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O R I G I N A L P A P E R

Competitive exclusion of Cyanobacterial species
in the Great Salt Lake

Hillary C. Roney Æ Gary M. Booth Æ
Paul Alan Cox

Received: 25 July 2008 / Accepted: 16 December 2008 / Published online: 8 January 2009

� Springer 2009

Abstract The Great Salt Lake is separated into different

salinity regimes by rail and vehicular causeways. Cyano-

bacterial distributions map salinity, with Aphanothece

halophytica proliferating in the highly saline northern arm

(27% saline), while Nodularia spumigena occurs in the less

saline south (6–10%). We sought to test if cyanobacterial

species abundant in the north are competitively excluded

from the south, and if southern species are excluded by the

high salinity of the north. Autoclaved samples from the

north and south sides of each causeway were inoculated

with water from each area. Aphanothece, Oscillatoria,

Phormidium, and Nodularia were identified in the culture

flasks using comparative differential interference contrast,

fluorescence, and scanning electron microscopy. Aphanot-

hece halophytica occurred in all inocula, but is suppressed

in the presence of Nodularia spumigena. N. spumigena was

found only in inocula from the less saline waters in the

south, and apparently cannot survive the extremely

hypersaline waters of the northern arm. These data suggest

that both biotic and abiotic factors influence cyanobacterial

distributions in the Great Salt Lake.

Keywords Competitive exclusion � Halophilic bacteria �
Aphanothece � Nodularia � Oscillatoria � Phormidium �
Gause’s principle

Introduction

Cyanobacteria are well-adapted for living in harsh condi-

tions, including photosynthetic areas beneath Antarctic ice,

hot springs and geysers in Yellowstone, and hypersaline

lakes, including the Great Salt Lake (Dyer 2003). Two

common cyanobacteria species that have been identified

from the Great Salt Lake are Aphanothece halophytica and

Nodularia spumigena (Brock 1976; Felix 1978; Felix and

Rushforth 1980), with N. spumigena episodically blooming

in Farmington Bay (Marcarelli et al. 2006). Since nitrogen

is the limiting nutrient in the Great Salt Lake (Oren 2002) it

is interesting to note that both A. halophytica and

N. spumigena can fix nitrogen.

The Great Salt Lake is a hypersaline remnant of the

Pleistocene Lake Bonneville (Oren 2002) which was

557 km long and 233 km wide with an area of 51,800 km
2

in what is now Utah, Idaho, and Nevada (Utah Geological

Survey 1990). As Lake Bonneville retreated, the lake lost

all outlets, so salinity increased.

After completion in the mid 19th century of the

transcontinental railway near Promontory Point, Utah,

trains had to traverse many additional rail kilometers

around the northern end of the Great Salt Lake. To reduce

this distance, the Union Pacific Railroad constructed a

19 km rail causeway across the Great Salt Lake in 1959

(Fig. 1), replacing an earlier trestle built in 1902 by the

Southern Pacific Railroad. Unlike the former wooden

trestle, which did not impact water flow, the 1959

causeway built with rock fill, hydrologically divided the

Communicated by L. Huang.

H. C. Roney � P. A. Cox (&)
Institute for Ethnomedicine, Box 3464,

Jackson Hole, WY 83001, USA

e-mail: [email protected]

G. M. Booth

Department of Plant and Wildlife Science,

Brigham Young University, Provo, UT 84602, USA

123

Extremophiles (2009) 13:355–361

DOI 10.1007/s00792-008-0223-1

waters of the Great Salt Lake into two portions, a northern

arm with negligible freshwater inputs, and a southern arm

with more than 90% of the freshwater flow (Butts 1980;

Oren 2000; Stephens and Gillespie 1976; Sturm 1980).

Three major rivers–the Bear, Weber, and Jordan–all flow

into the Great Salt Lake south of the railway causeway

(Gwynn 2002). The overall salinity of the Great Salt

Lake, which to that point had been linked solely to

changing water levels, quickly adjusted with the northern

arm becoming even more hypersaline, moving from an

average of 15% in the 1870s to as high as 28% salinity in

the 1960s (Sturm 1980). In 1970, the northern arm held

approximately 330–350 g salts per liter, while the south

arm held 120–130 g salts per liter (Oren 2002). While the

major cation in the water is Na, Mg, K, Ca, in decreasing

order of abundance are important as is the anion SO4
(Sturm 1980).

These habitat changes were later partially replicated

with construction of a second barrier to lake water flow.

A causeway for vehicular use has been periodically

constructed from Syracuse, Utah to Antelope Island, and

was most recently rebuilt in 1992 (Gwynn 2002). These

vehicular and railway causeways resulted in the parti-

tioning of the Great Salt Lake into three different salinity

regimes: the northern arm, with an average of 27%

salinity, the middle arm with average salinity of 10–16%,

and the southern arm, with average salinity of 6% or less

(Utah Geological Survey 1990). These three different

salinity regimes, any one of which would be considered

hypersaline, allowed species to sort according to eco-

logical tolerances. The results are striking: each large

area of the lake has different colored water, resulting in

part from different concentrations of Artemia fransiscana

brine shrimp cysts and microscopic green algae such as

Dunaliella salina and D. viridis as well as species of the

Archean genus Halobacterium (Post 1981), but also

perhaps due to different concentrations and species

compositions of cyanobacteria.

A study was designed to determine whether cyanobac-

terial distributions in the Great Salt are influenced by

abiotic factors, biotic factors, or both. To explore this

question, experiments were designed to examine two

hypotheses; Hypothesis 1: Cyanobacterial species abundant

north of the railway causeway are competitively excluded

from the south by other species, and Hypothesis 2:

Cyanobacterial species that thrive and bloom south of the

Antelope causeway cannot grow in high salinity waters

from the north.

Materials and methods

Experimental cultures

A total of 28 water samples from both the north and south

sides of Antelope Island causeway and the north and south

sides of the railway causeway (seven from each site) were

collected in December 2007. Water temperatures and GPS

coordinates were recorded at each site. To avoid pseu-

doreplication, six water samples from both sides of the

vehicular and railway causeways, approximately four liters

in volume, were used as inocula. The seventh jar from each

side was approximately eight liters in volume and used for

media after filtering and autoclaving. Approximately 30 ml

of the filtered media water was placed in autoclavable,

sterile 50 ml nalgene plastic flasks. In total there were 96

flasks inoculated with 10 ml of unsterilized water from

either the north or the south of the railway and the vehic-

ular causeways totaling six replicates of inoculum water for

each medium type. No nutrients or other growth media

were added to the water (Dyer 2003). In addition, 21

control flasks were prepared using autoclaved media and

autoclaved inocula. A random number table was used to

decide from which sample jars to draw the inoculum. In

addition, one control flask was prepared which consisted of

autoclaved distilled water inoculated into autoclaved dis-

tilled water medium. After all 129 flasks were inoculated;

they were placed in a heated green house with constant

8.5 h/day illumination. Each flask was gently shaken by

hand periodically for aeration. These liquid cultures were

incubated for 7 weeks.

Cyanobacterial identification

For aquatic cyanobacteria, identification by light micros-

copy (phase and/or interference contrast), and scanning

electron microscopy (SEM) are preferred (Cronberg and

Fig. 1 Great Salt Lake Rail Causeway with hypersaline water in the
north (left) and less saline water on the south (right)

356 Extremophiles (2009) 13:355–361

123

Annadotter 2006). Identification and abundance counts of

cyanobacteria from the culture flasks were performed using

differential interference contrast (DIC) and epi-fluores-

cence imaging, with SEM for verification.

Data analysis

To ensure arbitrariness of cyanobacterial counts, micro-

transects of water cultured from each flask, based on two

microscope slides were conducted. Each microtransect

was replicated twice for each slide, with data entered on a

six-cell mechanical lab counter. When mass colonies of

cyanobacteria where encountered precluding individual

counts, the colony were assessed as ‘‘large’’ or ‘‘very

large’’, with medians and nonparametric analyses used to

analyze qualitative data. In each of the 16 possible

combinations of 4 types of media (railway north, railway

south, Antelope north, and Antelope south) and 4 types of

inocula (railway north, railway south, Antelope north, and

Antelope south), the median counts of the 4 major

cyanobacterial species (A. halophytica, Oscillatoria sp.,

Phormidium tenue, and N. spumigena) were ranked.

A two-way Analysis of Variance (ANOVA) was cal-

culated for abundance of A. halophytica in the culture

flasks with ‘‘large’’ colonies scored as 500 and ‘‘very

large’’ colonies scored as 1,000 for this purpose. To reduce

impact about outliers and ensure consistency of distribution

across the observed range, all data were transformed with a

square root transformation prior to analysis as is standard

for count data. F statistics for the transformed data were

calculated to test three different pairs of hypotheses, with

the null hypothesis to be rejected at the P 0.05 level:

Hypothesis pair #1

H0: no variation in cyanobacterial counts exists due to

differences in media.

H1: variation in cyanobacterial counts exists due to

differences in media.

Hypothesis pair #2

H0: no variation in cyanobacterial counts exists due to

differences in inocula.

H1: variation in cyanobacterial counts exists due to

differences in inocula.

Hypothesis pair #3

H0: no variation in cyanobacterial counts exists due to

interactions.

H1: variation in cyanobacterial counts exists due to

media and inocula interactions.

For cyanobacterial taxa which proved to be of rare

occurrence in the culture flasks, exact logistic tests, rather

than an ANOVA, were calculated.

Results

Experimental cultures

When sampling for the experimental cultures of the Great

Salt Lake, profoundly different colors on either side of the

railway causeway were observed from the air (Fig. 1).

These differences were also apparent in water samples

taken from deep water on either side of the causeway rather

than evaporative ponds (Fig. 2). Salinity from the sample

sites was previously measured by hydrometers—south side

of Antelope causeway 20.6 ppt, north side of Antelope

causeway 75 ppt, south side of railway causeway 155 ppt,

and north side of railway causeway 195 ppt (Roney 2007).

Salinity values of the flasks were altered slightly by addi-

tion of inocula, except when the same inoculum was added

to the same medium.

At the time of sampling in December 2007, ambient air

temperature was -2.2�C on the railway causeway
(4181301600N1128320303400W) and -2.8�C on the vehicular
causeway (418404400N11281205700W), with water tempera-
tures north of the railway causeway at 2.0�C, south of the
railway causeway at 4.3�C. Water temperature north of the
vehicular causeway was 3.3�C, while the water tempera-
ture south of the causeway was 2.5�C. Analysis by light
microscopy showed no growth in any of the 21 control

flasks, which were found to be sterile.

Cyanobacterial identification

Four genera of cyanobacteria, Aphanothece, Oscillatoria,

Phormidium and Nodularia, were identified (Figs. 3, 4, 5,

6), with identifications confirmed by Dr. James Metcalf

Fig. 2 Water samples taken on north (left) and south (right) side of
railway causeway. Color differences primarily due to Dunaliella
distributions although the cyanobacterium Aphanothece flourishes in
the hypersaline waters in the north

Extremophiles (2009) 13:355–361 357

123

(University of Dundee, Scotland). In addition, a fifth

cyanobacterial genus, Spirulina, was observed, but was not

found in any transect through any of the microscope slides.

Because of its trichomes and its affinity for saline waters,

this species is referable to Spirulina labyrinthiformis

(Fig. 7), although Nübel et al. (2000) have placed a similar

salt-tolerant species into the new genus, Halospirulina.

Comparisons between differential interference contrast

microscopy, fluorescence microscopy, and scanning elec-

tron microscopy allowed different observations of

cyanobacterial morphology and size to be compared for

taxonomic identification.

Data analysis

Comparative medians of the four cyanobacterial genera for

each inocula type in the four media are shown in Fig. 8,

which demonstrates that A. halophytica appears throughout

all the four types of media and inocula, but that Nodularia

spumigena is abundant only in inocula from Antelope south

waters. Since there are 24 different permutations of the

ordered ranks of Aphanothece (A), Oscillatoria (O), Nod-

ularia (N), and Phormidium (P) plus an additional 16

permutations of three and two-way ties, as well as one

Fig. 3 Aphanothece halophytica: differential interference contrast
(left); fluorescence (middle); scanning electron microscopy (right)

Fig. 4 Oscillatoria sp.: differential interference contrast (left); fluo-
rescence (middle); scanning electron microscopy (right)

Fig. 5 Phormidium tenue: differential interference contrast (left);
fluorescence (middle); scanning electron microscopy (right)

Fig. 6 Nodularia spumigena: differential interference contrast (left);
fluorescence (middle); scanning electron microscopy (right)

Fig. 7 Spirulina cf. labyrinthiformis: differential interference con-
trast (left); fluorescence (middle); scanning electron microscopy
(right)

358 Extremophiles (2009) 13:355–361

123

possible case of a four-way tie in rank, there are 41 dif-

ferent possible rankings of the four cyanobacterial species.

These different rankings can perhaps most easily be por-

trayed as different colors (Fig. 8). A. halophytica was

dominant in all cultures flasks, except those in which N.

spumigena and Oscillatoria sp. occurred.

A two-way ANOVA for the distributions of A. halo-

phytica was performed as indicated in Table 1. The F

statistics for the ANOVA allows each of the null hypoth-

eses in the three pairs of hypotheses to be rejected at the

P 0.05 level. Therefore, it can be concluded that media
and inocula, as well as the interaction between media and

inocula significantly affected the growth of A. halophytica

in the culture flasks. Exact tests were calculated for

N. spumigena and Oscillatoria spp. using the exact option

of pro logistic in SAS. In these analyses, counts were

ignored, and instead, presence/absence data were used. For

Oscillatoria, the P value for the exact test of the medium

was 0.0993; thus the effect due to media differences was

not significant. However, the P value for the exact test of

inoculum was 0.0016; hence there was a significant inoc-

ulum effect in distribution of Oscillatoria. The exact test

for the inocula/media interactions was not significant with

a P value of 0.1963. Leaving interaction out of the model,

the additive model (with additive effects of inoculum and

medium) showed the odds of a positive response for

inocula from rail south, rail north, or Antelope north was

just 6.3% relative to inoculum from Antelope south (95%

confidence interval: 0.6–64.2%). Thus, Antelope south

inocula had a significant positive effect on the presence of

Oscillatoria in the culture flasks.

A similar analysis was conducted for the presence or

absence of N. spumigena in the culture flasks. The exact

test for the inocula/media interaction had a P value of

0.0032; hence the interaction was significant. The odds of a

positive effect were extremely high for the combination of

Antelope south inoculum with Antelope south medium. For

all other combinations, the odds of a positive response were

extremely low. For future studies of algal-cyanobacterial

interactions, counts were also made of the green alga

Dunaliella salina and D. virids in the flasks; an ANOVA of

square root transformed data count for Dunaliella showed

significant differences in distributions similar to Aphanothece

distributions; these data will be reported elsewhere.

Discussion

Both the ranking of median abundances in the culture

flasks and the results of the two-way ANOVA support the

overall hypothesis: cyanobacterial species abundant north

of the railway causeway (e.g. A. halophytica) are com-

petitively excluded from the south by other species, in this

case N. spumigena and Oscillatoria spp. It appears that the

cyanobacterium A. halophytica can grow in less saline

waters as well as the extreme saline waters north of the

railway causeway—since it is found in all inocula—but its

growth appears to be suppressed in the south by the pres-

ence of N. spumigena, which periodically blooms in the

Great Salt Lake.

In previous years, we have noted large N. spumigena

blooms in the low salinity regime of Farmington Bay, as well

as in water samples collected south of the railway causeway

(Roney 2007), particularly when winds have concentrated

blooms near the causeway. Rushforth and Felix (1982)

recorded N. spumigena as rare in the south arm; perhaps they

took their samples at a dormant season, as N. spumigena

blooms episodically. The absence of N. spumigena in the

southern arm of the Great Sale Lake may have influenced

the ability of A. halophytica to migrate and prosper in the

fresher water environment of the south arm instead of

thriving in the hyper-saline north arm. In all of our samples

south of the Antelope causeway (Farmington) since 2004,

N. spumigena was present in the water column.

The second overall hypothesis—that cyanobacterial

species that thrive and bloom south of the Antelope

Fig. 8 Rankings of cyanobacterial dominance by medians. Media/
Inocula in the upper left corner of the chart are extremely hypersaline,
while those in the lower right corner are far less saline

Table 1 ANOVA of Aphanothece halophytica distributions in
autoclaved media from the Great Salt Lake

Source Variation Degrees

free

Mean

square

F
Statistic

Significance

Media 64.6 3.0 21.5 11.4 P 0.01
Inocula 26.1 3.0 8.7 4.6 P 0.01
Interaction 105.6 9.0 11.7 6.2 P 0.01
Subtotal 196.3 15.0

Error 151.1 80.0 1.9

Total 347.4 95.0

Extremophiles (2009) 13:355–361 359

123

causeway cannot grow in the high salinity of the north—is

also supported by these experimental data. N. spumigena

was found only in inocula from the less saline waters south of

the Antelope Island causeway, and apparently cannot sur-

vive the high saline waters north of the railway causeway.

Experimental support for these two general hypotheses

helps shed light on our original question: are cyanobacte-

rial distributions in the Great Salt Lake influenced by

abiotic factors, biotic factors, or both? From these experi-

ments, it appears that both abiotic (salinity) and biotic

(interspecies competition) factors seem to affect distribu-

tions of cyanobacterial species. N. spumigena distributions

seem to be primarily influenced by salinity, since it can

only grow in fresher waters. By contrast, A. halophytica

distributions seem to be primarily influenced by competi-

tion from N. spumigena and Oscillatoria sp. There are, of

course, other geochemical processes which we did not

measure but which may affect distributions. We are also

interested in the relationship between the green alga

Dunaliella and cyanobacteria. Our initial analysis suggests

a commensalism with Dunaliella benefitting from the

presence of nitrogen fixing Apanothece: in our microscopic

analysis we often observed Dunaliella cells clustered

around mass colonies of Apanothece. It would be inter-

esting if nitrogen fixed by Apanothece in hypersaline

environments contributed to exceptional salt tolerance of

Dunaliella (Zamir et al. 2004).

These experimental results are consistent with Gause’s

principle, which predicts that no two species can

indefinitely occupy the same niche (Gause 1969; Hardin

1960), since there is a clear niche partitioning between

A. halophytica and N. spumigena in the Great Salt Lake.

These two species cannot occupy the same hypersaline

habitat north of the railway causeway, since N. spumigena

cannot tolerate hypersaline conditions, and A. halophytica

is suppressed in the presence of N. spumigena in the less

saline southern waters.

However, this leaves unanswered the question of why

A. halophytica is not totally excluded from the south, since it

occurs in all samples of inocula, regardless of salinity.

Perhaps A. halophytica is periodically excluded from

southern waters by N. spumigena blooms, but during

intervals between blooms, the extremely small A. halophytica

persists, albeit at lower levels. Thus, Gause’s Principle

should perhaps include a clarification: two species cannot

indefinitely occupy the same niche, except when that niche

is temporally partitioned, as occurs with episodic blooms of

Nodularia. This structuring of the cyanobacterial regimes by

salinity (Williams 1998) is consistent with the intermediate

salinity hypothesis of David Herbst (1999): ‘‘Abundance of

salt-tolerant organisms is limited by physiological stress at

high salinities, and by ecological factors, such as predation

and competition, in more diverse communities at low

salinities’’. Since N. spumigena distributions cannot survive

the high salinity stress of waters from the north arm of the

Great Salt Lake, and Aphanothece halophytica is competi-

tively excluded by N. spumigena at lower salinities, the

intermediate salinity hypothesis may apply.

The precise set of conditions that trigger episodic

N. spumigena blooms is unknown. Our data suggest that

these episodic blooms play a major role in excluding

A. halophytica from vast areas of the Great Salt Lake.

Being able to predict the occurrence of Nodularia blooms

would not only be of theoretical importance; it might also

lead to a better understanding of cyanobacterial blooms and

cyanotoxin impacts on wildlife and human health (Cox

et al. 2005; Metcalf et al. 2008).

Acknowledgments We thank J. Metcalf for assistance in cyano-
bacterial identification, J. Gardner for assistance in scanning electron

microscopy, and B. Schaalje for assistance with biostatistics. We are

grateful to the Wood Family Foundation for the Mus Views DIC/

Fluorescent Microscopy Facility at the Institute for Ethnomedicine,

and A. Fransiscana and R. Smithson for inspiration in our studies of

the Great Salt Lake.

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Extremophiles (2009) 13:355–361 361

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  • Competitive exclusion of Cyanobacterial species�in the Great Salt Lake
    • Abstract
    • Introduction
    • Materials and methods
      • Experimental cultures
      • Cyanobacterial identification
      • Data analysis
    • Results
      • Experimental cultures
      • Cyanobacterial identification
      • Data analysis
    • Discussion
    • Acknowledgments
    • References

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/ColorImageDownsampleType /Bicubic
/ColorImageResolution 150
/ColorImageDepth -1
/ColorImageDownsampleThreshold 1.50000
/EncodeColorImages true
/ColorImageFilter /DCTEncode
/AutoFilterColorImages false
/ColorImageAutoFilterStrategy /JPEG
/ColorACSImageDict <<
/QFactor 0.76
/HSamples [2 1 1 2] /VSamples [2 1 1 2]
>>
/ColorImageDict <<
/QFactor 0.76
/HSamples [2 1 1 2] /VSamples [2 1 1 2]
>>
/JPEG2000ColorACSImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/JPEG2000ColorImageDict <<
/TileWidth 256
/TileHeight 256
/Quality 30
>>
/AntiAliasGrayImages false
/DownsampleGrayImages true
/GrayImageDownsampleType /Bicubic
/GrayImageResolution 150
/GrayImageDepth -1
/GrayImageDownsampleThreshold 1.50000
/EncodeGrayImages true
/GrayImageFilter /DCTEncode

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