WATER POLLUTION CONTROL RESEARCH SERIES
18050 EEC i2/71
Acid Mine Pollution Effects
on Lake Biology
U.S. ENVIRONMENTAL PROTECTION AGENCY
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WATER POLLUTION CONTROL RESEARCH SERIES
The Water Pollution Control Research Series describes
the results and progress -in the control and abatement
of pollution in our Nation's waters. They provide a
central source of information on the research, develop-
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Inquiries pertaining to Water Pollution Control Research
Reports should be directed to the Chief, Publications
Branch, Research Information Division, Research and
Monitoring, Environmental Protection Agency, Washington,
D. C. 20460.
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ACID MINE POLLUTION EFFECTS ON LAKE BIOLOGY
by
Ronald W. Smith
and
David G. Frey
Indiana University
Water Resources Research Center
1005 East Tenth Street
Bloomington, Indiana 1+7^01
for the
ENVIRONMENTAL PROTECTION AGENCY
Project #18050 EEC
Contract #53-3^2-25
December 1971
For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402 - Price $1.28
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EPA Review Notice
This report has been reviewed by the Environmental Protection
Agency and approved for publication. Approval does not
signify that the contents necessarily reflect the views and
policies of the Environmental Protection Agency, nor does the
mention of trade names or commercial products constitute
endorsement or recommendation for use.
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ABSTRACT
Six coal stripmine lakes in southern Indiana encompassing a pH range
of 2.5 to 8.2 were studied from July 1969 to December 1970. Generally,
differences between the lakes indicated successional trends with
increasing pH. Environmental trends in the surface waters included
increasing levels of dissolved oxygen and decreasing concentrations
of dissolved substances. These tendencies were somewhat obscured by
differences in the annual cycles of stratification, four of the lakes
proving to be unexpectedly meromictic. Biological changes associated
with increasing pH included increasing diversity and increasing homeo-
stasis. Biomass was influenced by both pH and circulation patterns
(meromixis vs. holomixis), and bottom fauna was further limited by the
steep-sided basin form. All the stripmine lakes had much higher solute
concentrations and lower biological diversity than a small local non-
stripmine reservoir studied as a control. A fertilization program in
one lake has apparently produced elimination of all rooted aquatic
plants, violent oscillations of plankton, and low fish populations.
It is suggested that sport fishing in stripmine lakes, not presently
very satisfactory, could be improved by management techniques adapted
to their unique limnological nature.
111
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COIOTEKITS
Section
I Conclusions 1
II Recommendations 3
III Introduction 5
IV Description of Study Area 11
V Methods 21
VI Results and Discussion 27
VII Appendices 8?
VIII Acknowledgments 121
IX Literature Cited 123
X List of Appendices 131
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FIGURES
No. Page
1 Bathymetric map of Lake I (depths in m). lU
2 Bathymetric map of Lake II (depths inm). 15
3 Bathymetric map of Lake III (depths in m). 16
k Bathymetric map of Lake IV (depths in m). 17
5 Bathymetric map of Lake V (depths in m). 18
6 Bathymetric map of Lake VI (depths in m). 19
7 Complex stratification in Lake III, 15 July 1970. k6
8 Ranges and medians of surface pH for the Missouri
and Indiana stripmine lakes. 51
9 Maximum (A) and minimum (B) open surface (no ice
cover) dissolved oxygen; and percent saturation
(C) at overall maximums (any depth, ice present or
absent) in Pike County lakes. 5^
10 Maximum turbidity (A) and transparency (B) in the
Missouri lakes. 55
11 Maximum surface turbidity (A) and transparency
(B) in the Pike County lakes. 56
12 Maximum conductivity in relation to median surface
pH in the Missouri (A) and Pike County (B) lakes. 58
13 Double heterograde pH curves associated with mayimum
summer stratification in meromictic Lakes III, IV,
and V. 60
lU Number of animal taxa recognized in the Pike County
lakes. 65
15 Animal diversity and pH in the Missouri and Indiana
lake series. 67
16 Biomass and environmental stress in the Pike County
lakes. 69
17 Depth distribution of unweighted total mean zoo-
plankton standing crop (individuals/m^). 72
vi
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No
18 Depth distribution of unweighted total mean
"benthic standing crop (mg/m"). 73
19 Depth distribution of unweighted mean Euplotes
standing crop (individuals/an). 7^
20 Seasonal cycles of total zooplankton standing
crop (individuals/m^), September 1969 to December
1970. 75
21 Seasonal cycles of total benthos standing crop
(mg/m2), September 1969 to December 1970. 76
22 Increase of fish, benthic predators, and combined
higher consumers in Indiana lake series. 8l
23 Distribution of benthic biomass with depth and
area in the Pike County lakes. 8*f
2k Time-depth diagram of pH variation in Lake I. 88
25 Time-depth diagram of pH variation in Lake II. 89
26 Time-depth diagram of pH variation in Lake III. 90
27 Time-depth diagram of pH variation in Lake IV. 91
28 Time-depth diagram of pH variation in Lake V. 92
29 Time-depth diagram of pH variation in Lake VT. 93
30 Time-depth diagram of temperature variation (°C)
in Lake I. 9k
31 Time-depth diagram of temperature variation (°C)
in Lake II. 95
32 Time-depth diagram of temperature variation (°C)
in Lake III. 96
33 Time-depth diagram of temperature variation (°C)
in Lake IV. 97
3^ Time-depth diagram of temperature variation (°C)
in Lake V. 98
35 Time-depth diagram of temperature variation (°C)
in Lake VT. 99
vii
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No. Page
36 Time-depth diagram of dissolved oxygen variation
(mg/1) in Lake I. 100
37 Time-depth diagram of dissolved oxygen variation
(mg/1) in Lake II. 101
38 Time-depth diagram of dissolved oxygen variation
(mg/1) in Lake III. 102
39 Time-depth diagram of dissolved oxygen (mg/1) in
Lake IV. 103
kO Time-depth diagram of dissolved oxygen variation
(mg/1) in Lake V. 104
4l Time-depth diagram of dissolved oxygen variation
(mg/1) in Lake VI. 105
42 Time-depth diagram of specific conductance
in Lake I. 106
43 Time-depth diagram of specific conductance (^5 -10°)
in Lake II. 10?
44 Time-depth diagram of specific conductance (Kpc»10°)
in Lake III. fp 108
45 Time-depth diagram of specific conductance (Koc'lO")
in Lake IV. 109
46 Time-depth diagram of specific conductance
in Lake V. 110
47 Time-depth diagram of specific conductance
in Lake VI. Ill
viii
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TABLES
No.
1 Some morphometric characteristics of the Pike
County study lakes and control lake. 13
2 Observed ranges of selected chemical and physical
parameters in surface (S) and bottom (B) waters 28
3 Means and standard errors of surface (S) and bottom
(B) concentrations of selected ions based on six
determinations (see Appendix 2). 30
k Known animal taxa of the Pike County lakes. 33
5 Mean percent volume-weighted standing crop of
zooplankton organisms (individuals/up). 36
6 Mean percent area-weighted standing crop of
benthic organisms (mg dry weight/m"). 36
7 Standing crop of fish biomass. 37
8 Results of benthos sampling in Lake III, 15 July
1970. k7
9 Morphometry and physiochemical data of the five
stripmine lakes in Missouri studied by Campbell
et aL. (I965a, 1965b)*. 1*9
10 Morphometric differences between the Missouri
and Indiana lake series. 50
11 Observed annual ranges of bottom temperatures in
the Pike County study lakes. 53
12 Coal stripmine lake types proposed by Parsons
(19^). 63
13 Faunal changes in the Pike County lakes. 66
1^ Overall mean animal standing crop in the Pike
County lakes. 66
15 Percent of total standing crop in the Pike County
lakes. 68
16 Index of environmental stress in Pike County lakes. 68
ix
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No.
17 Depth distribution of mean percent area-weighted
total benthos biomass (mg/m"). 70
18 Depth distribution of mean percent volume-weighted
total zooplankton biomass (ind./nP). 71
x
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SECTION I
CONCLUSIONS
1. Six coal stripmine lakes in a series of increasing pH exhibited
sequential differences in both environmental parameters and community
structure.
2. These differences were analogous to some developmental trends
that characterize ecological succession.
3. Due to factors of morphometry and water chemistry some, perhaps
many, coal stripmine lakes are meromictic.
h. This pattern of incomplete circulation can have marked influences
on the biological community of a mine lake, including depression of
the fish population.
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SECTION II
RECOMMENDATIONS
1. Large areas of abandoned coal striplands that are not presently
utilized could and should be reclaimed for residential, agricultural,
wildlife, and recreational purposes.
2. Sport fishing in the stripmine lakes is quite compatible with
such uses and should be incorporated into development plans for
stripmine areas.
3. Stripmine lakes to be managed for sport fishing should be monitored
regularly to determine their annual patterns of circulation and general
limnological characteristics.
k. Fish production could probably be increased markedly in typical
stripmine lakes by increasing the relative area of shallow littoral
zone, increasing the habitat diversity and by the cautious use of
fertili zation.
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SECTION III
INTRODUCTION
Surface mining for coal is carried out extensively in the United States.
By 1965 approximately 3.2 million acres (1.3 million hectares) had been
surface mined, about kl percent for coal (U.S.D.I. 196?). Coal stripping
is carried out primarily in the Appalachian region, the South, and the
Midwest, including well over 100,000 acres (Uo,500 hectares) in Indiana
by 1971. One result of the Area Stripmining Method (as opposed to Con-
tour Stripping), usually practiced in flat terrain such as most of the
Midwest, is the creation of large numbers of small lakes and ponds in
the final cuts. These bodies of water vary in surface area from a few
hundred square meters to several hectares.
In Indiana coal stripmining is almost entirely confined to the south-
western third of the state where Pennsylvanian age strata are exposed
(Wier 1969, personal communication). This area is part of the "Eastern
Interior Coal Province," which also includes most of Illinois. Strip-
mining has left several thousand large and small stripmine lakes in
this area. In Pike County, the location of the present study, there
are no less than 950 (by actual count on U.S. Geological Survey topo-
graphic maps) of these lakes. These "strip pits" as they are called
are rightly regarded as having considerable recreational potential (Bass
1969> personal communication). Some old stripmine regions have already
been developed to a limited extent as recreational and wildlife areas
by the Indiana Department of Natural Resources, some coal mining com-
panies, and a few local communities. Such areas are used primarily for
picnicking, camping, hunting, and fishing. Unfortunately, the poten-
tially great recreational value of these areas has not, in my view, been
fully recognized and developed. Few natural wilderness areas remain in
Indiana. The striplands, though not natural in origin, can provide
semi-wilderness areas partially replacing those natural ones that have
been lost.
One aim of the present study is to provide certain fundamental limnolog-
ies! information about stripmine lakes in Indiana as a basis for sound
management of these lakes. Techniques developed for encouraging fish
production in farm ponds, natural lakes, and reservoirs are unlikely to
be very satisfactory for stripmine lakes. It is my hope that this work
will contribute to greater realization of the recreational potential of
stripmine lakes in Indiana and elsewhere. A second and more theoreti-
cal aspect of this work is discussed below.
In the majority of instances the water of newly formed coal stripmine
lakes is moderately to highly acid. High concentrations of various ions
occur in such lakes. This unique water chemistry results from the
leaching of substances contained in the cast overburden (materials such
as shale, clay, and sandstone overlying the coal seams) into the lakes
by surface runoff and ground water. The acid condition in particular is
a result of the formation of sulfuric acid by the oxidation of iron
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sulfide. Typically,the chemistry of these lakes comes gradually to re-
semble that of the more natural small lakes and ponds of the region. The
rate of this evolutionary process is quite variable and depends on such
factors as the nature of the cast overburden, kinds and amounts of materi-
als exposed, vegetative cover, nature of the lake bottom materials, and
patterns of ground water drainage and rainfall. Thus there is no good
correlation between the chronological age of a mine lake and its chemical
condition (Lewis and Peters 1955, Campbell et al. 19&5&) • Some ^i-116
lakes do not undergo this entirely successional sequence. When the over-
burden associated with the coal is low in toxic and acid-forming materi-
als or when modern mining and reclamation practices are followed, the
lake need not begin its existence as an environment hostile to most kinds
of aquatic life.
The biota of highly acid mine lakes (pH 2.5 or less) is quite restricted.
As the chemical conditions of the water and substrate become more moder-
ate, the diversity and abundance of organisms increases. Acid mine lakes
may properly be regarded as an example of gross industrial pollution.
Yet for the limnologist, they offer a rather unique opportunity to study
the adaptive mechanisms by which certain organisms do manage to survive
in these extremely hostile environments and the successional patterns
that emerge as the ecosystem gradually comes to resemble a more natural
aquatic situation.
A variety of studies have been carried out on coal stripmine lakes.
Most have emphasized either the chemical and physical aspects or the
biological aspects. Few investigators have dealt with both ecosystem
components. The chemical and physical attributes of stripmine lakes
have, perhaps, received more attention than their biological communi-
ties. The most important of previous stripmine lake studies are summariz-
ed below.
The physical and chemical conditions that can occur in stripmine lakes
are reasonably well known, at least in their broad outlines. Studies
such as those of Lewis and Peters (1955), Dinsmore (1958), Simpson
(1961), Parsons (196U), Campbell et al. (I965a, 1965b), and the series
by Waller (1967), Gash (1968), and Tobaben (1969) have contributed to
knowledge of these conditions. Nevertheless, a strikingly different
pattern has emerged in the Indiana stripmine lakes dealt with in the
present paper.
Perhaps the most dramatic feature of coal stripmine lakes is their
acidity. pH values of 2.0 to 2.5 are rather common, and lakes of pH less
than 2.0 are known to occur. The only parallels in nature are volcanic
lakes in Japan and Indonesia and bog lakes (Hutchinson 1957). Strip-
mine lakes with pH values below 2.5 are usually in basins that were used
for washing coal. These "tipples," as they are called, differ from final-
cut lakes in having very shallow basins and bottoms covered with thick
layers of coal and shale fragments. In these lakes total acid may ex-
ceed 6,500 mg/1 as CaCOg (Campbell et al. 1965a).
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Because of the low pH and the nature of the surrounding materials (cast
overburden or "spoil banks"), a great variety of substances is dissolved
in the water of acid mine lakes, often in very high concentrations.
Sulfate may reach 12,000 mg/1, and iron, aluminum, copper, zinc, lead,
arsenic, free carbon dioxide, and sometimes other substances are un-
usually high. Aluminum, for example, is usually present in lake waters
in concentrations less than 0.1 mg/1 (Hutchinson 1957), whereas in strip-
mine lakes it may reach 180 mg/1 or more (Parsons 196^). Oxygen is
usually present in stripmine lakes in amounts considered adequate for
aquatic life. Calcium and magnesium are typically quite high. Perhaps
the most striking physical characteristic of these lakes is their range
of apparent colors. Very acid lakes often appear a deep, clear red-
brown or red-black. These red hues are probably caused by a combination
of iron compounds and humic substances leached from the exposed coal and
shale. Lakes of higher pH exhibit many shades of blue, yellow, green,
brown, and various combinations of these. Many lakes undergo striking
color changes at various times during the annual cycle because of algal
blooms, upwelling of deep water leading to precipitation of iron com-
pounds, and perhaps other causes. Conductivity due to dissolved sub-
stances is ordinarily high in the very acid lakes, reaching values of
12,000 yjnphos/cm or more in some instances. As noted above, these chemi-
cal and physical features change as the mine lake matures, eventually
approaching those of natural small water bodies in the area.
Studies emphasizing the biological aspects of stripmine lake ecosystems
have been mainly qualitative descriptions of the organisms present. In
a pioneering study, Lackey (1938, 1939) surveyed 92 localities (mine
lakes and streams) influenced by acid-mine drainage. He found very few
species in the most acid waters, but observed that these were often very
abundant. Lackey's work, incidentally, is one of the two studies known
to the writer that deal to any extent with stripmine lakes in Indiana.
The other is Riley's 1952 survey of abandoned coal striplands as wild-
life habitats. Most of the subsequent studies of mine lake biota have
been limited to one or a few taxa or qualitative surveys of organisms
present. Examples of such work includes the papers of Yeager (19^2),
Galler (19^8), Levin (19^8), Myers (19^8), Ruhr (1951), Maupin, Wells,
and Leist (195M, Bell (1956), Dixon (1957), Brewer (1958), Arata
(1959), Ehrle (1§60), Stockinger and Hays (i960), Harp and Campbell
(1967), and Houde arid Forney (1970).
Few studies have attempted to deal with the entire ecosystems of strip-
mine lakes. Riley (1965) has studied four lakes in Ohio, and Dinsmore
(1958) studied 12 in Pennsylvania. Unfortunately, both these works
suffer from inadequate sampling programs (samples taken at long inter-
v als and only at the surface). The best description to date of the
annual cycle in an acid-mine water ecosystem is that of Parsons (1968),
which, however, is concerned not with mine lakes but with a stream sub-
ject to acid-mine drainage. Parsons sampled 11 stations on Cedar Creek
in central Missouri at monthly intervals over a 27-month period. He
has provided a good account of the responses of the stream ecosystem to
acid pollution and of recovery from heavy pollution. His data indicate
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a pattern of linear succession somewhat analogous to that which occurs
in streams undergoing organic pollution.
The most comprehensive body of work on ecosystem changes in coal strip-
mine lakes is contained in several studies carried out in central
Missouri. Four small lakes were investigated in 19^0 by Crawford
(19^2). In 1950 these lakes were restudied by Heaton (1950, 1951) in
order to document changes associated with aging. Since 1962 Campbell
and his co-workers (Campbell et al. 1965a, 1965&) have resumed investi-
gation of three of these lakes. The increase in diversity of organisms
and the modification of chemical and physical characteristics associated
with aging are well documented in these studies.
Some very basic aspects of coal stripmine lake ecosystem, however, have
not been adequately described. A few of the more important features
awaiting careful study include;
1. The seasonal cycles and stratification patterns of environmental
parameters such as dissolved oxygen, pH, temperature, light penetration,
and dissolved substances, especially the differences in these cycles and
patterns between lakes of different pH.
2. The seasonal cycles and depth patterns of occurrence and abundance
of the organisms inhabiting mine lakes at different stages of recovery
from acid pollution.
3. The relative importance of autochthonous primary production and
allochthonous organic matter input (dead leaves, etc.) as food sources
in lakes at different stages.
U. The physiological significance for aquatic organisms of the unique
water chemistry, including adaptive mechanisms that allow survival in
very acid lakes.
5. The importance to aquatic life, particularly benthos and fish, of
the low habitat diversity found in mine lakes.
6. The significance for benthic organisms and fish (in terms of
spawning for example) of the typical U-shaped basins of most stripmine
lakes.
7. The changes in community structure and ecosystem dynamics that
accompany recovery from acid pollution.
The present paper attempts to remedy certain of these deficiencies. In
addition, it provides information basic to the establishment of water
quality standards and fisheries management practices for coal stripmine
lakes.
Coal stripmine lakes provide a nearly unique opportunity to analyze the
maturation of evolution of an ecosystem type. A regular sequence of
changes occurs in these lakes during recovery with regard to both the
biotic and abiotic components of the ecosystem. A major aim in this
8
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work has "been to use a graded series of mine lakes as a natural model
for learning about ecosystem evolution in general. I have assumed that
a set of lakes at different pH levels closely approximates a series of
stages in the ecological succession of a single lake during recovery
from acid pollution. Beginning with the idea that there should be regu-
lar patterns of change or trends in the ecosystem associated with in-
crease in pH, I have attempted to identify and quantify them with regard
to:
1. The physical and chemical components of the ecosystem, especially
the differences in annual cycles of pH, temperature, conductivity, tur-
bidity, dissolved oxygen, and major ions.
2. The diversity of species in the various biological components, in-
cluding rooted plants, algae, zooplankton, benthos, fish, and other ver-
tebrates .
3. Annual cycles of biological production.
h. Overall ecosystem organization including community structure,
patterns of energy flow, etc.
An attempt to evaluate the significance of allochthonous organic mater-
ials relative to autochthonous primary production was not successful be-
cause of the difficulty of measuring primary production in lakes with
such a unique water chemistry and because of disturbance of installations
by beavers, muskrats, and humans. This comparison is of such fundamental
importance that I hope to make another attempt in the near future.
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SECTION IV
DESCRIPTION OF STUDY AREA
The stripmine area studied by Corbett (1965) was chosen for this study,
because it is the closest area to Indiana University with large numbers
of mine lakes having the desired features. This area lies in Pike
County, Indiana, and is contained on the Augusta and Oakland City quad-
rangle maps of the U.S. Geological Survey. A survey of this area was
conducted from October 1968 to July 1969 for the purpose of selecting
suitable lakes for extended study according to the following major
criteria:
1. Accessibility. Lakes that could not be reached by motor vehicle or
that had steep banks precluding the launching of a heavy workboat were
rejected. Two of the lakes selected were not fully accessible during
wet weather because of a clay road surface. At such times, limited work
was accomplished from a canoe portaged in.
2. pH. Hydrogen ion concentration was used as a measure of acidity
and as an indication of other chemical characteristics. The «-itn was to
select a reasonable number of lakes embracing the widest possible pH
range.
An effort was made to select lakes that were as uniform as possible with
regard to morphometry and other factors, differing mainly in their de-
gree of recovery from acid pollution. About 110 lakes were visited dur-
ing the survey. Most of them were excluded because of inaccessibility.
Many of the very acid lakes were rejected because they were, in fact,
abandoned mine tipples rather than final-cut lakes.
The six lakes finally selected for study range in pH from 2.5 to 8.2.
All are in basins formed as the final cuts of stripmining operations.
The lakes have been designated by Roman numerals I through VI in order
of ascending pH. Lakes II through V were under continuous surveillance
from July 19&9 through December 1970. Lakes I and VI were monitored
throughout 1970. Following is a brief description of each of the lakes.
Lake I (Fig. l). This lake varies in apparent color from a rather tur-
bid red-brown ("tomato soup") to a brilliant red-black. It is located
in the SW £ Sec. 17, T 2S, R. 7W in Pike County. Lake I is, at three
meters, the shallowest of the study lakes, and it also has the least
area and volume (Table l). Lake I was formed in 19^0.
Lake II (Fig. 2). This lake has the long narrow outline typical of
stripmine lakes. The water is generally a very clear green due to low
turbidity. The most striking thing about Lake II is the uniformity of
chemical and physical conditions that it exhibits. This lake tends to
be remarkably constant in pH, dissolved substances, etc., both through-
out the water column and throughout the year. Thus the ranges of the
various parameters given in Tables 3 and k are quite restricted for Lake
11
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II as compared to the others. Lake II was formed in I960 and is located
in the NE £ Sec. k, T. 3S, R. 8w in Pike County.
Lake III (Fig. 3). This lake is located adjacent to Lake IV on one sub-
unit of the Patoka Fish and Game Area of the Indiana Department of Natu-
ral Resources. Lakes I, V, and VI are on other subunits of this manage-
ment area. Lake III has greater flow-through than the other study lakes.
It has, apparently as a result of this, greater short-term changes in
such parameters as pH, temperature, and turbidity. This effect is con-
fined mainly to the surface layers because of a stratification situation
considered below. Lake III has greater variation in pH from inlet to
outlet, from surface to bottom, and over the annual cycle than any of
the other lakes. Lakes III and IV were completed in 1958.
Lake IV (Fig. k). This lake is directly connected to Lake III by a very
short channel approximately k meters wide and 20 centimeters deep. The
direction of water flow is from IV to III at all times. This lake has
the greatest maximum depth of the six (Table l). Lake IV has had a resi-
dent fish population for several years (Bass 196^), and during times when
water quality moderates, these fish may invade Lake III. During the win-
ter of 1969-70 the stream that formerly flowed into the upper end of Lake
III changed its course to the lower end of Lake IV (and thence into Lake
III). No water quality or biotal changes have been observed that can
be attributed to this change in stream channel. Lakes III and IV are
located in the NW £ Sec. 12, T. 3S, R. 8w, Pike County, Indiana.
Lake V (Fig. 5). This is the most irregular of the lakes in shape.
There are five small islands in the main body of the lake and three long
"fingers" pointing to the southwest. Lake V is being fertilized during
the summer months by the Department of Natural Resources in an attempt
to increase fish production. The advisability of this procedure is con-
sidered below. Lake V was formed in 19^0 and is located in the SW ^ Sec.
17, T. 2S, R. 7W in Pike County.
Lake VI (Fig. 6). This lake is one of the smaller of the study lakes
and is roughly Y-shaped. Construction of an access road in the autumn
of 1969 allowed this lake to be added to the series as a relatively high
pH lake that was not fertilized or otherwise managed. Lake VI was form-
ed in 1950 and is located centrally on the border between Sees. 3 and k,
T. 3S, R.7W in Pike County.
Control Lake. This lake was selected as representative of non-stripmine
small lakes of the region. It is owned jointly by three rural families
upon whose property it was constructed in 1963 for recreational purposes,
primarily for fishing by the owners and their guests. Because of time
limitations and relatively poor access this lake was visited less fre-
quently than the others, but it has provided some comparative data on
water chemistry and biota. It is located in the SW £ Sec. 31, T. 2S, R.
6w in Pike County.
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Table 1 (see p. 17) contains a summary of morphometrie data for the six
study lakes and the control lake.
TABLE 1. Some morphometric characteristics of the Pike County study
lakes and
Lake
I
II
III
IV
V
VI
Control
Year
formed
19^0
i960
1958
1958
19^0
1950
1963
control
Total
length
(»)
M5
1,300
975
900
960
510
850
lake.
Mean
width
(m)
U3
73
k9
56
31
Ul
95
Maximum
depth
(m)
3.0
6.0
8.0
10.5
5.5
7.0
7.0
Mean
depth
(m)
1-9
3.8
5.6
5.5
3.U
k.k
3.0
Surface
area
(10 m )
17.7
9^.5
^7.5
50. k
29.5
20.8
80.8
Volume
(10 m )
3^.5
355.3
265.0
275.^
100.6
90.8
242.3
13
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N
X Sanpling Station
FIG. 1. Bathynetric gap jaf lake I (depths in «).
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VJ1
0 100 M
I 1 1
X Stapling Station
FIG. 2. Bathymetric nap of Lake II (depths in m)«
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N
X Stapling Stations
FIG. 3. Bathyaetrlc map of Lake III (depths in a).
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X Sampling Stations
FIG. 4. Bathymetric sap of Lake IV (depths in «][.
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100 M
H
X Sampling Stations
FIG. 5. Bathymetric aap of Lake V (depths in m).
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VQ
X Sampling Station
FIG. 6. Bathymetrlc map of Lake VI (depths in m).
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SECTION V
METHODS
Standard procedures and equipment have been used in this study whenever
they were available and applicable. The selection and execution of
methods was guided by the publications of the American Public Health
Association (1965), Golterman (1969), Lagler (1956), Ricker (1968),
Vollenweider (1969), and Welch (19W). The specific instruments and
techniques used are briefly noted below.
A. Lake morphometry
Bathymetric maps of the study lakes were constructed as follows:
a. An outline map of each lake was prepared from aerial
photographs and U.S. Geological Survey topographic maps.
b. Each lake was then measured in situ with a 250 m cord
graduated in 5-meter intervals plus the workboat with 1-meter gradua-
tions marked on the starboard gunwale. For each lake the total length
and the width at several places (widest, narrowest, several intermediates)
were measured.
c. Final outlines were drawn to scale based on the field
measurements.
d. Depth contours were added based in each case on at least
100 soundings.
e. Total areas and the areas within each contour interval
were determined by the Cross-section Paper method (Welch 19^8) •
f. Other data (mean depth, mean width, volume) were calculat-
ed from the area data according to methods given by Welch (19W) •
B. Sampling program
1. Schedule. Initially an effort was made to visit each lake at
two-week intervals. This proved impractical, mainly because of the time
required to process the resultant biological collections. Hence with
the addition of Lakes I and VI to the series, the schedule was revised
to one of three-to four-week sampling intervals. Weather occasionally
interfered with sampling, especially when ice cover was too thick to
break through with the boat but too thin to walk and work on with safety.
2. Stations. During the first six months of the study (July to
December 1969) horizontal variation in environmental parameters and
plankton was monitored by studying two to four stations in each lake.
The magnitude of horizontal variation encountered was quite small in «-Ti
cases. It was, therefore, judged profitable to extend the overall range
of the lake series by adding Lakes I and VI at the expense of the multiple
21
-------
stations. Throughout 1970 a single station in the deepest area of
each lake was sampled (exclusive of benthos grab samples and various
other kinds of biological collections).
C. Physical and chemical parameters
1. Water samples were collected with two types of non-metallic
water samplers, a 3-liter van Dohrn bottle and (very briefly during the
winter of 1970) a ^-liter Kemmerer. The water samples were transported
to the laboratory in polyethylene bottles.
2. Water temperature was measured in situ by means of a Whitney
Underwater Thermometer (thermistor type), Model TC-5A.
3. Light penetration was measured in situ with a Whitney Under-
water Light Meter.
U. Turbidity was measured in the laboratory with a Hellige
turbidimeter.
5. pH was measured in the laboratory with a Beckman Model N pH
meter on water samples adjusted to 25 C in a water bath.
6. Specific conductance was measured at 25 C by means of a
conductivity bridge manufactured by Industrial Instruments, Inc. (Model
RC).
?' Dissolved oxygen was determined in the laboratory on samples
fixed in the field. The iodine-difference procedure of Ohle (1953) was
used, because it compensates well for all inorganic oxidizing and re-
ducing substances present. Two dissolved oxygen meters (Yellow Springs
Instruments, Models 51 and 5^ RC) gave such erratic and patently false
results in the stripmine lakes that their use was impossible.
8. Ionic determinations were made professionally by Calgon
Corporation of Pittsburgh, Pennsylvania. Ions monitored were calcium,
magnesium, sodium, potassium, total iron, dissolved iron, manganese,
aluminum, chloride, sulfate, silica, nitrate, and total phosphate. In
addition, total acidity, free mineral acidity, hardness, and dissolved
solids were determined. Ions were monitored for the surface and bottom
strata on a schedule of approximately two-month sampling intervals (see
Appendix 2 for details).
D. Biological parameters
1* Plankton was collected in three different ways: (a) by means
of a Wisconsin style tow net, (b) with a 10-liter plexiglass plankton
trap of the sort designed by Schindler (1969), and (c) by centrifugation
of 1-liter water samples in a Foerst centrifuge. Both the tow net and
the bucket of the trap were made of number-20 bolting silk. All plankton
samples were preserved with acid Lugol's solution (Edmondson 1959)
22
-------
immediately upon collection, except when live collections were required
to facilitate identification. Both trap and centrifuge samples were
taken at 1-meter intervals throughout the water column. Depending on
the density of organisms, either the entire sample or appropriate
aliquots were counted microscopically.
2. Benthos was collected quantitatively with a Ponar Grab
Sampler. Samples were taken from a-13 areas of each lake, and initially,
at all depths. Early experience indicated, however, that samples taken
from the deepest parts of Lakes III, IV, and V never contained organ-
isms. Therefore, later sampling was conducted mainly in the shallower
areas of these lakes (where the bottom was within the mixoll mm* on).
An effort was made to sample each 1-meter depth interval proportionately
to its percentage of total bottom area.
Each collection consisted of ten samples (grabs), which was about the
maximum that could be processed carefully within a reasonable period of
time. Including initial rough sorting, specific sorting, counting,
drying, weighing, and recording, a minimum mean processing time of two
hours per sample (not including time spent in taxonomic identification)
was required to reach the stage of raw data. Thus the 670 samples pro-
cessed required about 1350 hours. Additional time could have been
devoted to the bottom fauna only at the expense of other aspects of the
study.
The ponar sampled an area of 0.05 m^ and collected approximately 10
kilograms (clay) or a bit less (leaves or loose detritus) of bottom
materials per grab. The samples were placed individually in 20-liter
plastic buckets for transport to the laboratory. Ordinarily, initial
processing was completed within 2k hours of collection. Each sample
was washed through a graded series of three large (35 x U8 cm) screens
of mesh sizes: 121.0 mm^, 1.0 mm^, and 0.25 mm^. Organisms were picked
from the sorting screens and preserved in formalin. Subsequently, each
species was counted and weighed (constant weight at 105 C). In addition,
qualitative surveys of the bottom fauna were conducted at irregular
intervals with dip nets and artificial substrate samplers of the
condenser type (Hester and Bendy 1962).
3. Fish were collected with funnel traps constructed of hardware
cloth (12 x 12 mm mesh) and by means of seining in shallow water.
Specimens kept for analyses were preserved in formalin. Vertebrates
other than fish (frogs, snakes, muskrats, and beavers) were not studied
specifically.
U. Taxonomic identification. In any study such as this, in
which one attempts to deal with all major ecological categories, the
broad taxonomic spectrum dealt with makes the identification of organ-
isms one of the more difficult and time-consuming tasks. Some taxa are
relatively straightforward, while others are virtually impossible for
the nonspecialist. In many aquatic groups, the immature stages are
not sufficiently well known to allow identification to specific level.
23
-------
Although the compilation of exhaustive species lists was not a major
objective of this work, as many taxa as possible have been identified
or at least recognized. The following works have been useful in this
regard: Ahlstrom (19^0, 19^3), Burt and Grossenheider (19611-), Chu
(19^9), Eddy (1957), Edmondson (1959), Fassett (1966), Hotchkiss (196?),
Jahn and Jahn (19^9), Muenscher (19UU), Needham and Needham (1962),
Needham and Westfall (1955), Nelson and Gerking (1968), Pennak (1953),
Prescott (196U), Smith (1950), and Usinger (1963).
5. Morphometric correction. The following method was adopted as
the most straightforward approach to compensating for morphometric
differences between the study lakes. For each category of organisms
(zooplankton and benthos) dealt with, in each lake, the data were treated
by 1-meter depth intervals. The mean standing crop for each depth
interval was then multiplied by the percentage of the lake (percent area
for benthos and percent volume for zooplankton) at that depth. The sums
of these percent-weighted means are reported in Tables 5? 6, lU, 17,
and 18.
E. Experimental studies
1. Allocthonous organic matter entering the lakes in the form of
dead leaves was studied. Newly fallen leaves of the following four
types were collected in the autumn of 1969 from the shores of the study
lakes: River Birch (Betula nigra), Sycamore (Platanus occidentalis),
Cottonwood (Populus deltoides), and a mixture of the less common kinds
of leaves including Black Locust, Willow, Pines, Maples, etc. In each
group the leaves were thoroughly mixed, dried to constant weight at
105 C, and bagged in nylon mesh bags at 25 grams per bag. The bags
were sewn shut, and pieces of color-coded plastic tape were attached to
facilitate identification upon recovery. These bags were placed in
the lakes in Decmeber 1969 with the intention of recovering them after
1 to 1.5 years to obtain information concerning the relative decomposi-
tion rates in different lakes and of different kinds of leaves. These
leaf bags were apparently regarded by the resident aquatic mammals as
either food sources or hostile objects because during the first 6-8
months the bags in four of the lakes disappeared without a trace. Only
in Lake VI was there evidence of human interference with the experi-
mental installations of various sorts (rifle slugs in the fish trap
floats, artificial substrate samplers on the beach, etc.).
In conjunction with this work, an effort was made to estimate the amount
of annual input of leaves into each lake. Several trays of hardware
cloth (0.5 m area) were placed in each lake in early autumn of 1970 in
representative locations with respect to bank vegetation, prevailing
winds, etc. Together with their collections of fallen leaves they were
to have been recovered after completion of the autumn leaf fall. Un-
fortunately, the rate of deterioration of galvanized metal in acid mine
lakes was badly misjudged, so that when the time came to recover the
trays (late November and early December 1970) those in Lakes I, II, and
III were found to have been almost completely eaten away. It proved
-------
impossible to distinguish with any certainty between newly fallen
leaves and those from previous years. Hence the quadrat method
could not be used. Thus the attempt to study the relative import-
ance of allochthonous food sources came to naught.
2. Primary productivity was studied in situ by means of the
light-and-dark-bottle technique. Anomalous results such as an in-
crease of oxygen in the dark bottle, or remarkable increases or
decreases in both bottles, gave little confidence in the results.
Quite likely, these anomalies resulted from chemical oxygen demand
and the activities of chemosynthetic bacteria.
-------
SECTION VI
RESULTS AND DISCUSSION
The primary aim of this section is to analyze those trends in the study
lakes that seem to be associated with increasing pH. Such trends would
be expected in both the biotic and abiotic components of the ecosystems.
During the course of the study, however, it became apparent that a
second pattern, in addition to increasing pH, was present in the six
lakes. This was the persistent chemical stratification or meromixis of
Lakes III, IV, V, and VT. This pattern is clearly reflected in the
diagrams of annual variation in temperature and conductivity (Appendix
l) and in the differences between the surface and bottom waters in many
physical and chemical parameters (Tables 2 and 3). The temperature and
conductivity diagrams indicate that each of the meromictic lakes is be-
having as two separate lakes: an upper one that overturns twice annually
and a lower one that does not. This condition is substantiated by a
complete lack of overlap in the observed ranges of conductivity, and of
calcium, and other ions in the surface vs. bottom waters of Lakes III
through VT.
The influence of meromixis on the ecosystems of these lakes seems quite
marked in Lakes HI, IV, and V, but less so in Lake VI. It affects the
distribution of heat, turbidity, and dissolved substances, and the pene-
tration of light. These, in turn, have a profound influence on the bio-
logical communities of the lakes. The overall impact is such that mero-
mixis is probably as responsible as pH for the differences observed be-
tween the six ecosystems.
In the following discussion the ecosystem of each lake in turn is
briefly described. The emphasis in these descriptions has been placed
on annual cycles, the details of which may be found in the time depth
diagrams of Appendix 1. This is followed by a discussion of those
trends in the ecosystem series that seem attributable to the pH spectrum
or to the influence of meromixis. The pertinent data for this discussion
are presented in Tables 2 through 7.
A. Patterns within the lakes
Lake I was quite acid throughout the period of study, varying in surface
pH from 2.5 to 3.2 with 2.8-2.9 typical. The greatest vertical varia-
tion in pH observed in any of the lakes occurred in Lake I in March 1970
when a surface-to-bottom difference of two pH units (3.2-5.2) was noted
(Fig. 2b). Thermal stratification persisted throughout 1970 except for
periods of near homothermy in mid-August and from late September to mid-
October (Fig. 30). The integrity of stratification was maintained through
the warming period, indicating that the entire water mass was heated by
direct insolation. A maximum temperature difference of 10.2 C (surface-
to-bottom) was observed in mid-May 1970. Dissolved oxygen (Fig. 36)
was generally low, extremely so during July and August 1970 when the con-
centration was less than 1.0 mg/1 throughout the water column. This period
27
-------
TABLE 2. Observed ranges £f selected chemical and physical
parameters in surface (S) and bottom (B) waters.
Parameter
II
Lake
III IV
V
VI
Control
S 2.5 - 3.2 3.0 - 3.4 3.6 - 6.4 4.5 - 7.6 6.1 - 8.2 7.4 - 8.2 7.2 - 7.7
PH
B 2.8 - 5.2 3.0 - 3.4 4.0 - 6.4 5-9 - 6.5 6.3 - 7.0 6.6 - 7.2 6.9 - 7.1
Temperature
Dissolved
oxygen
(mg/1) *
S
B
1.4 -28.7 1.0 -31.7 5.6 -33.8 4.0 -29.9 0.8 -31.2 0.5 -28.9 7.0 -28.0
1.5 -24.3 6.3 -28.5 12.6 -14.4 9.7 -11.6 9.2 -14.7 9-0 -17.2 6.6 -10.1
S 0.2 - ^.9 2.8 - 9.1 3.9 - 9-9 6.2 -11.3 7.0 -14.6 7.2 -11.8 8.3 -11.8
(7.1) (10.4) (9.9) (12.7) (25.7) (16.8)
B 0.0 - Q.h 0.0 - 8.9 0.0 0.0 0.0 0.0 -11.6 0.0 -10.2
Total S 3615-^395 2515-27^5 965-1910 1140-28^0 1545-2660 1200-1570 90- 95
dissolved
solids (mg/1) B 4695-5^95 2570-3060 ij-7^5-5610 3290-5015 3015-3^35 1830-2751 100-135
* Numbers in parentheses indicate overall oxygen maximum
-------
TABLE 2. Continued
Parameter
Specific
conductance
(K25 ' 106)
Turbidity
(mg/1 Si02)
1 percent
light (m)
I
s 1868-4350
B 1*000-4350
S 2.3 - 7.6
B 12.2-60.0
2.5 - 3.0+
II
2273-2778
2273-2857
0.3 - 5.0
0.7 - 6.9
6.0+
III
1042-1887
2128-4651
0.8 -12.0
8.6 -69.0
1.5 - 7.0
Lake
IV
781-2778
29*41-4167
0.8 - 7.7
9.2 -20.0
2.0 - 7.5
V
1786-2500
2850-3636
1.7 -15.0
4.3 -55.0
1.0 - 3.0
VI
1250-1667
1786-2778
0.7 - 8.9
1.0 -11.0
5.5 - 7.0+
Control
132- 145
156- 213
2.9 - 9.3
26.0-47.0
2.5 - 4.0
-------
Mo1
Ald
TABLE 3. Means and standard errors of surface (S) and bottom (B) concentrations of
selected ions based on six determinations (see Appendix 2j.
Lake
Ion I II III IV V VI Control
S 425.7*23.0 334.0* 9.3 172.7*15.0 273.8*37.4 299-3*19-1 163.3* 5.4 12.0± 1.0
C&++
B 434.3*12.4 340.7*10.6 481.7* 6.5 443.0*10.1 405.3*12.2 298.7*16.4 17.0± 4.0
S 314.0*16.3 183.2* 5-1 128.3*13.0 170.5*22.8 192.3*22.2 120.7*19-1 6.50*0.71
Mg++
B 1*08.0*32.0 195.5* 9-7 501.2*16.2 339.2*16.8 325.0*12.6 213.0*13.3 7.00*3.00
S 81.7* 7.9 ^.75 ±1.05 5.68 ±2.18 1.09 * 0.6o 0.68 ±0.31 O.l4 *0.09 0.40*0.32
Fed
B 180.2*28.1 5.68 ±0.94 l8l.8±27.8 209.2*53.9 48.4 * 8.5 1.71 *0.74 9.25*8.25
S 50.5 * 2.3 33.6 * 0.6 8.78 ±0.69 5.68 ±l.o4 3.73 *1.46 o.l4 *o.o8 0.28*0.03
B 61.3 * 0.8 33.9 * 1.0 31.1 * 0.7 23.9 * 1.9 20.5 * 2.3 8.70 ±4.04 4.10*3-90
S 35.8 * 5.6 12.9 * 1.2 0.48 ±0.23 0.50 *0.39 0.095*0.103 0.083*0.098 0.037*0.042
B 32.0 * 8.0 12.6 * 1.3 0.54 ±0.24 0.12 ±0.17 0.10 ±0.10 0.037*0.045 1.26*0.72
d = dissolved
-------
TABLE 3. Continued
Lake
Ion I II III IV V VI Control
S 15.0 * 1.0 44.0 * 1.1 21.2 * 1.5 28-0 * 2.7 15-5 * 1.5 48.8 * 2.6 5.05*0.55
Na+
B 20.7 * 0.7 45.2 * 1.8 36.0 * 1.6 1*0.3 * 2.2 20.5 * 1.3 67.7 * 2.5 5.00*0.60
S 4.65 *0.24 5.32 ±0.15 3.90 *0.32 4.33 *0.35 3.88 ±0.19 4.45 ±0.16 1.70*0.50
K*
B 6.67 *o.4l 5.32 ±0.15 15.7 * 3.2 15-9 * 5.4 5.33 *0.24 6.62 ±0.45 2.30*0.50
S 2.87*0.14 0.83*0.01 0.20*0.01 0.13 *0.04 0.075*0.0^5 0. 0^1*0. 01^ 0.025*0.019
Znd
H B 2.68*0. iK) 0.91 *0.01 0.33*0.06 0.083*0.016 0.060*0.015 0.050*0.012 0.025*0.019
S 2726 * 103 1702 * 27 987 * 89 1397 * 178 151*1* * 117 912 * 35 55.0*5.0
T.H.
B 3315 * 166 1773 - 65 3651 - 69 292^ ± 159 2^67 4 65 1656 4 35 117.5117.5
S 2928 * 150 1671 i ^7 873 - 9^ I2l»6 t 193 1396 - 137 798 - 6k 32.5- 2.6
SO^
B 3to8 - 55 1775 - 83 3221 4 79 2365 ± 109 1996 ± 105 1267 * 101 15.0*10.0
S 10.0 * 6.1 4.67 4i.i6 17.2 ± 0.9 21.8 4 o.l 4.67 *0.52 3.33*0.45 2.50*1.50
Cl"
B 11.3 4 6.0 4.67 4i.i6 9-50 to. 76 21.7 4 5.4 4.25 ±1.08 4.33 4o.75 2.7542.25
T.H. = Total hardness, the stun of Ca, Mg, Fe, Mn, Al, and Zn as CaC
-------
to
IU
TABLE 3. Continued
Lake
Ion I II III IV V VI Control
S 43.2 ± 2.8 27.0 ± 1.2 11.2 ± 0.6 8.1? ±0.l6 10.2 ±0.70 1.88 ±0.15 5.50±2.U6
SiOg
B hk.O ± 2.2 27.0 ± l.o 15.0 ± 1.9 lU.o ± 3.1 15.8 ± l.l 5-95 ±0.83 7.00±0.89
-------
TABLE k. Known animal taxa of the Pike County lakes.
Taxon
II
Lake
III IV
V
VI
CILIOPHORA
Euplotes sp.
PLATYHELMINTHES
Turbellarian
ROTIFERA
Brachionus urceolaris
Keratella quadrata
Monostyla sp.
Keratella cochlearis
Testudinella patina
Filinia sp.
Hexarthra sp.
Unidentified rotifer 1
Unidentified rotifer 2
Chromogaster sp.
L€cane sp.
Brachionus angularis
Unidentified rotifer 3
Unidentified rotifer 4
Brachionus caudatus
B. calyciflorus
Filinia longiseta
Asplanchna sp.
Polyarthra sp.
Unidentified rotifer 5
Brachionus quadridentatus
ANNELIDA
Oligochaeta
Hirudinea
CRUSTACEA
Cyclops sp.
Calanoids
X
X
XXX
XXX
XXX
X X
X X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
33
-------
TABLE U. Continued
Lake
Taxon I II III IV V
Daphnia spp. x x x x
Ceriodaphnia sp. x x x x
Bosmina longirostris
Ostracods x
Cambarus (?) sp. x x
INSECTA
Collembola x
Ephemerid
Pantala flavescens x
Anax amazili x
Tramea Lacerata x x
Celithemis elisa x x x
Libellula sp. x x x
Gomphus consanguis x x x
Epicordulia princeps x
Erythemis sp. x x
Platythemis subornata x
Gomphus spicatus x
Ladona deplanata x
Tetragoneuria sp. x
Libellula pulchella x
Unidentified zygopteran x
Ischnura sp. x x
Argia sp. x x x
Enallagma sp. x
Sigara (?) sp. x x x x x
Gerrid
Sialis sp. x x x x
Unidentified trichopteran 1 x x
Unidentified trichopteran 2 x
Unidentified trichopteran 3 x
Berosus sp. x x x x
Dinetus sp. x x
Peltodytes sp. x x x x
Laccophilus sp. x
VI
x
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
-------
TABLE
Continued
Taxon
Ilybius sp.
Gyrimis sp.
Haliplus sp.
Heleidae
Tendipes sp.
Chironomid spp.
Unidentified dipteran
Chaoborus sp.
Unidentified tipulid
MDLLUSCA
Physa sp.
Lymnaea sp.
Helisoma sp.
Sphaerium sp.
VERTEBRATA
Lepomis cyanellus
L. macrochirus
Chaenobryttus coronarius
Micropterus salmoides
Minytrema melanops
Micropterus punctulatus
FunduLus notatus
Ondatra zibethica
Castor canadensis
Lake
I II III IV V VI
X
X
X
X X X X X X
X X X X X X
XXX
X
X X X X X
X X X X
X X X X
X
X
X
X X X X
XXX
X X
XXX
X
X
X
XX XX
X X
Total known taxa
12
23 32
53
35
-------
TABLE 5. Mean percent volume-weighted standing crop of
zooplankton organisms (individuals/m-*).
Lake
Taxon I II HI IV V VI
Rotifera 8,330 531,l4o 3,950 19,490 2,016,580 173,HO
Cladocera 54 0 1 23 52 12,530
Copepoda 0 0 1,365 3,564 4l,590 13,872
Euplotes 0 670 2,445 5,300 104,150 90
Total * 8,384 531,810 7,761 28,377 2,162,372 199,602
Percent of Lake V 0.4 24.6 0.4 1.3 100 9.2
Number of samples 40 112 l47 158 90 80
Vertical series 10 17 16 l4 15 10
*It is assumed for the purpose of rough comparisons that differential
size is approximately compensated by a higher reproductive rate of the
smaller organisms.
TABLE 6. Mean percent area-weighted standing crop £f
* _ _ . / O*
benthic
organisms (mg dry weight/m^) .
Herbivores
Predators
Total
Percent of Lake II
Percent predators
Number of samples
I
551
22
573
40.5
3.8
4o
II
1,373
43
l,4l6
100
2.9
120
III
5U
12
66
4.7
18.2
l4o
Lake
IV
74
21
95
6.7
22.1
130
V
37
183
220
15.5
83.2
130
VI
508
255
763
53.9
33.4
110
36
-------
TABLE 7. Standing crop of fish biomass,
Lake
Total trap hours
Specimens taken
Trap-hours/specimen
Live weight (g)
g/trap-hour
x weight (g/specimen)
III
528
7
75. ^
81^8.6
1.61
121.0
rv
1*98
23
21.6
1.3W.U
2.70
58.5
V
738
12
61.5
711.6
0.95
59.5
VI
180
te
*.3
1,399.7
7.78
33.0
of extremely low oxygen was associated with high water temperature and
partial overturn — a combination that almost certainly produced a
high chemical oxygen demand by upwelling of reduced solutes from the
deeper anaerobic strata. The highest observed concentration of dissolved
oxygen was 7.2 mg/1 at a depth of 1.0 m under a 15-cm ice cover in late
January 1970. This was only about 55 percent saturation at the in situ
temperature of 3.3 C. Despite relatively high turbidity, light was
probably adequate for photosynthesis throughout this shallow lake at
most times. During much of 1970 light reaching the bottom equaled or
exceeded 1 percent of surface illumination.
In general, the dissolved substances as indicated by total dissolved
solids and conductivity (Table 2) and many individual ions (Table 3)
were high in Lake I. The surface water in particular had a much greater
load of solutes than the other mine lakes. This was not universally
true, however, since some ions (e.g., Na"1", K% and Cl~) were higher in
some of the other lakes. The deep water of Lakes III and IV exceeded
those of Lake I in concentrations of the major cations and, in Lake
III, in total solutes.
Although it might be argued that Lake I is meromictic, it exhibited
virtual surfaee-to-bottom uniformity of water quality (pH 2.8-2.8,
temperature 20.6-20.1 C, conductivity ^350-^350 |j,mhos) on 29 September
1970. This lack of vertical stratification and the presence of
dissolved oxygen in relatively deep water (O.k mg/1 at 2.5 m) suggests
that autumnal overturn was complete although of short duration, as
indicated by the stratification of physical and chemical parameters on
the preceding and following sampling occasions (8 September, 1^ October).
On the latter date, the beginning of inverse thermal stratification was
observed (surface 7.9, bottom 9.5 C). Thus Lake I is probably best re-
garded as having a marked tendency toward meromixis that is imperfectly
realized because of its relatively shallow unprotected basin.
37
-------
The biological communities of Lake I were marked by very low fauna!
diversity (Table k). In the zooplankton, for example, only five taxa
were recognized compared to 20 and 21 species, respectively, in Lakes
V and VI. A total of 12 animal taxa were found in Lake I. Standing
crop of zooplankton (Table 5) was low, Lake I ranking fifth and only
slightly exceeding Lake III. In total benthic biomass, however, Lake
I ranked third, with markedly greater standing crop biomass than III,
IV, and V (Table 6). In both zooplankton and benthos a single species
dominated. These were the rotifer Brachionus urceolaris and the midge
larva Tendipes sp. which made up, respectively, about 99 percent and
85 percent of total biomass. Fish were not present in Lake I. Muskrats
maintained a lodge in a stand of cattails (Typha angustifolia) at the
northern end of the lake. Other aquatic vertebrates (amphibians, rep-
tiles) were never observed, even though several species were common
around the shores of Lake V a scant hundred meters distant.
Lake II differed in several ways from the other mine lakes. It was
marked by relatively great uniformity of physics and chemistry both over
the annual cycle and throughout the water mass. Total pH variation
during the 18-month study period was 3.0 to 3.^- (Fig. 25). A weak
stratification of pH (never exceeding a surf ace-to-bottom difference
of 0.3) sometimes developed during periods of thermal stratification.
Although this lake differed little from Lake I in surface pH, it was
quite different from Lake I in various limnological and biological
features. Many of these differences were at least partly related to
stratification differences.
The thermal regime of Lake II (Fig. 31) was marked by long periods of
homothermy and relatively small temperature differences during strati-
fication (the only observations of surface-to-bottom differences greater
than 5.0 C were on 3 May and 21 May 1970). The maximum observed dis-
solved oxygen was 10.5 mg/1 (at 1.0 m under 21 cm ice in late January
1970), which, at 3.2 C, constituted approximately 80 percent saturation
(Fig. 37). In midsummer dissolved oxygen values were quite low through-
out the water mass (3-5 mg/1 or less in July and August 1969 and again
in August 1970). Thus Lake II shared with Lake I the combination of
low pH, high water temperatures, and low dissolved oxygen throughout
the water mass in midsummer. This environmental combination is probably
strongly limiting to many species of aquatic organisms. Lake II had
greater transparency than the others. Light at the bottom (6.0 m) was
never less than 1 percent of surface illumination and frequently
exceeded 10 percent.
The surface and bottom ranges for total dissolved solids and for
conductivity were more similar than in the other lakes (Table 2).
The surface concentrations of total dissolved solids and of most ions
in Lake II ranked second to Lake I. In deep water, however, concentra-
tion values were exceeded by Lakes III, IV, and V and closely approached
by Lake VI.
38
-------
Although little different in pH, Lake II had almost twice as many
animal taxa (23) as Lake I (a 92 percent increase). This increase
was most striking for the insects, with 11 taxa added and but one
lost. It seems likely that this greater fauna! diversity is related
to the lower concentrations of solutes in Lake II.
Zooplankton standing crop was high, second only to Lake V. As in
Lake I, most of this biomass (over 98 percent) was due to Brachionus
urceolaris. Lake II had the greatest standing crop of benthos,
nearly twice that of second-ranked Lake VI, of which about 8U percent
was Tendipes sp. larvae. As in Lake I, over 95 percent of the total
benthic standing crop consisted of herbivores. There were no fish in
Lake II even though local fishermen reported having "stocked" it with
ttieir surplus catches from time to time. Because of its clear green
water, Lake II was popular locally for swimming and water skiing until
the area was closed for re-mining in January 1971.
Lake III had the most complex pattern of pH variation (Fig. 26) of the
six mine lakes studied. The water mass near the outlet was consistently
0.2 to 1.0 pH units lower than that near the inlet. This was the only
regular horizontal difference discovered in any of the lakes. Lake
III had a greater rate of flow-through than any other, and the higher
"downstream" acidity was probably due to the leaching of acid-forming
materials from a coal seam, mainly underwater, exposed in the old high-
wall. An acid heterograde pg^ curve with the minimum at intermediate
depth (Hutchinson 1957) was typical from July 1969 to August 1970.
This type of pH curve is usually associated with meromixis. Lake III
had a fairly well-marked trend toward higher pH during 1970, with no
values less than 4.5 observed after April.
The pattern of temperature variation (Pig. 32) indicates the meromictic
nature of Lake III very clearly. The high surface reading of 33.8 C
in late May 1970 was the highest temperature measured in any of the
lakes during the study. Temperature variation at the bottom encompassed
the very small overall difference of 1.8 C, and the maxima and the minima
lagged two to three months behind those at the surface. Dissolved
oxygen was restricted to the uppermost 2 m during July and August 1969,
leaving about two thirds of the water mass anaerobic (Fig. 38). This
was repeated in slightly less extreme form in the summer of 1970.
Even during such extreme midsummer conditions, in contrast to Lakes I
and II, oxygen at the surface rarely fell below 4.0 mg/1. The highest
observed concentration was 9-9 mg/1 under about 2 cm ice in late
December 1970 (about 87 percent saturation at 8.4 C). The deepest
observed penetration of dissolved oxygen was 6.0 m during the periods
of partial overturn in spring and autumn.
Turbidity values (Table 2) were generally higher in Lake III than in
the other lakes, especially in deep water. This was mainly due to the
precipitation of dissolved substances (as hydroxides of iron, etc.)
at the interface between oxygenated and anaerobic strata. Usually, the
turbidity maximum occurred at the level of this interface. During the
39
-------
partial overturns of spring and autumn, the surface water was colored
a dark red-brown by the precipitates that resulted from an upwelling
of anaerobic water with its heavy load of reduced solutes. The depth
of the 1 percent level of light penetration typically approximated
the depth of the oxygen interface. Thus the depth of the 1 percent
level was typically 2.0-U.Omin summer and U.5-6.0 m in winter. A
marked decrease in turbidity in May 1970 was accompanied by increased
light penetration to a 1 percent level of 7.0 m. This greater trans-
parency was maintained, except for periods of turbidity from clay
particles washed in by heavy rains throughout the remainder of 1970.
The contrast between the surface and bottom concentrations of solutes
in Lake III was extreme (Tables 2 and 3). Its deep water had the
highest conductivity and the greatest mean concentrations of total
dissolved solids, calcium, magnesium, and total hardness of all the
lakes studied. The surface water, however; had the lowest observed
concentration of total solids (965 mg/1 in March 1970) and ranked
fifth in mean total dissolved solids, calcium, magnesium, sulfate, and
total hardness (exceeding but slightly the surface values of Lake VI).
Faunal diversity (Table ^) was greater in Lake III than in Lake II
with nine more taxa (about a 39 percent increase) recognized. In Lake
II the zooplankton and benthos were quite unequal, contributing respec-
tively 26 percent and 70 percent of the known taxa. In Lake III,
however, the two groups were nearly equal with benthos constituting
^7 percent and zooplankton 50 percent. This sharp reduction in rela-
tive diversity of the benthos probably resulted from the anaerobic
bottom conditions and very steep-sided basin shape of Lake III. Some
major groups, including Mollusca, Annelida, and fish made their first
appearance in Lake III.
The standing crop of both zooplankton and benthos was lowest in Lake
III. Fish were not observed or trapped in that lake before June 1970.
This may have been due to the moderation of pH and other environmental
conditions as noted above. Although small (unidentified) fish were
observed on two or three occasions, the only specimens taken in over
500 trap hours were seven green sunfish caught on 30 October 1970. It
appears likely that colonization of Lake III by the established fish
populations of Lake IV will occur if the environment remains moderate.
Lake IV had, with the exception of a single low surface reading (^-.5
under ice cover on 26 January 1970), a quite moderate range of pH
variation (Fig. 27). Apart from this unusual value, the lowest observed
surface pH was 6.35- As in Lake III, the pH minimum (very rarely as
low as 5*7) frequently occurred at intermediate depth.
The pattern of temperature variation (Fig. 33) was also quite similar to
that in Lake III. Again there was a very slight overall annual varia-
tion at the bottom (1-9 C). Such uniformity of temperature in deep
water is typical of meromictic lakes, in general, but as in the
shallower lakes of this study, this may be modified by depth and
-------
transparency. Dissolved oxygen (Fig. 39) was generally greater than
in the previous lakes. Even in midsummer surface values were rarely
less than about 7.0 mg/1, with oxygen penetration at least 5.0 m.
The maximum observed penetration was 5.6 mg/1, at 8.0 m in December
1970. The overall maximum dissolved oxygen observed was 12.7 mg/1 at
1.0 m under about 1 cm of ice in December 1969, which at 9.1 C was
about 110 percent saturation. The highest surface concentration
(11.3 mg/1 in December 1970) was about 100 percent saturation. Lake
IV was the first in the series in which 100 percent saturation at the
surface was usual.
Turbidity was less and transparency slightly greater in Lake IV than
in Lake III. Again the 1 percent level of light penetration was often
associated with the oxygen interface.
As in Lake III, the ranges of such general indicators of dissolved
substances as total dissolved solids, conductivity, and total hardness
were non-overlapping for the surface and bottom strata. For most of
the major ions, Lake IV ranked fourth in surface concentration but
third or even second in bottom concentration. It had the highest mean
concentrations of dissolved iron, potassium, and chloride.
Forty-nine animal taxa were recognized in Lake IV (Table U), an increase
of 17 species or 53 percent over Lake III. There was a striking
increase in the number of vertebrate species, from one each in Lakes I,
II, and III to a peak of seven in Lake IV. Four more plankton species,
and seven more benthic species, were found in Lake IV than in Lake III.
Zooplankton biomass (Table 5) was greater in Lake IV than in Lakes I
and III but very much less than in the other three lakes. Similarly,
the biomass of benthic animals (Table 6) was much less than in Lakes
I, II, V, and VI and only slightly greater than in Lake III. Only
Lake VI exceeded Lake IV in fish biomass.
Lake V had a range of pH variation nearly encompassing those of Lakes
IV and VI. Thus a more distinct series might have been had by omitting
Lake V. It has been retained in the series, however, because it is
managed for sport fishing (i.e., fertilized in summer) by the Indiana
Department of Natural Resources. The other lakes, in contrast, have
not been disturbed since their formation except for the construction
of launching ramps.
Only a moderate overall range of pH variation (Fig. 28) was observed
in Lake V. The maximum vertical variation was found in late July when
pH decreased from 8.2 at the surface to 6.3 at 2.0 m, then increased
slightly to 6.6 at the bottom. As in Lakes III and IV, a pH minimum
at intermediate depth was not unusual.
Although the overall deepwater temperature variation was greater than
in Lakes III and IV (Fig. 3*0, Lake V must also be considered
meromictic. Some partial mixing obviously occurred at the times of
overturn, but the data for dissolved oxygen and conductivity (Figs. 40
-------
and U6) and for solutes (Tables 2 and 3) indicate rather complete chemical
separation of the surface and bottom water masses. Oxygen (Fig. ^0)
penetrated to the U.O m level in March 1970 but never reached the deepest
(6.0 m) strata. The maximum surface value observed was lU.6 mg/1 in
June 1970 (about 190 percent saturation). In September 1969 a maximum
concentration of 25.7 mg/1 was observed at 2.0 m, about 278 percent
saturation. Dissolved oxygen in excess of 20 mg/1 was observed at inter-
mediate depths on several other occasions. Surface oxygen was never
observed to be less than 7.0 mg/1. In contrast to the previous lakes,
the highest dissolved oxygen concentrations in Lake V occurred during
the warmest part of the year (April through October), probably because
of massive algal blooms that were apparently a response to the fertili-
zation of the lake.
Such blooms and the accompanying large populations of zooplankton were,
in part, responsible for the high turbidity and restricted light pene-
tration characteristic of Lake V (Table 2). There was, also, a marked
turbidity maximum at the lower limit of dissolved oxygen as in Lakes
III and IV. It was usual for the 1 percent level of light penetration
to occur within that boundary zone.
Again in Lake V, the general indicators of dissolved substances (total
dissolved solids, conductivity, total hardness) had non-overlapping
ranges for the surface and bottom water masses (Tables 2 and 3) • The
dichotomy is less well-marked than in Lakes III and IV, primarily due to
higher surface concentrations of major ions (Ca, Mg, SO^) in Lake V.
Although the total number of animal taxa known in Lake V is the same
as in IV (Table k), there are marked differences in fauna between the
two. Of the 15 taxa present in Lake IV but not in Lake V, only one was
replaced by a taxon found in one of the previous lakes in the series
(an unidentified rotifer species also found in Lake III). Four species
of fish were taken in Lake V compared to five each in Lakes IV and VI.
The standing crop of zooplankton in Lake V (Table 5) greatly exceeded
that of any other lake. The weighted mean value of more than two million
individuals per nr indicates a quite impressive level of production.
In contrast to plankton, benthic biomass (Table 6) was relatively low
in Lake V. Here, again, extensive areas of the bottom are anaerobic
throughout most or all of the annual cycle. One curious feature of
the benthos in Lake V is the apparent imbalance between the herbivores
and the predators. Usually, the generalization is made that each trophic
level is exceeded by the previous one upon which it feeds by a factor
of about 10. This will be reflected in the biomass unless a marked
difference exists in the rate of production. Since there is no very
good reason to think that the herbivore and predator benthos in these
lakes differ greatly in their rates of productivity (typically one
generation annually), the biomass of herbivores is expected to be
roughly ten times that of predators. It can, of course, be many times
greater. This expectation was reasonably well fulfilled in all the
lakes except Lake V, where the predators were about five times greater
-------
in biomass than the herbivores. It seems likely that these predators,
mainly immature Odonata, feed on the abundant small fish that feed in
turn on the heavy crop of plankton and seek shelter in the same beds
of dead leaves where the dragonfly naiads are most common. Many small
(2-5 cm) specimens of Lepomis and Micropterus were taken in the benthic
samples along with the dragonflies.
Fish biomass (Table 7) was quite low in Lake V. No very satisfactory
explanation for this has been found. The plankton populations could
certainly support very large numbers of small fish adequately. In
fact, large samples of immature fish were seined easily in shallow
water. It may be that the physical and chemical conditions, especially
in summer when oxygen is restricted to the upper 2 m, limit the survival
of more mature fish.
Lake VI had relatively little pH variation throughout the annual cycle
(Fig. 29). The surface water remained slightly alkaline, and there
was no overlap in pH ranges for the surface and bottom water masses
(Table 2). Temperature variation (Fig. 35) resembled the pattern
found in Lakes III-V, but was less extreme. A relatively wide tempera-
ture range (8.6 C) was measured in deep water. Dissolved oxygen (Fig.
Ul) varied moderately in the surface waters. The deepest stratum
(below 6.0 m) was essentially anaerobic from late August to mid-October
1970, but had reasonable oxygen concentrations during most of the
remainder of the year. The maximum observed concentration was 16.8 mg/1
at 5«0 m in September 1970 (about 205 percent saturation). This high
concentration at intermediate depth was due to photosynthetic oxygen
production by the dense growth of Potamogeton that carpeted Lake VI at
all depths from about 1.0 to 6.0 m in summer and early autumn. The
greatest observed open surface concentration was 11.8 mg/1 in December
1970 constituting about 110 percent saturation. In general, Lake VI had
lower turbidity and greater light penetration than any other except Lake
II. Light intensity at the bottom frequently equalled or exceeded 1
percent of surface illumination, except that in late summer and autumn
the plant growth mentioned above shaded out the deepest meter or two.
Solute concentrations in Lake VI, especially in deep water, were notice-
ably less than in the other lakes. This was true for total dissolved
solids, conductivity, and most ions (Tables 2 and 3). The observed
ranges of surface and bottom concentrations, however, were mostly nen-
overlapping, indicating rather rigid chemical stratification. Lake VI
was meromictic during 1970, but the monimolimnion was essentially
limited to the deepest meter. The stability of this stratification
was probably much less than in Lakes III-V.
Animal diversity was greatest in Lake VT, although habitat diversity
appeared to be no greater than, for example, in Lake IV. Four more
taxa were found than in Lakes IV and V. The fauna of Lake VI was
reasonably well balanced with 22 taxa of zooplankton, 25 of benthos,
and 5 of fish. Beavers and muskrats, which occurred in Lakes IV^and
V, were never observed in Lake VI.
-------
The standing crop of zooplankton in Lake VI (Table 5) ranked third, about
9 percent of that in Lake V. The weighted mean total of nearly 200,000
individuals per m^ was much greater than those of Lakes I, III, and-IV.
In benthic biomass (Table 6), Lake VT ranked second to Lake II. Although
the standing crop was only about half as great, benthic production in Lake
VI may have actually been even higher assuming fairly heavy predation by
its relatively large fish populations. The biomass of fish in Lake VI
(Table 7) was about three times that in second-ranked Lake IV. The average
weight of individuals caught, however, was markedly less than in Lakes
III, IV, and V.
The Control Lake had an observed pH range of 6.9-7.7 > within the ranges of
both Lakes V and VI. It must be noted, however, that this and the other
data for the control (Tables 2 and 3) are based on only two complete series
of samples (June and December 1970). In June, the lake was thermally
stratified and anaerobic below k.O m. In December, it was virtually homo-
thermal (7.0 C at the surface gradually shading to 6.6 at the bottom) with
10.0 mg/1 or more dissolved oxygen throughout. This lake probably follows
the temperature regime typical of small ponds and lakes in southern
Indiana. If so, it circulates twice annually in spring and autumn, with
the possibility of prolonged circulation in years of mild winters with no
ice cover.
Turbidity (Table 2) was higher in the control lake than in several of
the stripmine lakes. This turbidity was partly due to very dense plankton
concentrations on both sampling dates. The 1 percent level of light pene-
tration was at 2.5 m in June and 4.0 in December.
The most striking llmnological difference between the control and the
stripmine lakes was, of course, its very much lower concentrations of
dissolved substances (Tables 2 and 3). Even Lake VI exceeded the control
by a factor greater than ten in total dissolved solids and conductivity.
The control did have, however, higher concentrations of certain ions
(notably Fe, Al, and Si02) than Lake VI.
Because of the difficulty of access (it was necessary for the landowner
to hitch my boat trailer to his tractor and tow it to the lake and back
through his pasture) and limitations of time, extensive quantitative
sampling of the fauna was not undertaken. No absolute statements about
either the diversity or standing crop of animals are possible for the
control. A few general statements based on limited sampling and informal
observation can be made. There is no reasonable doubt that the diversity
of most ecological groups was greater than in any of the stripmine lakes.
A rapid survey of the plankton samples taken in June and December revealed
not fewer than 25 taxa of zooplankton, more than were found by careful
enumeration of 100 or more samples each (including qualitative samples)
taken at all seasons in any mine lake. In the few quantitative samples
taken, the number of individuals (total) was higher than in even those
from Lake V. The benthic fauna appeared not to exceed Lake VI in
biomass, but was probably more diverse. Fish were taken in the control
-------
lake by angling with rod and reel on several occasions. From personal
experience and the success of other anglers, it is clear that the
several species introduced have become well established.
B. Patterns between the lakes
The lakes were originally arranged on the basis of increasing pH but,
as previously noted, that simple series was unexpectedly complicated by
the meromictic nature of some of the lakes. Although the two factors
are superimposed in nature, an attempt has been made to distinguish
between the influences on the ecosystems of pH and of meromixis. The
very interesting patterns of environmental stratification that often
develop in a meromictic lake, illustrated by the following example
drawn from Lake III, can have pronounced effects on the biocoenosis of
the lake ecosystem.
1. The influence of meromixis. On the morning of 15 July 1970
the weather was pleasant~T27 C, overcast, light variable breeze) at Lake
III, and the green water was unusually transparent for that Lake
(1$ = 6.5 m). There was, in fact, no visible surface evidence of the
remarkable complexity that existed in the subsurface environment (Fig.
7) • The metalimnion was shallow (1-3 m), and the curve of dissolved
oxygen closely paralleled that of temperature, with anaerobic conditions
below 3-0 m. The vertical distribution of pH was quite complex with
minima at 2-3 m and 6-7 m and maxima at 0-1, U-5, and 8 m. Turbidity
increased abruptly from 6.6 mg/1 at the surface to 19 mg/1 at 3 m.
Dissolved substances, as indicated by conductivity, increased stepwise
with major steps (chemoclines) at 5-6 and 7-8 m. Thus even by these
rather crude measures, environmental stratification was indeed complex.
The influence of this kind of stratification on the benthic fauna is
evident in Table 8, which summarizes the results of bottom fauna samples
taken on the day in question. This particular example was chosen partly
because the benthic grab samples included a greater range of depths than
was usual in Lake III. At this time, only about 2k percent of the
bottom area was within the aerobic zone (shallower than 3 m).
Two kinds of benthic animals were taken, both of them immature stages
of Diptera. These two groups (the."true midges" of the family
Chironomidae and the "biting midges" or "no-see-ums" of the family
Heleidae) occurred in the benthos of all six lakes. Four of the six
samples taken above'the lower limit of dissolved oxygen (3.0 m) had at
least one organism, while none of the four samples taken deeper had any
macroscopic benthic animals. Densities of approximately 100 larvae per
grab sample (as in samples 3 and h) indicate a density in the lake
bottom sediments on the order of 2,000 larvae per m2. This was quite
high for Lake III, but would be only a moderate density for Lake II.
Nine taxa of zooplankton organisms were taken on 15 July 1970. Most
of these occurred at all depths, with greatest density at 2 m and least
at 7-8 m. Total rotifers, for example, decreased from 10,000 individuals
-------
pH 5
0
10
20 Turbidity
10
M
1.
20
2'
3-
U-
5.
6.
7-
8-
i
\
\
\
1000
10 D.O.
2000
3000
FIG. 7. Complex stratification in Lake III, 15 July 1970.
per m-3 at 2 m to 2300 per nP at 8 m. In contrast, one rotifer (Hexarthra
sp.) was most numerous at 8 m. Plankton animals are better able to
tolerate anaerobic conditions than benthos, or so it seems from these
results. The zooplankters in the deepest stratum can reach oxygen by
swimming or floating vertically upward a distance of a few meters.
Benthic animals, on the other hand, would have to crawl much longer
distances horizontally (assuming they do not swim very readily) . In
fact, however, the chironomid and heleid larvae were faculatively
pelagic in Lakes I, II, and III, occurring sporadically in plankton
samples at various depths and times without apparent relationship to
any of the environmental parameters measured. Nevertheless, these
larvae were recovered from only about 9»5 percent of the bottom samples
taken at anaerobic depths, always in relatively low densities. Thus
meromixis appears to have a much more limiting effect on bottom fauna
than on zooplankton (see p. 68-74 ).
It is not known how common meromixis is in coal stripmine lakes.
Campbell et, al. (I965a) presented evidence for meromixis in their shallow
acid Lake AjtpH 2.0-2.9, maximum depth 2.0 m) in central Missouri. Even
1*6
-------
TABLE 8. Results of benthos sampling in
Lake III, 15 July 1970.
Grab sample number Depth Benthic animals collected Number
(m)
•L 0«3 Chironomid larvae 6
Heleid larvae 1
2 0.5 None 0
3
U
5
6
7
8
9
10
1.0
1.5
2.0
2.5
U.O
5.0
5.0
, 8.0
Chironomid larvae
Heleid larvae
Chironomid larvae
None
Heleid larvae
None
None
None
None
91
1
99
0
1
0
0
0
0
when this lake was "thoroughly mixed" with an outboard motor boat, it
reverted within 2*J- hours to an extremely stratified condition (Campbell,
personal communication). Lake AX is not a typical stripmine lake but
rather a shallow abandoned mine "tipple," that is, an artificial lake
constructed for the purpose of washing coal. It differs from a final-cut
lake in having a saucer-shaped basin and also in having a bottom composed
of fragments of coal and shale (from fine powder to fairly large chunks)
to a depth of perhaps a meter or more. The other lakes studied by
Campbell's group showed no tendency toward meromixis, but rather exhibited
only limited and transient stratification (Crawford 19^2, Heaton 1951).
The stripmine ponds in southern Illinois studied by Lewis and Peters
(1955) were generally stratified in summer, but circulated throughout
the rest of the year. Similarly, Parsons (19&0 found stratification
to be absent or limited to summer in six acid mine lakes in Missouri.
Simpson (1961) found no thermal stratification in three stripmine lakes in
southeastern Kansas. Two other sets of Kansas mine lakes studied by
Stockinger and Hays (I960) and by Maupin et &L. (196*0 exhibited only
moderate summer stratification and circulated during the remainder of the
-------
year. Riley's (1965) incomplete data indicate rather weak and imperma-
nent stratification in the stripmine lakes he studied in southern Ohio.
Finally, the three-year study of six lakes in Kansas (Waller 196?, Gash
1968, Tobaben 1969) disclosed no evidence of meromixis. On the con-
trary, some of these lakes exhibited only brief or incomplete stratifi-
cation before treatment with lime, and typical stratification after
treatment.
Thus meromixis would seem to be somewhat uncommon in coal stripmine
lakes; the present study being the first to document prolonged stratifi-
cation in such lakes of moderate pH and depth. Had this situation been
anticipated, the meromictic lakes might have been avoided because of
their obscuring effect on the pH series.
Certain characteristics of stripmine lakes seemingly predispose them to
behave in meromictic fashion. The water typically has a heavy load of
dissolved substances. During summer thermal stratification, the concen-
tration of solutes in the surface water can be decreased by dilution
with rain water and the precipitation of reduced solutes by oxygen. At
the same time, the dissolved substances in deep water can be increased
by the addition of water with a very high concentration of solutes
(leached from the cast overburden or from exposures of coal and shale
in the highwall by surface and subsurface drainage) which flows downward
to its density level. Further, solubility is higher for some substances
in deep water due to anaerobic conditions, and thus some precipitates
(e.g., hydroxides of iron and manganese) can be redissolved when they
settle below the lower limit of oxygen. If the density difference
resulting from these processes becomes greater than that caused by
cooling of the surface water in autumn, the chemical stratification
will be perpetuated through the winter months. Inverse thermal stratifi-
cation will exist until the following spring when partial overturn
returns the lake to summer thermal stratification. The narrow deep
basin shape (small surface area) of typical stripmine lakes and the
sheltering spoil banks and highwalls combine to reduce the effect of
wind mixing to a minimum. Thus even a relatively small density difference
can be perpetuated. The processes noted above (dilution, precipitation
and resolution, leaching) are in operation throughout the year, continu-
ally increasing the density difference between surface and bottom
strata. The stability of stratification is directly related to this
difference and may be great as in Lake III, or relatively little as in
Lake VI. In greater or lesser degree, however, it is clear that mero-
mixis is not directly related to pH and therefore disturbs the simple
pattern of increasing pH in the series. Hence the influence of mero-
mixis must be taken into account in the analysis of trends that follows.
2. Other stripmine lake studies; Missouri. It is desirable to
compare trends in the Indiana lakes with those found in other stripmine
lake studies. The Missouri series studied by Campbell and others is the
only group of lakes with sufficient information available for detailed
comparisons. Table 9 includes morphometric and environmental data for
the Missouri series.
-------
TABLE 9« Morphometry and physiochemical data of the five
stripmine lakes in Missouri studied by Campbell
et al. (1965a. 19b5b)*.
Lake
p *,
Area (m )
Max. depth (m)
Mean depth (m)
Volume (nP )
pH
Conductance
(^.mphos)
Total solids
(120 C)
Turbidity (ppm)
Secchi disc (m)
Calcium (ppm)
Magnesium (ppm)
Ferric iron
Ferrous iron
Total iron (ppm)
Manganese (ppm)
Aluminum (ppm)
Sodium (ppm)
Potassium (ppm)
Zinc (ppm)
Sulfate (ppm)
*1
1,300
1.8
l.l
1,^30
2.10-2.9
2,000-
13,600
1,098-
12,220
10-28
0.5-1.1
63-U5
31-232
73-loUo
0-2580
86-2730
3.9-95
228-U75
If. 6-28. 2
0.06-1.6
6.5-100
A2
6,830
1.2
1.1
7,500
2.8-3.2
1,200-
3,990
1,290-
5,350
<7-22
0.5-1.1
9-79
59-159
28-35
0.5-2.2
28-te
7.7-17.2
71-2Mi
5.1-9-9
0.75-1.5
0.8-2.3
777-7600 ^99-1367
*3
2,160
U.2
3.0
6,U80
3.U-U.1
320-560
211-815
<7-9
0.6-U.o
29-131
10-22
0.9-1.6
0-0.2
0.9-1.7
1.0-3.5
0.5-1.7
2.U-U.5
5.3-9-2
0.6-2.5
1^3-339
B
6,^90
5.1
3.0
19,^70
6.U-7.6
3UO-500
281-520
<7-19
0.8-3.5
10-105
17-25
0.02-0.3
0.0
0.02-0.3
0.0-8.9
0.01-0.05
5.7-8.7
3-9-6.3
0.0-0.1
75-336
D
3,885
3.9
2.5
9,700
6.3-7.9
110-220
120-360
10- 2U
0.5-1.5
10-U6
6-16
0.03-O.if
0.0
0.005-0.5
0.0-6.8
0.008-0.07
2.9-7.2
3.8-5.3
0.0-0.1
1.8-155
-------
TABLE 9. Continued
Lake Al ^ A B D
Silica (ppm Si02) 5.2-92.5 l4.8 7.9-27-5 0.3-4.5 1.0-6.2
Phosphate (ppm) 0.0-0.01 0.15-0.5 0.0-0.9 0.0-0.59 0.0-0.47
*Morphometry converted to metric system
It is evident from a comparison of Tables 1 and 9 that as a group the
Missouri lakes were much smaller and shallower than those of the present
study. The Indiana lakes had about 10 times the area, twice the depth,
and 20 times the volume (mean values) as the Missouri lakes (Table 10).
A second significant difference between the two series is that the pH
spectra represented were not identical. The overall range covered by
the Missouri series (2.0-7.9) was slightly greater and slightly lower
on the pH scale than that of the Indiana series (2.5-8.2). Neither
group of lakes was uniformly distributed within its range and there
was, unfortunately, no Missouri lake in the middle range (about 4.0-6.5).
The surface pH ranges and medians are indicated for the two series in
Fig. 8.
These two basic differences between the Indiana and Missouri lakes must
be borne in mind, as comparisons between the two groups are made in the
following sections.
TABLE 10. Morphometric differences between the
Mean of series
Area (m2)
Mean depth (m)
Maximum depth (m)
Volume
Missouri and Indiana lake
Missouri
4,100
2.1
3.2
8,900
series.
Indiana
43,4oo
4.1
6.7
187,000
50
-------
8'
•• 7 •
B D 6 •
5 •
1
1 1 A3 3'
T ^i?
1 2 •
f
" ™ 0
V
rv
1 t in
T ii
i
Missouri pH Indiana
c
FIG. 8. Ranges and medians of surface pH for the
Missouri and Indiana stripmine lakes.
3. Environmental patterns
a. pH. Although the original selection of the lakes and
their arrangement into a series of increasing pH was based on relatively
few samples, the series remained valid after 18 months of study. In
view of the wide and overlapping ranges of pH in some of the lakes
this was, perhaps, primarily due to good fortune. The most striking
differences between the annual cycles of pH in the six lakes (Figs. 2^-
29) were in the vertical stratification and in the magnitude of varia-
tion with time. Vertical stratification of pH persisted throughout the
period of study in the four meromictic lakes (III-VT). These vertical
pH differences were greatest during summer thermal stratification, at
which times Lakes I and II also exhibited pH stratification. Lakes
III, IV, and V had the greatest overall surface pH variation and Lakes
I and II, the greatest overall bottom variation. The magnitude of pH
variation with time, like that of vertical stratification, appeared to
be related more to meromixis than to relative placement on the pH scale.
None of the Missouri lakes studied by Campbell et al. (I965a) had a
reported overall range of pH variation as great as~~those of Indiana
Lakes I, III, IV, and V. Information regarding vertical stratification
and patterns of annual variation in the Missouri lakes is not available
(except for Lakes A3, B, and D when studied by Crawford and by Heaton,
at which times their other characteristics were different).
-------
b. Temperature. The most obvious difference between the
patterns of temperature variation in the study lakes (Figs. 30-35) was
that between Lake II, with its prolonged periods of homothermy and
complete circulation, and the other five more or less permanently
stratified lakes. Lake I, while thermally stratified throughout most
of 1970, was essentially homothermous in mid-August and late September.
This lake is probably best regarded as transitional between typical
holomictic lakes (e.g., Lake II and Control) and the meromictic lakes.
The overall temperature differences in the bottom strata (Table 11)
provide a good indication of these differences in thermal patterns.
Certain anomalous thermal patterns usually associated with meromixis
were observed in the study lakes. For example, dichothermous temperature
distributions (the minimum temperature occuring at intermediate depth;
see Hutchinson 1957) occurred in Lakes I and II in March 1970, in Lake
III from February through June 1970, in Lake IV from February to early
August 1970, and in Lake VI during March and April 1970. Mesothermy
(maximum temperature at intermediate depth) was observed in Lake I in
August 1970, Lake IV in early November 1969 and 1970, and Lake V in
April 1970. The complex temperature distribution in which one or more
maxima and one or more minima occur at intermediate depth is known as
poikilothermy and, according to Hutchinson (1957), is very rare.
Poikilothermous curves were observed in Lake III throughout November
and early December 1969j and in Lake V in late October of 1969 and 1970.
As previously noted, in the Missouri series only Lake A]_ was meromictic,
while the other lakes exhibited limited stratification or none. The
holomictic nature of Lakes Ao, B, and D is almost certainly due to their
relatively (as compared with the Indiana mine lakes) low total solute
concentrations (Table 9) • Missouri Lake P^y like Indiana Lake II, was
holomictic. In both cases the reasons for the failure of meromixis to
develop are unknown (see p. 55).
c. Dissolved oxygen. In the six lakes studied,both the level
of dissolved oxygen and the percentage saturation increased with in-
creasing pH (Fig. 9)- This relationship was modified by meromixis in
that, with the exception of Lake VT, the deep waters of the chemically
stratified lakes remained anaerobic throughout the period of study
(Figs. 36-1*4). Only Lakes II and VI had appreciable amounts of dissolved
oxygen in deep water during a significant part of the annual cycle. The
only lake that did not fit into the trend of increasing oxygen with
increase in pH was Lake V which had unusually high levels of dissolved
oxygen and also very high relative saturation. The fact that the peak
occurred in Lake V (rather than Lake VI) probably resulted from the
photosynthetic oxygen production by massive blooms of algae that
occurred in Lake V in response to the fertilization program mentioned
previously.
Thus the differences between the various patterns of dissolved oxygen
variation observed in these lakes can be attributed, at least in part,
to the different pH levels, to meromixis vs. holomixis, and to differences
52
-------
TABLE 11. Observed annual ranges of bottom temperatures
in the Pike County study lakes.
Lake Annual temperature Temperature difference
range (°C) (°C)
I
II
III
IV
V
VI
9.5 -
6.3 -
12.6 -
9-7 -
9.2 -
9.0 -
2^.3
28.5
1UA
11.6
1U.7
17.2
1U.8
22.2
1.8
1.9
5-5
8.2
in photosynthetic activity. pH probably acts indirectly, acidity
increasing the total solutes (because many substances are more soluble
at low pH) and leading, in turn, to increased oxygen demand by the
reduced solutes.
Oxygen variation in the Missouri lakes was not discussed by Campbell
et al. (I965a). The earlier studies of Crawford (19^2) and Heaton
Tl95l)j however, indicated reasonably high levels of dissolved oxygen
at all depths in all three lakes then under study (not including Lakes
A]_ and Ag, which were added to the series later). On rare occasions
during thermal stratification, anaerobic conditions existed briefly in
the deepest strata. Even during mid-summer the dissolved oxygen levels
in the Missouri acid Lake Ag (Crawford 19^2, Heaton 1951) did not
approach the very low values observed in Lakes I and II of this study.
Lake A3 had much lower solute concentrations than Missouri Lakes A-^ and
Ag or than all of the Indiana stripmine lakes, and therefore it probably
had a correspondingly much lower chemical oxygen demand.
d. Optical properties. Turbidity and transparency (Table
2) did not appear to be related directly to pH, but were influenced by
the stratification patterns. As discussed previously (p. 39), high
turbidity was frequently observed in a stratum of water corresponding
in depth to the lower limit of oxygen in the meromictic lakes (except VI).
The two lakes with oxygen in deep water during the greater part of the
annual cycle (Lakes II and VI) had strikingly lower turbidity, especially
at the bottom. The control lake had quite high turbidity on the two
days that it was measured. Transparency, as indicated by the 1 percent
level, was approximately inversely related to turbidity. Lakes II and
53
-------
15
10-
rt
0)
i
w
ra
s
300
200
100
I
M
1
0)
Lake I
II
III
IV
VI
FIG. 9. Maximum (A) and minimum (B^ open surface (no ice cover)
dissolved oxygen; and percent saturation (Cj at overall
maximums (any depth, ice present or absent) in Pike County
lakes.
VI were the most transparent, Lakes III and V least transparent. Lake
I had greater transparency (as indicated by the shallowest observed 1
percent level) than expected on the basis of turbidity.
The methods for turbidity (platinum wire) and for transparency (secchi
disc) used by Campbell et al. (1965a) were not strictly comparable to
those used in this study. Unfortunately, the platinum wire turbidity
method lacks accuracy at low values (below 7 ppm). Missouri Lake Ao was
similar to Indiana Lake II in pH. These two relatively transparent lakes
may both be examples of Parsons' (I961f) Type III: Blue Lakes (pH 3.0-^.0,
no turbidity, no thermal stratification, iron less than 30 mg/1).
Parsons' classification scheme for stripmine lakes is compared with the
Indiana and Missouri lakes in a later section. Missouri Lakes Aj_, &*>,
B, and D had turbidity levels that were generally in the middle range
of the Indiana lakes. None had the very high upper limits observed in
the deep waters of Lakes I, III, and V. Relative turbidity and
-------
transparency within the Missouri series (Fig. 10) had an interesting
relationship to pH. Turbidity was lowest in Lake A3, the middle lake
in pH, and increased with both increasing and decreasing pH. A similar
relationship was present in the Indiana series (Fig. 11), turbidity
increasing both above and below Lake II on the pH scale. In the
latter series, however, this relationship was obscured by meromixis
and the attendant precipitates. It seems reasonable that stripmine lake.'.'
of about pH S.OA.O may be lowest in turbidity because below about pH
3 the solubility of iron increases abruptly (leading to turbidity from
precipitates), and above that pH k organic turbidity of various sorts
(plankton, increased decomposition, etc.) increases.
e. Dissolved substances. According to Hutchinson (1957» P.
552) "Normal fresh waters are dilute solutions of alkali and alkaline
earth bicarbonate and carbonate, sulfate, and chloride, with a variable
quantity of largely undissociated silicic acid ... which is often
present in excess of sulfate and chloride." In addition, there may be
a number of other ions (some quite significant biologically) and certain
organic and inorganic colloids. The stripmine lakes of this study cannot,
of course, be considered "normal" fresh waters. In most natural fresh-
water lakes the major cations are Ca++, Mg"*"1", Na+, and K1" and the major
anions are HCO^", 8014.', and Co^"" in order of decreasing concentration.
The proportions are much less constant than is the case in oceanic sea-
water.
Most acid
30H
Least acid
20
10'
B
O
01
•H
O
a)
CO
2 &
FIG. 10. Maximum turbidity (£ and transparency ££ in the Missouri
lakes.
55
-------
Most acid
Least acid
OJ
15
o
•rl
CQ
-P
•H
10
6
0)
s
w
* a
Pi
03
II
III
rv
v
VI
FIG. 11. Maximum surface turbidity (A) and transparency
(B) in the Pike County lakes.
Perhaps the most straightforward measure of ionization in a liquid is
electrical conductivity adjusted to a standard temperature, generally
25 C in North America. Results are reported in reciprocal meghoms or
micromhos. Since most of the substances dissolved in water ionize to
some degree, the values for conductivity vary with those for dissolved
solids (Tables 2 and 10). Overall variation in conductivity or specific
conductance (K2,_ • 10°) for the Indiana lakes is shown in Figs. U2-U6.
The range of conductivity values reported for freshwater lakes is about
9-^-00 micromhos (Hutchinson 1957? Reid 1961), but some saline lakes may
exceed 60,000 micromhos, and certain industrial effluents are much higher.
The range reported for the Missouri lakes (Table 9) was 110-13,600 |o,mhos,
probably approaching the extreme to be found in coal stripmine lakes.
The overall variation in conductivity in the lakes of this study was
much less than that of the Missouri lakes (Fig. 12). The curves for the
Indiana and Missouri lakes corresponded in conductivity only in the region
of pH 3. The very acid Lake A^_ had remarkably high conductivity, which
may have been due, at least in part, to the extensive areas of coal and
shale fragments in its basin. In both series, the surface conductivity
values tended to stabilize at pH values higher than 3.5 although at
different levels. These might be controlled by geological differences
(i.e., differences in the composition of the coal or associated materials)
or in part by differences in age. The latter seems unlikely, however,
since the pH and conductivity were not well correlated with age in
either series. For example, at the time of the study (Campbell &t al.
56
-------
1965a) Lakes A^_ and A2 were about 30 years old, and Lakes An, B, and
D were about U5 years old. Yet Lakes A± and Ap with similar pE differed
greatly in conductivity, while A? was much more acid than B and D but
relatively similar in conductivity. Indiana Lakes I and V are,
literally, but a stone's throw apart and were formed by the same mining
operation in 19^0, yet were markedly different in pH and to a lesser
degree in conductivity. Thus in both the Indiana and Missouri series,
there is a progressive but irregular decrease in dissolved substances
with increasing pH.
The overall difference in total solutes between the surface and bottom
strata of meromictic Lakes III-VT is apparent in Figs. kk-k7. The
difference was especially great in Lake III, accounting for its
relatively great stability of meromictic stratification. Holomictic
Lake II also had slightly higher maximum conductivity at the bottom,
but Lake I had identical maximum conductivities for the surface and bottom
waters.
f. Major ions. Unfortunately, there is no comprehensive
and universally accepted system of classifying lakes in terms of their
chemical nature. Of the various classificational or organizational
schemes proposed (reviewed by Hutchinson 1957)» perhaps the most useful
is the system of ionic diagrams proposed by Maucha (1932). This system
would be too complex in the present context, and hence I prefer to
discuss each of the major ions briefly. The data are summarized in
Table 3 and presented in detail in Appendix 2.
Ca++. In the lakes of the present study calcium varied from about 120
to 500 mg/1. The highest concentrations were found in Lake I and the
deep waters of Lakes III, IV, and V, and the lowest in the surface
waters of Lakes III and VI. Calcium in the Missouri lakes (Table 9) was
generally lower, but Lake A, had a maximum of ^55 mg/1. The range in
natural surface waters is from 0.13 or less to about 10,000 mg/1 (Dead
Sea), but most freshwater lakes contain 2.5-60 mg/1 (Hutchinson 1957).
The control lake, with an observed range of 11-21 mg/1, was well within
these limits.
Mg++. The overall range of magnesium concentration observed was from
B5~mg/l in the surface waters of Lake III to 537 mg/1 in the deep waters
of that lake. This range was much higher than that for natural fresh-
water lakes (0.14-9.9 mg/1), but much lower than typical values for
natural saline lakes (e.g., 5,600 mg/1 in Great Salt Lake and 38,000
mg/1 in the Dead Sea) according to Hutchinson (1957) • Magnesium concen-
trations in the Missouri lakes (Table 9) were well below those of the
Indiana lakes of similar pH. The control lake although having a maximum
observed value of 10 mg/1 in deep water during summer stratification
may reasonably be regarded as within the range for natural freshwater
lakes.
57
-------
1^,000 '
12,000 '
10,000 "
8,000 '
6,000
4,000 .
2,000 '
2,000
B
2 A-
B D
I II
8 pH
FIG. 12. Maximum conductivity in relation to median surface
pll in the Missouri (Ay"and Pike County (Bj lakes.
58
-------
Dissolved iron. In this section I have dealt with the total dissolved
iron without attempting to sort out the various ionic forms (Fe++, Fe+++,
FeOH+ , H2FeC>3~, etc.) that may be present. Hutchinson (1957) gives
the iron concentration for the upper circulating waters of lakes as
ranging from undetectable to 19.2 mg/1. The upper value is from a
volcanic lake with a high content of sulfuric acid and a pH of about
1.8. A much higher value of 50 mg/1 was reported from a bog of pH
1*.0 near Moscow, USSR. Iron in the surface waters of the study lakes
(Table 3) had the familiar inverse relationship to pH. The total
range was from <0.05-0.55 mg/1 in Lake VI to 60-105 mg/1 in Lake I.
Missouri acid Lakes A-j_ and A2 markedly exceeded Indiana lakes which
had much higher iron concentrations, especially in deep water. The
control lake exceeded Lake VI in iron, especially in deep water during
stratification.
Ferrous iron accumulates in the deep anaerobic waters of lakes during
stratification. Under such conditions, ferrous and manganous bicarbonates
in the deepest strata of the monimolimnion of a meromictic lake or the
hypolimnion of a thermally stratified lake can have a buffering action,
resulting in an acid heterograde curve of pH distribution with a
minimum value in the upper part of the anaerobic zone (Hutchinson 1957).
During summer stratification, meromictic lakes have four major layers
including the usual three (epilimnion, metalimnion, hypolimnion) and
themDnimolimnion. At such times, a sort of double acid heterograde
curve of pH distribution was observed in Lakes III, IV, and V (Fig. 13)
but not in Lake VI, which had oxygen in deep water during most of 1970.
Manganese. Like iron, manganese may exist in either a reduced or
oxidized form or both, depending on pH and redox potential. The total
manganese in the study lakes was clearly inversely proportional to pH.
The lowest values (<0.05-26.0 mg/l) occurred in Lake VI and the highest
(bO.O-6k.Q mg/1) in Lake I. Moderately high values also occurred in
deep water during stratification. The control lake was well within
the range of <0.005-15.0 mg/1 found in natural freshwater lakes (Hutch-
inson 1957). Values of manganese reported for the Missouri lakes were
generally comparable to those of the Indiana lakes of similar pH.
Aluminum. According to Hutchinson (1957), aluminum in natural lakes
ranges from undetectable to about 97 mg/1 (in a shallow saline lake in
a closed basin). In the present study a range of from 0.05 to Mf.O
mg/1 was found. The maximum (from Lake I) was rather low for an acid
mine lake. Missouri Lakes A]_ and &% were both much big*161"' Aluminum,
too, is much more soluble in acid waters (and also above pH 8).
Meromixis had little, if any, influence on aluminum concentration in
the monimolimnion as compared with the mixolimnion. In the control
lake the concentration in deep waters during summer stratification was
fifty times that in the surface waters (Appendix 2).
Na+. In naturally occurring salt lakes sodium may reach very high
values (e.g., 67,500 mg/1 in Great Salt Lake, Utah). In most natural
lakes, however, typical values range between 0.13 and 16.6 mg/1
59
-------
M
27-vii-70
10 J
FIG. 13. Double heterograde pH curves associated with maximum summer
stratification in meromictic Lakes III, IV, and V.
(Hutchinson 1957). The range in this study was from 11 mg/1 in the
surface waters of Lakes I and V to 75 mg/1 in the deep waters of Lake
VI. Thus the lakes of this study were moderately higher than "typical"-
freshwater lakes, but did not approach the natural saline lakes in
sodium concentration. Concentrations were only slightly higher in the
monimolimnion of the meromictic lakes than in the surface waters, and
showed little or no systematic variation with pH. The control lake was
moderate, and the Missouri lakes were generally somewhat lower in sodium
than the Indiana lakes of similar pH.
K+. The surface waters of the study lakes were all quite similar in
potassium concentration, varying from 2.5 to 5.5 mg/1. This is generally
within the range to be found in natural freshwater lakes (Hutchinson
1957). Saline lakes may contain much higher amounts. In the deep waters
of Lakes III and IV potassium reached 25 mg/1 during intense summer
thermal stratification. This, together with a similar but less dramatic
increase in magnesium, suggests a possible biogenic element in the
meromixis of these lakes. The control lake had lower potassium than any
mine lake, and the Missouri lakes were generally lower than the Indiana
series.
Zinc. According to Hutchinson (1957), the occurrence of zinc in natural
waters has received little attention. Reported values range from 0.0013
to 0.65 mg/1, the latter reputedly toxic to Cladocera. In the lakes of
this study zinc concentration was inversely related to pH, varying from
60
-------
0.1 mg/1 or less in Lake VI to as high as k.O mg/1 in Lake I. Die
latter concentration is probably toxic to many aquatic organisms.
There was no consistent difference between surface and bottom waters.
Missouri Lake A^ had as much as 100 mg/1, but the other lakes in that
series were similar to the Indiana lakes. The control lake had
moderate zinc concentrations.
Sulfate is by far the dominant anion in coal stripmine lakes.
in natural freshwater lakes the range encountered is about 6.2-20.5
mg/1, but values in excess of 60,000 mg/1 have been reported
(Hutchinson 1957) . The overall range found in this study was from
510 mg/1 (surface of Lake III) to 3650 (bottom of Lake I) . Values in
the surface waters were only moderately influenced by pH, and
significantly higher values were encountered in the monimolimnion of
the meromictic lakes. The control lake had a very much lower concen-
tration than any mine lake, but well within the upper part of the range
for natural freshwater lakes. The maximum value for Missouri Lake A-,
was quite high (?600 mg/1), but the other lakes in that series had
lower values than the Indiana lakes of similar pH.
Cl". Chloride in the study lakes ranged from less than 1.0 to about
40.0 mg/1. In general Lakes III and IV had the highest values, Lakes
II, V, and VT the lowest. These levels of chloride were somewhat high,
but were well within the range for natural lakes (Hutchinson 1957) • The
control lake had relatively low chloride. Chloride was not reported for
the Missouri lakes by Campbell et al. (1965a) .
HCO^" . Bicarbonate occurred in Lakes III through VI in increasing
amounts. The overall range was from 2k to 296 mg/1 (in the surface
waters of Lake III and bottom waters of Lake VI, respectively). This
was well within the range for natural freshwater lakes. The very low
ratio of bicarbonate to sulfate (from .03 in Lake III to .21 in Lake
VT) is probably met with, in nature, only in such uncommon situations as
volcanic lakes and peat bogs.
Other ions (nitrate, phosphate, and the forms of silica) of biological
significance were also measured. Silica varied inversely with pH,
but nitrate and phosphate were rather uniformly low. One doubts that
low levels of these substances are ever limiting factors in the mine
lakes because of other overriding chemical conditions.
Thus in the stripmine lakes of this study, the ionic situation is
complex. In typical freshwater lakes the dominant cations are considered
to be Ca++, Mg++, Na+, and K*" in that order. In the acid mine lakes,
iron, manganese, and aluminum tend to exceed sodium and potassium.
Iron and manganese are also high in the monimolimnion of the meromictic
lakes. In the surface waters of the less acid or slightly alkaline
mine lakes, the usual situation (sodium and potassium in third and
fourth rank) prevails. In "normal" lakes the dominant anions are HC03~,
S0j,=, ci~, and C0o= in decreasing order of abundance. The same ions,
but in different order of abundance, occur in the mine lakes, with
carbonate excluded at low pH values (less than 8.4).
61
-------
In summary, certain trends in the ionic composition of the study lakes
may be noted. pH influenced the concentration of several ions. Calcium,
magnesium, iron, manganese, aluminum, zinc, sulfate, and chloride varied
inversely with pH. Sodium and bicarbonate were directly related to pH.
Ions that were markedly higher in the monimolimnion of meromictic lakes
(probably due primarily to the redox situation) included calcium,
magnesium, iron, manganese, sodium, sulfate, and bicarbonate. Some ions
including calcium, magnesium, iron, manganese, sodium, zinc, and sulfate,
were much higher than in natural freshwater lakes.
The magnitude of seasonal variation in concentrations for most of the
ions measured was relatively small. The most prominent kind of trend
found was an increase in concentration—especially in deep water—during
maximum summer stratification. Such a pattern was observed for calcium,
magnesium, iron, sodium, potassium, manganese, silica, and sulfate in at
least some of the lakes, although not in Lakes I and II. No other marked
seasonal trends were observed.
The control lake had consistently lower values for nearly all ions than
any of the mine lakes. This lake fell well within the range of water
chemistry for natural freshwater lakes and it is probably representative
of "natural" (i.e., nonstripmine) ponds and small lakes of the region.
The most striking differences between the Missouri lakes and those of
the present study were (l) the much higher concentrations of several ions
in Lake AI than in any Indiana lake and (2) the relatively low total
solute concentrations in. Missouri Lakes A3, B, and D. These differences
may be related to the origin of Lake AI as a coal-washing basin and to
regional differences in geology and weather.
g. Lake types. Parsons (196*0 has suggested a classifica-
tion of stripmine lakes on the basis of certain physical and chemical
characteristics. The successive types in this series are thought to
represent sequential stages in the recovery of a mine lake from acid
pollution. The criteria upon which these types are based have been
arranged as Table 12. In this series acidity and iron decrease,
alkalinity increases, and turbidity and thermal stratification increase
initially and later decrease.
The lakes of this study do not, for the most part, fit very well into
this classification. On the basis of pH Lake I would be a Type II lake,
but it had greater iron, turbidity, and thermal stratification, and did
not show the seasonal variation suggested for that type. Lake II, as
previously noted, meets the specifications for Type III except that it
did have measurable turbidity and a limited degree of thermal stratifi-
cation. The remaining four lakes would all be Type IV on the basis of
pH, but all had prolonged stratification, high upper limits of turbidity,
and (except Lake VI) moderate to high dissolved substances including
iron.
62
-------
The Missouri lakes fit the scheme better than the Indiana series,
probably because Parsons' lake types were formulated mainly on the
basis of data from the same region. The striking differences between
the combinations of conditions found in the Indiana lakes of this study
and those of the Missouri lakes may, as noted below, reflect basic
differences in the geology and climate of the two areas.
Campbell et al. (I965a) arranged the five Missouri lakes into three
successions! stages: youthful (A^ , early recovery (A^, Ao), and late
recovery (B, D), which they characterized in terms of acidity, solutes,
transparency, thermal stratification, and biological diversity. The
same general problem is encountered when trying to fit the Indiana
lakes into this scheme as with Parsons' types (i.e., the sets of
conditions present in the individual lakes are not the same) . Thus
a given Indiana lake might rank as youthful in terms of stratification,
early recovery as to solutes and turbidity, and late recovery as to pH
and color (as in fact Lake VI does). Lake II was once again the only
Indiana lake that matched one of the categories (early recovery)
reasonably well.
TABLE 12. Coal stripmine lake types proposed by Parsons (196*0 .
Type I: Type II: Type III: Type IV:
Red lakes Transitional Blue lakes Grey lakes
PH
Titratable
acid
(mg/1 IL^SO^)
Turbidity
(ppm)
1.2-2,5
5500
AH seasons,
due to iron
2.5-3.5
3500
Spring only,
due to iron
• •* • a _ _
3.0-^.0
^ 3000
None
* IKO
Low
Low to
normal;
hydroxides hydroxides
Iron
High (£ 65)
Thermal
stratification
Alkalinity
(mg/1 as
CaC03)
Present in
summer
None
High (130)
in spring,
low (15)
other seasons
None
None
Low
(< 30)
None
None
organic and
inorganic
Low
Little to
normal
63
-------
No doubt a valid series of limnological stages through which all coal
stripmine lakes pass during recovery from acid pollution could be formu-
lated. To have general application, such a scheme would have to be based
on data from lakes in the various regions in which coal stripmining is
carried out and would probably have to be based on more (perhaps
different) criteria than those considered above. The formulation of such
a general successions! scheme for stripmine lakes is not an objective of
this paper.
k. Biological patterns
a. Diversity. There was clearly an increase in overall
fauna! diversity with increasing pH in the lakes of this study (Table
U). The rate of increase, however, was not uniform either for the total
fauna or for the ecological subgroups (Fig. 14). The overall rate of
increase was greater at low pH (Lakes I-IV) than at higher pH (Lakes
IV-VI). This was generally true of both zooplankton and benthos, and
vertebrates were most diverse in Lake IV. The increase in benthic taxa
between Lakes I and II was as great as the total additional increase
from Lake II to Lake VI. This may reflect the influence of meromixis
in Lakes III-VI on the conditions of life for the bottom fauna.
Similarly, the greatest increase in number of zooplankton taxa occurred
between Lakes II and III (nine taxa added of the total increase of 16
from Lake I to Lake VI).
It is of some interest to consider the relative importance of addition
and substitution in the fauna! changes that occurred through the lake
series (Table 13). The number of taxa carried over from one lake to
the next at each step was typically about 75 percent, but only about
half (56 percent) were carried over from Lake II to Lake III and nearly
all (9k percent) from Lake III to Lake IV. The increase in fauna!
diversity may reasonably be regarded as primarily a result of addition
of taxa. The total change from Lake I to Lake VI was from 12 to 53
taxa or an overall increase of Ul (3^2 percent). Eight of the 12 taxa
found in Lake I were also present in Lake VI. These groups must be
regarded as quite eurytopic with regard to pH, solutes, etc.
The Missouri lakes (Campbell et al. 1965a) had more animal taxa at
high pH and fewer at low pH than the Indiana lakes (Fig. 15). This may
be related to the relatively lower solutes in the high pH Missouri lakes.
Another contributing factor is the longer period of study (almost 30
years) and greater number of investigators of the Missouri lakes. This
does not, of course, explain the greater number of taxa found in the
Indiana lakes of low pH. It has been noted previously that Missouri
Lake As> and Indiana Lake II are environmentally quite similar, and it
is therefore surprising that Lake II had nearly four times as many
animal taxa (23 vs. 6) as Lake A£.
b. Standing crop. Two general features of the overall mean
biomass (Table 14) are of interest. First, the rank order of the lakes
was different for each of the three fauna! categories, no single lake
-------
50'
UO-
30
-p
o
-------
TABLE 13. Faunal changes in the Pike County lakes.
Lake Taxa maintained
interval No. %
I-II
II-III
III-IV
rv-v
V-VI
I-VI
TABLE I
9 75
13 56
30 9k
35 71
37 76
8 67
Taxa added
No. %
Ik 117
19 83
19 59
Ik 29
16 33
U5 375
k. Overall mean animal standing croj
Zooplankton
Lake Number/up
V
II
VI
IV
I
III
2,162,372
531,810
199,602
28,377
8,38U
7,76l
Benthos
Lake mg/m^
II 1,1,16
vi 763
I 573
V 220
iv 95
III 66
Taxa lost
No. $
3 25
10 kk
2 6
1^ 29
12 2k
k 33
) in the Pike
Net change
No. %
+11 +92
-1-9+39
+17 + 53
0
+ k + 8
+Ul +3^2
County lakes.
Fish
Lake g/trap-hour
VI
IV
III
V
7.78
2.70
1.61
0.95
as "fruit." On this basis, it appears that there was a trend toward
increasing biomass with increasing pH. This was not so straightforward,
however, as was the case with diversity. Lake II was much higher and
Lakes III and IV lower than would have been expected on the basis of pH
alone. The differences between lakes in the patterns of circulation are
probably responsible, at least in part, for this lack of agreement
between pH and biomass.
66
-------
8
6
pH
Indiana
Missouri
20
Number of known taxa
100
FIG. 15. Animal diversity and p_H in the Missouri
and Indiana lake series.
It is possible to construct a simple index of environmental stress by
assigning arbitrary values to the different levels of stress of the
various environmental factors. This has been done for the study lakes
using pH, dissolved oxygen, total solutes (as indicated by conduc-
tivity), and intensity of stratification (as indicated by surf ace-to-
bottom temperature differences). For each factor, the lakes were
simply ranked in order of increasing stress and assigned numbers from
one to six (Table l6). The total for each lake constitutes its index
of environmental stress. On this basis, Lake III was the most
successful and Lake VI the least so. The index of stress was inversely
67
-------
TABLE 15. Percent of total standing crop ±n the Pike County lakes.
Zooplankton
Benthos
Fish
Total
I II
0.3 18.1
18.3 U5.2
0 0
18.6 63.3
III
0.3
2.1
12.3
lU.7
Lake
IV
1.0
3.0
20.7
2^.7
V
73.6
7.0
7.3
87.9
VI
6.7
2k. U
59.6
90.7
TABLE 16. Index of environmental stress in Pike County lakes.
Lake
Factor I II III IV V VI
pH
Temperature
difference
Maximum
conductivity
Dissolved
oxygen
Index of
stress
6
2
5
6
19
5
1
2
5
13
k
6
6
k
20
3
5
U
3
15
2
k
3
1
10
1
3
1
2
7
related to overall standing crop (Fig. 16), indicating that a combination
of factors influenced animal biomass rather than mainly pH as seemed
true for diversity.
The influence of meromixis can be assessed by considering the distribu-
tion of biomass with depth (Tables 17 and 18). These data, however,
reflect the influence of relative area and volume. It is, therefore,
68
-------
1001
II
IV
III
Stress
10
15
20
FIG. 16. Biomass and environmental stress in the Pike County lakes.
more appropriate to use the original non-weighted values which have
been plotted as percentages of the total for each lake in Figures 17
and 18.
Clearly, the depth distribution of zooplankton was less influenced by
differences in stratification patterns than that of benthos. With
the single exception of Lake I, the zooplankton was distributed
remarkably uniformly throughout the water column. In Lake I the zoo-
plankton was concentrated near the bottom for unknown reasons.
Benthic biomass, on the contrary, had three distinct patterns of dis-
tribution with depth. The biomass tended to be greatest in deep water
in Lake II, in the shallow regions of Lakes I, III, V, and VI, and at
the surface and intermediate depths in Lake IV. More than half the
total benthic biomass in Lake II was concentrated in the two deepest
1-m depth intervals, while Lakes I and VI had relatively little, and
Lakes III, IV, and V no benthos in their deepest areas. The benthic
maximum between 5 and 6 m in Lake IV may have been partially due to
the presence of a moderate shelf between 5 and 8 m. However, it is
probably correct to view the lower limit of dissolved oxygen during
maximum summer stratification as very important in restricting benthos
to the shallower parts of these lakes. Thus it is not surprising to
find virtually no benthic production in the deepest areas of Lakes III,
IV, and V. Less readily accounted for is the low biomass below h m
in Lake VI where dissolved oxygen was always adequate to 6 m and
usually to 7 m.
69
-------
TABLE 1?. Depth distribution of mean percent area-
weighted total benthos biomass
Depth (m)
0-1
1-2
2-3
O_ii
ll C
5-6
6-7
7-8
8-9
9-10.5
I II III
^05 lUo 23
130 117 29
38 160 6
127 b
663 3
209 1
0
0
Lake
IV
27
6
3
3
3
51
2
0
0
0
V VI
162 328
52 109
5 116
0 101
1 22
0 ^9
38
Total 573 l,Ul6 66 95 220 763
The lower boundary of dissolved oxygen was also apparently important in
the distribution of the planktonic ciliate Euplotes (Fig. 19). Although
most of the zooplankton organisms were distributed more or less uniformly
as was the total, Euplotes tended to be concentrated at, or just below,
the lower limit of oxygen where it presumably fed on the bacteria
associated with that chemical boundary zone.
The mean biomass data plotted on a monthly basis (Figs. 20 and 21) pro-
vide a rough outline of the patterns of seasonal change. These data
are admittedly limited, especially for Lakes I and VI. The benthos
data for Lake I are inadequate for this purpose and have been omitted.
Zooplankton abundance (Fig. 20) was generally greatest in autumn and in
spring or early summer, least in midwinter and midsummer. There were,
however, exceptions such as the midsummer peaks of abundance in Lakes
III and V. The zooplankton biomass in Lake V fluctuated greatly during
70
-------
TABLE 18. Depth distribution of mean percent volume-
weighted total zooplankton biomass (ind./nr3).
Lake
Depth (m)
0 -0.5
0.5-1.5
1.5-2.5
2.5-3.5
3.5-^.5
^.5-5.5
5.5-6.5
6.5-7.5
7.5-8.5
8.5-9-5
9.5-10.5
Total
I II
1,31*0 56,1*50
l,66o 85,050
1*,580 81*,700
800 108,280
9!*, 060
70,070
33,200
8,380 531,810
III
1*1*5
730
1,060
975
1,705
1,550
650
1*25
220
7,760
IV
590
1,51*0
1,210
2,600
3,890
l*,830
3,770
5,080
3,070
1,290
510
28,380
V VI
nU, 290 8,680
866,950 Ui, U8o
599,^00 50,320
1*1*7,120 31,^90
98,250 25,860
36,360 22,390
ll*,5l*0
i*,8i*o
2,162,370 199,600
summer of 1970, perhaps in response to changing phytoplankton abundance
resulting from the artificial fertilization program. No other lake had
such pronounced short-term oscillations in biomass.
Benthic biomass in all of the lakes increased through the autumn to a
peak in November-December, then decreased abruptly to a low level in
March (Fig. 21). In Lakes II, III, and V the benthos remained low
during the summer, but in Lakes IV and VI a secondary peak occurred in
May-June. The rapid decrease in the benthic biomass in spring was
associated with the emergence of insects whose aquatic larvae consti-
tuted the bulk of the benthos in all of the lakes except Lake VI (where
gastropods made up over 50 percent).
The meromictic lakes did not have any characteristic pattern of
seasonal changes in biomass distinct from that of the nonmeromictic
lakes. The seasonal patterns do not appear to differ from those
71
-------
0
M
1
2 "
o ^_
I
o
M 1 —
1 -
2 •
3 '
k -
;
—
II
0
Mi _^_^__
i -
2 -
3 -
h •
5 ]
6 4
T? J
VI
—
^_
1
^^^^~""1 —
r
0
M
1 -
I
r
2 H
3 -
k •
5 -
"~f
6 •
7 •
8 -
9 -
10 ,
I l_
1
r
]
o -
M T .
1 \
- ,
1 2 J
IV rJ -
' '
\
r
i
1 3 -
1 i
-
L
t
i
5 -
6 -
7 -
8 J
w J
III I
1
r
1
. f
0
M^ — i— ^^
1 ~ • -,
1 -
1
r 2
«_ J 3 -
k -
^T
5 -
V I
1
J
S
1
FIG. 17. Depth distribution of -unweighted total mean
zooplankton standing crop (indivlduals/m^).
72
-------
M
1
o
^
0
M
1.
2.
3 •
h-
5 •
6
7 •
8 ,
0
M
1
^
3 '
*•
5 •
I
r
V
1
J
I
i:
r
f
E
J
0
M
1.
•
2 •
3 •
5 •
6
7 .
T<
n
\
M
-
2 •
3 •
h
5-
7 -
8 -
9 •
10 •
IV
J —
_J
J
0
M
II
1
- I 1
I
4 .
5 .
-1
FIG. 18. Depth distribution of unweighted total
mean benthic standing crop
73
-------
0
M •
1 -
2 '
3 -
k •
5 '
0
M
1.
2 •
3 '
h
5
£ .
r
V
i
TT
1
0
M
1 -
2 '
3 '
k '
5
7
n
J
1
/I
0
M •
-
•
3 •
k •
5 '
6 -
7 '
8 •
9 •
10 •
IV
M
1
2
3
k
5
6
7
8
.
•
•
•
•
III
1
\
_T
i
n
FIG. 19. Depth distribution of •unweighted mean Euplotes
standing crop (individuals/m-*).
-------
100
1,000,000
M A M J J
1,000,000 -
100,000 -
10,000
1,000
1,000,000 -
100,000 -
10,000
IV
1,000,000
100,000 •
10,000
VI
S ONDJFMAMJJASOND
FIG. 20. Seasonal cycles of total zooplankton standing crop
(indivlduals/m^), September 1969 to 'December 1970.
75
-------
I*, 000
3,000
2,000
1,000
1,500
S OND JFMAMJJAS 0 ND
1,000 '
500
2,000 i
1,500
1,000
500 '
VI
S ONDJ FMAMJJAS ON D
FIG. 21. Seasonal cycles of total benthos standing
September 1969 to December 1970.
(mg/m2)
76
-------
commonly observed in other kinds of lakes and ponds. Unfortunately,
no data are available regarding the relative biomass of the Missouri
lakes.
C. Patterns of ecosystem organization
!• The ecosystem concept. The term "ecosystem" was coined by
Tansley (1935) to designate a group of interacting organisms together
with their nonliving environment. The term embodies the idea that the
organisms and their surroundings constitute a functional as well as a
structural unit. This idea was eloquently stated and developed by
Forbes (188?) in his remarkable essay on the interconnectedness of life
within the microcosm of a lake.
When used without qualification, "ecosystem" designates any organism-
environment complex from a very small and impermanent unit, such as a
fallen log in a forest or a rain puddle on the prairie, to the entire
biosphere. In ordinary use, however, only those systems that have at
least moderate extension in space and time are intended. Hutchinson
(1967) has addressed himself to this problem and for him an ecosystem
is "... the entire contents of a biotope." A biotope in turn is
"... any segment of the biosphere with convenient upper and lower
boundaries, which is homogeneously diverse relative to the larger
motile organisms within it ...." The phrase "homogeneously diverse"
means that "... the relevant mosaic elements of the environment (sand
grains, chemical gradients, etc.) "are small compared with the normal
ranges of individuals of species S." If the mosaic elements are large
compared to these ranges, the area is "heterogeneously diverse." Thus
"The littoral of a lake will be heterogeneously diverse in relation to
a natommatid rotifer, homogeneously diverse in relation to a sun fish"
(Hutchinson 196?, p. 229).
As I conceive and use the term, an ecosystem ought to have enough size
and enough organizational stability (feedback mechanisms, alternate
energy and material pathways, etc.) to ensure its integrity over
periods of time measured in yeans (rather than in weeks or months) in
the absence of external catastrophe. Obviously, some ecosystems (e.g.,
temporary ponds) have seasonal cycles of activity.
No ecosystem on earth stands entirely alone. All life on the planet
depends ultimately for energy on solar radiation. The earth's larger
ecosystems such as the sea, the North African desert, the Taiga, and
the Tundra (sometimes distinguished by the term "Biomes") are all
linked by biogeochemical cycles, by the common atmosphere, water supply,
and store of chemical elements. In general the smaller an ecosystem
is, the greater will be the influence of neighboring systems upon it.
Small aquatic ecosystems, like oceanic islands are convenient to study
because it is relatively easy to define their spatial limits. The
small lakes of the present study were chosen partly for that reason.
They were also small enough to treat in a reasonably thorough manner,
77
-------
yet large enough to satisfy the criteria mentioned above. As Forbes •$..'
wrote (1887), "Nowhere can one see more clearly illustrated than in a ^
lake what may be called the sensibility of such an organic complex (i.e.,
ecosystem), expressed by the fact that whatever effects any species
belonging to it, must have its influence of some sort upon the whole
assemblage."
2. Some hypotheses of ecological succession. This enquiry began
with the idea that a series of coal stripmine lakes at different stages
of recovery from acid pollution could be regarded as a series of stages
in the primary succession of a single such lake. It was further suggested
that such a set of lakes could be used as an analogue to test hypotheses
about ecological succession in general. Odum (1963) suggested that
ecological succession has the following attributes: (1) It is an orderly
process of directional, and therfore predictable, community changes; (2)
It results from modification of the environment by the community; (3) It
culminates in the establishment of the maximum possible ecosystem •*';.
stability. Odum emphasized that the biological community (the "bioco- :!.-"
enosis" of Hutchinson 1967) controls ecological succession and that the |u-
physical environment "determines the pattern of succession but does not
cause it" (Odum 1963, p. 78).
Certain kinds of changes are regarded as generally characteristic of the
process of ecological succession (Odum 1959» 1963? Hutchinson 1959»
Margalef 1963). These are here set forth in their simplest form: (1) The
kinds of organisms present change continuously throughout succession; (2)
The number of taxa present increases initially, then stabilizes or
declines; (3) Total biomass increases; (U) Net production decreases as
a result of an increase in community respiration; and (5) Complexity Of
organization increases or, in Margalef's terms, total information in-
creases. All these changes are in the direction of increased homeostasis.
This pattern occurs only in the context of a relatively stable environ-
ment. If the environment is unstable, an ecosystem will be selected
that is composed of species with high reproductive potentials and broad *
environmental tolerances (Margalef 1963). 1:
./
Before considering the data in this context, one must recall that at
least three factors besides pH almost certainly have a strong influence
on the stripmine lake ecosystems. The .first is morphometry. Deevey
(19^1) failed to find a good correlation between mean depth and bottom
fauna in small lakes in New England. On the other hand, Rawson (1955)
and Rounsefell (196*0 have demonstrated a correlation between relative
extent of the shallow littoral zone and productivity of bottom fauna and
fish. Several authors including Burner and Leist (1953)> Arata (1959)
and KLimstra (1959) have suggested that the very restricted area of the
shallow shore zone in most stripmine lakes may be an important factor
limiting production of benthic organisms and organic production in
general. Prospective spawning sites are, also, thus limited. Verts
(1956) observed that muskrats living in stripmine ponds were forced to
burrow into the banks, because the restricted littoral areas were too
small to permit house building.
78
-------
The second factor, meromixis, is related to morphometry in two ways.
The narrow deep basin shape undoubtedly predisposes these lakes to
incomplete turnover. In addition, such narrow deep lakes necessarily
have a relatively small proportion of their volume in the upper
circulating mixolimnion.
The third factor is the relatively low habitat diversity present in the
mine lakes. Substrates present are limited almost entirely to clay,
fallen leaves, and in some cases tree branches. In the two lakes where
they occur in significant volume (IV and VI), rooted aquatic plants
provide a substrate for some organisms.
All three factors probably depress both production and diversity in the
•Study lakes. However, the first affects all the study lakes except I,
and the third affects all six. Only meromixis has a pronounced
differential occurrence—affecting primarily Lakes III, IV, and V (since
the monimolimnion of Lake VI is limited to the 6-7 m stratum and dissolved
oxygen is usually available at all depths).
The rate of ecological succession is highly variable, depending on
regional climate and geomorphology, kind of ecosystem, stage of the
evolutionary process, and many other factors. Generally change is slow
at first, then accelerated and finally slowed again in the late stages.
>.
3. Some evidence; system changes in the Indiana lakes. If the
six stripmine lakes do in fact represent a series of stages in the
successional development of an ecosystem, then some or all of the trends
listed above should be present in the series. In this section the data
are considered in relation to these hypothetical patterns of directional
change.
(l) The composition of the biocoenosis changes continu-
ously throughout succession. This was certainly true of the fauna
(Table 13). Considerable change occurred at each step in the series
including that from Lake IV to Lake V at which 1^ taxa were replaced with
no net change in number of taxa. Generally, the overall change in
composition of the fauna was greater in the earlier stages of the series.
(2) The number of taxa increases, especially in the early
§ages of succession. This, too, was generally true for the fauna. The
ta (Tables h and 13, Fig. I1*) show a progressive increase in number of
imal taxa except in Lake V. The rate of increase was greater in the
early stages (i.e., acid lakes) than in the later ones. Hall et al.
(1970) reported that predation by bluegills (Lepomis macrochirus) in a
series of artificial ponds increased diversity of prey species (zoo-
plankton and benthos), presumably by reducing competition between the
prey species. This kind of effect might account for the relatively great
increase in diversity from Lake III to Lake IV in the present work.
79
-------
(3) Biomass increases during the course of succession.
This was true only in a very general sense. A review of the data (Tables
Ik and 16, Fig. l6) will show that faunal biomass was not well correlated
with pH. Other factors, especially those associated with meromixis, were
also important.
(k) Net productivity decreases during the course of
successions. An increasingly greater fraction of the gross productivity
is used within the ecosystem as it matures leading to longer food chains
and a greater standing crop of secondary, tertiary, and higher order
consumers. The general increase in the proportion of predators among
the benthos (Table 6) suggests an increase in net productivity, as does
the increase in fish (Table 7) ,which are the top carnivores in those,
lakes where they occur. In Lake V, the increase in net productivity was
apparently translated into a relatively very high biomass of benthic
predators at the expense of the fish population (or perhaps a more
correct view would be that other factors limited the fish and thus allowed
invertebrate predators to increase). The overall increase in the relative
importance of higher order consumers is illustrated by the combined data
(as percent of maximum) for fish and benthic predators (Fig. 22).
(5) Ecosystem organizational complexity increases during
succession. This hypothesis implies such factors as increase in diversity
of pigments and other biochemicals, increase in ecosystem compartments
and alternative pathways, greater habitat diversity, and increase in the
total amount of information in the system. One historical approach to
assessing this aspect of ecosystems has been the study of the relation-
ship between the number of taxa ("biotic diversity") and the siz3 of the
area studied (mainly plant studies) or the number of individuals ((mainly
animal studies).
A much more comprehensive approach to the study of ecosystem complexity
is that of systems ecology. Among others, Van Dyne (1966) and Patten
(1966) have discussed the frame of reference and chief goals of what
might be called the "mainstream" of systems ecology in the U.S. In
general, a systems analysis includes the identification and quantifica-
tion of compartments (variables of habitat, resources, and biocoenosis)
and their interconnecting pathways of material and energy transfer. An
increase in the number of compartments implies more pathways, greater
redundancy, increased feedback, and hence increased homeostasis and
stability. In the lakes of this study, the only really significant
differences in compartments appeared to be those, of the biocoenosis.
That is, the environmental (habitat and resource) variables did not
differ appreciably. From this it follows that the observed increase in
biological compartments at various levels (i.e., the addition of major
taxa such as rooted plants and fish, or of minor taxa such as additional
species of rotifers or insect larvae) implies a corresponding increase
in organizational complexity.
80
-------
80
60
w
!
-------
One line of evidence that bears upon this question is that of seasonal
changes in biomass. If it is true that homeostasis, and hence stabil-
ity, increase with increasing pH in the six lakes, then this should be
"expressed" in terms of a sequential damping of the seasonal oscilla-
tions. An examination of the curves for zooplankton (Fig. 20) suggests
that the more "mature" systems (Lakes IV through VT) do, in fact, have
seasonal variations of somewhat less dramatic magnitude than the less
mature Lakes I and II. As noted previously, the fertilization of Lake
V in summer apparently resulted in dramatic oscillations between May and
September 1970. The bottom fauna cycles (Fig. 21) are expected to be
more oscillatory because of the emergence of aquatic insects at certain
seasons. It appears, however, that Lake VT had much higher minima than
the others.
Thus it would seem that the hypothesis of increasing organizational
stability is supported, in a somewhat limited way, by the data.
D. Alternative patterns of stripland utilization
1. General utilization. The general question of the best use
for abandoned coal stripmine areas has been much discussed (Arata 1957,
Birkenholz 1958, Bowden 1961, Burner 1956, Burner and Schoonover 1953,
Funk 1962, Hall 19^0, Harman 1957, Holmes 19^4, Lewis and Nickum
Limstrom 19^8, Limstrom and Merz 19^9, Roseberry 1963, Riley 19^7,
1957, Sawyer 19^9, Touenges 1939, USDI 1967, Wells 1953, Yeager
and many others). A rather large percentage of the abandoned coal
striplands in southern Indiana and Illinois and in central Missouri that
I have visited are not presently utilized in any fashion, but rather
ignored beyond nominal replanting. Of the many possible uses for old
coal stripmine areas, the following would seem to be among the most
practical.
a. Wildlife habitat. Some areas of old stripmine land in
the Patoka Fish and Game Area have been designated as wildlife areas.
Even with rather extensive planting of food plots, the density of most
species is extremely low. Perhaps improvements in reclamation proce-
dures leading to increased plant growth, and hence in improved cover and
food supply, would increase wildlife populations.
b. Farmlands. In many areas, including some of southwestern
Indiana, the land stripped for coal was never very good farmland. Thus
even if sources of acid pollution were covered and the topsoil restored
to the surface, it would be suitable for little more than rangeland or
pasture. In some areas old striplands have been restored to satisfactory
crop production (see USDI 1967). Recently, sludge from Chicago's
treatment plants has been used to fertilize old stripland in Fulton
County, Illinois, with striking results (Anonymous 1971).
c. Residential. Abandoned stripmine lands can provide space
for human habitation, either in the form of single-family dwellings or
as multiple-dwelling complexes. This use is compatible with a and b
82
-------
above. While some very attractive farm homes, summer cabins, and
permanent residences have been observed by the writer on abandoned
striplands in Indiana, much greater utilization would be possible
without overcrowding or "despoliation" of the landscape.
d« Recreational areas. Some very attractive recreational
sites have been developed on old stripmine areas. Such areas may
include golf courses, camping and picnicking facilities, water sports
areas, etc. (see USDI 1967). Again, the potential for this sort of
development on abandoned striplands has been little realized.
2. Sport fishing in the final cut lakes is quite compatible with
all of these uses. In the course of numerous conversations with local
anglers, it has become apparent to the writer that even the nonacid
stripmine lakes in the study area do not usually provide very good
fishing success. Some results of this study having implications for
the formulation of management techniques that could improve fish produc-
tion in such lakes are here briefly discussed.
a. Meromixis in these lakes has several implications for
their successful management. Therefore, a regular schedule of
limnologies! sampling should be undertaken on any lakes to be manipu-
lated for sport fishing. Such a program should, at a minimum, include
depth series for temperature, dissolved oxygen, pH, conductivity, and
transparency. These data would indicate the presence or absence of
meromixis and the gross effects of any management procedures undertaken.
The two main direct effects of meromixis are: (l) Solutes become
"trapped" in the monimolimnion. This might be desirable in the case
of high concentrations of inorganic ions, but unfortunate in that of
essential nutrients. (2) Dissolved oxygen is usually not present in
the monimolimnion at any time of the annual cycle. Further, during
summer thermal stratification the oxygenated epilimnion may be shallower
than if the lake were holomictic. This confines many groups of animals
to the surface strata where other factors (e.g., temperature) may be
sub optimal.
Under some circumstances it would probably be possible to convert
meromictic stripmine lakes to holomixis by pumping deep water up to the
surface, most effectively at the time of partial overturn. If all
sources of excess solutes could be stopped, a permanent change should
be possible.
b. Basin shape is quite important to the production of benthic
animals, which are, in turn, an important food source of some medium-
and large-size game fishes. In the stripmine lakes, the steep-sided
form of the basin restricts the shallow area of greatest benthic produc-
tion. This effect, as we have seen for Lake III (pp. 55-56), can be
compounded by meromixis. A comparison of bottom fauna distributions
with area and depth (Fig. 23) illustrates this relationship in the
lakes of the present study. The concentration of bottom fauna in the
shallows was especially marked in Lakes III and V, but only in Lake II
was the distribution very uniform.
83
-------
00
09
CO
O
fH
0)
ft
-P
0)
o
o
PH
50
Q
100
CO
03
•H
f
1
O
fH
d)
CM
100
1 2 3
II
o 50
Percent area
OM 1 2 3
100
50
Percent area
100
lOOi
ra
1
O
»H
0)
50
0
III
I T
0
50
Percent
OM
, 2
,
^
area
6
100
T
VI
50 100
Percent area
FIG. 23. Distribution of benthic biomass with depth and area in the Pike County lakes.
-------
Thus one obvious way to improve bottom fauna production in such lakes
•would be to increase the extent of the shallow littoral zone. The
stripmine lake most productive of fish in the Patoka Fish and Game
Area is one in which the littoral was extended in the mid-1960's by
construction of a low earth-fill dam. It is apparently the only strip-
mine lake in the region that supports fish populations approaching
those of typical local nonstripmine ponds. Whenever feasible, the use
of this technique should materially improve fish production in stripmine
lakes.
c. Fertilization, if practiced, would require careful
monitoring and regulation. The resultant high algal production, such
as observed in Lake V, can lead to heavy mortality among fishes and
other animals because of (l) oxygen depletion during the dark period
(or even on cloudy days) resulting from algal decomposition and the
respiration of both algae and the dense zooplankton populations that
accompany such blooms, and (2) toxic substances released by decomposing
algal masses. Excessive algal production can also reinforce meromixis
by adding a biogenic component to the crenogenic element already present.
One intentional effect of lake fertilization is the "shading out" of
rooted aquatic plants (Bass 19&0. While excessive growth of the
macrophytes may be undesirable, their complete and lasting eradication
(as has been accomplished in Lake V) seriously reduces habitat diversity,
which is already quite low in stripmine lakes. The rooted plants provide
food as well as shelter for benthic animals, and are important in
recycling nutrients from the bottom sediments.
-------
SECTION VII
APPENDICES
Appendix 1. Time-depth diagrams
Page
PH 88
Lake I 88
Lake II 89
Lake III 90
Lake IV . 91
Lake V 92
Lake VI 93
Temperature gh
Lake I 9^
Lake II 95
Lake III 96
Lake IV 97
Lake V 98
Lake VI 99
Dissolved oxygen 100
Lake I 100
Lake II 101
Lake III 102
Lake IV 103
Lake V 10^
Lake VI 105
Specific conductance 106
Lake I 106
Lake II 10?
Lake III 108
Lake IV 109
Lake V
Lake VI
87
-------
Ice
00
oo
M AMJ
A'S r o ' N D
FIG. 24. Time-depth diagram of jgH variation in Lake I.
-------
OO
VO
0
M
1 -I
2 -
3 -
5
6
3.3
3.3
J'A'SO'ND'J'F'M'A
1969 1970
nljl-jIAlsroIiTD
PIG. 25. Time-depth diagram of jaH variation in Lake II.
-------
Ice
A ' S ' 0 ' N ' D
FIG. 26. Time-depth diagrm of j>H variation in Lake III.
-------
VO
H
SONDJFMAM
J A S 0 N D
1970
FIG. 2?. Tine-depth diagram of j« variation in Lake IV.
-------
Ice
VQ
to
O'N'DJFMAMJJAS
0 N D
1970
PIG. 28. Time-depth diagram of jgH variation in Lake V,
-------
Ice
(JO
J ' F ' M ' A ' M ' J
'J'A'S'O'N'D
FIG. 29. Time-depth diagram of jgH variation in Lake VI.
-------
Ice
M ' A ' M ' J
N
FIG. 30. Tiae-depth diagram of teaperature variation (°c) in Lake I.
-------
Ice
VJl
J
1969
S ' 0 N ' D
J P
1970
M'A'MJJ'A'S
FIG. 31. Time-depth diagram of temperature variation (°c) in Lake II.
-------
Ice
r. i _ i .. I . i .. I . I . I .—i _ i „ i „ i
DJFMAMJJASOND
J ' A ' S ' 0 ' N
1969
1970
PIG. 32. Time-depth diagram of temperature variation (°c) in Lake III.
-------
Ice
M A M J
J A S 0 N D
1970
PIG. 33. Time-depth diagram of temperature variation (°C) in Lake IV.
-------
Ice
O'ND'J'F'M'A
5 -
M'J'J'A'S'O'H'D
FIG. 3*f. Time-depth diagram of temperature variation (°C) in Lake V.
-------
Ice
J ' P
1970
M'A'M'J'J'A'S'O'N'D'
PIG. 35. Time-depth diagram of temperature variation (°c) in Lake VI.
-------
loe
ML?
7 65^ 321 nw
J'F'MA'M'J'J'A'S'O'N'D
FIG. 36. Tiae-dcpth diagram of dissolved oxygen variation («g/l) IB Lake ^.
-------
Ice
s
FIG. 37. Time-depth diagram of dissolved oxygen variation (ng/l) in Lake II.
-------
Ice
O
ro
8
J^ A FSr 0
1970
-------
Ice
s-
J'ASO'ND'J
10 _
P • M A
M'J'J'A'S'O'N'D
1969
1970
PIG. 39. Time-depth diagram of dissolved oxygen (mg/l) in Lake IV.
-------
Ice
'S'O'N'D'J'FMA
PIG. 40. Time-depth diagram of dissolved oxygen variation (mg/l) in Lake V.
-------
Ice
6
VJl
H ' D
FIG. 41. Time-depth diagram of dissolved oxygen variation (ag/l) in Lake VI.
-------
O
ON
2000
loe
J
1970
M ^ j rj 'A s ' o
FIG. 42. Time-depth diaaaa of apeoific conductance (K»g*10 ) In Lake I.
-------
Ice
J ' A ' S r 0 ' N
1969
D'J'F
1970
MA'M'J'J'AS'O'N'D
FIG. 43. Tiae-depth diagraa flf apeciflc conductance (Kg<«10") in Lake II,
-------
Ice
1970
FIG. *»4. Time-depth diagram of specific conductance (Kog»10°) in Lake III.
-------
Ice
O
VO
DJPMAMJJASOND
FIG. **5. Time-depth diagram of specific conductance
in Lake IV.
-------
Ice
O
J AS O
1970
FIG. 46. Time-depth diagram of specific conductance (foe'10") in Lake V.
-------
Ice
N ' D
PIG. 4?. Time-depth diagram of specific conductance (foe;* 10°) in Lake VI.
-------
APPENDIX 2. Seasonal variation in surface (S) and bottom
(Bj concentrations
Lake I
29 Jan
Sampling 27 Mar
Dates 12 June
(1970) 11 Aug
ll* Oct
17 Dec
Calcium S 520
392
1*60
1*21*
368
390
B 1*80
W
1*1*0
1*32
1*16
390
Magnesium S 3^0
281*
317
II
28 Jan
1* Apr
1* June
18 Aug
27 Oct
2 Dec
380
320
320
328
328
328
390
328
32i*
322
31.8
332
178
190
189
III
19 Feb
31 Mar
3 June
17 Aug
12 Oct
2l* Dec
1M
120
160
212
212
181*
500
1*90
1*88
1*88
1*61*
1*60
110
85
122
of selected ions (mg/1).
IV
26 Jan
31 Mar
3 June
17 Aug
12 Oct
2l* Dec
196
ll*l*
268
368
372
295
1*32
1*90
1*16
1*1*0
1*1*0
1*1*0
ll*l*
100
193
V
28 Jan
31 Mar
3 June
11 Aug
12 Oct
ll* Dec
320
201*
312
320
320
320
392
1*00
360
1*1*8
l*2l*
1*08
105
ll*6
210
VI Control
28 Jan
1* Apr
1* June 10 June
15 Aug
13 Oct
9 Dec 9 Dec
160
161*
11*8 11
152
172
181* 13
236
281*
300 21
336
31*8
288 13
122
117
10l* 6
112
-------
APPENDIX 2. Continued
Iron
(Dissolved)
356
337
250
B 380
1*68
to
1*88
386
275
S 105
77
65
60
78
105
B 2to
21*1*
2to
130
90
137
S 52
to
50
200
161*
178
2to
198
190
191*
173
178
5.1
8.8
3.5
3.7
3.5
3.9
l*.l*
9.8
6.9
5.0
3.8
1*.2
31*
33
32
163
168
122
525
1*50
527
537
517
to
10.0
6.1*
0.65
0.9
2.1*
13.7
130
122
135
170
270
261*
8.5
5.8
9.5
221*
237
125
322
300
307
1*03
376
327
i*.o
1.0
0.3
0.2 '
0.25
0.8
100
80
93
350
350
282
5.9
5.5
10.5
220
239
231*
298
293
378
327
337
317
2.0
1.1
0.2
0.05
0.05
0.65
51
H.5
70
63
55
1*0
-
8.2
5.8
117
131*
130
161*
195
207
231*
258
220
<0.05
0.55
-------
APPENDIX 2. Continued
Aluminum
Sodium
0.2
J( JT ~'S -' "
-. r. _l «-• r- )l A
B
63 34 29 19.5 13.5 6.5
8.0
52
52
57
60
63
64
60
62
59
36
25
34
37
4o
43
27
25
23
32
44
41
17
11
13
32.5
36
34
38
34
31
32.5
35
33
15
13.8
12.4
11.5
11.8
12.8
15
13
3J..4
11.2
12.1
12.8
47
41
41
9.7
10.7
8.5
34
29
32
30
31.5
30
0.5
0.7
0.7
0.05
0.5
0.4
0.7
0.6
0.8
0.6
0.5
0.05
23
16
19
114
4.8
3.3
4.1
21.5
19.5
18.5
29
29
26
1.1
0.4
0.05
0.05
0.5
0.9
<0.05
0.05
0.05
0.05
0.5
0.05
24
20
28
0.05
1.7
2.9
-
13.5
16.5
21
24
27.5
-
0.3
0.05
0.05
0.05
<0.05
-
0.1
-------
APPENDIX 2. Continued
5.6
B
4.4
5.6
Potassium ._ ... „.„ —
3.7 5.2 3.5 3.2 3.2 4.6
1.2
Zinc
16
17
16
2k
20
19
23
19
19
4.4
3.7
4.5
5.3
5.2
4.8
6.3
7.1
7.1
8.2
5.7
5.6
3.0
t
2.2
2.9
45
47
43
53
Ul
41
45
46
45
5.5
5.2
4.8
-
5.5
5.6
5.6
5.2
4.8
-
5.5
5-5
0.75
0.95
0.80
24
26
19
37
32
31
36
40
40
4.1
3.5
2.5
4.5
4.5
4.3
11.0
9.0
9.4
14
25
26
0.15
0.20
0.20
35
37
24
38
35
34
45
45
45
4.0
3.2
3.7
M
5.5
4.7
9.0
8.4
8.7
22
25
22
0.20
0.25
0.05
15
19
19
20
16
18
21
24
24
3.5
3.2
4.0
3.9
4.5
4.2
4.7
4.6
5.5
5-9
6.0
5.3
-
0.25
0.05
**9
54
58
60
65
62
75
73
71
4.2
4.6
3.9
4.3
4.7
5.0
5.9
6.9
6.8
7.8
5.0
7.3
0.10
0.05
<0.05
2.2
B
2.3
2.3
<0.05
115
-------
Total
Hardness
(Ca, Mg,
Fe, Mn, Al,
and Zn as
Sulfate
APPENDIX 2. Continued
3.0
3.1
3.0
B 2.2
l.U
2.0
3.0
k.O
3.5
S 3105
2^50
2790
2865
2690
2^55
B 3kOO
3685
3585
35^5
3065
2610
s 2750
2365
3^50
0.80
0.90
0.8o
0.80
1.00
0.95
0.85
0.95
0.90
1805
1710
1690
1730
1610
1665
2095
1760
1700
1710
1700
1675
1650
1550
1700
0.25
0.20
0.20
0.35
0.50
o.ho
o.Uo
0.20
0.10
860
675
920
1220
12^5
1000
3705
3375
3685
3785
3825
3530
750
510
875
0.10
0.05
0.05
0.05
0.05
0.10
0.15
0.10
0.05
1100
785
iWo
1850
1900
1265
2615
2680
2500
3^35
3325
2990
925
600
1300
<0.05
0.05
<0.05
-
0.10
<0.05
0.10
0.05
<0.05
1235
1130
1650
1700
1785
1765
2315
22l*5
2605
2610
2585
2W)
1250
775
1600
<0.05
<0.05
<0.05
0.10
0.05
<0.05
0.05
0.05
<0.05
900
890
800
860
980
loto
1270
1525
1615
1900
1985
16^0
750
750
675
<0.05
<0.05
<0.05
50
60
135
100
35
116
-------
Chloride
APPENDIX 2, Continued
2900 16?5 1075 1625 1500 61*0
3150 1575 1150 1900 1650 1050
2950 1875 875 1125 1600 925 30
B 3650 2125 2850 2050 2250 1000
3^00 1550 3200 2150 1550 1025
3^50 1850 3275 2190 2200 1250 5
3350 1700 3300 2500 1875 1350
3350 1625 3300 2650 2000 1675
3250 1800 3^00 2650 2100 1300 25
B
8
3
if
3
ifO
2
lif
3
5
3
ifO
3
if
5
if
6
5
if
if
5
if
6
5
if
19
15
16
18
20
15
8
10
8
9
13
9
18
21
23
26
25
18
15
22
19
37
18
19
5
if
6
3
6
if
8
3
3
<1
6
5
3
3
2
3
5
if
3
if
3
if
8
if
Bicarbonate S
2lf 51 n^ 160
<1
117
-------
APPENDIX 2. Continued
Nitrate
47
B
45
Silica
(Si02)
B
0
0
0
39
39
38
50
45
48
47
43
36
±3
52
43
<5
5
<5
0
0
0
32
25
23
27
27
28
31
25
24
28
27
27
5
<5
<5
8
233
0
10
9
12
13
12
11
13
12
17.1
23
15
10
5
<5
<5
71
177
-
8
8
8
9
8
8
12
9-2
11.5
29
12
10
<5
<5
<5
181
284
366
12
11
12
9
9
8
15
14
14
21
15
16
<5
<5
<5
213
296
528
2.1
2.4
1.6
1.4
1.7
2.1
4.5
5.0
4.2
5.0
9.0
8
<5
<5
<5
<5
118
-------
APPENDIX 2. Continued
10
<5
<5
B 5
5
<5
10
<5
<5
Total S <0.1
Phosphate <0.1
-------
SECTION VII
ACKNOWLEDGMENTS
•A large number of people have contributed in various ways to the
realization of this project, including the following to whom I am
especially indebted:
My advisor, Dr. D. G. Frey; Dr. Allen F. Agnew, former Director of
the Indiana University Water Resources Research Center; my Project
Officer, Quentin Pickering of the Environmental Protection Agency;
Richard E. Bass and Donald Mann of the Indiana Department of
Natural Resources; Lowell Oxley for permission to work on his
property; my research assistant, Miss Nancy Tormohlen; Mrs.
Betty Lucas of the Indiana University Water Resources Research
Center; Dr. W. R. Breneman, Frank N. Young, Craig E. Nelson,
Charles B. Heiser, and Richard C. Starr of Indiana University;
Dr. R. S. Campbell of the University of Missouri; and Dr.
Charles Krebs of the Institute of Animal Resources Ecology,
University of British Columbia. My wife, Christina C. Smith, has
helped from time to time with virtually all phases of the work
from the planning stage through the typing of the final manuscript.
I am deeply indebted for her help and encouragement.
This project was made possible by financial assistance from the
Indiana Department of Natural Resources (A.R.P. No. 3^2-303-721)
and from the U.S. Environmental Protection Agency (Grants 18050
EEC and 18050-2 EEC; FWQA: 53-3^2-26).
121
-------
SECTION VIII
LITERATURE CITED
1. Ahlstrom, E. H. 19^0. A revision of the Rotatorian genera
Brachionus and Platyias with descriptions of one new species
and two new varieties. Bull. Amer. Mus. Nat. Hist. 77:lk3-lQk.
2. Ahlstrom, E. H. 19^3- A revision of the rotatorian genus Keratella
with descriptions of three new species and five new varieties^
Bull. Amer. Mas. Nat. Hist. 80:Ull-l*57.
3. American Public Health Association. 1965. Standard methods for
the examination of water and wastewater. 12th ed. A.P.H.A. ~New
York, New York. 769 p.
k. Anonymous. 1971. The value of sludge. Time. 27 September, p. 93.
5. Arata, A. A. 1957. Trapping the striplands. Illinois Wildlife.
12:5-6.
6. Arata, A. A. 1959- Ecology of muskrats in strip-mine ponds in
Southern Illinois. J. Wildlife Manage. 23:177-186.
7. Bass, R. E. 196^. Report of the strip pit investigations and
management on the Patoka Fish and Game Area, Pike County, Indiana.
Unpublished report. Indiana Department of Natural Resources,
Indianapolis.
8. Bell, R. 1956. Aquatic and marginal vegetation of strip mine waters
in southern Illinois. Trans. Illinois Acad. Sci. 1*8:85-91.
9. Birkenholz, D. E. 1958. Reclamation of a spoil bank area for
wildlife purposes. M.S. Thesis. Southern Illinois University,
Carbondale.
10. Bowden, K. L. 1961. A bibliography of strip-mine reclamation: 1953-
1960. University of Michigan Department of Conservation. 13 p.
11. Brewer, R. 1958. Breeding bird populations of strip-mined land
in Perry County, Illinois. Ecology. 39:5^3-5^5.
12. Burner, C. C. 1956. Summary of fish growth and fishing success
in managed strip-pits through spring 1956. Mimeographed report.
13. Burner, C. C. and C. Leist. 1953. A limnological study of the
college farm strip-mine lake. Trans. Kansas Acad. Sci. 56:78-85.
lif. Burner, C. C. and R. Schoonover. 1953. Progress report on fisheries
management projects in Linn County, Kansas. Mimeographed report.
Forestry, Fish and Game Commission, Pratt, Kansas.
123
-------
15. Burt, W. H. and R. P. Grossenheider. 196U. A field guide to the
mammals . 2nd ed. Houghton Mifflin, Boston. 284 p.
16. Campbell, R. S., 0. T. Lind, G. L. Harp, W. T. Gelling, and J. E.
Letter Jr. 1965a. Water pollution studies in acid strip-mine
lakes: changes in water quality and community structure associated
with aging. In: Symp. on Acid Mine Drainage . Mellon Institute,
Pittsburgh, Pa. 11 p.
17. Campbell, R. S., 0. T. Lind, W. T. Geiling, and G. L. Harp. 1965b.
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129
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SECTION IX
LIST OF APPENDICES
Page
A. Appendix 1. Time-depth diagrams 8?
Fig. 2k'. Time-depth diagram of pH variation in Lake I. 88
Fig. 25: Time-depth diagram of pH variation in Lake II. 89
Fig. 26: Time-depth diagram of pH variation in Lake III. 90
Fig. 2?: Time-depth diagram of pH variation in Lake IV. 91
Fig. 28: Time-depth diagram of pH variation in Lake V. 92
Fig. 29: Time-depth diagram of pH variation in Lake VI. 93
Fig. 30: Time-depth diagram of temperature variation
(°_C) in Lake I. 9^
Fig. 31: Time-depth diagram of temperature variation
(°C) in Lake II. 95
Fig. 32: Time-depth diagram of temperature variation
(°C) in Lake III. 96
Fig. 33: Time-depth diagram of temperature variation
(°C) in Lake IV. 97
Fig. 3^-: Time-depth diagram of temperature variation
(°C) in Lake V. 98
Fig. 35: Time-depth diagram of temperature variation
(°C) in Lake VI. 99
Fig. 36: Time-depth diagram of dissolved oxygen
variation (mg/l) in Lake I. 100
Pig» 37: Time-depth diagram of dissolved oxygen
variation (mg/l) in Lake II. 101
Fig. 38: Time-depth diagram of dissolved oxygen
variation (mg/l) in Lake III. 102
Fig. 39: Time-depth diagram of dissolved oxygen
variation (mg/l) in Lake IV. 103
Fig. 40: Time-depth diagram of dissolved oxygen
variation (mg/l) in Lake V. 104
Fig. Ul: Time-depth diagram of dissolved oxygen
variation (mg/l) in Lake VI. 105
Fig. k2: Time-depth diagram of specific conductance
(K25»10°) in Lake I. 106
Fig. ^3: Time-depth diagram of specific conductance
(K25-10&) in Lake II. 107
Fig. ¥f: Time-depth diagram of specific conductance
(K^5-10°) in Lake III. 108
Fig. k$: Time-depth diagram of specific conductance
(Kp5-10°) in Lake IV. 109
Fig. k6: Time-depth diagram of specific conductance
(K25'10&) in Lake V. 110
Fig. ^7: Time-depth diagram of specific conductance
in Lake VI. HI
131
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B. Appendix 2. Seasonal variation in surface (S) and bottom
(B) concentrations of selected ions (mg/1). 112
132
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1
5
Accession Number
f\ Knbiert Field St. Group
Acid-Mine Pollution
Lake Biology
SELECTED WATER RESOURCES ABSTRACTS
INPUT TRANSACTION FORM
Organization
Indiana University Foundation
Indiana University, Bloomington, Indiana
Title
"Acid Mine Pollution Effects on Lake Biology"
1 Q Authors)
Smith, Ronald. W.
Frey, David G.
16
21
Project Designation
EPA WQO Contract No. 53-3^2-26
Project No. l8050-EEC
Note
22
Citation
23
Descriptors (Starred First)
*acidic water
*strip-mine lakes
benthic fauna
plankton
aquatic productivity
ecosystems
limnology
water properties
25
Identifiers (Starred First)
02. Water cycle
2H. lakes
2K. chemical processes
05. Water quality management & protection
*5C. effects of pollution
27
Abstract
Six coal stripmine lakes in southern Indiana encompassing a pH range of 2.5 to 8.2
were studied from July 1969 to December 1970. Generally, differences between the
lakes indicated successions! trends with increasing pH. Environmental trends in
the surface waters included increasing levels of dissolved oxygen and decreasing
concentrations of dissolved substances. These tendencies were somewhat obscured by
differences in the annual cycles of stratification, four of the lakes proving to
be unexpectedly meromictic. Biological changes associated with increasing pH
included increasing diversity and increasing homeostasis. Biomass was influenced
by both pH and circulation patterns (meromixis vs. holomixis), and bottom fauna
was further limited by the steep-sided basin form. All the stripmine lakes had
much higher solute concentrations and lower biological diversity than a small local
non-stripmine reservoir studied as a control. A fertilization program in one lake
has apparently produced elimination of all rooted aquatic plants, violent oscilla-
tions of plankton, and low fish populations. It is suggested that sport fishing in
stripmine lakes, not presently very satisfactory, could be improved by management
techniques adapted to their unique limnological nature.
Abstractor
Ronald W. Smith
Institution
Indiana University Foundation
WR:I02 (REV. JULY I98SI
WRSIC
SEND TO: WATER RESOURCES SCIENTIFIC INFORM*
U.S. DEPARTMENT OF THE INTERIOR
WASHINGTON. D. C. 20240
TION CENTER
*U.S. GOVERNMENT PRINTING OFFICE: 1972 484-4*3/107 1-3
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