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OCR for page 300
8
Fish Population Trends
in Response to
Surface Water Acidification
Terry A. Haines
INTRODUCTION
The most widely reported consequence of acid deposition
is the reduction or elimination of fish populations in
response to surface water acidification (Overrein et al.
1980, Haines 1981, Altshuller and Linthurst 1984, Dillon
et al. 1984). To demonstrate conclusively that fish
population trends in a body of water are related to
atmospheric acid deposition, temporal associations between
surface water chemistry and fish populations are required
It must be shown that the water body formerly supported a
viable fish population, either self-sustaining or
hatchery-maintained; that one or more fish species
formerly present have been reduced or eliminated; that
the water body is more acidic now than when f iSh were
present and that the increased acid level was not caused
by local factors; and that other adverse factors are
either absent or unimportant. The number of available
data sets meeting these criteria is very small, and most
have not been published in the peer-reviewed literature.
A larger number of data sets exist that demonstrate
spatial associations between surface water chemistry and
fish populations. In some cases, laboratory or field
experiments serve to confirm the spatial associations
between fish species with water chemistry parameters
linked to acidification (e.g., pH and aluminum). In
other cases temporal data on fish populations are
available, but there are no temporal data on water
quality; or the converse may be true. Other than
temporal association studies with data on both fish
populations and water quality, the strongest cases for
demonstrating adverse effects of acidification on fish
population trends are those that combine spatial data
300
1
OCR for page 301
301
with one or more of the above types of evidence (Magnuson
et al. 1984). These studies are also very scarce. In
this chapter I review available North American data sets
that purport to relate fish population status or trends
to acid deposition.
LABORATORY INVESTIGATIONS
Laboratory experiments have confirmed that chemical
parameters associated with acidification, such as pH,
dissolved organic carbon, levels of aluminum, and calcium
affect the survival of fish. These laboratory studies
have been reviewed in Altshuller and Linthurst (1984) and
Haines (1981).
Exposing fish to various pH levels reveals intra-
specific and interspecific differences in acid tolerance
(Tables 8.1 and 8.2). The environment, however, is much
more complex than this, and acid tolerance is also
affected by other factors. Varying concentrations of
toxic trace metals, such as aluminum and cadmium, can act
synergistically with acid, while divalent cations,
especially calcium, and organic ligands can act antago-
nistically with acid and trace metals. In general,
reduced pH and increased trace metal concentrations are
toxic to fish, and this toxicity is reduced by increased
levels of dissolved organic carbon and divalent cations.
Toxicity varies in a complex manner with changing propor-
tions of the toxic and mediating factors, fish life his-
tory stage, and fish species. Given the complexity of
the interactions it is generally not possible to predict
the response of a fish species population to acidification
in the absence of considerable experimental evidence.
Such evidence is currently available for only a few fish
species, such as brook trout (Salvelinus fontinalis) and
white sucker (Catostomus commersoni), although data are
now being collected on additional species.
F IELD EXPERIMENTS
A few researchers have studied the effects of
acidification on fish using manipulation experiments.
Hall et al. (1980) acidified a small stream in New
Hampshire, but the stream did not support a permanent
fish population. A few adult brook trout were present in
the acidified reach during part of the year, but they did
not exhibit morphological symptoms of acid stress.
OCR for page 302
302
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OCR for page 304
304
Mills (1984) acidified Lake 223 of the Experimental
Lakes Area, Ontario, by direct addition of sulfuric
acid. The lake pH was reduced from 6.49 (1976) to 5.17
(1983). Lake trout (Salvelinus namaYcush), white sucker
(Catostomus commersoni), fathead minnow (Pimephales
promelas), and slimy sculpin (Cottus cognates) were
abundant at the start of the experiment, and pearl dace
(Semotilus margarita) were rare. Fathead minnow and
slimy sculpin abundance declined in 1979 (pH 5.64), lake
trout declined in 1980 (pH 5.59), and white sucker
declined in 1981 (pH 5.16). Pearl dace first increased
rapidly in 1980, following the decline of fathead minnow,
then declined in 1982 (pH 5.11). - ~ -I
fully reproduced in 1982.
NO clan species success-
Adults of some species survived
DUt were In generally poor condition because of the loss
of forage species. It is difficult, however, to extrapo-
late these results to areas that receive acidic deposi-
tion. Inasmuch as the acid was added directly to the
lake, the dissolved organic carbon and trace metal
(especially aluminum) concentrations in the lake may not
reflect those in lakes in which the watershed also
receives acid inputs.
SPATIAL ASSOC IATION S
South Central Pennsylvania
Personnel from the Institute for Research on Land and
Water Resources, Pennsylvania State University, have
conducted a number of studies in the Laurel Hill area of
south central Pennsylvania. In one, researchers
conducted a spatial association study in 61 streams on
Laurel Hill (W. Sharpe, Pennsylvania State University,
personal communication, 1985; Figure 8.1). For 12 of the
streams on-site surveys identified significant cultural
disturbances in the watershed that were expected to
affect water quality, such as agricultural, highway, and
industrial discharge, and these streams were therefore
excluded from further consideration. The remaining
streams were located in watersheds with only minor land
use activities. These streams were surveyed for water
chemistry and fish population (by electrofishing a 100-m
section) and were divided into three groups: those with
reproducing fish populations (young-of-the-year fish of
nonhatchery origin present), those with remnant fish
populations (only 2-year classes), and those with no fish
(Table 8.3).
OCR for page 305
305
79o158
0 5 10 kilometers
4oo 1 5
Ligonier
17
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2 1
Johnstown
~O~Ddi
3 1
79°1 5'
~3
-
- _ Stream
O Basin Boundary
. 4ooool
Reproducing cold water fishery absent (12
Remnant cold water fishery remaining
( 2 year classes ) (4)
Confounding cultural impacts (12)
Reproducing cold water fishery present (3
FIGURE 8.1 Location and fish population status of 61
streams on Laurel Hill, south central Pennsylvania.
Source: W. Sharpe, Pennsylvania State University.
The 33 streams that contained reproducing fish
populations all were of pH 6.0 or higher, and aluminum
concentrations were less than 100 ~g/L. Of the 12
streams lacking reproducing fish populations, only one
(Beaver Dam Run, pH 6.22) had a pH above 6.0. Eight of
the 12 had pH values less than 5.0 and aluminum
OCR for page 306
306
TABLE 8.3 Fish Population Status and Water Chemistry of 49 Streams on
Laurel Hill, South Central Pennsylvania
Fish Population Number Mean pH Mean Alkalinitya Mean Aluminumb
Status of Streams (range) (range) (range)
ReproducingC 33 6.73 139 21
(6.0-7.2) (27-340) (5-97)
Remnants 4 6.36 88 49
(5.7-7. 1) (6-262) (18-100)
Not reproducinge 12 4.97 7 454
(4.3-6.2) (0-43) (15-1 ,000)
a Units are microequivalents per liter.
b Units are micrograms per liter total filterable.
C Young-of-the-year fish present.
Only two year-classes present.
eNo young-of-the-year fish present.
SOURCE: W. Sharpe, Pennsylvania State University.
concentrations of 400 ug/L or more. The 4 streams with
remnant fish populations were chemically quite variable,
with pH values ranging from 5.7 to 7.1. Of the streams
that contained reproducing fish populations, 10 contained
only one species (brook trout), 18 contained two species
(brook trout plus mottled sculpin in 17, brook trout plus
blacknose dace in 1), 4 streams contained three species
(brook trout, mottled sculpin, and brown trout in 2, and
rainbow trout in place of brown trout in the remaining
2), and 1 stream contained four species (the three
salmonids plus mottled sculpin).
Fish population status in these streams is thus
generally related to stream chemistry. Although there
are exceptions, the more acidic streams are generally
those with sparse or no fish population. This does not
prove cause and effect, although other explanations
involving factors other than acidification have been
ruled out.
Vermont
Langdon (1983, 1984) conducted a spatial association
study of fish populations and water chemistry in 29
Vermont lakes. The lakes selected for the fisheries
survey were chosen from a larger set of lakes (Clarkson
1982) and represented the lowest alkalinity lakes in the
OCR for page 307
307
survey.
_ Fish were collected with experimental (variable
mesh size) gill nets, baited minnow traps, and seines.
Two of the lakes were fishless, and 27 contained 1 to 10
fish species.
For this report I conducted a stepwise multiple-
regression analysis for physical and chemical factors
that could potentially affect the number of fish species
in these lakes. The variables considered were pH,
specific conductance, color, sum of divalent cations
(calcium and magnesium), surface area, maximum depth, and
elevation. The only significant (p < 0.05) variables
were pH-(p = 0.0001), sum of divalent cations (p = 0.026),
and surface area (p = 0.028). The regression equation is
, ~ ~
log(number of species + 1) = -1.33 + 0.41 (pH)
-0.004 (sum of divalent cations) + 0.0004 (surface
2 area);
r = 0.62, p = 0.0001, n (number of lakes) = 29.
I also compared fish abundance, expressed as catch per
day (a standard unit of effort), with the same physical
and chemical variables. Similar to the above analysis
the three statistically significant variables of fish
abundance were pH (p = 0.048), sum of divalent cations (p
= 0.006), and surface area (p = 0.070). The regression
equation is
log(catch per unit effort + 1) = -1.17 + 0.57 (pH)
-0.023 (sum of divalent cations) + 0.001 (surface
area);
r2 = 0.51, p = 0.001, n = 29.
Generally, smaller lakes have lower pH and contain
fewer numbers of fish species and individuals. Sur-
prisingly, the relationship with divalent cations was
reversed; larger lakes were lower in divalent cations
than expected. Attempts to partition the variability
between pH and surface area were unsuccessful. Apparently
these two variables are highly correlated.
The lakes surveyed were generally not affected by
cultural disturbances. Most contained no permanent
structures in the watershed, although five contained
seasonal dwellings and one contained year-round dwellings.
Many of the lakes, however, were stocked with salmonid
fish, but only one lake contained stocked salmonids
alone. All the lakes were accessible to fishing and,
fishing pressure was not believed to be excessive for any
lake.
OCR for page 308
308
The previous presence of fish populations in the two
lakes that are now fishless is equivocal. One lake is
known to have contained fish previously but may have been
affected by beaver activity that blocked fish access to a
spawning stream. There are no reliable data to confirm
that the other lake ever contained fish, but there are
anecdotal records that suggest that it did (Langdon 1983).
Maine
Haines (1985) conducted a spatial association survey
of water chemistry and fish population in 22 lakes in
Maine. The lakes were all low in color and contained no
human habitation or other recent land disturbance in the
watershed. None had been stocked, reclaimed, or otherwise
manipulated, and they were sufficiently remote from
vehicle access as to make casual introduction of fish
species or intensive fishing pressure unlikely. A survey
team visted each lake at least three times. The team
collected water samples in spring, summer, and fall to
analyze for pH, alkalinity, specific conductance, color,
and all major ions. Fish populations were surveyed once,
in summer, using a standard collecting protocol. Two
experimental gill nets (6 ft deep containing five 25-ft
long panels of 0.5-, 0.75-, 1-, 1.25-, and 1.5-inch mesh
square measure) were set from late afternoon until
midmorning the following day. One net was set in shallow
water and one in the deepest area.
In addition, six
standard wire mesh (0.25 inch square measure) minnow
traps were set in various locations in the littoral area
for the same time period. The team counted and
identified the species of all fish collected.
A stepwise multiple-regression analysis was conducted
following the same procedure used with the Vermont
spatial association data. For number of fish species the
significant variables were sum of divalent cations (p =
0.045), and surface area (p = 0.042).
important variable was pH (p = 0.21).
equation (including pH) is
log(number of fish species + 1) = -0.659
The next most
The regression
+ 0.003 (sum of divalent cations) + 0.008 (surface
2area) - 0.124 (pH);
r = 0.66, p = 0.0002, n = 22.
OCR for page 309
309
Fish abundance was also analyzed as for Vermont. The
only significant variable was sum of divalent cations (p
= 0.066). The next most important variable was pH (p =
0.23). The regression equation (including pH) is
log(catch per unit effort + 1) = -1.27
+ 0.011 (sum of divalent cations) + 0.353 (pa);
r2 = 0.53, p = 0.0007, n = 22.
The number of significant variables for the Maine data
was less than for the Vermont data, but the same variables
were identified as most important in both cases. In the
Maine lakes the relationship of divalent cations was
direct, as expected, rather than inverse. In general,
small lakes were more acidic, lower in divalent cations,
and contained fewer numbers of species and individuals.
Attempts to partition the variability among pH, divalent
cations, and area were unsuccessful, apparently because
of the high degree of correlation among these variables.
Wisconsin
Several authors have reported results of fish popula-
tion and water chemistry surveys in northern Wisconsin
lakes. Rahel and Magnuson (1980) reported no relation-
ship between lake pH and number of fish species present
in lakes that were similar in size and habitat. Con-
versely, the same authors (Rahel and Magnuson 1983)
reported that cyprinids and darters were generally absent
from Wisconsin lakes of pH ~ 6.2. However, oligotrophic
lakes with pH near 7.0 also lacked acid-sensitive fish
species because of species interactions and biogeo-
graphical factors.
Wiener et al. (1984) surveyed fish populations in two
groups of lakes in northern Wisconsin. All lakes were
seepage lakes, but one group was acidic with pH levels
near 5.0 and the other consisted of lakes with near-
neutral pH levels. The acidic group contained signifi-
cantly fewer fish species than the near-neutral group,
and the absent species consisted primarily of cyprinids
and darters. However, it is doubtful that these lakes
have been acidified by atmospheric deposition. (See
Chapter 7.) These data confirm that fish species
distribution is affected by water chemistry factors
related to acidification but do not demonstrate trends
related to atmospheric deposition.
OCR for page 310
310
TEMPORAL ASSOCIATIONS
Ontario
Lakes located in the LaCloche Mountain region of
Ontario constitute one of the best North American data
sets documenting the adverse effects of acid deposition
on fish populations, although the major source of acid is
certainly the sulfur dioxide emissions from metal smelters
located 65 km to the northeast in Sudbury, Ontario
(Beamish 1976). Acidification has occurred with suf-
ficient rapidity that declines and in some cases
extinctions of fish populations have been documented
since the late 1960s (Beamish and Harvey 1972, Beamish
1974a,b, Beamish et al. 1975, Harvey 1975, Beamish 1976,
Beamish and Van Loon 1977, Harvey and Lee 1982). The
acidic deposition is accompanied by increased deposition
of certain heavy metals (e.g., copper, nickel), which may
also be toxic to fish (Beamish 1976, Beamish and Van Loon
1977). Of the 212 lakes in the LaCloche Mountains, 68
have been surveyed for fish populations (Harvey 1975).
Of these, 38 lakes are known or suspected to have had
reductions in the number of fish species, and 54 fish
populations are known to have been lost (Harvey and Lee
1982). The number of fish species per lake was signifi-
cantly (p = 0.005) correlated with lake pH, even after
correction for differences in lake area. Explanations
for declines in fish populations other than changes in
water chemistry related to acid deposition have been
ruled out (Altshuller and Linthurst 1984).
Adirondack Mountains, New York
The first reports of acid-deposition-induced acidifi-
cation of lakes and concomitant reductions in fish
populations in North America were for the Adirondack
Mountain region (Schofield 1965, 1973, 1976a,b). Several
recent survey reports (Pfieffer and Festa 1980; Colquhoun
et al. 1984) summarized recent data concerning water
chemistry and fish populations in lakes within the
Adirondack ecological zone. Schofield (1976b) surveyed
40 lakes that had been initially surveyed in the 1930s.
In the 1930s three lakes had pH < 5.0 and no fish, and
one lake with pH between 6.0 and 6.5 also had no fish.
In 1975, 19 of these 40 lakes had pH < 5.0 and had no
fish. Two additional lakes with pH between 5.0 and 5.5
OCR for page 324
324
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325
1967. Inasmuch as pH was measured calorimetrically both
times, the difference seems unlikely to be a result of
methodological differences. (See Chapter 7.)
Two streams, Wilder and Templeton, had pH < 5.0 in
1983 and had lost all fish species by 1967. These streams
were of different character than the others surveyed,
being of lower gradient (i.e., drop in elevation per
stream mile), and higher color (organic acids), and con-
taining primarily warm-water fish species. Nevertheless,
pH apparently has declined and fish communities have been
lost in these streams. There is no apparent source of
Acid other than atmospheric deposition.
Nova Scotia
Watt et al. -(1983) compared temporal trends in angler
harvest of Atlantic salmon with current water chemistry
in 22 rivers in Nova Scotia. Angler harvest data were
available from 1936 to the present and were nearly
continuous. Earlier data exist, but the methodology used
is not comparable with that used subsequent to 1936.
Previous (1954-1955) water chemistry data of high quality
(multiple pH values measured electrometrically) were
available for five rivers. Twelve of the r i Ye c now Hal
pH > 5.0 (Table 8.9). Three of these have previous pH
data available; two (St. Marys and Musquodobit) have not
changed significantly, and one (Medway) declined 0.8
unit. Ten of the twelve rivers had no significant change
in salmon harvest, one (Gold) increased significantly,
and one (Musquodobit) decreased significantly although
its pH did not change appreciably.
Ten of the rivers now have pH < 5.0. Two of these
(Clyde and Tusket) have previous pH data and both declined
in pH, but statistical tests of the decline could not be
performed because of inadequate data. (There is only a
single historical value in one case, and there was a
change in sample collection site in the other.) Nine of
the ten rivers with low pH exhibit a significant decline
in angler harvest of salmon, which generally began in the
1950s; the other's decline (Lisomb) was not significant.
The rivers included in this comparison were carefully
selected to eliminate any that were affected by other
pollution sources, fish stocking, or change in physical
barriers to fish migration. It is presumed that fish
from rivers with geographical proximity have similar
migratory patterns and are subjected to similar levels
~ ~ .. ~ ~ ~
OCR for page 326
326
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OCR for page 327
327
and sources of mortality away from the home rivers
Other studies indicate a significant mortality of early
feeding fry of Atlantic salmon in a Nova Scotia river
with pH < 5.0 (Lacroix et al. 1983)
SUMMARY
.
.
The hypothesis that increased acidity of surface
waters caused by long-range atmospheric transport has
reduced or eliminated fish populations in northeastern
North America was evaluated by examining fishery survey
data. The number of statistically valid data sets located
was remarkably low. The strongest evidence in support of
the hypothesis consists of temporal association data from
Adirondack Mountain lakes and Nova Scotia rivers. Both
data sets clearly demonstrate declines in acid-sensitive
fish species populations over the past 20 to 40 years.
Limited water chemistry data indicate that the lakes and
streams in question are currently more acidic than they
were previously, and fish population status is clearly
correlated with present pH. Waters that are now acidic
(pH < 5.0) support few or no fish populations. A
temporal association study in Massachusetts streams
provides similar results, although the number of streams
is small and the data are of lesser quality. The
LaCloche Mountain area data set is of high quality but
does not reflect effects from long-range transport.
The remaining data sets consist largely of spatial
associations of surface water chemistry and fish popula-
tions, supported in some cases by temporal association
data or field experiments. Data from Pennsylvania
streams, Vermont lakes, and Maine lakes demonstrate that
fish population status is related to present water
chemistry. Generally, waters with summer pH less than
5.0 to 5.5 support few or no fish populations. Limited
data suggest that at least some of these waters formerly
supported fish. Limited field experimental data
demonstrate that fish will not now survive in these
acidic waters and that addition of acid to surface waters
will eliminate fish populations. Although these data
demonstrate a relationship between surface water acidity
and fish population status, they cannot be used to
determine the source of the acidity.
Critics have suggested that other factors could reduce
or eliminate fish populations. These factors include
chemical pesticides use, change in fish hatchery
OCR for page 328
328
production, change in angler pressure, and increased
beaver activity Direct effects of human activities
(obstruction of fish migrations by dams, degradation of
water quality by agricultural and urban runoff, municipal
sewage, and industrial wastes, for example) greatly
reduced fish populations in accessible waters in the
colonial and post-Civil War periods.
However, remote lakes were not directly affected by
these factors, and while lumbering and subsequent burning
of the watersheds undoubtedly affected fish populations
in less accessible lakes, these factors generally were
most important in the late 1800s and early 1900s. Concern
for declining fish resources resulted in developing
artificial propagation of fish in hatcheries and led to
the founding of the American Fisheries Society to advance
knowledge concerning fish resources (Kendall 1924,
Thompson 1970).
Early attempts to supplement fish
populations by introducing hatchery-reared fish generally
failed and fell into disfavor, however. For example,
Smallwood (1918) documents the failure of hatchery
introduction in Lake Clear, New York. Lake Clear now has
a pH > 7.0 and contains at least five species of fish
(Colquhoun et al. 1984).
Beaver activity can either enhance or degrade fish
populations, depending on the particular circumstances.
Beaver were reintroduced into the Adirondack Mountains of
New York beginning in 1905 and by the 1920s had reached
population densities high enough to raise concern about
beaver damage to various resources (Johnson 1927).
Chemical pesticides have been used in remote areas of
northeastern North America for control of spruce budworm
and blackfly populations, with detrimental effects on
fish populations (Burdick et al. 1964, Anderson and
Everhart 1966, Elson 1967, Kerswill and Edwards 1967,
Locke and Havey 1972). Organochlorine compounds are
generally no longer used for these purposes, and most
affected fish populations have recovered (Dean et al.
1979). Analysis of brook trout from a series of remote
lakes of varying pH in northern New England failed to
detect significant organochlorine residues in any fish
(Haines 1983).
It is impossible to rule out factors other than
acidification as important in the decline or loss of fish
populations except by intensive, case-by-case investiga-
tions. Such investigations have seldom been made, and in
fact the quantity of data on which to base estimations of
trends in fish populations is exceedingly sparse. How-
OCR for page 329
329
ever, a few such data sets do exist, and in the cases
cited here the involvement of factors other than acid~-
fication in the decline of fish populations appears to be
negligible.
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Representative terms from entire chapter:
brook trout
