
in August. The autumn maximum generally occurs in late
September to October, and the winter minimum from February
to March.

Reference to Figure 25 reveals that general pat-
terns of fluctuation in phytoplankton quantitative abun-
dance in Reach A1 closely approximated the plankton pro-
duction patterns outlined above by Nelch. welch, however,
has stated that the spring maximum commonly exceeds the
autumn maximum, and the winter minimum usually is some-
what "smaller" than the summer minimum. Figure 25 indi~
cates that although patterns of quantitative minima in
Reach A1 are similar to the generalizations made by welch,
the autumn September-October maximum was considerably
greater than to be expected.

Two possible explanations are apparent for the
Reach A1 anomaly of a greater autumn maximum than that
of April. One is that by definition, if not entirely in
fact (see pages 129-132), the phytoplankton data reported
in this study are of net plankton and by definition, if
not in fact, must be considered to exclude the contribu-
tions of nannoplankton. Welch (£22. £52.) has stated
that net plankton data alone exhibit only a mere hint
of spring maxima, the autumn maximum being the only one
indicated in well-defined form.

The second possibility for the larger autumn
maximum with respect to that of spring is the declining

water level in Reach A1 from June to October. This

 

 

 

433
concentration of water and its dissolved solutes in the
isolated basin of Reach A1 may possibly be the explana-
tion for the fact that values for hardness, alkalinity,
silicates, orthophosphates, and nitrates and nitrites
were as great or greater than those reported current dur-

ing the May maximum.

Psammon algae: Qualitative aspects

In contrast to the extensive literature relative
to the ecology of soil and aquatic algae, the literature
of investigations of the psammolittoral is minuscule and
that relative to the algal component of the psammon is
limited in the extreme.

Pearse _t al. in 1942 noted that although algae.
in the marine beach are only rarely numerous or concen-
trated enough to be visible to the unaided eye, they are
probably much more significant in the economy of the
beach than previously supposed. In generalizing with
reference to psammon algae they stated that the algal
psammonflora is composed predOminantly of members of the
Bacillariophyceae and, to a much lesser extent, the
Chlorophyta, Pyrrhophyta, and Cyanophyta. Most are found,
they stated, as individuals rather than as colonial forms
and often lack sheaths. Most are very small, i.e. usually
from three to ten microns in greatest dimension. The
green algae of the psammon, they generalized, are usually

non-colonial, coccoid forms, and the blue-green algae

 

 

 

 

 

 

434
are in general single, non-colonial forms lacking
sheaths.

It has been previously noted in this discussion
that Sassuchin at al. (1927) recognized four vertical
strata or zones in the eupsammon and hygropsammon, the
"green horizon” being the first subsurface stratum. The
character of this green horizon they described as being
determined by the large numbers of algae, particularly
diatoms, therein. These investigators listed approxi-
mately 95 algal taxa present in their collections from
the hygro- and eupsammon zones. Although they presented
their data in semi-quantitative form, the subjective
nature of their quantitative categories suggests the data
should be best considered qualitative. Of the taxa listed,
18 were assigned to the Cyanophyta, 27 to the
Bacillariophyceae, 10 were autotrophic flagellates, and
40 were classified as members of the Chlorophyta. Twenty-
one algal species listed by Sassuchin at al. were also
present in the substrate of the Reach A1 transect at one
or more collection periods during the 12-month study.

Pennak (1939) has generalized that the most
common algae in the psammolittoral are members of the
Cyanophyceae, Chlorophyceae, and Bacillariophyceae. In
the beaches of soft-water lakes, he stated, desmids are
abundant, and in hard-water lakes great concentrations
Of diatoms are common as far back as 250 cm. from the

edge of the water.

 

 

 

 

435

Ruttner-Kolisko (1953, 1954) has also noted that
algae, particularly diatoms, are important contributors
to the psammon. Although she included numerous tables
relative to quantitative abundance of psammon algae in
her reports of investigations of a number of Scandanavian
freshwater beaches, it is not entirely clear how she has
assembled her data. Thus, it is perhaps best to consider
it from a qualitative aspect in this discussion.

Ruttner-Kolisko's data relative to quantitative
aspects of psammon algal abundance along a number of
transects investigated are grouped into applicable cate-
gories of diatOms, flagellates, blue—green algae, and
green algae. Although her tables indicate the obviously
dominant position of diatoms in quantitative counts,

there are apparently no consistent patterns in distri-

 

bution in hydro-, hygro-, and eupsammon zones for any

of the four algal classification categories listed in
her tables. Unfortunately, she has included in her re—
ports the scientific names of but five taxa, those which
apparently were most abundant in collections from the

transects investigated. These include Tabellaria

 

fenestrata, Navicula mutica, Ceratoneis arcus, meridion

 

circulars, and Diatoma hiemale.

Janika (1954), in a brief report relative to the
study of a lotic psammolittoral environment, noted that
of BO algal species present in the psammon the great

majority were diatoms. Also present, but rare, were

 

 

436
flagellates, conjugates, and cyanophytes. The species
Pinnularia borealis and Navicula gothlandica she noted
to be especially common.

Neel (1948) has published what is to date apparent-
ly the most extensive presentation of data relative to
the algal component of the psammon flora in the litera-
ture of psammolittoral investigations (see page 396 for
an evaluation of quantitative data presented by Neel).
Neel lists 14 species assigned to the Chroococcales,
eight assigned to the Hormogonales, one species placed
in the Heterococcales, five representatives of the
Centrales, 115 species of Pennales, three members of the
Volvocales, one species assigned to the Oedogoniales,

13 species of Chlorococcales, 16 members of the
Zygnematales, and three species of dinoflagellates.
These species were present in the substrate of at least
one of six transects studied.

Neel (lgg. git.) concluded that diatoms out-
numbered all other groups of psammon organisms, and be-
cause of their abundance were a major component of the
psammon of the lake studied. Next in abundance, Neel
stated, were cyanophytes, particularly representatives
of the Chroococcales. Members of the Hormogonales he
found present but too few in number to warrant their in—
clusion in tables. Species belonging to the genera

Chroococcus, Microcystis, Aphanocapsa, Gomphosgheria,

Coelosphaerium, and flerismogedia he found to be common

 

   

 

437
in both the lake and psammolittoral. Green algae, he
stated, were far less common in the psammon than
cyanophytes or chrysophytes.

It should be noted that of the 173 species of
algae reported by Neel as present in the psammolittoral
of six transects and of the 179 algal species found in
the beach of the Reach A1 transect during the 12~month
study, only 30 species were shared in common in the two
investigations. Two species were members of the
Chroococcales, six of the Chlorococcales, one of the
Zygnematales, and the remaining 21 of the SO species
shared in common were members of the Pennales. Also
noteworthy is the fact that 26 of the 30 psammon species
shared in common were in the Reach A1 study of minor
quantitative importance. Only the species Navicula
radiosa var. tenella, fl. gupula var. ca itata, Cymbella
microcephala, and Rhogalodia ibba, shared in common in
both studies, were of relative numerical importance in
the Reach A1 transect.

The qualitative structure of the Reach A1 beach
transect psammon algae community during the 12-month
study has been discussed in considerable detail on pages
262-301 (see also pages 149-160, 171-185, 199-213, and
229-242 for data and observations relative to qualitative
psammon samples taken each month at the 300, 200, 100,
and 25 cm. transect intervals respectively). Further

discussion and interpretation of the qualitative data at

 

 

438

this point is perhaps an exercise in redundancy. At the
risk of introducing serious inaccuracies in an attempt to
generalize upon a relatively large mass of highly variable
data by other than statistical means, certain generali-
zations may be made with respect to characterization of
the Reach A1 beach transect on a qualitative basis.

Qualitatively, and on a 12-month overall basis,
the psammon algal flora of the Reach A1 beach transect
was essentially one comprised of members of the
Hormogonales and Pennales. At each of the transect in-
tervals, species assigned to these two orders contri-
buted 57 per cent or more of the total species count
recorded at that interVal during the 12-month study. The
dominance of the qualitative flora by cyanophytes and
chrysophytes is made even more prOnounced if it is noted
that species assigned to the divisions Cyanophyta and

Chrysophyta together contributed from 69 to 84 per cent

 

of the total species recorded at the separate beach in-
tervals during the entire 12-month study period, an aver-
age contribution of 78 per cent of the species present I
in the beach transect as a whole.

Considered on the basis of their contribution to
the 179 algal species found present in the beach transect
during the 12-month Reach A1 transect study, members of
the Cyanophyta comprised 34 per cent and members of the l
Chrysophyta comprised 40 per cent, a total of 76 per J

cent of the total species list. The Hormogonales, with

   

 

raw-way, A, a, 3

PR.

439
25 per cent, and the Pennales, with 35 per cent, together

contributed 60 per cent of the total species list con-

structed from samples taken at the 25, 100, ZOO, and 300

cm. intervals of the Reach A1 beach transect during the

12-month study.
Members of the Chlorophyta comprised but 23 per
cent of the 179 algal species reported from the Reach A1

beach transect. The green algal orders Chlorococcales

and Zygnematales together contributed 19 per cent of
the total species list, members of the Chlorococcales
alone comprising 15 per cent of the 12—month list.
Examination of tables 20, 24, 28, 32, and 38
does not indicate any conspicuous exception on a month
by month basis to the generalization that the qualita-
tive psammolittoral algal flora of the Reach A1 beach

transect was essentially one dominated by filamentous

blue—green algae and diatoms. with respect to the quali-

tative algal flora at each of the four transect inter-
vals, each also was dominated throughout the 12-month

sampling period by members of the Hormogonales and

Pennales. It should be noted that members of the

Chlorophyta in any one month at any one beach interval
never exceeded 24 per cent of the total species present

at that interval that month. As a generalization, with-

in the green-algal component of the qualitative algal

flora at each transect interval sampled, members of

the Chlorococcales tended to be more common than members

 

 

 

 

440
of the Zygnematales except at the 25 cm. interval. 3e—
yond the 25 cm. interval, coccoid green algae were quali~
tatively more abundant than desmids.

If the criterion of presence throughout the 12—
month sampling period at each of the four beach transect
intervals be considered, then the qualitatively character-
istic algae of the Reach A1 beach transect were Cymbella

microce hala, Navicula decussis, H. heufleri var.

 

legtocephala, fl. radiosa var. tenella, and Rhopalodia
q'bba. Also more or less ubiquitous throughout the beach
during the 12-month sampling period in that they were
absent at most two months from at most two beach inter-
vals were §ygyzdl-_ggcus

huqi;~la, and Eymbella a

...-.—

 

worthy that of the eight species which might be con—
sidered qualitatively characteristic of the Reach A1
beach transect, six are diatoms and not one is a member
of the Hormogonales. Also of note is the fact that at
least four of the eight (éyggchococcus aeruginosus,
Chlorococcqm humicola, Nayicula heufleri var. lsotoceohglg,
and Cymbella microgeghala)are described in the literature

as being commonly found in soil (Deskikachary, 1951;

Lund, 1945, 1946).

Psammon algae: quantitative aspects

 

Probably one of the greatest problems encountered

in the study of the algae of the psammolittoral environment

 

 

 

 

441
is that of obtaining an accurate measure of the quanti-
tative abundance of the algae per unit volume or unit
weight of the substrate. Although the psammolittoral
zone has been referred to frequently in the literature as
a unique, highly eutrophic aquatic environment, the
psammolittoral differs from the aquatic environment in
one very major respect: the water has a great deal of
sand in it. Perhaps the thought has arisen in the mind
of every investigator of the psammolittoral zone that,
rather than the environment being considered aquatic with
a great deal of sand in it, could it not also be con—
sidered a sandy terrestrial environment with -- at times --
a great deal of water in it? Should not, perhaps, the
hydropsammon zone be considered a benthic environment
harboring an epipelic (motile) or epfimammic (attached)
algal community and the hygro- and eupsammon zones be
considered terrestrial or aerial epilithic environments?
For a description of an epknammic algal community, see
Round (1965b).

The fact that the psammolittoral environment has
been studied by taxonomically—oriented limnologists or
limnologically—oriented taxonomists to the almost com—
plete exclusion of biologists conversant with other
aspects of ecology may be partially responsible for the
almost absolute absence of any meaningful quantitative
data relative to the algal component of the psammon.

In order to obtain a measure of the quantitative abundance

 

 

  

 

442
of the algae within the psammolittoral environment it
becomes necessary to turn to techniques utilized by in—
vestigators of algae of the soil.

Considerable controversy exists in the literature
with respect to accuracy of direct counts of soil samples,
involving examination of the original sample or a diluted
aliquot thereof, in contrast to indirect counts, which
involve culturing prior to examination of the original
sample or a diluted, cultured aliquot (Forest, 1962;
Tchan and Hhitehouse, 1953; Round, 1965a; Lund, 1945).
Insofar as can be determined in a review of the methods
utilized in quantitative study of the algal component of
the psammon, all investigators apparently have used tech-
niques involving separation of the algae from the sand
through repeated stirring and decanting of a sand sample
of given volume or weight. None seemingly have made
direct counts upon portions of the substrate sand itself.
The decantation techniques must be considered "direct”
in that culturing is not ordinarily involved, counts
being made of organisms present in the wash water and
results extrapolated to the original volume or weight of
the sample.

Forest (£92. git.) has reviewed in length the
literature relative to quantitative analysis of the soil
algal community and has conducted a detailed study of
the relative accuracy of the direct examination technique

in comparison to various indirect culture techniques. In

 

 

 

 

443
the direct count technique utilized by Forest, the counts
involved averages from a total of three examinations inas-
much as it was discovered that a total of three readings
was sufficient to give similar results in a series of
determinations. Forest concluded that direct examination
misses a considerable portion of the flora and that what
actually is gained from direct examination is not the
number of algae per unit sample volume but rather an in—
dication of the relative abundance of the most abundant
species. Forest further concluded that the moist plate
culture method (essentially the addition of sterile water
to a Petri plate containing the soil sample which is then
exposed to light for two weeks prior to counting) gave

the most satisfactory results.

 

It should be noted, however, that his data from
moist plate cultures were reported as plate-frequency,
i.e. whether a given alga is present or absent On a given
plate. Since plate—frequency is a relative index, it
cannot be stated in numbers of algae per sample volume.
One cannot but wonder if the culturing of an initially
dry soil presumably containing spores, cysts, and other
survival units as well as vegetative cells presents an
accurate picture of the soil flora at the time of col-
lection.

Tchan and fihitehouse (£92. git.), on the other
hand, reported that culture techniques as opposed to

direct counts of soil algae ordinarily produced results

 

 

 

 

444
25 to 40 per cent less than direct counts. when culture
techniques did, rarely, give higher counts than did the
direct method, the difference was only on the order of
12 per cent or less. Tchan and whitehouse also reported
that when moisture content of dry soil is increased, the
number of soil algae per unit volume or weight of dry
soil may double within two hours or less. They found
five-fold variations in the number of algae per gram of
soil during 24-hour periods which they attributed to
fluctuations in water content through day evaporation
and night condensation.

If relatively minor variation in moisture content
within the soil may induce the rapid fluctuations in
algal abundance described by Tchan and whitehouse, it
would appear that any quantitative figures resulting
from the direct count techniques must be considered as
reflective of relative densities within the soil at the
time of collection rather than an absolute measure of
algal abundance.

Pennak (1950) has reported that algal populations
in the freshwater beach commonly reach concentrations of
10,000 to 50,000 cells per cc. of sand and may occasional-
ly reach concentrations as high as 325,000 cells per cc.
He states also that in quantitative counts of psammon
algae almost invariably 95 per cent or more are diatoms.

Alth0ugh Heel (1948) made few generalizations

with regard to quantitative distribution of algae in six

 

 

 

 

445

freshwater beach transects studied, his data show values
for total algal individuals per cc. of sand ranging from
zero to 325,900. In most instances, at any given inter-
val on any given transect, the vast majority of algal
units enumerated are diatoms. His tables indicate no
conspicuous general tendencies for any taxon to exhibit
pronounced "preference" for a particular position along
any given transect.

deber (1963) has included data relative to quan-
titative determinations of algal abundance in a report
of a study of the psammon fauna of a freshwater lake.
He lists 15 genera of green algae, seven of blue—green
algae, and one member of the Euglenophyta. Members of
the Chrysophyta were not enumerated to genus. weber
noted that counts of diatoms were extremely high when
the total number of frustules per cc. were considered,
but the counts became greatly reduced when only frustules
with chloroplasts were included (see Lund, 1945, for a
discussion of the problems involved in taxonomic and
ecologic study of soil diatoms). deber concluded that
generalizations in the literature relative to the abun-
dance of diatoms in respect to the other algal groups
are suspect inasmuch as previous investigators had not
indicated whether or not only living cells were included
in their counts.

Blum (1956) has concluded that shifts in the

”climax communities” of lotic algal communities may occur

 

 

446
in a matter of days or even hours. He states that the
length of time needed for such a change in the community
structure is not dependent upon the life spans of the
dominant algae but rather upon the length of time it
takes the new dominants to establish themselves among
their predecessors. It is possible, Blum states, that
the establishment of a complex algal community and the
rapid succession to a climax can be within a period of
time characterized by diurnal changes.

Thus, the investigator of psammolittoral algae
must recognize that any quantitative data gathered are
suspect from the outset. Neither direct nor indirect
counting methods are entirely satisfactory in determina-
tion of the absolute composition and magnitude of the
flora at the time of collection. The separation of the
algae from the sandy matrix undoubtedly leads to frag-
mentation and dispersal of multicellular forms, and un-
doubtedly some individuals are lost despite the utmost
care in separation of water and sand. It is frequently
difficult to determine if a diatom frustule or desmid
semi—cell was alive at the time of collection. And the
investigator apparently is dealing with a highly ephemeral
biota which may undergo major change in relative compo—
sition and total magnitude within a matter of hours.

The quantitative structure of the Reach A1 beach
transect algal community during the 12~mcnth study has

been discussed in considerable detail on pages 299-315

 

 

'3», ,4

; {4‘5“

 

 

447
(see also pages 159-171, 184—199, 212-229, and 244-262
for data and detailed relevant observations derived from
quantitative psammon samples taken each month at the 300,
200, 100, and 25 cm. transect intervals respectively).
Thus, only very broad generalizations are presented at
this point in the discussion.

It is even more difficult to generalize about the
quantitative aspects than it was to generalize upon the
qualitative characterization of the algal component of
the Reach A1 beach transect psammon. It is apparent,
however, that certain generalizations made in the litera-
ture with respect to psammon algae were not entirely
substantiated during the Reach A1 beach transect study.

The psammolittoral has been repeatedly character-
ized in the literature as dominated by diatoms. ‘

Four beach intervals were each sampled once a
menth for twelve months giving a total of 48 quantita—
tive sample units investigated during the Reach A1 beach
transect study. In only 16 instances did the number of
diatoms present in a sample unit exceed the number of
individuals assigned to the Hormogonales, and in only 13
instances did the number of individuals assigned to the
Pennales contribute more than 50 per cent of the total
algal individuals present per cc. of sand of that sample
unit. The maximum contribution of the Pennales to the
total count of algae present in quantitative samples at

any interVal during the 12—m0nth sampling period was

 

 

 

 

 

448

the 70.84 per cent recorded at the 25 cm. interval on
May 6.

0n the other hand, in 32 instances the number of
individual members of the Hormogonales exceeded the num-
ber of diatoms in a sample unit, and in 20 instances
members of the Hormogonales alone contributed more than
50 per cent of the algal individuals present in a sample
unit. In 44 of the 48 sample units, members of the
Hormogonales and Pennales together contributed more than
50 per cent of the total number of algal individuals
present. The four exceptions to the complete dominance
by filamentous blue—green algae and diatoms of the quan-
titative algal flora present in the sample units described
occurred in May, July, August, and September at the 300
cm. interval and may be attributed to the appearance of

relatively large numbers of Botrydiopsis arhiza,

Chlorococcum humicola, and Synefiococcus aeruginosus at

 

those times.

0nly at the 300 cm. interval in key and July
through September did coccoid forms (Chroococcales,
Chlorococcales, and Heterococcales) collectively exceed
50 per cent of the algal individuals present in a sample
unit. It should be noted, however, that in 23 of 48
sample units diatoms alone, or ceccoid algae alone, or
diatoms and coccoid algae together contributed at least
50 per cent of the total number of algal individuals pre—

sent.

 

 
  

 

iv“; »

449

The psammon Flora oF the Reach A1 transect as a
whole could be characterized at any month oF the year as
quantitatively basically dominated by Filamentous blue-
green algae and diatoms. It would appear, however, that
insuFFicient data have been presented in the literature
and in this report relative to the quantitative character-
ization oF the algae oF the psammolittoral to justiFy a
brOad generalization as to the quantitative nature oF
the algal component oF the psammon biota in any Fresh—
water beach.

It is evident that generalizations relative to
characterization 0F the quantitative structure of the
algal community oF the psammolittoral must include the
diFFerentials in expression oF the community exhibited
in the hygro- and eupsammon zones. It has been previous—
ly noted (pg. 315) that the Hormogonales exhibited a
clear quantitative dominance oF the algal community at
the 300 cm. interval except For the Hay-September period
when the Heterococcales and Pennales established a tran—
sitory dominance (see Table 22). And at the 200 cm.
interval, members oF the Hormogonales maintained a clear
quantitative dominance throughout the year (see Table 26).
In the sand at the 100 and 25 cm. beach transect inter-
vals, however, distinct seasonal Fluctuations in relative
dominance oF the algal community were apparent. At the
100 cm. interval, Filamentous blue-green algae dominated

quantitative samples From November to February and in

 

 

450

August. The diatoms were most abundant march to July and
September to October (see Table 30). Conversely, at the
25 cm. interval, diatoms were relatively most abundant
November to March, in June, and again in September and
October. Filamentous cyanophytes were at the 25 cm. in-
terval most abundant in April and May and in July and
August (Table 34).

Perhaps one 0F the best illustrations oF the
heterogeneity oF the quantitative character oF the algae
present in the Reach A1 beach transect is the Fact that
oF the 179 species 0F algae encountered during the 12—

month study, only six species -- Gleeothece rupestris,

 

fiyneehococcus aeruginosus, Chlorococcqm humicola,

Navicula decussis, N. heuFleri var. leptocephala and fl.

 

 

radiosa var. tenella -- were present at least six months

 

0F the twelve in quantitative samples taken at each in-
terval.

There is no evidence that the trends in dominance
exhibited by the algal taxa in the Reach A1 transect are
exhibited in other Freshwater beaches. weber (1953) in
a report relative to the ecology eF the micreFauna oF
several psammolittoral zone transects noted that the algae
as a group exhibited no consistent population Fluctua—
tions or patterns. Members oF the Cyanophyta, he stated,
were most abundant in October and may and at lowest num-
bars in April, June, July, and December. No samples were

taken during the January—march period. He Found the

 

 

 

 

 

 

 

 

451

Chlorophyta to be at lowest population density in December
and April and to be most abundant in May through June.
It is not entirely clear whether he is reFerring to all
the transects he investigated during a year-long study
inasmuch as his collections apparently were alternated
between transects in diFFerent months.

So Far as can be determined, only Baklanovskaya
(1963) has traced the monthly quantitative variation in
psammon algae through the course oF one entire year. His
data are not available For comparison, but he has noted
that the seasonal dynamics oF the algae within the psammo—
littoral zone he investigated reFlected the seasonal
changes in the phytoplankton eF the adjacent reservoir.
It has been previously noted, however, that in the Reach
A1 beach transect investigation there was a more or less
inverse relationship in quantitative algal abundance
evidenced between the adjacent lake and each beach tran—
sect interval. The Further From the lake, the more pro-
nounced this inverse relationship became (see pages 303-
305 and 369-370 For a summary description oF the character
eF Fluctuations in quantitative algal abundance in the
beach substrate, and see also pages 402-406 For a possible
explanation For the pattern oF this Fluctuation).
Obviously, much more work must be done beFore generali-
zations may be proposed For the seasonal Fluctuations in

algal abundance in the psammolittoral zone.

 

 

 

452

Indices oF diversity oF lake
and beach transect intervals

In recognition 0F the Fact oF the somewhat sub-
jective nature oF even the most careFully gathered quan—
titative data relevant to the algal component oF the
Reach A1 beach transect psammon, indices oF diversity
were utilized in an attempt to identiFy evidence eF
physical or chemical Factor relationship to the algal
community (see pages 315 to 345 For discussion 0F the
utilization oF the index 0F diversity in community
ecology and in the Reach A1 beach transect study). The
literature does not indicate that heretoFore use oF
index oF diversity has been made in psammolittoral
studies.

The ratio between the number oF species and the
number oF individuals present in a community or sample
thereeF is termed the index oF diversity eF that com-
munity. The usage oF index oF diversity in the Reach A1
beach transect study was based on the premise that strong
physicochemical limiting Factors or interspeciFic com—
petition reduces the diversity within a community. Severe
physical or chemical limiting Factors, according to Odum
(1959), result in a low diversity index as does a period
oF high productivity Followed by competition For space
and nutrients.

Through utilization oF indices oF diversity and

calculation oF standard correlation (r) values,

 

 

 

 

T ‘ T’T‘ 1 1i T ‘ Ti

453
relationships were identiFied between psammon algal com-
munity structure at each transect interval and the physical
Factors 0F water content in both per cent saturation and
wet gross weight basis, sand grain size uniFormity coeF-
Ficient, harmonic mean grain size diameters, and percent—
age velatile matter.

A positive relationship within 95 per cent con-
Fidence limits was identiFied between algal community
indices oF diversity at each transect interval and water
content 0F the substrate on wet gross weight basis (see
pages 340—342 and 404—405). Computed r values For har-
monic mean sand grain size diameters in relationship to
algal community indices oF diversity at each interval
and For sand grain size uniFormity coeFFicients in rela—
tionship to diversity indices at the same intervals also
closely approached, but did not reach, 95 per cent con-
Fidence limits (see pages 343, 362, and 363 For Further
discussion oF indices oF diversity in relationship to
these physical Factors).

Ancillary to these results, but perhaps Far more
signiFicant in Future investigations oF the psammo-
littoral, was the discovery that indices oF diversity For
the beach collection sites -- with the exception oF the
period oF Frozen substrate and subsequent disruptions
thereaFter because eF Fluctuating lake levels -- monthly
exhibited a distinctly decreasing gradation From the

waters oF Reach A1 to the 300 cm. beach transect interval.

 

 

 

 

 

 

454

With the exceptions noted, each collection site maintained
a clearly deFined characteristic index 0F diversity that
when plotted against time exhibited patterns 0F Fluctua-
tion essentially parallel to those 0F the other transect
intervals (again, see pages 319-325 For discussion oF this
phenomenon).

Several major conclusions can be drawn From the
material presented on pages 319 to 325. One is that
with respect to the algal Flora the primary limiting
Factor in the psammolittoral is apparently water content,
all other physicochemical limiting Factors seemingly be-
ing subordinate to water content. The second is that
utilization oF indices oF diversity may be a very use-
Ful process in study oF the ecology oF the psammolittoral

environment.

Community coeFFicients

Inasmuch as the index oF diversity does not lend
itselF to quantitative comparison and identiFication oF
communities as well as other comparison techniques, com—
munity coeFFic'ents were utilized in an attempt to
analyze the algal community structure oF the Reach A1
beach transect (see pages 325—3AD For discussion oF the
use oF coeFFicient oF community as a technique in quanti—
tative community ecology and in the Reach A1 beach tram-
sect study). So Far as is known, community coeFFicients

.. . ._ J. ,.1 t 1
have not been utilized in the study oF the psammolit ora~

 

 

 

,_ ,; emanates see, ,
this {we afimmenmentss Aditfiemal evadeseélifiaee -
the decreasing index eF diversity with iaeneaeeag dis-
tanee shoreward From waterline. And fiurther amidenee was
the Fact that, on the basis oF community coeFFicient
determinations, the algal community at the 25 cm. interval
was considerably more similar to those at the 100 and
200 cm. intervals than to that in Reach A1. The algal
community at each transect interval was more similar to
any other given transect interval on the basis oF com-
munity coeFFicients than any was to the community in
Reach A1.

Frequent reFerence has been made also in this

 

discussion to the Fact that many oF the most qualitatively
and quantitatively abundant species in the Reach A1 beach
have been characterized in the literature as soil algae.
It has also been pointed out that eight 0F 23 species
listed by Durrell (1962) as characteristic oF the severe
environmental conditions in Death Valley were commOn in

the Reach A1 beach transect. And Microcoleus vaginatus,

 

 

 

 

W’“

456
one oF the most quantitatively important algae in the
Reach A1 beach transect, has been cited by Durrell and
Shields (1961) as one 0F the most important algae in—
volved in the Formation oF algal crusts on the moist
surFace 0F soils 0F the arid southwestern United States.
0F interest too is the previously cited Fact that 0F 60
algal species listed by Cameron (1964) as characteristic
0F arid soils, almost one-third were also present in
samples taken From the Reach A1 beach transect.

IF two distinct communities are assumed, it is
anticipated that somewhere along the beach an ecotone
must exist (see pages 323-325 For discussion oF the
ecotone concept and results oF attempts to idsntiFy the
ecotone along the beach transect). There is evidence
that the algal community at the 25 cm. interval was some-
what transitional between lake and beach communities, but
there was no indication that the 25 cm. interval could
be considered as, or present in, an ecotone. No ecotone
could be identiFied at any collecting site along the beach
transect nor could evidence be presented that the entire
beach From the 308 cm. interval to the shoreline be con~
sidered transitional between two distinct communities,
aquatic and terrestrial.

The Fact that the algal community at each inter—
val on the basis 0F community coeFFicients and the number
oF species shared in common was more similar to that

interval lakeward than to that landward suggested that

 

 

 

 

 

 

 

457

there was no transition in evidence From an aquatic to a
terrestrial environment along the beach transect. It
should be noted that approximately 44 per cent oF the 261
species collected during the Reach A1 transect study were
Found both in the lake and in the beach. Also to be
considered was the Fact that decreasing indices oF di-
versity shoreward along the transect indicated an in-
creasingly severe environment with increasing distance
landward From waterline. And reFerence to Table 35, in
which a summary tabulation is presented For each algal
taxon with respect to its quantitative and qualitative
contributions at each sampling site in the lake and beach
transect, suggested that the algae at the most landward
intervals along the transect were basically an expres-
sion oF a "survivors" community oF more or less aquatic
Forms associated with species adapted to severe terres—
trial environmental conditions. That is to say, data
presented in Tables 35, 41, 42, and 43 were interpreted
to conclude that an increasingly severe environment
developing in a shoreward direction along the beach tran—
sect induced a gradual attenuation oF a distinct beach
algal community, a community essentially characterized
as qualitatively and quantitatively cyanophyte-diatom in
composition.

Listed among the most prominent members 0F the

beach algal community were Cloeothece rugestris,

 

Synechococcus aeruginosus, Aulosira laxa, Cylindrospermgm

 

 

 

 

 

   

 

 

minimum, Lyngbya aerugineo-caerulea, L. aestuarii, L.

 

hieronymusii, microcoleus chthonoplastes, E. vs inatus,
Nostoc paludosqm, Botrydiopsis arhiza, Chlorococcum

humicola, Phormidiufl angustissimqm, Cymbella microce hala,

 

Navicula decussis, fl. heuFleri var. leptggeghala, fl.
radiosa var. tenella, and Rhopalodia gigga.

Monthly Fluctuations in quantitative abundance
oF the algae in Reach A1 and along the beach transect
have been summarized and discussed in some length on
pages 301 to 315. It has been noted that total numbers
eF individual algae per cc. oF sand sample ranged From
75,955 at the 25 cm. interval on December 8 to 1,369 at
the 300 cm. interval on August 6. These Figures may be
somewhat meaningless in themselves, but their signiFi-
cance may be in that the relative density 0F the 75,955
algae per cc. oF sand at the 25 cm. interval was 114,562
times that oF the phytoplankton per cc. oF lakewater re—
ported in December 8 samples, and the 1,369 algae per
cc. oF sand at the 300 cm. interval were 233 times as
dense as the density oF phytoplankton per cc. 0F lake
water reported on August 6 (see Table 49).

An environment that is as physically hostile as
is apparently the psammolittoral zone with respect to
Fluctuations in Factors such as temperature and water
content and that can support such an amazingly dense

algal growth is surely worthy oF more attention by the

ecologist and phycologist.

CHAPTER VII
SUMMARY

This investigation involved a 12—month study oF
the ecology oF the psammon algae 0F the psammolittoral
zone 0F a Michigan hardwater lake. The investigation was
undertaken to ascertain the practicability oF adaptation
oF techniques oF soil science and limnology to study an
environment that is neither aquatic nor terrestrial, to
establish iF there is evidence oF zonation oF algal com-
munities in a sandy beach, to establish iF there is evi-
dence oF seasonal quantitative and qualitative variation
in algal communities within a sandy beach, to attempt
correlation oF beach algal zonation and seasonal varia-
tion with physicochemical conditions prevailing in the
habitat, and to compare the algal Flora oF the psammo-
littoral zone with that oF the adjacent lake.

During the 12~month study, the opportunity was
presented to examine the Fate oF algal populations in
the psammolittoral during periods when the substrate
was Frozen, to trace modiFication 0F physicochemical
conditions within the beach substrate as Falling lake
water levels produced a gradual migration oF the water-
saturated beach zone, and to note the eFFect oF ice

459

 

 

 

460
erosion upon the substrate and the algal component oF the
psammon therein.

The transect method 0F community investigation
was utilized in study oF the hygro- and eupsammon por-
tions oF the psammolittoral zone selected. Each collec-
tion date, quantitative and qualitative phytoplankton
samples were taken From the lake adjacent to the tran-
sect. SurFace lake water samples were also taken oFF-
shore From the transect For subsequent chemical examina-
tion. Interstitial water was withdrawn From the hygro-
psammon zone oF the beach transect From April through
October. The Following physicochemical determinations
were made upon lake and interstitial water samples:
temperature, pH, alkalinity, hardness, dissolved oxygen
and carbon dioxide, total and volatile dissolved solids,
nitrate and nitrite nitrogen, orthophosphate, total iron,
silica, and sulFates.

Each month, at each 25 cm. interval shoreward
FrOm the November-June waterline to a point 300 cm. in-
shore, quantitative and qualitative sand samples were
taken From the beach psammolittoral zone. Only algal
samples From the 25, 100, ZOO, and 300 cm. intervals were
included in this study. In addition, at each 25 cm.
interval along the beach transect, monthly determinations
were made oF temperature, pH, alkalinity, substrate per-
centage water content, ”black layer” position, and total

organic content. In July through September, location oF

 

 

 

:9'55EJM ‘, 7“, p

461

phreatic levels beneath the substrate was investigated.
At each 25 cm. interVal along the transect, determinations
were made 0F substrate harmonic mean sand grain diameter,
grain size distribution, eFFective grain size, grain uni-
Formity coeFFicient, porosity, and unit weight.

According to criteria utilized by the American
Bureau oF Soils, the substrate 0F the transect could be
characterized as Fine sand. Average harmonic mean grain
size oF the entire transect was 0.109 mm. Mean eFFective
grain size For the beach transect was 0.094 mm., and mean
uniFormity coeFFicient For the transect as a whole was
2.43. Mean porosity oF the intervals examined along the
transect was 46.8 per cent. An overall tendency was
evidenced For harmonic mean grain size, eFFective grain
size, and uniFormity coeFFicient to increase with in-
creasing distance From waterline. Porosity decreased
with distance From shoreline. Sand samples taken at
interVals closest to the normal shoreline showed a great-
er water retention capacity than those located Further
inland.

Positive Film exposures made along the transect
indicated eFFective light penetration into the sand to
be somewhat less than 0.5 cm. with no essential diFFerence
in light penetration in wet or dry sand.

Transect temperature determinations made during
the 12-month study indicated a strong tendency For tempera—

tures oF the most lakeward stations to approximate within

 

 

"W452

 

462

3°C. those oF the lake surFace. Temperatures oF the most
landward intervals tended to approximate, within 3°C.,
breast height air temperature. Temperature variation be—
tween extremes oF the transect tended to be lowest during
winter months and greatest in spring and Fall.

The black layer was poorly deFined ahd sporadic
in appearance during those months it was present. At no
time during the study did the black layer reach the depth
penetrated by the sampling cores.

An inverse relationship was apparent between the
12-month mean percentage water content and distance From
shoreline. Twelve-month mean percentage water content
on wet gross weight basis ranged From 3.0 per cent at
the 300 cm. interval to 24.6 per cent at the 25 cm. in-
terval. Maximum diFFerentials in percentage water con—
tent between shoreline and the 300 cm. interval occurred
on those collection periods during which the substrate
was Frozen. 0n the basis oF 12-month means, the sand oF
the 300 cm. interval contained only 12.2 per cent as much
water as did the 25 cm. interval.

An inverse relationship was apparent between 12-
month mean water retention at each interval, expressed
as percentage oF theoretical maximum water retention
capacity, and the distance oF sample interval From shore—
line. Fluctuations in substrate organic content apparent-

ly resulted in monthly irregularities in water content

 

 

463
at certain intervals which could not be explained on the
basis oF granulometric conditions alone.

It was concluded percentage water content at col-
lection intervals along the transect was directly related
to porosity and distance From the shoreline and/or phreatic
surFace up to a certain critical distance, and beyond the
critical distance it became secondarily related to the
amount oF organic matter present.

Determinations oF pH monthly at each 25 cm. in-
terval along the transect were made both directly with
pH meter probe and with standard soil pH determination
techniques. Closely similar results between the two pH
determination techniques were observed until sand per-
centage water content Fell below 10 per cent. In general,
interstitial water tended to be more acidic than that in
the adjacent lake. In the transect, pH tended to shiFt
toward more acidic values with increasing distance From
shoreline. Monthly shiFts in pH 0F lakewater tended to
be mirrored by similar shiFts in pH at any given inter-
val along the transect.

Alkalinity tended to decrease markedly with dis-
tance From shoreline. Because oF buFFering eFFects oF
iOns producing alkalinity, it was concluded organisms
present in interstitial water oF more landward portions
oF the transect were more subject to pH variations than

those in the more shoreward portion.

 

 

464

As a result oF Field experimentation, it was con-
cluded interstitial water withdrawn From the beach was
not the water within which the algae oF the psammolittoral
live. There was no evidence that interstitial water is
derived From the adjacent water basin or that chemical
conditions within the interstitial water may inFluence
those within the adjacent water basin.

In interstitial water, mean values For total and
bicarbonate alkalinity, calcium hardness, carbonate hard—
ness, total hardness, dissolved carbon dioxide, nitrate
and nitrite nitrogen, total iron, sulFates, and silicates
were appreciably higher than those in the adjacent lake.
Only mean values For carbonate alkalinity, non—carbonate
hardness, dissolved oxygen, and orthophosphate were higher
in the lake than in interstitial water. Similar patterns
in monthly Fluctuations in abundance oF silicates, ortho~
phosphates, total iron, and nitrate and nitrite nitrogen
were observed in interstitial water and lake water. No
eFForts were made to correlate Fluctuations in qualita-
tive and/or quantitative abundance oF the algae oF the
beach transect with concomitant Fluctuations in the
chemistry oF the interstitial water. Apparent relation—
ships did exist, however, between Fluctuations in quali-
tative and quantitative abundance oF phytoplankton and
those oF silica, orthophosphates, and nitrate and nitrite

nitrogen. Mean values For non—volatile dissolved solids

 

 

465
in lake water was only 57.3 per cent that in interstitial
water.

During the 12-month study, 187 species oF algae
were identiFied present in the lake adjacent to the beach
transect. Qualitative collections were dominated by
members oF the Chrysophyta in all months but January and
June. Greatest phytoplankton species diversity was ex-
hibited in Fall, spring, and summer and least diversity
during the December-Karch period when the lake was sub-
jected to ice cover. Quantitatively, the phytoplankton
was dominated by chrysophytes and cyanophytes in all
months but June. A clear seasonal progression in quane
titative dominance oF the phytoplankton appeared during
the study. Members oF the Chroococcales and Hormogonales
quantitatively dominated the phytoplankton From November
through January and in July and August. members oF the
Chrysophyta, particularly the Chrysomonadales, dominated
phytoplankton samples February through may and again in
September and October. The phytoplankton population
Fluctuation pattern was bimodal during the 12-month study.
Minima in numbers oF phytoplankters occurred in winter
and mid-summer, and maxima occurred in spring and early
Fall.

During the 12-month study, 179 species oF algae
were identiFied present in samples collected From the
psammolittoral zone transect. OF these, 74 species were

Found only in the beach, and 105 were present both in

 

 

 

 

 

the psammon and in the phytoplankton. Qualitatively,

the psammon algal Flora was essentially one comprised oF

members oF the Hormogonales and Pennales. At each tran-

sect interval, species assigned to these two orders con-

tributed 57 per cent or more 0F the total species count

recorded at that interval during the 12-month study. A

total 0F 76 per cent 0F the psammon algae species list
was contributed by members oF the Chrysophyta and
Cyanophyta. There was no conspicuous exception on a
month by month basis to the generalization that the quali-
tative algal Flora oF the beach transect was dominated

by Filamentous blue-green algae and diatoms. Each oF

the Four transect intervals was qualitatively dominated
throughout the 12-month sampling period by members oF

the Hormogonales and Pennales.

The direct count method was utilized in collec-
tion oF quantitative data relative to the algae oF the
beach transect. Data were reported both in numbers oF

individuals per cc. oF sand and as a relative abundance

index in compariSOn to the density oF phytoplankton per

cc. oF lake water on each collection date. The psammon

Flora was quantitatively dominated throughout the year

by Filamentous blue-green algae and diatoms. Throughout

the year, the Hormogonales dominated the algal Flora at
the 200 and 300 cm. intervals, except For the may-September
period when the Heterococcales and Pennales established

transitory dominance at the 300 cm. interval. At the

 

 

 

467
100 cm. interval, the Hormogonales dominated winter and
late summer samples, and the Pennales dominated spring
and early summer and late Fall. At the 25 cm. interval,
diatoms dominated winter and late Fall, Filamentous blue-
green algae dominated spring and summer collections.

The 12-month Fluctuation in total number oF algal
individuals in the beach was essentially bimodal in
character at all beach intervals. Total number oF algal
individuals at all beach intervals reached maxima during
the December-March period when the south—Facing beach
was solidly Frozen except during late aFternoon hours,
and secondary maxima occurred in the beach intervals at
some point during the Nay-July period. Algal population
minima in the psammolittoral zone occurred in the March-
April period aFter thaw and during a time oF Fluctuating
lake levels. Subsequent to the summer maximum, a general
decline in number 0F algal individuals occurred at all
intervals until the October termination oF the study. An
inverse relationship thus was evidenced in quantitative
algal abundance in the lake and at each beach transect
interval.

Indices oF diversity were calculated For the
algal c0mmunity at each beach interval each month oF the
12-month sampling period in order to identiFy evidence
oF physical or chemical Factor relationship to the algal
community. Relationships were evidenced between psammon

algal community structure at each transect interval and

468
the physical Factors oF substrate water content, sand
grain size uniFormity coeFFicients, harmonic mean grain
size diameters, and percentage volatile matter recorded
at each interval. A direct relationship within 95 per

cent conFidence limits was identiFied between algal com—

munity indices 0F diversity at each transect interval and

water content oF the substrate. Inverse relationships

between algal community indices oF diversity and harmonic
mean grain size diameters and between community indices
oF diversity and grain size uniFormity coeFFicients ap—
proached but did not reach 95 per cent conFidence limits.
Decreasing magnitudes oF indices 0F diversity with in-
creasing distance From shoreline indicated an increasing—
ly severe environment shoreward From waterline in the
psammolittoral zone.
Community coeFFicients were calculated For the
algal communities within the lake and each beach tran-
sect interval each month oF the 12-month study. A Funda-

mental diFFerence in community structure was indicated

between the lake and beach. It was considered signiFi—

cant that many oF the most quantitatively and qualitative-
ly abundant algal species in the beach have been character-
ized in the literature as soil algae and that a relatively
large number are characteristic oF arid soil environments.

No ecotone could be identiFied in the beach as a

whole or at any interval. It was concluded there was no

transition in evidence From an aquatic to a terrestrial

 

 

469
environment along the beach transect. Rather, it was
concluded that a beach algal community exhibited a quali-
tative and quantitative attenuation shoreward From water-

line in response to an increasingly severe environment.

 

 

 

 

 

 

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AND

 

 

 

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like mm
a Wm,n.

 

 

 

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Chroococcales .
Hormogonales .
Euglenales . .
Volvocales . .
Tetrasporales .
Chlorococcales
Oedogoniales .
Zygnematales .
Peridiniales .
Heter0coccales
Chrysomonadales
Chrysotricales
Centrales . . .

Pennales . . ,

489

PLATES

Plate I
Plate IV
Plate XI
Plate XVI
Plate XX

Plate
Plate
Plate
Plate
Plate
Plate
Plate
Plate
Plate
Plate
Plate
Plate
Plate

Plate

IV

XI

XI

XI

XV

XV
XVII
XVIII
XVIII
XVIII
XVIII
XIX
XXVI

   

 

Figure

Figure
Figure

Figure

Figure
Figure

Figure

498

Plate I

 

Agmenellum quadruplicatum (Meneghini)
Brébisson X988 '

Agmenellum thermale (KUetz.) D. and D. X900

Anacystis marginata Meneghini X988

Anacystis montana (Lightfoot) Droust and
Daily X988

Aphanocapsa elachista Nest and Meet X980

Aphanothece saxicola Naegeli X988

Chroococcus limneticus Lemmermann X450

 

 

491
PLATE. I .--

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@9 65 -" $2". . . E
i oer. 9"" .§§

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“WM" a
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Figure

Figure
Figure

Figure

Figure
Figure
Figure

Figure

Figure

Plate II

Chroococcus limneticus var. distans G.M.

Smith X458

Chroococcus minutus (Kfietz.) N59-

xgno \

Chroococcus turgidus (KUetZ.) Naegeli X458 ‘

Coccochloris peniocystis (Kfietzing) Drouet ‘

and Daily X988
Coelospherium dubium Grunow X458
Coelospherium Naegelianum Unger X
Coelospherium pallidum Lemmermann
Dactylococcopsis fascicularis Lemm
X988

DaCtYlococcopsis raphidioides Hans

458
X988

ermann

girg X900

 

 

493
PLATE II

 
 

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,, 8 s Q. ,9 .ee...e
see a ex ...... aaoew .
/ see e\\ ..,..eese_e.....

Illul\ —|Il._ ......... .

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9
n

 

 

 

 

 

 

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Figure

Figure

Figure

Figure

Figure

 

5.

6.

Plate III

Gloeothece rupestris (Lyngb.) Bornet X458

Gomphosphaeria aponina Kbetzing X900

Gomphosphaeria aponina var. gelatinosa
Prescott X458

Gomphosphaeria aponina var. multiplex
Nygaard X458

Gomphosphaeria lacustris Chodat X900

Microcystis elabens (Bréb.) Kfitz. X980

 

 

495

PLATE III

 

.lli

.I.
A. a

L
L
1.

V

 

 

 

 

 

 

 

Figure
Figure
Figure
Figure
Figure
Figure

Figure

496

Plate IV

  

Microcystis incerta Lemmermann X1888
Peloglea bacillifera Lauterborn X2888
Synechococcus aeruginosis Naegeli X100
Xenococcus Ksrneri Hansg. X1888
Anabaena affinis Lemmermann X1888
Anabaena sp. 1 X1888

Anabaena sp. 2 X1888

 

 

497
' PLATE IV

 

13'

135'

 

 

 

 

 

 

Figure
Figure
Figure
Figure
Figure

Figure

U N
O .

4.
5.
6.

Plate V
Anabaena sp. 3 X438
Anabaena sp. 4 X458
Aulosira laxa Kirchner X438
Calothrix membranacea Schmidle X438
Cylindrospermum minimum 8.3. West X458

Gloeotrichia sp. X438

 

 

 

r
/
t
/
[Ills 3.1.25 @ ..v .ro...¢.&....vr. 00......
8. 9% gee ......r. eta... ..
. 0

 

 

Figure

Figure
Figure
Figure
Figure
Figure
Figure
Figure

Figure

‘ Lyngbya

X2888
Lyngbya
Lyngbya
Lyngbya
Lyngbya
Lyngbya
Lyngbya
Lyngbya

Lyngbya

Plate VI

aerugineo-caerulea (KUetz.) Gomont

aestuarii (Mertens) Liebmann X458
Birgei G.M. Smith X438'
contorta Lemmermann X2888
epiphytica Hieronymus X438
hieronymusii Lemmermann X988
putealis Montagne X438
rivularianum Gomont X2888

versicolor (Martm.) Gomont X2888

 

 

581

 

PLATE VI

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

0's"
. .

,. . 17.83.17.

 

 

[W'WW “- '} ii; 4‘

 

 

Figure
Figure
Figure
Figure
Figure
Figure

Figure

'\I
0

Plate VII

Microcoleus acutissimus Gardner X988
Microcoleus chthonoplastes Thur. X988
Microcoleus vaginatus (Vauch.) Com. X900
Nodularia harveyana (Thw.) Thuret X988
Nostoc linkii var. arvense Rao, C.B. X450
Nostoc paludosum KUetzing X988

Nostoc sp. 1 X988

 

 

,
Lw'.-=aw , .

 

583

PLATE VII

 

 

 

 

 

 

 

1

Tlll

 

 

 

 

., ....w. e, ...,rur.
. Erasers},

Figure
Figure
Figure

Figure

Figure
Figure
Figure
Figure
Figure
Figure

Figure

Oscillatoria
Oscillatoria
Oscillatoria
Oscillatoria
X988
Oscillatoria
Oscillatoria
Oscillatoria

Oscillatoria

Phormidium africanum Lemmermann X2000
Phormidium ambiguum Gomont X1940

Phormidium angustissimum w. at 8.5. West

X2888

584

 

Plate VIII

Agardhii Gomont X1948
amoena (KUetz.) Gomont X988
formosa Bory X988

limosa (Roth) C.A. Agardh

okeni Ag. X458
sancta (Kfietz.) Gomont X458
splendida Greville X1978

tenuis C.A. Agardh X1888

 

. . -_ ._
l_-v-~wse~_.’u‘.m‘gm we,

 

 

 

 

 

 

 

 

 

 

 

 

585
PLATE VIII

 

 

 

 

 

 

 

 

 

 

 

 

 

 
 
 
 

 

Figure
Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

&

Plate IX

Phormidium anomala Rao, C. B. X1888

Phormidium jadinianum Gom. X988

Phormidium luridum (Kbetz.) Gomont
X2000

Phormidium microtomum Skuja X988

Phormidium minnesotense (Tilden)
Drouet X1888

Phormidium molle (Kfitz.) Grun. X450

Phormidium uncinatum (C.A. Ago)
Gomont X458

Plectonema notatum Schmidle X450

Plectonema puteale (Kirchn.) Hansg-
X458

Plectonema tenue Thuret X458

p0rphyrosiphon fuscus Gomont X988

 

 

 

    

587

 

     

 

LATE IX

p

 

 

 

 
 
 

,, but} 5151;..11.

 

 

 

 

 

Figure
Figure
Figure

Figure

Figure
Figure

Figure

&

 

Plate X

Schizothrix friesii (A9.) Gomont X988

Schizothrix fuscescens KUetzing X1888

Schizothrix lacustris A. Braun X988

Schizothrix rivularis (Molle) Drouet
X450

Spirulina subsalsa Oersted X988 ‘
|
Symploca Kieneri Drouet X988 ‘

T01ypothrix tenuis Kfietzing x450

 

 

509

 

 

 

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PLATE X

 

 

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Figure
Figure
Figure
Figure

Figure

Figure
Figure
Figure
Figure

Figure

Figure

Plate XI

Phacus sp. X988

Trachelomonas

Trachelomonas

_Ch1amydomonas

Chlamydomonas
X988
Chlamydomonas

Chlamydomonas

granulosa Playfair X988
Playfairii Deflandre X988
Frankii Pasch X908

polypyrenoideum Prescott

Snowii Printz X908

sp. .X1588

Gloeocystis gigas (KUetz.) Lagerheim X90U

Gloeocystis vesiculosa Naegeli X900

Ankistrodesmus Braunii (Naeg.) Brunnthaler

X458

Botryococcus Braunii Kfietzing X450.

 

 

firm... . -.w’}'M

 

PLATE X I

 

..."

 

 

Figure

Figure

Figure

Figure
Figure
Figure

Figure

Figure

Figure

1.
2.

3.

4

0

6.

7

Plate XII

Chlorella vulgaris Beyerinck X988

Chlorococcum humicola (N589.) Rabenhorst
X988

Franceia Droescheri (Lemm.) G.M. Smith
X458

Nephrocytium obesum Nest and Meet X458

Oocystis crassa wittrock X458

Oocystis elliptica w. west X458

Oocystis elliptica var. minor Nest and
west X1888

Oocystis submarine Lagerheim X988

Pediastrum Boryanum (Turp.) Meneghini X450

 

 

 

m
_

 

 

 

 

3

ll
PLATE XII

5

 

 

   

Figure

Figure

Figure

Figure

Figure

Figure

Plate XIII

Pediastrum Boryanum var. undulatum Mills
X458

Pediastrum duplex var. rugulosum
Raciborski X458

Pediastrum integrum Naegeli X988

Pediastrum integrum fa. glabra Raciborski
X458

Pediastrum tetras (Ehrenb.) Ralfs X450

Pediastrum tetras var. tetraodOn (Cords)

Rabenhorst X988

 

 

 

 

 

 

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

8.

516

Plate XIV

Planktosphaeria gelatinosa G.M. Smith X98.

Scenedesmus arcuatus var. platydisca
C.m. Smith X988

Scenedesmus bijuga (Turp.) Lagerheim X988

Scenedesmus incrassatulus var. mononae
G.M. Smith X988

Scenedesmus opoliensis var. contracts
Prescott X458

Scenedesmus quadricauda var. quadrispina
(Chodat) G.M. Smith X988

Scenedesmus quadricauda var. longiSPina
(Chod.) G.M. Smith x450

Scenedesmus quadricauda var. Mestii
G.M. Smith X988

Schroederia Judayi G.M. Smith X988

Tetraedron caudatum (Corda) HanSQiPQ x988

Tetraedron duospinum Ackley X980

Tetraedron minimum (A. Braun) HanSQiPQ
X900

Tetraedron muticum (A. Braun) Hansgirg

X908

  

 

 

 

 

517
PLATE XIV

, , level...

 

 

 

 

Figure

Figure

Figure
Figure
Figure
Figure

Figure

Figure

A
O

 

Plats XV

Trochiscia aspera (Reinsch) Hansgirg
X1080

Trochiscia granulata (Reinsch) Hansgirg
X988

Trochiscia sp. 1 X1888

Trochiscia sp. 2 X1888

Bulbochaetae sp. 1 X2888

Oedogonium rufescens mittrock X458

Oedogonium rufescens wittrock antheridia
X450

Oedogonium sp. 1 X458

 

 

 

 

 

 

Figure 1.

Figure 2.

Figure 3.

Figure 4.

Plate XVI

Euastrum hypochondrum fa. decoratum
Scott and Prescott X458

Staurastrum Manfeldtii Delp. X988

Staurastrum orbiculare var. extensum
Nordst. X1888

Staurastrum sp. 1 X988

 

 

 

 

 

1.?“er .,

 

 

 

Figure

Figure
Figure
Figure

Figure

Figure

Figure

Figure
Figure

Figure

Figure

Figure

Figure
Figure
Figure
Figure

Figure

2.
3.
4.
5.

12.

_\

3.
14.
15.
16.
17.

522

Plate XVII
Cosmarium angulare var. canadense
Irénee—Marie X225
Cosmarium angulosum Brébisson .X225
Cosmarium,81ytii Mills X225
Cosmarium Botrytis Meneghini X225
Cosmarium geometricum var. suecium
Borge X225
Cosmarium granatum DeBreb X225
Cosmarium impressulum fa. suborthogonum
(Racib.) w. at 5.5. West x450
Cosmarium pseudoretusum Ducellier X225
Cosmarium punctulatum Brébisson X225
Cosmarium reniforme var. elevatum
w. and w. X225
Cosmarium venustrum (Bréb.) Arch X500
Closterium abruptium var. africanum
Fritsch and Rich X188
Cosmarium vexatum West X588
Cosmarium sp. 1 X225
Euastrum insulare (Mittr.) Roy X225
Staurastrum sp. 2 X225
Micrasterias mahabuleshwarensis var.

ringens (Bai1.) Krieg X225

 

 

 

 

523
PLATE XVII

 

 

 

 

 

 

 

 

 

524

Plate XVIII
Figure 1. Dinobryon divergsns Imhof X225
Figure 2. Dinobryon sociale Ehrenberg X225
Figure 3. Dinobryon sociale var. americanum
(Brunn) Bachmann X225
Figure 4. Uroglenopsis americana (Calkins)
Lemmermann X458
Figure 5. Botrydiopsis arhiza Borzi X458
Figure 6. Chlorallanthus oblongus Pascher X450
Figure 7. Chlorobotrys simplex Pascher X225
Figure 8. Phaeothamnion Borzianum Pascher X225

Figure 9. Glenodinium pulvisculus (Ehrenb.) Stein

 

 

Figure 14. Peridinium sp. X225

X225

Figure 18. Hemidinium ochraceum Lebander X225 ‘

Figure 11. Peridinium cinctum (Mue11.) Ehrenberg X225

Figure 12. Peridinium inconspicuum Lemmermann X225 ‘

Figure 13. Peridinium willei Huitfeld-Kaas x225 ‘
T
|
1

 

 

PLATE XVIII

 

 

 

Figure
Figure
Figure
Figure
Figure
Figure

Figure

526

Plate XIX
Coscinodiscus radiatum Ehren. X588
Coscinodiscus sublineatus Grun. X1888
Cyclotella meneghiniana Kfietzing X1888
Cyclotella compta (Ehr.) Kbetzing
Cyclotella ocellata Pant. X1888
Melosira italics (Ehr.) Kfietzing X450

Melosira varians C.A. Ag. X450

 

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PLATE XIX

 

 
 
   

    
  
 
    

  

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Achnanthes
Achnanthes
Achnanthes
Achnanthes
Achnanthes
X1888
Achnanthes

X1080

Amphora veneta Kfitz X1888
Anomoeoneis vitrea Grun. X1888
Asterionella formosa Hass X758
Caloneis sp. 1 X1888

Cocconeis pedicularis Ehr. X1888

Cocconeis sp. 1 X1888

528

Plate XX
exigua Grun. X1888
flexella (K‘otz.) Brun. x1000
minutissima KUtz X1888
linearis (w. Sm.) Grun. x1000

linearis fa. curta H.L. Smith

lanceolata var. dubia Grun.

 

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530

Plate XXI
Cocconeis placentula var. lineata (Ehri)
V.H. X1888
Cymbella cistula var. maculata (Kbtz.)
Van Heurck X588
Cymbella affinis Kbtz. x1000
Cymbella turgida Gregory X1888
Cymbella microcephala Grun. X1888
Cymbella ventricosum Kfietzing X1000
Cymbella sp. 1 X1888
Eunotia pectinalis var. minor (KUstz.)
Rabenhorst X1888
Eunotia praerupta Ehr. X1888
Fragilaria brevistriata Grun. X1888

Fragilaria construens (Ehr.) Grun. X1880

 

 

 

 

   

 

:1: I
/ 22,

 

531
PLATE XXI

 

 

 

 

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure
Figure

Figure

Figure

Figure

12.

532

Plate XXII
Fragilaria crotonensis Kitten X588
Fragilaria pinnata var. lancettula
(Schum.) Hust. x1000
Frustulia rhomboides var. crassinervia
(Breb. ex W. Sm.) Ross X1888
Gomphonema angustatum var. producta
Grun. X1888
Gomphonema angustatum var. sarcophagus
(Gregory) Grun. X1888
Gomphonema olivaceum (Lyngbye) KUtZ X1000
GPmPhonema olivaceum var. vulgaris
Grun. X1888
Gomphonema parvulum (Kfitz.) Grunow X1888
Gomphonema sp. 1 X1888
Meridion circulars var. COHStFiCtum
(Ralfs) V.H. x1000
Navicula capitata Ehr. X1888

Navicula cryptocephala KUtz X1800

 

 

 

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Figure

Figure

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Figure

Figure

Figure

Figure

Figure

Figure

Figure

Plate XXIII

Navicula cryptocephala var. veneta

(KUtz.) Rabh.

Navicula cuspidata Kfitz x500
Navicula decussis Ostr. X1888

Navicula heufleri var. heufleri Grun.

X1000

Navicula heufleri var. leptocephala
(Breb. ex Grun.) Patr. x1000

Navicula lanceolata (A9.) KUtz x1000

Navicula minima
Navicula mutica
Grun. X1888
Navicula mutica
Navicula mutica
Grun. X1888
Navicula mutica
Navicula pupula
Hust X1888
Navicula pupula

Meyer X1888

a

534

X1808

Grun. X1888

var. Cohnii (Hilse)

 

KBtz x1000

var. undulata (Hilse)

var. tropica Hust X1888

var. mutica (Krasske)

var. capitata Skv. and

|
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535

PLATE XXIII

 

 

 

 

 

 

 

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Figure

Figure

Figure

Figure

Figure

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Figure

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Figure

Figure

Figure

Figure

3‘4.

18.

12.
13.
14.

15.

536

Plate XXIV

Navicula pupula var. pupula Kfitz X1888

Navicula pupula var. rectangularis
(Greg.) Grun. x1000

Navicula radiosa var. tenella (Breb.
ex KUtz.) Grun. x1000

Navicula radiosa KUtz x750

Navicula hungarica var. linearis
Ostrup X1888

Navicula secreta var. apiculata Patr-
X1088

Navicula tripunctata (O.F. Mull.)
Bory X1888

Navicula virudula var. rostellata
(KUtz.) Clere X1888

Navicula sp. 1 X1888

Navicula sp. 2 X1888

Navicula sp. 3 X1888

Navicula sp. 5 X1888

Navicula sp. 4 X758

 

 

 

 

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537

PLATE XXIV

-

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Figure
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Neidium affine

Neidium dubium (Ehr.) Cl.

Nitzschia
Nitzschia
Nitzschia
Nitzschia
Nitzschia
Nitzschia
Nitzschia
Nitzschia
Nitzschia
Nitzschia
Nitzschia

Nitzschia

538
Plate XXV
(Ehr.) Pfitz. x1000
x1000

acicularioides Hustedt X1888.
angustata var. acuta Grun. X758
clausii Hantzsch X1888
filiformis (w. Sm.) Hust x1000
frustulum (KUtz.) Grun. x750
kutzingiana Hilse X1888
linearis (A9.) Mm. Smith X588

palea (KGtz.) w. Smith x1000

romana Grun. X1888
subcapitellata Husteds X588
sp. 1 X1888

sp. 2 X588

 

 

 

 

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PLATE XXV

.g,.__,.,.__._..=,5.552%

 

 

 

 

 

 

 

 

Figure
Figure
Figure

Figure

Figure

Figure

Figure

Figure
Figure
Figure

Figure

Figure

Figure

9.

10.

Plate XXVI

Nitzschia sp. 3 X1888

Nitzschia sp. 4 X1888

Pinnularia sp. 1 X758

Pinnularia biceps Greg X1888

Pinnularia borealis var. rectangularia
Carlson X1888

Rhopalodia gibba (Ehr.) o. Mull. x500

Stauroneis phoenicenteron fa. gracilis
(Ehr.) Hust x500

Surirella angusta KUetzing X1888

Synedra famelica KUtZ. X1888

Synedra incisa Boyer X1888

Synedra rumpens var. familiaris (KUth)
Hust X1008

Synedra ulna (Nitz.) Ehr. X588

Tabellaria fenestrata (Lyngb.) Kbtz. X5DO

 

 

 

PLATE XXVI

       

assesses

 

 

 

 

 

 

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31

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