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4
Biological System of Mono Lake
INTRODUCTION
Mono Lake is a productive aquatic ecosystem but with
very few species. The lake has two major habitats--an
open water pelagic region and a nearshore littoral region.
Trophic structure, the linkages within the food web, is
different in these two habitats.
In the pelagic waters, phytoplankton are the primary
producers, using sunlight to reduce inorganic carbon to
organic matter. These algae are grazed by the brine
shrimp, Artemia monica, which are preyed upon mainly by
eared grebes (Podiceps nigricoZlis) and California gulls
(Laws californicus). No fish live in Mono Lake. The cur-
rent combination of high salinity and alkalinity makes it
impossible for fish to survive. Inputs of organic material
to the profundal sediments, those sediments under the
pelagic zone, consist largely of fecal pellets and cysts of
brine shrimp and detritus. No zoobenthos has been record-
ed in the profundal sediments, which are anoxic much or
all of the year. The role of protozoans and bacteria as
food for brine shrimp or as decomposers of organic matter
remains undetermined, but is likely to be of importance.
In the littoral region, the overlying waters have the
same planktonic organisms as the pelagic zone and an
intermittent complement of organisms associated with the
bottom. The benthic habitat is highly variable as a func-
tion of depth and substrate (Herbs", 1986; Pelagos Corpora-
tion, 1987~. The principal constituents are a microbial
69
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70
The Mono Basin Ecosystem
community, an algal flora, and a brine fly, Ephydra hians,
which feeds upon the benthic algae and probably bacteria
and detritus derived from a number of sources, some likely
to be terrestrial. The brine fly is prey to a variety of
birds including phalaropes, and to a lesser extent eared
grebes and gulls.
Although the trophic structure of Mono Lake is simple
in comparison with that in many aquatic ecosystems, the
lack of sufficient information on key components such as
bacteria and protozoans precludes the formulation of a
complete, quantitative description of carbon or nitrogen
_~ ~ ~ . Two trophic links that
have received some quantitative attention are the algae-
brine shrimp and brine shrimp-bird links. Grazing by brine
shrimp contributes to a decline in phytoplankton during the
spring and maintains a low algal abundance during the
summer (Lenz, 1982, Jellison, 1985~. The regeneration of
ammonium by the brine shrimp, in turn, sustains the
growth of the phytoplankton (Jellison and Melack, 1986~.
The decline in brine shrimp in the autumn can be, in part,
attributed to predation by the grebes (Cooper et al., 1984~.
The remainder of this chapter discusses the ecological
and physiological aspects of the components of the food
web--and primary producers and decomposers (bacteria,
phytoplankton, and phytobenthos), primary consumers (brine
shrimp and brine fly), and secondary consumers (aquatic
bird populations).
flow through the whole food web
ECOLOGICAL ASPECTS OF AQUATIC PELAGIC AND
LITTORAL ORGANISMS
Primary Producers and Decomposers
Bacteria
The abundance and significance of bacteria in alkaline,
saline lakes are not well-known. Bacteria probably func-
tion as both decomposers and primary producers in the
food web of Mono Lake.
Recent research by R. S. Oremland and his associates
indicates that the same major processes that are carried
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Biological System of Mono Lake
71
out by anaerobic bacteria in fresh water and marine habi-
tats also occur in alkaline, saline lakes. They have exam-
ined, for example, methanogenesis, sulfate reduction, and
other anaerobic processes in Big Soda Lake, Nevada (Orem-
land et al., 1982, 1985; Iversen et al., in press). In Mono
Lake, R. S. Oremland (U.S. Geological Survey, Menlo Park,
personal communication) has discovered that large quanti-
ties of methane are leaving the sediments even though
relatively small amounts of methane are produced at the
sediment-water interface. He argues that most of the
methane-rich gas seeps in the lake produce biogenic meth-
ane that is derived from the anaerobic decomposition of
fossil organic matter by bacteria. However, the methane
from one seep associated with a hot spring had a more
thermogenic character, indicating a chemical process that
does not involve bacteria. The presence of numerous gas
seeps on the floor of Mono Lake is supported by the dis-
covery that large areas of bottom sediments are disturbed
by gas bubbles (Pelagos Corporation, 1987~.
Pelagic, aerobic bacteria are often abundant in alkaline,
saline lakes. In freshwater lakes and in the ocean, con-
centrations of pelagic bacteria between 105 and 1 o6 bac-
teria/ml are commonly observed.
. . . , . ~ . . . ..
However, alkaline, saline
lakes In east At~r~ca contain from 107 to 108 bacteria/ml
(Kilham, 1981~. These large concentrations presumably rep-
resent a balance between the availability of organic sub-
strates in these highly productive lakes and the abundance
of heterotrophic organisms that consume bacteria (e.g.,
ciliates). Pyramid Lake in Nevada is the only alkaline,
saline lake in the Great Basin in which a detailed study of
bacteria has been carried out. Hamilton-Galat and Galat
(1983) found from 5.1 x 105 to 2.5 x 107 bacteria/ml in
Pyramid Lake.
Bacterial numbers more or less tracked
periods of algal production. One reason that bacterial
numbers are not higher in Pyramid Lake is that the lake is
only moderately productive (i.e., mesotrophic). For Mono
Lake, R. S. Oremland (personal communication) and R. W.
Harvey (U.S. Geological Survey, Menlo Park, personal com-
munication) have observed bacterial concentrations of
between 1.4 and 2.0 x 107 bacteria/ml. On average, these
concentrations are considerably higher than most found in
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72
The Mono Basin Ecosystem
Pyramid Lake and generally similar to those observed in
the lakes in east Africa.
Phyto plank ton and Phyto b e nthos
The algal community of Mono Lake includes few species,
as is typical of hypersaline waters. The phytoplankton is
dominated by a coccoid green alga, NannochZoris sp., cyan-
obacteria, and diatoms (Mason, 1967; Lovejoy and Dana,
1977; Melack, 1983~. The benthic algae are composed of
Nitzschia frustum, other less common diatoms, filamentous
cyanobacteria, and the green alga, Ctenocladus circinnatus
(Herbs", 1986~.
The seasonal dynamics of the phytoplankton in Mono
Lake are unusual (Mason, 1967; Lovejoy and Dana, 1977;
Melack, 1983, 1985; Jellison and Melack, in press) (Figure
4. 1~. During the winter, the phytoplankton are abundant
throughout the lake, and after the onset of the seasonal
thermocline in early spring, the algae increase in the upper
water. This increase was reduced during 1984, 1985, and
1986 after the initiation of meromixis. As described in
chapter 3, the chemical stratification reduced vertical mix-
~ -- ~ ~ ~ ~~ ~~ nutrient,
nitrogen, to the euphotic zone. A rapid decrease in algal
abundance occurs in late May and June above the thermo-
cline. During the summer, the phytoplankton are sparse in
the upper waters and abundant in the deeper, cold and dim
or dark waters. In midsummer, higher chlorophyll concen-
trations occur in a layer coinciding with the chemocline.
In autumn, algal concentrations increase in the upper
waters as thermal stratification weakens and brine shrimr,
sing, wn~cn reoucecl the supply of the limiting
numbers decline (Figure 4.1~.
Primary productivity measurements spanning the period
from 1983 to 1985 vary from 340 to 540 g carbon/m2/yr
(Jellison and Melack, in press). Mono Lake would thus be
classified as eutrophic. Production was higher during the
spring of 1983 than in 1984 and 1985; the difference may
be at least partially attributed to meromixis.
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Biological System of Mono Lake
60~
-
CL
O 30
o
I
O
82 /~`
86
73
, i,
. ,
i, I
`, ,
I
1
1
1
_ 'v' WN I~ · ·-
I I~ at/ 1
J F M A M J J A S O N D
TIME (months)
FIGURE 4.1 Mean mixolimnetic chlorophyll a for 1982,
1983, 1984, 1985, and 1986.
Primary Consumers
Zooplankton
The Mono Lake brine shrimp, Artemia monica, is the
major zooplankton species (Mason, 1967; Lenz, 1980, 1982~.
A. monica, a member of the A. franciscana superspecies, is
now considered a sibling species (Bowen et al., 1985~. The
zooplankton also includes protozoans and has included roti-
fers (Mason, 1967~.
The abundance of brine shrimp in Mono Lake varies
seasonally (Lenz, 1982, 1984; Figure 4.2~. The brine shrimp
hatch from overwintering cysts from January through May.
By mid-May, the first adult brine shrimp are present. For
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74
The Mono Basin Ecosystem
50
45
40
35 - Standard Enora
55 ~ Naupill ~ ~ ~ I I I~Adults
O~ in
J F M A M J J A S O N
MONTH
. ,
D
FIGURE 4.2 Seasonal abundance of brine shrimp at Mono
Lake in 1985. Lakewide mean of 10 stations (three vertical
net tows per station).
approximately one month females bear live young, which
mature rapidly in the warm upper mixed layer. In June
females switch to oviparous reproduction. The diapause
eggs lie dormant on the bottom of the lake until the fol-
lowing winter. During the summer, brine shrimp are abun-
dant in the oxygenated upper waters and very sparse or
absent in the anoxic deeper waters. By September, the
brine shrimp begin to decline in numbers and are almost
absent from the plankton by December.
Studies conducted with similar methods since 1978 per-
mit interannual comparisons of brine shrimp abundances and
reproductive characteristics (Lenz, 1984; G. Dana, R.
Jellison, and J. M. Melack, University of California, Santa
Barbara, unpublished). Statistically significant interannual
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Biological System of Mono Lake
75
differences in abundances of first-generation adults (late
May to June populations) occurred. In 1979, 1984, and
1986 brine shrimp numbered between 19,000 and 31,000 ani-
mals/m2. whereas from 1980 to 1983 numbers were only
. . ~ · · . .
2,400 to 5,700 an~ma~s/m~.
dances (first- and second-generation adults) were much
higher in 1981 and 1982 than in other years. A number of
factors are associated with these variations. First-
generation adult abundances denend on the number of cysts
Maximum midsummer aoun-
. —
available for hatching, hatching success, and survival to
adulthood. In laboratory experiments, Dana and Lenz
(1986) determined that salinities in the period from 1979 to
1986 are not indicated as a cause for changes in hatching
success. Emergence trap trials in spring 1985 showed that
~ ~ . ~ . ~ . ~ . .
~7 ~ J
very low hatching occurred In sediments In anoxlc water
below the chemocline (Dana et al., in press). In contrast,
large numbers of cysts lying in sediments under the oxy-
genated mixolimnion hatched. The number of cysts
available depends on the production of cysts during the
previous year and possibly past years and on the viability
of the cysts. Cyst production is related to brood size,
numbers of ovigerous females, the percentage of those
females producing cysts, and the time interval between
broods. Brood size varied from 30 to 140 eggs per brood
from 1983 to 1986 and is explained primarily by differences
in female length and algal abundance. Second-generation
abundances depend on the abundance, percent ovoviviparity,
and fecundity of the first-generation females. Recruitment
to adults depends on survival of naupliar and juvenile
stages. Differences in all these factors occurred from 1982
to 1986. The switch from ovoviviparity (live bearing) to
oviparity (cyst production) occurred at the time of dec-
reasing phytoplankton in all years studied. In years with a
substantial spring hatch, the first generation dominates the
population. When the spring hatch is relatively low, first-
generation adults are less abundant, algal densities remain
higher later into the spring, and a large second generation
can occur.
The spatial distribution of brine shrimp is heterogeneous
on large (square kilometers) and small (square meters) spa-
tial scales and varies on time scales from hours to days to
seasons (Lenz, 1982; Melack, 1985; Lenz et al., 1986; Conte
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76
The Mono Basin Ecosystem
et al., in press). These differences in concentrations of
brine shrimp result in variable profitability of foraging for
Small-
spr~ngs
where upwelling varies widely in strength and hence in the
entrainment of brine shrimp. Mason ( 1966) hypothesized
that the very dense plumes that formed near shore, but not
in association with springs, result from thermal currents
and behavioral responses of the animals. Foam lines con-
tain concentrations of living ant! dead brine shrimp as well
as other debris and can stretch for hundreds of meters.
These features seem to delimit water masses. Large-scale
patchiness has been documented by sampling transects and
lakewide grids. Abundance differences between the eastern
and western halves of the lake are common. The degree of
variability differs seasonally and appears greater during
transition periods such as spring and autumn.
the birds (see section on bird populations below).
scale patches are associated with sublacustrine
. . .
· ~ .
-
- .,
Current sampling programs are designed to assess the
lakewide abundance of brine shrimp and include biweekly or
monthly samples from 10 pelagic stations. Regular sampling
is not performed in water overlying the littoral region or
at sites of aggregation such as springs. Therefore, while
providing statistically sound estimates of the overall abun-
dance of brine shrimp, the sampling does not include sites
that may be of particular importance to some birds some of
the time. No efforts are in progress to sample zooplankton
other than brine shrimp.
Zoobenthos
The benthic community of Mono Lake includes several
species of dipteran insects, as is typical of hypersaline
waters. The predominant dipteran is the brine fly, Ephydra
hians, but other species are present, such as the deer fly
(Chrysops spy and the long-legged fly (Hydrophorus plum-
beus). The biting midge (Culicoides occidentalis) is also
found among the macroinvertebrates (Herbs", 1986~.
The seasonal dynamics of the macroinvertebrates are not
well-known. However, recent research on brine flies by
Herbst (1986) using the third instar and pupae as popula-
tion~ indices showed a phase of rapid population growth
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Biological System of Mono Lake
77
occurring in the spring (May and June), a summer maximum
(July through September), a gradual decline in the autumn,
_ _ : ~1 ~ ~ late winter through early
spring. Since seasonal dynamics of the phytobenthic bac-
terial and algal populations (diatoms and filamentous algae)
are unknown, one cannot determine if the zoobenthic com-
munity, as reflected by numbers of brine flies, is tracking
periods of algal production.
The spatial distribution of brine flies is heterogeneous
on large (square kilometers) and small
scales (Herbs", in press).
. ~ · ~
and minimal abundance ~ rom
, (square meters)
Small-scale patches are associ-
ated with tufa pinnacles and nearshore grasses, which are
excellent substrates for larval and pupal attachment.
Large-scale patchiness has recently been documented by
video and lakewide bathymetric transects (Pelagos Corpora-
tion, 1987~. Large mats of pupae have been found on dead
submerged grasses along the eastern shore and on under-
water tufa and hard-surface sediments at depths sometimes
greater than 10 m in the central and eastern provinces, as
shown in Figure 4.3. Abundance differences observed be-
tween tufa-hard rock shoal regions and soft mud-sand lake
bottom sediments common to the eastern and central prov-
inces are probably due to the larvae's inability to attach to
smooth surfaces. The placement of eggs by brine fly
females at depths greater than a few meters has been ob-
served (Pelagos Corporation, 1987~. This observation raises
questions about the typical mechanism of oviposition re-
ported for other ephydran flies, including whether females
utilize respiratory mechanisms other than gas bubble
entrapment for vertical descent. If the lake level dropped,
the loss of hard-surface sediments would reduce brine fly
habitat.
PHYSIOLOGICAL ASPECTS AND SALINITY TOLERANCES
OF AQUATIC PELAGIC AND LITTORAL ORGANISMS
Primary Producers
Two kinds of evidence are available to evaluate the
effects of increased salinity on phytoplankton: (1) algal
responses to experimental increases in the salinity of Mono
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Representative terms from entire chapter:
brine shrimp
78
The Mono Basin Ecosystem
/~
EphydrahiansMats
' 1 Paoha _\ z / \
\` Island I /
\~ \\ ~/O~,N
Biological System of Mono Lake
79
arsenic, and fluoride contribute to these reductions in
growth. Overall, Chapman's results indicate that a salinity
of about 175 g/1 results in fairly large decreases in growth
of the two currently dominant algae.
A number of species of phytoplankton, not currently
extant or dominant in Mono Lake, are known to grow well
at salinities reaching 200 g/1 (Hammer, 1986; Melack, in
press). In particular, a green alga, DunalieZ/a sp., now
occurs in low numbers in Mono Lake, and a related species,
D. parka, grew best in Mono Lake concentrated to 150 g/1
. · .
and still grew slowly at 235 g/1 (Chapman, 1982).
Herbst (1986) isolated a clone of Ctenocladus circinnatus
from Mono Lake and determined its growth rate and yield
at salinities of 25, 50, 75, 100, and 150 g/1. The solutions
were obtained by dilution or low-temperature evaporation
of Mono Lake water. The experiments indicated decreased
growth and yield at 75 and 100 g/1 and no growth at 150
g/l.
Primary Consumers
· —
A fundamental requirement for aquatic organisms living
in saline lakes is to have sufficient "free water," or water
that is available to sustain vital cellular activities. (In
aquatic organisms, water molecules forming hydration shells
are termed "bound water"; water molecules not associated
with these shells are termed "free water.") This biological
axiom is most evident in the osmotic effects of salinity
upon the growth and larval development of brine shrimp
and brine flies. If the salinity of the medium in which the
organisms reside is sufficiently high, the thermodynamic
forces responsible for the osmotic gradient will not allow
sufficient free water to remain inside the organism. The
loss of free water will in turn cause an inhibition or ces-
sation of metabolic processes. This physical relationship
sets an absolute upper limit on the salinity of salt lakes in
which a self-sustaining population of halophilic organisms
such as brine shrimp and brine flies can persist.
Physiological solutions within and surrounding cells are
primarily dilute aqueous solutions. These solutions contain
a large amount of free water and behave in a manner
110
The Mono Basin Ecosystem
major refuse dump within 15 km of the colonies and exten-
sive agricultural fields nearby, Mono Lake has only a few
small refuse dumps nearby and has no nearby agricultural
activity that would sustain a large gull population twinkler,
1 983b). There is a lack of information on the types and
availability of prey taken by gulls early in the breeding
season, and additional sampling is required to determine the
importance of foods other than brine shrimp throughout the
season.
Summarizing the information on birds, large numbers of
eared grebes, red-necked phalaropes, Wilson's phalaropes,
and California gulls use the lake and depend on the brine
shrimp and brine flies for their food. Additionally, the
gulls require safe refuges on which to nest. Present cen-
sus techniques are statistically inadequate to detect small
but possibly biologically significant changes in avian num-
bers or temporal patterns of use of Mono Lake. Likewise,
we do not know critical food densities below which the
invertebrates taken by birds are not sufficient to provide
needed body weight, or how the density of available prey
will change with overall prey densities. These data are
needed if we are to quantitatively predict how bird popula-
tions will respond to changing food supplies with changing
lake salinity. Clearly, however, if the invertebrate popula-
tions are drastically reduced, there will be reductions in
the number of birds visiting and using Mono Lake.
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