-------
TABLE 38. LAKE ERIE WESTERN PLUS CENTRAL BASIN
TOTAL PHOSPHORUS BUDGET (METRIC TONS)
MARCH 27-DECEMBER 14, 1975
Cruise Interval
Cruise
No.
1 -2
2 3
3 "4
4 »5
5* 6
1 »6
Days
51
33
49
28
65
226
In
+2614
+ 1531
+2016
+ 1318
+2640
10127
Out to
E.B.
- 531
- 245
- 297
- 206
- 781
-2060
Change
In
Water
Mass
-3150
+ 711
- 22O
+ 1351
+3325
+2017
Net
Settling
-5233
- 575
- 1939
-6050
Net
Regeneration
+ 239
+ 1458
Retalnec
o/
/o
79.69
84.00
85.27
84.37
70.51
79.66
In the
Water Mass
Cruise
No.
1
2
3
4
5
6
Metric
Tons
8439
5289
6000
5781
7132
10458
CO
-------
TABLE 39. LAKE ERIE CENTRAL BASIN
1975 PHOSPHORUS BUDGET, METRIC TONS
CRUISE 1-
In from
^
In from V.'.B.
SOP- 237
SRP- 121
PP = 852
TP -1210
Cruise 1
In Water:
SOP- 648
SRP» 588
PP. 5865
TP . 7101
Point Diffuse Total
Sources Sources In
Sp" 57
IP " H4 27 141
TC " 21°
, TP ' 277 131 408
Cruise 2
In Water:
SOP. 650
SRP- 770
PP - 3109
TP - 4529
Cruise l-» 2
SOP-
SRP-
2
182
PP - -2756
TP . -2572
Out to E.B.
t> ^t-
SOP. 99
SRP- eg
pp = 377
TP - t31
3659
In from W. B.
In from Point & Diffuse Sources
Out to E.B.
Accumulated in Water
Settled
Imbalance
PP
+852
+210
-374
-2756
-3659
-215
SOP
+237
+ 57
- 99
4 2
-
4193
SRP
+121
+ 141
- 58
+ 182
-
+ 22
SOP+ SRP
+358
+ 198
-157
+ 184
+215
TP
+ 1210
+ 408
- 531
-2572
-3659
0
159
-------
TABLE 40. LAKE ERIE CENTRAL BASIN
1975 PHOSPHORUS BUDGET, METRIC TONS
Cruise 2
In
From:
Point Diffuse Total
Sources Sources In
SOP.
.^n from W.B.
' *
SOP. 39
SRP. 82
PP. 384
TP. 505
SRP
PP " ^
v TP! lft,
EPI & MESO
Cruise 2
T U_A
In Water:
SOP. 430
SRP- 528 Cruise 2 >3
PP - 1932 SOP. 28
TP - 2890 SRP. .173
PP - 249
TP . 104
Hypo Volune |
Incorporation: Exchange:
SOP- 29 SOP. o
SRP- 32 SRP. 7
PP. 155 PP. 51
TP. 216 TP. 58
WPO
Cruise 2
In Water:
SOP. 220
SRP. 242
PP . 1176
TP . 1638 Cruise 2 »3
SOP. -29
SRP. .39
PP. 350
TP . 282
32
9 77
117
66 226
Cruise 3
In Water:
SOP- 453
SRP. 355
PP - 2181
TP- 2994
V
Settling
PP- 656
Cruise 3
In Water:
SOP- 191
SRP- 203
PP- 1526
TP . 1920
^Out to E.8.
*
SOP. 22
SRP. 60
PP- 163
TP. 24-5
P.
160
-------
TABLE 41 . LAKE ERIE CENTRAL BASIN
1975 PHOSPHORUS BUDGETS, METRIC TONS
CRUISE 2-3
EPILIMNION AND MESOLIMNION
In from W.B.
In from Point & Diffuse Sources
Loss or Gain of Volume
Exchange
Out to E.B.
Settled
Accumulated in Water
Imbalance
HYPOLIMNION
In from EPI
Loss or Gain of Volume
Exchange
Settled
Accumulated in Water
Imbalance
TOTAL
In from W.B.
In from Point & Diffuse Sources
Out to E.B.
Accumulated in Water
Settled
Imbalance
PP
+384
-1-117
+ 155
+ 51
-163
-656
+249
-360
+556
-155
- 51
- 99
+350
0
+384
+117
-163
+598
- 99
-360
SOP
+39
+32
+29
0
-22
-
+28
+50
_
-29
-
-
-29
0
+39
+32
-22
-
-
+49
SRP
4 82
+ 77
+ 32
+ 7
- 61
-173
+310
_
- 32
- 7
-
- 39
0
+ 82
+ 77
- 60
-212
-
4311
SOP&
SRP
+ 121
+109
+ 61
+ 7
- 83
-
-145
+360
- 61
- 7
-
- 68
0
+ 121
+ 109
- 82
-212
-
+360
TP
+505
+226
+216
+ 58
-246
-656
+ 104
G
+656
-216
- 58
_ 99
+282
0
+505
+ 226
-245
+387
_ 99
0
161
-------
TABLE 42. LAKE ERIE CENTRAL BASIN
1975 PHOSPHORUS BUDGET, METRIC TONS
Cruise 3
In
From:
Point Diffuse Total
Sources Sources In
In from W.B. ^
SOP. 63
SRP- 103
PP - 520
TP- 686
>
4
1
1
1
1
A
SOP-
SRP- 94
PP -
v TP 231
EPI & MESO
Cruise 3
in Water:
SOP. 458
SRP- 355 Cruise 3 > 4
PP - 2181 SOP- 51
TP . 2994 SRP. 317
PP - 341
TP- 709
f A 7T
Hypo Volume
Incorporation: Exchange:
SOP. 38 SOP- 0
SRP- 55 SRP- 23
PP - 240 PP . 68
TP - 333 TP . 91
A- JL
HYP3
Cruise 3
In Water:
SOP. 190
SRP. 209
PP . 1521
TP . 1920 Cruise 3 »4
SOP. -51
SRP. 64
PP _
rr--748
TP--736
I f v
I
1
1 PP
SRP. 62 ! SRP. 129 "840
1
A
4
15 10
14
64 29
Cruise
T_ L/ai-
in Wats
SOP-
SRP.
PP . 2
TP- 3
w
Settling
PP- 300
4
Cruise 4
In Water:
SOP- 139
SRP- 273
PP- 772
TP . 1184
|
1
4
I
I
1
|
1
i
0
9
6
5
4
r:
510
672
522
704
V
1
1
1
r
*- 62
r
Out to E.B.
SOP. 42
SRP. 83
PP - 172
TP -296
SRP- 191
162
-------
TABLE 43. LAKE ERIE CENTRAL BASIN
1975 PHOSPHORUS BUDGETS, METRIC TONS
CRUISE 3-4
EPILIMNION AND MESOLIMNION
In from W.B.
In from Point & Diffuse Sources
Loss or Gain of Volume
Exchange
Out to E.B.
Settled or Regenerated
Accumulated in Water
Imbalance
HYPOLIMNION
In from EPI
Loss or Gain of Volume
Exchange
Settled or Regenerated
Accumulated in Water
Imbalance
TOTAL
In from W.B.
In from Point & Diffuse Sources
Out to E.B.
Accumulated in Water
Settled or Regenerated
Imbalance
PP
+520
+146
+240
+ 68
-172 ^
-399~62
+341
-
+399
-240
- 68
-840
-749
-
+520
+ 146
-172
-407
-901
SOP
+63
+40
+38
-
-42
-
+ 51
+48
-
-38
-
-51
+ 13
+63
+40
-42
0
-
+61
SRP
+ 103
+109
+ 55
+ 23
- 83
+ 62*
+317
- 48
-
- 55
- 23
+ 129
+ 64
- 13
+ 103
+109
- 83
+381,
+ 129+62
- 61
SRP
+ 166
+ 149
+ 93
+ 23
-125
+62*
+368
-
-
- 93
- 23
+ 129
+ 13
+ 166
+ 149
-125
+381
J-CO*
+ 1 29+62
TP
+686
+295
+333
+ 91
-296
-399
+710
+399
-333
- 91
-711
-736
+686
+295
-297
- 26
-71 0
^Regenerated from and settled to unstratified areas.
163
-------
TABLE 44. LAKE ERIE CENTRAL BASIN
1975 PHOSPHORUS BUDGET, METRIC TONS
CRUISE 4> 5
In from Point Diffuse Total
CAO Sources Sources In
sc°. 39
*? 67 23 90
PP- £
^ TP . 15! 12j 274
In from W.8.
SOP. 30
SRP. 97
PP . 321
TP . 448
Cruise 4
In Water:
SOP. 648
SRP- 945
PP. 3295
TP - 4888
Cruise 4
SOP. .3
SRP. -133
PP . 1382
TP » 1246
1
Cruise c
In Water:
SOP. 645
SRP. 812
PP- 4677
TP «= 6134
-»5
Out to E.B.
b *-
SOP. 28
SR?= 24
PP- 153
TP . 205
p =
SR
729
In from W.B.
In from Point & Diffuse Sources
Out to E.B.
Accumulated in Water
Settled or Regenerated
Imbalance
+ 321
4 145
- 153
+1382
-1069
SOP
+ 30
-r 39
- 28
- 3
-f 44
+ 97
+ 90
- 24
- 133
+ 729
+ 1025
+ 127
+ 129
- 52
- t36
+ 729
+ 1069
TP
+ 448
+ 274
- 205
41246
+ 729
0
164
-------
TABLE 45. LAKE ERIE CENTRAL BASIN
1975 PHOSPHORUS BUDGET, METRIC TONS
CRUISE
n .rom
^
In fror. W.B.
SOP. 69
SRF. 283
PP. 801
TP. 1153
Cruise 5
Ir. Water:
SOP- 645
SRP. 812
PP-4677
TP - 6134
Point Diffuse Total
^j Sources Sources *nco
SRP- 127 22 149
PP- 195
TP- 312 85 397
t
Cruise 6
In Water:
SOP- 638
SRP- 2481
P° - 6504
TP- 9623
Cruise 5~»6
SOP.
SRP-
PP -
TP .
-7
l6&8
1827
3488
Out to E.B.
SOP- 77
SRP- 120
PP- 585
TP- 782
P = 2720
SR
In from W.B.
In from Point &. Diffuse Sources
Out to E.B.
Accumulated in Water
Settled orRegsnerated
Imbalance
PP
4-801
4195
-585
+1827
?
-1416
SOP
+69
+53
-77
- 7
+ 52
SRP
+283
+ 149
-120
4-1668
4-272O
+1364
SOP -|- SRP
+352
+202
-197
+1661
42720
+ 1416
TP
+1153
+ 397
- 782
+3488
+2720
0
165
-------
TABLE 46. LAKE ERIE CENTRAL BASIN
1975 PHOSPHORUS BUDGET, METRIC TONS
CRUISE
In from
In from W.3.
SOP. 69
SRP. 283
PP = 801
TP. 1153
Cruise «j
In Water:
SOP. 645
SRP. 812
PP - 4677
TP = 6134
Point Diffuse
,,r.0 Sources Sources
*
312 85
Total
In
53
149
397
Cruise £
In Water:
SOP. 638
SRP. 2481
PP - 6504
TP « 9623
Cruise
SOP. -7
SRP. 1668
PP . 1827
TP . 3488
PP - 914
SRP & SOP . 1806
Out to E.B.
SOP. 77
SRP. 120
PP. 585
TP = 782
In Prom W. B.
In from Point & Diffuse Sources
Out to E.B.
Accumulated in Water
Settledor Regenerated
In-, balance
PP
+ 801
4 195
- 585
+ 1827
+ 914
- 502
SOP
+69
+53
-77
- 7
+52
SRP
+ 283
+ 149
- 120
+1668
+ 1806
+ 450
SOP+ SRP
+ 352
+ 202
- 197
+ 1661
+1806
+ 502
TP
+ 1153
+ 397
- 782
+3488
+2720
0
166
-------
TABLE 47. LAKE ERIE CENTRAL BASIN
1975 PHOSPHORUS BUDGET, METRIC TONS
CRUISE
1 ->6
In from
_
SOP
SRP- 470
PP -
TP - 1204
Point Diffuse
Sources Sources
396
Total
In
221
566
813
1600
In fro-: W.B.
SCF= -*37
s=p= 685
pp 2878
Cruise I
In Water:
SOP. 648
SRP- 588
PP . 5864
TP - 7100
Cruise (,
In Water:
SOP- 638
SRP- 2481
PP - 6504
TP - 9623
Cruise 1 >6
SOP. -9
SRP- 1887
PP. 644
TP-2522
A
1
3674
Out to E.B.
SOP. 268
SRP- 344
PP - 1447
TP - 2059
1021
4695
In from W. B.
In from Point & Diffuse Sources
Out to E.B.
Accumulated in Water
Settled °r Regenerated
Imbalance
PP
+2878
f 813
-1447
+644
-4695
-3095
SOP
+437
+221
-268
- 9
+399
SRP
f 685
f 566
- 344
f1887
f3674
f2694
SOP+ SRP
+ 1122
+ 787
- 612
+ 1878
+3674
+3095
TP
+40OO
+ 1600
-2059
+2522
-1019
o
167
-------
The preliminary budget calculations for the 3 to 4 cruise interval
showed a net imbalance of +191 metric tons of SRP + SOP and an
imbalance of -191 metric tons of PP. Since the calculated excess of
191 metric tons of SRP + SOP was assumed to be regenerated from
the sediments, then the excess was thought to originate in the hypolim-
nion. The hypolimnion budget showed only 129 metric tons of SRP +
SOP regenerated, 62 metric tons less than the total excess. The epi-
limnion budget indicated a deficit of 62 metric tons of SRP + SOP;
consequently it was postulated that the 62 metric tons of SRP, or 32.5
percent of the total 191 metric tons regenerated, were transported from
the sediment directly into the epilimnion in unstratified areas. During
cruise 4 the hypolimnion covered an area of 9,6OO km2 in the central
basin, while the remaining 6,49O km2, or 40 percent, were not strati-
fied. The relationship between the quantity of SRP regenerated in the
unstratified portion of the central basin to the actual (unstratified) area
appeared to substantiate the theory that SRP is directly transported
from the sediments to the unstratified overlying water. The cruises
3 to 4 budget showed a net 191 metric tons of SRP + SOP regenerated
and converted to PP. Comparison of the budgets for the 2 to 3 and
3 to 4 cruise periods indicates that the conversion of SRP + SOP to
PP was greater during the 3 to 4 cruise interval. This increase in
biological activity was also substantiated by an increase of 4.6 juig/l
of uncorrected chlorophyll a during the 3 to 4 cruise interval.
The addition of 729 metric tons of SRP resulting from anoxic
and oxic regeneration in the hypolimnion was shown in the budget
between cruises 4 and 5. The imbalance of +1,069 metric tons of
SRP + SOP was converted to PP by biological assimilation. High bio-
logical activity was also confirmed by a 2.3 xjg/l increase in corrected
chlorophyll a between the cruises and represented the highest chloro-
phyll a concentration during the 1975 cruise season. It should be men-
tioned that the estimated figure, 1,O69 metric tons of SRP + SOP as-
similated, was high. If a fraction of the net regenerated phosphorus
was attributed to resuspended PP, the quantity of SRP converted to PP
would have been smaller. Nevertheless, the general trend of phos-
phorus conversions to various forms in the central basin would be very
similar even if more detailed data were available.
Table 45 shows the net phosphorus forms budget for cruises 5 to
6. The regeneration of 2,720 metric tons of SRP + SOP and the
calculated quantity of SRP + SOP converted to PP appeared to be ex-
cessive. The overestimation of these quantities was primarily attri-
buted to resuspension of PP, as was discussed as part of the 1975
total phosphorus budget. Table 46 shows the net phosphorus forms bud-
get for cruises 5 to 6, corrected for resuspended PP. The corrected
168
-------
budget for cruises 5 to 6 indicated a regeneration of 1,806 metric tons
of SRP with 502 metric tons of SRP + SOP converted to PP. This
estimate suggests there was a considerable decrease in biological activ-
ity during the 5 to 6 cruise interval, as compared to cruises 4 to 5.
A 3.1 pg/\ decrease in corrected chlorophyll a concentration was further
evidence of decreased biological activity.
The phosphorus forms budget for the entire 1975 season appears
as Table 47. The conversion of 3,095 metric tons of SRP + SOP to
PP was not corrected for resuspension of PP and, therefore, appears
higher than the actual conversion.
Figures 57 and 58 show the SRP + SOP> PP conversion rates
and sedimentation rates for the 1975 cruise season. Anoxic regenera-
tion of phosphorus and peak biological activity during the cruise 4 to 5
period is apparent from both curves.
CONCLUSIONS
The 1973 through 1975 measurement of total phosphorus were com-
pared with 1970 CCIW measurements (Table 48). The external phos-
phorus loadings to both the western and central basins were considerably
greater during 1970. The concentrations and quantities of TP in the
western basin during 1970 were not significantly greater and the quantity
and concentration in the central basin was only slightly greater. The
percent of TP retained in the western basin during 1970 was the greatest
of the three years examined. In 1974 and 1975, 59.2 and 53.1 percent,
respectively, of the western basin external TP load was retained in the
basin, as compared with the 77 percent retained in 1970. The percent
of the quantity of phosphorus retained in the central basin was greatest
in 1974, but not much different from 1970 or 1975. The quantity of TP
retained in the combined western and central basins was greatest in
1970 due to the large fraction of external loading retained in the western
basin. The estimated 197O total external loadings of P to the western
and central basins used by Burns (1976) were 23,580 and 5,220 metric
tons, respectively (Table 48). The updated U.S. Army Corps of Engi-
neers (1975) estimated loadings to the western and central basins for
1970 were 17,043 and 4,501 metric tons, respectively. The revised
estimates, particularly for the western basin, indicate that little change
In total loading has taken place from 197O to 1973. Based on the
revised loading values for the western basin, the percent of P re-
tained within the basin Is estimated to be 61 percent, which is similar
to the value for 1974 (see Overview section). The Canada Centre for
Inland Waters has not recalculated the 1970 phosphorus budget based on
the revised loading estimates (Noel Burns, personal communication).
169
-------
SOF - Soluble Organic Phosphorus
SRP - Soluble Reactive Phosphorus
PP - Paniculate Phosphorus
SOP + SRP -* PP TRANSFORMATION CALCULATED
FROM NET PHOSPHORUS BUDGET
40 r
>-
<
o
Of
10 -
_ _
APR | MAY | JUN | JUL | AUG | SEP | OCT NOV DEC
Figure 57. Lake Erie central basin - 1975,
phosphorus transformation.
Figure 58.
Regeneration and sedimentation rates of particulate
phosphorus in the central and western basins of
Lake Erie - 1975.
170
-------
TABLE 48. LAKE ERIE WESTERN AND CENTRAL BASIN 1970-1975
MAY THROUGH SEPTEMBER (5/1-9/30) MEAN TOTAL PHOSPHORUS
CONCENTRATION AND QUANTITIES, "ANOXIC" REGENERATION,
EXTERNAL LOADING, AND SINK
YEAR
1970
1973
1974
1975
WESTERN BASIN
Mean 5/1 - 9/30
TP
M-Tons
895*
-
840
980
TP
Cone.
JJg/1
38.3*
-
35.4
41.8
TP
External
Loading
M-Tons
per Year
23580***
17500
166OO
14900
TP
Retained
Percent
of Input
77
-
59.2
53.1
CENTRAL BASIN
Mean 5/1 - 9/30
TP
M-Tons
5500*
-
5250
5150
TP
Cone.
17.9*
-
16.6
17.0
Area of
Anoxlc
Hypollm-
nlon km
6600
11270
10250
400
TP
Anoxlc
Regener-
ation
M-Tons
1900
2570
1720
730
TP
External
Loading
M-Tons
per Year
5220**
4000**
42OO**
3850**
TP
Retained
Percent
of Input
65
-
67.4
63.3
w.B. -t- C.B.
TP
External
Loading
M-Tons
per Year
28800
21500
20900
18800
TP
Retained
Percent
of Input
87.7
-
83.8
79.7
* C.C.I.W. data recalculated with the volumes used by CLEAR for 1973, 1974 and 1975.
** Does not Include TP entering from western basin.
*** Does not Include TP from precipitation. The 1970 estimated phosphorus loading appears excessive.
-------
The area of anoxia was highly variable during the study period
(Table 49). In 1970 the area of anoxia was nearly half that which oc-
curred in 1973 and 1974, but the quantities of TP in the central basin
were not significantly different. This may be attributed to the extension
of the anoxic period into late September 1970, while in 1973 and 1974,
turnover occurred early in September. In 1973 anoxic regeneration
was overestimated, since the increase in quantity of TP also included
resuspended phosphorus from the sediments, attributed to stormy weath-
er following turnover in 1973.
The 1970 and 1974 data are most comparable on the basis of
sampling schedule. The quantity of phosphorus regenerated in 1970
was slightly greater than in 1974 despite the nearly twofold increase
of anoxic area in 1974. As previously mentioned, this was attributed
to the duration of the anoxic period.
The 1975 estimate of anoxic area and the quantity of phosphorus
regenerated associated with anoxia was underestimated. During the
late summer anoxic period, the hypolimnion was reoxygenated prior
to turnover; so the area and the phosphorus associated with anoxia were
difficult to estimate.
The 1975 central basin phosphorus budget shows that 8.1 percent
of total phosphorus loading (internal plus external) was attributed to
anoxic regeneration and 30.0 percent to oxic or physical regeneration
(resuspension). Similar conditions were observed in 1970. In 1973
and 1974 no measurements were taken during December, but similar
conditions probably existed in these years. If the 1975 western basin
and central basin phosphorus budget was extended over the whole year,
the percentage of total phosphorus loading (internal plus external) con-
tributed by anoxic regeneration would decrease considerably.
The following major conclusions were reached during the course
of this study:
1. The 1970, 1974 and 1975 nutrient monitoring surveys
do not show significant change in concentration and
quantity of phosphorus in the Lake Erie western and
central basin water.
2. Physical regeneration (resuspension) appears to be
the major source of internal phosphorus loading.
172
-------
TABLE 49. TOTAL PHOSPHORUS "ANOXIC" REGENERATION
ASSOCIATED WITH ANOXIC HYPOLIMNION
LAKE ERIE WESTERN AND CENTRAL BASIN
1973, 1974 AND 1975
Year
1973
1974
1975
External Total Phosphorus
Loading (Metric Tons* )
W.B.
1750Q
166QO
14900
C.B.
4OOG
42 OO
3850
W.B.&C.B.
21495
20869
18775
Total Phosphorus ''Anoxic Regenerc
W.B.
0
O
0
C.B.
2570
1720
730
W.B.&.C.B.
2571
1723
731
ation (Tone)
% of External Loading
W.B.&.C.B.
12.0
8.3
3.9
* Corps of Engineers, Buffalo District, Data.
-------
3. Anoxic regeneration of phosphorus was considered of
secondary importance as a source of internal loading
and only contributed during the late summer in the
central basin.
4. During 1970-1975, the western and central basin
served as phosphorus sinks.
REFERENCES
Burns, N. (ed.) 1976. Lake Erie in the early seventies. J. Fish
Res. Bd. Can. 33(3):349-645.
U.S. Army Corps of Engineers. 1975. Lake Erie water manage-
ments study, Vol. 3. U.S. Army, Corps of Eng., Buffalo
Dist. Buffalo, N.Y.
174
-------
SECTION 7
CHLOROPHYLL a AND PHEOPIGMENT DISTRIBUTION IN THE
CENTRAL AND WESTERN BASINS OF LAKE ERIE
Laura A. Fay and David E. Rathke
Center for Lake Erie Area Research
The Ohio State University
INTRODUCTION
Chlorophyll in Lake Erie's central and western basins was moni-
tored by the Center for Lake Erie Area Research (CLEAR) from 1973 to
1975 as a portion of the Lake Erie Nutrient Study sponsored by the U.S.
Environmental Protection Agency's Large Lake Research Station at
Grosse He, Michigan (Grant No. R-802543-01). Chlorophyll pigments,
along with other eutrophication indicators, were measured to determine
if recent attempts to impede eutrophication by reducing nutrient loading
into Lake Erie have been effective. Water samples collected on 15
general cruises during these three field seasons were analyzed for
chlorophyll pigments. Sampling and analysis in Lake Erie's eastern
basin was undertaken by the Great Lakes Laboratory (GLL) at Buffalo
(SUNY).
The major objective of this portion of the survey was to quantify
chlorophyll a. and pheopigment a in the central and western basins and
to observe seasonal and yearly trends of chlorophyll concentrations.
Shoreline versus open lake concentrations were also examined. To
analyze the data, several analyses of variance and regression models
were used.
MATERIALS AND METHODS
Water samples were collected from 51 stations, 37 in the central
basin and 14 in the western basin, with Niskin water bottles set in series.
Depth selections were identical to those for nutrient analysis. Water
was filtered through Whatman GF/C glass fiber filters to which a few
drops of MgCOs suspension were added. The samples were placed
in plastic petri dishes and frozen (-8°C) until analysis.
175
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Filters were ground with a teflon homogenizer at 300 rpm for less
than one minute in 4 ml of 90 percent acetone. The samples were
diluted to 12 ml and placed in a dark refrigerator for 4 hours before
analysis.
After 10 minutes of centrifugation at 3000 rpm, the samples were
analyzed spectrophotometrically. A Varian Techtron model 635 automatic
spectrophotometer in conjunction with a computer was used for sample
analysis and calculations. Table 50 presents the equations used to cal-
culate chlorophyll concentrations. Three different equations for chloro-
phyll a were used: Odum et al. (1958); SCOR/UNESCO (1966) and
Lorenzen (1967), chlorophyll a corrected for the amount of pheopigment.
Calculations of chlorophylls b and c_, and pheopigment a were made
for all samples. In addition, the ratio of absorbances at 663 nm before
and after acidification (Fo/Fa ratio, Lorenzen, 1967) was determined.
An Fo/Fa ratio of 1.7 indicates that the sample contained pure chloro-
phyll a, while a ratio of 1.0 indicates that the sample was pure pheo-
pigment a.. Ratios falling within this range represent varying percentages
of both chlorophyll a and pheopigment a. This ratio was helpful in deter-
mining the physiological condition of the phytoplankton.
The chlorophyll data are expressed as jug/I of corrected chlorophyll a
or as metric tons of corrected chlorophyll a. All of the data presented ~
were volume weighted in accordance with the procedure outlined in the
overview section.
Chlorophyll Standards
Chlorophyll standards were acquired from the U.S. Environmental
Protection Agency, Environmental Monitoring and Support Laboratory at
Cincinnati, Ohio. Comparisons of estimated concentrations by the E.P.A.
and those determined by CLEAR on the Varian Techtron are as follows:
Equation
SCOR/UNESCO
Chi a
Chi b
Chi c
Lorenzen
Corrected Chi a
pheopigment a
: , 1
EPA
8.40 + 0.40
2.55 + 0.20
1 . 69 + 0 . 40
6 . 09 + 1.10
4.17 + 1 . 40
CLEAR
8.21 + 0.04
2.85 + 0.03
1 .50 + 0.02
5.83
4.30
176
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TABLE 50. EQUATIONS USED FOR LAKE ERIE
1973-1975 CALCULATIONS
SCOR/UNESCO (1966)
Chi a (vg/l)=11 .64(0.D.663-O.D.750)-2.16(O.D.645-O.D.750)
+0.10(O . D. 630-O . D. 750)
Chi b (Mg/l)=20.97(O.D.645-O.D.750)-3.94(O.D.663-O.D.750)
-3.66(O . D. 663-O . D . 750)
Chi c (pg/l)=54.22(O.D.630-O.D.750)-14.81(O.D.645-O.D.750)
-5.53(O . D. 663-O . D. 750)
Lorenzen (1967)
Corrected Chi a (Mg/l)=26.73(O.D.663-O.D.750)
-(O.D.663ac.d-O.D.750ac.d)
Pheopigment a (pg/l)=26.73x1 .7(0 .D.663acid-O.D.750acid)
-(O . D. 663-O . D. 750)
Fo :Fa=O . D. 663-O. D. 750/(O . D. 663acid~° D 750acid)
Odum et al. (1958)
Chi a (pg/l)=13.4(O.D.665)
* All values must take into account the volume of the extract (ml),
the volume of water filtered (liters) and cuvette cell length (cm).
These factors must all be taken into account for the final deter-
mination. An example of a complete SCOR/UNESCO equation
follows:
11 .64(O . D. 663-O . D .750)-2.16(O . D. 645-O .D .750)
Chi a = +0.1O(O.D.63O-Q.D.750)xVolume of extract
Volume filtered (liters) x cuvette length (cm)
O.D. = Optical Density.
177
-------
Chlorophyll b_ was found to be slightly higher than the EPA estimate
Both chlorophylls a and £ fell within the estimated range. Comparisons
of corrected chlorophyll a_ and pheopigment a were also positive.
Measurement Errors
A single sample of chlorophyll was scanned 20 times at 750, 663,
645 and 630 nm during a 30-minute period and the results analyzed for
repeatibility. Chlorophyll a concentrations had a standard deviation of
0.01 with 0.12 percent standard deviation. Chlorophyll t) appeared to be
the second most reliable determination with 0.02 standard deviation
(0.70 percent). Chlorophyll £ had a 0.04 standard deviation or 3.60
percent.
An error resulting from an inaccurate absorbance reading of 0.001
was calculated for each of the three chlorophyll absorbance maxima.
A theoretical measurement error of less than one percent was calculated
for chlorophylls a_ and JD, while the estimated error for chlorophyll c
was 4.4 percent. In addition the chlorophyll £ value was also influenced
by an error at the chlorophyll ts absorbance.
Less than one percent of the chlorophyll ^ and c observations had
negative values. It was felt that the negative concentrations resulted
from errors in absorbance readings when low chlorophyll concentrations
were encountered. Errors resulting from extended periods of freeze
storage (greater than 60 days) for corrected chlorophyll a and pheopig-
ment a were estimated. Due to the variability of the length of sample
storage, plus the need for further investigations into the effects of
storage on chlorophyll degradation and pheopigment development, no
attempt was made to correct for this source of error.
RESULTS AND DISCUSSION
When analyzing the chlorophyll a data for the 1973, 1974 and 1975
seasons the major emphasis was placed on generalized and long-term
effects. Due to the three-year duration of the study and lengthy inter-
vals between sampling cruises, it was felt that this overview would
yield the most useful information concerning the overall trends and
regional effects since the 1970 Canada Centre for Inland Waters study
(Glooschenko et al. 1974).
178
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Sub-Basin Comparisons
Geologically, Lake Erie is divided into western, central and east-
ern basins by glacial moraines. The western basin is the shallowest
of the three basins with a mean depth of 7.4 m and represents only 5
percent of the total lake volume. The central basin has a mean depth
of 18.5 m and contains 63 percent of the total lake volume (Verber,
1950). The relationship of basin morphometry to the rate of eutrophi-
cation has been discussed by many investigators. Vollenweider (1971)
demonstrated that similar nutrient loadings to a shallow lake (10 m)
and to a deeper lake (100 m) would result in a greater increase in
the rate of eutrophication of the shallower lake. Sakamoto (1966) also
emphasized that phytoplankton production was influenced by basin depth
and attributed this influence to differences in availability of light and
nutrients in deep versus shallow basins.
The shallow western basin of Lake Erie is continuously supplied
with nutrients from the Detroit, Maumee, Raisin and Portage Rivers
as well as non-point sources. In addition to external loading, nutrients
are regenerated from decomposition of organic matter, i.e. plankton.
Thus the nutrient-rich western basin maintains high concentrations of
chlorophyll a throughout the year.
The deeper central basin is less productive than the western basin
throughout the year and especially during the summer months. Summer
standing crops in deep lakes are generally smaller than they are during
vernal and autumnal circulation periods (Sakamoto, 1966). This can be
attributed to the accumulation of nutrients in the hypolimnion which are
largely prevented from mixing with the nutrient-depleted waters of the
epilimnion.
The basin effect on chlorophyll a concentrations for the western
basin and the central basin was found to be of first order importance.
Western-central basin ratios of corrected chlorophyll a were calculated
using volume-weighted cruise averages (Table 51) in order to evaluate
the difference between the basins. The average concentrations of chloro-
phyll a in the western basin were usually two to three times those found
in the central basin. The largest basin ratios were observed during
the summer months when the western basin reached its maximum chloro-
phyll a concentrations and the central basin its lowest. The lowest
western to central basin ratios were observed in the fall and winter
months. For example, during the fall of each year, chlorophyll a con-
centration was reduced to approximately 1 .5 times that of the central
basin.
179
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TABLE 51. CORRECTED CHLOROPHYLL a
VOLUME WEIGHTED CONCENTRATIONS, 1973-1975
Year
1973
1974
1975
Cruise
17-23 July
29 Aug-4 Sept
14-24 October
Yearly Average
25 April-4 May
1-10 June
28 June-7 July
12-19 August
26 Aug-7 Sept
21 Oct-1 Nov
Yearly Average
27-31 March
21-25 April
9-19 June
13-21 July
30 Aug-5 Sept
27 Sept - 6 Oct
2-14 December
Yearly Average
Western
8.34
11 .74
11 .95
10.68
8.75
10.03
16.50
17.14
13.48
14.65
13.43
10.34
21.07
17.59
16.31
12.34
4.66
13.72
Central
2.40
3.38
7.87
4.55
3.47
2.63
2.41
3.82
3.76
9.45
4.26
5.09
2.71
3.11
7.72
10.05
6.92
5.93
W :
Basin
3.48 :
3.47 :
1 .52 :
2.34 :
2.52
3.81
6.85
4.49
3.59
1 .55
3.15
2.03
7.78
5.66
2.11
1.23
0.67
2.31
C
Ratio
1 .0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
180
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For more detailed chlorophyll a. comparisons within the central
basin, it was subdivided into eastern and western segments. The divi-
sion was demarcated by a line running south from Pointe Aux Pins,
Ontario to slightly east of Cleveland, Ohio (Figure 59). The eastern
central basin included stations 23-29 (grids 19-31) and the western basin
stations 40-54 plus stations 65 and 74 (grids 32-43). Cruise average
corrected chlorophyll a concentrations for the three sub-basins were
examined (Table 52). The western basin normally had the highest aver-
age chlorophyll a values ranging from 1 to 9 times the eastern central
basin average concentrations, although exceptions to the west-east trend
of decreasing chlorophyll a_ concentrations did occur. For example, in
December of 1975 the western basin was observed to have lower chloro-
phyll a values than the eastern central basin (0.7:1.O). The western
centraT basin generally had the second highest corrected chlorophyll a
concentrations with the exception of early June 1974 and early October
1975. The three-year average corrected chlorophyll a_ concentrations
for each station (Table 53) also substantiate the west to east trend
(Figures 60 and 61). Two anomolies existed: (1) a low concentration
area in the western basin east of the Detroit River mouth and (2) a
high concentration area in the central basin extending from the island
area eastward along the southern shore of the central basin.
Due to high and low areas of concentration within the central basin,
it was further subdivided into 3 regions (Figure 62): (1) shore (s), (2)
mid-lake (m) and (3) the Sandusky sub-basin (b), Thomas, et al. (1976).
The average corrected chlorophyll a concentrations (ug/l) of the western
basin and the designated sub-basins of the central basin for each of the
three years were:
1973 1974 1975
Western (wb) 12.05 13.45 14.80
Sandusky sub-basin (b) 6.55 9.O8 9.77
Shore (s) 5.40 4.52 6.71
Mid-lake (m) 3.60 3.3O 5.O8
The trend throughout the three years was very consistent. Chlorophyll
a was found in highest concentrations in the western basin followed by
the Sandusky sub-basin and the shore area, with the mid-lake having
the lowest values, as demonstrated by the three-^year basin ratio:
western Sandusky
basin sub-basin shore mid-lake
3.37 : 2.18 : 1.36 : 1.00
181
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BASIN SUBDIVISIONS USED FOF
CHLOROPHYLL ANALYSIS
1973-1975
Figure 59. Basin subdivisions used for chlorophyll
analysis 1973-1975.
CENTRAL BASIN
SUBDIVISIONS USED FOR
CHLOROPHYLL ANALYSIS
1973-1975
Figure 60.
Central basin subdivisions used for chlorophyll
analysis, 1973-1975.
182
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TABLE 52. CORRECTED CHLOROPHYLL a (jug/I)
VOLUME WEIGHTED CONCENTRATIONS, 1973-1975
oo
CO
Cruise
1973
Late July
Late August
Mid October
1974
Early May
Early June
Early July
Mid August
Early September
Late October
1975
April
Mid June
Mid July
Early September
Early October
Early December
Western
Basin
8.34
11.74
11.95
8.75
10.03
16.50
17.14
13.48
14.65
10.34
21.07
17.59
16.31
12.34
4.66
Central E
West
3.69
5.22
8.27
4.32
2.03
3.74
5.77
5.44
12.56
6.28
3.28
4.49
10.26
9.89
7.47
Sasin
East
1.75
2.37
7.65
3.01
2.54
1.68
2.76
2.84
7.74
4.44
2.40
2.36
6.33
10.14
6.62
Basin
WB :
4.77 :
4.95 :
1 .56 :
2.91
3.95
9.82
6.21
4.75
1 .89
2rtrt
.33
8.78
7.45
2.58
1.22
0.70
Ratio
WCB
2.05 :
2.20 :
1.08 :
1 .44
0.80
2.23
2.09
1 .92
1 .62
4 A A
1 ,°t 1
1.37
1.90
1.62
0.98
1.13
: ECB
1.0
1.0
1 .0
1.0
1.0
1.0
1 .0
1 .0
1.0
In
(J
1 .0
1 .0
1 .0
1.0
1 .0
-------
TABLE 53. CORRECTED CHLOROPHYLL a (jug/I)
STATION MEANS FOR THE 15 CRUISES, 1973-1975
oo
Station
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
Means
5.44
3.94
3.65
3.54
3.93
4.55
4.73
4.45
3.56
3.72
4.24
6.05
6.37
4.01
3.41
3.96
4.26
5.21
5.81
3.80
4.46
9.03
5.74
5.87
4.79
6.06
6.84
Standard
Deviation
4.28
3.13
2.76
2.75
3.06
2.89
2.92
3.51
2.86
2.52
3.23
4.13
3.98
3.02
2.25
2.41
3.26
2.97
3.98
3.07
3.60
4.99
4.41
4.24
3.77
3.49
4.76
Number of
Measurements
46
46
50
47
51
49
45
54
52
52
45
48
43
53
51
52
46
49
51
51
52
44
49
43
50
48
46
Station
50
51
52
53
54
55
56
57
58
59
60
61
65
66
67
68
69
70
71
72
73
74
75
76
78
79
81
Means
8.36
7.35
8.53
10.53
10.16
11.43
11 .98
14. 8O
13.66
12.93
14.30
7.47
9.21
13.29
11.15
11 .51
14.42
24.60
2.63
4.04
3.87
8.55
21 .23
20.79
3.84
3.02
20.14
Standard
Deviation
3.63
4.16
6.38
5.80
6.98
6.60
4.57
13.31
8.67
7.63
12.71
4.69
5.57
5.36
8.10
8.17
7.62
14.93
1 .72
3.53
3.50
5.15
14.25
10.56
2.99
2.41
11 .43
Number of
Measurements
43
46
46
45
45
42
45
44
39
45
45
41
46
39
46
45
20
32
10
10
51
45
31
22
45
42
39
-------
Corrected Chlorophyll a
Corrected Chlorophyll a (ug/l) _.
0 10 * °> 00 0
1 ' 1 » f
n I
to 8-
c
(D
0)
ro
*.
ui
0) ^S
O T
r1-«§
T (D
j» ^
i* O" -
00 Q) -T
S jg«
fn (D o
w n 3
rf " "i oo
O) w -
H- O
! T
O ~
D 0
-w 3
^T)
(0 "^
CO " 0
ilo> °
^£
Ol (O
l""^
or ~
m
D>
a -
3 s:
.----' 3 «Q
/ c
^-^" ro -
m****^ O»
\. 0) "^
^» _h
op f
w m »^^^
n n o>-
(D 2 > tj ^^
C T ^ x*' ^ <
j T ~L ST y> ii
o o ^- -. __ S; (Q s*:
a a ^X*3 O*
/ _* 2 in »~
X 5 8 1 :
N-^^ " ro^ t
~~7i»
- ^.-__ , Jl
^~^-~.ui a
r+
C S
1 1 1 \\ 1 1
\\
«
9
»
* ^
0
_
A
_
./
* .
:*
«
9*
m
*
0*
*
,*
*
A
^
-------
The three-year average chlorophyll a concentrations for each station
in the central basin showed a shore versus mid-lake effect (Figure 63).
For each transect across the lake, i.e. stations 23 to 29, the south
shore (Station 23) exhibited the highest concentrations with a mid-lake
low (Station 25) and higher concentrations on the north shore (Station 29).
The south shore concentrations were higher than the north shore concen-
trations with the shore stations in the western central basin having
higher concentrations than those in the eastern central basin. Regres-
sion analysis showed the basin subdivisions to be of first order signifi-
cance, accounting for the major variability in the three years of data.
The chlorophyll data indicated that areas designated as sub-basins were
signficantly different and represented areas reflecting "real" differences
in standing crop. These differences demonstrate the need for a synoptic
lake survey.
Seasonal Trends
Seasonal variations of chlorophyll a_ concentrations have been docu-
mented by previous studies on the Great Lakes. Although each of the
Lake Erie sub-basins exhibited somewhat different seasonal trends, two
major effects were evident. The mid-lake central basin and to a lesser
extent the north and south shoreline exhibited an early spring maximum,
a summer minimum and a rapid increase following fall turnover. The
western basin and Sandusky sub-basin both had highest average concen-
trations during the summer months and lower concentrations in both
spring and fall. The percent of the yearly total chlorophyll a and pheo-
pigment present during each cruise substantiates these western and
central basin trends (Tables 54 and 55). Contour maps of corrected
chlorophyll a concentrations averaged for all depths from each cruise
of 1973 through 1975 further depict seasonal trends (Figures 64-78).
Volume-weighted cruise tonnages and concentrations of corrected chloro-
phyll a_ and pheopigment a are presented for the central (Table 56) and
western (Table 57) basins.
The factors responsible for the seasonal responses of the phyto-
plankton are complex. Seasonal variations in temperature and light
are of primary significance in the development of phytoplankton standing
crop. As the vernal equinox approaches, the sun moves directly over-
head resulting in a greater percent of the sun's light penetrating the
water column. This provides more available light for photosynthesis
while increasing the water temperature. During the late spring and
summer months low turbidity in the western and central basins further
enhances light penetration. The shallow western basin warms more
rapidly than the deeper central basin and maintains a higher temperature
throughout the summer. These conditions plus the availability of nutrients
186
-------
00
CORRECTED CHLOROPHYLL a (JJG/L)
STATION MEANS FOR 1973-1975
CONTOUR INTERVAL = 5 pG/L
Figure 63. Corrected chlorophyll a (xjg/l) station means for 1973-1975.
-------
TABLE 54. PERCENT OF ANNUAL TOTAL CORRECTED CHLOROPHYLL a AND
PHEOPIGMENT a CONTRIBUTED AT EACH CRUISE INTERVAL - CENTRAL BASIN
Yoar Llmnlon
1973 epl
me so
hypo
total
1974 epl
meso
hypo
total
1979 ept
meso
hypo
total
Unstratified
Apr 1 1 -May
Chi a
13.9
14.4
Pheo a
13.0
26.4
Stratified
June
Chi a
5.9
0.7
3.0
9.6
3.6
0.7
3.3
7.6
Pheo a
4.3
0.6
3.6
8.5
3.0
0.7
3.0
6.7
July
Chi a
12.6
1.5
3.6
17.7
8.2
0.2
1.2
9.6
4.4
1.1
3.2
8.7
Pheo a
19.8
2.3
3.3
25.4
9.4
0.4
2.7
12.5
2.2
0.6
1.9
4.7
Aug- Sept
Chi a
20.8
1.8
3.0
25.6
12.6
0.9
1.6
15.1
17.3
1.0
3.5
21.8
Pheo a
24.1
3.6
3.8
31.5
10.1
0.9
3.5
14.5
9.5
0.7
3.0
13.2
Sept-Oct
Cht a
13.3
0.5
1.1
14.9
28.2
Pheo a
23.9
0.9
2.7
27.5
13.3
Unstratified
Oct-Dec
Chi a
56.7
36.8
19.2
Pheo a
43.1
23.6
35.7
CO
CO
-------
TABLE 55. PERCENT OF ANNUAL TOTAL CORRECTED CHLOROPHYLL a AND
PHEOPIGMENT a CONTRIBUTED AT EACH CRUISE INTERVAL - WESTERN BASIN
Year Llmnlon
1973
ept
meso
hypo
total
1974
epl
meso
hypo
total
1975
epl
meso
hypo
total
April-May
Chi a
11.1
12.7
Pneo a
8.0
18.3
June
Chi a
12.7
25.8
Pneo a
10.7
16.6
July
Chi a I
26.7
20.8
21.4
Pheo a
43.6
24.8
11.6
Aug-Sept
Chi a
37.2
21.2
19.8
Pheo a
25.1
22.6
18.8
Sept-Oct
Chi a
16.6
14.8
Pheo a
24.2
20.5
Oct-Dec
Chi a
36.1
17.5
5.5
Pheo a
31.3
9.8
14.1
00
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! CORRECTED CHLOROPHYLL a (JJG/U
AVERAGE FOR ALL DEPTHS
SAMPLED
CRUISE 2 JULV 17-JULV 24, 1973
CONTOUR INTERVAL = ?-10
Figure 64. Corrected chlorophyll a (jjg/\~) average for all depths sampled,
cruise 2, July 17-July 24, 1973.
-------
CORRECTED CHLOROPHYLL a (JJG/U
AVERAGE FOR ALL DEPTHS
SAMPLED
CRUISE 5 AUG. 29-SEPT. 4, 1973
CONTOUR INTERVAL » 2-1O JLK3/L
Figure 65.
Corrected chlorophyll a (Mg/l) average for all depths
sampled, cruise 5, August 29-September 4, 1973.
-------
ro
CORRECTED CHLOROPHYLL a (LIG/U
AVERAGE FOR ALL DEPTHS
SAMPLED
CRUISE 7 OCT. 14-OCT. 24, 1973
CONTOUR INTERVAL = 2-10 JJG/L
Figure 66.
Corrected chlorophyll a (;ug/l) average For all depths
sampled, cruise 7, October 14-October 24, 1973.
-------
UD
CO
CORRECTED CHLOROPHYLL a (JJG/L)
AVERAGE FOR ALL DEPTHS
SAMPLED
, CRUISE 2 APR. 25-MAY 4, 1974
j CONTOUR INTERVAL = 2-5 XJG/L
Figure 67.
Corrected chlorophyll a (ug/l) average for all depths
sampled, cruise 2, April 25-May 4, 1974.
-------
vo
CORRECTED CHLOROPHYLL a (JJG/L)
AVERAGE FOR ALL DEPTHS
SAMPLED
CRUISE 4 JUNE 1-JUNE 10, 1974
CONTOUR INTERVAL = 2-5 JUG/L
! I j
Figure 68.
Corrected chlorophyll a (jug/I) average for all depths
sampled, cruise 4, June 1-June 10, 1974.
-------
in
CORRECTED CHLOROPHYLL a (JJG/U
AVERAGE FOR ALL DEPTHS
SAMPLED
, CRUISE 5 JUNE 28-JULY 7, 1974
1 CONTOUR INTERVAL = 2-10
Figure 69.
Corrected chlorophyll a Oug/l) average for all depths
sampled, cruise 5, June 28-July 7, 1974.
-------
vo
CORRECTED CHLOROPHYLL a (JJG/L)
AVERAGE FOR ALL DEPTHS
SAMPLED
CRUISE 7 AUG. 12-AUG. 19, 1974
CONTOUR INTERVAL * 2-10 L'G/L
Figure 70.
Corrected chlorophyll a C/ug/l) average for all depths
sampled, cruise 7, August 12-August 19, 1974.
-------
10
CHLOROPHYLL
AVERAGE FOR ALL DEPTHS
SAMPLED
CRUISE 8 AUG. 26-SEPT. 7, 1974
CONTOUR INTERVAL =2=10 JJG/L
Figure 71.
Corrected chlorophyll a (jug/I) average for all depths
sampled, cruise 8, August 26-September 7, 1974.
-------
00
CORRECTED CHLOROPHYLL a (JJG/U
AVERAGE FOR ALL DEPTHS
SAMPLED
CRUISE 10 OCT. 21-NOV. 1, 1974
CONTOUR INTERVAL = 2-5 JUG/L
Figure 72. Corrected chlorophyll a (;ug/l) average for all depths
sampled, cruise 10, October 21-November 1, 1974.
-------
10
CORRECTED CHLOROPHYLL a UJG/L)
1.0 METERS BELOW LAKE SURFACE
CRUISE 1A & 1B MARCH 27-APR. 2E
1975
CONTOUR INTERVAL = 2-5 JJG/L
Figure 73. Corrected chlorophyll a (jug/I), cruise 1A and 1B,
March 27-Aprll 25, 1975.
-------
t\>
o
o
CORRECTED CHLOROPHYLL a (JJG/L)
1.0 METERS BELOW LAKE SURFACE
CRUISE 2 JUNE 9-JUNE. 19, 1975
I CONTOUR INTERVAL = 2-10 JJG/i,
Figure 74. Corrected chlorophyll a (jug/I), cruise 2,
June 9-June 19, 1975.
-------
ro
o
CORRECTED CHLOROPHYLL a (JJG/L)
1.0 METERS BELOW LAKE SURFACE
CRUISE 3 JULY 13-JULY 21, 1975
CONTOUR INTERVAL = 2-10 JUG/L
Figure 75. Corrected chlorophyll a (AJg/l), cruise 3,
July 13-July 21, 1975.
-------
ro
o
CORRECTED CHLOROPHYLL a (JUG/L)
|1 .0 METERS BELOW LAKE SURFACE
' CRUISE 4 AUG. 30-SEPT. 5, 1975
CONTOUR INTERVAL = 2-5 1JG/L
Figure 76. Corrected chlorophyll a (xjg/l), cruise 4,
August 30-September 5, 1975.
-------
ro
o
CO
CORRECTED CHLOROPHYLL a (JJG/L)
1.0 METERS BELOW LAKE SURFACE
CRUISE 5 SEPT. 27-OCT. 6, 1975
CONTOUR INTERVAL = 2-5
Figure 77.
Corrected chlorophyll a Qug/l), cruise 5,
September 27-October 6, 1975.
-------
ro
o
-P.
CORRECTED CHLOROPHYLL a (LJG/L)
1.0 METERS BELOW LAKE SURFACE
CRUISE 6 DEC. 2-DEC. 14. 1975
CONTOUR INTERVAL = 2 JJG/L
Figure 78. Corrected chlorophyll a (ug/l), cruise 6,
December 2-December 14, 1975.
-------
TABLE 56. CORRECTED CHLOROPHYLL- a AND
PHEOPIGMENT a - CENTRAL BASIN
Limnion
1973
Epi tons
M9/1
Meso tons
M9/1
Hypo tons
M9/1
Total tons
M9/1
1974
Epi tons
(J9/1
Meso tons
M9/1
Hypo tons
M9/1
Total tons
M9/1
1975
Epi tons
M9/1
Meso tons
pg/1
Hypo tons
M9/1
Total tons
M9/1
Unstratified
April-May
Chi a
1082.73
3.47
1571 .02
5.09
Pheo a
231 .82
0.79
584 . 20
1 .89
Stratified
June
Chi a
458.06
2.34
58.29
2.56
233.22
2.49
749 . 57
2.40
395 . 1 1
2.27
81 .48
2.65
362.84
3.47
839.43
2.71
Pheo a
78.05
0.40
11 .05
0.49
65.54
0.70
154.64
0.50
66.90
0.38
14.78
0.48
67.18
0.64
148.86
0.48
July
Chi a
535 . 1 8
2.34
63.97
2.86
152.21
2.48
751 .36
2.40
641.82
2.58
18.28
1.46
92.21
1 .84
752.31
2.41
482.69
2.64
121 .26
3.41
351 .75
3.89
955 . 70
3.10
Pheo a
248.53
1 .08
28.90
1 .29
41 .21
0.67
318.64
1 .02
1 68 . 82
0.68
6.34
0.51
48.73
0.79
223 . 89
0.72
49. 08
O.27
13.96
0.39
41 .41
0.46
104.45
0.34
(continued)
205
-------
TABLE 56 (continued)
Limnion
1973
Epi tons
M9/1
Meso tons
M9/1
Hypo tons
M9/1
Total tons
M9/1
1974
Epi tons
M9/1
Meso tons
M9/1
Hypo tons
M9/1
Total tons
M9/1
1975
Epi tons
M9/1
Meso tons
pg/i
Hypo tons
jjg/l
Total tons
M9/1
Stratified
Aug-Sept
Chi a
884 . 99
3.72
75.90
3.05
1 29 . 40
2.75
1 090 . 29
3.52
985.20
4.19
72.81
3.05
127.02
2.51
1 1 85 . 03
3.82
1894.42
8.33
108.98
7.16
380.98
5.77
2384.38
7.72
Pheo a
302.90
1 .27
45.64
1.84
48.07
1.02
389 . 61
1 .26
181 .01
0.77
17.01
0.71
63.18
1.25
261 .20
0.84
210.04
0.92
14.80
0.97
66.06
1.00
290.90
0.94
Unstratified
Sept-Oct
Chi a
1036.03
4.26
41.61
2.38
85.16
1.76
1 1 62 . 80
3.76
3086.02
10.05
Pheo a
429.46
1.77
17.05
0.98
49.02
1 .01
495 . 53
1.60
293 . 58
0.96
Oct-Dec
Chi a
2409 . 1 2
7.87
2871.40
9.45
2103.32
6.92
Pheo a
541.78
1.77
423.44
1 .39
790.16
2.60
206
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TABLE 57. CORRECTED CHLOROPHYLL a AND
PHEOPIGMENT a - WESTERN BASIN, 1973-1975
Limnion
1973
Epi tons
M9/1
Meso
Hypo
Total
1974
Epi tons
H9/1
Meso
Hypo
Total
1975
Epi tons
H9/1
Meso
Hypo
Total
1973
Epi tons
jjg/1
Meso
Hypo
Total
1974
Epi tons
pg/l
Meso
Hypo
Total
1975
Epi tons
M9/1
Meso
Hypo
Total
Chi a Pheo a
April-May
210.OO 26.17
8 . 75 1 . 09
244.52 66.01
1 0 . 34 2 . 79
Aug-Sept
277.61 81.74
11.74 3.46
403.22 73.83
17.14 3.14
380 . 86 67 ..99
16.31 2.91
Chi a Pheo a
June
241.74 34.81
10.03 1.44
495.14 6O.36
21.07 2.57
Sept-Oct
316.09 78.60
13.48 3.35
284.58 74.21
12.34 3.22
Chi a Pheo a
July
199.28 141.82
8.34 5.93
395. O9 80.78
16.50 3.37
410.76 42.19
17.59 1.81
Oct-Dec
269.37 101.77
11.99 4.52
331.43 31.70
14.65 1 .40
105.01 51.23
4 . 66 2 . 27
207
-------
in the western basin provide an environment favorable for algal growth
evidenced by the high concentrations of chlorophyll a_ observed throughout
the basin during the summer.
The central basin maintained high chlorophyll a concentrations in
the spring prior to stratification and during the fall "following turnover.
The maximum central basin concentrations were found in the fall just
after turnover when the nutrient-rich hypolimnion became mixed with
the nutrient-depleted epilimnion. In the late fall, storms cause resus-
pension of sedimented material, which increases the turbidity. At the
same time, water temperatures are decreasing, and, thus, chlorophyll
concentrations decline during this period.
Year to Year Trends
In order to facilitate a year to year comparison of chlorophyll a
in Lake Erie, volume-weighted quantities (metric tons) were calculated.
The cruise by cruise volume-weighted quantities were plotted for 1973
through 1975, and points equidistant between the cruises were used to
estimate chlorophyll a. values for periods between cruises. The between-
cruise estimates and the cruise values were averaged for each year to
give estimated chlorophyll a tonnages for the western and central basins.
This procedure was used to compensate for irregular sampling intervals
and varying number of cruises over the three-year period. For compar-
ative purposes only, the between-cruise estimates of chlorophyll a from
June through November of each year were used due to the great vari-
ability found in early spring and late fall data.
From 1973 to 1975, the mean tonnage of corrected chlorophyll a
increased in the central basin: ~"
Central Basin
1973 1201.25 tons
1974 1291 .78 tons
1975 1795.08 tons
From 1973 to 1975, an increase of 49 percent was estimated and the
greatest year to year difference, a 38 percent increase in corrected
chlorophyll a_, occurred between 1974 and 1975. This large increase was
attributed to a greater hypolimnion concentration of chlorophyll a in 1975
than in 1974. The establishment of the thermocline higher in the water
column in 1975 resulted in a larger hypolimnion volume. Since a great-
er portion of the hypolimnion was in the photic zone and nutrients were
less likely to become limiting, the hypolimnion was able to support a
larger standing crop of phytoplankton during this year. In addition, the
extensive anoxic conditions which developed in the hypolimnion in 1973
208
-------
and 1974 were not encountered in 1975. Lack of extensive anoxia also
provided a more favorable hypolimnion environment than was previously
observed.
From 1973 to 1975 the mean tonnage of corrected chlorophyll a
increased in the western basin, with the greatest increase in mean tons
occurring between 1973 and 1974. The three-year data is as follows:
Western Basin
1973 236.96 tons
1974 342.51 tons
1975 395.01 tons
The corrected chlorophyll a. tonnages were estimated to be 44.5 percent
greater in 1974 than 1973, and a further increase of 15 percent was
calculated for 1975 and 1974.
The significance of year to year differences of chlorophyll a ton-
nage in Lake Erie is somewhat difficult to evaluate over a short time
period. Yearly changes in standing crop can be expected due to sea-
sonal variation, different meteorological effects and cruise scheduling.
Only a sampling program spanning several additional years will provide
the necessary information to evaluate changes in the trophic status of
the lake.
Historical Trends
Few previous studies on Lake Erie have examined chlorophyll
concentrations (Table 58). Tucker (1949) utilized 1939 western basin
chlorophyll data supplied by Chandler and Weeks and reported a good
correlation (r = 0.78) between phytoplankton counts and plankton pig-
ments using the technique developed by Harvey (1934). Brydges (1971)
reported western basin and north shore data collected between 1967 and
1969. Spectrophotometric determinations were made on pigments ex-
tracted in 90 percent acetone and concentrations were calculated using
the equations developed by Richards and Thompson (1952). Brydges
found chlorophyll a to be positively correlated with total phosphorus,
with the highest concentrations of both chlorophyll a_ and total phosphorus
found in the western portion of the lake. Only two entire Lake Erie
chlorophyll surveys have been undertaken, one by the Federal Water
Pollution Control Administration (FWPCA, 1968) in May, July and Octo-
ber of 1967 and January and August of 1968, and another by Glooschenko
et al. (1974) based on 10 cruises from April to December of 1970.
209
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TABLE 58. SIMILAR STATIONS FROM CHLOROPHYLL
SURVEYS UNDERTAKEN BY THE FEDERAL WATER
POLLUTION CONTROL AGENCY (1968) AND THE
CENTER FOR LAKE ERIE AREA RESEARCH
1974-1975
STATION NUMBERS
FWPCA. 1967 - 1968 CLEAR, 1974 - 1975
D9-1 81
E2-1 70
G12-1 50
D13-1 51
F16-1 47
H20-1 73
L3O-1 78
L36-1 79
210
-------
The FWPCA sampled 30 stations along a longitudinal transect of
the entire lake (Figure 79). Eight FWPCA stations are comparable to
CLEAR western and central basin stations (Table 58) and were used for
comparison. The original FWPCA data utilized Richards and Thompson's
equations (1952), and the data was reported as chlorophylls a plus b.
The raw data absorbencies were obtained and recalculated using the
SCOR/UNESCO chlorophyll a equation (1966). The averages (surface
and mid-depths) for the eight stations on each of the May, July and
October cruises were recalculated and plotted (Figure 80). The FWPCA
1967 May to October average chlorophyll a was estimated to be 4.86
pg/1. Cruise averages based on the eight similar CLEAR stations dur-
ing corresponding time periods were plotted (Figure 80). The CLEAR
data for 1973 have not been included in this analysis because of missing
data prior to July. The yearly SCOR/UNESCO chlorophyll a averages
for 1974 and 1975 were 9.24 ,ug/I respectively. The 1974 and 1975
chlorophyll a concentrations were higher than the 1967 averages which
may be only"partially attributed to differences in sampling schedules.
There was an increase of 7.08 jug A of chlorophyll a from 1967 to 1975
for the western and central basins representing an increase of 146 per-
cent.
Problems were encountered in utilizing the 1967 FWPCA data.
The original FWPCA 645 nm absorbences (chlorophyll b_) were found to
be as high or higher than the absorbences at 663 nm (chlorophyll a).
Absorbences for chlorophyll b should be consistently lower than chloro-
phyll a absorbences. It has been pointed out that the Beckman D
spectre-photometer used by the FWPCA may have been miscalibrated
(Cornelius Weber, U.S. EPA, Cincinnati, personal communication).
Thus, the FWPCA chlorophyll data lost much of its validity.
Glooschenko, Moore and Vollenweider (1974) sampled 5 western
basin and 12 central basin stations during 10 cruises in 197O. Samples
were collected at 1 m and 5 m depths and integrated. The analytical
procedures were similar to those used by CLEAR, 1973 through 1975,
making the data comparable. Five western basin and twelve central
basin CLEAR stations similar in location to Glooschenko were selected
for comparison (Figure 81 and Table 59). Basin averages were cal-
culated using only surface and mid-depth or lower epilimnion values
for comparison with the 1970 study. Only SCOR/UNESCO chlorophyll a
values were compared since the 1970 report did not include corrected
chlorophyll a. Yearly averages were calculated for 1970 and 1973
through 1975~ based on cruises from June to November (Tables 60 and
61).
211
-------
1967-1968 F\APCA
STATION LOCA1 ION
Figure 79. FWPCA station locations, 1967-1968.
-------
14 -
r\>
ii
CO
10 -
O)
BJI
a
2
o
6
1975^
6 -
^- 11.94 jug/I, 1975
(June to November)
mean
9.24 jug/I, 1974
(June to November)
mean
4.86 jug/l, 1967
(May to October)
mean
1967
Figure 80.
Comparison of central and western basin chlorophyll a values
(average of surface and mid-depth values) from the FWPCA,
1967 data and corresponding 1974 and 1975 data.
-------
r>o
CLEAR STATIONS
ENCLOSED IN BOXES USED
FOR COMPARISONS WITH" '
GLOOSCHENKO'S 1970
CHLOROPHYLL STUDY
KILOMETERS
0 '0
STATUTE MILES
Figure 81.
79"
CLEAR stations enclosed in boxes used for comparisons with
Glooschenko's 1970 chlorophyll study.
-------
TABLE 59. SIMILAR STATIONS FROM CHLOROPHYLL
SURVEYS IN GLOOSCHENKO (1970) AND CLEAR (1973-1975)
Glooschenko, 1970 CLEAR, 1973 - 1975
Station Number Station Number
Western Basin Western Basin
23 50
25 61
27 59
29 55
31 53
Central Basin Central Basin
7 79
9 31
14 37
18 44
22 46
34 40
35 42
37 35
38 78
41 29
44 26
47 25
215
-------
TABLE 60. AVERAGE CHLOROPHYLL a (ug/l) IN LAKE ERIE
1970 BY MONTH AND WESTERN TO CENTRAL BASIN RATIOS
Cruise
April 7-11
May 6-10
June 2-6
July 3-7
July 28 - August 1
August 25 - 29
September 23 - 27
October 21-25
November 25 - 30
December 14 - 18
April 1 - December
(Yearly Average)
June-November Average
Western
5.9
4.4
5.9
11 .9
9.8
19.3
10.2
7.8
7.6
3.3
8.9
10. 8
Central
5.3
2.9
2.7
2.5
3.7
4.1
9.2
3.0
5.4
5.7
4.4
4.2
W
1 .11
1 .52
2.19
4.76
2.65
4.71
1 .11
2.60
1.41
0.58
2.02
2.58
: C
: 1 .0
: 1 .0
: 1 .0
: 1 .0
: 1 .0
: 1.0
: 1.0
: 1.0
: 1.0
: 1.0
: 1 .0
: 1 .0
Data source:
Taken from Glooschenko et al., 1974 (Eastern Basin data deleted).
216
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TABLE 61 . CLEAR SCOR/UNESCO CHLOROPHYLL a
CONCENTRATIONS BASED ON STATIONS SIMILAR TO
GLOOSCHENKO'S 1970 SURVEY (SURFACE AND
MID-VALUES ONLY)
Year
1974
1975
Cruise
1
2
3
4
5
6
Yearly
Average
(n = 6)
June-Nov
Average
(n = 5)
1
2
3
4
5
6
Yearly
Average
(n = 6)
June-Nov
Average
(n = 4)
Western
5.36
7.47
15.16
15.19
11 .09
12.05
11 .05
12.34
13.27
14.10
12.05
15.87
10.74
8.28
12.39
13.19
Central
4.68
2.30
1 .94
3.63
5.13
8.73
4.40
4.35
6.20
1 .82
2.85
8.20
10.34
8.82
6.37
5.80
Basin
W
1.15
3.25
7.81
4.18
2.16
1 .38
2.51
2.84
2.14
7.75
4.23
1 .94
1 .04
0.94
1 .95
2.27
Ratio
C
1 .00
1 .00
1 .00
1 .00
1 .00
1 .00
1 .00
: 1 .00
1 .00
1.00
1 .00
1 .00
1 .00
1 .00
1 .00
: 1.00
217
-------
The central basin chlorophyll a_ trends for 1970, 1974 and 1975
were similar (Figure 82). The bimodal peaks occurred during the spring
and early fall, and a summer minimum was evident all three years.
The magnitude of the high and low values found each year were similar,
with differences due to cruise scheduling and fluctuations in the standing
crop. The greatest increase in the average June to November chloro-
phyll a concentration occurred between 1974 (4.35 ;ug/l) and 1975 (5.80
;ug/l). The central basin showed an estimated concentration increase
of 38 percent from 1970 to 1975.
The western basin chlorophyll a. concentrations were high through-
out the summer months all three years (Figure 83). The average June
to November chlorophyll a_ concentration was lowest in 1970 (10.82 jug/l),
while in 1975 the average was 13.19/jg/l. This represented an esti-
mated increase of 2.37 jug/1 or 22 percent from 1970 to 1975.
If all CLEAR western and central basin stations and all depths
were utilized for comparison with Glooschenko's 1970 study, the increase
is even greater (Table 62). The 1975 June to November western basin
chlorophyll a average would be 18.26 jjg/l, representing a 7.44 jjg/l
or 84 percent estimated increase in the western basin chlorophyll a con-
centration from 1970 to 1975. An estimated increase of 2.04 jug/l~or
46 percent was calculated for the central basin from 1970 to 1975 based
on this comparative technique. This large increase can be partially
attributed to a more intensive lake coverage in 1974 and 1975 particular-
ly along the shorelines of the central basin and a more thorough coverage
of the western basin. The observed increase can be attributed to sevei
al factors such as year to year fluctuations, meteorological influences
and continued enrichment of the lake. The factors most responsible
for the observed increase can best be determined from continued yearly
studies. Comparisons of chlorophyll a data taken from several investi-
gations are difficult to evaluate because of differing analytical techniques,
sampling schedules and sampling locations (Tables 58 and 59). There-
fore, trends based on such comparisons must be viewed conservatively.
Trophic Status
Various trophic classification systems have been proposed based on
chlorophyll a concentrations. Three such classification systems were
examined to evaluate the degree of eutrophication of Lake Erie. Gloos-
chenko and Dobson (1975) established eutrophication classifications based
on chlorophyll a_ data from the Great Lakes. Tailing (1961) devised a
classification system applicable to marine Investigations. Sakamoto
(1966) studied a series of Japanese lakes ranging from 4.5 to 65 m in
depth from which he developed a method for determination of trophic
status. The three classifications were as follows:
218
-------
10
6 -
ol
jj 4
5
2-
1975 f
1970
5.80 pg/l=1975 mean
X*
:4.35 ;jg/l=1974 mean
4.20 jug/t=1970 mean
J rF ' M ' A ' M ' J ' J ' A ' S ' O n N ' D '
Month
Figure 82.
Comparison of central basin chlorophyll a^ values
from Glooschenko's 1970 data and corresponding
1974 and 1975 data.
20 1
16 -
12
ral
.c
a
o
L
O
12.34 xjg/l=1974 rr.ean /__:
10.82 ,ug/1=1 970 mean /_/\J T !-_.__
13.19 >ug/l=1975
mean
Figure 83.
'F'M'A'M'J'J'A'S'O'N'D'
Month
Comparison of western basin chlorophyll a values
from Glooschenko's 1970 data and corresponding
1974 and 1975 data.
219
-------
TABLE 62. CLEAR SCOR/UNESCO CHLOROPHYLL a
VOLUME WEIGHTED CONCENTRATIONS. BASIN
VALUES INCLUDE ALL STATIONS AT ALL DEPTHS
Year
1973
1974
1975
Cruise
1
2
3
Yearly
Average
1
2
3
4
5
6
Yearly
Average
(n = 6)
June-Nov.
Average
(n = 5)
1
2
3
4
5
6
Yearly
Average
(n = 6)
June-Nov.
Averages
(n = 4)
Western
10.42
13.70
16.15
13.42
9.78
10.93
18.24
18.84
15.45
14.46
14.62
15.58
11 .92
22.53
18.14
18.09
14.26
6.O3
15.16
18.26
Central
2.93
3.91
8.97
5.27
3.85
2.80
2.73
4.34
4.70
10.33
4.79
4.98
6.19
2.95
3.26
8.23
10.53
8.21
6.56
6.24
Basin
W :
3.56
3.50
1 .80
2.55
2.54
3.90
6.68
4.34
3.29
1 .40
3.05 :
3.13
1.93
7.64
5.56
2.20
1 .35
0.73
2.31 :
2.93 :
Ratio
C
1 .00
1 .00
1 .OO
1 .00
1 .00
1 .00
1 .00
1 .00
1 .00
1 .00
1 .00
1 .00
1 .00
1 .00
1 .00
1 .00
1 .00
1 .00
1 .00
1 .00
220
-------
Glooschenko
Sakamoto Tailing and Dobson
Eutrophic 5-140 jug/I 30 >ug/l 5-10 ;ug/l
Mesotrophic 1-15 ;ug/l 1-30 jug/I 5 jug/I
Oligotrophic 0.3-2.5 >ug/l 1 jug/I
The yearly average volume-weighted corrected chlorophyll a^ concen-
trations for each sub-basin were used to evaluate the three classification
systems. When Glooschenko and Dobson's system was applied, the west-
ern basin, Sandusky sub-basin and western central basin were found to
be eutrophic, while the eastern central basin was judged mesotrophic.
According to Sakamoto's classifications the western basin and Sandusky
sub-basin were eutrophic and the entire central basin was considered
mesotrophic. With the criteria developed by Tailing, both the western
basin and central basin were considered mesotrophic.
Of the types of investigations used to establish each of the classi-
fication systems, the Glooschenko-Dobson system, developed from using
Great Lakes data, seems to be the most applicable to Lake Erie.
Particulate Organic Carbon versus Chlorophyll a
During the last cruise of 1973 and all cruises in 1974 and 1975,
calculations and tonnages of particulate organic carbon (POC) were
determined at all stations sampled. The western basin concentrations
(Figure 84) and tonnages (Figure 85) of POC and corrected chlorophyll
a both demonstrated an increase during the summer months and a de-
c~rease in the fall. In the central basin both chlorophyll a_ and POC
decreased from the spring phytoplankton pulse through mid-summer and
increased in the late summer and early fall. During the December
cruise of 1975 an inverse chlorophyll a - POC relationship was evident
in the central basin. It was felt that as a result of the high winds
occurring in the late fall, detrital POC was resuspended from the sedi-
ments. During this period, chlorophyll a^ concentrations decreased due
to the suboptimal conditions for phytoplankton growth. Central basin
surface and bottom area-weighted concentrations of particulate phos-
phorus (PP), POC and corrected chlorophyll a_ were examined (Figures
86 and 87). It was apparent especially in the surface waters that de-
creases and increases in chlorophyll a were accompanied by corres-
ponding changes in PP and POC until December (Cruise 6).
Ratios of pheopigment to corrected chlorophyll a, corrected chloro-
phyll a to POC and PP to POC were used to examine the effect of resus-
pension on chlorophyll a concentrations (Figures 88 and 89). The spring
221
-------
l\5
ro
ro
1500
1000
o
500
LEGEND
» -Western Basin POC
-Western Basin Chi a
x-Central Basin Chi a
*Central Basin POC
|4 15 |7|8
|2 |3 |4 |5 |6
J'J'A'S'O'N'D|J 'F 'M'A'M'J 'j 'A'S 'o 'N ' D I j ' F MA M j j 'A 's 'o 'N '6T
1973 1974 1975
2O
15
10
Figure 84. Particulate organic carbon and corrected chlorophyll a
concentrations - Lake Erie central and western basins,
1973-1975.
-------
ro
PO
co
200 -
150 -
100 -
p
LEGEND * Western Basin POC
* Western Basin Chi a
Central Basin Chi a
» Central Basin PCC
o
0
Q.
'A 's 'o ' N ID JJ 'F i MI AIM' j ' j ' A1 s ]o 'N 'c | j >F 'WA 'M'J ' j 'A is 'O ' N'D
1973 1974 1975
4000
3OOO
20OO O
IB
1000 2
o
z
Figure 85. Particulate organic carbon and corrected chlorophyll a
tonnages, Lake Erie central and western basins, 1973-
1975.
-------
IN3
01
Z 0-
o Q.
12
i- 24
22
- 20
18
- 16
14
- 12
10
- 8
6
21- 4
r600
-500
U400
-300
-200
- 100
. POC
pp
\
Chla
_ , CRUISE |l
MAR I APR I MAY
|4 la
JUN j JUL | AUG | SEP | OCT NOV DEC
Figure 86. Lake Erie central basin surface water - 1975: corrected
chlorophyll a, partlculate phosphorus and partlculate or-
ganic carbon concentrations, area weighted.
-------
ro
ro
2 0. O
O.
O O.
I2p24
IOJ-20
18
8
-600
- 16
14
- 12
-500h pp
x
N.
-400
10-
- 8
- 4
L 0
-300
-200
h- 100
POC
Chlo
CRUISE I
-|%.nwTjt. [I [^ [3
MAR I APR | MAY | JUN ( JUL
|4 |S !*
"AUGI SEPI OCT|NOV|DEC
Figure 87. Lake Erie central basin bottom water - 1975: corrected
chlorophyll a, participate phosphorus and particulate or-
ganic carbon concentrations, area weighted.
-------
0.5
0.4
0.3
0.2
O.I
Ratios of: Pheophytin vs. Corrected Chlorophyll a
Corrected Chlorophyll a vs. Particulate" Organic Carbon
Particulate Phosphorus vs. Particulate Organic Carbon
(Area Weighted)
CRUISE
-10.05
0.05
0.03
0.02
POC_
-li
li.
MAR I APR ' MAY ' JUN'JUL ' AUG ' SEP ' OCT ' NOV
DEC
0.01
Figure 88. Lake Erie central basin bottom water - 1975.
o
<
0.5
0.4
0.3
0.2
O.I
Ratios of: Pheophytin vs. Corrected Chlorophyll a
Corrected Chlorophyll a vs. Particulate Organic Carbon
Particulate Phosphrous vs. Particulate Organic Carbon 005
(Area Weighted!
CRUISE 11
_LL
0.04
0.03
0.02
0.01
MAR i APR I MAY ' JUN ' JUL ' AUG '' SEP '' OCT ' NOV '
Figure 89. Lake Erie central basin surface water - 1975
226
-------
and fall were periods of the highest pheopigment to corrected chlorophyll
a ratios, indicative of a resuspension of detritus containing pheopigment.
During the same periods, the PP to POC ratio was also high. It was
assumed the PP as well as POC was resuspended from the sediments;
however, the large quantities of PP were not necessarily of an organic
nature. The chlorophyll a to POC ratio was low during the fall due to
the decrease in chlorophyll a and an increase in POC. The highest
chlorophyll a to POC ratio occurred during the late summer and cor-
responds to "to the highest standing crop in the central basin. During
this same period, the pheopigment to corrected chlorophyll a ratio was
the lowest, indicative of a larger portion of the chlorophyll a associated
with viable cells. It was evident that the correlation between corrected
chlorophyll a and POC, and to a less degree PP, was quite strong
during the sTratified period, but not during unstratified periods. This
poor correlation was attributed to resuspension of previously settled
plankters which had undergone various degrees of decomposition. The
increase of pheopigments and POC strengthens this conclusion. It is
also possible that chlorophyll a not associated with viable cells is re-
suspended. If this occurs, the biomass estimates based on chlorophyll
a and POC may be misleading during periods of extensive resuspension.
Summary and Conclusions
1. As a segment of the Lake Erie nutrient study 1973, 1974
and 1975, corrected chlorophyll and pheopigment concen-
trations were determined at 51 stations in the western
and central basins. Of the 2,332 corrected chlorophyll a
measurements taken, 98 percent of the values ranged be-
tween 0.34 and 29.12 jug/I. The mean of these observa-
tions was 7.50 /ug/l.
2. The spectrophotometric technique recommended by SCOR/
UNESCO (1966) was utilized for the analytical procedure.
Concentrations of chlorophylls a, b_ and £ were calculated
according to SCOR/UNESCO along with corrected chloro-
phyll a^ pheopigment a^ and the Fo/Fa ratio described by
Lorenzen (1967). Only corrected chlorophyll a and pheo-
pigment £* were discussed in detail.
3. The quantitative effects of basin, station, cruise, sampling
depth and stratification have been determined with analysis
of variance models. Of these variables, basin was con-
sidered a first order effect, while station within basin and
cruise had second order effects.
227
-------
4. The seasonal trends of chlorophyll a differed in the
western and central basins. Due to different physical,
chemical and biological factors influencing these two
basins, they exhibit different seasonal patterns. The
western basin maintained highest chlorophyll a con-
centrations during the summer months, while~the cen-
tral basin exhibited two peaks, one in the early spring
and the other following turnover in the late summer
(early fall).
5. The June to November mean concentration of corrected
chlorophyll a was shown to increase yearly from 1973 to
1975. The maximum increase in tons for the central
basin was 38 percent, which occurred in 1975, while the
maximum increase for the western basin of 44 percent
was observed in 1974.
6. Long-term chlorophyll trends are difficult to evaluate.
Lake Erie chlorophyll surveys have been undertaken
by the Federal Water Pollution Control Administration
(FWPCA, 1968) in 1967 and 1968 and by Glooschenko
in 1970 (Glooschenko, et al. 1974). The FWPCA
survey substantiated the year to year chlorophyll in-
creases observed in the 1970's. The increase in
chlorophyll a concentration over the nine-year period
(June to November 1967 to 1975) was 7.08 ug/l or
146 percent. Due to an apparent analytical problem
the FWPCA data was of questionable reliability. When
Glooschenko's 1970 survey was compared with the cor-
responding CLEAR survey a 38 percent increase in the
central basin chlorophyll a concentrations was estimated
between 1970 and 1975. Thirty-three percent of the in-
crease occurred between 1974 and 1975. The western
basin showed an estimated 22 percent increase between
1970 and 1975, with the greatest Increase observed from
1974 to 1975.
7. The western basin and the western half of the central
basin were considered eutrophic based on criteria of
the chlorophyll trophic classification system established
by Glooschenko and Dobson (1975). The eastern half
of the central basin was considered mesotrophic.
228
-------
8. The relationships of participate organic carbon to
chlorophyll a and pheopigment to chlorophyll a
were examined during 1975, in order to evaluate
the effects of resuspension. Particulate organic
carbon and pheopigments were found to increase
during periods of maximum resuspension, while
chlorophyll concentrations decreased. This would
indicate that during the spring and fall periods of
high winds, detrital material is resuspended con-
tributing to the particulate organic carbon and
pheopigments in the water column.
9. The drawbacks involved in the chlorophyll technique
have been discussed throughout the text. Errors
involved in the analytical portion of the technique
have been calculated. Measurement errors are
negligible, and error due to extended periods of
storage present the greatest analytical error.
Determination of chlorophyll concentrations from
samples freeze stored for more than two months
may yield concentrations that underestimate the
actual values.
In calculating volume-weighted tons of chlorophyll
it was assumed that the horizontal distribution of
chlorophyll was consistent within a grid. Patchi-
ness of phytoplankton does occur, thereby resulting
in patchy chlorophyll concentrations. Error in
chlorophyll analysis as a result of patchiness seems
inevitable but has not been quantified.
REFERENCES
Brydges, T.G. 1971. Chlorophyll a Total phosphorus relationships in
Lake Erie. Proc. 14th Conf. Great Lakes. Res. 185-190.
Federal Water Pollution Control Administration. 1968. Lake Erie Sur-
veillance data summary 1967-1968. U.S. Dept. Interior. 65 p.
Glooschenko, W.A. and H.F.H. Dobson. 1975. Water quality in the
Great Lakes. Nature Canada. 4:3-6.
229
-------
Glooschenko, W.A., J.E. Moore and R.A. Vollenweider. 1974. Spa-
tial and temporal distribution of chlorophyll a and pheopigments in
surface waters of Lake Erie. J. Fish. ResT Bd. Can. 31:265-274.
Harvey, H.W. 1934. Measurement of phytoplankton population. Mar-
ine Biology Assoc. U.K. 19:761-773.
International Joint Commission. Great Lakes Water Quality Board.
1975. Great Lakes water quality, 1974 annual report, appendix B.
Lorenzen, C.J. 1967. Determination of chlorophyll and pheo-pigments
spectrophotom etr ic equations. Limnol. Oceanogr. 12:343-346.
Odum, H.T., W. McConnell and W. Abott. 1958. The chlorophyll a
of communities. Publ. Inst. Mar. Sci. Texas 5:65-96. ~
Richards, F.A. and T.G. Thompson. 1952. The estimation and chai
acterization of plankton populations by analyses II: a spectrophoto-
metric method for the estimation of plankton pigments. J. Mar.
Res. 11:156-172.
Sakamoto, M. 1966. Primary production by phytoplankton community
in some Japanese lakes and it's dependence on lake depth. Arch.
Hydrobiol. 62:1-28.
SCOR/UNESCO. 1966. Monograph on oceanographic methodology I.
Determination of photosynthetic pigments in sea water. Paris. 69 p,
Strickland, J.D.H. and T.R. Parsons. 1968. A practical handbook of
seawater analysis. Fish. Res. Bd. of Can. Bull. 167 p.
Tailing, J.F. 1961. Photosynthesis under natural conditions. Ann.
Rev. Plant Physiol. 12:133-154.
Thomas, R.L., J.-M. Jaquet and A.L.W. Kemp, 1976. Surficial
sediments of Lake Erie. J. Fish. Res. Bd. Can. 33:385-403.
Tucker, A. 1949. Pigment extraction as a method of quantitative
analysis of phytoplankton. Amer. Micros. Soc. Trans. 68:21-33.
Verber, J.L. 1950. Percent and area of contour levels of Lake Erie.
mimeo. 4 p.
Vollenweider, R.A. 1971. Scientific fundamentals of the eutrophication
of lakes and flowing waters, with particular reference to nitrogen
and phosphorous as factors in eutrophication. Organization Econom.
Coop. Develop. Paris. 159 p.
230
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SECTION 8
PRIMARY PRODUCTIVITY SURVEY OF THE CENTRAL
AND WESTERN BASINS OF LAKE ERIE
Clifford T. Sheffield
Center for Lake Erie Area Research
The Ohio State University
Walter E. Carey
Department of Nuclear Engineering
The Ohio State University
INTRODUCTION
Within the last twenty years biologists have observed that man has
increased the nutrient input of Lake Erie to cause significant changes in
its flora and fauna (Beeton and Edmondson 1972). From 1938-1964, to-
tal phytoplankton concentration in the island area increased by 3.5 times,
and the dominant phytoplankton changed from diatoms to blue-green algae.
From 1920-1962 in central Lake Erie, phytoplankton numbers increased
several fold during spring and autumn pulses.
Primary productivity is a measure of the rate of buildup of organ-
ic compounds from the energy that is transferred to successive trophic
levels. Ths photosynthetic process is responsible for the greater part
of this buildup in most environments (Goldman 1963). Estimates of pri-
mary productivity for Lake Erie first appeared in 1949, when Verduin
(1951) measured the rates of CO2 removal per unit of phytoplankton
volume per hour in western Lake Erie. Verduin concentrated phyto-
plankton by passing lake water through bolting cloth and placed these sam-
ples in bottles under an optimal light intensity of 4OO foot candles (fc).
However, he estimated that about two-thirds of the photosynthetic organ-
isms escaped concentration (Verduin 1956). Consequently, since 1957 he
has used natural phytoplankton densities and has estimated rates of Co2
removal in open-lake conditions. He found that under these conditions
photosynthetic rates exceed those rates obtained from bottle experiments
by two times (Verduin 1960 and 1962). For these reasons his more
recently published rates of 3070 mg absorbed per m per day for net
231
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photosynthesis and 616O mg C absorbed per m2 per day for gross photo-
synthesis are used for comparison to the results of subsequent studies,.
Saunders (1964) made the first estimates of primary productivity
in Lake Erie with 14c at a. single station in the western basin in 1957;
estimates were 68.3 mg C/m3 per day for net and 142.9 mg C/m3 per
day for gross photosynthesis. He converted these estimates to 97.6
and 371.5 mg C/m per day, respectively, for net and gross daily
integral photosynthesis by using a formula derived by Rodhe, Vollen-
weider and Nauwerck (Saunders 1964).
Further primary productivity work was done on Lake Erie in 1968
when Porkso et al. (1969) took a one-day cruise along the entire lake
to make the first estimates of primary productivity for central Lake
Erie. With the 14C technique and a shipboard incubator (light intensity
= 1000 foot candles, temperature = 60°F), Parkos et al. (1969) found
surface rates of 127.7 mg C/m3 per hour at one station in the western
basin and 42.4-54.7 mg C/rr>3 per hour at two stations in the central
basin. In 1969 and 1970, Cody (1972) used 14C and found extremely
variable rates, from 11-5470 mg-atoms C [132-65,640 mg C] per m2
per day, in western Lake Erie during m situ studies at ten stations.
Glooschenko et al. (1974) made intensive measurements in Lake Ontario
and all three basins of Lake Erie (25 stations) by calibrating a ship-
board incubator (light intensity = 8000 lux, temperature = surface water
temperature) with simultaneous in situ experiments. Surface water
productivity obtained from the incubator for each station was converted
to an in situ estimate for the entire water column. Mean values for
ten cruises in 1970 were 30-4760 mg C/m2 per day and 120-1690 mg
C/m per day for the western and central basins respectively.
This report includes data for the central and western basins of
Lake Erie that were obtained during July-October 1974. The objectives
of the study were to investigate, by means of the 14c method and ship-
board incubation of samples, the vertical, horizontal and temporal dis-
tribution of primary productivity in central and western Lake Erie and
to compare data obtained in 1974 with data from previous investigations.
Samples were taken from five western basin stations and eleven central
basin stations during cruises abourd the R/V Hydra-26 July-6 August
(Cruise 6), 12 August-19 August (Cruise 7), 26 August-7 September
(Cruise 8), and 21 October-1 November (Cruise 10, Figure 90).
Between cruises, a location approximately 250 m offshore, south
of The Rattles of Rattlesnake Island in western Lake Erie was chosen
for in situ experiments. Cody (1972) reported ranges of 16.7-185.4
232
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CO
CO
R - .11. of jg glU productivity eKp.rlm.nt.
near Rattlesnake Icland
Detroit
[Cleveland
Figure 90. Map of Lake Erie showing station locations in the central
and western basins.
-------
mg-atoms C [200-2225 mg C] per m3 per hour, 36.1 - 654.9 mg-atoms
C [433-7859 mg C] per m2 per hour, and 240-5470 mg-atoms C [288O-
65,640 mg C] per m2 per day for optimal, integral, and daily integral
primary productivity, respectively, at Rattlesnake Island in 1970.
PROCEDURES
Field Methods
At each station submarine photometer readings were taken at one-
meter intervals from lake surface to bottom, or until the light meter
read zero. The reading made when the photometer was suspended just
above the water represented surface light intensity. Samples of lake
water were collected from one meter and from a depth of about one per-
cent of the surface light intensity, which represented the lower limit
of the euphotic zone. Samples were also collected from intermediate
levels (commonly three meters and five meters), depending upon the
thickness of the euphotic zone.
Sample preparation follows the method of Goldman (1963). A sam-
ple of lake water, taken with a five-liter Niskin sampling bottle, was
transferred to 300-ml BOD bottles, two light and one dark for each
depth. The dark bottles were wrapped with a double layer of black
electrical plastic tape. A layer of aluminum foil covered the tape to
prevent heat buildup. Two layers of foil covered the stopper to keep
out light. Between cruises both dark and light bottles were washed
with concentrated HNOg and thoroughly rinsed with tap water with a
final rinse of distilled water. The bottles were inverted and air dried.
A fresh stock solution of 14C was prepared daily by combining the
contents of ampules of sodium bicarbonate - 14C (NaH14CO3) in sterile
aqueous solution with pH = 9.5, which was supplied in the amount of
5.0 pd/5.0 ml per ampule by New England Nuclear Corp. Approxi-
mately 1.0 ml of the stock solution was added to each BOD bottle by
means of a 2-ml hypodermic syringe (accuracy = + 5 percent) with
a 15-cm 18-gauge needle. The stock solution was added to the bottom
of the bottle, and the syringe was rinsed gently with water from the
top of the bottle. The activity of the NaH14COs, added via the syringe,
was taken to be 1.0 jjd (2.22 x 106 + 10 percent disintegrations per
minute) as determined by New England Nuclear Corporation.
After innoculation, the bottle contents were thoroughly mixed, and
the bottles were placed in a shipboard incubator for three to five hours.
Light intensity of approximately 9OOO lux was provided by a bank of six
234
-------
daytime fluorescent lamps. Surface water was continuously circulated
through the incubator. After incubation the bottle contents were mixed,
10 ml of material was removed with a repipet, and 10 ml merthiolate
solution was added to stop further 14C uptake. The bottle contents were
thoroughly mixed.
Since the relative activity of the plankton was unknown when exper-
iments were begun, the investigator concentrated as much material as
possible from a bottle onto a membrane filter. As more material is
added, sample counting time decreases linearly, unless the additional
material decreases detection of the -particles emitted from 14C. In
areas with high concentrations of suspended particles, filter pores
clogged within approximately ten minutes. In some areas (e.g., Sta-
tions 30, 47, and 73), the entire bottle content was passed through the
filter within ten minutes; in other areas (notably Stations 61, 75, and
76) aliquot was used since the filtration time for the entire content
would have exceeded ten minutes. Material from each BOD bottle was
vacuum-filtered onto a 25-mm diameter Gelman GA-6 Metricel membrane
filter (pore size 0.45 u). The filtration apparatus aboard the R/V Hydra
consisted of a vacuum manifold that held four filter funnels so that the
contents of as many as four BOD bottles could be filtered simultaneously.
Vacuum pressure did not exceed 300 mm Hg. Since the membrane fil-
ters differ in rates of water absorption, all filters were presoaked in
distilled water to decrease filtration time. Once the material had been
concentrated on the membrane filter, the filter was rinsed with approxi-
mately 25 ml of 0.003 N HCl to remove any 14C that may have accumu-
lated on the surface of the filtered material; a rinse of 25 ml filtered
lake water (if available) or distilled water followed. The membrane
filter was then attached with household glue to a 32-mm diameter stain-
less steel planchet. Water retained by the filtered material absorbs
the -particles emitted from 14C, so filters were stored in pillboxes
and assayed several weeks later.
Radioassay
Filters were assayed with a thin end-window gas-flow detector;
gas composition was approximately O.9 percent isobutane in 99.1 pei
cent helium. Planchets were loaded onto a Nuclear Chicago Corp.
Model No. C-110B automatic sample changer. A Nuclear Chicago
Corp. Model 181A sealer recorded the counts per measurement, and
a Nuclear Chicago Corp. Model No. C.111B printing timer recorded
the time required for the measurement. A 0.0875 AJCI14C standard
235
-------
source was used to calculate counter yield by the formula of Arena
(1971):
Y = cpm/dpm,
where Y is counter yield, cpm is observed count rate of the standard
(counts per minute), and dpm is the calibrated activity of the standard
(disintegrations per minute). Counter yield was 16-17 percent. Light
bottles were counted a minimum of 10,000 counts (+2.1 percent pro-
bable error at P = 0.05) and dark bottles a minimum of 1OOO counts (+
(+6.4 percent probable error at P = 0.05). One thousand counts per"
measurement for dark bottles was chosen to reduce sample counting
time to about ten minutes. The discrepancy between light and dark
bottle counting statistics is discussed later.
Primary Productivity Calculation
Incubator productivity was calculated by an expanded formula
developed by Saunders, et al. (1962). Dark bottle incubator proauc-
tivity was subtracted from light bottle incubator productivity to correct
for nonphotosynthetic uptake of 14C. The corrected incubator produc-
tivity (mg C/m h) was converted to integral productivity (mg C/m2. h)
by the relationship reported by Glooschenko et al. (1974):
mg C/m2th _
mg C/m3.h = 1-85x>
where mg C/m2.h is integral productivity, mg C/m3.h is incubator
productivity, and x is Secchi disc depth in meters; the relationship is
solved for mg C/m2*h. Since the incubator productivity of Glooschenko
et al. (1974) is based on a composite sample from one-meter and five
meter depths, incubator productivities from one-meter and five-meter
depths in this report were averaged, and the mean value was inserted
into the above relationship. Integral productivity was converted to daily
integral productivity (mg C/m2« day) by the method of Glooschenko et
al. (1974).
In situ Productivity
Procedure for in situ productivity experiments near Rattlesnake
Island was similar to the shipboard method except incubation. Photo-
meter readings and Secchi disc depth were determined as previously
described. Temperature was measured by means of a YSI Model 51A
dissolved oxygen meter equipped with a YSI 5450 probe. Lake water
236
-------
was collected with a three-liter non-metallic Kemmerer water sampling
bottle. For the one-meter depth, one intermediate depth, and the bot-
tom sampling depth, pH was measured in the laboratory on South Bass
Island within two hours of collection, and total alkalinity was determined
within four hours of collection. Upon innoculation with one microcurie
of NaH14CO~, one light and one dark bottle were suspended in the lake
at the depth from which the water was taken. The incubation appara-
tus consisted of an anchored nylon line with an attached float on the
lake surface. Aluminum strips were tied to the line at the proper
depths. The BOD bottles were attached to the strips by shower cui
tain hooks that fastened to hose clamps at the necks of the bottles. If
possible, the samples were fixed when the bottles were retrieved from
the line; when the weather was rough, they were fixed at the laboratory
approximately half an hour after retrieval. Samples were filtered at
the South Bass Island laboratory or aboard the R/V Hydra, and the
membrane filters were prepared for gas-flow counting.
ln_ situ primary productivity (mg C/m3- h) at each sample depth
was calculated. These estimates were then plotted against depth on
graph paper. Integral in situ productivity (mg C/m h) was calculated
by determining the area beneath the productivity-depth curve with a
planimeter.
Correction Factors
The filtered material absorbs energy from the 3 -particles emitted
from 14C and prevents the escape and subsequent detection of some of
the particles by counting equipment. Lower sample count rate and,
therefore, lower productivity result. An experiment was performed to
determine whether this self-absorption of 3-particles occurred in fil-
tered material during the 1974 season. Lake water from a depth of
one meter in Put-in-Bay Harbor was collected with a Kemmerer water
sampling bottle on 5 October 1974. The water was transferred to six
clear BOD bottles, and approximately one microcurie of NaH14CO3 was
was added to each bottle. After the contents were thoroughly mixed, the
the bottles were incubated approximatly four hours in the shipboard
incubator. After incubation the contents were fixed as previously des-
cribed. Aliquots of 25, 50, 75, and 100 ml from each bottle were
filtered onto membrane filters as described. Because filtration of the
100-ml aliquots required approximately ten minutes, these volumes
were assumed to represent the denser concentrations of material
collected over the season. Filters were rinsed and processed for gas-
flow counting as in the productivity experiments. Filters were counted
10,000 counts minimum. Sample count rate was corrected to actual
237
-------
C uptake. Carbon-fourteen uptake (dpm) was divided by aliquot
volume (ml) and duration of incubation (h), and the result (dpm/ml
per hour) was plotted against aliquot volume.
Strickland and Parsons (1968) stated that the fixative formalde-
hyde at a concentration of 0.3 percent volume may influence the
excretion or loss of fixed 14C from the more delicate marine algae.
The effect of merthiolate upon sample activity was studied, as mer-
thiolate may also cause 14C loss from phytoplankton. Lake water
collected on 16 September from one meter below the surface of
Put-in-Bay Harbor was added to six clear 300-ml BOD bottles.
About one microcurie NaH14CO>3 was added and th° bottles were
incubated approximately four hours in the shipboard incubator.
Samples were fixed as in the productivity experiments. At intervals
of 1, 2, 4, 8, 16, and 24 hours after- fixation 5O-ml ali.quots from
each BOD bottle were filtered onto membrane filters. The filters
were prepared for gas-flow counting. The experiment was run again
on 19 October 1974, and aliquots were filtered at intervals of 0, 1,
2, 4, 8, and 16 hours after fixation. During both experiments bot-
tles were stored overnight and filtered the next day. Filters were
counted 10,OOO counts minimum. Graphs were drawn of carbon-fourteen
uptake per ml per hour vs. time after fixation for the twelve bottles.
Relative light intensity in the incubator was measured while
the research vessel was docked at Put-in-Bay on 19 September 1974.
The incubator was divided into nine horizontal zones, and several
photometer readings were taken at each zone to represent light in-
tinsity at clear water, mid-lake central basin stations, as the water
had been standing for several days. Photometer readings were re-
peated after the water had been circulated for two hours, which
approximated turbid stations. Whether or not relative light intensity
was reduced by use of a clear plexiglass top when the lake was
rough was also determined. A mean relative light intensity value
for any position in the incubator during each set of conditions was
determined by averaging individual measurements at each location
in the incubator.
238
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RESULTS AND DISCUSSION
Incubator, integral, and daily integral productivity in 1974
were, respectively, 4.1-124 mg C/m3 per hour, 16-364 mg C/m2
per hour, and 140-4370 mg C/m2 per day for western Lake Erie
and 5.1-51.2 mg C/m3 per hour, 44-242 mg C/m2 per hour,
and 53O-2180 mg C/m2 per day for central Lake Erie (Table 63).
Results of calculations using Glooschenko's relationship are present-
ed in Table 64.
Maximum incubator productivity in 1974 agrees to within 3
percent of that reported by Parkos et al. (1969) and 16 percent of
the maximum mean productivity per cruise reported by Vollenweider
et al. (1974) for western Lake Erie (Table 63). Maximum central
basin incubator productivity in 1974 is 2.4 times greater than the
mean productivity per cruise reported by Vollenweider, but the dif-
ference may be in the comparison of mean values per cruise to a
range of values for the entire season of 1974. Parkos et al. (1969)
report productivity 7 percent greater than the maximum incubator
productivity found during this study.
Daily integral productivity reported by Saunders (1964) is
relatively low when compared with other western basin estimates
in Table 63, but this probably is explained by his sampling one site
late in the year. The lower limit of Verduin's estimates fall with-
in the daily integral productivity range calculated in 1974. Cody's
(1972) mean estimate of 10,OOO mg C/m2 per day is 2.3 times
greater than the maximum daily integral productivity in the west-
ern basin in 1974.
Central Lake Erie incubator productivity for late July averaged
1.7 times mid-August values and 0.7 times late October values
(Table 65). Nutrient release upon fall overturn (Gachter et al. 1974)
probably resulted in the October maximum. Mean incubator produc-
tivity for western Lake Erie decreased from late July to late October
with the July value being 1.4 and 1.6 times greater than September
and October values respectively. Integral productivity for central Lake
Erie followed, the trend of incubator productivity with the late July value
being 1.2 times the August value and 0.9 times the October value.
Western Lake Erie integral productivity in July was 2.4 and 1.7
times September and October integral productivity, respectively.
239
-------
TABLE 63. COMPARISON OF PRIMARY PRODUCTIVITY MEASUREMENTS IN
CENTRAL AND WESTERN LAKE ERIE FROM 1957-1974
Date
1957-1962
1957 Oct.
1968 July
1969-1970
1970 Api Dec.
1974 July-Oct.
1968 July
1970 Apr-Dec.
1974 July-Oct.
Method
pH-CO2
4 A
14C
14
C
14
14C
14
C
14
C
4 A
14C
14C
14
C
\ A
14C
Incubation
natural conditions
In situ
shipboard
_ln situ
[_n_ situ, shipboard
In sttu
shipboard
shipboard
^n_ situ, shipboard
shipboard
mg C/m3.h mg C/m .h
Western basin
127.7
13.2-2225 18.0-7859
4 . 8- 1 46 . 9a 5 . 0-397a
3.6-152 95-342
4.1-124b 16-364,
Central basin
42.4-54.7
5.5-21.4a 17-141a
5.1-51.26 44-242
mg C/m .day
net 3,070
gross 6, 16O
net 96.7
gross 371 .5
132-65,640
mean 10,OOO
30-4, 760a
140-4,370
1 20-1 , 69Oa
530-2,180
Source
Verduln (1962)
Saunders (1964)
Parkos, Olson &
Odlaug (1969)
Cody (1972)
Vollenwelder, Muns
war & Stadelmann
(1974)
Sheffield (1975)
Parkos, Olson &
Odlaug (1969)
Vollenwelder,
Munawar, & Stadel-
man (1974)
Sheffield (1975)
a range of mean values per cruise
b range of station means for productivity within cuphotlc zone
-------
TABLE 64. CARBON ASSIMILATION RATES BY STATION
AND CRUISE FOR CENTRAL AND WESTERN LAKE ERIE IN 1974
Mean Incubator Rate3 Integral Rate Daily Integral Rate
Date Station
1 Aug 30
30 July 47
4 Aug 48
27 July 52
26 July 6l
28 July 66
27 July 67
30 July 73
26 July 75
26 July 76
13 Aug 23
13 Aug 25C
14 Aug 30
15 Aug 34C
15 Aug 3^
17 Aug 44
12 Aug 47
16 Aug 48
6 Sept 53
7 Sept 6l
6 Sept 67
7 Sept 75
7 Sept 76
23 Oct 23
23 Oct 25
26 Oct 30
26 Oct 32
27 Oct 36
27 Oct 39
29 Oct 44
21 Oct 47
28 Oct 48
30 Oct 53
1 Nov 6l
29 Oct 66
30 Oct 67
21 Oct 73
l NOV 75
31 Oct 76
a average of all sample
b incubator rates from
found in Glooschenko
c includes sample from
d includes sample from
(ng C/m^.h + SE)
Cruise 6
19.8 + 1.5
16.4 7 1.6
11.1 7 1.2
20.8 7 3.3
12.4 7 1.3
33.0 7 3.1
33.5 + 1.4
17.3 + 2.2
124 7 6
98 7 2
Cruise 7
8.3 + 0.9
5.1 7 0.4
8.1 7 0.9
10.1 7 1.0
7.271.3
13.6 + 0.8
14.0 7 0.7
15.7 7 0.9
Cruise 8
33.3 + 2.2
30.4 + 1.1
31.0 7 1.6
32.3 7 2.3
76.8 + 2.6
Cruise 10
23.1 + 0.9
14.5 + 0.5
18.4 7 1.1
10.9 7 0.6
18.6 7 1.0
17.0 ~ 1.2
51.27 1.2
29.9 7 1.6
23.2 7 0.9
34.8 7 0.9
4.1 + 0.1
28.1 7 1.0
46.0 7 1.4
30.371.7
77.6 + 2.4
26.4+ 1.2
depths.
1 m and 5 m depths were averaged
et al (1974)
metalimnion.
hypolimnion.
2
(mg C/m .h)
130
180
83
109
37
194
157
133
364
166
94
84
84
82
73
124
154
135
44
87
75
57
85
87
82
80
69
160
60
242
216
172
123
16
136
141
223
171
73
and the mean
2
(mg C/m .day)
1560
2160
1000
1310
440
2330
1880
1600
4370
1990
1130
1010
1010
980
880
1490
1850
1620
530
1040
900
680
1020
780
740
720
620
1440
540
2180
1940
1550
1110
140
1220
1270
2010
1540
660
value inserted into the relationship.
241
-------
TABLE 65. MEAN PRODUCTIVITY BY BASIN FROM JULY-OCTOBER 1974
No. of
Stations
Cruise 6
5
5
Cruise 7
8
Cruise 8
4
Cruise 10
11
5
Basin
central
western
central
western
central
western
Incubator
Productivity
(mg C/m3.h + SE)
17.1 + 1 .7
60 +22
10.2 + 1 .3
42.6 + 11 .4
24.7 + 3.4
36.4 +12.3
Integral
Productivity
(mg C/m2 .h)
127
184
104
76
138
107
Daily
Integral
Productivity
(mg C/m2. day)
1530
2200
1250
910
1240
96O
ro
-fa.
ro
-------
Horizontal distribution of incubator productivity follows that of
previous investigators. Parkos et al. (1969) and Glooschenko et al.
(1974) report a west to east gradient in Lake Erie with the highest
productivity in the western end. A north to south gradient in both
basins during 1970 (Glooschenko et al. 1974) is not quite as evident
during 1974. The maximum productivity in mid-summer in western
Lake Erie reported by Glooschenko occurred in 1974. However,
upon examination of his monthly distribution maps (Glooschenko
et al. 1974, Figure95), one finds that the estimates of 124 mg
C/m3- h reported in July 1974 for Stations 75 and 76 are nearly
double those reported for the summer 1970.
Results of three in situ experiments near Rattlesnake Island
show that when productivity"^ mg C/nrP-h) is plotted against relative
light intensity, three distinct curves result (Figure 91> Primary
productivity on 10 September remained linear through a relative
light intensity at least 2.5 times greater then that for 21 July and
6 October. In October, when surface light intensity and water
temperatures are lower, phytoplankton survive at lower light inten-
sities than in July and September. Primary productivity in October
leveled off more rapidly than in July and September.
For each of the three in situ experiments the primary produc-
tivity vs. depth curve has a slope similar to its corresponding
light transmission curve (Figure 92> Maximum productivity occur-
red at approximately one meter. Cody (1972) reported maximal
productivity in western Lake Erie from one to three meters and
rarely at depths shallower than one meter.
For a series of increasing volumes of filtered phytoplankton,
self-absorption of 3-particles from 14C exists if a decrease in
dpm/ml per hour occurs. Results of the self-absorption experiment
(Figure 93) indicate that there was no decrease in dpm/ml per hour,
although results from individual bottles varied. No correction of
sample count rate was made on the basis of this evidence.
Effect of merthiolate fixation on sample activity experiments
indicate that loss of activity occurs non-linearly for approximately
the first eight hours (Figure 94> Approximately 14 percent loss
occurs during the first four hours, another five to seven percent
during e§ht additional hours. On the average, approximately 21
243
-------
200
160
o
|120
o
I 80
40
o 21 July 1974
10 Sept 1974
A 6 Oct 1974
10
15
20
25
30
Relative Light Intensity (juA x 10 )
Figure 91. Relationship of in situ productivity to relative light
intensity near Rattlesnake Island.
-------
ro
-t=.
tn
% Surface Light Intensity
10
light transmission
A 21 July
o 10 Sept
6 Oct
0.1
Figure 92.
1.0 10
Relative lin Situ Productivity
Relationship of relative in situ productivity and light
transmission of depth near Rattlesnake Island.
100
-------
20
18
14
I
12
ERROR BARS REPRESENT ±SE
25 50 75
Aliquot Volume (ml)
100
Figure 93. Relationship of sample activity to aliquot volume.
ERROR BARS REPRESENT ± SD
12 16 20
Time After Rxation (h)
Figure 94. Relationship of sample activity to time after fixation.
246
-------
percent of the original activity is lost per sample within eight hours
after fixation. Again individual bottles vary in loss rates. Hourly
loss rates become similar after eight hours.
Relative incubator light intensity in clear water averaged
900 y A (range of 775-1075 yA) and 775 ~v A (range of 675-975 yA) in
turbid water. When the clear plexiglass top covered the top of the
incubator during rough weather, relative light intensity in clear
water was reduced from 900 to 825 y A (range of 725-925 yA), a
reduction of 9 percent. Since no records were kept of the horizon-
tal position of the bottles in the incubator, no correction was applied
to the carbon assimilation rates.
The estimated average range of total error of a single produc-
tivity measurement determined by the methodology presented in
this report is +18 percent to -32 percent. Counting statistics,
syringe calibration, ampule activity calibration and fixation effects
were factors considered. Dark bottle 14C uptake and light bottle
14C uptake can both be expressed in relative units in which light
bottle uptake equals 100 and dark bottle uptake its corresponding
percentage. From such consideration the relative dark uptake
(49 percent) that yields the greatest uncertainty to the net sample
count rate contributes a total probable error in counting statistics
of 10 percent (P = 0.05). Mean relative dark bottle uptake of
14 percent (n = 182) yields a probable error of 3 percent (P = 0.05),
and thus the mean probable error caused by counting statistics
alone is +3 percent. Syringe accuracy contributed+5 percent error.
Accuracy of ampule calibration by New England Nuclear Corp. as
determined by ampule lables and technical information is assumed
to be +10 percent of the stated value of 5.0 yCi per 5.O ml
[1.0 PCi/1.0 ml] aqueous solution. Samples were assumed to have
been filtered within four hours after fixation. Therefore, samples
lost 14 percent of their original activity (Figure 94, curve B). This
estimate is probably conservative since filtration of some samples
was not completed for at least 24 hours. No accurate records
were kept of the time between sample fixation and filtration, as the
fixation effect was not discovered until about two-thirds of the samples
had been taken through the sampling season. For this reason primary
productivity measurements remain uncorrected for sample activity
loss. The fixation effect contributes the greatest uncertainty to the
methodology used during this study.
247
-------
Agreement between duplicate light bottles (n = 183 pairs) av-
eraged 21 percent +2 percent (SE) for all samples over the entire
season. This measured value, falls within the predicted range of
+ 18 percent to -32 percent. Saunders et al. (1962) reported that
+20 percent is the normally accepted error for estimates of photo-
synthesis. The positive side of the estimated error for 1974 is
10 percent less than Saunders' figure, while the negative side is
60 percent greater, since sample fixation causes loss in addition to
the three other sources of error considered. The amount of error
introduced by the relationship of Glooschenko et al. (1974) is un-
known. Error bars in figures accompanying the text represent the
estimated error of +18 percent to -32 percent unless otherwise
indicated.
Graphs of the vertical distribution of incubator productivity
within the euphotic zone reveal that the wide range of error may
reduce productivity maxima and minima of raw data. For example,
the graph of incubator productivity vs. depth for Station 47 on
30 July (Figure 95) shows that peaks at 1 m, 5 m, and 14 m reduce
to a small increase (approximately 20 percent) from 1 m to a
plateau at about 7 m. Superficially, a doubling of incubator pro-^
ducttvity is observed from 2 m to 5 m and from 7 m to 14 m.
Vertical profiles where maximum productivity was £ 1 .32 times the
minimum productivity were compared. Maximum to mintmum pro-
ductivity ratios _> 1.32 were considered significant, since -32 per-
cent was the average maximum error involved in making the
productivity estimates. The number of significant productivity
profiles was 15 of 39 or 38 percent of the total incubator produc -
tivity experiments.
Central Lake Erie incubator productivity profiles had a wider
range of carbon fixation rates than western Lake Erie profiles.
Maximum productivity to minimum productivity ratios for central
Lake Erie averaged 1.80, 1.83, and 1.17 for early August, mid-
August, and late October, respectively. SAfestern Lake Erie ratios
averaged 1.33, 1.16, and 1.07 for late July, early September, and
late October, respectively. For central Lake Erie at Station 23
during both mid-August and late October maximum productivity at
1 m decreases with depth (Figure 95> The profile at Station 25 in
mid-August showed increasing productivity from about 5 m to a
plateau at 15 m to the bottom of the euphotic zone at 18 m (Figure
248
-------
^10
a
0)
D
15
20
10
20
Incubator Productivity (mg C/m3-h)
30 40 n 4
8
12
16
-i
STA 47
30 JULY 1974
METALIMNION
STA 23
13 AUG 1974
BOTTOM
Figure 95. Vertical profile of incubator productivity at station 47 on
30 July 1974 and station 23 on 13 August 1974.
-------
96). For Station 30 in early August incubator productivity increased
linearly from 1 m to 11.5 m (Figure 96); in mid-August productivity
remained the same from 1 m to 4 m, where it increased and reached
a plateau at 15 m in the metalimnion (Figure 97). Station 34 in mid-
August had a profile similar to Station 30 in Figure 97 although the
euphotic zone extended to only 11 mm. Station 36 in mid-August had
a nearly linear profile sinilar to Station 30 (Figure 96) except it started
at 3 m. Station 39 on late October had minimum productivity at 5 m
which increased at 7 m to nearly the level at 1 m (Figure 97). Pro-
ductivity at Station 48 in early August had a maximum at 4 m and
decreased to nearly the same level as at 1 m (Figure 98); in mid-
August productivity remained nearly the same for the first five meters,
increased slightly at 9 m, and it decreased to a minimum level at 13
m (Figure 98). Productivity at Station 52 in late July had a sharply
defined maximum at 3 m (Figure 99). In mid-lake areas phytoplahkton
accumulated in the lower epil imnion and metal imnion. Station 30
(Figures 96 and 97) showed this trend in incubator productivity pro-
files. The profile for 14 August is supported by chlorophyll a (uncor-
rected) data (Table 66). Productivity at Station 36 on 15 Aug~ust
showed a marked increase toward the metalimnion. However, chloro-
phyll a. concentration and productivity do not agree.
TABLE 66. VERTICAL DISTRIBUTION OF UNCORRECTED
CHLOROPHYLL a CONCENTRATION AT STATIONS
30 AND 36 IN AUGUST 1974
Station 30
Depth
(m)
1
14
16
20
14 August
Concentration
(mg/m3)
2.7
3.9
3.8
3.5
Station 36
Depth
(m)
1
16
19
23
15 August
Concentration
(mg/m3)
3.9
3.9
3.6
5.5
250
-------
£10
Q.
0)
Q
15
8
Incubator Productivity (mgc/rrf3- h)
12 16 n 10 20
STA 25
13 AUG 1974
20
METALIMNION
30
40
STA 30
1 AUG 1974
Figure 96. Vertical profile of incubator productivity at station 25 on
13 August 1974 and station 30 on 1 August 1974.
-------
INJ £10
en QL
ro fli
15
20
8
M ETA LIMN I ON
Incubator Productivity (nig C/nr'-h)
J2 16 Of 10
STA 30
14 AUG 1974
20
BOTTOM
30
40
STA 39
27 OCT 1974
Figure 97. Vertical profile of incubator productivity at station 30 on
14 August 1974 and station 39 on 27 October 1974.
-------
ro oi
en _ '
co -E
Q.
0)
Q
15
20
10
p.
20
i
l ' l
Incubator Productivity (mg C/m3- h)
30
~T~
40
STA 48
4 AUG 1974
METALIMNION
10
20
30
40
STA 48
16 AUG 1974
METALIMNION
Figure 98. Vertical profile of incubator productivity at station 48 on
4 August 1974 and 16 August 1974.
-------
-t=> 0)
o
15
20
10
20
Incubator Productivity (mg C/m3- h)
_30 40 n^ 10
STA 52
27 JULY 1974
BOTTOM
20
30
STA 73
30 JULY 1974
METALIMNION
Figure 99.
Vertical profile of incubator productivity at station 52 on
27 July 1974 and station 73 on 30 July 1974.
-------
In western Lake Erie productivity increased linearly with depth
at Station 61 in late July similar to Station 30 in Figure 96. Produc-
tivity at Station 66 in late July had a profile similar to Station 52
(Figure 99) with a maximum at 5 m, but the maximum was not
quite as sharply defined. Productivity at Station 75 in early Sep-
tember resembled Station 30 (Figure 96) although the increase was
not quite linear.
Central Lake Erie showed higher relative dark bottle 14C
uptake (dark bottle activity) than western'Lake Erie. Central basin
mean dark uptake increased during thermal stratification from 18
percent in early August to 24 percent in mid-August and decreased
after fall overturn to 12 percent in late October. Relative dark
uptake ranged from 3-34 percent and 3-58 percent for early and
mid-August, respectively. Relative dark uptake of 560 percent and
630 percent at Station 23 on 13 August and 77 percent at Station 73
on 30 July is considered aberrant and unexplainable. Western Lake
Erie relative dark uptake remained at the same levels throughout
the season, as mean rates were 6 percent, 8 percent, and 7 per-
cent (excluding Station 61) for late July, early September, and late
October, respectively. The mean relative dark uptake for all sam-
ples during the season was 14 percent. For incubator studies in
the eastern tropical Pacific Ocean, Jones et al . (1953) reported
relative dark 14C uptake of 16-22 percent for productivity experi-
ments lasting four to eight hours. Larson (1972) reported relative
dark uptake from 4-70 percent during i_n situ productivity studies
in Crater Lake. Relative dark 14C uptake of 6 percent has been
obtained in western Lake Erie (Verduin, personal communication );
the western basin value found in 1974 agree with this estimate.
Obviously, higher relative dark uptake drastically reduces light
bottle uptake when corrected by subtraction.
Primary productivity at one meter in central and western
Lake Erie during 1974 showed high correlation (r = 0.92) with
uncorrected chlorophyll ji concentration, which is a phytoplankton
biomass indicator (Figure 100). The correlation exceeds by three-
fourths that one reported by Glooschenko et al. (1974) for mean values
per cruise of chlorophyll a_ and incubator productivity in surface
water in all three basi n s. Glooschenko et al. called the slope of
the graph the assimilation number, the units of which are mg C/mg
chlorophyll a-h. Glooschenko's mean assimilation number of 1.93
mg C/mg chlorophyll a-h agreed to within 10 percent of that for
1974. ~
255
-------
100
80
o
01
60
o 40
"ra
.Q
3
U
20
Figure 100.
y - 2.13 x + 2.04
r - 0.92
0 10 20 30 40 50
Uncorrected Chlorophyll a Concentration (mg/m3)
Relationship of incubator productivity to uncorrected
chlorophyll a concentration at 1 M depth.
100
20 40 60
Total Phosphorus Concentration
80
100
Figure 101 . Relationship of incubator productivity to total phos-
phorus concentration at 1 M depth.
256
-------
The relationship between incubator primary productivity and
nutrient concentration in central and western Lake Erie during 1974
is not clear. From the concentrations of total and soluble reactive
phosphorus and total inorganic ammonia, and nitrate-nitrite nitro-
gen in the two basins, only total phosphorus correlated with incuba-
tor productivity (r = 0.68) for lake water samples from the one
meter depth (Figure 101> Brydges (1971) found that in western
Lake Erie station averages of chloropyll a from a composite
sample of 1.5 m and 7.5 m water were positively correlated with
total phosphorus at 1 .5 m for the years 1967-1969. Megard (1972)
reported that for eutrophic Lake Minnetonka , Minnesota total phos-
phorus in the range of 0-170 mg/m3 (Hg/1) was linearly correlated
with daily maximum photosynthesis per unit volume of water during
the months of July, August, September. Brydges (1971) also found
that soluble phosphorus, nitrate-nitrite nitrogen, and ammonia
nitrogen did not correlate with chlorophyll si.
Light energy penetrating the surface waters in central Lake Erie
was transmitted more readily than in western Lake Erie during ther-
mal stratification in August (Table 67> The mean euphotic zone in
central Lake Erie was 2.4 times deeper than in western Lake Erie. In
mid-August light transmission improved slightly. Upon lake overturn
the euphotic zone in central Lake Erie was only 30 percent deeper than
in western Lake Erie. Verduin (1954) reported that dissolved pigments
have not been observed in western Lake Erie, and that differences in
light transmission were attributable to suspended particles. The same
is assumed for central Lake Erie.
Incubator productivity represents maximal carbon assimilation
rates that result from optimal light intensities. A rough approxi-
mation of incubator light intensity assumes that full light intensity
measured by the photometer (10.5 mA) equals 10,000 foot candles
(Saunders et al. 1962) during the brightest days. Therefore,
since an average of O.9 V amps of relative light intensity reaches
the tops of the bottles in the incubator under clear water condi-
tions, approximate light intensity is 9000 lux. Tailing (1966)
stated that the light-saturating range of Asterionella communities
in English lakes is 8000-10,000 lux. Tailing (1966), using an
incubator equipped with daylight fluorescent lamps, converted incu-
bator light intensity to quantity of photosynthetically active radiant
energy (400-700 nm) by the conversion factor of 4.1 kerg/cm
per sec per kilolux. Therefore, 37 kerg/cnrf7 per sec (9 x 4.1)
approximates the radiant energy received by phytoplankton samples
in the incubator during the 1974 study. If the areal and time unite
are changed to m2 and h, respectively, the resulting radiant
257
-------
TABLE 67. MEAN SECCHI DISC AND EUPHOTIC ZONE DEPTHS
FOR CENTRAL AND WESTERN LAKE ERIE FROM JULY-OCTOBER 1974
No. of Stations
Basins
Secchi disc (m)
Euphotic Zone (m)
01
oo
Cruise 6
5
5
Cruise 7
8
Cruise 8
4
Cruise 10
11
5
central
western
central
western
central
western
4.5
2.0
6.0
1 .0
3.0
2.0
12.0
5.0
14.5
3.5
8.0
6.0
-------
energy of approximately 1 x 1012 erg/nr£ per hoir falls near the
maximum carbon uptake rates on the light-saturated portions of
photosynthesis-light intensity curves published by Stadelmann,
et al. (1974) for Lake Ontario phytoplankton communities.
Photosynthetic productivity per unit volume of water reflects
phytoplankton densities in the water column at a particular depth,
but the relationship is not a simple one to one correspondence.
Tailing (1966) found for English lakes a broad correlation from
April to June between population density (cell number) and gross
photosynthetic rate per unit volume of water. Blelham Tarn had
population densities and photosynthetic rates two to three times
greater than Windermere North Basin. In both lakes phytoplankton
from below the surface showed greater population densities and
greater photosynthetic rates than surface phytoplankton. Verduin
(1960) found that a five-fold increase in population density accom-
panied a doubling of net photosynthetic rate per volume of water,
when the phytoplankton community changed from June to August,
1957. An explanation may lie in the specific photosynthetic rate
of the phytoplankton. Megard (1972) defined the specific photosyn-
thetic rate mg C/m3-da^mg Ch1or a/m3 which is the equivalent of the
assimilation ratio when calculated per hour. Tailing (1966) used
both cholrophyll a and cell number per unit volume of water as
measures of population density, and Verduin (1960) used phytoplank-
ton volume per unit volume of water. Specific photosynthetic
rate decreased with increasing depth during in situ productivity
studies on Lake Tahoe (Kiefer et al. 1972). The 1974 incubator
studies re-veal that at the same light intensity at Station 47 on 12
August the 1-m and 13.5-m samples yielded specific photo-
synthetic rates of 2.5 and 3.7 mg C/mg chlor a^h, respectively.
This increase of 48 percent resulted from a 15 percent increase
in the incubator productivity from 13 to 15 mg C/m3
-------
density ( mg chlor a/m3, Megard, 1972). For these reasons incu-
bator productivity is a metabolic index of population density as
Saunders et al. (1962) believed when they exposed samples from
different depths to a light intensity of 500 foot candles. The
strong correlation between incubator productivity and chlorophyll
a concentration at the one meter depth in 1974 emphasizes their
contention.
Both incubator and i_n situ productivity data for 1974 remain
uncorrected for excretion of organic matter, respiration losses,
and grazing because they were not quantified by experiment. Ex-
cretion of organic compounds may cause significant underestimates
of productivity. Fogg et al. (1965) found that during a four-hour
experiment approximately 34 percent of the initially assimilated
was excreted by diatom dominated communities in English
Wtth thS 14C tecnniclue no way exists to measure loss of
during breakdown of organic compounds in respiration
(Kiefer et al. 1972). The reaction involved is:
"CHgO + 02 » *C02 + HgO.
Significant grazing of phytoplankton standing crop.may reduce pro-
ductivity. Parkos et al. (1969) felt that grazing may be impor-
tant in localized areas of lakes but should not cause the large
variations in productivity they observed in each of the four Great
Lakes they investigated. Verduin (1952) felt that grazing was in-
significant in western Lake Erie. Glooschenko et al. (1974) felt
that the zooplankton in Lake Erie, which are characterized by
protozoans, rotifers, and cladocerans, consume detritus and bac-
teria more than phytoplankton.
For the 1974 incubator studies 14C uptake was assumed to be
linear at constant light intensities. The literature gives varying
results. Barnett and Hirota (1967) found that at constant light
intensity the rate of 14C uptake was constant during a two-hour
incubation. In a second experiment of four hours duration,
Barnett and Hirota used phytoplankton concentrations ten times
greater, and light intensity was half of the first experiment.
They found that 14C uptake rate decreased to 62 percent of the
rate at one hour.
Fogg et al. (1965) found that photosynthesis was proportional
to incubation time under constant light intensity in situ and in
the laboratory, but organic matter was excreted at an almost
260
-------
constant percentage of the photosynthetic uptake.
Extracellular 14C adsorbed to phytoplankton leads to overesti-
mates of productivity. McMahon (1973) showed that rinsing filters
with 100 ml of distilled or filtered lake water removed extra-
cellular C, which had led to erroneous conclusions concerning
self-absorption. The investigator assumed that the acid and fil-
tered lake water rinses used in 1974 adequately removed
extracellular 14C.
Estimation of phytoplankton productivity at several stations
during different times of the day may exaggerate productivity
relative to each other. Vollenweider and Nauwerck (1961) found
during In situ studies on Lake Erken, Sweden the interval of
0530-0930 yielded productivity per unit area of water one third
of the daily total of a series of five four-hour time blocks.
Holmes and Haxo (1958) demonstrated changes in photosynthesis
throughout the day in incubator studies at a station in the tropical
eastern Pacific Ocean. Maximim photosynthesis occurred at
0800-1000. Verduin (1957) found that the period 0700-1000
in western Lake Erie yielded maximal net photosynthetic rate per
unit volume of water two-thirds greater than during the intervals
1000-1300 and 1300-1600. All times are given in EST.
Whether the 14C method estimates net or gross productivity
remains unsolved. Saunders (1964) felt that because the sum of
a series of short experiments over a day exceeded the estimate of
a single full day exposure, the series estimated' gross productivity
and the single exposure net. Vollenweider and Nauwerck (1961)
suggested that productivity measured by 14C lies between net and
gross.
Both incubator and in situ productivity estimates made by
the 14C method include combined contributions by phytoplankton
and bacteria. Cody (1972) states that the measurement of carbon
assimilation is complicated by autotrophic bacteria, both chemo-
synthetic (aerobic) and photosynthetic (nonsulfur which are aerobic
and sulfur which are anaerobic). The relative amount of bacterial
assimilation is unknown.
Phytoplankton, zooplankton, and bacteria all play a role in
dark 14C uptake (Gerletti 1968). Sorokin (1965) described
three groups of bacteria involved in aerobic decomposition of organic
261
-------
matter which account for the major part of dark CO2 assimilation in
surface layers of eutrophic and mesoeutrophic lakes. Another group
of bacteria intermediate between heterotrophic and chemosynthetic
oxidizes simple organic products of anaerobic decomposition. A
third group are chemoautotrophs that utilize CO2 in the syntheses of
all their organic compounds.
During ir, situ studies on Lake Maggiore, Italy, Gerletti
(1968) found that below the optimal level of illumination (where
maximum photosynthesis is expected to occur) relative dark uptake
of CO2 (dark bottle activity expressed as percent of the cor-
responding light bottle activity) increased. In situ experiments
at Rattlesnake Island during 1974 support GeFlettT. Relative dark
14C uptake on 21 July was 2 percent at one meter, increasing
to 14 percent at five meters; for 1O September, 2 percent at one
meter, increasing to 33 percent at six and eight meters; and on
6 October, 6 percent increasing to 35 percent at four meters.
However, in the incubator studies at an optimal light intensity,
relative dark bottle rates ranged from 1-58 percent and were
especially high in the central basin during late July to mid-
August. On the hottest summer days incubator temperature
remained within 2°C of the lake surface temperature. Since
the temperature from surface to lower epilimnion did not vary
by more than 2 C on the average over the sampling season, the
temperature differential between incubator and sample depth is
negligible. Even though light intensity in the incubator isoptimal,
a significant increase in relative dark bottle activity occurs.
The source of this uptake is not specifically known, although the
bottle surface effect on bacteria may stimulate their growth
(Gerletti 1968). Menon et al. (1972) reported bacterial densities
of 3.3-3700 x I03/ml and 3.4-79O x 104/ml for central and
western Lake Erie, respectively. Vertical distribution was uni-
form until stratification, when hypolimnion bacterial densities
increased more than epilimnion densities. Bacteria in late
August utilized organic matter from excretion and degradation
products of blue-green and green algal blooms to attain maximum
densities.
Morris et al. (197V) found that with marine phytoplankton
relative dark uptake of 14C became greater with smaller popula-
tion densities. Smaller phytoplankton densities in both central
and western Lake Erie in 1974 yielded higher relative dark
uptake, but the relationship is non-linear (Figure 102). Relative
262
-------
140
120
2-100
o
D)
E
~ 80
2
D.
n
.0
o
a>
60
40
20
0' 6/i 0.2 073 0-4
Ratio of Dark Bottle Activity to Light Bottle Activity
0.5
Figure 102.
Relationship of mean relative dark 14C uptake
to mean incubator productivity at each station
in central and western Lake Erie.
263
-------
dark uptake increased with decreasing turbidity (Figure 1O3), which
supports the previous relation.
No relationship was noted between dark 14C uptake and tempera-
ture. For both relative and absolute values equally high and low
rates were observed at temperature ranges of 10-13eC and 18-24*0.
In situ productivity vs, depth profiles show the effect of decreas-
ing light intensities on carbon assimilation rates, whereas incubator
productivity profiles at constant light intensity show relatively little
change in carbon assimilation rates. The shipboard counterpart to
Rattlesnake Island, Station 67, did not have significant changes in
vertical distribution of productivity. In situ profiles show that pro-
ductivity at one meter and deeper closely parallels light
transmission in all three experiments down to a relative light
intensity of approximately 10 percent of the surface light intensity.
Such parallelism indicates that phytoplankton populations are
homogeneously distributed, at least down to the level of 10
percent of the surface light intensity. Rodhe (1965) found that
the slopes of assimilation curves of nine lakes closely agreed
with the slope of the most penetrating component (usually green)
of subaquatic light at a level from 1-12.5 percent of the sub-
surface intensity of the component.
Incubator data showed that the phytoplankton in both basins
survived the light intensity of the water bath. For example, a
sample of lake water from four meters at Station 53 on 6
September from a relative light intensity (775 y A) had a pro-
ductivity potential nearly 20 percent greater than a sample from
one meter, which received about a 370 times greater relative
light intensity. This suggests that the water at Station 53 circulates
completely down to four meters so that the cells at Four meters
do not remain in total darkness, otherwise they may have been
injured by the incubator light intensity and would not have func-
tioned photosynthetically, or if so, then at a reduced rate.
Water temperature and oxygen concentration remain unchanged
from one meter to four meters which supports this hypothesis.
Keifer et al. (1972) reported that phytoplankton from 100-400
m in Lake Tahoe survived light intensity increases of 107, as
When the samples were incubated at a light intensity 40 percent
of the surface, they had photosynthetic potentials per unit of
chlorophyll a_ slightly less than phytoplankton at 75 m in the
euphotlc zone (85).
The incubator productivity-total phosphorus relationship
264
-------
O
o
x
X
o
30
5
x
il20
1
o
o
co
x
o
Q
«4-
o
o
10
DATA FROM STATIONS 30. 47.48, 67, 75. 76
JULY - OCTOBER 1974
0.8
1.0
0.2 0.4 0.6
Light Transmission
RATIO OF PHOTOMETER READING AT SAMPLING DEPTH TO PHOTOMETER
READING ONE METER ABOVE SAMPLING DEPTH
14
Figure 103. Relationship of relative dark C uptake
to light transmission.
265
-------
may indicate that total phosphorus is an indicator of phytoplankton
densities. Brydges (1971) suggested that if total phosphorus concen-
trations were reduced, there would be less algae. The 1974 incu-
bator data support Brydges' argument.
The parellelism in the _in situ productivity-light transmission
curves in Figure 92 suggests that light is a controlling factor in
primary productivity in western Lake Erie. Since total phosphorus
concentration is lenearly related to incubator productivity at the
one-meter depth, then both light and total phosphorus may together
limit primary productivity. If the light-in_ situ productivity re-
lationship holds for central Lake Erie, then both factors may be
limiting in central Lake Erie also.
SUMMARY
Western basin incubator productivity was 3.5 times greater
than central basin incubator productivity in the summer and 1.5
times greater in the fall. Incubator productivity estimates were
comparable to those in 1970. Central basin incubator productivity
decreased in late summer and increased upon fall overturn. West-
ern basin incubator productivity declined throughout the season.
In situ productivity-light intensity curves for western Lake
Erie showed that phytoplankton are photosynthetically active at
lower light intensities in October, as opposed to July and Septem-
ber, and that maximal productivity is reached within a narrower
range of light intensities. Maximum productivity occurs at ap-
proximately one meter. In all three m_ situ experiments relative
productivity curves lay parallel to light transmission curves down
to a level of approximately ten percent of the surface light intensity.
Estimated errors for the 1974 methodology showed that a
slightly greater total error existed than was normally accepted
in other studies cited. Each sample lost an average of 14 percent
of its initially assimilated 14C when merthiolate fixative was used.
This fixation effect had the greatest uncertainty in the estimate of
total error.
Incubator productivity profiles showed several forms. Maxi-
ma were located at one meter, slightly below one meter, and
some mid-lake central basin profiles increased toward the
266
-------
metaHmnion. Lower maximum to minimum incubator productivity
ratios indicate that western Lake Erie waters are mixed more
thoroughly than central Lake Erie waters.
14
Central basin relative dark bottle C uptake averaged three
times greater than western basin relative dark bottle uptake.
Central basin relative dark bottle uptake increased during thermal
stratification, yet western basin relative dark bottle uptake re-
mained rather constant. Higher relative dark bottle uptake
occurred in areas of less turbidity and phytoplankton densities.
The effect of temperature on relative dark bottle uptake was not
evidenced.
Incubator productivity correlated with uncorrected chlorophyll
a concentration and total phosphorus concentration at the one meter
depth. The incubator productivity-total phosphorus, and relative
in situ productivity-light transmission relationships suggest that
light and phosphorus together operate as limiting factors in central
and western Lake Erie.
267
-------
REFERENCES
Barnett. A. M. and J. Hirota. 1967. Changes in the apparent rate of
14C uptake with length of Incubation period in natural phytoplankton
populations. Limnol. Dceanogr. 12:349-353.
Beeton, A. M. and W. T. Edmondson. 1972. The eutrophication
ptoblem. J. Fish. Res. Bd. Can. 29:673-682.
Brydges, T. G. 1971. Chlorophyll a-total phosphorus relationships in
Lake Erie. Pages 185-190 un Proc. 14th Conf. Great Lakes Res.,
Int. Assoc. Great Lakes Res.
Cody, T. E. 1972. Primary productivity in the western basin of Lake
Erie. Ph.D. Thesis (published). Ohio State Univ., Columbus,
Ohio. 113 p.
Fogg, G. E., C. Nalewajko, and W. D. Watt. 1965. Extracellular
products of phytoplankton photosynthesis. Proc. Roy. Soc.
Lond., Ser. B, 162:517-534.
Gachter, R., R. A. Vollenweider, and W. A. Glooschenko. 1974.
Seasonal variations of temperature and nutrients in the surface
waters of lakes Ontario and Erie. J. Fish. Res. Bd. Can.
31:275-290.
Gerletti, M. 1968. Dark bottle measurements in primary productivity
studies. Mem. 1st. Ital. Idrobiol. Dott. Marco de Marchi
Pallanza Italy 23:197-208.
Glooschenko, W. A., J. E. Moore, M. Munawar, and R. A. Vollen-
weider. 1974. Primary production in lakes Ontario and Erie:
a comparative study. J. Fish. Res. Bd. Can. 31:253-263.
Glooschenko, W. A., J. E. Moore, and R. A. Vollenweider. 1974.
Spatial and temporal distribution of chlorophyll a and pheopigments
in surface waters of Lake Erie. J. Fish. Res.~Bd. Can.
31:265-274.
Goldman, C. R. 1963. The measurement of primary productivity and -
limiting factors in freshwater with carbon-14. Pages 103-113 in
M. S. Doty(ed.), Proceedings of the conference on primary pro-
ductivity measurement, marine and freshwater. USAEC TID-7633.
268
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Holmes, R. W. and F. T. Haxo. 1958. Diurnal variation in the photosyn-
thesis of natural phytoplankton populations in artificial light. U.S.
Fish Wildlife Serv. Spec. Sci. Rep. Fish. 27:73-76.
Jones, G. E., W. H. Thomas, and F. T. Haxo. 1958. Preliminary
studies of bacterial growth in relation to dark and light fixatation
of C4C2 during productivity determinations. U.S. Fish Wildlife
Serv. Spec. Sci. Rep. Fish. 279:79-86.
Kiefer, D. A., O. Holm-Hansen, C. R. Goldman, R. Richards, and
T. Berman. 1972. Phytoplankton in Lake Tahoe: deep-living
populations. Limnol. Oceanogr. 17:418-422.
Larson, D. W. 1972. Temperature, transparency, and phytoplankton
productivity in Crater Lake, Oregon. Limnol. Oceanogr. 17:410-
417.
McMahon, J. W. 1973. Membrane filter retentiona source of error
in the 14c method of measuring primary production. Limnol.
Oceanogr. 18:319-324.
Megard, R. O. 1972. Phytoplankton, photosynthesis, and phosphorus
in Lake Mi nnetonka, Minnesota. Limnol. Oceanogr. 17:68-87.
Menon, A. S., W. A. Gollschenko, and N. M. Burns. 1972. .Bacteria-
phytoplankton relationships in Lake Erie. Pages 94-101 (n Proc.
15th Conf. Great Lakes Res., Int. Assoc. Great Lakes Res.
Morris, I., C. M. Yentsch, and C. S. Yentsch. 1971. Relationships
between light carbon dioxide fixation and dark carbon dioxide
fixation by marine algae. Limnol. Oceanogr. 16:854-858.
Parkos, W. G., T. A. Olson, and T. O. Odlaug. 1969. Water quali-
ty studies on the Great Lakes based on C-14 measurements on
primary productivity Univ. Minn., Water Resour. Res. Center
Bull. 17. 121. p.
Rodhe, W. 1958. The primary production in lakes: some results
and restrictions of the 1^C method. Rapp* P.-V. Reun. Cons.
Int. Exolor. Mer. 144:122-128.
1965. Standard correlations between pelagic photosynthesis
and light. Pages 365-381 In C. R. Goldman (ed.) Primary
productivity in aquatic environments. Mem. 1st. Ital. Idrobiol.
269
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Dott. Marco de March! Pallanza Italy, 18 Suppl., Univ. Calif.
Press, Berkeley , Calif.
Saunders, G. W. 1964. Studies of primary productivity in the Great
Lakes. Univ. Mich. Great Lakes Res. Div. Publ. 11:122-22Q.
Saunders, G. W., F. B. Trama, and R. W. Bachmann. 1962. Evalu-
ation of a modified C technique for estimation of ohotosynthesis
in large lakes. Univ. Mich. Great Lakes Res. Div. Publ. 8.
61 p.
Sorokin, J. I. 1965. On the trophic role of chemosynthesis and
bacterial biosynthesis in water bodies. Pages 365-381 in C. R.
Goldman [ed.] , Primary productivity in aauatic environments.
Mem. 1st. Ital. Idrobiol. Dott. Marco de March! Pallanza Italy,
18 Suppl., Univ. Calif. Press, Berkeley.
Stadelmann, P., J. E. Moore, and E. Pickett. 1974. Prirrary production
in relation to temperature structure, biomass concentration and
light conditions at an inshore and offshore station in Lake Ontario.
J. Fish. Res. Bd. Can. 31:1215-1232.
Strickland, J. D. H. and T. R. Parsons. 1968. A oractical hand-
book of seawater analysis. Fish. Res. Bd. Can. Bull. 167.
311 p.
Tailing, J. F. 1966. Photosynthetic behavior in stratified and unstra-
tified lake populations of a planktonic diatom. J. Ecol. 54:Q9-127.
Verduin, J. 1951. Photosynthesis in naturally reared aquatic commu-
nities. Plant Physiol. 26:45-49.
. 1952. Photosynthesis and growth rates of two diatom
communities in western Lake Erie. Ecology 33:163-168.
. 1954. Phytoplankton and turbidity in western Lake Erie.
Ecology 35:550-561 .
. 1956. Energy fixation and utilization by natural communities
in western Lake Erie. Ecology 37:40-50.
. 1957. Daytime variations in phytoplankton photosynthesis.
Limnol. Oceanogr. 2:333-336.
270
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1960. Phytoplankton communities of western Lake Erie and
the CO2 and O2 changes associated with them. LimnoT. Ocenangr.
5:372-380.
1962. Energy flow through biotic systems o^ western Lake
Erie. Pages 107-121 in H. J. Pincus(ed'), Great Lakes basin.
Am. Assoc. Adv. Sci. Publ. 71.
Vollenweider, R. A., M. Munawar, and P. Stadelmann. 1Q74. A
comparative review of phytoplankton and primary production in
the Laurentian Great Lakes. ). Fish. Res. Bd. Can. 31:73Q-
762.
271
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SECTION 9
ZOOPLANKTON DISTRIBUTION IN THE CENTRAL
AND WESTERN BASINS OF LAKE ERIE
Donna D. Larson and David E. Rathke
Center for Lake Erie Area Research
The Ohio State University
INTRODUCTION
Zooplankton samples were collected during the 1974 field season in
the western and central basins of Lake Erie as part of the EPA Nutrient
Control Project funded by the U.S. Environmental Protection Agency and
administered by The Ohio State University, Center for Lake Erie Area
Research. This phase of the nutrient study was designed to provide a
comprehensive collection of information on the species and numbers of
the major zooplankton groups in the western and central basins, in addi-
tion to relating biomass of these animals to other physical, chemical
and biological parameters.
METHODS
During six cruises between May and November of 1974, 51 stations
throughout the western and central basins were visited (see Overview,
Section 1, Figure 1). Zooplankton samples were collected by vertical
tow using a 0.5 meter, 80-micron mesh net which was raised mechani-
cally at a constant rate. Organisms were relaxed with club soda and
then preserved in five percent MgCOs buffered formalin.
At the Columbus laboratory, samples were counted using a strati-
fied method allowing for enumeration of approximately 200 adult crust-
aceans and 250-300 immature crustaceans and rotifers. With a Wild M5
binocular dissecting microscope at 50x in conjunction with a Ward zoo-
plankton counting wheel, copepods, cladocerans and rotifers were identi-
fied to species, excepting immature forms. The open chamber permitted
manipulation and dissection of individuals when necessary for identification.
272
-------
Distinction was made between male and female adult crustaceans, and
all gravid or egg-carrying female animals were enumerated separately and
numbers of young or eggs noted. Nauplii of the cyclopoid and calanoid
species were counted together, but the nauplii were grouped into three
size classes. The cyclopoid and calanoid copepodids were each separated
into three size classes. Adult cladocerans were also placed into three
size classes. The rotifers were identified to species whenever possible.
All rotifers carrying eggs are listed separately, and number of eggs per
individual recorded.
For routine zooplankton identification the taxonomic references
used were: Ward and Whipple (1959), Pennak (1953)', Jahoda (1948),
Voigt (1957), Deevey and Deevey (1971), Brooks (1957), Eddy and Hod-
son (1961) and Ahlstrom (1940, 1943). Occasionally problem species
were sent to other taxonomists for confirmation.
Numbers per cubic meter and milligrams per cubic meter of dry
weight were calculated and basin averages determined for each species
and major group. Biomass values for the rotifers were obtained by
assigning each species a geometric configuration approximating its shape,
determining a volume and assuming a specific gravity of one. Dry weight
was calculated by taking 10 percent of the wet weight as recommended
by Ruttner-Kolisko (1974). Biomass estimates for the cladocerans and
copepods were obtained from Dumont, et al. (1975), Makarewicz (1974),
Nauwerck (1963), Wilson and Roff (1973) and through personal communica-
tion of unpublished data by J.B. Wilson of the Canada Centre for Inland
Waters.
Based on depth and chemical parameters the western and central
basins were further divided into the western basin, Sandusky sub-basin,
western central basin and eastern central basin (Figure 104). These sub-
basins proved valuable in evaluation of distribution of major zooplankton
groups and individual species as the seasons progressed.
NET EFFICIENCY
Differences in net mesh size may result in disproportionate numbers
of zooplankters being captured. If the mesh is large many small organisms
are lost, but small mesh nets may cause serious clogging problems result-
ing in an inaccurate representation of numbers or organisms per cubic
meter of water. Many zooplankton surveys have assumed a net efficiency
of 100 percent, and therefore numbers of organisms presented are prob-
ably lower than actually occur in the water column. According to
McNaught, et al. (1975), estimates of zooplankton blomass based on 100
percent net efficiency are usually low by a factor of two and often by
273
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Figure 104. Sub-basin map.
ZOOPLANKTON
COLLECTING STATIONS
1967 - 1968 :
DATA SOURCE: Davis, 1969
Figure 105. Zooplankton collecting stations 1967-1968,
274
-------
as much as a factor of ten. The average net efficiency during this
study, as determined by two flow meters mounted on the inside and
outside of the net, was approximately 70 percent based on spring,
summer and fall measurements. Numbers per cubic meter were ad-
justed based on this percentage.
PREVIOUS STUDIES
During the past ten years there have been three extensive Lake
Erie zooplankton surveys. Davis (1969) made two transects of the en-
tire lake in July and October of 1967 and a partial transect in January
of 1968. He visited 30 stations located mid-lake (Figure 105) and iden-
tified and counted copepods, cladocerans and rotifers. Watson and
Carpenter (1974) sampled crustacean zooplankton at 30 stations through-
out the western, central and eastern basins from April to December of
1970 (Figure 106). Patalas (1972) traversed the entire lake to sample
crustacean zooplankton at 34 stations during two cruises between June
and August of 1972 (Figure 107).
A methods comparison of recent Lake Erie zooplankton surveys is
presented as Table 68. Differences in sampling techniques, length of
field season, cruise frequency, net size, counting techniques and the
plankton groups examined are apparent when the surveys are contrasted.
These differences in technique make comparison of surveys difficult.
RESULTS
Total Zooplankters
Peak concentrations of total zooplankters were observed during early
June and early September in the western central and eastern central sub-
basins, but this bimodal pattern did not occur in the western basin and
Sandusky sub-basin (Figure 108, Table 69). Zooplankton numbers per
cubic meter increased rapidly in the western and central basins after a
spring low, and remained stable during June and July. Populations in
the two western sub-basins maintained this high level from mid-August
through the end of October, while levels in the central basin dropped
sharply in mid-August. This low was followed by an increase in early
September and a decline at the end of October.
Discussion of the three major zooplankton groups will be presented
in the following sections. A list of species identified from the western
and central basins during 1974 appears as Table 70.
275
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1970 ZOOPLANKTON
COLLECTING STATIONS
DATA SOURCE: \Aatson, 197
Figure 106. 1970 zooplankton collecting stations.
20 0 20 40 60 80 j
KILOMETERS
1968 ZOOPLANKTON
COLLECTING STATIONS
ATA SCJRCE: Patatas, 1972
Figure 107. 1968 zooplankton collecting stations.
276
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TABLE 68.
LAKE ERIE ZOO PLANKTON STUDIES
COMPARATIVE METHODS
ro
i
Investigators)
Engel ("1962*)
Davis (1969)
Watson
and
Carpenter (1974)
f>atalas (1972)
Rolan,
Zack and
Prttschau (1973)
CLEAR
(unpublished)
Great
Lakes
Laboratory
(unpublished)
Year
1961
1967-
1968
1970
1972
1973
1974
1974
Season
June-August
July, October,
1967, January,
1968
April -
December
June -
August
September, 1971
to
January, 1973
May -
November
March-
November
Basin,
# Stations
# Cruises
Western Basin
6 cruises
6 stations
All Basins
30 stations
3 cruises
All basins
30 stations
1O cruises
All basins
34 stations
2 cruises
Cleveland Harbor
10 stations
monthly
CB, WB
50 stations
6 cruises
Eastern Basin
25 stations
6 cruises
Collecting
Method,
Net Size
Juday Trap
64 M net
VT, 0.5m, 64M
VT, 0.4m, 64P
VT, 0.12m, 90M
VT, 0.5m, 76 M
VT, 0.5m, 80M
VT, 0.5m, 64M
Groups
Considered
rotifers,
crustaceans
rotifers,
crustaceans
crustaceans
crustaceans
crustaceans
rotifers,
crustaceans
crustaceans
Counting
Method
Sedgewlck
Rafter
cell
Sedgewlck
Rafter
cell
Inverted
microscope
__
open
counting
chamber
open
counting
chamber
open
counting
chamber
-------
1 x 10 _
1 x 10'
AVERAGE ZOOPLANKTON CONCENTRATIONS
IN N0./m3 BY SUB-BASIN
A qj M P J f J
* WB
SSB
WCB
* ECB
N
1974
Figure 108. Average zooplankton concentrations in
no./m3 by sub-basin.
278
-------
TABLE 69. AVERAGE 1974 ZOOPUANKTON NUMBERS/m3
BY BASIN AND CRUISE
1
2
3
/
B
6
Rotifers
Cladocerans
Copepods
Total
Rotifers
Cladocerans
Copepods
Total
Rotifers
Cladocerans
Copepods
Total
Rotifers
Cladocerans
Copepods
Total
Rotifers
Cladocerans
Copepods
Total
Rotifers
Cladocerans
Copepods
Total
Western
1.5 x 1O4
3.0 x 102
3.9 x 1O4
5.43 x 104
2.1 x 105
4.8 x 104
8.6 x 104
3.44 x 105
1.2 x 10^
5.5 x 104
1.2 x 105
2.95 x 105
2.5 x 105
1.4 x 104
5.8 x 10
3.22 x 105
1.6 x 1 O5
3.8 x 1O4
3.2 x 10
2.3 x 1O5
1.9 x 105
1.5 x 104
1.6 x 10*
2.21 x 105
"B"
3.6 x 104
1.1 x 103
5.7 x 104
9.41 x 104
1 . 0 x 1 05
1.0 x 105
1.2 x 1O5
3.4 x 105
4.1 x 104
4.3 x 104
1 .7 x 1Q5
2.53 x 105
1.8 x 1 O5
1.1 x 104
2.7 x 104
2.18 x 105
1.1 x 1 O5
5.6 x 1O4
4.5 x 104
2.11 x 105
1.1 x 1O5
4.5 x 104
3.6 x 104
1.91 x 1O5
WCB
9.3 x 1O4
2.5 x 103
8.6 x 1O4
1.815X 105
3.8 x 1O4
6.4 x 104
2.0 x 1O5
3.02 x 1O5
3.5 x 1O4
5.1 x 104
1.9 x 1 O5
2.76 x 1O5
6.7 x 104
1.7 x 104
4.5 x 1O4
1 . 29 x 1 05
2.4 x 105
6.7 x 1O4
4.1 x 104
3.48 x 1O5
3.6 x 1O4
3.7 x 104
4.7 x 104
1.2 x 10
ECB
3.6 x 1O4
7.6 x 102
3.5 x 1O4
7.176 x 104
6.6 x 1O4
5.3 x 104
2.0 x 1O5
3.19 x 1O5
4.2 x 1O4
8.2 x 1O4
1.8 x 105
3.04 x 1O5
4.9 x 1O4
1.9 x 1 O4
7.6 x 104
1 . 44 x 1 O5
1.3 x 1O5
2.8 x 1O4
4.6 x 104
2.O4 x 1O5
4.0 x 104
5.0 x 104
5.7 x 1O
1.47 x 105
Shore
5.7 x 104
1.7 x 103
7.9 x 1O4
1.377 x 105
7.2 x 104
8.9 x 104
2.0 x 105
3.61 x 1O5
4
4.6 x 10
7.8 x 104
1 .9 x 1O5
3.14 x 105
7.7 x 104
2.3 x 104
6.5 x 104
1.65 x 1O5
2.4 x 1O5
5.9 x 104
4.4 x 104
3.43 x 105
6.0 x 104
5.5 x 104
5.5 x 104
1.7 x 1 O5
279
-------
TABLE 70. ZOOPLANKTON SPECIES FOUND IN WESTERN
AND CENTRAL BASINS OF LAKE ERIE, 1974 AND 1975
Rotifers
Asplanahna priodonta
Brachionus angularis
B. budapestiensis
B. calyciflorus
B. oaudatus
B. havanaensi-s
B. diversicornus
B. patulus
B. quadridentatus
B. ureeolaris
Chromogaster ovalis
Cephalodella sp.
Collotheca pellagioa
Conochiloides dossuarius
Conochilis unicornus
Euchlanis dilitata
Filinia longiseta
Kellicottia longispina
Keratella ooehlearis
K. coahlearis fo. tecta
Keratella arassa
K. quadrata
K. hiemalis
Leaane luna
Rotaria sp.
Hexarthra mira
Notholaa foliaaea
N. aanminata
N. laurentia
N. squamula
Pleosoma lentiaulare
P. hudson-ii,
P. trunoatwn
Pomphlyx suleata
Polyapthra vulgaris
P. doliahoptera
P. ewryptera
P. longiremis
P. major
P. remata
Synahaeta lakowitziana
5. oblonga
S. pectinata
Synahaeta sp.
Tri-chooepoa oyl-Lndriaa
T. multicrinis
T. similis
T. longiseta.
Cladocerans
Alona affinis
Bosmina longirostris
Ceriodaphnia lacustris
Chydorus sphaerious
Daphnia ambiqua
D. galeata
D. longiremis
D. parvula.
D. retroauzva
Polyphemus pediaulus
Sida crystallina
Diaphanosoma leuahteribergianwn
Eubosmina ooregoni
Euryaeraus lamellatus
Holopediim gibberum
Ilyoaryptus sordidus
Latonia setifera
Ledigia quadrangularis
Leptodora kindtii
Moina braehiata
Pleuroxus procurvas
Copepods
Cyclops bieuspidatus thomasi
C. vernalis
Mesoayolops edax
Euayclops agilis
E. speratus
Tropocyclops prasinus
Paracyclops fimbriatus poppei
Diccptomus ashlandii
D. oregonensis
D. sicilis
D. siciloides
D. minutus
Eurytemora af finis
Epischupa lacustris
Limnocalanus macrurus
Canthocamptus robevtcokevi
280
-------
Rotifers
Due to their relatively low biomass and taxonomic problems, many
plankton investigators have chosen to exclude the rotifers from zooplankton
population studies. Small in size compared to the copepods and cladocerans,
rotifers were found by Schindler and Noven (1971) to comprise no more
than 12 percent of the zooplankton biomass in several lakes characterized
as oligotrophic to eutrophic. In contrast, during 1974 rotifers in the
enriched western basin of Lake Erie comprised 6 to 40 percent of the
total zooplankton biomass. Based on studies in Lakes Michigan and
Huron, Gannon and Stemberger (1975) concluded that such high popula-
tions of rotifers may be considered indicative of eutrophic waters. Be-
cause their rapid reproductive rates make them quick to respond to
environmental changes, rotifers may be valuable as indicator organisms.
Only two investigators since 1960 have included rotifers in Lake Erie
surveys. Secoy (1962) identified 2O1 species of rotifers from the island
region of Lake Erie in 1960. When comparing her study to those of
Ahlstrom (1934), Chandler (1940), Kellicott (1896, 1897) and Jennings (1901)
she found that 106 of the 201 species had been found by more than one of
the previous workers, and only six species were common to all surveys.
Davis (1969) sampled in July, October and January and found highest
rotifer concentrations and number of species in the western basin during
July and October.
The species list which appears as Table 70 includes 47 rotifer taxa
identified from 1974 collections. Most of the species are planktonic and
ubiquitous in distribution. The littoral-benthic organisms which are listed
were found only occasionally as accidental members of the plankton com-
munity.
Lake Erie Western and Central Basin Rotifers
During the 1974 May to November field season, rotifer populations
demonstrated bimodal increases with a mid-summer decline (Figure 1O9).
Numbers were low in early May in all but the western central basin, where
large populations of Notholca spp. were found, primarily N. laurentiae.
By early June, rotifer numbers were high in the western and Sandusky
sub-basins and the community was dominated by Keratella quadrata, K_.
cochlearis, Cgnochilus unicornus and Polyarthra vulgaris. Fewer rotifers
were collected in the central basin during early June. Dominant members
of the central basin community were K. quadrata, Polyarthra dolichoptera
and P. vulgaris.
281
-------
no./m-1
1 x 103
1 x 10
AVERAGE ROTIFER CONCENTRATIONS
IN NO./m3 BY SUB-BASIN
1974
WB
SS B
WC B
EC B
N
Figure 109. Average rotifer concentrations in
no./m3 by sub-basin.
282
-------
The first half of July brought a drop in rotifer populations, although
levels remained comparatively high in the western basin. The high
western basin concentrations were mainly due to peak populations of
C. unicornus and Brachionus budapestiensis.
The second pulse in the rotifer community occurred during mid-
August in the western basin and Sandusky sub-basin, and early September
in the western central and eastern central sub-basins. The mid-August
pulse was the result of large numbers of Brachionus angularis in the
western basin and a peak concentration of Asplanchna priodonta in the
Sandusky sub-basin. The high levels in early September in the western
central and eastern central basins resulted from large numbers of C.
unicornus, K. cochlearis, Trichocerca spp. and Polyarthra major.
By late October of 1974 rotifer populations had declined in the
central sub-basins but remained high in the western basin and Sandusky
sub-basin. The western basin community was dominated by Synchaeta
spp. and Brachionus calyciflorus and the Sandusky sub-basin primarily by
Synchaeta spp.
Rotifers were abundant in both basins during 1974. The western
basin is the shallowest and most nutrient-rich area, and had the highest
numbers of rotifers during most of the 1974 sampling season.
Due to high turnover rates and rapid restructuring of the rotifer
community resulting from environmental changes, sampling during 1974
may not have been frequent enough to reflect seasonal successional patterns
within this group.
Cladocerans
Patalas (1972), as well as other zooplankton ecologists, consider large
numbers of cladocerans to be indicative of eutrophic conditions. In 1961
Engel (1962) noted an increase in cladoceran populations in western Lake
Erie since Chandler's 1938-39 survey, and especially a rise in the num-
bers of Chydorus sphaericus, which is considered an indicator of eutrophic
conditions. An increase in Lake Erie cladoceran populations since 1939
was also noted by Bradshaw (1964), who compared his 1949 data to those
of Chandler (1940) and Hubschman (1960).
Watson and Carpenter (1974) found cladocerans more numerous in Lake
Erie than in Lakes Ontario and Huron, with populations especially high in
the western and central basins of Lake Erie. Patalas (1972) compared
Lakes Superior, Huron and Erie and noted that as eutrophication progressed,
percentages of calanoid copepods declined and percentages of cladocerans and
cyclopoid copepods increased. Numbers of cladocerans were found to be
highest in the western basin of Lake Erie during Patalas1 survey.
283
-------
Lake Erie Western and Central Basin Cladocerans
From May to November of 1974, cladoceran concentrations in the
four sub-basins were very similar and bimodal peaks were concurrent
in all basins (Figure 110). In early May cladocerans occurred at low
concentrations. By the early June cruise high numbers had been reached,
remained stable through the July cruise, and declined sharply by mid-
August in all sub-basins. Early September brought a second pulse, nearly
equalling that which occurred in the early summer. Populations remained
high in all but the western basin until November.
The major component of the early summer cladoceran pulse was
Bosmina longirostris. This cladoceran reached its highest concentration
in the Sandusky sub-basin and was found in lowest numbers in the western
basin. Eubosmina coregoni was the most numerous cladoceran in the
western basin, but occurred in low concentrations elsewhere, Daphhia
retrocurva was also abundant during the first half of the summer, and
occurred in highest concentrations in the western and Sandusky sub-basins.
The early September pulse was dominated by Bosmina longirostris,
Eubosmina coregoni and Daphnia galeata mendotae. Eubosmina coregoni
was found in greatest numbers in the western and Sandusky sub-basins,
Bosmina longirostris reached highest concentrations in the western central
basin, and D. galeata mendotae was restricted to the western central and
eastern central basins.
By the end of October E. coregoni, B. longirostris and Chydorus
sphaericus dominated the cladoceran community. The seasonal peak of
C. sphaericus was observed during the late October cruise. This species
was found in high concentrations in all but the western basin, where it was
declining after a mid-August maximum. Eubosmina coregoni populations
were highest in the eastern central basin and lowest in the western ba-
sin In October. Bosmina longirostris was found in highest numbers in
the Sandusky sub-basin and western basin, with lower concentrations
observed in the central basin.
Cladocerans comprised less than 10 percent of the total zooplankton
biomass (mg/m3) during early May. By early September they made up
over 55 percent of the zooplankton biomass in all of the sub-basins and
were of greatest importance in the western and Sandusky sub-basins
where enriched conditions seem to be most favorable for their reproduction
and growth.
Copepods
Patalas (1972) considered the differences in the crustacean populations
of Lakes Superior, Erie, Huron and Ontario to be due to differences in
284
-------
1 x 10s _
no./m _
1 x 10 _
1 x 10 _
1 x
AVERAGE CLADOCERAN CONCENTRATIONS
IN NO./m3 BY SUB-BASIN
MFTfj
N
1974
Figure 110. Average cladoceran concentrations in
by sub-basin.
285
-------
nutrient loading. With increased nutrients, a decrease in calanoid
copepods and an increase in cyclopoid copepods and cladocerans was
apparent. According to McNaught (1975) this succession may be better
explained by the fact that calanoid copepods are nannoplankton feeders,
while many of the species most abundant in eutrophic waters are gen-
eralist feeders. Therefore animals such as Bosmina longirostris and
Eubosmina coregoni, which are able to utilize a wide range of particle
sizes, are better adapted to the enriched conditions of the central and
western basins of Lake Erie than are the diaptomids, which utilize par-
ticles in the nannoplankton range.
Gannon and Stemberger (1975) stated that in the oligotrophic off-
shore waters of Lake Michigan calanoids made up 50 percent of the
total microcrustaceans. But onshore where nutrient concentrations were
high, calanoids made up only five percent.
In 1928 and 1929 Diaptomus was the most abundant copepod in the
eastern half of Lake Erie in July (except for Limnocalanus located in
the deeper waters, Wilson, 1960) and in the island region of the west-
ern basin in 1930 (Wright, 1955). Since then, the Diaptomus copepods
have decreased in relative abundance, and Limnocalanus has nearly
disappeared (Gannon and Beeton, 1971). In the central and western ba-
sins of Lake Erie the calanoids were greatly outnumbered by the cyclo-
poids and cladocerans throughout the 1974 season, and this trend was
also observed by Davis (1969), Patalas (1972), and Watson and Carpenter
(1974).
Lake Erie Western and Central Basin Copepods
The average copepod populations were small in early May, peaking
in June and July and gradually declining from August through October
(Figure 111). This general trend was apparent in all four sub-basins.
The copepods found in the western and central basins of Lake Erie were
mainly cyclopoids. Although calanoids at times made up about 30 per-
cent of the copepod biomass, they were never as numerous as the cyclo-
poids. One harpactocoid copepod, Canthocamptus robertcokeri, was
found only in low numbers early in the season in the western portion of
the lake.
Of the cyclopoid copepods, Cyclops bicuspidatus thomasi was most
numerous during June and July, especially in the western basin and
Sandusky sub-basin. This copepod nearly disappeared from the western
and Sandusky sub-basins by August, but remained common in the central
basin through October. The Cyclops vernal is population reached a peak
in early July in the western basin and outnumbered C. bicuspidatus
thomasi. Cyclops vernal is was also abundant in the Sandusky sub-basin,
286
-------
no./m
1 x 10
1 x 10'
ttVERaGE COPEPOD CONCENTRATIONS
IN NO./m3 BY SUB-BASIN
A M J J
1974
* WB
SS B
WC B
a 1C B
N
Figure 111. Average copepod concentrations in
no./m3 by sub-basin.
287
-------
although It was greatly outnumbered by C_. bicuspidatus thomasi, and
was nearly excluded from the central basin throughout the sampling
period.
Mesocycloos edax was abundant in the central basin during the
second half of the summer, with maximum populations observed in the
eastern central basin in late August. Mesocyclops edax was seldom
collected from the western portion of the lake.
Tropocyclops prasinus var. mexicanus was found in relatively low
numbers throughout the summer. Its highest populations occurred in
September and October with a maximum in the eastern central basin
during late October.
The most important calanoid copepods in 1974 were the diaptomids:
Diaptomus oregonensis, D^ siciloides, D_. ashlandii, D_. minutis, and
D_. sicilis. During spring and early summer, D_. ashlandii and D.
siciloides were the most abundant calanoids in the western basin~and
the Sandusky sub-basin. Diaptomus ashlandii was also found in the
western central basin and was the only diaptomid common in the central
basin at this time. After July, few diaptomids were collected in the
western basin or Sandusky sub-basin. Diaptomus oregonensis was the
most abundant calanoid during August and September and was restricted
to the central basin. Diaptomus siciloides was a secondary compenent
of the calanoid community at this time and was also confined to the
central basin.
Three other calanoid copepods were found: Limnocalanus macrurus
in the western and western central basins early in the summer; Eury-
temora affinis in the western basin in early summer and in the eastern
central basin in the fall; and Epischura lacustris in the Sandusky sub-
basin in early June. These copepods were never found in abundance.
Immature copepods made up a significant portion of the zooplankton
community. High numbers of naupii were found in June in the central
basin and July in the western and Sandusky sub-basin. During June and
July the cyclopoid copepodid population was high, with the greatest con-
centrations found in the central basin. Calanoid copepodids occurred in
relatively low concentrations in both basins throughout the summer.
Copepodid numbers were usually higher than nauplii numbers. This
would indicate that many nauplii were lost through the 80-micron mesh
of the net. When comparing levels of copepodids to adult copepods, it
was evident that a large number of these immature forms did not de-
velop into adults, probably due to predation.
288
-------
FUTURE OBJECTIVES
Further analyses and comparisons of 1974 and 1975 data are yet to
be completed. Biomass values for the most frequently encountered
microcrustacean species will be obtained directly in the laboratory. Dry
weight (mg/m3) may therefore be calculated more accurately than by
use of literature values alone.
In 1966, Williams stated that numbers of rotifers correlate directly
to phytoplankton abundance. Glooschenko, et al. (1974) did not consider
zooplankton numbers to be in phase with chlorophyll a_ or pheopigments,
but claimed that the zooplankton food chain in Lake Erie is chiefly
detrital and that living algae are more important as food in oligotrophic
lakes. Patalas (1972) related crustacean abundance with degree of
trophy, especially particulate phosphorus loading rates.
Attempts will be made to investigate some of these reports by corre-
lating zooplankton biomass and community structure with parameters such
as temperature, chlorophyll, particulate carbon, particulate phosphorus,
particulate nitrogen and turbidity. Statistical analyses will hopefully
provide insight into community structure within the basins and through-
out the season and also determine variability due to sampling and count-
ing methods.
SUMMARY
1. Zooplankton samples collected from 51 western and central
basin stations during the May to November field season of
1974 were analyzed to determine species composition and
seasonal succession of rotifers, cladocerans and copepods.
2. In spite of their small size, rotifers comprised a signifi-
cant percentage of the zooplankton biomass (6-40 percent
in the western basin). Numerical differences in rotifer
concentrations from basin to basin were quite dramatic.
3. During 1974, high numbers of cladocerans were observed
in both basins, with early summer and autumn pulses.
Cladocerans comprised the highest percentage of the
zooplankton biomass during the late summer and fall.
4. Cyclopoid copepods were dominant over calanoid cope-
pods throughout the 1974 field season. The greatest
numbers of copepods occurred during June and July
in both basins, and numbers declined steadily through
the remainder of the sampling season.
289
-------
High numbers of rotifers occurred during 1974 in the
western and central basins of Lake Erie, and such
high concentrations are considered indicative of enriched
conditions. Cladoceran numbers have increased steadily
since 1939, and this group currently has higher concen-
trations in the western basin of Lake Erie than else-
where in the Great Lakes. Through the years, cladocerans
and cyclopoid copepods have gradually outnumbered the
once dominant calanoid copepods. The trend from a cala-
noid-dominated to a cyclopoid and cladoceran-dominated
community has been observed during studies of other
lakes which have undergone nutrient enrichment.
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Davis, C.C. 1969. Seasonal distribution, constitution and abundance of
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Gannon, J.E. and A.M. Beeton. 1971. The decline of the large zoo-
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293
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SECTION 10
BENTHIC MACROIN VERTEBRATE DISTRIBUTIONS IN
THE CENTRAL AND WESTERN BASINS OF LAKE ERIE
N.W. Britt
Department of Entymology
The Ohio State University
A.J. Pliodzinskas and E.M. Hair
Center for Lake Erie Area Research
The Ohio State University
INTRODUCTION
Lake Erie has been a discussion topic for a number of years due
to its status as a polluted body of water. Changes in the lake's water
quality and its biota have been noted throughout the last century (Beeton,
1961; Brinkhurst et al., 1968; Britt, 1955; Britt et al., 1973; Brown,
1953; Burns and Ross, 1972; Carr and Hiltunen, 1965; Federal Water
Pollution Control Administration, 1968; Herdendorf et al., 1974; Hiltunen,
1969; Langlois, 1954; Shelford and Boesel, 1942; Osburn, 1926; Wood,
1953 and 1963; Wright, 1955). In most cases, the conclusions of their
studies indicated degradation of Lake Erie's waters.
Aspects of water quality degradation involve: 1) increased sediment
loadings, 2) increased turbidities, 3) higher bacterial counts (particularly
of the coliform bacteria), 4) increase in the number and frequency of
nuisance algal blooms, 5) an increasingly larger area of the bottom
becoming anoxic, 6) decreasing fish populations, 7) shifts in the bottom
fauna from sensitive forms to tolerant forms, and 8) Increased loading
and regeneration of major algal nutrients (particularly phosphorus and
nitrogen). To alleviate degradation, both the United States and Canada
began to implement control measures designed to reduce the level of
waste and nutrient inputs into Lake Erie. To evaluate the efficacy of
these measures upon Lake Erie waters, a multidisciplinary approach
project was initiated by the Center for Lake Erie Area Research (1974).
This assessment consists of monitoring several physical, chemical and
biological components of the Lake Erie ecosystem. This particular
subproject was designed to examine Lake Erie's benthic macroinverte-
brate communities. 294
-------
METHODS
Duplicate Ponar grab samples (0.555 m2) were taken at each of 52
stations in Lake Erie's western and central basins during cruises aboard
Ohio State University's research vessel, R/V Hydra. During the initial
stages of the project several research vessels were used including:
R/V Dambach (Great Lakes Laboratory, SUNY), R/V Maple (Great Lakes
Research Division, University of Michigan), and P/B Bluewater (Gross
lie Laboratory, USEPA).
Samples were taken during ice-free months June 1973 through
December 1975 for a total of 15 benthological cruises (Table 71). The
results of nine cruises, June 1973 through October 1974, are discussed
here.
TABLE 71. SAMPLING PERIODS FOR LAKE ERIE
NUTRIENT STUDY - BENTHOS CRUISES
Cruise Number Date
1 1973 28 June - 12 July
2 17 July - 23 July
5 29 August - 4 September
7 14 October - 24 October
2 1974 25 April - 4 May
4 1 June - 10 June
6 26 July - 4 August
8 26 August - 7 September
10 21 October - 1 November
The Ponar samples were taken with a power winch and wet seived
on board through a U.S. Soil Series No. 49 screen. The retained
material was washed into plastic jars and preserved. The samples
initially were preserved with 10 percent formalin, but the 1975 samples
were preserved with AGW (alcohol:glycerine:water in a 7O:5:25 mixture).
The samples were then labeled and stored until analyzed.
The stored samples were stained with Rose Bengal solution and
sorted into Oligochaeta and non-Oligochaeta fractions. Non-ol igochaetes
were identified by Dr. N.W. Britt. The oligochaetes were initially
identified by Dr. E.M. Hair and later by A.J. Pliodzinskas. Enumera-
tion and identification proceeded simultaneously.
295
-------
ORGANISM DISTRIBUTION, DENSITY AND DIVERSITY
Organism distributions and their abundances are the results of vari-
ous interactions of the biotic and abiotic components of the environment.
In the open waters of Lake Erie's western and central basins during the
1973 and 1974 sampling seasons, the benthic macroinvertebrate fauna
is composed of at least 66 taxa, identified to species where possible.
Thirty-six of these taxa are found throughout both basins, while eleven
and nineteen are limited to the western and central basins, respectively
(Tables 72 and 73).
The number of different taxa per square meter or diversity found
throughout the lake varies among stations and with time (Table 74).
Average diversity of samples (1973-1974) indicates that the lake bottom
can be divided into four general zones of diversity (Figure 112). The
boundaries of these zones presented herein should not be considered
absolute as they are interpretations of the combined data.
Zone I, areas of lowest diversity, is to be found: 1) along the
northeastern edge of the central basin (stations 28 and 29), 2) south of
Pelee Point (station 50), and 3) just west of Fairport Harbor (station
35). The Zone I stations were the least successfully sampled stations
during the 1973 and 1974 seasons due primarily to substrate type and
secondarily to adverse weather conditions. These stations have either
sand or sand/gravel veneered clay substrates which reduce the effective-
ness of the Ponar sampler (Figure 113). Generally, these types of sub-
strates are inhospitable to sedentary organisms due to the grinding action
of the sediments during current activity (Purdy, 1967, Figure 114). In
addition, sandy regions are relatively turbulent and result in little de-
tritus settling out and remaining in the sediments; subsequently, there
is little food resource available. It is much more difficult for small
organisms to burrow in sand than in mud because of the difference in
particle size and in the ability of the organism to make cohesive bur-
rows in each sediment type.
Zone II (4-7 taxa) lies primarily within the 20-25-meter depth zone
(Figure 115) and substrates consisting of soft, silty, grey mud with the
major fraction consisting of clay. The Zone II region consists of four
separate areas. The largest area encompasses much of the south side
of the western half of the central basin up to about midway between
Cleveland and Fairport Harbor. Two other areas of Zone II are located
in the eastern half of the central basin, one in the middle-eastern sec-
tion and one in the north-central section of the sub-basin. Along the
western edge of the lake is the fourth Zone II region. This area lies
in depths of six meters and has a substrate consisting of soft grey mud
296
-------
TABLE 72. TAXANOMIC GROUPS FOUND IN LAKE ERIE'S
WESTERN AND CENTRAL BASINS DURING LAKE ERIE
NUTRIENT STUDY (1973-1974)
ANNELIDA
Hi rudinea
Helobdella elongata
E. stagnalis
E. fusca
Glossiphonia complanata
G. heteroclita
Erpobdella punatata
Polychaeta
tfanayunkia speciosa
01igochaeta
Aulodrilus americanus
A. lirrmobius
A. pluriseta
Branchiura sowerbyi
Ilyodrilus templetoni
Lirmodrilus cervix
L. cervix variant
L. claperedeianus
L. hoffmeisteri
L. maianeensis
L. spiralis
L. udekemianus
Peloscolex ferox
P. multisetosus
Potamotkrix moldaviensis
P. vejdovskyi
Tubifex tubifex
Ar-steonais lomondi
Dero digitata
Nais sp.
Ophidonais serpentina
Paranais foreli
P. longiseta
P. menoni
Stylaria lacustris
Vejdovskyella intermedia
Stylodrilus herringianus
ARTHROPODA
Crustacea
Asellus r. racovitzai
Garmarus fasciatus
Pontoporeia affinis
Mysis relecta
Insecta
Oecetis sp.
Oecetis eddlistoni
Chironomus plumosus
C. riparius
C. (Cryptochironomus) sp.
Coelotanypus sp.
Metriocnemus sp.
MicTOpsectra sp.
Microtendipes sp.
Procladius sp.
Tanypus sp.
Tanytarsus sp.
MOLLUSCA
Gastropoda
Physa sp.
Heliosoma sp.
Amnicola sp.
Bulimus tentaculata
Valvata sincera
V. tricarinata
Campeloma sp.
Goniobasis sp.
Somatogyra sp.
Pelecypoda
Ligumia recta
Lampsilis radiata luteola
Leptodea sp.
Pisidiiari sp.
Pisidium compression
Sphaerium sp.
COELENTERATA
Hydra sp.
PLATYHELMINTHES
Planaridae
Phabdocoela
297
-------
TABLE 73. BENTHIC FAUNA FOUND IN LAKE ERIE'S
WESTERN AND CENTRAL BASINS
vo
oo
Western Basin
Oeaetie ep.
Oecetie eddliatoni
Helobdella fueoa
Glossiphonia heteroclita
G. aomplanata
HelioBoma ep .
Campeloma ep .
Lampsilie r. luteola
Leptodea ep.
Planaridae
LirmodriluB udekemianuB
Central Basin
Tony tarBUB ep.
Micropnectra ep.
MetriccnemuB ep .
Miorotendipes ep .
Pontoporeia affinis
AulodriluB americanus
Ophidonais serpentina
Arcteonais lomondi
Paranaia foreli
P. menoni
Nais ep.
Stylaria laouetTiB
VejdovBkyella intermedia
Paranais longiaeta
Amphichaeta ep.
Stylodrilus herringianus
Manayunkia speaiosa
MyBie reliata
Hydra ep.
Both Basins
Helobdella stagnalis
H. elongata
Erpobdella punctata
Chironomue plumoeue
C. ripariue
C. (Cryptochironomus) ep.
Proeladiue ep.
Coelotanypus ep.
TanypuB ep .
GanrnaruB faaiatus
Asellus r. raaovitzai
Armicola ep.
Valvata Bincera
V. tricarinata
BulimuB tentaoulata
Somatogyra ep.
Goniobasia ep.
Sphaerium ep.
Pisidium ep.
P. aompreBBwn
Rhabdoooela
Physa ep.
Aulodrilus lirmobius
A. pluriseta
Branchiura eowerbyi
LinmodriluB cervix
L. cervix variant
L. claperdeianus
L. hoffmeisteri
L. mawneensis
L. BpiraliB
Peloscolex ferox
P. multieetosuB
Potamothrix moldaviensis
P. vejdovskyi
Tubifex tubifex
-------
TABLE 74. BENTHIC MACROINVERTEBRATE DIVERSITY*
IN LAKE ERIE'S WESTERN AND CENTRAL BASINS DURING
LAKE ERIE NUTRIENT STUDY (1973-1974)
Station Nuaber
23
24
?
2
5
I
27
28
29
^
}
1
32
33
!
<
3
4
'
\
A
5
6
&
J
40
41
1
4
4
2
3
»
Cruiae Number
1 2' 5
17
J4
1
5
12
0
3
ll
q
12
11
13
7
13
15
I1?
12
12
4
6
^
45
46
47
48
12
8
49 l*>
50
51
52
53
!
I
4
6
56
57
58
53
bO
61
>5
^
67
fa
2
1
12
13
13
23
13
19
11
17
17
14
7
in
16
7
12
1
9
2
1
11
14
13
10
13
8
12
14
13
13
8
7
in
4
IP
9
9
l
11
6
7
8
8
8
11
6
8
14
11
8
11
1
ll
i
9
2
11
10
7
2
7
10
6
10
1
11
8
£
3
q
7
11
6
8
I
8
13
16
11
90
12
10
q
7
3
10
11
8
^
9
10
10
6
2
7
S
10
7
4
7
b
8
9
7
7
8
4
10
5
8
6
8
^
1
6
"5
10
(a *
TO
71
10
5
8
9
72
73
;
1
i
B
r5
f8
3
laan
Stations Sanpled
}4
7
10
«3
8
a
6.
9
*3
3
8
8
5
12
5
8
39
4
2
6
8
5
7
38
2
12
6
9
13
12
11
9
8
6
6
11
5
8
7
11
8
11
11
5
11
14
11
1?
6
n
2
10
4
6
8
9
30
4
13
7
8
4
17
ll
7
7
9
12
b
4
4
9
7
1
b
5
7
4
4
6
2
12
f,
12
q
9
8
^
6
4
1
7
3«
6
ll
12
B
4
6
12
12
10
B
12
11
12
11
8
5
7
6
c
8
b
8
11
9
12
2
7
7
4
5
8
5
6
7
7
4
8
35
8
8
9
11
12
12
8
8
7
V
3
'}
14
10
4
9
7
10
6
7
15
16
16
11
14
11
11
IS
q
li
11
11
5
10
5
10
9
10
3t
10
17
10
12
9
9
7
11
&
7
12
8
7
4
8
9
7
9
9
5
7
13
11
ll
11
7
14
13
10
n
9
6
14
6
9
34
X
10
12
8
4
10
2
2
^
8
8
12
11
3
9
9
10
11
8
10
6
5
7
q
7
8
B
10
2
7
7
y
8
10
12
12
10
10
8
13
9
11
8
11
4
7
b
7
7
9
6
6
5
9
* Diversity is expressed as number of different
taxa present/m .
299
-------
KILOMETERS
ZONE EZ[ 12-15 TAXA]
£££% ZONE in [ 8-n]
u*SS!j ZONEH[ 4-7J
CTCC! ZONE I [ 0-3]
Figure 112. Average benthic macro invertebrate diversity
(Taxa number/m2, 1973-1974).
Post-glacial mud
Soft mud «ith some sand
Sand and/or gravel
Glacial sediments, clay till
vvith lag sand
Bedrock, snaies in centra! basin
and carbonates in western ba^-n
o
ml
o>l
Figure 113. Distribution of surficial sediments.
300
-------
40
40 80
KILOMETERS
f r~= PROBABLE PREVAILING
BOTTOM FLOW
Data Source: FWPCA, 1S68
Figure 114. Prevailing annual bottom flow.
20 0 20 40 60 80
I 1III 1 ' i i i '
KILOMETERS
Figure 115. Lake Erie bathymetry.
301
-------
with sand. These nearshore stations are at the mouth of the Maumee
River, near the mouth of the Raisin River; at Locust Point; and in the
western arm of Catawba Island.
Zone III (8-11 taxa) extends throughout most of the western basin
and along the margins of the central basin. The largest section of Zone
III lies along the northern part of the central basin. Depths in Zone III
range from a shallow eight meters (station 60) to the deepest station
(37) at 24 meters. The zone contains the greatest variety of substrates
ranging from rock through sand to mud.
Zone IV (12-15 taxa), the zone of greatest diversity, has two com-
ponents, one at each end of the study area. Farther west is the area
just east of the Detroit River mouth and farther east is the area from
just west of Ashtabula towards the northeast (Figure 112). The sediment
type in both regions is identical, i.e., soft, grey, silty mud with sand,
A cursory examination of Figure 112 would give an incomplete assess-
ment of macro invertebrate diversity. Most of this diversity is the re-
sult of the presence of one major group, the Oligochaeta (Figure 116).
The oligochaetes make up 40.9 percent of all taxa and numerically aver-
age 61.9 percent of all individuals (range: 20.2-96.1 percent).
Organism abundance levels (density) are considered to indicate the
relative degree of environmental stress. This stress is the frequency
with which the physiological tolerance limits of an organism to some
environmental factor are exceeded (Menge and Sutherland, 1976). Aver-
age density levels for the Lake Erie macro invertebrate fauna are divided
into two categories (2000-3000/m2 and 3000-4000/m2), both of which are
present in the Zone I diversity category. If greater abundance reflects
a lower stress environment for the species present, then the margins of
the central basin, the mouth of the Detroit River, and the region just
lakeward of Sandusky Bay may be considered as more favorable environ-
ments than other parts of the lake sampled (Figure 117).
The high densities are due to the presence of large numbers of
oligochaetes (Figures 118 and 119). The two other faunal components,
which together with the oligochaetes, make up 92 percent, numerically,
of all the benthic macroinvertebrates are the Chironomidae and the
Sphaeriidae. Chironomid larvae contribute 11.8 percent toward density
and Sphaeriids, 18.3 percent. In terms of their contribution to diversity,
the chironomids contribute 15.2 percent and the sphaeriids, 4.5 percent.
Most of the lake has average chironomid densities up to 500/m2 with
scattered regions of higher density (Figures 120 and 121). Average
sphaeriid densities indicate that there are two regions of low density,
the western basin and the northeastern corner of the central basin
302
-------
KILOMETERS
L*f/2:3 ^'CO
Figure 116. Average percent oligochaeta in the bottom fauna.
KILOMETERS
i 4000-5000
3000-4000
2000-3000
cC'aj-J 1000-2000
O-IOOO
Figure 117. Average benthic macro invertebrate densities
(indiv./m^) for central and western basins of
Lake Erie 1973-1974.
303
-------
0-IOOO/m2
[II ] IOOO-2000/m2
[Ipllfij 2000-3000/m2
~_~r_"] 3000-4000/m2
4000-5000/m2
Figure 118. Average oligochaete density for central
and western basins of Lake Erie 1973-1974.
Figure 119. Average percent oligochaetes in the bottom
fauna.
304
-------
0- 500/m2
500-lOOO/m*
ni IOOO-1500/m*
Figure 120. Average chironomid density in central and western
basins of Lake Erie 1973-1974.
0 40
c
KILOMETERS
%
L:;:"J o-io
V&M "1-20
31-40
Figure 121. Average percent of chironomidae in the bottom fauna.
305
-------
(Figures 122 and 123). Likewise most of the central basin has shaperiid
densities in the 500-1000/m2 range with scattered regions of higher den-
sity.
When the overall diversity is separated into its component groups,
the diversities within each component may be examined and the distribu-
ions of each individual taxa may be studies. Oligochaete diversity
varies throughout the lake bottom, but most of the lake bottom has from
five to eight oligochaete species (Figure 124). When referring to oligo-
chaetes in this paper we are, for all intents, referring to the Tibifici-
dae, although both the Lumbriculidae and the Naididae are represented
(infrequently).
The most important tubificid group is the genus Limnodrilus, in
which there are seven species. The most ubiquitous is L. hoffmeis.teri
and in order of decreasing distribution they are: L_. cervix, L. cervix
variant, L. maumeensis, L. claperedeianus, L_. sprialis, and~L. udeke-
mianus (Figure 125).
In the western basin, the Limnodrilus species are the dominant
oligochaetes. In the central basin, Peloscolex ferox and P. multiseto-
sus become dominant. P_. ferox oftentimes is the only oligochaete in
some of the stations. Both Peloscolex species are widely distributed
throughout the central basin and in large areas of the western basin
(Figure 126).
Relative to these two genera, the other oligochaete species are
much less important, except locally, and less broadly distributed.
Branchiura sowerbyi seems to be limited to the western basin and to
the southwestern corner of the central basin (Figure 126). However,
it has been collected at one site along the north central basin margin.
Other tubificids, including Potamothrix moldaviensis, P. vejdovskyi,
Tubifex tubifex, Ilyodrilus templetoni, Aulodrilus americanus, A.
Limnobius, and A. pluriseta are scattered throughout both basins as
are the Naididae (Figure 127).
Potamothrix moldaviensis seems to be found along the northern and
southern central basin margins. Potamothrix vejdovskyi, however, is
found in a number of areas. It is very probable that the distribution
of these two species is much more widespread. Tubifex tubifex is
found scattered along the margins of both basins, but is probably found
throughout the western basin and the western half of the central basin
(Figure 127). Ilyodrilus templetoni has been found only off the tip of
Pelee Point. Aulodrilus americanus and A. limnobius are found in both
basins, but so far have been collected from only three stations (Figure
3Q6
-------
20
20 40 60 80
i'iiI 1I
KILOMETERS
JQlUIUfll 0- 500/m2
r--n 500-IOOO/m2
EiMi3 IOOO-1500/m2
^~~J 1500-2000/m2
2000-2 500/ m2
Figure 122. Average sphaeriid density in the central and western
basins of Lake Erie 1973-1974.
0 40
>
KILOMETERS
80
Figure 123. Average percent of sphaeridae in the bottom fauna.
307
-------
20
Co
O
CO
0-4TAXA
5-8 TAXA
9-13 TAXA
Figure 124.
Average oligochaete diversity in the central and
western basins of Lake Erie 1973 - 1974.
-------
O.)u. hoffnwisteri
CO
o
f.)L. spirolis
L. udakemianus
(x)
Figure 125. Distribution of Limnodrilus species in the central and western
basins of Lake Erie 1973-1974.
-------
Q.) Peloscolex ferox
d.) Stylodrilus
herringianus
b.) Peloscolex
multisetosus
oo
i«
o
C.) Bronchiura
sowerbyi
Figure 126.
Distribution of Oligochaete species in the central and western
basins of Lake Erie 1973-1974.
-------
OJ
0.) Potamothrix moldoviensij
(j.) Ilyodrilus templetoni
+ Audrilus
omericanus
a A. I imnobi
Figure 127. Distribution of Qligochaete species in the central and western
basins of Lake Erie 1973-1974.
-------
127). Aulodrilus pluriseta has been found in three regions of the central
basin (Figure 127).
The lumbriculid, Stylodrilus herringianus. generally considered to
be intolerant of enriched conditions, is limited to the far east and mid-
dle region of the central basin (Figure 126).
Brinkhurst el al. (1968) divided the oligochaetes of Lake Erie into
three associations which approximated the three basins. They reported
that the oligochaetes were very abundant at river mouths and in the
western basin with Limnodrilus hoffmeisteri being the most abundant
species. Found with L_. hoffmeisteri were other Limnodrilus species
(L- claperedeianus. L_. cervix, and L^ maumeensis). Tubifex tubifex
was found only at the Detroit river mouth. Further out into the western
basin, they found Aulodrilus and Potamothrix species together with
Branchiura sowerbyj^ and Peloscolex ferox or multisetosus. The central
basin had Fj, ferox as its most abundant oligochaete. In addition Both-
rioneurum vejdovskyanum. Ilyodrilus templetoni, Potamothrix. and"^uTo-
drllus sPecies were found. Aulodrilus species were widely distributed"
and of greatest abundance in the shallower regions of the eastern basin.
The next most important faunal component is the Mollusca, repre-
sented by 16 taxa, of which the Sphaeriidae are the most important.
The shpaeriids are distributed ubiquitously throughout both basins with
the density pattern described earlier. The Unionidae are limited to the
open waters of the western basin and occur in relatively low numbers.
The Gastropoda contribute the most toward mollusc diversity with nine
taxa. Most of these snails, however, are limited to the western basin
and the southern margin of the central basin (Figure 128). Valvata
sincera is the most widely distributed snail. It occurs throughout both
basins but doesn't occur in the middle reaches of the central basin
(Figure 128). The distribution of V_. sincera represents the maximum
distribution of all the Gastropoda.
The gastropod distribution in Lake Erie is most probably due to
oxygen processes in the lake. The snails collected were almost exclu-
sively prosobranch (internal gill) types which must utilize dissolved
oxygen in the water for respiration. Although these organisms can be
found at oxygen concentrations as low as 2 ppm (Hart and Fuller, 1974)
a complete lack of oxygen for such time periods as in the Lake Erie
central basin during summer is prohibitive. During the mixed well-
oxygenated part of the year, there appears to very little immigration to
these areas. Valvata sincera seems to be the only snail which has an
extended range well into the central basin, but it too is not found in the
middle reaches of the central basin. Snail abundance levels in the central
basin are very low (10-20/m2).
312
-------
Q.) Sphoeriidae
to
I"
oo
Figure 128. Distribution of mollusca in the central and western basins of
Lake Erie 1973-1974.
-------
Untonid clams are likewise limited in their distribution. This study
found their abundance to be very low in the open waters (1 or 2/m^)
when collected. However, the frequency with which they were collected
was very low (maximum of 4 stations/cruise - usually 2 stations/cruise).
The distribution of the unionids and their abundance is probably due to
several factors; the most important of which are probably dissolved
oxygen and siltation (which may occur synergistically).
Following the Mollusca in importance are the Chironomidae with 10
taxa. The density levels of the chironomids have been examined earlier
and will not be discussed here. The most common chironomid is
Chironomus plumosus, and in order of decreasing distribution they are:
Procladius sp., Chironomus riparius, Micropsectra sp., Coelotanypus
sp., Chironomus (Cryptochironomus) sp., Microtendipes sp., Metrioc-
nemus sp., and Tanytarsus sp. (Figure 129).
Throughout both basins C_. plumosus and Procladius sp. are the dom-
inant chironomids. In the western basin, Coelotanypus sp. and C.
(Cryptochironomus) sp. are significant members of the chironomld" as-
semblage. In the central basin these two genera become insignificant
except near the basin margins. Coelotanypus sp. is represented along
the southwestern edge of the central basin and C_. (Cryptochironomus)
sp., along the northwestern margin of the central basin and the southern
margin off Fairport Harbor and Conneaut (Figure 129).
Along the northern shore, Micropsectra sp. and Tanytarsus sp. are
well distributed (Figure 129). Rare or uncommon genera include Micro-
tendipes sp., Tanypus sp., and Metriocnemus sp., which are found along
the margins of the central basin (Figure 129).
The distributions of the various taxa comprising relatively minor
positions in the benthic fauna are shown in Figure 130. These faunal
units contribute 15 taxa toward diversity, but only eight percent numer-
ically toward density. These fauna include the six leech species, two
amphipod species, one isopod species, one opposum shrimp species,
one polychaete species, one coelenterate, two flatworm families, and
one genera of caddisfly.
High sediment organic content in bodies of water, such as Lake
Erie (Figure 131) are accompanied by a high oxygen demand by these
sediments and their microflora. In shallow waters this demand may be
met at the mud-water interface by water containing dissolved oxygen
being driven downwards by wind action. The dissolved oxygen content
of the bottom water at the interface, however, will be lowered. In
waters deep enough to develop a summer hypolimnion, wind circulation
314
-------
0.) Chironomus
plumosus
C.) Coelotanypus sp
f.) Chironomus (Cryptochironomu s)
Tanytarsus sp
-t- Mlcrotendipes sp
a Metrlocnemus sp
Figure 129.
Distribution of chlronomldae In the central and western basins of
Lake Erie 1973-1974.
-------
oo
0.) Hirudinea
f.) + Manoyunkia speciosa
Mysis relicta
Figure 13O. Distribution of minor elements in the central and western basins
of Lake Erie 1973-1974.
-------
Figure 131. Distribution of organic carbon in surficial
sediments (% dry weight).
BOTTOM DISSOLVED OXYGEN
CRUISES AUG29-SEPT4.I97
Contour tntorvol: 2 PP">
Figure 132. Bottom dissolved oxygen, cruise 5, August 29-
September 4, 1973.
317
-------
is insufficient to renew the hypolimnetic oxygen stock. As a result,
unless some other mechanism operates, the hypolimnion may become
anoxic. For the hypolimnion to become anoxic during the summer,
it must be relatively thin and have a sufficient oxygen depletion rate
as in the case in Lake Erie's central basin (Blanton and Winklhofer,
1972). Thermal stratification may persist in Lake Erie for up to 110
days, which is sufficient for anoxia to develop (Dobson and Gilbertso"^,
1972).
During summer stratification the dissolved oxygen content of the
bottom waters becomes critical for the bottom fauna. Only the faunal
elements tolerant of low oxygen levels or complete anoxia for relatively
long time periods can survive. Faunal groups generally found under
these conditions are the tubifid oligochaetes, the sphaeriid clams, and
the midge larvae (particularly those of Chironomus sp.). These are
the major faunal elements of Lake Erie's central basin.
The survival of these organisms and their relative success in areas
exhibiting anoxic conditions is due at least in part to the ability of these
organisms to shift their metabolism from aerobic respiration to anaero-
bic respiration. Anaerobic respiration involves obtaining energy by
splitting carbohydrates into lactic acid, fatty acids or mixtures of the
two. The process stops at this point. Aerobic respiration is similar
except that the splitting of the carbohydrates is carried further to pro-
duce carbon dioxide and water.
In 1973 the central basin was anoxic to its greatest extent by late
August (Figure 132). At this time the average bottom fauna density
for the central basin was 3235/m2 with an average diversity of seven.
These values were the lowest for the 1973 sampling year. Prior to
this time the central basin densities averaged 4120 and 5457/m2
during June and July 1973, while diversities were 10 and 9 for the
same period (Table 75) . As the season progressed towards anoxia in
the central basin, changes in both density and diversity were noted
(Figures 133 and 134). Upon the return of oxic conditions there was
also an increase in the average density but no change in the diversity
for the 1973 year.
In 1974, the bottom went anoxic in the southwestern edge of the
central basin at stations 51 and 46 about 12 August, while maximum
anoxia extent was reached about 26 August (Figures 135 and 136).
Lowered benthic densities were observed for the 26 August-7 September
cruise. In the central basin density levels as well as diversity levels
dropped significantly in June (Figures 133 and 134). Recovery occurred
during July but dropped during anoxia. June is the month when a large
emergence of midges occurs, and this may be the primary cause for
the density and diversity drop.
318
-------
TABLE 75. COMPARISON OF DENSITY AND DIVERSITY
(TAXA NUMBER/M2) FOR THE WESTERN AND
CENTRAL BASINS, 1973 - 1974
U)
i>
UD
Cruise Number
Average Density
Central Basin
Western Basin
Central and
Western Average
Average Diversity
Central Basin
Western Basin
Central and
Western Average
1973
1
4120
5460
4494
1O
13
1O
2
5457
3805
5551
9
9
9
5
3225
3615
3323
7
11
8
7
3669
2337
3248
7
6
7
1974
2
4338
1618
3250
9
10
9
4
1865
674
4480
7
7
7
6
391O
1443
3135
9
7
8
8
3471
2592
3202
9
1O
10
10
3409
2017
3000
9
12
9
-------
CO
f>0
o
n
^«
(Q
DIVERSITY
->J 00 if) O f\) Oi
£> a-
2. * 2*,
S «g §«
C * d T-,
(o a
vl -
co <
_
(0 to
O
(D
to
o
(D
SL
(U
(D
I
t:
51
OJP
i 2
to <
3! 3)
^!2
S^
O .
»Z
> c
01
m
CT
to
^-»
8
(D
CO
L
(D
c
(D
CO
CO
(D
(1)
(Q
(D
a
(D
to
a
O
?
(D
(n
" 3
O
0)
I
(D
(D
0)
(D
(f
DENSITY (IND/Mf)X 10s
-------
FIGURE 24
BOTTOM DSSOLVED OXYGEM
CRUSE? 4U8UST 12-19, !S7«
Ccntoi* Mtrvot: I ppm
Figure 135. Bottom dissolved oxygen, cruise 7,
August 12-19, 1974.
FIGURE 25
BOTTOM OSSOLVEO OxrSEN
CRUISE 8 AUG 26- SEPT. 7, IS74
Contour Inntvoll
Figure 136.
Bottom dissolved oxygen, cruise 8,
August 26-September 7, 1974.
321
-------
HISTORICAL TRENDS
The open water fauna of Lake Erie has definitely undergone a num-
ber of changes since at least 1930. Prior to and during the 1930's, the
bottom fauna of the western basin was dominated by the burrowing may-
fly, Hexagenia (Carr and Hiltunen, 1965; Chandler, 1963; Shelford and
Boesel, 1942, Wright, 1955). Data for the central basin bottom fauna
was not available for this time period. Other taxanomic groups were
relatively unimportant in comparison, although the tubificid oligochaetes
became very important in river mouths and harbors (Table 76). Wright
(1955) categorized the western basin into several pollution categories
based upon the number of tubificid oligochaetes present. This was an
arbitrary scale which seemed proper for the time in the western basin.
Light pollution was at oligochaete densities of 1OO to 999/m2. Moderate
pollution was at densities 10OO-5OOO/m2. Heavy pollution was at oligo-
chaete densities greater than 5000/m2. Most of the western basin at
this time was considered unpolluted, while the mouths of the Maumee,
Raisin and Detroit Rivers were considered polluted (Figure 137).
From 1930 through 1961, Carr and Hiltunen (1965) reported that:
(1) Hexagenia had decreased to less than one percent of its previous
abundance, (2) oligochaetes had increased ninefold in density, (3) the
extent of areas defined as polluted by Wright/s index had increased,
(4) chironomid densities had increased four times, (5) sphaeriid levels
had magnified twice, and (6) the snail densities increased sixfold.
Britt (1955) reported a drastic decline in the mayfly population dur-
ing 1953 due to thermal stratification and bottom water oxygen depletion
in the western basin. The population recovered by by 1959 had been
very much reduced (Beeton, 1961). By 1965, Britt et al. (1973) con-
sidered Hexagenia almost extinct in the western basin. In 1973 and
1974 no Hexagenia was collected. In the spring, summer and fall
of 1963 and 1964, the U.S. Public Health Service conducted surveys
of the bottom fauna of Lake Erie (FWPCA, 1968). This survey was
also conducted in 1967 and 1968. They concluded that most of the
western and central basins were characterized by a lack of the pollution
sensitive scud populations and a preponderance of pollution tolerant
species of sludgeworms, bloodworms, fingernail clams and nematodes
(Figure 138). They identified 34 species in the lake. The survey
conducted during the 1963-1964 seasons indicated that the average bot-
tom fauna density was 1666/m2. Average densities for the western and
central basins were 1164 and 1861/m2, respectively (Figure 139).
The 1973 and 1974 sampling seasons discussed here indicate
average bottom faunal densities to be 3409/m2. Average densities
for the western and central basins are 2612 and 3719/m2, respectively,
double the levels found in 1963 and 1964 (Figure 140).
322
-------
TABLE 76.
LAKE ERIE WESTERN BASIN BOTTOM FAUNA
1929-1930 - WRIGHT (1955)
u>
i\i
U)
Amnlcola
Blthynta
Chlronomtdaa
Gastropoda
Qonlobnsla
Hexagenla
Hlrudlnea
Sphaerllda*
Trtchoptera
Tublflcldne
Valvata
laland Region
1929 1930
11
56
2U3
24
23
18
12
9
21
21
610
14
14
1
5
1
Maumae Bay - Toledo Harbor
1 929 1 930
50
31
15O
22
59
107
72
31
100
14
5
98
03
70
332
0
504
16
Portage
1929
18
0
19
River Area
193O
?
few
0
Detroit Rtver
1930
Raisin River Section
192Q 1930
Amnlcola
Blthynla
Chlronomldae
Gastropoda
Gonlobaala
Hexaganla
HlrucJlnoa
Sphaerlldae
Trlchoptera
TubiflcUlae
Valvnta
9
ei
0
374
1500
10
107
1H
57
7
125
70
i!1
4
88
55
18
0
3
46
0
1 04
1
-------
Eastern Limits of
Study Ar«a
loin
[ '"~J I I(£AVY FfJI .1.1 ITION
Figure 137. Extent of organic pollution Indicated by tublflcld Index
(Wright, 1955).
-------
ro
20 0 20 40 60 80
I 1I 1I
KILOMETERS
Data Source: FWPCA, 1S68
PONTOPOREIA AFFINIS
PLUS OTHER AMPHIPODA
TUBIFICIOAE ISOPODA
U;UJ:-:^:- NEMATODA
PROSOBRANCHIA
TUBIFICIOAE
TENDIPEDIOAE
SPHAERIIDE
NEMATODA
INSUFFICIENT
SAMPLING
ONLY
Figure 138. Lake Erie benthic population distribution 1963-1964,
-------
JI777
.1067
20
20
40 60 80
I - 1 I - il
KILOMETERS
542
5749-
^847
146*
1086 «I097
913
^
91
369
8
443
«7Ofi
\
3544
574
.1073
1056
926
/295»
CU
^=
991
ISO
.1001 .£
860
.4934
*805
__ .3675
.1741
'97S9*332 .1036
944 »806 S
864*"67>^7
556,-^
/^
^
772 '
437
1310. /
6.63^
r
331
2623
1717
61* 2391
787
1096
454
345
483*
.12543
434 »5262
216
1883
145*
64
2833
3725
809
Figure 139.
665
762
459
788
163
348
694
Data Source: FWPCA, 1S68
Lake Erie benthic population distribution
1963-1964.
Density = lndividuals/rr2
Figure 140.
Average density Lake Erie benthic populations
1963-1964.
326
-------
If Wright's (1955) pollution index utilizing the oligochaetes was
applied to the 1973 and 1974 field data, almost all of the western and
central basin would be considered moderately polluted. This index,
however, is an arbitrary one and caution should be applied in placing
absolute faith in it.
CONCLUSIONS
Lake Erie's bottom fauna has changed since 1930. During the per-
iod prior to 1953, the western basin bottom fauna was dominated by the
burrowing mayfly, Hexagenia sp. Tubificid oligochaetes were insignifi-
cant members of the fauna except near river mouths and harbors. Cui
rently Hexagenia is considered extinct in the western basin. Greater
than 61 percent of the bottom fauna is dominated primarily by the tubi-
ficid oligochaetes of the genus Limnodrilus. Chironomid larvae comprise
the next most abundant group with less than 20 percent of the bottom
fauna. The sphaeriid clams make up less than 10 percent of the bottom
fauna, except near the Detroit River where they comprise 11 to 20 pel
cent of the bottom fauna.
The central basin fauna is likewise dominated by oligochaetes. The
dominant oligochaete is Peloscolex ferox. However, tubificids of the
genus Limnodrilus frequently make up significant components of the
fauna and are especially important along the northern shore (11 to 30
percent of the bottom fauna). The sphaeriid clams are major components
throughout the central basin, and in many cases are co-dominant with
the oligochaetes. The sphaeriid clams comprise from 21 to 50 percent
of the bottom fauna.
Eight percent of the bottom fauna, numerically, is comprised of
the 15 "minor" taxa. These include leeches, amphipods, isopods, poly-
chaete worms, mysids, coelenterates, flatworms, and caddisflies.
Most of the numerical contributions of these taxa is due to the aquatic
isopod, Asellus racovitzai racovitzai.
If Wright's (1955) pollution index is applied to the 1973-1974 sam-
pling period, most of the study area would be considered moderately
polluted. Caution must be exercised in trusting this index. The
average density of the bottom fauna has doubled since the USPHS 1963-
1964 survey with most of this change due to increases in oligochaete
density.
327
-------
REFERENCES
Beeton, A.M. 1961. Environmental changes in Lake Erie. Trans.
Am. Fish. Soc. 90(2): 1 53-159.
Blanton, J.O. and R.S. Winklhoffer. 1972. Physical processes affecting
the hypoHmnion of the central basin of Lake Erie. Pages 9-38 in
Project Hypo, N.Burns and C. Ross (eds.). CCIW Paper No. 6~T~
USEPA Tech. Rept. TS-05-71-208-24. 182 p.
Brinkhurst, R.O., A. L. Hamilton, and H.B. Herrington. 1968. Com-
ponents of the bottom fauna of the Great Lakes. Univ. Toronto.
Great Lakes Inst. PR 33. 49 p.
Britt, N.W. 1955. Stratification in western Lake Erie in summer 1953:
effects on the Hexegenia (Ephemeroptera) population. Ecology
36(2):239-244.
Britt, N.W., J.T. Addis and R. Engel. 1973. Limnological studies
of the island area of western Lake Erie. Bull. Ohio Biol. Surv.
4(3): 1-89.
Brown, E.H. 1953. Survey of the bottom fauna at the mouths of ten
Lake Erie south shore rivers: its abundance, composition, and
use as an index of stream pollution. Ohio Dept. Nat. Res., Div.
Water, Lake Erie Poll. Surv. Final Rept. p. 156-170.
Burns, N. and C. Ross (eds.). 1972. Project Hypo. CCIW Paper No.
6. USEPA Tech. Rept. TS-05-71-208-24. 182 p.
Carr, J.F. and J.K. Hiltunen. 1965. Changes in the bottom fauna of
western Lake Erie from 1930 to 1961. Limnol. Oceanogr. 1O(4)
55-1 -569.
Center for Lake Erie Area Research. 1974. Lake Erie nutrient control
program: an assessment of its effectiveness in controlling lake
eutrophication. Prog. Rept. 1973 Field Season. The Ohio State
Univ. 129 p.
Chandler, D.C. 1963. Burrowing mayfly nymphs in western Lake Erie
previous to 1947. Proc. 6th Conf. Great Lakes Res., Univ. Mich.
Great Lakes Res. Div. Pub. 10:267-268.
328
-------
Dobson, H.H. and M. Gilbertson. 1972. Oxygen depletion in the hypo-
limnion of the central basin of Lake Erie 1929 to 1970. Pages 3-
8 In Project Hypo, N. Burns and C. Ross (eds.). CCIW Paper
No~7~6, USEPA Tech. Rept. TS-05-71-2O8-24. 182 p.
Federal Water Pollution Control Administration. 1968. Lake Erie
Environmental Survey 1963-1964. U.S. Dept. Interior FWPCA
Great Lakes Region 1968. 170 p.
Federal Water Pollution Control Administration. 1968. Lake Erie
Surveilance Data Summary 1967-1968. U.S. Dept. Interior
FWPCA Great Lakes Region 1968. Cleveland Program Office.
65 p.
Hart, C.W., Jr. and S.L. Fuller, (eds.)- 1974. Pollution ecology of
freshwater invertebrates. Academic Press, N.Y. 389 p.
Herdendorf, C.E., S.M. Hartley, and L.J. Charlesworth. 1974. Lake
Erie bibliography in environmental Sciences. Bull. Ohio Biol.
Surv. 5(5):119.
Hiltunen, J.K. 1969. Distribution of Oligochaetes in western Lake
Erie 1961. Limnol. Oceanogr. 14(2):260-264.
Langlois, T.H. 1954. The western end of Lake Erie and its ecology
J. Edwards Publ. Co. Ann Arbor, Mich. 479 p.
Menge, B.A. and J.P. Sutherland. 1976. Species diversity gradients:
synthesis of the roles of predation, competition and temporal
heterogeneity. Am. Nat. 110(973):351-369.
Osburn, R.C. 1926. A preliminary study of the extent and Distribution
of sewage pollution in the west end of Lake Erie. Unpublished
report to Ohio State Fish and Game Division, (manuscript availa-
ble at F.T. Stone Laboratory, Ohio State University, Put-in-Bay,
Ohio). p. 238-271.
Purdy, E.G. 1964. "Sediments as substrates" in Approaches to
Paleoecology. J. Lumbrie and N. Newall. 1964. John Wiley
and Sons, N.Y. 432 p.
Shelford, V.E. and M.W. Bosel. 1942. Bottom animal communities
of the Island area of western Lake Erie in the summer of 1937.
Ohio J. Sci . 43(5): 179-190.
329
-------
Thomas, R.L., J.M. Jaquet, L.A. Kemp and C.F. Lewis. 1976.
Surficial sediments of Lake Erie. J. Fish. Res. Bd. Can.
33(3):395-403.
Wood, K.G. 1953. Distribution and ecology of certain bottom living
Invertebrates of the western basin of Lake Erie. Ohio State
Univ. Ph.D. Dissertation. 72 p.
Wood, K.G. 1963. The bottom fauna of western Lake Erie. 1951-
1952. Proc. 6th Conf. on Great Lakes Res. Univ. Mich. Great
Lakes Res. Div. Pub.
Wright, S. 1955. Limnological survey of western Lake Erie. U.S.
Fish and Wildlife Serv., Spec. Sci. Rept. Fisheries No. 139
341 p.
330
-------
SECTION 11
STATISTICAL ANALYSIS OF THE 1975 INTER-
COMPARISON STUDY ON LAKE ERIE
Paul I. Feder
Department of Statistics
The Ohio State University
John E. Zapotosky
Center for Lake Erie Area Research
The Ohio State University
INTRODUCTION
In recent years a number of laboratories and government agencies
have maintained extensive environmental monitoring programs on the
Great Lakes. As part of these efforts, the Canada Centre for Inland
Waters (CCIW), the Great Lakes Laboratory (GLL) of the State Univer-
sity of New York at Buffalo and the Center for Lake Erie Area Research
(CLEAR) at The Ohio State University have ongoing monitoring programs
on Lake Erie. As in the IFYGL intercomparisons (Robertson, Elder and
Davies, 1974) this study attempts to assess the differences that arise
between these organizations due to sampling and subsequent analysis, to
determine the feasibility of combining past and future data obtained by
the laboratories.
On 10 June 1975 a number of physical, chemical and biological
parameters were taken at two stations four miles northeast of Conneaut,
Ohio. This report discusses the statistical results of a split sample
analysis on six chemical properties of the water; in particular, the
results for the measures of soluble reactive phosphorus (SRP), total
phosphorus (TP), total soluble phosphorus (TSP), nitrate plus nitrite
(N+N), ammonia (NH3) and total reactive silicate (SiOg) are considered.
BACKGROUND
Three research groups are presently monitoring physical, chemical
and biological parameters in Lake Erie. These groups are the Canada
Centre for Inland Waters (R/V Northern Seal), Great Lakes Laboratory,
331
-------
State University of New York at Buffalo (R/V Dambach) and the Center
for Lake Erie Area Research, Ohio State University (R/V Hydra).
On 10 June 1975 these groups sampled two stations approximately
seven miles east of Conneaut, Ohio. Stations sampled included an
inshore station, (3 miles off shore) designated 023 (lat. 42°02'4811, long.
80°27'08") and a more lakeward station (7 miles off shore ) designated
024 (lat. 42°05'54", long. 88O29'00"). Station depths were 14-14.7 m
(023) and 22-22.7m (024). Sampling depths were determined via a cast
bathythermograph (model OC-10/3, range 0-200').
The levels of stratification of the lake at Stations 23 and 24 at the
time of the intercomparison were approximately as indicated below (Fig-
ure 141). The dots at each station represent the (nominal) sampling
depths. At the time of sampling, the lake level (as measured at Erie,
Pa.) was approximately 1.25 m above low water datum (LWD).
Sampling began at Station 23 at 0930 and Station 24 at 1130. The
R/V Hydra anchored at both sites while the other ships did not. Apart
from drift at Station 24 the ships were within 1/4 mile of each other.
At Station 23 the winds were from the ENE at about 18 knots. These
winds diminished to 15 knots and switched to the NE at Station 24.
Waves increased from approximately one foot at Station 23 to two to
three feet at Station 24 and hampered sample transfer between boats.
The wind conditions at Station 24 caused the unanchored ship to drift
shoreward. The study was truncated after sampling at this station and all
ships returned to Conneaut Harbor to exchange samples from Station 24.
At each sampling horizon of each station 18 liters of water were
taken. This volume was split into three (six liter) subsamples and ex-
changed among the participating groups. Containers were acid-washed
and rinsed in distilled and double-distilled deionized water. Exchanges
occurred immediately after sampling on Station 23 (about 103O) and ap-
proximately one hour after sampling on Station 24. All times were EST.
A list of the parameter determinations made by each of the three
groups is given in Table 77. The following were selected for statistical
analysis: ammonia-nitrogen, nitrate-nitrate nitrogen, soluble reactive
phosphorous, total filtered phosphorous, total phosphorous (unfiltered) and
total reactive silicate. Among the nutrient parameters, the phosphorous
parameters are the only ones for which determinations were obtained by
all three groups. The nitrogen and silicate parameters were felt to be
of considerable intrinsic interest.
A summary of the operational details of the intercomparison is
given in Table 78. This table contains information concerning sites,
332
-------
CO
GO
CO
PORT BURWELL, ONT.
CONNEAUT, OHIO
24 23
Figure 141. Temperature profile (°C) Secchi depths and sampling sites,
-------
TABLE 77. PHYSICAL AND CHEMICAL PARAMETER
DETERMINATIONS MADE BY GLL, CCIW, CLEAR
Parameter
Temperature
Dissolved Oxygen
Conductivity
PH
Secchi Transp.
Transmittance
c
Chlorophyll a
GLL
X
X
X
X
X
-
CCIW
X
X
X
X
X
X
X
CLEAR
X
X
X
X
X
X
Parameter
Alkalinity
Ammonia(NH0)
O
Nitrate-Nitrite
(NANI)
Total N
Kjeldahl N
(Organic)
Nitrate (NOg)
Total Reactive
Silicate (SiOp)
GLL CCIW CLEAR
X - X
X X
- x x
LL| Vv'
X -
X -
- x x
oo
CO
Part. Organic
Nitrogen
Part. Carbon
(Total)
Part. Organic
Carbon
X
X
X
X
X
Soluble Reactive X X
Phosphorus (SRPH)
Total Soluble X X
Phosphorus (TSPH)
Total Phosphorus X X
(TOPH)
Chloride - X
X
X
X
-------
TABLE 78. SUMMARY OF OPERATIONAL DETAILS OF
1975 LAKE ERIE INTERCOMPARISON
co
co
en
Sampling 023
Sites and
Sampling
Depths
024
Sampling
Apparatus
Storage
N+N
NH3
SRP
TP
TSP
SiO0
GLL
1
7
11
1
11
21
Water Bottle
-
HgCl2-4°C,
plastic bottles
4°C plastic
bottles
HgCl2-4°C,
plastic bottles
4°C plastic
bottles
CCIW
1
7
13
1
11
19
Submersible
Pump
None*
-
None*
Sulfuric Acid-
Glass bottles
Sulfuric Acid-
Glass bottles
?
CLEAR
1
7
13
1
11
21
Water Bottle and
Submersible Pump
None*
None*
None*
1OO ml glass
bottles
4°C glass bottles
4°C plastic bottles
(continued)
-------
TABLE 78 (continued)
GO
CO
CTl
Analysis**
N+N
NH3
SRP
TP
TSP
Kjeldahl-
Nessler's
Ascorbic Acid
Ammonium Molybdate
Ascorbic Acid
Ammonium Molybdate
Ascorbic Acid
Ammonium Molybdate
SulFanilamide, Phosphoric Acid,
N apthy I ethy I en e Diamide Dihydro-
chloride
Sodium Nitroprissuide
Stannous Chloride
Ammonium Molybdate
Stannous Chloride
Ammonium Molybdate
Stannous Chloride
Ammonium Molybdate
Oxalic Acid, Ascorbic Acid
Ammonium Molybdate
* Samples filtered then analyzed as quickly as possible - shipboard.
** Both CCIW & CLEAR used similar equipment (Technicon AA11) and
reagents, GLL employed wet lab analytical methods.
-------
depths, sampling methodology, storage techniques, and analysis methods.
Terminology
Some terminology that *s referred to in subsequent sections of the
report is discussed below.
Sampling process includes overall handling of water samples from
the time the ship arrives on station until the samples are divided among
the groups for analysis. This consists of all aspects of obtaining the
water samples, i.e., determination of sampling sites and sampling hori-
zons, type and utilization of sampling devices and sampling containers,
and conditions and duration of storage of water samples on board ship
until they are subdivided for analysis.
Analysis process refers to the treatment of water samples from
the time they have been subdivided and transferred to the analysis
groups' ships until the final laboratory results are reported and recorded.
This includes duration and methods of handling and storage of samples on
board ships, methods and duration of storage and handling of samples in
the laboratories, reagents and procedures utilized in the analysis process,
and formulas and procedures for calculating and recording the analysis
results. The mixing of reagents and execution of the laboratory tests is
only a small portion of the entire analysis process. Disagreements in
analysis results can be due to differences in procedures anywhere within
the analysis process (for example, due to storage of samples or due to
ship vs. shore laboratory analyses). A much more detailed experiment
than the Lake Erie intercomparison would be necessary to separate the
individual effects of the various components of the analysis process.
Steps in that direction are reported by Robertson, Elder, and Davies
(1974).
Sampling group is defined as that group (CLEAR, CCIW or GLL)
which obtained and subdivided the water samples.
Analysis group is defined as that group (CLEAR, CCIW or GLL)
which analyzed the subsamples of all samples obtained by the three
sampling groups.
Subsample variation represents that portion of the variability
in analytic results due only to variability in the analytic process. It
excludes all effects due to variability among water samples. It is
estimated from differences among the various analytic results on the
same water samples (analysis group by sampling group by sampling
depth mean square). Subsample variation is sometimes referred to
in subsequent discussion as "subplot" variation.
337
-------
Whole sample variation is the variability that enters into compari-
sons among sampling groups or among sampling horizons. The whole
sample variation reflects both variability among different water samples
as well as variability among individual analyses performed on the same
water sample. The variability among water samples is due both to
inherent variability in the lake water as well as to differences among
the sampling processes. Estimates of whole sample variation are based
on comparisons among the averages of all determinations on the same
water sample (in particular, sampling group by sampling depth mean
square divided by number of analysis groups). It is sometimes referred
to as "whole plot" variation.
SUMMARY OF MAJOR FINDINGS
A considerable number of statistical analyses were performed on
the intercomparison data. Among the conclusions arrived at are the
following:
1 . For each parameter but ammonia, there are statistically signi-
ficant differences among analysis group average determinations.
Both analysts groups report similar average determinations for
ammonia, however the variabilities are rather different. At each
station, the analysis group differences are especially large for
soluble reactive phosphorous. (Note that a "statistically significant"
difference is not necessarily a "practically significant" difference
and vice versa. What is considered to be a "practically significant"
difference depends heavily on the uses to which the data are to
be put. A 10 percent discrepancy might be quite acceptable for some
purposes but totally out of the question for others.)
2. No significant sampling group differences are evident.
3. Sampling depth effects are important especially at Station 23.
4. No systematic differences are evident between measurements
on samples obtained via submersible pump or water bottles.
5. The average nutrient levels at Station 23 are consistently
higher than those at Station 24. Station 23 is more shoreward
thus more susceptable to shoreline processes (tributary inputs,
mixing, resuspension).
6. The subsample variability is about the same at stations 23 and
338
-------
24 but the whole sample variability is greater at Station 24 than at
Station 23. This is probably due to the rougher surface conditions
at Station 24 and the shoreward drift of two of the vessels. Both
factors would contribute to heterogeneity among the water samples.
7. The variability between replicate pump determinations is gener-
ally smaller than the whole sample variability. The variation
between replicate pump measurements is probably an underestimate
of variability. Silicate is an exception.
Results 1 and 2 are in direct agreement with results reported by
Robertson, Elder, and Davies (1974) based on a similar intercomparison
that was carried out in 1972 on Lake Ontario.
Experimental Data
The experimental data are given in Table 79, which contains informa-
tion about the analysis group (ANLGRP1 = CLEAR, 2 = CCIW, 3 =
GLL), sampling depth (in meters), sampling site (23 or 24), method by
which CLEAR collected comparable water samples (OSUCODW = water
bottle, A = pump replicate 1, B = pump replicate 2), soluble reactive
phosphorous (SRPH), total soluble phosphorous (TSPH), total phosphorous
(TOPH), nitrate-nitrite (NANI), ammonia (NH3), and total reactive silicate
(SiO ). Of the 66 observations contained in Table 79, the 54 observations
corresponding to OSUCOD = W represent the 18 CLEAR water bottle sam-
ples and the samples collected by CCIW (18) and GLL (18). These 54
observations contain all the information pertaining to comparisons among
analysis groups and most of the information pertaining to comparisons
among sampling groups. The primary data analyses were carried out on
these 54 observations.
The CLEAR replicate pump measurements correspond to OSUCOD =
A or B. There are three pairs of such replicate pump measurements at
each sampling site. Comparisons between the replicate pump measure-
ments yield information about pump sample to pump sample variability.
Comparisons between the pump and water bottle measurements yield
information about the presence of any systematic discrepancies between
the two methods of sampling. Comparisons of CLEAR pump and water
bottle measurements were carried out on these 12 observations.
Blank spaces in the various data columns correspond to missing
data. For example, CCIW did not perform ammonia analyses, so that
when ANLGRP = 2 the Nh^ column is blank.
339
-------
TABLE 79.
BASIC DATA ON SIX CHEMICAL PROPERTIES
OF WATER SAMPLES
OBS ANLGRP
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
1
1
1
2
2
2
3
3
3
1
1
1
1
1
1
1
1
1
2
2
2
3
3
3
1
1
1
SMPGRP
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
2
2
2
2
2
2
2
2
2
3
3
3
DEPTH
1
7
13
1
7
13
1
7
13
1
7
13
1
7
13
1
7
13
1
7
13
1
7
13
1
7
13
SITE
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
23
OSUCOD SRPH
W
W
W
W
W
W
W
W
W
A
A
A
B
B
B
W
W
W
W
W
W
W
W
W
W
W
W
11 .7
10.9
7.6
12.0
10. 0
7.8
14.2
14.2
15.6
11 .4
10.7
5.9
11 .0
11.6
6.2
10.9
11 .3
9.1
9.6
9.6
7.5
14.9
15.6
9.8
11.7
11 .8
7.7
TSPH
15.0
13.3
11 .0
18.0
15.0
15.0
14.2
19.2
16.3
14.8
15.4
12.1
15.0
14.0
12.0
14.9
20.6
12.0
15.6
14.8
11 .0
TOPH
28.0
19.8
26.6
30.0
31 .0
27.0
36.1
29.5
25.1
25.1
26.5
23.7
26.0
25.1
24.7
25.3
25.3
21 .8
26.0
26.0
26.0
33.2
32.4
30.2
27.8
32.2
25.0
NAN I
302
295
233
326
318
264
300
290
240
295
295
235
296
300
254
319
317
270
302
288
240
NH3
58.0
61 .3
51 .0
88.0
55.0
55.0
60.5
60.2
50.0
58.0
56.0
50.0
54.0
53.4
49.7
53.0
60.0
28.0
59.0
56.5
51 .0
SiO2
340
320
320
310
290
280
430
435
435
310
325
405
320
320
340
300
270
270
300
300
300
(continued)
-------
TABLE 79 (continued)
OBS ANLGRP SMPGRP DEPTH SITE OSUCOD SRPH TSPH TOPH NANI NH3 SiO2
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
1
1
1
2
2
2
3
3
3
1
1
1
1
1
1
1
1
1
2
2
2
3
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
2
2
2
2
2
2
2
13
1
11
21
1
11
21
1
11
21
1
11
21
1
11
21
1
11
21
1
11
21
1
23
23
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
24
W
W
W
W
W
W
W
W
W
W
W
A
A
A
B
B
B
W
W
W
W
W
W
W
11 .0
11 .0
7.5
15.6
14.2
15.6
4.6
1.6
2.5
3.8
1.3
1.8
6.3
7.0
5.6
5.0
2.1
1 .4
5.5
2.3
2.5
2.7
2.1
2.2
2.3
1.7
2.0
5.6
18.0
19.0
14.0
15.6
15.6
15.6
8.6
5.6
11 .6
8.9
6.5
8.0
6.3
7.0
7.7
7.9
6.2
6.2
8.0
6.9
8.0
5.6
31 .0
30.0
29.0
30.2
33.2
23.6
19.4
14.6
22.6
19.0
17.0
23.0
17.8
10.7
16.4
19.4
16.3
17.2
15.3
19.5
17.2
18.2
13.6
14.6
17.0
17.0
15.0
17.1
324
319
252
201
207
347
216
223
381
2O2
172
248
210
207
305
162
206
245
171
232
268
61 .0
59.0
42.0
36.5
31.3
45.7
39.0
28.0
44.0
38.1
27.3
46.5
36.5
31 .4
57.7
29.0
34.2
35.0
39.0
340
310
270
230
175
370
220
170
310
325
325
255
450
270
195
240
190
215
180
150
200
(continued)
-------
TABLE 79 (continued)
OBS ANLGRP SMPGRP DEPTH SITE OSUCOD SRPH TSPH TOPH NANI NHg SiO2
oo
-pi
ro
56
57
58
59
60
61
62
63
64
65
66
3
3
1
1
1
2
2
2
3
3
3
2
2
3
3
3
3
3
3
3
3
3
11
21
1
11
21
1
11
21
1
11
21
24
24
24
24
24
24
24
24
24
24
24
W
W
W
W
W
W
W
W
W
W
W
7.0
6.3
4.9
2.0
2.3
4.7
2.0
2.0
6.6
7.7
5.6
7.7
7.0
8.3
5.3
5.6
12.0
7.7
5.6
7.2
7.7
5.6
16.4
18.4
49.5
17.1
16.7
21.0
15.0
12.0
17.8
10.7
16.4
208
195
242
230
212
264
24.0
15.0
35.5
33.8
36.0
44.0
58.0
28.0
300
225
325
260
170
170
-------
Experimental Results
A considerable number of statistical analyses and data displays
were carried out on the intercomparison data. The main conclusions
from these analyses were summarized in the section titled "Summary of
Major Findings". This section contains a more extensive, but still rela-
tively brief and non technical discussion of the bases of the conclusions.
The principal result of this section is statistical evidence of differ-
ences among analysis groups but not among sampling groups. This is in
direct agreement with results reported by Robertson, Elder, and Davies
(1974) based on a similar intercomparison that was carried out in 1972
on Lake Ontario.
The intercomparison experiment was carried out at two sampling
stations (23 and 24) on Lake Erie. Because the horizon depths and
nutrient levels at the two sites were quite different and since the whole
sample variability at Station 24 exceeded that at Station 23 (for some
considerably), the data from each of the two stations was analyzed sepa-
rately. Thus, no variance estimates have been pooled across stations.
Some comparisons (particularly those involving analysis groups) might
be reexamined with variance estimates pooled across stations, for in-
stance where pooling seems sensible. The increased degrees of freedom
might increase the sensitivity of the tests.
In the course of several of the analyses, it appeared from the
data that certain determinations might be out line relative to the rest.
In such cases the data were reanalyzed after omitting the suspect value
or values. The results of both the original and the reanalyses are
reported.
Overall Average Values. Table 80 contains the overall average
values by station of each of the six nutrients. The averages are taken
over analysis group, sampling group, and sampling depth. It is immed-
iately evident that the average values are consistently higher at site 23
than at site 24. The discrepancies between the two sites vary from 19
percent to 66 percent of the site 23 average values.
This observation is in agreement with other data collected. Sta-
tion 23 is more shoreward, thus more susceptable to shoreline processes
(tributary inputs, mixing, sediment resuspension). Indeed, it appears
that station 23 is located deeper within a nutrient rich surface plume
(see Figure 141 and section on sampling depth effects) than Station 24.
343
-------
TABLE 80. OVERALL AVERAGE DETERMINATIONS BY SITE
Parameter (ppb)
Site 23 Avg.
Site 24 Avg.
Difference
co
-ti
Soluble reactive phosphorus
Total soluble phosphorus ffiltered")
Total phosphorus (unfiltered)
Nitrate - nitrite
Ammonia
Silicate
11.42 (11.48*)
15.07
28.57
289.94
55.27 (54.93**)
305.56
3.86
7.36
17.93 (16.71**)
233.89
35.33
227.78
7.56 (7.62*)
7.71
10.64 (11.86**)
56.05
19.94 (19.6***)
77.78
Reanalysis of site 23 soluble reactive phosphorus data after deleting observation 24.
** Reanalysis of site 24 total phosphorus data after deleting observation 58.
*** Reanalysis of site 23 ammonia data after deleting observation 7 and 24.
-------
Whole Sample, Subsample, and Pump to Pump Variability. The
intercomparison is an example of a split plot experiment. Comparisons
of analysis groups are made within the same water samples whereas
comparisons of sampling groups, depths, or sites are made among diffei
ent water samples. Thus comparisons of sampling groups or sampling
depths reflect whole sample variation whereas comparisons of analysis
groups reflect only subsample variation. This implies that different com-
parisons require different error yardsticks.
In addition to whole sample and subsample variability, another vari-
ability estimate can be obtained by comparing the determinations based
on the replicate CLEAR pump samples. Conceptually, these pump to
pump variability estimates should approximately agree with or exceed the
whole sample errors, but except for silicate they turn out to be about
half the value. Table 81 contains the estimated whole sample, subsample,
and pump to pump variances of each of the six chemical properties, by
site. Beside each variance estimate is the degrees of freedom on which
it is based. A number of comparisons can be made from the table.
1 . The whole sample variance estimates are consistently greater
at Station 24 than at Station 23. Although it is difficult to compare
variances based on just four degrees of freedom, there is some evi-
dence of greater whole sample variation at Station 24 than at Station
23.
2. Except for silicate, there is no evidence of greater subsample
variation at one station than at another.
3. There is some suggestion that pump to pump variation is
greater at Station 24 than at Station 23, although the evidence is
weaker than that for whole sample variations.
4. With the exception of silicate, the whole sample variation is
one and one half to three times the pump to pump variation.
Analysis Group Effects. The average determinations for the three
analysis groups (averaged over sampling group and depth) were obtained
and compared at each site. Table 82 contains, for each of the six chemi-
cal properties, the three analysis group averages, the estimated standard
errors (based on the subsample variation) appropriate for assessing dif-
ferences between pairs of averages, and the significance levels at which
the averages are significantly diffferent.
At site 23 the analysis group averages differ significantly for all
parameters but ammonia. (The ammonia determinations differ in varia-
bility but not in average value.) At site 24 there is statistical evidence
345
-------
TABLE 81 .
SUBSAMPLE, WHOLE SAMPLE, PUMP-TO-PUMP
VARIANCES - BY SITE
Station 23
Property
Subsample
Variation
d.f.
Whole Sample
Variation d.f.
Pump to Pump
Variation d.f.
soluble reactive phosphorous
total soluble phosphorous (filtered)
total phosphorous (unfiltered)
nitrate-nitrite
ammonia
silicate
2.62(0.24*)
3.16
3.71
27.47
82.31
(40.20***)
122.22
8(7*)
8
8
4
4
(2 )
4
0,27(0.31*)
0.60
1 .55
36.24
23.02
(6.84***)
77.78
(4*)
4
4
4
4
(4***)
4
0.18 3
no pump samples
0.63 3
12.5 3
3.98 3
4566.67 3
CO
-Pi
CTl
Station 24
Property Subsample Whole Sample Pump to
Variation d.f. Variation d.f. Variation
soluble reactive phosphorous
total soluble phosphorous (filtered)
total phosphorous (unfiltered)
nitrate-nitrite
ammonia
silicate
0.10 8
0.98 8
27.47(4.38**)(7**)
28.22 4
65 . 72 4
899.31 4
0.36 4
1 .87 4
22.71(4.80**)(4**)
1441.35 4
59 . 23 4
1 949 . 66 4
0.25
no pump
4.51
756.33
24.14
3708.33
* Reanalysis of site 23 soluble reactive phosphorus data after deleting observation
** Reanalysis of site 24 total phosphorus data after deleting observation 58.
*** Reanalysis of site 23 ammonia data after deleting observation 7 and 24.
Pump
d.f.
3
samples
3
3
3
3
24.
-------
TABLE 82. ANALYSIS GROUP AVERAGES - BY SITE
Property
Station 23
sol . react . phos .
tot. sol. Phos. (filtered)
tot. phos. ('unfiltered)
nitrate-nitrite
ammonia
silicate
station 24
sol . react . phos .
tot. sol. phos. (filtered)
tot. phos. (unfiltered)
nitrate-nitrite
ammonia
silicate
Analysis Group Averages
CLEAR CCIW GLL
10.30
(10.30*)
13.67
26.87
278 . 89
54.88
(54.88***)
317.78
2.77
7.26
20.70
(17.89**)
223 . 67
35.22
252 22
9.56
(9.56*)
15.56
28.44
301.00
^~^~
293 . 33
2.40
7.96
17.33
(17.33**)
244 . 1 1
203.33
14.41
(15.31*)
16.00
30.39
55.67
(54.75***)
6.41
6.87
15.74
(15.74**)
35.44
Std. Err. of
Difference d.f.
.765 8
(+) (7*)
.838 8
.905 8
2.47 4
4.27 4
(3. 54+++) (2***)
5.22 4
.147 8
.466 8
2.47 8
(++) (7**)
2.50 4
3.82 4
14.14 4
Significance
Level
.0008
(.0001*)
.05
.01
.002
.86-
(.97***)
.01
.0001
.12
8
( .20**)
.002
.96
.03
CO
*
**
***
Reanalysis of site 23 soluble reactive phosphorus data after deleting observation 24.
Reanalysis of site 24 total phosphorus data after deleting observation 58.
Reanalysis of site 23 ammonia data after deleting observations 7 and 24.
After reanalysis std. err. of CLEAR, CCIW diff = 0.232; std err of CLEAR, GLL
and CCIW, GLL diffs = 0.253
After reanalysis std. err. of CLEAR, CCIW and CLEAR, GLL diffs = 1.075; std. err. of
CCIW, GLL diff = 0.986
After reanalysis
-------
of analysis group differences for soluble reactive phosphorous, nitrate-
nitrite, and silicate. At both sites the analysis group differences are
especially large for soluble reactive phosphorous.
Several of the discrepancy patterns are consistent between sites.
GLL's soluble reactive phosphorous levels are relatively high at both
sites. CLEAR's nitrate-nitrite levels are smaller than CClW's and
CLEAR's silicate levels are greater than CCIW's at both sites. CLEAR's
ammonia levels have the same mean but are less variable than GLL's
at both sites.
Sampling Group Effects. The average determinations for the three
sampling groups (averaged over analysis group and depth) were obtained
and compared at each site. Table 83 contains, for each of the six chem-
ical properties, the three sampling group averages, their estimated stan-
dard errors (based on the whole sample variation), and the significance
levels at which they are significantly different.
As can be seen from Table 83 no significant sampling group effects
are evident at either site.
Sampling Depth Effects . The average determinations for the three
sampling depths (averaged over analysis group and sampling group) were
obtained and compared for each site. Table 84 contains, for each of the
six chemical properties, the three sampling depth averages, their esti-
mated standard errors (based on the whole sample variation), and the
significance levels at which they are significantly different.
At site 23 significant depth effects are evident for all six chemicals
properties. There is consistently a sharp drop in nutrient levels between
7 and 13 meters.
At site 24 the depth effects are either nonsignificant or at best
marginally significant. This is due to a combination of higher standard
errors at site 24 (especially for nitrate-nitrite, ammonia, and silicate)
and less pronounced changes with depth.
It is interesting to note the drop in concentration levels between 1
meter and 13 meters at Station 23 and (with the exception of nitrate-
nitrite) a similar drop in concentration levels between 1 meter and 11
meters at Station 24. However, at Station 24 the concentration levels
remain the same or actually rise between 11 meters and 21 meters.
Comparison of Pump and Water Bottle Samples Collected and
Analyzed by CLEAR. Water samples can be collected either by a sub-
mersible pump or by water bottle samplers. In order to determine
348
-------
TABLE 83. SAMPLING GROUP AVERAGES - BY SITE
Property
Station 23
sol . react . phos .
tot. sol. phos. (filtered)
tot. phos. (unflltered)
nitrate-nitrite
ammonia
silicate
Station 24
sol. react, phos.
tot. sol. phos. (filtered)
tot. phos. (unfiltered)
nitrate-nitrite
ammonia
silicate
Sampling Group Averages
CLEAR CCIW GLL
11 .56
(11 .56*)
15.22
29.23
289 . 67
61 .38
(56.23***)
310.00
3. 83
7.80
17.83
(17.83**)
262.50
37.42
245 . 83
10.92
(11 .82*)
14.53
27.36
292 . 67
49.68
(53.46***)
303.33
3.54
7.O6
16.37
(16.37**)
214.00
29.37
195.83
11 .79
(11 .79*)
15.47
29.11
287.50
54.75
(54.75***)
303 . 33
4.20
7.22
19.58
(17.21**)
225.17
39.22
241 .67
Std. Ern of
Difference d.f.
0.42 4
*
(+) (4*)
0.64 4
1 .02 4
4.92 4
3.92 4
(+++) (-4***^
7.20 4
0.48 4
1 .12 4
3.89 4
(-K-) (4**)
31.00 4
6.28 4
36.05 4
Significance
Level
/^/"\
.22
(.75*)
.40
.23
£+ 4
.61
.10
(.70***)
.61
.53
.79
.73
(.70**)
.36
.35
.40
* Reanalysis of site 23 soluble reactive phosphorus data after deleting observation 24.
** Reanalysis of site 24 total phosphorus data after deleting observation 58.
*** Reanalysis of site 23 ammonia data after deleting observations 7 and 24.
+ After reanalysis std err of CLEAR, GLL diff = 0.452; std err of CLEAR, CCIW and
CCIW, GLL diffs = 0.492
++ After reanalysis std err of CLEAR, CCIW diff = 1.79; std err of CLEAR, GLL, and
CO
-P>
UD
CCIW, GLL diffs = 1 .95
After reanalysis std err of CLEAR, CCIW diff
CCIW, GLL diffs = 2.53
3.02; std err of CLEAR, GLL, and
-------
TABLE 84. SAMPLING DEPTH AVERAGES - BY SITE
Property
Station 23
sol. react, phos.
tot. sol. phos. (filtered)
tot. phos. (unfiltered)
nitrate-nitrite
ammonia
silicate
Station 24
sol . react . phos .
tot. sol phos. (filtered)
tot. phos. (unfiltered)
nitrate-nitrite
ammonia
silicate
Sampling Depths Averages
1 meter
12.40
(12.40*)
15.68
29.73
311 .50
62.17
(57.01**)
318.33
1 meter
4.61
8.09
21 .87
(19.06**)
198.00
37.17
238.33
7 meter
12.07
(12.07*)
16.32
29.93
3O6.17
57.53
(57.53***)
301 .67
7 meter
3.60
6.73
14.68
(14.68**)
212.50
34.88
1 80 . 00
13 meter
9.80
(10.70*)
13.22
26.03
252.17
46.12
(49.90**)
296.67
13 meter
3.37
7.26
17.23
(17.23**)
291 .17
33.95
265.00
Std Err of
Difference d.f.
0.42 4
(+) (4*)
0.64 4
1 .02 4
4.92 4
3.92 4
(+++) (4***)
7.20 4
0.48 4
1.12 4
3.89 4
(++) (4**)
31.00 4
6.28 4
36.05 4
Significance
Level
0.008
(0.05)
O.O2
0.03
0.002
0.04
(O.O9***)
0.08
0.12
0.53
0.28
(0.20**)
0.08
0.87
0.17
oo
en
o
**
***
Reanalysts of site 23 soluble reactive phosphorus data after deleting observation 24.
Reanalysis of site 24 total phosphorus data after deleting observation 58.
Reanalysis of site 23 ammonia data after deleting observations 7 and 24.
After reanalysis std err of CLEAR, GLL diff = 0.452; std err of CLEAR, CCIW, and
CCIW, GLL diffs » 0.492
After reanalysis std err of CLEAR, CCIW diff = 1.79; std err CLEAR, GLL and CCIW,
GLL diffs = 1 .95
After reanalysis std err of CLEAR, CCIW diff = 3.02; std err CLEAR, GLL and CCIW,
GLL diffs = 2.53
-------
whether any systematic effects are introduced by the method of sampling,
CLEAR collected two pump samples and a water bottle sample at each of
the six site-depth combinations. Comparison of the replicate pump mea-
surements provides direct information about pump sample to pump sam-
ple variability. Comparison of pump and water bottle measurements
provides information concerning the existence of systematic effects due
to sampling method. The analysis results are summarized in Table 85.
This table shows that with the exception of silicate, the pump to pump
variability is consistently smaller than the whole sample variation and the
pump to pump variation at site 23 is smaller than that at site 24.
The "t-ratio" column summarizes the statistics to test for systema-
tic differences between the two sampling methods. The t-ratio is formed
by dividing the difference between average pump determination and water
bottle determination by an estimate of its standard error based on pump
to pump variation. Where the pump to pump variation is similar between
sites, the 3 degrees of freedom variance estimates at each site are
pooled to form a more precise 6 degrees of freedom variance estimate.
Generally speaking, no significant systematic differences between the two
sampling methods are apparent. However inferences based on just three
degrees of freedom or even six are not very powerful.
DISCUSSION
A number of questions and issues arose in the process of analyzing
the data. Several of these are indicated below:
1 . A number of determinations were identified that seemed to be
out of line. These were sometimes isolated determinations, but
also sometimes involved all the determinations from the same
water sample. Where possible outliers are present, the basic data
should be reviewed to determine whether or not the suspect data
are erroneous and if so, why. Are the outlying values due to
clerical errors, error in the analysis procedure, errors in mixing
or calibrating reagents, or mixups in samples? Can the appropri-
ate values be determined? Are these "out of line" observations
just normal variation?
2. Why is the variation between replicate pump samples generally
smaller than the whole sample variation? Which, if either, is a
true estimate of sample to sample variation? Why is this relation-
ship reversed for silicate?
3. There was some confusion about sampling depths at the bottoms
of both stations. At Station 23, both CLEAR and CCIW report
351
-------
TABLE 85. COMPARISON OF CLEAR PUMP AND
WATER BOTTLE MEASUREMENTS - BY SITE
Properties
Station 23
Soluble reactive phosphorus
Total soluble phosphorus (filtered)
Total phosphorus (unfiltered)
Nitrate-nitrite
Ammonia
Silicate
Station 24
Soluble reactive phosphorus
Total soluble phosphorus (filtered)
Total phosphorus (unfiltered)
Nitrate-Nitrite
Ammonia
Silicate
Pump
Average
9.4667
Water
Bottle
Msmnt.
10.0667
Pump to
Pump
Var.
0.18
Whole
Sample
Variation
0.31*
t-ratio
-1 .82
d.f.
6
Signifi-
cance
Level
0.12
no pump sample determinations made
25 . 1 833
275.8333
55.78
390.00
3 . 1 333
28.133
276.6667
56.77
326.67
2 . 900O
0.63
12.5
3.98
4566.67
0.25
1.55
36.24
6.84***
77.78
0.36
-5.27
-O.33
-0.70
1 .39
0.706
3
3
3
6
6
0.014
0.76
0.54
0.22
0.50
no pump sample determinations made
1 7 . 4833
224 . OOOO
39.58
303 . 33
1 8 . 8667
251 .6667
37.83
258.33
4.51
756.33
24.14
3708.33
4.80**
1441 .35
59.23
1949.66
-0.92
-1 .422
0.50
0.99
3
3
3
6
0.44
0.26
0.66
0.36
tn
Reanalysis of site 23 soluble reactive phosphorus data after deleting observation 24.
Reanalysis of site 24 total phosphorus data after deleting observation 58.
Reanalysis of site 23 ammonia data after deleting observations 7 and 24.
**
***
-------
sampling at 13 meters whereas GLL reports sampling at 11 meters.
At Station 24, both CLEAR and GLL report sampling at 21 meters
whereas CCIW reports sampling at 19 meters. These discrepancies
in sampling depths were ignored for the purpose of this analysis,
however they could possibly introduce bias or variability into the
data. This should be kept in mind when preparing protocols for
future intercomparisons.
4. Generally speaking, the sampling variation at Station 24 was
larger than that at Station 23, sometimes substantially so. Does
heterogeneity of samples vary substantially from station to station?
If so, this should be taken into account when designing future sam-
pling procedures and when analyzing the data.
5. It would be useful to repeat the intercomparison at several dis-
tinct points in time, for instance, weeks or months apart, and to
incorporate more replication at each time. This would provide
information as to whether there is time to time random variation of
the results within laboratories and as to whether the discrepancies
between laboratories are random or systematic. The separation of
systematic and random sources of variation might give clues about
their nature, thereby enabling sources of variation to be eliminated.
See Robertson, Elder, and Davies (1974) for an example of this.
REFERENCE
Robertson, Andrew, Floyd Elder and Tudor Davies. 1974. IFYGL chem-
ical intercorYparisons. Proc. 17th Conf. Great Lakes Res. Internat.
Assoc. Great 'Lakes Res. p. 682-696.
353
-------
TECHNICAL REPORT DATA
(Please read Instructions on the reverse before completing)
i. REPORT NO.
EPA-600/3-80-062
4. TITL
2.
ITLE AND SUBTITLE
Lake Erie Nutrient Control Program - An Assessment
of its Effectiveness in Controlling Lake
Eutrophication
6. PERFORMING ORGANIZATION CODE
3. RECIPIENT'S ACCESSI ON-NO.
5 REPORT DATE
JULY 1980 ISSUING DATE.
7. AUTHOR(S)
8. PERFORMING ORGANIZATION REPORT NO
Charles Herdendorf
9. PERFORMING ORGANIZATION NAME AND ADDRESS
The Ohio State University
484 West 12th Avenue
Columbus, Ohio 43210
10. PROGRAM ELEMENT NO.
11. CONTRACT/GRANT NO.
R802543
12. SPONSORING AGENCY NAME AND ADDRESS
Environmental Research Laboratory
Office of Research and Development
U. S. Environmental Protection Agency
Duluth, Minnesota 55804
13. TYPE OF REPORT AND PERIOD COVERED
Final 1973-76
14. SPONSORING AGENCY CODE
EPA/600/03
15. SUPPLEMENTARY NOTES
Large Lakes Research Station, ERL-Duluth
9311 Groh Road, Grosse He, Michigan 48138
16. ABSTRACT ~~~ " ~ ~~
A three-year assessment of nutrient control efforts was conducted in the western
and central basins of Lake Erie during the period June 1973 to June 1976. The
objective of the study was to determine recent trends in lake eutrophication and water
quality which may be related to recent attempts to control nutrient loadings to these
basins. The assessment was accomplished by visiting approximately 50 stations at
nearly monthly intervals during the ice-free periods. Over 25 water quality,
meteorologican and biological parameters were routinely determined shipboard or on
samples collected at a typical station. Measurements were taken at several depths
in order to characterize the various strata of water in the lake and to permit
volume-weighted calculation of nutrient concentrations and quantities. Data from
previous limnological surveys as far back as 1928 were compared with the results of
the present study to establish longterm trends, as well as recent trends since the
last comprehensive survey in 1970.
The fundamental conclusion of this assessment is that during the first half of
this decade no significant decrease in the loading of nutrients to Lake Erie has taken
place. Therefore, during this period the concentrations and quantities of nutrients
within the waters of the lake have remained relatively stable. An encouraging sign
of nutrient control is that although no decreases have been observed, the constant
increases which have taken place in preceeding decades have been stopped.
7.
KEY WORDS AND DOCUMENT ANALYSIS
DESCRIPTORS
b.IDENTIFIERS/OPEN ENDED TERMS C. COSATI Field/Group
Algae, Nutrient,
Benthos
Hydrodynamic
Lake Erie
06
3. DISTRIBUTION STATEMENT
Release Unlimited
19. SECURITY CLASS (This Report)
Unclassified
21. NO. OF PAGES
20. SECURITY CLASS (This page)
Unclassified
22. PRICE
380
EPA Form 2220-1 (9-73)
354
a U.S. GOVERNMENT PRINTING OFFICE 1980-657-165/0117
-------