AN ABSTRACT OF THE THESIS OF
Raymond Zenon _Riznyk
(Name)
in
Title:
for the
Doctor of Philosophy
(Degree)
presented on
Botany (Phycology)
(Major)
ci(Date)
/767
ECOLOGY OF BENTHIC MICROALGAE OF ESTUARINE
INTERTIDAL SEDIMENTS
Abstract approved:
Redacted for Privacy
The benthic microalgae of sediments of the two tidal flats in
Yaquina Bay, Oregon were investigated to determine the environmental
factors limiting the abundance and the hdrizonta1:ansi vertical distribution of these organis'ins. The Southbeachtidal Tiat'which is under the
marine realm of deposition consists of fine to medium grained sand.
The Sally's Bend tidal flat is under the fluviatild realm of
deposition and consists of silt.
Measurements were made of interstitial temperatures,
movements of sand, turbidity, pH, salinity, depth of light penetration through the sediments, and the water content of the substrate.
Samples of the benthic microalgal community were collected by
using a piston corer. Sections of the cores were used for estimating
the biomass: (1) by making direct counts of live microalgae, (2) by
estimating chlorophyll a concentration and (3) by measuring ash-free
dry weight.
The greatest biomass of microalgae was found to be in
cores from the lower intertidal zone while cores from the upper inter­
tidal zone had the lowest biomass. This distribution probably results
from the greater fluctuations in temperature, salinity, water content,
and oxygen content, which are more variable in the upper intertidal
zone. The greater biomass in cores from the lower intertidal zone
may be the result of less fluctuation in environmental factors as well
as the fact that this area is exposed to nutrient-laden water for
longer periods of time than the upper intertidal zone. The greatest
biomass of microalgae was found in the upper centimeter of cores col­
lected at all levels of the intertidal zone, because light can penetrate
no more than a few millimeters through sediment. Occurrence of
algae below the photic zone is thought to result from vertical migra­
tion, sedimentation, or the activity of burrowing animals.
It was found that the Southbeach tidal flat had a significantly
greater biomass than Sally's Bend at all intertidal levels and in the
various layers of the cores. This was attributed to differences in
environmental conditions peculiar to each tidal flat which is the
result of the hydrography of the bay.
Estimates of the rates of potential gross production were made
using a Gilson Differential Respirometer. The community from the
Southbeach tidal flat had a greater potential gross rate of production
than the Sally's Bend community. This was partially the result of
high rates of bacterial respiration in cores from the Sally's Bend
tidal flat. This tidal flat had significantly greater amounts of organic
matter than Southbeach and the abundance of bacteria in sediment is
related to the amount of organic matter.
Measurements of the concentrations of chlorophyll a were
corrected for the percentage of pheophytin present. Significantly
greater amounts of pheophytin were found in cores from the Sally's
Bend tidal flat which probably resulted from the large amounts of
allochthonous detrital chlorophyll deposited in these sediments.
The microflora consisted almost exclusively of diatoms. One
hundred and fifty-four species and varieties were identified. Most of
the species found in the lower intertidal zone were found in the mid
and upper intertidal zones as well. Many of the species identified
have never been reported from Oregon prior to this investigation.
Ecology of Benthic Microalgae of
Estuarine Intertidal Sediments
by
Raymond Zenon Riznyk
A THESIS
submitted to
Oregon State University
in partial fulfillment of
the requirements for the
degree of
Doctor of Philosophy
June 19 69
APPROVED:
Redacted for Privacy
Pro
otany
in charge of majo
Redacted for Privacy
Chatiziman of the Dep j ment of Botany and Plant Pathology
Redacted for Privacy
Dean of Graduate School
Date thesis is presented
Typed by Mary Jo Stratton for
Raymond Zenon Riznyk
ACKNOWLEDGEMENTS
I would like to extend my gratitude to my major professor, Dr.
Harry K. Phinney, for his helpful advice and encouragement and also
for his editorial efforts in the preparation of this thesis.
I would also like to thank Dr. C. David McIntire for his
assistance in diatom systematics and for the verification of many of
the diatom species identified.
Finally, I would like to express my sincere appreciation to my
wife, Natalie, who assisted with the field, the lab, and the photo­
graphic work and to whom I shall always be indebted.
This investigation was supported by a National Aeronautics and
Space Administration pre-doctoral traineeship.
TABLE OF CONTENTS
Page
INTRODUCTION
Literature Review
Description of the Study Area
Geology
1
3
9
9
Climate
Temperature
Currents
Turbidity
Dissolved Qxygen
Salinity
10
12
13
13
13
14
MATERIALS AND METHODS
16
Procedures for Measuring Various Environ­
mental Parameters
Interstitial Temperatures
Movement of Sand
Turbidity
pH
Salinity
Light Penetration through Sediments
Water Content of the Substrate
Analysis of Sediment Texture
Sampling Procedures
Direct Counts of Epipelic Microalgae
Direct Counts of Epipsammic Microalgae
Biomass Determination of Chlorophyll a
Content
Percentage of Degradation of Chlorophyll a
Measurements of Production in the Gilson
Differential Respirometer
Determination of Biomass as Ash-Free
Dry Weight
16
16
16
17
17
17
18
18
18
19
22
24
25
26
27
30
Preparation of Permanent Slides for a
Taxonomic Account
Photomicrographs
RESULTS
Interstitial Temperatures
Observations on the Movement of Sand
Turbidities
31
32
34
34
36
36
Page
pH of the Sediment
Salinities
Light Penetration through the Sediment
Water Content of the Sediments
Sediment Analysis
Direct Counts of Interstitial Microalgae
Estimate of Chlorophyll a
Southbeach Study Area
Sally's Bend Study Area
Comparison of the Concentrations of Chlorophyll
a at the Two Study Areas
Determination of the Percentage of Pheophytin
Southbeach Study Area
Sally's Bend Study Area
Comparison of Percentage of Pheophytin at the
Two Study Areas
Experimental Gross Production of Oxygen
Southbeach Study Area
Sally's Bend Study Area
Comparison of Production of Oxygen at the
Two Study Areas
Analysis of Ash-Free Dry Weight
Taxonomic Study
36
39
39
41
41
42
50
50
57
62
63
63
73
74
76
76
84
85
90
90
DISCUSSION
123
SUMMARY
149
BIBLIOGRAPHY
152
APPENDICES
162
Appendix A
162
Appendix B
180
LIST OF FIGURES
Page
Figure
1
Map showing location of the two study areas
and transects sampled (U. S. C. & G. S.
2
3
4
5
6
7
8
9
10
Chart No. 6058. 1965).
11
Piston corer showing piston, core
sectioning device, and plastic coring
tubes containing sediment.
20
Variation in the ratios of optical densities
at 430 nm to that at 410 nm as related to
the percentage of pheophytin in mixtures
of a standard chlorophyll extract and
pheophytin.
28
Effect of waves or tides on the temperature
of surface and subsurface sediments at
Southbeach measured at different times
of the year.
35
Seasonal variations in the absolute turbidity
of the water at the two sampling sites.
37
Seasonal variations in pH at the two
sampling sites.
38
Seasonal variations in salinity at the
two sampling sites.
38
Variation with intertidal level in numbers
of cells/cm3 of sediment at Southbeach as
related to season and depth in the sediment.
44
Variation with intertidal level in numbers of
cells/cm3 of sediment at Southbeach as
related to season and depth in the sediment.
45
Variation with intertidal level in numbers of
cells/cm 3 of sediment at Southbeach as
related to season and depth in the sediment.
46
Figure
11
12
13
14
15
16
17
18
19
20
Page
Variation with depth in the number of cells/
cm3 of sediment at Southbeach as related
to season and position in the intertidal
zone.
48
Variation with depth in the number of cells/
cm3 of sediment at Southbeach as related to
season and position in the intertidal zone.
49
Variation with intertidal level in concentra­
tion of chlorophyll a as related to season
and depth in the sediment.
52
Variation with intertidal level in concentra­
tion of chlorophyll a as related to season and
depth in the sediment.
53
Variation with intertidal level in concentra­
tion of chlorophyll a as related to season and
depth in the sediment.
54
Variation with intertidal level in concentra­
tion of chlorophyll a as related to season
and depth in the sediment.
55
Variation with intertidal level in concentra­
tion of chlorophyll a as related to season
and depth in the sediment.
56
Variation with depth in the concentration
of chlorophyll a in the sediments as
related to position in the intertidal zone
and season.
58
Variation with depth in the concentration of
chlorophyll a in the sediments as related
to position in the intertidal zone and
season.
59
Variation with depth in the concentration of
chlorophyll a in the sediments as related to
position in the intertidal zone and season.
60
Page
Figure
21
22
23
24
25
26
27
28
Variation with intertidal level in pheophytin
expressed as percentage of total chlorophyll
(chlorophyll a and pheophytin) as related
to season and depth in the sediment.
65
Variation with intertidal level in pheophytin
expressed as percentage of total chlorophyll
(chlorophyll a and pheophytin) as related to
season and depth in the sediment.
66
Variation with intertidal level in pheophytin
expressed as percentage of total chlorophyll
(chlorophyll a and pheophytin) as related to
season and depth in the sediment.
67
Variation with intertidal level in pheophytin
expressed as percentage of total chlorophyll
(chlorophyll a and pheophytin) as related to
season and depth in the sediment.
68
Variation with intertidal level in pheophytin
expressed as percentage of total chlorophyll
(chlorophyll a and pheophytin) as related to
season and depth in the sediment.
69
Variation with depth in pheophytin expressed
as percentage of total chlorophyll (chlorophyll
a and pheophytin) in the sediments as related
to the position in the intertidal zone and
season.
71
Variation with depth in pheophytin expressed
as percentage of total chlorophyll (chlorophyll
a and pheophytin) in the sediments as related
to the position in the intertidal zone and
season.
72
Variation with depth in pheophytin expressed
as percentage of total chlorophyll (chlorophyll
a and pheophytin) in the sediments as related
to the position in the intertidal zone and
season.
73
Figure
29
30
31
32
33
34
35
36
37
38
39
Page
Variation with intertidal level in potential
gross production of oxygen as related to
season and depth in the sediment.
79
Variation with intertidal level in potential
gross production of oxygen as related to
season and depth in the sediment.
80
Variation with intertidal level in potential
gross production of oxygen as related to
season and depth in the sediment.
81
Variation with intertidal level in potential
gross production of oxygen as related to
season and depth in the sediment.
82
Variation with intertidal level in potential
gross production of oxygen as related to
season and depth in the sediment.
83
Variation with depth in potential gross pro­
duction of oxygen as related to position in
the intertidal zone and season.
86
Variation with depth in potential gross
production of oxygen as related to position
in the intertidal zone and season.
87
Variation with depth in potential gross
production of oxygen as related to position
in the intertidal zone and season.
88
Variation with intertidal level in organic
matter as related to season and depth in
the sediment.
92
Variation with intertidal level in organic
matter as related to season and depth in
the sediment.
93
Variation with intertidal level in organic
matter as related to season and depth in
the sediment.
94
Figure
40
41
42
43
44
Page
Variation with intertidal level in organic
matter as related to season and depth in
the sediment.
95
Variation with intertidal level in organic
matter as related to season and depth in
the sediment.
96
Variation with depth in organic matter in
the sediments as related to season and
position in the intertidal zone.
97
Variation with depth in organic matter in
the sediments as related to season and
position in the intertidal zone.
98
Variation with depth in organic matter in
the sediments as related to season and
position in the intertidal zone.
99
LIST OF TABLES
Page
Table
1
2
3
4
5
6
7
8
9
10
11
Interstitial water in g/cm 3 of the
sediment layer 0.0-1.3 cm below the
surface.
40
Mechanical analysis of the sediments by
the Bouyoucos hydrometer method.
41
Seasonal variation in number of cells/cm 3
of sediment.
43
Seasonal variation in concentrations of
chlorophyll a in pg/cm3 corrected for
pheophytin content.
51
Seasonal variation in pheophytin expressed
as percentage of total chlorophyll
(chlorophyll a and pheophytin).
64
Seasonal variation in potential gross
production expressed as pg 02/cm3/hr.
77
Seasonal variation in ash-free dry weight
of organic matter in mg/cm3 of sediment.
91
Species of diatoms collected from the
Southbeach and Sally's Bend study areas
from July 6, 19 67 to July 14, 1968.
100
Diatom species collected from the lower
intertidal zone at the Sally's Bend and
Southbeach study areas.
105
Diatom species collected from the mid-
intertidal zone at the Sally's Bend and
Southbeach study areas.
111
Diatom species collected from the upper
intertidal zone at the Sally's Bend and
Southbeach study areas.
117
LIST OF APPENDIX TABLES
Table
1
2
3
4
5
6
7
8
9
Page
Number of cells/cm3 in the sediment
layers 0. 0-1. 3 cm and 1.3-2. 6 cm below
the surface at Southbeach.
162
Number of cells/cm3 in the sediment
layers 5. 2-6. 5 cm and 7. 8-9. 1 cm below
the surface at Southbeach.
163
Biomass expressed as concentration of
chlorophyll a in p,g/cm 3 in the sediment
layer 0. 0-1.3 cm below the surface.
164
Biomass expressed as concentration of
chlorophyll a in p.g/cm3 in the sediment
layer 1.3-2. 6 cm below the surface.
165
Biomass expressed as concentration of
chlorophyll a in p,g/cm 3 in the sediment
layer 5.2-6.5 cm below the surface.
166
Biomass expressed as concentration of
chlorophyll a in p.g/cm3 in the sediment
layer 7. 8-9. 1 cm below the surface.
167
Concentration of pheophytin expressed as
percentage of total chlorophyll (chlorophyll
a and pheophytin), in the sediment layer
0. 0-1. 3 cm below the surface.
.168
Concentration of pheophytin, expressed as
percentage of total chlorophyll (chlorophyll
a and pheophytin), in the sediment layer
1. 3-2. 6 cm below the surface.
169
Concentration of pheophytin, expressed as
percentage of total chlorophyll (chlorophyll
a and pheophytin), in the sediment layer
5.2 -6. 5 cm below the surface.
170
Page
Table
10
11
12
13
14
15
16
17
18
Concentration of pheophytin, expressed as
percentage of total chlorophyll (chlorophyll
a and pheophytin), in the sediment layer
7. 8-9. 1 cm below the surface.
171
Rates of gross production in pg 02/cm3/hr
in the sediment layer 0. 0-1. 3 cm below the
surface.
172
Rates of gross production in pg 02/cm3/hr
in the sediment layer 1. 3-2. 6 cm below the
surface.
173
Rates of gross production in pg 02/cm3/hr
in the sediment layer 5. 2-6. 5 cm below the
surface.
174
Rates of gross production in pg 02/cm3/hr
in the sediment layer 7. 8-9. 1 cm below the
surface.
175
Biomass expressed as ash-free dry weight
in mg/cm3 in the sediment layer 0. 0-1. 3
cm below the surface.
176
Biomass expressed as ash-free dry weight
in mg/cm3 in the sediment layer 1. 3-2. 6 cm
below the surface.
177
Biomass expressed as ash-free dry weight
in mg/cm3 in the sediment layer 5.2-6. 5
cm below the surface.
178
Biomass expressed as ash-free dry weight
in mg/cm3 in the sediment layer 7. 8-9. 1
cm below the surface.
179
LIST OF PLATES
Plate
Page
Achnanthes brevipes - Actinoptychus
marmoreus
180
II.
Amphiprora alata
Amphora proteus
181
III.
Amphora angusta
Biddulphia aurita
182
IV.
Biddulphia dubia - Campylosira
cymbelliformis
183
Cerataulus turgidus - Cocconeis scutellum
184
Coscinodiscus curvatulus - Coscinodiscus
nitidus
185
VII.
Coscinodiscus radiatus - Diploneis smithii
186
VIII.
Epithemia sorex - Gomphonema angustatum
187
Gyrosigma balticum - Hantzschia marina
188
Isthmia nervosa - Navicula cancellata
189
Navicula complanata - Navicula humerosa
190
Navicula lyra - Navicula sp. 4
191
I.
V.
VI.
IX.
X.
XI.
XII.
Nitzschia sigma
XIII.
Nitzschia acuminata
XIV.
Opephora schwartzii - Pleurosigma
XV.
XVI.
XVII.
192
angulatum
193
Raphoneis amphiceros - Rhabdonema
arcuatum
19 4
Stephanopyxis nipponica - Terpsinoe
americana
195
Trachyneis aspera - Tropodeneis vitria
196
ECOLOGY OF BENTHIC MICROALGAE OF
ESTUARINE INTERTIDAL SEDIMENTS
INTRODUCTION
Interest in estuaries and bays has increased sharply during the
past few years. This is due in part to mass migrations of people to
coastal areas where residents wish to live, swim, boat, and fish.
The recreational value of such areas has long been recognized but
the satisfaction of living at the seashore has resulted in the urbani­
zation of great stretches of coastline throughout the world. Of the
ten largest metropolitan areas in the world, seven border estuarine
areas, i. e. New York, Tokyo, London, Shanghai, Buenos Aires,
Osaka, and Los Angeles. Dredging, diking, and filling to create
waterfront real estate has permanently changed the ecology of many
estuaries and has adversely effected estuarine dependent organisms.
In this country California has suffered the greatest rate of destruction
with 67% of its estuarine areas being filled for housing, roads,
airports, parking lots, factories, dumps, and baseball stadiums. On
the eastern seaboard New Hampshire, Connecticut, New York, and
New Jersey have lost between 10 and 15% of their estuarine areas to
dredging and filling (Last ditch fight... 1967).
Even areas with less population pressure and minimum
urbanization are not immune to plans for developing industrial
2
facilities. For example, the city of Newport, Oregon is considering
plans for the economic development of Yaquina Bay and estuary,
including construction of a deep water shipping facility adjacent to
Oregon State University's Marine Science Center. Because of the
possible threat to the future of the Marine Center, the University of
Oregon Bureau of Governmental Research has recommended that
Southbeach be kept for marine science and that industry, including
deep water shipping, be located on the north side of the bay. How­
ever, when the objectives are the creation of new jobs and the
relief of a depressed economy, the general populace does not want to
be reminded of the benefits of conservation.
In the face of this continuing and extensive development of
coastal areas it is evident that knowledge of estuarine and coastal
environments is not keeping pace with the problems arising from their
exploitation.
There is an ever increasing need for more comprehen­
sive understanding of the estuarine environment. With this objective
in mind it was a major purpose of this study to provide information
concerning the ecology of benthic microalgae of intertidal estuarine
sediments. The study includes the taxonomy and distribution of the
estuarine benthic microalgae present, it provides an estimate of the
biomass and potential productivity of this community, and describes
the environmental factors which may control the distribution and
abundance of the benthic microalgae in the Yaquina Bay estuary.
3
Literature Review
Although the existence of diatoms and other microalgae in
marine intertidal areas has long been recognized, research on the
marine benthic intertidal flora has most commonly been restricted to
study of the macrophytic seaweeds while the microphytes such as the
diatoms have been neglected.
This benthic flora can be considered
a part of the benthos which has been defined by Round (1964) as the
assemblage of organisms living on the bottom of freshwater or brack­
ish ponds, lakes, rivers, and the sea bed. The benthic microalgae
include the epilithic flora on rock or stone surfaces, the epiphytic
flora attached to macrophytic benthic plants, the unattached epipelic
flora in sediments, and the epipsammic flora attached to sand grains.
Conspicuous films of benthic microalgae have been reported
from Britain by Aleem (1949, 1950a, b), Bracher (1929), Carter
(1932, 1933a, b), Ghazzawi (1933), Hendey (1951, 1964), Herdman
(1921, 1922, 1924), Hopkins (1963), Hustedt and Aleem (1951),
Meadows and Anderson (1968), Perkins (1960), Round (1960), and
Smyth (1955).
In France, Cal lame and Debyser (1954) have studied
diatoms of the intertidal zone. In Germany, Hustedt (1939) has
reported heavy growths of diatoms along the coast. Such growths are
so frequent in this area they have been given the vernacular names
"Wattenpapier" and "Wattblute". Williams (1962) cites numerous
other investigations of the benthic microalgae in Germany.
4
Castenholz (1967) has noted the occurrence of non-planktonic diatoms
on the western coast of Norway. Some Danish investigations of the
benthic microalgae have been cited by Pamatmat (1968). In India,
Ganapati, Lakshmana Rao, and Subba Rao (1959) have reported on the
occurrence of benthic microalgae and in Africa, Cholnoky (1968) and
Hendey (1958) have studied the benthic microflora.
In this country abundant growths of microalgae have been found
on tidal flats in Massachusetts (Faux4-Fremiet, 1951; Moul and
Mason, 1957; Lackey, 1961; Drum and Webber, 1966), New Jersey
(Nelson, 1947), North Carolina (Pearse, Humm and Wharton, 1942;
Hustedt, 1955), Georgia (Pomeroy, 1959; Williams, 1962), Texas
(Oppenheimer and Wood, 1962; Wood, 1963), and in Washington
(Phifer, 1929; Pamatmat, 1968).
The first practical method of estimating the extent of algal
populations based on extraction and determination of the concentra­
tion of chlorophylls and carotenoids from plankton samples was
developed by H. W. Harvey (1934). This method of estimating algal
standing crops has been further developed and extended by several
workers, notably Gardiner (1943), Tucker (1949), Richards and
Thompson (1952), and more recently by Parsons and Strickland
(1963). Estimation of standing crops of benthic microalgae by
extraction and measurement of chlorophyll concentrations have been
relatively few even though this method has been much used in
5
estimating the phytoplankton biomass.
Determinations of the density of benthic microalgae have been
made by Wetzel (1964) who measured the pigment content of an
epipelic population by extracting the upper centimeter of 400 cm 2
cores. Moul and Mason (1957) obtained estimates of diatoms from
marine sand and mud flats. Eaton and Moss (1966) and Moss and
Round (1967) used the cellulose tissue trapping technique to obtain
pigment estimation of epipelic microalgae. Moss and Round (1967)
also extracted pigments from the epipsammic algae and Steele and
Baird (1968) made extracts of sediment cores which were collected
from the sublittoral zone of a sandy beach.
The equations used to calculate the amount of pigment depend
for their validity on the absence of pheo-pigments in the extracts.
According to Strickland and Parsons (1965), pheophytins are generally
absent in extracts from open ocean phytoplankton. However, it has
been shown by Yentsch and Menzel (1963), Lorenzen (1965), and
Yentsch (1965) that chlorophyll degradation products may at times
constitute a significant fraction of the pigments present in seawater.
A method for determining the pheo-pigments has been developed
recently by Lorenzen (1967). Moss (1967) has also developed a method
for the estimation of the percentage of degradation of chlorophyll to
pheo-pigment.
Non-functional chlorophyll and its degradation products are
6
dominant components of sediments both at the surface and in the
deeper strata. Vallentyne (1955) has shown that extracts of sedimen­
tary pigments contain chlorophyll degradation products as well as
chlorophyll a , all having absorption peaks at 665 nrn. Gorham
(1960) found large amounts of degraded pigments in surface muds in
the English Lake District. Patterson and Parson (1963) observed
that 93% of the pigments in a littoral mud sample was pheo-pigment.
Moss (1967) noted that extracts of fresh-water muds from two English
ponds contained 53 and 55% degraded pigment. Tietjen (1968) found
that a major fraction of chlorophyll in estuarine sediments was
detrital chlorophyll. These chlorophyll degradation products are
important because they can cause errors in the estimation of
chlorophyll a by trichromatic spectrophotometry.
Manometric estimates of oxygen production by benthic micro-
algae also have been few although manometric methods for estimating
photosynthesis and respiration have been available for many years.
The usage and theory behind the differential respirometer has been
completely described by Umbreit, Burris, and Stauffer (1964).
Manometry has been frequently used to elucidate the physiology
of photosynthesis in marine algae. Ogata and Matsui (1965b) review
earlier investigations utilizing manometry to determine rates of
photosynthesis of some marine algae. More recently Haxo and
Clendenning (1953), Ogata and Matsui (1965a, b), and Kjeldsen (1967)
7
have utilized manometry to study photosynthetic rates of marine algae.
A few studies on production by benthic microalgae in situ have
utilized modifications of the light and dark bottle technique and have
analyzed for oxygen by Winkler titration. Pomeroy (1959) utilized
bell jars pressed down into the sediments and filled with sterile,
filtered, estuarine water. The Winkler method was used to estimate
initial and final oxygen concentrations. Pamatmat (1968) also
utilized the bell jar technique to measure oxygen production of
sediments in situ. He also took centimeter layers of sediment,
placed them in bottles and attempted to measure the potential photo­
synthetic activity of subsurface sediment with an oxygen electrode.
Other investigations concerning various facets of intertidal
benthic productivity have been carried on by Griz(ntved (1960, 1962 as
cited in Pamatmat, 1968) who studied production by the microbenthos
in some Danish fjords estimating assimilation by the C14 method, and
by Taylor and Palmer (1963) and Taylor (1964) who have investigated
the relationship between light and photosynthesis in intertidal benthic
diatoms also by measuring C14 uptake.
The distribution of the microflora has been studied in
the Woods Hole area of Massachusetts by Moul and Mason (1957) and
Lackey (1961); in the Port Aransas, Texas area by Oppenheimer and
Wood (1962); in the Friday Harbor area of Washington by Phifer
(1929) and Pamatmat (1968); in the Coos Bay area on the Oregon
8
coast by Castenholz (1963), who was primarily interested in the
vertical distribution of the epilithic microflora; along the Scottish
coast by Meadows and Anderson (1968), who studied the distribution
of epipsammic microorganisms, and on the south coast of England by
Aleem (1949, 19$.0a, b).
Studies pertaining primarily to the taxonomy of the benthic
intertidal diatoms were performed by Drum and Webber (1966) on a
Massachusetts salt marsh; by Wood (1963) on the sediment of south
Texas bays; by Hustedt (1939, 1955) on the mud-flats of the North Sea
and at Beaufort, North Carolina; by Hustedt and Aleem (1951) on the
mud-flats of Salstone, England; and by Hendey (1951, 1964) in
Chichester Harbor and coastal British waters.
One aspect of benthic microalgal ecology that has attracted
much concentrated attention is the rhythmic vertical migration of these
organisms in the sediment. Many diatoms move to the sediment
surface only during the day or in some areas only at low tide during
the day. The surfacing of tidal flat diatom communities on exposure
at low tide and their disappearance just before the incoming tide was
first described by Fauvel and Bohn (1907, as cited in Hopkins, 1966).
Bracher (1929), Faux4-Fremiet (1951), and Pomeroy (1959) found
that both illumination and the tidal rhythm influenced the vertical
migration of the algae. Aleem (1950a) and Perkins (1960) concluded
that illumination alone controlled the rhythm in the areas that they
9
studied. Cal lame and Debyser (1954) and Hopkins (1965) described
the rhythm as persisting under laboratory conditions for only one day
and discovered that it needed to be reenforced each day by the tidal
rhythm if synchronization with the tides was to persist. Palmer and
Round (1965), Round and Happey (1965), Round and Palmer (1966),
and Round and Eaton (1966) have shown that a distinct migratory
rhythm is not necessarily connected with a daily tidal cover but may
be an expression of an inherent biological activity of both motility
and photo-sensitivity, and possibly may also involve a positive
geotaxy.
Description of the Study Area
Geology
Yaquina Bay and its estuarine drainage basin is located on the
central coast of Oregon on the western flank of the Coast Range. The
bay and estuary were formed when the river mouth was drowned as
sea level rose during post-Pleistocene deglaciation (Baldwin, 1964).
The estuary extends 23 miles inland along the Yaquina River but has
its fresh-water head 10 miles inland near Elk City.
Ku lm (1965) divided Yaquina Bay into three realms of deposi­
tion based on sediment texture and mineralogy. These realms have
been termed marine, fluviatile, and marine-fluviatile. The marine­
fluviatile realm lies between the fluviatile and marine realms and
10
includes the Southbeach and Sally's Bend tidal flats (Figure 1). The
sediments of the Southbeach tidal flat are characteristic of the marine
realm while those of the Sally's Bend tidal flat resemble those of the
fluviatile realm. For a comprehensive treatment of the geomorphology
and sedimentology of the estuary see Ku lm (1965).
Climate
Cool, dry summers and mild, wet winters are characteristic
of the Yaquina Bay area. The mean annual air temperature is 51. 1F
but ranges from an average of 64.5 F in summer to 48.9 F in winter.
The mean annual precipitation recorded at Newport is 64.9 inches.
Daily precipitation ranges from extremes of 0. 00 inches on a summer
day to 3. 09 inches on a winter day. Between 80 to 90% of the
precipitation usually falls from September to May. Run-off is
closely related to fluctuations in precipitation. During summer
months the average run-off is approximately 150 cu ft/sec whereas
in winter it is about 1000 cu ft/sec (Kjeldsen, 1967).
Fog frequently occurs in summer and fall when cold water
brought to the surface by upwelling cools the warm surface air (Ku lm,
1965).
Cooper (1958) has reported the velocity of onshore wind at
Newport in summer to be greater than 4 mph and from a north or
northwesterly direction. The occurrence of winds of more than 16
11
SALLY'S BEND
TIDAL FLAT
SOUTHBEACH
TIDAL FLAT
111.1:11K
YAQUINA BAY AND RIVER
l'FiraTe4=­
Figure 1. Map showing location of the two study areas and transects
sampled (U. S. C. & G. S. Chart No. 6058. 1965).
12
mph have also been reported for the summer. Winter winds are
offshore with velocity of more than 4 mph but usually less than 16
mph.
However, also during winter, infrequent high velocity onshore
winds from the south to southwest occur. Fall and spring winds are
transitional with periods in which winter-type and summer-type
winds alternate.
Temperature
Kjeldsen (1967) assembled data from a Georgia Pacific Corpora­
tion study by Waldemar Seton and determined the average annual
variation in water temperatures in winter and summer for a period of
four years. He found them to be 5. 6 C off the north jetty; 8 C at
Marker 15; 11 C at Markers 25 and 26; 13 C at Marker 47; and 14 C
at the Toledo bridge. Frolander (1964) in his 28 hour study of
Yaquina Bay found that the stations furthest upstream had the highest
average temperatures and also the greatest temperature ranges when
compared with the stations near the mouth of the estuary. A
decrease in the temperature of the surface water occurs in November
and the low temperatures continue until March.
The lowest
temperatures of the bottom water occur in summer and are asso­
ciated with offshore upwelling that carries cold water into the estuary
by tidal currents. Generally speaking, however, the temperatures
of bottom water are highest in spring and late fall with the highest
13
surface water temperatures occurring in summer and early fall
(Ku lm, 1965). For more complete information on the temperature
regimes within the bay see Frolander (1964), Ku lm (1965), and
Kjeldsen (1967).
Currents
Ku lm (1965) estimated that the velocity of the tidal current
averaged 77 cm/sec with a maximum value of 109 cm/sec at the
entrance of the bay between the two jetties. Burt (as noted in Ku lm,
1965) measured current velocities at channel localities 1.1, 2. 9, and
9. 7 miles from the bay entrance with current drags. At 1. 1 miles
from the entrance the current velocity average was 40 cm/sec; at
2. 9 miles, it was 30 cm/sec; and at 9. 7 miles from the entrance the
velocity decreased to 20 cm/sec.
Turbidity
According to Kjeldsen (1967) the occurrence of turbid conditions
within the bay is irregular and not related to precipitation and run-off
but to the occurrence of onshore winds. These winds cause a wave
action which resuspends settled particles.
Dissolved Oxygen
During summer when the water is completely mixed in the
14
channel, Ku lm (1965) found low concentrations of dissolved oxygen of
4. 5 m1/1 at Coquille Point. In winter, when there is a lack of tidal
mixing, the concentrations of dissolved oxygen increased to more
than 6 m1/1, there being little difference between values in surface
and bottom waters. Ku lm (1965) states that the increase in dissolved
oxygen is probably due to the cold fluviatile waters which are
saturated with oxygen.
Salinity
Burt and McAlister (1959) described Yaquina Bay as being
partly mixed in winter and spring and showing a salinity difference of
4 to 19%obetween waters at top and bottom. In summer and fall the
Bay is completely mixed with a salinity difference of less than 3%0
between top and bottom.
Frolander (1964) made hourly measurements of salinity at the
bottom over a 28 hour period during August, 1963. He showed that in
late summer when the estuary is well mixed there is little variation
in salinity near the mouth of the estuary but further up the channel he
found increased differences in salinity between high and low tide
readings.
Kjeldsen (1967) cited a study of salinities by Raymond Jones of
Georgia Pacific Corporation.
The mean salinity of weekly samples of
surface water at the jetties was 33%owith a mean variation of 5 %o
15
between winter and summer. Further up the channel at Marker 25 %o
the mean salinity was 28%owith an average variation of 26%obetween
winter and summer.
Ku lm (1965) noted a correlation with precipitation and the
average monthly differences in salinity of waters at surface and
bottom due to a lack of tidal mixing. Salinities at the bottom of the
estuary reached their lowest values in winter. For more comprehen­
sive treatments of salinity regimes in the Bay see Burt and
McAlister (1959 ),Frolander (1964), Ku lm (1965), and Kjeldsen (1967).
16
MATERIALS AND METHODS
Procedures for Measuring Various
Environmental Parameters
Interstitial Temperatures
The temperature of the interstitial water of the sediment was
measured at several depths down to 20 cm for varying lengths of time
at different times of the year in order to assess the change in
temperature that occurred as the incoming tide covered the sediment.
A YSI Model 41 Tele-Thermometer (Yellow Springs Instrument Co.,
Yellow Springs, Ohio) and general purpose vinyl tip thermistor
probes were utilized for all temperature measurements.
The probes
were inserted in the sediment to the desired depths and taped to a
cement block. Readings were taken at regular intervals.
Movement of Sand
The effect of tidal current on sand movement was investigated
at the Southbeach study area by using sand coated with a fluorescent,
red plastic (Great American Color Co.
,
Los Angeles, Calif. ).
According to Ingle (1966) the hydraulic properties of the sand grains
are unaffected as long as a wetting agent is used prior to release. A
hollow, tin cylinder, 16 cm in length and 13 cm in diameter, was
pushed down into the sediment until the lip of the cylinder was flush
17
with the sediment surface. Sand was scooped out of the cylinder.
Dyed sand that had previously been wetted by a wetting agent was
poured into the cylinder. The cylinder was carefully removed leav­
ing a plug of dyed sand. Observations were made for four months from
July 14, 1968 to November 2, 1968.
Turbidity
Samples of estuarine water were collected at regular intervals
throughout the sampling period and the absolute turbidity of this
water was measured with an Elko II Electrophotometer (Carl Zeiss
Inc. , New York, N. Y. ).
pH
The pH of the sediment at the two study areas was determined
at various times of the year during the sampling period. An Orion
Digital pH Meter (Orion Research Inc. , Cambridge, Mass. ) was used
in the laboratory on samples brought in from the field.
Salinity
Samples of water from pools on the tidal flats were collected
regularly throughout the sampling period and analyzed for salinity
with a hydrometer. Conversions were made from density to parts
per thousand by the use of a U. S. Coast and Geodetic Survey table
18
(U. S. Department of Commerce, 1941).
Light Penetration through Sediments
The effect of depth of substrate on light penetration was in­
vestigated on sediments collected from the two tidal flats. A
radiant source of incandescent and fluorescent light providing 260 foot
candles was utilized. Sediment layers of varying thickness from the
two study areas were placed in petri dishes. These in turn were
placed directly on the photocell of a Weston Model 756 light meter
and the percentage of light penetration determined.
Water Content of the Substrate
An analysis of the amount of water the sediment from each
station contained at low tide was performed by oven drying the
sediment at 70 C. The loss in weight represented the amount of
moisture present in the sediment.
Analysis of Sediment Texture
The sediment from the two tidal flats was determined by using
the Bouyoucos (1933) hydrometer method. The sediment was
classified according to the Wentworth (1922) scheme of classification.
19
Sampling Procedures
Samples of the benthic microalgal community were collected by
use of an acrylic plastic piston corer 23 cm in length with an 0. D. of
1. 9 cm and an I. D. of 1.5 cm. A rubber piston was devised consist-.
ing of a metal handle with a loop for gripping, two no. 1 rubber
stoppers, a 3 in. (7. 6 cm) long x 1/8 in. (0. 32 cm) diameter bolt with
three nuts and two washers. The two nuts immediately above and
below the rubber stoppers could be tightened or loosened so that a
proper fit inside the acrylic plastic tube could be obtained (Figure 2).
Samples were collected at each study area approximately once a
month during a low tide. A transect from low tide line to mean high
tide line was marked off and samples taken at 30 m intervals. This
resulted in the collection of 28 cores on each collection date: four
each at low tide level, 30 m from low tide level, 60 m from low tide
level, 90 m from low tide level, 120 m from low tide level, 150 m
from low tide level, and at mean high tide level.
Due to the consistency and high water content of the sediment
at the Sally's Bend study area, it was necessary to construct and
utilize a pair of mud skis that allowed the investigator free access to
the tidal flat. In this area as the plastic corer was being inserted
into the sediment the piston was pulled upward simultaneously. This
held the sediment core in place. After the corer was removed from
20
Figure 2. Piston corer showing piston, core sectioning device, and
plastic coring tubes containing sediment.
21
the sediment, the piston was removed very gently so as not to disturb
the core, and the plastic tube was corked at the top and at the
bottom.
The sampling of the sandy sediment at Southbeach did not
require the use of the mud skis o r the rubber piston. The empty
plastic tube was simply pushed down into the substrate. A cork was
inserted at the top of the tube while it was still buried in the sediment.
The plastic corer was withdrawn together with the enclosed sediment
core and a cork inserted at the bottom. All cores from Southbeach
and Sally's Bend were kept upright in a Styrofoam ice chest at the
time of collection and during the time of transporting them back to the
Marine Science Center. As soon after collection as possible the
sediment cores were pushed out of the plastic tubing with a piston
and immediately sliced into 1 cm 3 or 1/2 cm 3 sections corresponding
to depths of 0. 0-1. 3 cm, 1. 3-2. 6 cm, 5. 2-6. 5 cm, 7. 8-9. 1 cm and
0. 0-. 65 cm, 1. 3-1.95 cm, 5.2-5. 85 cm, and 7. 8-8.45 cm,
respectively.
These sections were placed in glass vials, capped,
and returned to Corvallis. The 1 cm 3 core sections were used for:
1) measuring the chlorophyll a concentration; 2) measuring the
percentage of degradation of chlorophyll a; 3) measuring the ash-free
dry weight by ashing in a muffle furnace; 4) making direct counts of
live microalgae under the compound microscope; and 5) a taxonomic
investigation of the microalgae and their distribution at the two study
22
areas. The 1/2 cm 3 core sections were used for measuring potential
oxygen production and consumption in a Gilson Differential
Respirometer.
Direct Counts of Epipelic Microalgae
Direct counts of live microalgae were made by observing wet
mounts using the 43x objective of a compound microscope. Counts
were made of the organisms in sections of cores from 0.0-1.3 cm,
1.3-2. 6 cm, 5.2-6.5 cm, and 7. 8-9.1 cm depths taken every 30 m
along a transect from low tide level to mean high tide line. A 1 cm 3
portion of sand with its algal community was placed in a 40 ml test
tube.
Two ml of aged, Millipore-filtered sea water was added to the
sediment in the tube. The tube was agitated on a vortex-type mixer
for approximately 10 seconds.
This procedure served to separate the
epipelic algae from the interstices of the sand grains. Only the
epipelic community could be counted following this treatment because
the epipsammic algal community still adhered to the sand grains.
After a few seconds the sand grains settled leaving the microalgae in
suspension. This 2 ml algal suspension was decanted into a graduated
10 ml centrifuge tube. This procedure was repeated five times until a
10 ml volume of algal suspension accumulated in the tube. The
suspension was centrifuged at 7000 rpms for a period of 3 minutes
leaving a visible algal pellet at the bottom of the tube. Nine mls of
23
of supernatant were pippetted off very carefully without disturbing the
algal pellet. The remaining water and pellet were agitated again on
the mixer to obtain a uniform suspension of microalgae. To this
suspension was added three drops of a concentrated rose bengal dye
solution and the suspension agitated again. The tubes were left for
12 hours. It was found that the rose bengal stained the chloroplasts
a deep pink and this facilitated the counting of live organisms and
prevented the counting of any dead frustules that were present.
A Whipple disc was inserted in the eyepiece of the compound
scope to facilitate counting. All the algae in the field of the micro­
scope enclosed by the grid were counted. Under the 43x objective
the Whipple grid was equal to 0.289 mm2. Cover slips 22 mm2 were
used throughout the study. Therefore, there were 76 fields present
per coverslip under the 43x objective. One quarter of the coverslip
or 19 fields were counted and the total number of algae present were
recorded. Fields were selected for counting by randomly choosing 19
fields along four separate strips near the center of the coverslip. A
0. 05 ml aliquot of the 1 ml algal suspension was used. This is 1/20
of the suspension.
Therefore, the total number of algae in 1 cm 3 of
sand when 19 fields are counted is equal to 20 (dilution factor) x 4
(microscope factor) x total no. of algae counted in 19 fields. A
coefficient of variation performed on a series of replicate samples
was 4. 9%.
24
Only samples from the Southbeach study area were counted. It
was found that the samples from Sally's Bend always had so much silt
and organic matter present that it obscured the dyed algae and
accurate counts could not be obtained.
Direct Counts of Epipsammic Microalgae
After the epipelic microalgae were separated from the sediment
by repeated agitation and decanting, the sediment itself was treated
to remove the algal community which remained attached to the sand
grains. Ten ml of a 30% hydrogen peroxide solution was added to the
40 ml test tubes containing the sediment. These were left to sit at
room temperature for two days while oxidation of the organic matter
present occurred. This process served to oxidize the sheath of
pectinous material which attached the epipsammic diatoms to the
sediment. After this treatment the hydrogen peroxide was pipetted
off very carefully without disturbing the sediment and diatom frustules
remaining on the bottom. Two ml of distilled water were added and
the test tubes agitated on the mixer as before. This procedure was
repeated five times until 10 ml of diatom suspension was obtained.
This suspension was centrifuged and the supernatant pipetted off
until 1 ml remained. The 1 ml suspension was agitated on the mixer
to distribute the diatomfrustules evenly throughout the suspension.
A 0. 05 ml drop of suspension was added to a slide and 19 fields
25
counted as previously to determine the number of cells present in 1
cm
3.
This also was done only for the Southbeach study area because
of the problems with silt from Sally's Bend.
Biomass Determination by Chlorophyll a Content
The 1 cm 3 core sections of sediment from depths of 0.0-1.3 cm,
1.3-2.6 cm, 5.2-6.5 cm, and 7.8-9.1 cm from low tide level, 60 m
from low tide level, 90 m from low tide level, 120 m from low tide
level, 150 m from low tide level and mean high tide level were placed
in small glass vials. Five ml of 90% acetone in distilled water were
added and the vials capped. These were put in a dark, 50 C cold
room for a period of 18 to 20 hours. During this period pigment was
extracted from the chloroplasts by the acetone. The acetone extract
was centrifuged at 7000 rpm to rid it of suspended sediment. The
clear supernatant was decanted into a 1 cm light path quartz cuvette
and readings were taken at 630 nm, 645 nm, and 665 nm using a
Beckman DB-G Grating Spectrophotometer. The tungsten filament
light source was used at these wavelengths. The chlorophyll a content
was calculated using Strickland and Parsons' (1963) modification of the
Richards and Thompson method. The chlorophyll a content was ex­
pressed as µg of chlorophyll a /cm 3 of sediment. No turbidity correc­
tions were made at 750 nm as would be necessary when working with ma­
terial filtered through cellulose ester filters. A series of analyse s were
26
made on identical samples from each study area. Good reproducibility
was obtained in each case, the coefficient of variation being 8. 4% at
Southbeach and 9. 15% at Sally's Bend.
Percentage of Degradation of Chlorophyll a
The precentage of degradation of chlorophyll a was determined
using the same acetone extracts that were used for determining
chlorophyll a concentration. Optical densities were measured at
wavelengths of 410 nm and 430 nm using a technique developed by
Moss (1967).
An algal culture of epipsammic and epipelic organisms was
made by adding about 130 grams of sediment to an enriched seawater
medium. An extract of the algal culture containing epipelic and
epipsammic algal populations in an active phase of growth was pre­
pared in 90% acetone in distilled water so that the optical density was
not above 0. 6 in a 1 cm light path. Two 100 ml aliquants of the
extract were placed in 250 ml flasks. Five ml of 5% HC1 was added
to one flask and 5 ml of distilled water was added to the other.
These
flasks were left in the dark at room temperature for 10 minutes. The
chlorophyll extract with HC1 added is considered to be completely
oxidized to pheophytin while the flask with the distilled water added is
considered to be all chlorophyll a. Magnesium carbonate was added
to the flask of acidified extract until the solution was neutral. A
27
series of mixtures of chlorophyll extract and pheophytin were made
and their optical densities measured on the Beckman DB-G Grating
Spectrophotometer. The values from this series of mixtures were
plotted on a graph and a calibration curve was drawn (Figure 3).
The percentage of degradation of chlorophyll a in acetone
extracts of sediment samples was determined by reading optical
densities at wavelengths of 410 nm and 430 nm. A ratio of the 430:
410 0. D. values was obtained. The percentage of pheophytin was
then read from the standard curve.
Measurements of Production in the Gilson Differential Respirometer
Estimates of potential primary production and community
respiration were based on measurement of the amount of oxygen re­
leased by photosynthesis and amount utilized by respiration. To get
adequate light penetration to the sediment in the flask only a 1/2 cm
3
core section was used. These core sections were placed in the flasks
and 3 ml of aged, Millipore-filtered sea water with a salinity of 33%0
was added.
The "one vessel" method involving only oxygen exchange between
the gas and liquid phase was utilized for measuring oxygen production.
Two ml of a buffer developed by Pardee as described in Umbriet,
Burris, and Stauffer (1964) were added to the center well of each flask
along with a filter paper wick. This buffer is capable of maintaining
1.4
1.3
1. 2
ftf t
1. 1
19
ro
ro
o 1.0
a 444
a)
0
0
om
444
0.9
0.8
0.7
10
20
30
40
50
60
70
80
100
Percentage of Pheophytin
Figure 3. Variation in the ratios of optical densities at 430 nm to that at 410 run as related to the percentage of pheophytin in mixtures of
a standard chlorophyll extract and pheophytin.
29
a virtually constant pressure of 1% carbon dioxide in the gas phase.
Any carbon dioxide released is absorbed by the buffer and any carbon
dioxide taken up is replaced by the buffer. Therefore, only oxygen
production is measured.
Respiration was measured by replacing the buffer in the center
well with 3 ml of 10% KOH solution and a filter paper wick. Any
change in volume was the result of oxygen uptake since any carbon
dioxide produced was absorbed by the 10% KOH.
Measurements of changes in gas volume were taken every hour
for a period of four hours following an equilibration period of one
hour. Temperature for all productivity measurements was 15 C.
Light intensity was 800 foot candles as measured by a Weston Model
756 Sunlight Illumination meter. Flasks were shaken at 100 oscil­
lations per minute. The measured net rate of photosynthesis was
converted to an estimate of gross photosynthesis by adding to the
amount of oxygen evolved during the period of illumination the amount
of oxygen consumed during an equivalent period of darkness. Results
are expressed as p.g 0 2 /cm 3 of sediment/hr.
All results obtained with the Gilson Differential Respirometer
were corrected to S. T. P. Data concerning barometric pressure for
the periods of experiments were obtained from the U. S. Weather
Bureau on the Oregon State University campus. The following formula
was utilized:
30
(273)(Pb-3-Pw)
(t +273)(760)
where Pb = barometric pressure
Pw= water vapor pressure
t = temperature
Respirometer flasks were cleaned in white gasoline to remove
silicone grease and washed in aqua regia after each run to assure
cleanliness and non-contamination of the flasks.
A coefficient of variation was performed on identical samples
from each station. At Southbeach the C. V. was 8.23% and at Sally's
Bend, 14.9 %.
Determination of Biomass as Ash-Free Dry Weight
3
The 1 cm sections of the sediment cores were placed in tared
crucibles and weighed on a Mettler balance. These were placed in a
70 C oven for 24 hours. When removed from the oven the crucibles
were placed in a desiccator containing anhydrous Ca 2 SO4 and cooled
to room temperature. The crucibles were reweighed and the ovendry weight recorded. The crucibles were placed in a muffle furnace
at 50& C for 24 hours. When removed from the furnace the crucibles
were again placed in the desiccator until they cooled to room tempera­
ture. They were weighed and the ash-free dry weight recorded.
A coefficient of variation was determined for identical samples
from each station. At Southbeach the C. V. was 16. 8% and at Sally's
31
Bend the C. V. was 9. 6%.
Preparation of Permanent Slides for a Taxonomic Account
Permanent slides of diatoms from each study area and sampling
date were made in order to observe presence or absence of the various
diatom species composing the community at different times of the
year as well as at the different tidal levels at the two study areas.
The 1 cm 3 cores of sediment were placed in crucibles, ashed in a
muffle furnace for 24 hours to combust all organic matter present
leaving only inorganic sediment and diatom frustules. The sediment
and cleaned frustules were transferred from the crucible to a 40 ml
test tube. Two ml of distilled water were added to the sediment in the
tube. The tube was agitated on a vortex-type mixer for approximately
10 seconds. The cleaned diatom frustules take longer to settle than
the sediment and remain suspended in the supernatant water which was
decanted into a 10 ml centrifuge tube. Another 2 ml of water were
added, the sediment agitated, and the diatom suspension poured off.
This procedure was repeated until 10 ml of diatom suspension in
distilled water was obtained. The diatom suspension was spun at
7000 rpm for 3 minutes. After centrifugation, 9 ml of water were
pippetted off and 1 ml remained. This suspension was washed twice
with distilled water to remove traces of salt. A 1 ml sample of
water and algal pellet remained after this procedure. This was
32
agitated again to insure an equal distribution of diatoms throughout
the suspension. A drop of the diatom suspension was added to a 22
mm2 cover slip and dried on a hot plate at 100 C.
These cover slips
with dried diatom suspension were then inverted over a glass
microscope slide on which had been placed a drop of Hyrax mounting
medium. The slide was heated on an electric hot plate to dispel the
benzene solvent and then removed from the plate and allowed to cool.
These slides were observed under a compound microscope for the
taxonomic study.
Photomicrographs
Photomicrographs (Appendix B, Plates I through XVII) of all
diatom species identified in this study were taken from the permanent
slides. A Leitz Micro-attachment with Leica model M-1 camera body
and integral Microsix-l-model exposure meter was attached to the
phototube of a Zeiss RA model microscope. Either a Zeiss Neofluar,
N. A. 1. 30, 100x .obj.ective, a Zeiss Neofluar N. A. 1. 25, 63x objec­
tive, or a Zeiss, N.A. 0. 85, 40x objective was used with a Leitz
Periplan 10x ocular. All photomicrographs were taken at a one
second exposure without filters at a value of 10 on the light meter.
Photomicrographs were enlarged to 1500 times their normal size
except as noted. The scale figured on Plates I through XVII is equal
to either 10p.if the photomicrograph was enlarged to 1500 times, 19p, if
33
enlarged to 800 times, or 2511 if enlarged to 600 times. Kodak 35mm
high contrast copy film (ASA 64), developed in Eastman D- l9
developer, was used.
34
RESULTS
Interstitial Temperatures
It was found that the rise and fall of the tide either decreases
or increases the sediment temperature depending on season (Figure
4).
In summer lower low tide occurs during the daylight hours and
the tidal flats are heated by the incident radiation. The sediments
are therefore cooled by the encroaching tide. In winter when the air
temperatures are low and lower low tide occurs during the evening
hours the sediments are warmed by the encroaching tide. In fall and
spring the flooding of the tide may either increase or decrease the
temperature of the sediment depending on the time of low tide and the
amount of incident radiation. If low tide occurs in the early morning
hours the sediment has not had a chance to be heated by the incident
radiation and the encroaching tide will increase the sediment tempera­
tures. Conversely, if low tide occurs in late afternoon the flooding
of the tide will lower the sediment temperatures.
Changes in sediment temperature with the flooding tide are also
a function of depth. Differences in amplitude at greater sediment
depths are slight in the summer whereas in winter the differences in
amplitude are much broader. Closer to the surface of the sediment
the differences in amplitude as the area is covered and uncovered by
the tide are more pronounced both in winter and in summer. In
June 28, 1968
25
July 13, 1968
25
by tide
20
uncovered
covered
by wave
covered
20
att 8
0
p 15
15
ti
ti
a
a)
a.
tu 10
4) 10
uncovered by
tide
1200
1300
1400
1500
1600
1700
1800
1100
1130
Time
covered
by tide
20
1200
1230
1300
Time
20
September 23, 1968
February 15, 1969
0
as 15
covered
iby wave
I?, 15
ca
uncovered
v
10
uncovered
by tide
H
I
I
I
1030
1130
1230
I
1330
Time
I
1430
I
1530
a
v 10
I
1630
0800
1815
0820
0830
0845
Time
Figure 4. Effect of waves or tides on the temperature of surface and subsurface sediments at Southbeach measured at different times of
the year. o
o = surface, o
= 2 cm in depth, a -a = 7 cm in depth, 0 Q = 20 cm in depth.
36
winter the temperature fluctuation near the surface and deeper in the
sediment are nearly identical.
Observations on the Movement of Sand
Coring of the plug of red-dyed sand at intervals throughout a
four month period indicated that only the top 2.5 cm of sediment was
agitated by current and wave action.
Turbidities
The highest turbidities were recorded during fall, winter, and
spring at both study areas with marked decreases in turbidity
occurring during the summer months (Figure 5). Samples from the
Sally's Bend study area showed much higher turbidities than those
collected from the Southbeach study area at all times of the year.
pH of the Sediment
Due to the strong buffering action of seawater the pH does not
vary to any great degree. It was found that the pH varied from 7. 1 to
8. 0 at Sally's Bend and from 7. 3 to 7.7 at Southbeach (Figure 6).
The
pH of the sediment from Sally's Bend was generally lower than that of
Southbeach, probably as a result of the larger amount of organic
matter which is present in the sediment at Sally's Bend. Lower pH
values can result from the CO2 arising from the decomposition of this
37
0.0400
0.0375
0.0350
0.0325
0.0300
0.0275
0.0250
0.0225
0.0200
0.0175
0.0150
0.0125
kr­
0.0100
0.0075
I
Summer
'67
I
Fall
'67
i
Winter
'67 -'68
I
Spring
'68
I
Summer
'68
Season
Figure 5. Seasonal variations in the absolute turbidity of the water at the two sampling sites.
= Southb each.
0
0 Sally's Bend, Z1%
38
8. 5
8.0
7. 5
X
7.0
6.5
Summer
'67
Fall
'67
Winter
Spring
'67-'68
'68
Summer
'68
Season
Figure 6. Seasonal variations in pH at the two sampling sites.
A-­
= South beach.
= Sally's Bend,
45
40
8
'4 30
25
Summer
'67
Fall
'67
Winter
'67-'68
Spring
'68
Summer
'68
Season
Figure 7. Seasonal variations in salinity at the two sampling sites.
= Southbeach.
O
0
=Sally's Bend,
39
organic matter.
Salinities
The salinities, as measured by a hydrometer, ranged from 22
to 36 %o at Southbeach and from 24 to 36. 6 %o at Sally's Bend (Figure 7).
The lowest salinities occurred during the winter months and these low
salinities are correlated with the winter precipitation and runoff and
also with the low rate of evaporation of water from the sediments.
Conversely, the highest salinities were found during the summer
months when there is scarcely any precipitation or runoff and the rate
of evaporation of water on and in the sediment is likely to be greater.
The Southbeach tidal flat had higher salinities than Sally's Bend
throughout the entire sampling period. Since the Southbeach tidal
flat is under the influence of the marine realm whereas the Sally' s
Bend tidal flat is under the influence of the fluviatile realm
this is as would be expected.
Light Penetration through the Sediments
It was found that there was a definite difference in the two
sediment types with regard to light penetration. Ten percent of
available light penetrated a 2 mm layer of sand from Southbeach and
only a 0.5 mm layer of silt from Sally's Bend.
3
Table 1. Interstitial water in g/cm of the sediment layer O. 0-1. 3 cm below the surface.
Lower
Date
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1 -67
12-27-67
1 -25 -68
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
Intertidal
.6552
.9602
1.1543
1.3192
1.2105
1.0307
.8868
1.3872
.9950
1.2854
1.0588
1.1264
1.0438
Southbeach
Mid
Intertidal
.9940
.9606
.9968
1.4120
.7540
1.0041
.7210
.9718
1.0708
.9909
.9403
.8542
.8450
Sally's Bend
Upper
Intertidal
Lower
Date
.5192
.9101
1.0142
8-23-67
9-20-67
10-18-67
1.161S
11 -15 -67
.8084
.9789
1.2290
1.3037
.8424
.8650
.8758
.8407
.8824
12-13-67
1 -11 -68
2- 9-68
3- 9-68
4- 9-68
5- 7-68
6-10-68
7-14-68
Mid
Upper
Intertidal
Intertidal
Intertidal
1.1737
3.6073
1.0801
1.5885
2.6985
1.3462
1.2710
1.4221
1.2505
1.4615
1.3195
1.1189
1.0993
1.0246
1.6643
1.5625
1.6686
1.6019
1.6972
1.3357
1.3760
1.3921
1.1848
1.2602
1.2032
1.4765
1.4259
1.3833
1.0881
1.7802
1.2462
.9672
1.1451
1.1689
1.3013
1.2731
41
Water Content of the Sediments
The medium to fine sand at Southbeach had an average water
3
content of 1. 001 g/cm whereas the silt from Sally's Bend had an
average water content of 1. 397 g/c
3
(Table 1). This represents an
average difference of approximately 400 mg of water/cm3 between
the two study areas.
Sediment Analysis
The sediment at Southbeach consists of sand whereas at Sally's
Bend the sediment is a heavy clay loam (Table 2). These results are
not in accord with Ku lm (1966) who used a different scheme of
classification. However, the important fact is that the sediments of
the Sally's Bend tidal flat are significantly finer .grained than those of
the Southbeach tidal flat.
Table 2. Mechanical analysis of the sediments by the Bouyoucos
hydrometer method.
Total percent sand
Total percent coarse silt
Total percent fine silt
Total percent clay
Sediment type:
*Sand
* *
Heavy clay loam
Southbeach *
Sally's Bend
95. 7%
37.0%
2. 3%
28. 0%
0. 3%
1.0%
1. 7%
34. 0%
J.
42
Direct Counts,of Interstitial Microalgae
Accurate counts could not be obtained from the samples
collected at the Sally's Bend study area because of the large amounts
of fine silt that obscured the cells during counting. Therefore, cells
were counted only in samples from the Southbeach study area.
The data are summarized in Table 3 and Appendix A, Tables 1
and 2.
Table 3 contains data concerning variation in numbers of
cells/cm 3 of sediment on a seasonal basis from summer 1967 to
3
summer 1968 at the various tidal levels. The number of cells/cm
of sediment ranged from a mean of 51,760 in summer 1968 in the
5.2-6.5 cm core layer from the lower intertidal zone to a mean of
380 in summer 1967 in the 7. 8-9.1 cm core layer from the upper
intertidal zone.
3
The number of cells/cm in cores from the lower, mid, and
upper intertidal zones showed a significant difference (p < 0. 005) at
the 99. 5% level of confidence in a one-way analysis of variance.
Cores from the lower intertidal zone contained significantly more
cells/cm 3 of sediment than cores from either the mid-intertidal or
upper intertidal zones (Figures 8-10). Conversely, cores from the
3
upper intertidal zone contained significantly fewer cells/cm of
sediment than cores from either the mid-intertidal or the lower
intertidal zones. This relationship was found during every season of
the one year sampling period. If the number of cells counted at the
43
3
Table 3. Seasonal variation in number of cells/cm of sediment. Each value is the mean of
2, 3 or 4 samples.
Southbeach
Depth in cm
Mid Intertidal
Lower Intertidal
Upper Intertidal
Summer 1967
0.0 - 1. 3
1.3 - 2.6
5.2 - 6. 5
7. 8 - 9. 1
22,533*
20, 480*
26, 960*
31, 240*
20, 200*
13, 960*
8, 226*
11, 666*
20, 380*
15, 740*
770*
380*
Fall 1967
0.0 - 1. 3
1. 3 - 2.6
5. 2 - 6. 5
7. 8 - 9. 1
20, 889
17, 609
45, 146
34, 360
32, 507
23, 827
9, 706
7, 653
17, 253
11, 160
5, 608
6, 360
Winter 1967-1968
0.0 - 1. 3
1. 3 - 2.6
5. 2 - 6. 5
7. 8 - 9. 1
46, 340
42, 280
35, 610
24, 320
42, 146
24, 413
15, 493
17, 081
36,
29,
14,
6,
43, 092
43, 546
32, 346
32, 519
42, 460
28, 700
38, 079
40, 346
25, 783
20, 639
41, 080
39, 100
5, 660
4, 080
940
030
940
380
Spring 1968
0.0 - 1. 3
1.3 - 2.6
5.2 - 6. 5
7. 8 - 9. 1
44, 540
49, 200
41, 100
36, 720
7, 500
5, 680
Summer 1968
0.0 - 1. 3
1.3 - 2.6
5.2 - 6. 5
7. 8 - 9. 1
Epipelic counts only.
48,
39,
51,
40,
520
360
760
760
60
50
Fall 1967
40
U
co
co
pI
fl)
U
30
011
20
10
Lower
Lower
Upper
Mid
Mid
Upper
Intertidal Level
Intertidal Level
3
Figure 8. Variation with intertidal level in numbers of cells/cm of sediment at Southbeach as related to season and depth in the sediment.
= 0.0-1.3 cm layer,
12 = 1.3-2.6 cm layer,
CI =5.2-6.5 cm layer,
= 7.8-9.1 cm layer.
60
60
Winter 1967-1968
Spring 1968
50
50
M
u 30
O
20
\ \\
10
10
1
1
1
Mid
Lower
Upper
Lower
Mid
Upper
Intertidal Level
Intertidal Level
3
Figure 9. Variation with intertidal level in numbers of cells/cm of sediment at Southbeach as related to season and depth in the sediment.
= 0. 0-1. 3 cm layer,
p = 1. 3-2. 6 cm layer,
0= 5.2 -6. 5 cm layer,
= 7.8 -9. 1 cm layer.
60
Summer 1968
50
40
rl
30
O
20
10
I
Lower
I
Mid
Upper
Intertidal level
3
Figure 10. Variation with intertidal level in numbers of cells/cm of sediment at Southbeach
as related to season and depth in the sediment. L. = 0.0 -1. 3 cm layer,
= 7.8 -9. 1 cm layer.
= 1. 3-2.6 cm layer, 0 = 5. 2-6. 5 cm layer,
47
various depths sampled is averaged for the entire sampling period it
is found that the lower intertidal zone averaged 36,436 cells/cm
3,
the
mid-intertidal zone averaged 24,712 cells/cm 3, and the upper inter­
tidal zone averaged 16,960 cells/cm
3.
3
The number of cells/cm at various depths in the cores showed
a significant difference (p <0.01) at the 99% level of confidence in a
3
one-way analysis of variance. The greatest number of cells/cm of
sediment was found in the upper 2. 6 cm of cores and the lower two
depths sampled in the cores had fewer cells/cm3 (Figures 11 and 12).
This relationship was especially evident in cores from the upper
intertidal zone where there is a rapid decrease in the number of cells/
cm 3 at greater core depths. If one averages the number of cells
counted throughout the entire sampling period it is found that there is
a significant decrease in the number of cells as the depth increases.
The 0.0-1.3 cm core layer below the surface averaged 35,491 cells/
cm
3,
the 1.3-2. 6 cm core layer averaged 31, 104 cells/cm 3, the
5.2-6.5 cm core layer averaged 20,481 cells/cm3, and the 7. 8-9.1
3.
cm core layer averaged 16, 802 cells/cm
No distinct seasonal trend could be ascertained at the Southbeach study area from the data available.
Mid-Intertidal
Lower Intertidal
50
50
_
en
40
s 40
-1
U
m
30
m
O
u 30
O
20
20
10
10
0. 0-1. 3
1. 3-2. 6
5. 2-6. 5
7. 8-9. 1
Depth in cm
0. 0-1. 3
1.3 -2.6
5. 2-6. 5
Depth in cm
3
Figure 11. Variation with depth in the number of cells/cm of sediment at Southbeach as related to season and position in the intertidal
= Spring 1968, 0 = Summer 1968.
zone.
P= Summer 1967, 0 = Fall 1967, 0 = Winter 1967-1968,
7. 8-9. 1
60
Upper Intertidal
en
U
U 30
0
20
. ,
10
°
0.0 -1. 3
1. 3-2. 6
5. 2 -6.5
7. 8-9. 1
Depth in cm
3
Figure 12. Variation with depth in the number of cells/cm of sediment at Southbeach as
related to season and position in the intertidal zone. L= Summer 1967,
0 = Summer 1968.
0 = Fall 1967, 0 = Winter 1967-1968,
= Spring 1968,
50
Estimate of Chlorophyll a
Southbeach Study Area
The variations in concentrations of chlorophyll a are summa­
rized in Table 4 and Appendix A, Tables 3 through 6. Table 4
contains data concerning concentrations of chlorophyll a corrected for
pheophytin on a seasonal basis from summer 1967 to summer 1968.
The estimates of chlorophyll a in pg/cm3 of sediment range from a
mean of 31.9 in summer 1968 in the 0.0-1.3 cm core layer from the
mid-intertidal zone to a mean of 0.2 in summer 1967 in the 7. 8-9. 1
cm core layer from the upper intertidal zone.
The concentrations of chlorophyll a in cores from the lower,
mid, and upper intertidal zones showed a significant difference
(p <0.01) at the 99% level of confidence in a one-way analysis of
variance. There was a definite tendency for cores from the lower
intertidal zone to have the highest concentration of chlorophyll a/cm
3
(Figures 13-17). This is especially evident in the core layers 5. 2­
6. 5 cm and 7. 8-9. 1 cm below the surface. This relationship was
not always found in the two upper layers sampled. However, if the
amount of chlorophyll a present throughout the sampling period at the
various depths is averaged it is found that the cores from the lower
intertidal zone had the highest concentration averaging 19.9 p.g/cm
the mid-intertidal zone averaged 13. 3 pg/cm
3,
3,
and the cores from the
51
3
Table 4. Seasonal variation in concentrations of chlorophyll a in µ g/cm corrected for
pheophytin content. Each value is the mean of 2, 3 or 4 samples.
Depth
in cm
Lower
Southbeach
Mid
Upper
Lower
Sally's Bend
Mid
Intertidal
Intertidal
Intertidal
Intertidal
Intertidal
Upper
Intertidal
Summer 1967
0. 0 - 1. 3
1.3 - 2.6
5. 2 - 6. 5
7. 8 - 9. 1
18.0
21.7
13.8
10.5
14.5
15. 7
5.8
7.7
13. 8
5. 2
0. 4
0. 2
5. 1
5. 9
8. 6
4. 3
4. 4
5. 1
4.0
2. 4
3.2
4. 8
3. 9
6. 1
3.6
4. 5
5.0
4. 8
4. 1
5. 2
4. 2
3.0
2. 8
1. 8
1.8
Fall 1967
0.0 - 1.3
1.3 - 2.6
5. 2 - 6. 5
7. 8 - 9. 1
20.2
21.7
18.4
11.6
13. 5
12.0
3.2
2.3
12. 2
8, 9
4. 2
4. 3
2. 8
Winter 1967-1968
0.0 - 1.3
5.2 - 6. 5
18.1
19.3
18.7
7. 8 - 9. 1
17.5
1. 3 - 2. 6
19. 7
13.9
5.2
7.2
17. 8
9. 6
5. 5
4. 2
4.0
4. 2
3. 8
3. 5
2. 8
4.0
2. 6
2. 2
2. 6
2.0
2.4
2. 8
3. 9
3. 8
3.6
3.6
2.6
3.4
4.0
Spring 1968
0.0- 1.3
1. 3 - 2. 6
5.2 - 6. 5
7. 8 - 9. 1
21.7
24.2
24.1
22.2
21.5
31.2
22. 5
24. 9
11. 1
2.4
3.9
4.0
3.2
15.9
4. 7
2. 7
3. 1
Summer 1968
0.0 - 1. 3
1. 3 - 2.6
5. 2 - 6. 5
7. 8 - 9. 1
26.5
26.7
24.3
18.8
No data available.
29.9
16.3
4. 7
3. 7
3. 7
9. 8
2. 7
2. 9
9.4
3.2
3.0
3.4
5. 4
0. 7
2. 8
3.2
2.5
31.9
23.0
*
40
Summer 1967
10
Lower
Upper
Mid
Lower
Mid
Upper
Intertidal level at Sally's Bend
Intertidal level at Southbeach
Figure 13. Variation with hit ertidal level in concentration of chlorophyll a as related to season and depth in the sediment.
P= 0. 0-1. 3 cm layer,
0 = 1. 3-2. 6 cm layer,
0 = 5. 2-6. 5 cm layer,
= 7, 8-9. 1 cm layer.
40
Fall 1967
Fall 1967
30
30
E
U
di
cc)
ta
2
0 20
20
b0
bo
10
10
\
Lower
r0
Upper
Mid
Lower
Mid
Upper
Intertidal level at Sally's Bend
Intertidal level at Southbeach
Figure 14. Variation with intertidal level in concentration of chlorophyll a as related to season and depth in the sediment.
Q = 0.0-1. 3 cm layer,
p = 1.3-2.6 cm layer,
0 = 5.2-6.5 cm layer,
= 7.8-9.1 cm layer.
Winter 1967-1968
Winter 1967-1968
30
30
8
U
8
cal
cal
U
>
LL
2
O
o 20
20
U
bO
b0
10
10
0
Lower
Upper
Mid
Intertidal level at Southbeach
0
Lower
Mid
Upper
Intertidal level at Sally's Bend
Figure 15. Variation with intertidal level in concentration of chlorophyll a as related to season and depth in the sediment
= 0.0-1.3 cm layer,
= 1.3-2.6 cm layer,
0 = 5.2-6.5 cm layer,
= 7.8-9.1 cm layer.
40
Spring 1968
30
Spring 1968
30
0,
U
U
co
co
0
0
a.
2 20
20
\
\.\
\\
\\
10
10
I
Lower
I
I
Mid
Upper
Intertidal level at Southbeach
Lower
Mid
Upper
Intertidal level at Sally's Bend
Figure 16. Variation with intertidal level in concentration of chlorophyll a as related to season and depth in the sediment.
= 7. 8-9, 1 cm layer.
p= 0.0 -1, 3 cm layer, 0 = 1. 3-2. 6 cm layer, 0 = 5. 2-6. 5 cm layer,
40
Summer 1968
Summer 1968
30
30
10
10
Lower
Mid
Intertidal level at Southbeach
Upper
Lower
Mid
Upper
Intertidal level at Sally's Bend
Figure 17. Variation with intertidal level in concentration of chlorophyll a as related to season and depth in the sediment.
= 1. 3-2. 6 cm layer, o = S. 2 -6.5 cm layer,
o = O. 0-1. 3 cm layer,
= 7. 8-9. 1 cm layer.
57
upper intertidal zone averaged the lowest with 10.1 pg/cm 3.
The concentrations of chlorophyll a at various depths in the
cores showed a significant difference (p <0. 01) at the 99% confidence
level.
The greatest concentration of chlorophyll a was in the upper
core layer of sediment (Figures 18-20). This relationship was
especially clear in cores from the upper intertidal zone where there
is a rapid decrease in chlorophyll a at greater depths. If the amount
of chlorophyll a present throughout the entire sampling period is
averaged it is found that there is a significant decrease in the amount
of chlorophyll a as the depth increases. The 0.0-1.3 cm core layer
averaged 20.7 µg /cm3, the 1.3-2.6 cm core layer averaged 17.7 pg/
cm
3,
the 5 2-6.5 cm core layer averaged 10.0 pg/cm 3, and the 7.8­
9. 1 cm core layer averaged 9. 3 pg/cm 3.
No distinct seasonal trend could be established at the Southbeach
study area based on variations in the concentration of chlorophyll a.
Sally's Bend Study Area
The concentrations of chlorophyll a are summarized in Table
4 and in Appendix A, Tables 3 through 6. Table 4 contains data
concerning the concentrations of chlorophyll a corrected for the
percentage of pheophytin on a seasonal basis from summer 1967 to
summer 1968. The estimates of chlorophyll a in µg /cm3 of
sediment range from a mean of 8.6 in summer 1967 in the 0.0-1.3
40
40
Lower Intertidal
Lower Intertidal
30
30
ci
U
cs
o 20
O
10
10
0.0-1.3
1.3-2.6
5.2-6.5
Depth in cm at Southbeach
7.8-9.1
0.0-1.3
1.3-2.6
5.2-6.5
7.8-9.1
Depth in cm at Sally's Bend
Figure 18. Variation with depth in the concentration of chlorophyll a in the sediments as related to position in the intertidal zone and season.
p = Summer 1967,
= Fall 1967, 0 = Winter 1967-1968,
= Spring 1968, 0 = Summer 1968.
40
40
Mid-Intertidal
Mid-Intertidal
30
30
U
U
cal
a.
0 20
20
1)0
ao
10
10
1
0.0-1.3
1.3-2.6
5.2-6.5
Depth in cm at Southbeach
7.8-9.1
0.0-1.3
1.3-2.6
1
1
5.6-6.5
7. 8-9.1
Depth in cm at Sally's Bend
Figure 19. Variation with depth in the concentration of chlorophyll a in the sediments as related to position in the intertidal zone and season.
p= Summer 1967, 0 = Fall 1967, 0 = Winter 1967-1968,
= Spring 1968, 0 = Summer 1968.
Upper Intertidal
Upper Intertidal
30
U
tot
Nit
0ti
o
o
"9
20
cU
U 20
00
-b0
10
0.0-1.3
1.3-2,6
5.2-6,5
Depth in cm at Southbeach
7.8-9.1
0.0 -1.3
1.3-2.6
5,2-6.5
7.8-9.1
Depth in cm at Sally's Bend
Figure 20. Variation with depth in the concentration of chlorophyll a in the sediments as related to position in the intertidal zone and season.
= Spring 1968, 0 = Summer 1968.
Summer 1967, 0 = Fall 1967, 0 = Winter 1967-1968,
61
cm core layer from the upper intertidal zone to a mean of 0.7 in
summer 1968 in the 1.3-2.6 cm core layer from the upper intertidal
zone.
No significant difference in the concentrations of chlorophyll a
in cores from the lower, mid, and upper intertidal zones was shown
by a one-way analysis of variance. There was no significant increase
or decrease in chlorophyll a from the lower intertidal to the upper
intertidal zone (Figures 13-17).
However, the concentrations of chlorophyll a at various depths
in the cores showed a significant difference (p <0.10) at the 90%
confidence level. Figures 18 through 20 depict the estimated amounts
of chlorophyll a at each depth sampled at every season. Averages
of the amounts of chlorophyll a present throughout the entire sampling
period show decreasing amounts of chlorophyll a as the depth increases.
The 0.0-1.3 cm core layer below the surface averaged 4.7 pg/cm 3,
the 1.3-2.6 cm core layer averaged 3.7 pg/cm3, the 5.2-6.5 cm core
layer averaged 3.3 pg/cm 3, and the 7.8-9.1 core layer averaged 2.9
pg/cm3.
The data does not indicate the presence of any distinct seasonal
pattern at the Sally's Bend station as far as chlorophyll a concentration
is concerned.
62
Comparison of the Concentrations of
Chlorophyll a at the Two Study Areas
The cores from the lower intertidal, mid-intertidal, and upper
intertidal zones of the Southbeach study area all had significantly
greater (p < 0. 005) concentrations of chlorophyll a than corresponding
cores from the Sally's Bend study area at the 99. 5% level of confidence
in a t-test. This relationship held for every season at each tidal
level (Table 4, Figures 13-17). If the entire sampling period is taken
into consideration and the chlorophyll a content averaged at each
intertidal level for each study area one finds that the Southbeach lower
intertidal level cores averaged 19. 9 lag/cm 3 as compared to an
estimated average of 4.0 p.g/cm 3 at Sally's Bend, the Southbeach
mid-intertidal level cores averaged 13.3 pg/cm 3 as compared to 3.5
3
p.g/cm at Sally's Bend, and the Southbeach upper intertidal level
cores averaged 10.0 pg/cm 3 as compared to an estimated average of
3.3 pg/cm3 at Sally's Bend.
The concentrations of chlorophyll a at various depths in cores
from the Southbeach study area were significantly greater (p <0.005)
than concentrations from the corresponding core depths from the
Sally's Bend study area at the 99. 5% confidence level in a t-test. This
relationship held for every season of the sampling period (Figures
18-20).
If the entire sampling period is taken into consideration it is
found that the 0.0-1.3 cm core layer at Southbeach averaged 20.7
63
3
p.g/cm of chlorophyll a as compared to an average of 4.7 pg/cm
3
at
Sally's Bend, the 1.3-2. 6 cm core layer at Southbeach averaged 17. 7
3
p.g/cm as compared to 3.7 pg/cm3 at Sally's Bend, the 5.2-6.5 cm
3
,
core layer at Southbeach averaged 10. 0 p.g/ cm as compared to 3. 3
p.g/ cm
3
at Sally's Bend, and the 7. 8-9.1 cm core layer at Southbeach
3
3
averaged 9. 3 p.g/cm as compared to 2.9 pg/cm at Sally's Bend.
Determination of the Percentage of Pheophytin
Southbeach Study Area
The percentages of pheophytin or chlorophyll degradation
pigment in cores collected during the sampling period are summarized
in Table 5 and in Appendix A, Tables 7 through 10. The percentage
of pheophytin present in the core samples range from a mean of 87%
in summer 1967 in the 7. 8-9. 1 cm core layer from the upper inter­
tidal zone to a mean of 1. 5% in summer 1967 in the 1. 3-2. 6 cm core
layer from the lower intertidal zone.
The percentages of pheophytin in cores from the lower, mid,
and upper intertidal zones showed a significant difference (p <0. 01)
at the 99% level of confidence in a one-way analysis of variance. A
definite relationship existed between the position in the intertidal
zone and the percentage of pheophytin present (Figures 21 through 25).
The highest percentage of pheophytin was located in cores from the
upper intertidal zone and cores from the lower intertidal zone had the
64
Table 5. SeasonaLvariation in pheophytin expressed as percentage of total chlorophyll
(chlorophyll a and pheophytin). Each value is the mean of 2, 3, or 4 samples.
Depth
Lower
Southbeach
Mid
Upper
Lower
Sally's Bend
Mid
Upper
in cm
Intertidal
Intertidal
Intertidal
Intertidal
Intertidal
Intertidal
31.0
40.0
73.0
67.6
64.0
62.0
53. 7
63. 0
87. 0
64. 7
58.8
57.5
54.7
58.6
60. 1
57. 3
56. 3
66.0
53. 8
59.1
59.1
61.4
62.5
60.0
60. 7
72.1
60. 5
57.0
57.0
67.0
Summer 1967
0.0 - 1.3
1.3 - 2.6
12.0
5.2 - 6. 5
8.0
10.0
7. 8 - 9. 1
1. 5
13.0
17.5
33.0
27.0
70.0
74. 5
Fall 1967
0.0 - 1.3
2. 7
1. 3 - 2. 6
5. 2 - 6. 5
7. 8 - 9. 1
4.5
4.8
15.5
14.3
12.3
46.6
53.6
22.0
18. 8
41.6
51.0
Winter 1967-1968
0.0- 1.3
4.1
3.9
2.0
5.6
1. 3 - 2. 6
5. 2 - 6. 5
7. 8 - 9. 1
11.0
11.0
38.2
30.2
5. 9
14. 4
43. 5
36. 8
58. 3
59. 1
64. 3
70. 3
67. 1
60. 5
61.4
61.6
59. 5
59. 5
62. 6
65. 0
58.6
59.7
59.6
64.3
60. 5
61.0
59.0
63.0
62.5
61.0
57.0
87.0
Spring 1968
0.0 - 1.3
1.3 - 2.6
11.5
5.2 - 6. 5
11.2
16.0
7. 8 - 9. 1
6.0
14, 5
12. 5
19. 5
9.0
25.2
18.5
47.7
27.2
63. 6
58. 8
65. 1
Summer 1968
0.0 - 1. 3
28. 5
29. 5
10. 7
1.3 - 2.6
23.0
15.2
5.2 - 6. 5
7.8 - 9. 1
11. 7
20.0
27.0
22.0
*No
data available.
6.2
48. 5
44.7
*
63. 3
64.3
67. 5
66. 3
90
90
Summer 1967
Summer 1967
80
80
70
70
/
//
I/
/
60
/,
50
/
/
40
/
30
20
30
,41
20
G/'//
10
10
Lower
Mid
Upper
Lower
Mid
Upper
Intertidal Level at Sally's Bend
Intertidal Level at Southbeach
Figure 21. Variation with intertidal level in pheophytin expressed as percentage of total chlorophyll (chlorophyll a and pheophytin) as
related to season and depth in the sediment.
= 7. 8-9. 1 cm layer,
p = 0,0-1, 3 cm layer,
0 = 1. 3-2.6 cm layer,
0 = 5. 2-6. 5 cm layer,
80
80
Fall 1967
70
70
Fall 1967
60
60
_
-4)
50
50
40
40
30
30
20
20
10
10
a.
Lower
Mid
Upper
Lower
Mid
Upper
Intertidal Level at Sally's Bend
Intertidal Level at Southb each
Figure 22. Variation with intertidal level in pheophytin expressed as percentage of total chlorophyll (chlorophyll a and pheophytin) as
related to season and depth in the sediment.
= 7. 8 -9. 1 cm layer.
Q = 0.0-1. 3 cm layer,
0 = 1. 3-2.6 cm layer,
0 = 5. 2-6. 5 cm layer,
80
80
Winter 1967-1968
Winter 1967-1968
70
70
60
60
50
40
30
20
0
10
10
Lower
Mid
Upper
Lower
Upper
Mid
Intertidal Level at Sally's Bend
Intertidal Level at Southbeach
Figure 23. Variation with intertidal level in pheophytin expressed as percentage of total chlorophyll (chlorophyll a and pheophytin) as
related to season and depth in the sediment.
= 7.8-9.1 cm layer.
A = 0.0-1.3 cm layer,
= 1.3-2.6 cm layer,
0 = S. 2-6.5 cm layer,
80
80
Spring 1968
Spring 1968
70
70
60
60
11- ------ --o- ----­
.6.-----­
0---7-----­
-11
---10in,
50
50
ra
.-.
Pi
..0
.0
a.
o 40
a.
oa) 40
xa.a,
a.
g
c`'
30
30
20
20
10
10
I
I
Lower
Mid
L._
I
I
Upper
1
Lower
1
Upper
Mid
Intertidal Level at Sally's Bend
Intertidal Level at Southb each
Figure 24. Variation with intertidal level in pheophytin expressed as percentage of total chlorophyll (chlorophyll a and pheophytin) as
related to season and depth in the sediment.
= 7.8-9.1 cm layer.
A = 0.0 -1. 3 cm layer,
p = 1.3-2.6 cm layer,
0 = 5.2-6.5 cm layer,
90
90
0
Summer 1968
Summer 1968
80
80
70
70
60
60
50
50
40
40
30
30
20
20
10
10
mr
Lower
Mid
Upper
Lower
Mid
Upper
Intertidal Level at Sally's Bend
Intertidal Level at Southbeach
Figure 25. Variation with intertidal level in pheophytin expressed as percentage of total chlorophyll (chlorophyll a and pheophytin) as
related to season and depth in the sediment.
= 7.8-9.1 cm layer.
p = 0.0-1.3 cm layer,
= 1.3-2.6 cm layer,
0 = 5.2-6.5 cm layer,
70
lowest percentage of pheophytin. If the entire sampling period is
considered one finds that the upper intertidal cores averaged 35. 1%
pheophytin, the mid-intertidal cores 23. 1 %, and the lower intertidal
cores 9.4%.
The percentage of pheophytin at various depths in the cores also
showed a significant difference (p <0.001) at the 99.9% level of
confidence in a one-way analysis of variance. The highest percentage
of pheophytin was in the 5.2-6.5 cm and 7. 8-9. 1 cm core layers
(Figures 26 through 28). This was especially evident in cores from
the mid-intertidal and upper intertidal zones whereas the cores from
the lower intertidal zone did not show this relationship. On the basis
of the entire sampling season the average percentage of pheophytin
present in the 0. 0-1. 3 cm core layer was 15.7%, in the 1.3-2. 6 cm
core layer 14. 6%, in the 5.2-6.5 cm core layer 30.4%, and in the
7. 8-9.1 cm core layer 29. 4 %.
Sally's Bend Study Area
The percentage of pheophytin in cores collected during the
sampling period are summarized in Table 5 and in Appendix A,
Tables 7 through 10. The percentage of pheophytin present in the
core samples from Sally's Bend ranged from a mean of 87% in
summer 1968 in the 1.3-2. 6 cm core layer from the upper intertidal
zone to a mean of 53.7% in summer 1967 in the 0. 0-1. 3 cm core layer
80
Lower Intertidal
70
60
41 50
a.
a)
a. 40
30
20
10
0.0 -1. 3
1. 3 -2.6
5. 2-6. 5
Depth in cm at Southbeach
7. 8-9. 1
0. 0-1. 3
5. 2-6. 5
Depth in cm at Sally's Bend
1. 3 -2.6
7. 8-9. 1
Figure 26. Variation with depth in pheophytin expressed as percentage of total chlorophyll (chlorophyll a and pheophytin) in the sediments
as related to the position in the intertidal zone and season. A = Summer 1967, 0= Fall 1967, 0 = Winter 1967-1968,
= Spring 1968,
0 = Summer 1968.
80
80
Mid-Intertidal
Mid-Intertidal
70
70
60
60
/171
V. 50
0
50
0
a.
a.
a.
4°
40
30
30
//1
20
20
10
10
0. 0-1. 3
5. 2-6, 5
1.3 -2. 6
Depth in cm at Southbeach
7. 8-9. 1
0. 0-1. 3
1. 3-2. 6
5. 2-6. 5
7. 8-9. 1
Depth in cm at Sally's Bend
Figure 27. Variation with depth in pheophytin expressed as percentage of total chlorophyll (chlorophyll a and pheophytin) in the sediments as
= Summer 1967, CI = Fall 1967, 0 = Winter 1967-1968,
related to the position in the intertidal zone and season.
0 = Spring 1968, 0 = Summer 1968.
90
90
Upper Intertidal
80
70
60
4ra, 50
40
30
20
10
0. 0-1. 3
1.3-2.6
Depth in cm at Southbeach
7. 8-9, 1
0.0 -1. 3
1,3-2.6
5. 2-6. 5
7.8 -9. 1
Depth in cm at Sally's Bend
Figure 28. Variation with depth in pheophytin expressed as percentage of total chlorophyll (chlorophyll a and pheophytin) in the sediments as
related to the position in the intertidal zone and season. A = Summer 1967, 0= Fall 1967, 0 = Winter 1967-1968,
= Spring 1968, 0 = Summer 1968.
74
from the upper intertidal zone.
The percentages of pheophytin in cores from the lower, mid,
and upper intertidal zones showed a significant difference (p <O. 01)
at the 99% level of confidence in a one-way analysis of variance. The
highest percentage of pheophytin was located in cores from the upper
intertidal zone, this percentage being slightly greater than that from
the mid-intertidal and lower intertidal zones (Figures 21 through 25).
If the entire sampling period is considered, one finds that cores from
the upper intertidal zone had an average of 65. 3% pheophytin, the
mid-intertidal 59. 5 %, and the lower intertidal 61. 1%.
The percentages of pheophytin at various depths in the cores
also showed a significant difference (p <0.025) at the 97.5% level of
confidence in a one-way analysis of variance. The 5.2-6.5 and 7. 8­
9. 1 cm core layers had a slightly higher percentage of pheophytin
(Figures 26 through 28). On the basis of the entire sampling period
the average percentage of pheophytin present in the O. 0-1. 3 cm core
layer was 60. 5%, in the 1. 3-2. 6 em..--cort +titriit it was 57.9 %, in the
5.2-6.5 cm core layer 62. 1%, and in the 7. 8-9.1 cm core layer
64. 6 %.
Comparison of Percentage of Pheophytin
at the Two Study Areas
The cores from the lower intertidal, mid-intertidal, and upper
intertidal zones of the Sally's Bend study area all had significantly
75
greater (p <0.005) percentages of pheophytin than corresponding cores
from the Southbeach study area at the 99. 5% level of confidence in a
t-test (Table 5, Figures 21-25). When the entire sampling period is
taken into consideration and the percentage of pheophytin is averaged
for each intertidal level at each study area, the cores from the Sally's
Bend lower intertidal zone are found to have had 61. 1% compared to
9.4% for Southbeach, cores from the Sally's Bend mid-intertidal
zone 59. 5% as compared to 23. 1% at Southbeach, and cores from the
Sally's Bend upper intertidal zone 65.3% as compared to 35. 1% at
Southbeach.
The percentages of pheophytin at various depths in the cores
from the Sally's Bend study area were significantly greater (p <
0. 005) than percentages from the corresponding core depths from the
Southbeach study area at the 99. 5% level of confidence in a t-test.
The percentages of pheophytin found at each depth in the intertidal zone
at each study area are compared graphically in Figures 26 through 28.
When the values for percentages of pheophytin for the entire sampling
period are averaged it is found that in the 0.0-1.3 cm core layer at
Sally's Bend 60.5% of pheophytin is found compared to 15.7% at
Southbeach, in the 1.3-2.6 cm core layer 57.9% pheophytin at Sally's
Bend as compared to 14. 6% at Southbeach, in the 5.2-6.5 cm core
layer 62. 1% at Sally's Bend as compared to 30.4% at Southbeach, and
in the 7.8-9.1 cm core layer 64. 6% at Sally's Bend as compared to
76
29. 4% at Southbeach.
Experimental Gross Production of Oxygen
Oxygen production by the autotrophic organisms in each core
layer was estimated using the Gilson Differential Respirometer.
Results are expressed as pg 0 2 /hr/cm 3 which represents the potential
rates of gross production of the plants in the various core layers at
the experimental level of illumination intensity.
Southbeach Study Area
The data for the potential rates of gross production for the
various seasons during the sampling period are summarized in Table
6 and in Appendix A, Tables 11 through 14.
The estimates of potential
gross production of oxygen range from 188.4 tig/hr/cm 3 in summer
1968 in the 0.0-1.3 cm core layer from the lower intertidal zone to
-1.4 pg/hr/cm3 in summer 1968 in the 5.2-6.5 cm core layer from the
upper intertidal zone.
The rates of potential oxygen production in cores from the lower,
mid, and upper intertidal zones showed a significant difference (p <
0. 01) at the 99% confidence level in a one-way analysis of variance.
Cores from the lower intertidal zone had a higher potential oxygen
production than cores from either the mid-intertidal or upper inter­
tidal zone and the cores from the upper intertidal zone produced
77
Table 6. Seasonal - variation in potential gross production expressed as p g 02/cm
Each value is the mean of 2, 3, or 4 samples,
3/hr.
Sally's Bend
Southbeach
Depth
Lower
Mid
Upper
Lower
in cm
Intertidal
Intertidal
Intertidal
Intertidal
Intertidal
20.7
Mid
Upper
Intertidal
Summer 1967
0.0 - 1.3
1. 3 - 2. 6
5. 2 - 6. 5
7. 8 - 9. 1
101.4
85.9
50.9
42.4
81.6
86.9
29.1
19.8
66.9
14.0
- 5. 1
2. 5
S. 0
5.6
- O. 8
74.2
44.3
44.8
32.7
36. 8
31.2
3. 8
-11.4
Fall 1967
0.0 - 1.3
1.3 - 2.6
124.6
76. 8
97.9
- 7.7
41.0
59.4
170. 1
55. 5
- 4.7
108.3
44.0
-21.6
-10.4
17. 4
5. 2 - 6. 5
7. 8 - 9. 1
75.0
33.8
17.3
22.3
13.5
13.9
22.9
42.6
44.9
20.6
14.6
4.6
41.6
22.4
- 2.3
Winter 1967-1968
0.0 - 1.3
1.3 - 2.6
129.5
133.9
5. 2 - 6. 5
7. 8 - 9.1
88.4
64.4
97.6
107. 3
34, 8
107.3
62.3
33.0
42.3
40, 5
-10.1
-15.0
-14, 2
27. 1
27, 3
- 3, 5
14.9
-33.6
-38.7
-11.2
-16.9
48.4
27.1
- 51.3
Spring 1968
0.0 - 1.3
1.3 - 2.6
5.2 - 6. 5
7.8 - 9.1
137.2
197.2
108.2
114.7
153.8
135.7
156. 5
79. 4
59, 9
8.7
84.9
15. 4
11.8
7.0
66.2
6.6
-11.9
9, 1
- 7.0
51.8
40.8
23.8
31.9
40.4
23.2
Summer 1968
0.0 - 1.3
1.3 - 2.6
5.2 - 6.5
7. 8 - 9. 1
188.4
130.5
121.0
87.5
170.3
106.1
51.0
45.4
148. 8
69. 5
-
1.4
27.6
5. 1
- 4.0
- 0.7
- 8.0
78
significantly less oxygen than either the lower intertidal or midintertidal zone cores (Figures 29-33). This was especially evident in
the 1. 3-2. 6 cm, 5.2-6.5 cm, and 7. 8-9.1 cm core layers. Occasion­
ally the 0.0-1. 3 cm layer had a higher oxygen production in cores from
the upper intertidal zone than this layer from the mid-intertidal zone.
Averages of the amount of oxygen produced per hour during the
entire sampling period show that the lower intertidal zone cores pro­
duced the largest amounts of oxygen of 114. 1 pg /hr/cm3 and the
upper intertidal cores had the lowest oxygen production of 50. 1
µg /hr /cm3.
The rates of potential oxygen production at various depths in
the cores also showed a significant difference (p < 0. 01) at the 99%
confidence level in a one-way analysis of variance. The highest rate
3
of oxygen production was by the upper cm of sediment and there was
a significant decrease as the depth increased (Figures 34-36).
The
1. 3-2. 6 cm core layer from the lower intertidal zone occasionally had
a higher rate of oxygen production than the 0. 0-1. 3 cm core layer.
If the rates of oxygen production are averaged for the entire sampling
period one finds that the upper cm 3 was most productive with 121. 3
p.g/hr, the 1. 3-2. 6 cm core layer produced 72.2 pg/hr, the 5.2-6.5
cm core layer produced 52. 6 pg/hr, and the 7. 8-9. 1 cm core layer
produced 35. 8 pg/hr.
200
200
Summer 1967
160
Summer 1967
160
120
120
80
80
en
U
O
40
40
bo
0
0
-40
-40
I
Lower
Mid
Upper
Lower
Intertidal Level at Southbeach
Mid
Upper
Intertidal Level at Sally's Bend
Figure 29. Variation with intertidal level in potential gross production of oxygen as related to season and depth in the sediment.
= 0.0-1.3 cm layer,
= 1.3-2.6 cm layer,
0 = 5.2-6.5 cm layer,
= 7.8-9.1 cm layer.
200
200
Fall 1967
Fall 1967
160
Ni
120
80
160
:IN
120
N..... N.
.z`"
m---- 8 0
----__
E
0
__- -0
II- .....
'e" --------
--­
0" 40
b0
=­
0
-40
-40
Lower
Mid
Upper
Lower
Mid
Upper
Intertidal Level at Sally's Bend
Intertidal Level at Southbeach
Figure 30. Variation with intertidal level in potential gross production of oxygen as related to season and depth in the sediment.
= 0.0-1.3 cm layer,
0= 1.3-2.6 cm layer,
0 = 5.2-6.5 cm layer,
= 7.8-9.1 cm layer.
200
200
Winter 1967-1968
Winter 1967-1968
160
160
120
120
0 80
80
cri
0
oN
b0
b0
40
40
0
0
-40
-40
Lower
Mid
Upper
Lower
Intertidal Level at Southbeach
Upper
Mid
Intertidal Level at Sally's Bend
Figure 31. Variation with intertidal level in potential gross production of oxygen as related to season and depth in the sediment.
= 0.0-1.3 cm layer,
0 = 1.3-2.6 cm layer, 0 = 5.2-6.5 cm layer,
= 7.8-9.1 cm layer.
200
EJ
200
Spring 1968
Spring 1968
160
160
120
120
80
E
80
ty
IND
0
"s.
O
IUD
40
40
0
0
-40
-40
I
Lower
Upper
Mid
Intertidal Level at Southbeach
Mid
Lower
Upper
Intertidal Level at Sally's Bend
Figure 32. Variation with intertidal level in potential gross production of oxygen as related to season and depth in the sediment.
= 0.0-1.3 cm layer,
= 1.3-2.6 cm layer,
p = 5.2-6.5 cm layer,
= 7.8-9.1 cm layer.
200
160
r.
120
en
0
-40
Lower
Mid
Intertidal Level at Southbeach
Figure 33.
Upper
Lower
Mid
Upper
Intertidal Level at Sally's Bend
Variation with intertidal level in potential gross production of oxygen as related to season and depth in the sediment.
= 1.3 -2. 6 cm layer, 0 = 5. 2 -6.5 cm layer,
=7. 8-9. 1 cm layer.
p = 0.0 -1. 3 cm layer,
84
Sally's Bend Study Area
The data for rates of gross oxygen production are summarized
in Table 6 and Appendix A, Tables 11 through 14. The estimates of
gross oxygen production ranged from 74.2 µg /hr /cm3 in summer
1967 in the 0.0-1.3 cm core layer from the mid-intertidal zone to
-51.3 µg /hr/cm3 in summer 1968 in the 1.3-2.6 cm core layer from
the lower intertidal zone.
The rates of potential oxygen production in cores from the lower,
mid, and upper intertidal zones showed a significant difference (p
0.001) at the 99.9% confidence level in a one-way analysis of variance.
Less oxygen was produced by cores from the lower intertidal zone
than from either the mid-intertidal or upper intertidal zone (Figures
29 through 33). Generally more oxygen was produced by cores from
the mid-intertidal level than from the upper intertidal level. This
relationship was especially evident atthe three lower depths.
Occasionally, the upper cm 3 of the cores from the upper intertidal
zone produced more oxygen. If one averages the rate of oxygen
production during the entire sampling period, the cores from the lower
intertidal zone produced -8.3 µg /hr /cm3, cores from the mid-intertidal
produced 31.3 pg/hr/cm3, and cores from the upper intertidal zone
produced 17.3 µg /hr /cm3.
The rates of potential oxygen production at various depths in the
85
cores also showed a significant difference (p <0.001) at the 99. 9%
confidence level in a one-way analysis of variance. The highest rate
3
of oxygen production was by the upper cm of sediment and the 7. 8­
9. 1 cm core layer usually had the lowest rate (Figures 34 through
36).
Production by the 1. 3-2. 6 and the 5.2-6.5 cm core layers varied
depending on whether the cores were from the lower, mid, or upper
intertidal levels. In cores from the lower intertidal zone the 1. 3-2. 6
cm core layer produced less oxygen than the 5. 2-6.5 cm core layer,
but in the cores from the mid-intertidal zone the production of these
two cores were almost the same. In cores from the upper intertidal
zone the 5.2-6.5 cm core layer produced less oxygen than the 1. 3-2. 6
cm core layer, If the rates of oxygen production are averaged for the
entire sampling period the 0. 0-1. 3 cm core layer is found to have
produced 33.5 pg/hr, the 1. 3-2. 6 cm core layers 5. 8 µg /hr, the
5.2-6.5 cm core layers 9. 4 p.g/hr, and the 7. 8-9.1 cm core layers
5. 0 p.g/hr.
Comparison of Potential Production of
Oxygen at the Two Study Areas
The Southbeach study area had a much higher potential for
oxygen production than the Sally's Bend study area (Table 6).
The cores from the lower intertidal, mid-intertidal, and upper
intertidal zones of the Southbeach study area all had significantly
greater (p <.0. 005) rates of potential production than corresponding
200
."
..,
,
,
160
.
.../
.....
.
.s.
..,
Lower Intertidal
Lower Intertidal
N.
\
-­
120
120
M
m
0 80
N
O
80
0
0(.1
b°
=.
40
0
-40
I
0. 0-1. 3
I
I
1.3 -2. 6
5.2 -6. 5
Depth in cm at Southbeach
I
7. 8-9. 1
0. 0-1. 3
5. 2-6. 5
1. 3-2. 6
Depth in cm at Sally's Bend
Figure 34. Variation with depth in potential gross production of oxygen as related to position in the intertidal zone and season.
0 = Summer 1968.
= Spring 1968,
= Fall 1967, 0 = Winter 1967-1968,
A = Summer 1967,
7. 8-9. 1
200
200
Mid-Intertidal
160
160
120
120
.
80
0
0eq
Mid-Intertidal
.........
...... -0
80
en
U
40
bO
0
0
-40
-40
0. 0 -1. 3
5. 2 -6.5
1.3 -2.6
Depth in cm at Southbeach
7.8-9. 1
0. 0-1. 3
1. 3 -2.6
5. 2-6. 5
Depth in cm at Sally's Bend
Figure 35. Variation with depth in potential gross production of oxygen as related to position in the intertidal zone and season.
A= Summer 1967, 0 = Fall 1967, 0 = Winter 1967-1968,
= Spring 1968, 0 = Summer 1968.
7. 8-9. 1
200
200
Upper Intertidal
160
Upper Intertidal
160
120
120
80
ON
40
to 40
0
0
-40
-40
1
0.0-1.3
I
1.3-2.6
I
5.2-6.5
Depth in cm at Southbeach
l
7.8-9.1
i
0.0-1.3
1.3-2.6
5.2-6.5
Depth in cm at Sally's Bend
Figure 36. Variation with depth in potential gross production of oxygen as related to position in the intertidal zone and season.
=Spring 1968, Q = Summer 1968.
A = Summer 1967, 0 = Fall 1967, 0 = Winter 1967-1968,
l
7.8-9.1
89
cores from the Sally's Bend study area at the 99.5% level of confi­
dence in a t-test. The lower intertidal zone at the Southbeach study
area always had a higher potential rate of oxygen production than
this zone at Sally's Bend (Figures 29-33). The yearly average poten­
tial production of oxygen by cores from Southbeach was 114.4 µg /hr/
3
cm whereas the yearly average production at Sally's Bend was -8. 3
µg /hr /cm3. Cores from the mid-intertidal and upper intertidal zones
from the Southbeach study area also had higher potential rates of
oxygen production than cores from the Sally's Bend study area. The
yearly average potential production at Southbeach in cores from the
mid-intertidal zone was 73. 9 µg /hr/cm3 as compared to 31.3 pg/hr/
cm
cm3ini cores from Sally's Bend and in upper intertidal cores from
Southbeach the yearly average was 50. 1 µg /hr/cm3 as compared to
17. 3 µg /hr /cm3 for Sally's Bend.
The rates of potential production at various depths in the
cores from the Southbeach study area were significantly greater
(p <0. 005) than rates from the corresponding core depths from the
Sally's Bend study area at the 99. 5% level of confidence in a t-test.
The rates of potential production found at each depth in the intertidal
zone at each study area are compared graphically in Figures 34
through 36.
If yearly averages are compared one finds that the 0. 0-1.3 cm
core layer from Southbeach produced 121. 3 dig/hr while core layers
90
from Sally's Bend produced 33.5 pg/hr. The 1.3-2. 6 cm core layer
from Southbeach produced 72.2 pg /hr compared to 5. 8 pg/hr from the
Sally's Bend cores, the 5.2-6.5 cm core layer produced 52. 6 pg/hr
as compared to 9 .4 p.g/hr at Sally's Bend and the 7. 8-9. 1 cm core
layer produced 35. 8 pg/hr as compared to 5.0 µg /hr at Sally's Bend.
Analysis of Ash-Free Dry Weight
Values obtained by combusting the organic matter in the core
layers are recorded in Table 7 and Appendix A, Tables 15 through 18.
No significant difference in ash-free dry weight from the lower, mid,
or upper intertidal zones or from the various depths in the cores from
the same study area was shown by a one-way analysis of variance.
However, a significant difference (p <0.005) between the
organic matter content of the cores from the Sally's Bend and the
Southbeach study areas was shown at the 99. 5% level of confidence in
a t-test. In all core layers examined, the Southbeach study area had a
lower organic matter content than cores from the Sally's Bend study
area (Figures 37 through 44).
Taxonomic Study
One hundred and fifty-four species and varieties of diatoms
representing 56 genera were identified from the two study areas
(Table 8). The largest number of species was found at the Southbeach
91
3
Table 7. Seasonal variation in ash-free dry weight or organic matter in mg/cm of sediment.
Each value is the mean of 2, 3, or 4 samples.
Depth
Lower
in cm
Intertidal
Southbeach
Mid
Intertidal
Sally's Bend
Upper
Intertidal
Lower
Mid
Upper
Intertidal
Intertidal
Intertidal
Summer 1967
0.0 - 1.3
36.1
51.6
41.0
144.8
168.7
379.9
1. 3 - 2. 6
32.7
40.3
32.0
46.4
42. 1
29. 2
59. 8
200. 7
166, 3
190. 9
163. 1
145.2
206.7
245. 4
410. 2
210. 4
106.7
83.4
36.5
75.0
51.6
89.9
100.2
79.3
101.1
113.7
111. 7
108. 5
92. 7
107. 5
104. 7
27. 7
33. 3
80.6
86. 3
100. 1
98. 1
46.8
47.8
94, 0
111.1
84. 4
107. 8
105. 9
91.9
105.9
114.3
39. 0
89. 2
105. 4
45.1
45.1
52.4
94.2
121.3
108.2
102.2
115.7
103.9
99. 9
115. 7
72.7
82.2
96.3
105.0
99.2
5.2 - 6. 5
7. 8 - 9. 1
43.1
41. 1
Fall 1967
0.0 - 1.3
1. 3 - 2. 6
5. 2 - 6. 5
7. 8 - 9. 1
102.0
28.9
46.1
72.0
33.0
55.3
48. 1
108. 5
102.6
Winter 1967-1968
0.0 - 1.3
37, 5
1. 3 - 2. 6
5.2 - 6. 5
26.8
36.6
49.8
28.3
42.0
7. 8 - 9. 1
37, 0
38. 5
Spring 1968
0, 0 - 1.3
1. 3 - 2. 6
5.2 - 6.5
7. 8 - 9. 1
33.6
31.8
41.3
33.1
38.7
29.3
41.7
35.3
154.2
110.9
Summer 1968
0.0 - 1.3
1. 3 - 2. 6
5. 2 - 6. 5
7. 8 - 9. 1
32.8
27.0
33.7
28.8
39.7
25.4
42.5
30.9
37.5
33.5
37.5
47.7
57, 8
95.1
96.3
86.4
90.2
85.0
105. 7
450
Summer 1967
rn
400
400
350
350
x300
8 300
yxY
pQ
a 250
250
r
0 200
eo
200
ISO
150
100
100
50
SO r-
Mid
Upper
Intertidal Level at Southbeach
Lower
Lower
Figure 37. Variation with intertidal level in organic matter as related to season and depth in the sediment.
0 = 1.3 -2.6 cm layer,
p = 5.2-6. 5 cm layer,
Upper
Mid
Intertidal Level at Sally's Bend
= 7.8 -9.1 cm layer.
A= 0.0 -1.3 cm layer,
40°
400
Fall 1967
350
350
300
300
250
a 250
200
I
Fall 1967
200
eo
Ii
150
150
100
100
50
50
Lower
Mid
Upper
Lower
Intertidal Level at Southbeach
Figure 38. Variation with intertidal level in organic matter as related to season and depth in the sediment.
0 = 1.3 -2.6 cm layer,
p = 5.2 -6.5 cm layer,
Upper
Mid
Intertidal Level at Sally's Bend
= 7.8-9,1 cm layer.
L
:!1. 0-1. 3 cm layer,
W inter 1967-1968
Winter 1967-1968
350
350
u 300
u 300
yyd
yM
te)
F
aM
250
250
a
a
to 200
I* 200
150
150
100
100
50
Lower
Mid
Upper
Intertidal Level at Southbeach
Lower
Upper
Mid
Intertidal Level at Sally's Bend
Figure 39. Variation with intertidal level in organic matter as related to season and depth in the sediment.
= 7. 8-9. 1 cm layer,
0 = 1. 3-2. 6 cm layer, o = 5. 2-6. 5 cm layer,
A = 0.0-1. 3 cm layer,
400
400
Spring 1968
Spring 1968
350
350
on, 300
300
m
cd 250
as
250
200
IND
;.4
O 200
1:0
150
150
100
100
50
50
_L
Lower
I
Upper
Mid
Intertidal Level at Southheach
40,
Varlatlora witi.
0 =1. 3-2. 6 cm layer,
lew.1 i orgardr. Irnatt
C; =5.2-6. 5 cm layer,
I
Upper
Mid
Lower
Intertidal Level at Sally's Bend
elated to 2ea3on and depth
=7. 8-9. 1 cm layer..
he
dirne.
=0. 0-1. 3 cm layek,
400
400
Summer 1968
350
350
ro 0 300
Pd
Summer 1968
300
250
a
td
250
b0
200
8 200
bo
bo
a
a
150
150
100
100
50
50
Lower
Mid
Intertidal Level at Southbeach
Upper
Ord
Lower
Mid
Upper
Intertidal Level at Sally's Bend
Figure 41. Variation with intertidal level in organic matter as related to season and depth in the sediment.
=7. 8-9. 1 cm layer.
0 =1. 3-2.6 cm layer, 0 =5, 2-6. 5 cm layer,
A -0.0 -1. 3 cm layer,
400
400
350
350
0 300
U
$4
it
yN
yyd
300
td
E 250
250
U
cd
CO
BO
bo° 200
0 200
150
150
100
100
50
50
b0
I
0.0 -1. 3
1. 3-2. 6
I
5. 2-6. 5
Depth in cm at Southbeach
1
7. 8-9. 1
1
0. 0-1. 3
1. 3 -2.6
5. 2-6. 5
Depth in cm at Sally's Bend
Figure 42. Variation with depth in organic matter in the sediments as related to season and position in the intertidal zone.
A =Summer 1967, 13 =Fall 1967, 0 =Winter 1967-1968,
=Spring 1968, O =Summer 1968.
7.8 -9, 1
400
400
Mid-Intertidal
Mid-Intertidal
350
350
300
300
U
U
c.
250
250
U
bo
s.
N
'if 200
200
O
bo
bO
150
150
100
100
50
50
I
0.0-1.3
I
1.3-2.6
t
5.2-6.5
Depth in cm at Southbeach
7.8-9.1
­
0.0-1.3
1.3-2.6
5.2-6.5
Depth in cm at Sally's Bend
Figure 43. Variation with depth in organic matter in the sediments as related to season and position in the intertidal zone.
= Spring 1968, 0 = Summer 1968.
A =Summer 1967,
=Fall 1967,
0 = Winter 1967-1968,
7.8-9.1
450
450
Upper Intertidal
Upper Intertidal
400
400
350
350
300
300
250
250
on
s­
0
200
b° 200
150
150
100
100
50
50
I
0. 0-1. 3
1. 3-2. 6
5. 2-6. 5
Depth in cm at Southbeach
7. 8-9. 1
.,
, , .0
0.0 -1, 3
1. 3-2. 6
,,
I
5. 2-6. 5
Depth in cm at Sally's Bend
Figure 44. Variation with depth in organic matter in the sediments as related to season and position in the intertidal zone.
Summer 1967, 0 = Fall 1967, 0 = Winter 1967-1968,
= Spring 1968, 0 = Summer 1968.
7. 8-9, 1
100
Table 8. Species of diatoms collected from the Southbeach
and Sally's Bend study areas from July 6, 1967
to July 14, 1968.
Achnanthes brevipes Agardh
Achnanthes haukiana Grunow
Achnanthes lanceolata (de Brebisson) Grunow
Achnanthes sp. 1
Achnanthes sp. 2
Actinoptychus marmoreus Brun
Actinoptychus senarius Ehrenberg
Amphiprora alata (Ehrenberg) Kiitzing
Amphora angusta var. typica Cleve
Amphora elegans Peragallo
Amphora granulata Gregory
Amphora lineolata Ehrenberg
Amphora ocellata Donkin
Amphora ostrearia de Brebisson
Amphora proteus Gregory
Amphora truncata Gregory
Amphora sp. 1
Anaulus balticus von Riemer Simonson
Auliscus coelatus Bailey
Aulocodiscus probabilis A. Schmidt
Bacillaria paxillifer (0. F. Willer) Hendey
Biddulphia alternans (Bailey) Van Heurck
Biddulphia aurita (Lyngbye) de Brebisson
Biddulphia dubia (Brightwell) Cleve
Biddulphia longicruris Greville
Caloneis brevis (Gregory) Cleve
Caloneis westii (Wm. Smith) Hendey
Campylodiscus echeneis Ehrenberg
Campylosira cymbelliformis (A. Schmidt) Grunow
Cerataulus turgidus (Ehrenberg) Ehrenberg
Chaetoceros didymus Ehrenberg
Cocconeis clandestina A. Schmidt
Cocconeis costata var. pacifica (Grunow) Cleve
Cocconies decipiens Cleve
Cocconeis dirupta Gregory
Cocconeis placentula var. euglypta (Ehrenberg) Cleve
Cocconeis scutellum Ehrenberg
Cocconies sp. 1
Cocconeis sp. 2
Coscinodiscus curvatulus Grunow
Coscinodiscus eccentricus Ehrenberg
101
Table 8. (con't.)
Coscinodiscus lineatus Ehrenberg
Coscinodiscus marginatus Ehrenberg
Coscinodiscus nitidus Gregory
Coscinodiscus radiatus Ehrenberg
Coscinodiscus sp. 1
Cymbella mexicana Ehrenberg
Cymbella ventricosa Kiltzing
Diatoma hiemale var. mesodon (Ehrenberg) Grunow
Diatoma vulgare Bory
Dimmerogramma minor (Gregory) Ralfs
Dimmergramma sp. 1
Donkinia recta (Donkin) Grunow ex Van Heurck
Diploneis bombus Ehrenberg
Diploneis crabro Ehrenberg
Diploneis interrupta (Kiltzing) Cleve
Diploneis smithii (de Bre'bisson) Cleve
Epithemia sorex Kiftzing
Epithemia turgida Kiitzing
Eunotogramma laeve Gi-unow
Eunotogramma marinum (Wm. Smith) Peragallo
Fragillaria pinnata Ehrenberg
Grammatophora angulosa Ehrenberg
Grammatophora marina (Lyngbye) Ktitzing
Grammatophora maxima Grunow
Gomphonema angustatum var. subcapitata Kiitzing
Gomphoneis herculiana var. robusta Grunow
Gyrosigma balticum (Ehrenberg) Cleve
Gyrosigma fasciola (Ehrenberg) Cleve
Gyrosigma obliquum Grunow apud. Cleve and Grunow
Gyrosigma wansbeckii (Donkin) Cleve
Gyrosigma sp. 1
Hantzschia marina (Donkin) Grunow
Hantzschia virgata (Roper) Grunow
Isthmia nervosa Ktftzing
Licmophora abbreviata Agardh
Licmophora ehrenbergii (Ktitzing) Grunow
Licmophora gracilis (Ehrenberg) Grunow
Licmophora tenuis (Ktitzing) Grunow
Mastogloia exigua Lewis
Melos.ira moniliformis (Willer) Agardh
Melosira sulcata (Ehrenberg) Kiitzing
Navicula abscondita Hustedt
Navicula arenaria Donkin
102
Table 8. (con't.)
Navicula cancellata Donkin
Navicula clavata Gregory
Navicula complanata Grunow
Navicula comoides (Agardh) Peragallo
Navicula digito-radiata (Gregory) Ralfs
Navicula directa var. remota Grunow
Navicula diversistriata Hustedt
Navicula finmarchica Cleve
Navicula forcipata var. densestriata A. Schmidt
Navicula granulata Bailey
Navicula grevillei (Agardh) Heiberg
Navicula humerosa (de Brebisson) ex Wm. Smith
Navicula jamalinenses Cleve
Navicula litoricola Hustedt
Navicula lyra Ehrenberg
Navicula lyra var. elliptica A. Schmidt
Navicula lyra, var. subelliptica Cleve
Navicula normaloides Cholnoky
Navicula palpebralis (de Brgbisson) ex Wm. Smith
Navicula peregrina (Ehrenberg) Kiitzing
Navicula pseudony Hustedt
Navicula pusilla Wm. Smith
Navicula rhynchocephala Kiitzing
Navicula sp. 1
Navicula sp. 2
Navicula sp. 3
Navicula sp. 4
Nitzschia acuminata Wm. Smith
Nitzschia angularis Wm. Smith
Nitzschia apiculata Gregory
Nitzschia granulata Grunow
Nitzschia marginulata Grunow
Nitzschia navicularis (de Brebisson) Grunow
Nitzschia punctata (Wm. Smith) Grunow
Nitzschia punctata var. coarctata Grunow
Nitzschia sigma Wm. Smith
Nitzschia socialis Gregory
Nitzschia spathulata var. hyalina Gregory
Nitzschia vivax Wm. Smith
Nitzschia sp. 1
Opephora marina (Gregory) Petit
Opephora schwartzii (Grunow) Petit in Pellatin
Pinnularia cruciformis Donkin
103
Table 8. (con't. )
Plagiogramma staurophorum (Gregory) Heiberg
Plagiogramma brockmanii Hustedt
Pleurosigma affine var. normanii Ralfs
Pleurosigma angulatum var. quadrata Wm. Smith
Pleurosigma decorum Wm. Smith
Raphoneis amphiceros (Ehrenberg) Ehrenberg
Raphoneis sp. 1
Rhabdonema arcuatum (Lyngbye) Kiitzing
Rhoicosphenia curvata (Kiitzing) Grunow
Rhopalodia musculus Kazing
Skeltonema costatum (Greville) Cleve
Stauroneis salina Wm. Smith
Stephanopyxis nipponica Gran and Yendo
Synedra fasciculata (Agardh) Kiitzing
Synedra fasciculata var. truncata (Greville) Patrick
Surirella fastuosa var. cuneata Witt
Surirella gemma (Ehrenberg) Kiitzing
Surirella ovata Kiitzing
Tabellaria flocculosa (Roth) Ku-tzing
Terpsinoe americana Baily
Thalassionema nitzschioides Grunow
Thalassiosira aestivalis Gran and Angst
Trachyneis aspera var. aspera (Ehrenberg) Cleve
Trachysphenia australis Petit
Trigonium arcticum (Brightwell) Cleve
Tropodeneis lepidoptera var. minor Cleve
Tropodeneis vitria (Wm. Smith) Cleve
104
study area which is under the influence of the marine realm (Tables
9-11). The Sally's Bend study area which had far fewer species is
considered to be in the marine-fluviatile realm (Tables 9-11). This
study does not pretend to be an exhaustive taxonomic account of the
diatom flora but does record the most abundant and prominant species
together with some of the rarer species.
Photomicrographs representing the species identified are
included to provide a partial means of identifying the various taxa and
to provide a useful record of the entities encountered in the course of
this study (Appendix B, Plates I-XVII).
Diatoms were found to constitute almost the entire flora of
microalgae. On various occasions several species of Silicoflagellates
were encountered but these were presumed to be filtered out from
the phytoplankton. Very rarely, one or two unicellular specimens
from the Chlorophyta, Euglenophyta, or Pyrrophyta were seen.
No
filamentous Chlorophyta or Cyanophyta were ever encountered. Since
the taxonomy of the unicellular green algae and the Euglenoids is
difficult without cultures, to enable observation of various aspects of
the life cycle, no effort was made to determine the species of these
taxa.
Table 9. Diatom species collected from the lower intertidal zone at the Sally's Bend and Southbeach study areas.
Sally's Bend
N
to
N
to
N.
I
I
I
00
11
I
.-1
,.1
NI
N
0\
00
a1
m
Species
Achnanthes brevipes
Achnanthes haukiana
Achnanthes lanceolata
Achnanthes sp. 1
Achnanthes sp. 2
Actinoptychus marmoreus
Actinoptychus senarius
Amphiprora alata
Amphora anzusta
Amphora elegans
Amphora granulata
Amphora lineolata
Amphora ocellata
Amphora ostrearia
Amphora proteus
Amphora truncata
Amphora sp. 1
Anaulus balticus
Auliscus coelatus
Aulocodiscus probabilis
Bacillaria paxillifer
Biddulphia alternans
Biddulphia aurita
Biddulphia dubia
Biddulphia longicruris
Caloneis brevis
I
I
X
to
0
N
to
I
N
to
I
Le)
I
M
1-1
I
to
I
,-I
%1
I
Southbeach
00
00
I
I
I
A
to
al
N.--I
,-1
x
00
to
00
to
CO
00
00
N
i
i
I
3.
I
to
01
4A
to
0
to
N1
aI
.
.
,c
p.
to
O1
!
x
to
00
x
x
x
x
x
x
x
x
x
x
x
x
to
to
I
.
01
N.
,1
I
q\N""A
N.
N
I
I
to
I
to
r-
00
to
I
U)
co
to
I
M
00
to
I
CI
1
-
oo
to
I
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
x
x
x
x
x
x
x
x
x
x
I
00
to
I
00
tO
C
4
A
, 0I
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
00
to
x
x
x
N.
d'
v.4.4
x
x
to
N
.
x
x
N.
x
x
x
x
x
x
x
x
x
x
x
XXXXXXX
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
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
(continued)
Continued.
9.
Table
Southbeach
Bend
Sally's
00
00
00
1.0
CO
C7)
00
00
00
CA
C
,
rq
to C
m
I A
I .
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
x x
x
x x
x x
x
x x x x
x
x x
x
x x
x
x x x
00
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
cI1
x
x
x x x
eI
x
x
x
x
x
x
x x x x
1-1
x
0
x
'I'
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
1
x
x
scutellum
x
sp.
Cocconeis
sp.
Cocconeis
x
x
x x
lineatus
x
radiatus
x x
nitidus
vulszare
x
x
x
placentula
Cocconeis
Coscinodiscus
Coscinodiscus
Coscinodiscus
sp.
Coscinodiscus
Cymbella
mexicana
Cymbella
ventricosa
Diatoma
Diatoma
Dimmerojzramma
minor
Dimmerogramma
sp.
1
x x
dirupta
Cocconeis
Coscinodiscus
x
decipiens
Cocconeis
x
costata
Cocconeis
Coscinodiscus
x
clandestina
Cocconeis
x
Cocconeis
Coscinodiscus
x
1 2
hiemale
x x
x
x
x
turgidus
Cerataulus
Chaetoceros
x
Campylosira
x
echeneis
Campylodiscus
cymbelliformis
marginatus
x x x x
v.1
eccentricus
x
C
curvatulus
x
a%
-
didymus
x x x x
0,
0
C e-I
4
4
I
I
4
0
westii
x
N
N.
N.
N.
N1
N. t.0
N.
00 t.0
00
IC
00 1.0
00
1 0)
0)
00
I
-1
00
00
m1
117
00
0
n csj
csj
N.
N t.0
Species
Caloneis
recta
Donkinia
(
continued)
Table 9. Continued.
Sally's Bend
N.
l0
Diploneis bombus
Diploneis crabro
Diploneis interrupta
Diploneis smithii
Epithemia sorex
Epithemia turgida
amotogramma laeve
Eunotogramma marinum
Fragillaria pinnata
Grammatophora angulosa
Grammatophora marina
Grammatophora maxima
Gomphonema angustatum
Gomphoneis herculiana
Gyrosigma balticum
Gyrosigma fasciola
Gyrosigma obliquum
Gyrosigma wansbeckii
Gyrosigma sp. 1
Hantzschia marina
Hantzschia virgata
Isthmia nervosa
Licmophora abbreviata
Licmophora ehrenbergii
Licmophora gracilis
Licmophora tenuis
N.
0
cv
1
1
00
1.11
kt)
1
1
M
Species
N.
CV
I
00
x
kl)
.-I
0,
I
0
...
x
x
I
N.
N.
..-o
l0
in
-4
I
4-I
CV
kr)
.-I
1
I
00
00
kr)
l0
1
1
-
I-I
4
c.4
Southbeach
00
00
kr)
kr)
Gil
A
A
-4
x
x
x
x
00
k0
00
t.1..
0
.1.
N-I
-I
4
u%
x
x
VD
1
I
to
x
00
k0
1
1
N
x
N.
kr)
o
t..!....
x
x
x
x
N.
00
kr)
kr)
kr)
4
,t,
1
1
1
(11
0
N.
01\
00
I
x
x
x
N.
k0
N.
kr)
N.
k0
N.
lO
..-1
x
I
-I
.-i
I
Ckl
N-4
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
g
N.1
_4
x
x
x
x
1
t....
CV
x
x
x
x
x
x
x
x
x
1
Lt.)
Cs]
.
'-4
x
x
x
x
x
x
x
x
x
00
k0
1
en
CV
A
x
00
kr)
00
k0
1
1
cr,
00
,-I
,
ce1
-,
4
00
00
l0
kr)
1
1
kt)
,,.
00
14
kr)
CM
.
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
(continued)
Table 9. Continued.
Sally's Bend
N
N.
I
I
0
N
VD
M
N
Species
Mastogloia exizua
Melosira moniliformis
Melosira sulcata
Navicula abscondita
Navicula arenaria
Navicula cancellata
Navicula clavata
Navicula complanata
Navicula comoides
Navicula digito-radiata
Navicula directa
Navicula diversistriata
Navicula finmarchica
Navicula forcipata
Navicula granulata
Navicula grevillei
Navicula humerosa
Navicula jamalinenses
Navicula litoricola
Navicula lvra
Navicula norm aloides
Navicula palpebrialis
Navicula peregrina
Navicula pseudony
Navicula pusilla
Navicula rhynchocephala
to
01Cl I
N.
tO
I
00
-1
x
x
x
N.
tf)
,-I
CO
I
x
x
'o
I
00
VD
i
-1
.1
..-
e.-II
N
N-II
-
x
x
x
I I
0
1
x
N.
tO
Southbeach
00
00
00
VD
to
to
I
01
I
N
A
04
00
to
1
Ck
!
J)
00
to
00
tO
0
V
I NI
i
I
N.
N.
to
N.
tO
t 1
0
CI
I
4
to
I
I
VD N
N-
CO
A
N
to
4I
0
..-I
1
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
N.
to
I
N.
tO
N.
I
1.10
I
C N. 1
I
N
vI.
Cs1
,-.
N
I
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
N.
C1
x
x
CO
to
I
x
x
x
x
x
x
x
x
-II
00
to
I
.-I I
c'
x
x
x
00
c,
00
to
I
l.i
x
x
x
00
00
m . N.4 m
x
I
x
x
x
VD
I
I
x
x
x
x
00
to
I
s
ko
00
-II
r
,-+
x
x
x
x
x
x
00
tO
N
I
tO
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
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
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
( continued)
Table 9. Continued.
to
N.
to
oI
I
00
I
I
I
N
Navicula sp. 1
Navicula sp. 2
Navicula sp. 3
Navicula sp. 4
Nitzschia acuminata
Nitzschia angularis
Nitzschia apiculata
Nitzschia granulata
Nitzschia marginulata
Nitzschia navicularis
Nitzschia punctata
Nitzschia sigma
Nitzschia socialis
Nitzschia spathulata
Nitzschia vivax
to
en
I
Species
N.
00
at
N.
to
I
N.
to
I
Sally's Bend
00
to
I
00
to
I
00
to
I
00
to
I
00
to
I
Lr)
O
I
N
I
I
"-t
to
I
e-I
I
%1
I
00
to
e0
00
to
N.
Tr
to
to
I
I
-1
x
01
Nt.
cn
00
at
0
I
x
x
x
to
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
C..
to
x
I
N.
NN
N
Cl
x
x
00
NIA
I
x
00
x
x
x
x
x
x
x
Nitzschia sp. 1
Opephora marina
Opephora schwartzii
Pinnularia cruciformis
Plagiograrnm a staurophorum
Plagiogramm a brockrnanii
Pleurosigma affine
Pleurosigma angulatum
Pleurosigma decorum
Raphoneis amphiceros
x
Raphoneis sp. 1
Rhabdonem a arcuatum
x
x
x
x
x
x
x
x
x
x
x
00
to
to
to
oo
to
00
00
to
x
x
x
x
x
x
x
x
x
x
x
x
00
NM
x
x
x
x
x
x
x
x
x
00
x
x
x
II
N.
N.
x
x
Southbeach
nto
N
to
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
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
(continued)
Table 9. Continued.
Sally's Bend
N.
Species
t./0
N.
10
n
z
CO
CO
01
8
Rhoicosphenia curvata
Rhopalodia musculus
Skeltonema costatum
Stauroneis salina
Stephanopyxis nipponica
x
Synedr a. f asci culata
x
N.
x
N.
tO
If)
.4
N.
00
00
t4D
M
I
C4
1NI2
eI
x
A
00
.0
0,!
A
Southbeach
00
VD
CO
VD
CO
VD
0
4.
00
k0
x
LO
N.
N.
N.
N.
N.
N.
00
VD
VD
VD
VD
VD
VD
VD
I
4.
I
LA
N
4
N.
0.1
N
1
00
CT1
x
t\1
1-1
CV
I
00
tO
00
t.0
00
00
00
VD
VD
VD
en
NI.,
If)
I
00
C1
I
tN)
1.-1
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
Surirella fastuosa
x
Surirella gemm a
Surirella ovata
Taheliarla-flocculosa
Terpsinoe americana
Thalassionema nitzschioides x
Thalassiosira aestivalis
Trachyneis aspera
Trachysphenia australis
Trigonium arcticum
Tropodeneis lepidoptera
Tropodeneis vitria
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
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
Table 10. Diatom species collected from the mid-intertidal zone at the Sally's Bend and Southbeach study areas.
Sally's Bend
N Nto
c.0
I
en
01
Species
Achnanthes brevipes
Achnanthes haukiana
Achnanthes lanceolata
Achnanthes sp. 1
Achnanthes sp. 2
Actinoptychus marmoreus
Actinoptychus senarius
Amphiprora alata
Amphora angusta
Amphora elegans
Amphora granulata
Amphora lineolata
Amphora ocellata
Amphora ostrearia
Amphora proteus
Amphora truncata
Amphora sp. 1
Anaulus balticus
Auliscus coelatus
Aulocodiscus probabilis
Bacillaria paxillifer
Biddulphia alternans
Biddulphia aurita
Biddulphia dubia
Biddulphia longicruris
Caloneis brevis
I
0
ty
I
I
00
01
x
N
to
N
00
I
I
,I
I
0
II
x
t/D
I
in
I
N
k0
I
en
4
NI
I
N
I-1
x
x
00
00
VD
V:,
I
,-4
00
to
A
011
00
to
0%
00
to
N
mr.1
I
I
I
I
x
I
00
to
c.0
Southbeach
N
N
to
N
VD
to
4
8
4NI
,I0
C!,
I
N
I
I
t.0
I
tA
00
x
x
k0
4
I
CT
x
x
x
N
x
0
v.
x
x
N
N
VD
VD
.
m
I
,-o
4-1
x
.
..-1
N
k0
NI
N
N
e4
,I N
I
x
I
00
to
I
in
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
00
to
.
00
00
00
,
VD
NI..
in
to
x
x
x
to
I
NN
00
,..,
,-.4
ci
en
ti
I
xx
I
x
I
x
I
to
I
to
I
x
x
x
x
x
x
x
x
x
xx
x
x
x
x
x
x
x
xx
xx
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
x
x
xxxxxxxx
xx
xxxxxxx
xx
x
x
x
x
xx
xx
xx
xx
xx
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
I
x
x
x
CM
x
x
x
x
x
I
00
x
x
x
x
I
en
01
x
x
00
VD
x
x
x
x
(continued)
Table 10. Continued.
Sally's Bend
N.
N.
N.
to
N
to
I
M
I
I
I
ty
001
N.
tO
I
I
to
Species
Caloneis westii
Campylodiscus echeneis
Campylosira cymbelliformis
Cerataulus turgidus
Chaetoceros didymus
Cocconeis clandestina
Cocconeis costata
Cocconeis decipiens
Cocconeis dirupta
Cocconeis placentula
Cocconeis scutellum
Cocconeis sp. 1
Cocconeis sp. 2
Coscinodiscus curvatulus
Coscinodiscus eccentricus
Coscinodiscus lineatus
Coscinodiscus marginatus
Coscinodiscus nitidus
Coscinodiscus radiatus
Coscinodiscus sp. 1
Cymbella mexicana
Cymbella ventricosa
Diatoma hiemale
Diatoma vulgare
Dimmero,vamma minor
Dimmerogramma sp. 1
Donkinia recta
00
to
I
01
00
.-I
0wi
I
1.0
Il
,I4-1
I
M
N.1
I
M
CO
to
I
1.4
1.1
!
vl
00
I
I
01
01
I
I
to
N
to
n
00
to
I
al
4
CO
00
00
gi.
01
I
to
to
I
1)
;
to
Nt.
-1
I
I
to
N.
N.
N.
N.
N.
41.
ell
4
,:t.
d,
8
e-I
NI
x
x
x
to
N.
x
x
x
x
x
x
x
x
x
x
x
x
x
III III
Southbeach
00
x
x
x
x
to
I
to
x
x
x
x
x
x
x
x
to
I
N.
to
I
N.
to
I
co
-1
I
I
x
N.
to
I
N.
e-I
N
I
N
1A
x
x
.-N
00
CO
00
00
00
NN
al
00
'-4
t.o
CO
,4
en
't
I
to
x
x
x
x
00
to
Lo
to
en
I
x
x
i
x
x
x
x
x
to
-II
to
I
x
x
x
to
,-I
In
to
N
I
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
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
(continued)
Table 10. Continued.
Sally's Bend
N.
CD
N.
D
I
m
0I
N
csa
Species
Dip loneis bombus
Diploneis crabro
Diploneis interrupta
Diploneis smithii
Epithemia sorex
Epithemia turgida
Eunotogramma laeve
Eunotogramma marinum
Fragillaria pinnata
Grammatophora angulosa
Grammatophora marina
Grammatophora maxima
Gomphonema angustatum
Gomphoneis herculiana
Gyrosigma balticum
Gyrosigma fasciola
Gyrosigma obliAuum
Gyrosigma wansbeckii
Gyrosigma sp. 1
Hantzschia marina
Hantzschia virgata
Isthmia nervosa
1
1
00
DI
x
x
x
N.
t.0
1
00
.-I
0
,r1
1
x
N.
t.0
1
Co
.-1
1
1-1
1.-1
x
C..
.0
I
en
-t
1
N
I-I
x
CO
ID
I
DD
00
I
CO
I
1.4
1
1
1
.-1
x
x
x
x
x
Southbeach
00
tO
I
al
00
CO
to
g.
VD
I
0
,1
I
1
V'
x
V
4/
.4-1
1
x
l0
I
1
,0
N.
x
N.
00
tO
N.
N.
N.
4
Nt.
D
to
0;
.0
I
N.
D
I
N.
ID
I
Cr)
N.
VD
I
0
1
00
Cj1
.-I
x
1
,-.1
11
1
N
-4
x
x
x
00
ki0
D
NN
CO
CV
er)
N ,4
1
00
at
I
1
I
I
,,
1
..-.1
x
x
00
VD
tO
ef)
CN3
1
00
If)
N.
x
x
x
I
-,
1
Nr
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
.0
I
N
1
VD
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
Licmophora . abbreviata
Licmophora ehrenbergii
Licmophora gracilis
Licmophora tenuis
1
if)
00
00
x
x
x
x
x
I
tO
'.-,
x
x
x
x
x
tta
x
x
x
x
CO
x
x
x
x
x
(continued)
Table 10. Continued.
N.
kO
I
en
Species
Mastogloia exigua
Melosira moniliformis
Melosira sulcata
Navicula abscondita
Navicula arenaria
Navicula cane ell ata
Navicula clavata
Navicula complanata
Navicula comoid es
Navicula digito-radiata
Navicula directa
Navicula diversistriata
Navicula finmarchica
Navicula forcipata
Navicula granulata
Navicula grevillei
Navicula humerosa
Navicula amalinenses
Navicula litoricola
Navicula lyra
Navicula normaloides
Navicula palpebrialis
Navicula peregrina
Navicula pseudony
Navicula pusilla
Navicula rhynchocephala
N
N.
(0
0
I
es.1
I
00
I
Crl
N.
to
I
00
-II
N.
Sally's Bend
00
00
t.0
cI
I
VD
N.
ka
I
I
en
0,-I
,-,
I
.-i
.-,
x
x
x
cn
-I
I
N
.-.
-,
-I
I
I
-.
Southbeach
00
t.10
00
t.0
I
I
CA
CA
I
I
en
'I.
00
VD
00
o
OD
VD
t.......
0.-I
V
I
I
I
I
.0
N.
,.0
I
-
N.
N.
VD
011
N.
,.0
N.
N.
N.
N
00
00
00
VD
VD
1/40
i
I
I
I
VD
I
k.0
I
VD
i
00
t.0
CO
k.0
I
I
I
VID
.4.
N.
0
I
00
CIN
e-I
en
I
-I
N-I
I
N
-I
N.
N
I
N
N-I
N Nenell
to
I
-,
VD
cn
00
c0
I
I
I
-I
-1
Co
el.
.-I
In
00
t.0
I
on
N
I
ko
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
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
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
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
(continued)
x
x
x
x
Table 10. Continued.
N.
to
ty
nto
0(V
o
00
I
o
Species
Navicula sp. 1
Navicula sp. 2
Navicula sp. 3
Navicula sp. 4
Nitzschia acuminata
Nitzschia angularis
Nitzschia apiculata
Nitzschia granulata
Nitzschia marginulata
Nitzschia navicularis
Nitzschia punctata
Nitzschia sigma
Nitzschia socialis
Nitzschia spathulata
Nitzschia vivax
Nitzschia sp. 1
Opephora marina
Opephora schwartzii
Pinnularia cruciformis
Plagiogramma staurophorum
Plagiogramma brockmanii
Pleurosigma affine
Pleurosigma angulatum
Pleurosigma decorum
Raphoneis amphiceros
Raphoneis sp. 1
Rhabdonema arcuatum
Sally's Bend
N.
N.
t.13
e-I
I
N.
to
m
I
N
00
to
,r4
I
00
to
00
to
d,
dA
00
CO
00
1/40
0
e-4
4
00
to
x
to
N.
tO
r.
to
4
11
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
N
to
x
x
x
N.
to
N.
N
1,1
CO
00
Er)
rn
to
N
CO
to
to
00
00
01
00
tO
00
CI)
Nt.
tf)
to
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
N
N
00
to
-1
x
x
to
N
N
x
x
x
Southbeach
N
to
41.
xx
xx
xx
x
N.
to
8
Oli
LA
x
x
N.
to
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
xx
xx
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
xx
x
x
x
x
xx
x
x
x
x
x
x
x
xx
xx
xx
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
(continued)
Table 10. Continued.
Southbeach
Sally's Bend
N.
N.
to
to
to
I
001
I
m
ty
Species
I
CO
Rhoicosphenia curvata
Rhopalodia musculus
Skeltonema costatum
Stauroneis salina
Stephanopyxis nipponica
Synedra fasciciilata
Surirella fastuosa
Surirella gemma
Surirella ovata
Tabellaria flocculosa
Terpsinoe americana
Thalassionema nitzschioides
Thalassiosira aestivalis
Trachyneis aspera
Trachysphenia australis
Trigonium arcticum
Tropodeneis lepidoptera
x
Tropodeneis vitria
I
I
01
N.
00
1..1
0.-1
I
N.
ID
I
.,,
.
4-1
I
.-i
.-o
N.
to
I
m
T-I
I
N
.-I
00
tO
00
00
to
to
r1
4,
c,l
I
A
I
-1
,-I
A
00
00
l0
c,l
i
N.
4
A
to
CO
to
I
I-I
I
tO
00
to
,
,- t
I
N
N.
to
to
l
C..
to
01
4
N.
to
,
v.
02A 0
I
1
,-1
N.
N.
N.
00
00
00
00
to
to
.
to
to
.
to
.
00
CO
to
I
to
.
to
.
to
01
00
n
-I2
I
.
.
..-1
I
e-I
N
I
Li)
e0
c.-r
N
A
x
x
x
x
I
x
x
x
N.
to
N
I
-1
i
m
to
-1
,-I
I
I
4
.
00
N
I
111
x
tO
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
xxxxx
x
x
x
x
x
xx
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
XX
x
x
X
X
X
X
X
X
x
x
x
x
x
X
x
x
X
X
x
x
X
x
Table 11. Diatom species collected from the upper intertidal zone at the Sally's Bend and Southbeach study areas.
N
V:,
I
m
0.1
Species
I
00
Achnanthes brevipes
Achnanthes haukiana
Achnanthes lanceolata
Achnanthes sp. 1
Achnanthes sp. 2
Actinoptychus marmoreus
Actinoptychus senarius
Amphiprora alata
Amphora angusta
Amphora elegans
Amphora granulata
Amphora lineolata
Amphora ocellata
Amphora ostrearia
Amphora proteus
Amphora truncata
Amphora sp. 1
Anaulus balticus
Auliscus coelatus
Aulocodiscus probabilis
x
x
Bacillaria paxillifer
Biddulphia alternans
Biddulphia aurita
Biddulphia dubia
x
x
Biddulphia longicruris
Caloneis brevis
N.
N.
V0
I
140
o03
I
01
x
x
I
00
.-1
I
N.
c0
I
If)
..-1
I
0
.-.
x
x
x
x
x
.-1i
,I
N
Sally's Bend
00
00
CO
CO
CO
m
.-I
.-I
I
,I
I
0.,
1.
I
.­
I
01
A
00
'.0
CO
CO
71
C1
1
I
00
'.0
I
d
x
4
.
x
x
00
CO
I
0
eI
I
00
c0
I
V
...I
.
v0
N
x
x
x
x
N.
140
t2
g.
x
x
x
x
x
x
x
x
x
x
x
N
N.
'.0
CO
A
4
A
A
x
x
x
N.
N.
I
.
0
10
I
4.
m
I
0
/I
I
.-'
,1
N.
0
CO
I
N.
.
A
v,
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
c0
00
c0
x
x
00
CO
00
c0
I
I
I
I
I
01
00
CO
CO
A4
V)
(N)
.
(NI
.1
x
x
x
1-1
eq
.
.
VD
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
xx
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
xx
xx
x
x
x
xx
x
x
x
x
x
00
CO
I
CO
x
x
x x
x
x
.
00
x
x
x
x
i
x
xx
x
x
x
x
0
121
x
x
x
x
00
...1
x
x
x
x
Southbeach
N.
xx
xx
x
x
x
x
x
x
x
x
x
(continued)
(continued)
a
1
sp.
erogramm
Dimm
Coscinodiscus
x
lineatus
Coscinodiscus
eccentricus
Coscinodiscus
x
curvatulus
Coscinodiscus
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
x
x
x
x
x
clandestina
Cocconeis
x x
costata
Cocconeis
x
decipiens
Cocconeis
x
x
x
x
x
x
placentula
dirupta
Cocconeis
x
x
x
x
x
x
scutellum
Cocconeis
x
x
x
x
x
sp.
2 1
sp.
x
marginatus
Coscinodiscus
x
x
nitidus
Coscinodiscus
Cocconeis
x
x
radiatus
Coscinodiscus
Cocconeis
x
1
sp.
Cocconeis
x
x
mexicana
Cymbella
x
x
ventricosa
Cymbella
x
hiemale
Diatoma
x
minor
Dimmerogramma
vulgare
Diatoma
x x
x
x
x x
recta
Donkinia
x
cymbelliformis
Campylosira
x x
x x
x x
x x
x x
x
x
x x
x
x
x
x
x
x
x
turgidus
Cerataulus
x
x x x
didymus
Chaetoceros
echeneis
Campylodiscus
x
x
x
x
westii
Caloneis
Cs.1
M
VD
I
.-1
..-1
kr) 00
kr) 00
. ul I
.q I
CO
00
00
.0
CID
kr) CO
.C I
.
. I
00
k0 00
CO
. .I I
01
II I
N
N.
kr)
kt, N.
. I
C
ID N.
N
C,
N.
N.
. ,, I
m
VD
VD
VD
t.0 N.
VD CO
00
00
k0
VD
ul
ct.
CO
NI
C
..-1
Cs.1
C1
.-1 -.4
4
4
01
Z
-I 7.
VD
VD
I
. I
ck
N-
:
:
1-1
L, 0I
t I
. g
1
01 .0 00
:4
00
00
01
01
N.
k.0 00
VD
CO
1-1
t.0 N.
VD
C
.1 lr)
I
I 0I
4 .I , I
CI
I
,I
.I 0
.-.4
4
VD
N.
00
I .I I
,t 0
N.
t.0 N.
VD
I 0I
0.1
I I
01
00
Species
Southbeach
Bend
Sally's
Continued.
11.
Table
Continued.
11.
Table
Southbeach
Bend
Sally's
00
kr,
10
k0 00
I
00
CO 00
m
VD
t0
-00
00
t0
V
e-I
I
tD
Li,
M
N
N
00
00 k0
N
0
g0)
!
01
I
N N e-I
m
-1
N.
I
I
00
o
1-I
-1
t
v4
A
I 0 I to
I 4. I N
1
4,
4.
4.
N. t0
k0
N.
t0
t.0
1.0
t0
N k0
00
00 t.0
00 t0
00tD
t0
00t0
00
00 1.0
V)
M
0
CO
I I
lilt
N. t0
N l0
tD
N. to
N.
Species
x
x
x
smithii
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
Gomphonema
Gyrosigma
Gyrosigma
Hantzschia
marina
x
x
x
x
x
x
x
obliquum
Gyrosigma
sp.
x x
x
fasciola
Gyrosigma
wansbeckii
x
Gyrosigma
balticum
x x
x
X
x x
x
X
x
x
marina
Grammatophora
x x
x x
x
x x
x
x
X
x
x
x
x
x
x
x
x
x
x
x
x
x x
angulosa
Grammatophora
x
x
x
x
1
x
x
x
x
marinum
pinnata
Fragillaria
maxima
herculiana
x
x
x
laeve
Eunotogramma
angustatum
X
turgida
Eunotogramma
Gomphoneis
x
sorex
Epithemia
Grammatophora
x
Epithemia
x
x x x
x
x
x
x
x
x
x
x x
x
interrupta
Diploneis
x
x
x
x
x
x
x
crabro
Diploneis
x
bombus
Diploneis
x
Diploneis
virgata
Hantzschia
nervosa
Isthmia
x
abbreviata
Licmophora
ehrenbergii
x
Licmophora
x
x
x
.gracilis
Licmophora
tenuis
Licmophora
r
x
x
x
x
exigua
Mastogloia
(continued)
Table 11. Continued.
Sally's Bend
N.
N.
N.
N.
00
00
00
00
00
00
00
N.
N.
N
N.
1/40
1/40
1/40
1/40
1/40
1/40
1/40
1/40
1/40
1/40
1/40
I
0
N
CO
I
I
01
CO
I
I
01
1/40
I
I
1/40
I
CO
I
I
I
cm
I
CO
Species
Melosira moniliformis
Melosira sulcata
Navicula abscondita
Navicula arenaria
Navicula canc ellata
Navicula clavata
Navicula complanata
Navicula comoides
Navicula digito-radiata
Navicula directa
Navicula diversistriata
Navicula finmarchic a
Navicula forcipata
Navicula granulata
Navicula .grevillei
Navicula humerosa
Navicula amalinenses
Navicula litoricola
Navicula lyra
Navicula normaloides
Navicula palpebrialis
Navicula peregrina
Navicula pseudony
Navicula pusilla
Navicula rhynchoc ephala
Navicula sp. 1
Southbeach
N.
N
I
00
x
x
CO
.-4
I
01
I
0
e.-I
x
x
x
x
IY1
..-I
I
m
.-i
I
-I
,I
N
7-I
x
x
x
...-1
.-I
I
-1
x
x
x
en
x
01
4
x
tI,
Li;
0
..-1
to
NII
-i
r".
CO
N-
x
x
x
x
x
x
x
x
x
x
x
CO
x
x
x
x
x
Cj1
x
I
Oil
c0
N.
N.
00
00
CO
CO
CO
I
CO
I
I
I
,I,
0
1-1
I
I
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
N.
-4
e-I
t,
N
I
NI
CM
.-I
..-1
x
x
x
x
x
x
v-I
I
x
x
x
x
x
x
x
x
x
x
,I
N
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
.-1
-i
x
x
x
I
I
x
x
x
00
k0
I
x
x
I
CA
x
x
x
x
CO
I
x
x
x
x
00
CO
cc)
x
x
x
x
00
rn
x
x
x
x
I
I
x
x
x
tO
CM
In
x
x
x
00
I
CO
x
x
x
x
x
I
cn
x
x
x
x
x
cr
VD
to
CO
cs.)
I
to
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
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
(continued)
N.)
O
Table 11. Continued.
Sally's Bend
N
N
I
0
to
tO
I
M
Species
Navicula sp. 2
Navicula sp. 3
Navicula sp. 4
Nitzschia acuminata
Nitzschia apiculata
Nitzschia granulata
Nitzschia marginulata
Nitzschia navicularis
Nitzschia punctata
Nitzschia sigma
Nitzschia socialis
Nitzschia spathulata
NI
CM
I
00
N
VD
tO
N
I
I
I
N
00
NI
0N..
I
0.1
1cf,
I
,-I
11
tO
00
tO
00
tO
00
tO
00
tO
I
I
I
I
1..1
-
I
I
en
e-I
N1
(Z1
al
al
I
I
en
'7'
00
tO
CO
l...I
I
N
A
00
tO
tO
0-.1
N
tO
I
.7
41
I
I
N
V.')
V.)
t
N­
VD
01
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
x
OIC,
x
x
x
x
I
tO
r.
'40
00
tO
00
tO
CO
110
tO
00
tO
00
tO
I
I
I
I
I
I
I
I
I.
en
011
04I
.-+
xx
Southbeach
N
N
4.
I
x
x
N
NtO
tO
I
1...1
x
I
N
cM
In
en
I
I
I
N
11
N
,4
x
x
x
CM
',.-1
CM
N
x
x
x
x
1al
I
en
00
4
4
x
x
Nitzschia vivax
x
Nitzschia sp. 1
Opephora marina
Opephora schwartzii
Pinnularia cruciformis
Plagiogramma staurophorum
Plagiogramma brockmanii
x
Pleurosigma affine
Pleurosigma angulatum
x
Pleurosigma decorum
Raphoneis amphiceros
Raphoneis sp. 1
Rhabdonema arcuatum
Rhoicosphenia curvata
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
vL
x
x
x
x
x
x
x
x
N
I
00
x
x
x
01:3
%i
x
x
x
co
00
l0
x
x
x
x
x
x
x
xx
x
x
x
xx
xx
xx
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
(continued)
N.)
Table 11. Continued.
Sally's Bend
N.
N.
N.
CO
VD
I
%.0
I
m
0!al
N
Species
Rhopalodia musculus
x
Skeltonema costatum
Stauroneis salina
x
Stephanopyxis nipponic a
Svnedr a fasciculata
Surirella fastuosa
Surirella gemm a
Surirella ovata
Tabellaria flocculosa
Terpsinoe americana
0
N
x
i
x
I
00
-II
0
x
N.
tO
e
tf)
.-4
e
-
N.
LO
I
rn
.-1
I
N
00
00
v-i
cA
,1.
A
(.0
I
,-I
1
to
x
Southbeach
00
00
tl)
d,
k0
I
00
lt)
Ci1
4
!
00
t.0
00
tO
N.
VD
N.
l0
d,
l0
t 'ot
I
0
.-1
I
I
V
to
e
X
x
x
x
x
x
x
x
x
x
x
x
N.
N.
tO
4
4
0I
l0
A
X
x
x
X
X
N.
k0
C..
k.0
.
..-,
I
I
x
x
x
x
x
x
x
x
x
x
x
x
x
x
I
N.
t.0
I
4
N.
N
N
I
N
x
00
VD
I
LO
N
e
..-1
x
x
x
x
CO
00
00
I
00
0-II
q)
00
t.0
I
00
tO
I
00
VD
00
V
(1)
t.0
tO
N
N
x
,0
en
x
I
m
e
x
x
x
x
X
x
x
x
x
x
X
x
x
X
x
X
x
x
x
x
Thalassionem a nitzschioides
x
Thalassiosira aestivalis
Trachyneis aspera
Trachysphenia australis
Trigonium arcticum
Tropodeneis lepidoptera
Tropodeneis vitria
x
x
X
x
X
x
x
x
x
x
x
x
x
x
x
123
DISCUSSION
On bright, sunny days during the summer, the microalgae
frequently formed brownish-green mats covering considerable areas
of the exposed surface of the sediments at the Southbeach and Sally's
Bend study areas. The estimates of the concentrations of chlorophyll
a and the direct counts of cells demonstrated that the microalgae were
concentrated in the uppermost layers of sediment throughout the
intertidal zone.
The chief advantage of occupying the uppermost layers
of sediment is the amount of light energy available for autotrophic
growth when the rapid extinction of the available light by the sediment
is considered. This extinction is the result of reflection, refraction,
scattering, and absorption of light by sediment, detritus, and inter­
stitial organisms.
Light penetration through the substrate has been measured by
various investigators. Oppenheimer and Wood (1962) found that light
penetrates 11 mm through sandy silt. Hopkins (1963) found 1% of the
incident radiation penetrating up to 4 mm. Taylor (1964) found 10% of
solar radiation penetrating approximately 1. 5 mm while 1% reached
a depth of 3 mm. Experiments performed on sediments from Yaquina
Bay indicated that 10% of incident radiation penetrated to a depth of
2 mm in sand but only 0. 5 mm in silt. Because of the larger numbers
of viable microalgae found in the upper centimeter and the percent­
age of incident radiation penetrating through the sediments being quite
124
low, it is probable that these microalgae have a high photosynthetic
efficiency at low light levels. Taylor and Palmer (1963) found that
intertidal benthic diatoms can photosynthesize above 90% of their
maximum capacity at 1.5 mm in depth and at 35% of their maximum
capacity at 3 mm in depth which would be at approximately 1% of
mid-day solar radiation.
Although the bulk of the algal biomass is located in the upper
3
cm of sediment it was found that significant numbers of microalgae
are present at considerable depths. Live algae below the sediment
surface have also been observed elsewhere. Moul and Mason (1957)
reported up to 40 live diatom cells/mm 3 of mud between 5 to 6 cm in
depth. Oppenheimer and Wood (1962) observed live diatoms to a
depth of 12 cm in the sediment. Gridntved (1962
,
as cited in Pamat­
mat, 1968) demonstrated photosynthetic activity of subsurface
sediment down to 5 cm by means of C14 uptake. Steele and Baird
(1968) found viable diatom populations at a depth of 20 cm in a sandy
beach. Pamatmat (1968) demonstrated the presence of live algae by
photosynthetic activity 10 cm beneath the sediment surface. In
Yaquina Bay estuary diatoms were found to be present 18 cm below
the sediment surface which was the greatest depth sampled.
The question arises as to the mode of subsurface existence of
these microalgae. A possible explanation is that these diatoms
assimilate carbon heterotrophically. The heterotrophic mode of
125
nutrition has been demonstrated by Lewin (1953) and Lewin and
Lewin (1960) for marine littoral diatoms in culture. Pamatmat (1968)
has reported that Dr. Joyce C. Lewin found that all species of
pigmented diatoms that are capable of heterotrophic growth can form
chlorophyll in the dark when supplied with a utilizable organic sub­
strate and have the ability to photosynthesize immediately upon
exposure to light. Another possible explanation is that these organ­
isms are especially adapted to spend long periods in the dark at low
metabolic rates with negligible pigment degradation. Culture
experiments performed by Steele and Baird (1968) have shown that an
axenic culture of a diatom could exist in the dark for 23 days without
any measurable loss of carbon or any pigment degradation.
On the tidal flats in Yaquina Bay two communities of microalgae
are present, i, e. epipelic and epipsammic. Epipsammic algae are
particularly subject to burial, being attached to sand grains, whereas
the epipelic algae are known to be able to migrate back to the surface.
Sand movements on the tidal flats may bury some of the algae and this
might explain the vertical distribution. It would be clearly advanta­
geous for the epipsammic algae to be able to detach themselves after
burial and to migrate upwards and reattach themselves to sand
grains at the sediment surface. Most of the epipsammic algae
observed, as Cocconeis and Achnanthes, have a raphe valve and these
species have shown movement under laboratory conditions (Moss and
126
Round, 1967). However, burial by sediments in Yaquina Bay may not
be as prevalent as reported elsewhere. Studies on the movement of
the sediment have shown no mixing below 2. 5 cm in depth over a four
month period. This may result in part from the stabilization of the
sediment by the microalgal community. A mucus cementing material
has been shown to be produced by diatoms. Williams (1962) cited
various investigators who consider this slime important in marine
sedimentation. They suggest that it gives cohesiveness to sediment,
thus preventing its removal by waves and currents.
A more probable explanation of the vertical distribution of the
microalgae is the occurrence of a diurnal migratory rhythm present
in the epipelic community. There are many papers on vertical
migration and this phenomenon has been reported from many areas
throughout the world. The physiological mechanisms of these diurnal
algal migrations are not yet completely understood. The advantages
of a migratory mechanism is obvious in the tidal habitat. A diurnal
migratory rhythm would allow the motile algae to cope with any
sedimentation by vertical movement upwards to remain within the
photic zone. Aleem (1950a) has shown that motile diatoms migrate
during a diurnal cycle and in this way can exist in anaerobic sediments
using energy stored while in the photosynthetic zone.
Various mechanisms have been proposed regarding these diurnal
migrations. Perkins (1960) suggested that the diurnal rhythm is
127
dependent on the intensity of light and as long as the incident light
exceeds a certain intensity the microalgae will remain near the
surface. Palmer (1960) also showed that light was an important
factor in vertical migration since he found that placing opaque shields
on the sediment surface prevented vertical migration. Hopkins (1963)
also found that migration of diatoms was stimulated by light but not by
pH, dissolved oxygen, or soluble organic matter. This light depend­
ency was seen in Yaquina Bay also. No mats of microalgae were
observed at low tide during evening hours or during cloudy days.
Only when the day was very sunny and bright were characteristic
patches of microalgae seen scattered over the surface of the tidal
flats. Recently Round and Happey (1965) and Round and Eaton (1966)
have found rythmic vertical migration in microalgae in non-tidal
waters of streams and ponds and therefore believe that these move­
ments are not directly related to tidal exposure or submersion. They
suggest that the upward rhythm is due to photosensitivity and motility
and the downward movement is due to a positive geotaxis.
Whatever
the mechanism, it has been shown that a vertical migration does
exist and the observations in Yaquina Bay tend to support the conclu­
sion that emergence of algae on the sediment surface occurs as the
tide recedes and the sediment surface is directly exposed to incident
radiation. The fact that the upper intertidal zone had a higher biomass
which decreased sharply at the lower depths is most likely the result
130
in the lower intertidal zone may also be explained by better oxygenated
conditions in this area.
Sediment temperatures are more variable with increasing
elevation on the beach, coincident with the longer period of exposure
to incident radiation. Organisms living near the surface of the
sediment are exposed to a highly variable temperature regime.
Johnson (1965) found that organisms living in the upper cm are exposed
to a temperature range three times as great as that experienced by
subtidal individuals of the same species. In the upper intertidal
zone of Yaquina Bay most of the microalgae were found in the upper
centimeter. This is the zone that undergoes the greatest variation in
temperature. The sediment nearer the surface at Southbeach under­
goes a much greater fluctuation in temperature than sediment at
greater depths. Pamatmat (1968) also found this relationship at
Friday Harbor. This greatly fluctuating environment may prevent a
large population from developing if temperature conditions are sub­
optimal for the species.
The effect of changes in salinity have also been measured by
various investigators. Johnson (1967) noted that in the upper inter­
tidal zone evaporation may increase the interstitial salinity during
periods of prolonged exposure. In the lower intertidal zone, inter­
stitial salinities are fairly constant. During the winter season, when
most of the precipitation occurs, the exposed tidal flat may be near
131
saturation with rain water and very low salinities could result.
Williams (1962) noticed that after rain at ebb tide salinities of 2 to 3%c
resulted . In warm weather, when evaporation was maximal,
salinities of 39 to 45 %o were recorded.
This greatly fluctuating
salinity may also prevent a community from becoming established in
the upper intertidal zone.
Another factor that may enhance the biomass of the lower inter­
tidal zone is that the microalgae in this zone are being exposed to sea
water for longer periods of time than those in the upper intertidal
zone. Organic and inorganic nutrients present in seawater are
necessary for growth and production of microalgae. Nutrients may
become limiting in the upper intertidal zone because sandy beaches
have few bacteria to regenerate nutrients necessary for microalgal
growth (ZoBell, 1946).
In Yaquina Bay, in sandy intertidal areas, the maximum number
of species and individuals are found in the lower intertidal zone.
This phenomenon cannot be attributed to any single environmental
factor but may result from the entire environmental complex which
varies with the ebb and flood of the tide.
It is obvious from the concentrations of chlorophyll a and direct
counts that the sand substrate supported a larger community of
microalgae than the mud substrate. No horizontal distributional trend
was found on the Sally's Bend tidal flat. This may be due to several
132
environmental variables which differ on the two tidal flats.
The most obvious difference between the two tidal flats is the
nature of the substrate. The Southbeach substrate consists primarily
of medium and fine sand whereas the Sally's Bend tidal flat consists
primarily of silt. On the basis of particle roundness the sediments of
Sally's Bend appear to be related more to sediments of fluviatile
origin and those of Southbeach to a marine origin (Ku lm and Byrne,
1966).
It was found that the substrate from Sally's Bend allowed 10% of
the available light to penetrate 0.5 mm whereas the Southbeach
sediment allowed this amount of light to penetrate 2.5 mm. These
findings agree with those of Gomoiu (1967) who found that light
penetrated in a different manner according to the granulometric
character of the sediments. As the sediments become finer, light
penetration is r educed.
The fact that Sally's Bend sediment has a
shallower photic zone than the Southbeach sediment certainly affected
the size of the standing crop.
Another factor that may prevent a sizable standing crop from
developing at the Sally's Bend study area is sedimentation. Kulm
(1965) found that due to the geometry of the bay, much of the fine
sediment carried by the river is transported over the Sally's Bend
flat and then sediments out. The fine silt, clay, and organic detritus
that is abundant on the Sally's Bend tidal flat can clog the interstices
133
of the sediment and restrict the development of a population.
The comparatively large amounts of organic matter in the
sediment at Sally's Bend may also be a factor preventing the establish­
ment of sizeable standing crops. It has been found by ZoBell (1940)
that the bacteria consume an average of 14.9x10
-12
3
cm of oxygen
per cell per hour in sea water at 22 C and considerably more in bay
mud which is usually richer in oxidizible organic matter. The
potential productivity experiments indicated low rates of oxygen
production by cores from Sally's Bend due to high rates of respiration
attributed to the bacteria. That bacterial respiration occurs to a
larger extent in sediment from the Sally's Bend tidal flat than in that
from Southbeach may limit the oxygen content of the silt.
It has also been found that bacteria have the tendency to alter
the pH of their environment (ZoBell, 1942). In Yaquina Bay the pH
of the sediment from Sally's Bend had a wider fluctuation than the
sediment from Southbeach. ZoBell (1942) has reported that certain
bacteria can lower the pH to 4.5 and others can raise it to 9.5. In
general however, the pH at Sally's Bend was lower than the pH at
Southbeach which is most likely the result of the activity of bacteria
decomposing the organic matter and producing CO2.
Castenholz (1961) observed that growth of epilithic diatom
populations are held in check by grazing of littorines and limpets and
noted that diatoms can increase in density when the grazing pressure
134
is removed. Extensive grazing activity on the Sally's Bend tidal flat
may also keep the population of microalgae at a minumum.
The nature of the sediment, bacterial activity, and grazing seem
to be the factors most probably influencing the standing crop of
microalgae at Sally's Bend. Bacteria, by virtue of their large num­
bers, rapid rate of multiplication, and intense biochemical activity,
have a pronounced effect on the physical, chemical, and biological
conditions of a tidal flat.
Estuarine sediments contain large amounts of chlorophyll
degradation products because portions of plant detritus, both from
plankton and benthos, are deposited in these areas. In Yaquina Bay
both the Southbeach and Sally's Bend tidal flats had significant amounts
of pheophytin present. However, the Sally's Bend sediment contained
much more pheophytin than the Southbeach sediment. This large
amount of detrital material is most probably allochthonous in origin
since the plant biomass at this station is relatively low. On the basis
of particle roundness, Kulm (1966) found that the sediments of the
Sally's Bend tidal flat appear to be related more to sediments of a
fluviatile origin. Due to the geometry of the bay plant detritus
washed down from upstream areas may be deposited on this tidal
flat along with the fluviatile sediment. It has also been shown that in
Yaquina Bay there is a decrease in tidal current velocity as one
proceeds from the mouth of the estuary upstream. Lack of vigorous
135
tidal currents may also be contributing to the large amount of
detrital chlorophyll present on the Sally's Bend tidal flat.
The Southbeach tidal flat sediment consists of well sorted sand
and is considered by Ku lm (1966) to be under the influence of the
marine realm of deposition and is characterized by vigorous tidal
action.
These tidal currents would tend to remove detrital material
from the Southbeach sandflat. The relatively low concentrations of
pheopigments at Southbeach could also result from the detrital
material being primarily autochthonous since this area is not a site
of deposition for fluviatile sediment and detritus. However, some
allochthonous material may accumulate in the form of detrital
seaweeds. At the Southbeach station the highest percentage of pheo­
phytin occurs in the upper intertidal zone. This area undergoes the
greatest fluctuation of physical and chemical factors and has the
lowest biomass. The high percentage of pheophytin which occurs in
this area of the intertidal may originate from detrital seaweed
washed up at the high tide line.
Another factor that may contribute
to the higher percentage of pheophytin in this sediment is a higher
mortality rate of the microalgae resulting from desiccation and other
environmental extremes which do not affect the microalgae closer to
the low tide line.
In considering vertical distribution of pheophytin the lower two
layers of the cores had a higher percentage whereas the upper two
136
layers had a lower percentage of pheophytin. Studies on sediment
movement over a four month period at Southbeach indicated that only
the upper 2.5 cm were disturbed by currents. This current action in
the sediment near the surface may remove much detrital material
and may account for the low percentage of pheophytin found in the
upper layers. Yentsch (1965) found that pheophytin forms when
microalgae are placed in the dark. Due to sedimentation, burrowing
invertebrates, and vertical migration, the microalgae may be carried
below the photic zone and over a period of time the chlorophyll might
degrade into pheophytin. This may account in part for the presence of
more pheophytin at the lower depths.
Bacterial action and the passage of microalgae through the gut
of mieofauna (Currie, 1962) have also been shown to degrade
chlorophyll to pheophytin. Microalgae and other detritus deeper in
the sediment come under the influence of biological activity which
causes the conversion of chlorophyll to pheo-pigments.
The pheo-pigments on the Sally's Bend tidal flat do not show any
significant trend as to vertical and horizontal distribution as is
evidenced on the Southbeach tidal flat. This is most likely due to a
difference in the sedimentary environments within the bay. At Sally's
Bend the sediment and allochthonous detritus carried in suspension by
the Yaquina River is deposited on the tidal flat. Since the primary
source of pheo-pigment is allochthonous detritus no distinct trend in
137
horizontal distribution is shown and the pheo-pigment will be evenly
deposited throughout the intertidal area.
On both the tidal flats studied there was no noticeable seasonal
variation in standing crop of microalgae as determined by cell counts
and concentrations of chlorophyll a. This absence of a seasonal
variation has also been noted by Williams (1962) on a mudflat in
Georgia. Steele and Baird (1968) found a relative constancy of
chlorophyll a seasonally although there were occasionally higher
values in the spring and summer.
In contrast to a more or less constant standing crop of benthic
microalgae, the standing crop of phytoplankton has been shown to
fluctuate seasonally. It has been suggested that a spring increase
occurs when mixing is reduced by temperature stratification and that
earlier in the year the cells did not remain in the photic zone long
enough to build up a large population (Round, 1965a). Subsequent to
spring growth nutrient replenishment is at a minimum and the
decrease in midsummer and subsequent recovery in the autumn may
result from partial depletion followed by replenishment as autumnal
mixing occurs (Round, 1965a).
The situation in an intertidal sand or mud flat is quite different
than the open ocean. Hendey (1951) noted that littoral diatoms do not
follow the course of seasonal occurrence usually found in the plankton
species which is characterized by two sharply marked peaks in spring
138
and autumn, but are found more or less evenly distributed throughout
the year. In estuarine situations the amount of nutrients may be
much larger depending on the amount of drainage from agricultural
land and from urban areas. Nutrients would therefore be available
to the benthic microalgae at all times of the year. This may account
for the relatively non-fluctuating standing crop of benthic microalgae
of estuarine sediments. Smyth (1955) states that the bottom diatom
flora, in contrast to the phytoplankton, appears to be a remarkably
stable population.
This is due to the apparent absence of great
fluctuations in the availability of nutrient salts. Also the large
numbers of bacteria which have been reported for marine sediments
may provide the microflora with a rich supply of ammonium, nitrate,
and phosphate through the mineralization of organic matter.
However, some investigators as Aleem (1950b) found that littoral
diatoms show a definite periodicity. In a study on the south coast of
England he found a period of maximum abundance in spring and a less
conspicuous period of abundance in early autumn. Castenholz (1967)
has also found seasonal patterns in nonplanktonic marine diatoms on
the western coast of Norway.
Whether a seasonal pattern exists or not seems to be dependent
on the substrate of the benthic microflora. Both Aleem (1950b) and
Castenholz (1967) were investigating epilithic diatoms whereas the
investigations showing no seasonal fluctuations were concerned with
139
either epipelic or epipsammic microalgae. Castenholz (1967) noticed
that attached diatoms in permanently submerged situations were more
continuous in their seasonal distribution without the hiatuses shown
by the epilithic flora on intertidal transects.
The differences in the seasonal pattern between the epilithic
flora and the flora on sediments may be due to differing nutrient
conditions.
The epipelic and epipsammic flora are continuously
exposed to nutrients which are probably present interstitially to a
large degree being constantly replenished by bacterial action. The
epilithic flora on the other hand is not continually surrounded by
nutrients and probably their seasonal abundance is regulated by the
amount of nutrition brought in with the tide. In time of upwelling or
deep mixing if the radiant energy is adequate a bloom similar to a
phytoplankton pulse may occur.
The experiments with the Gilson Differential Respirometer
supported the results obtained by estimation of chlorophyll a concen­
tration and by direct counts at the Southbeach study area. However,
the potential oxygen production at the Sally's Bend study area did not
correlate well with the chlorophyll a concentration.
In the Southbeach study area cores, the photosynthetic rate
decreased with an increase in depth below the sediment surface.
The
upper cubic centimeter had a potential rate of production approximately
3. 5 times as great as the 7. 8-9. 1 cm core layer. What is remarkable
140
is the fact that these lower depths had any photosynthetic activity at
all. Gridntved (1962 as cited in Pamatmat, 1968) found photosynthetic
activity down to a depth of 5 cm. Pamatmat (1968) demonstrated
photosynthetic activity up to 10 cm in depth below the sediment
surface
.
Is it possible that these intertidal microalgae are able to
form chlorophyll beneath the sediment surface and then photosyn­
thesize immediately upon exposure to light? The investigation of
Yentsch (1965) indicating that light is important for the maintenance
of chlorophyll in Phaeodactylum tricornutum would not seem to support
this theory. However, Harvey's (1953) experiments indicate that
this species can form chlorophyll in the dark. Pamatmat (1968)
has reported that the pennate littoral diatom Cylindrotheca fuciformis,
when grown in an inorganic medium plus thiamine and sodium lactate iu
the dark for three weeks can photosynthesize immediately when exposed
to light. These results may indicate that some diatoms can form
chlorophyll in the dark for short periods of time but after periods of
prolonged darkness the chlorophyll may eventually be degraded into
pheophytin. Since the surface sediment has been shown to be disturbed
by tidal currents only to a depth of 2.5 cm yet live chlorophyll bearing
algae capable of photosynthesis were present at depths of 9 cm, the
results suggest a vertical migration of the microalgae.
The greater rates of oxygen production from the cores taken
at the various intertidal levels coincided with the higher concentrations
141
of chlorophyll a and larger numbers of microalgae as determined by
direct counts.. The low tide level had the greatest potential rate of
oxygen production per hour whereas cores from the mid-intertidal
and upper intertidal zones had less oxygen produced per hour.
The most obvious difference between cores from the Sally's
Bend study area and the Southbeach study area is the low potential
production values at Sally's Bend. In many instances more oxygen
was consumed than produced. In all likelihood this is the result of the
smaller standing crop of benthic microalgae and the greater number of
bacteria that would exist in an area as rich in organic detritus as
the Sally's Bend tidal flat. ZoBell (1946) states that relatively few
bacteria are found in coarse sand and indicates that the size of the
bacterial population is closely related to the character of the
sediments. As a rule, the greater abundance of bacteria in the finer
sediments is attributed primarily to the higher organic content and
the fact that small particles offer much more surface area than
larger particles. Bacterial respiration is largely responsible for the
low rates of potential oxygen production.
At Sally's Bend there was no obvious pattern of distribution of
organic matter but a large variation existed in the amount present at
various locations. Since the distribution of bacteria is related to
organic matter this would account for the patchiness of production in
cores from the Sally's Bend study area.
142
Rivers entering most estuaries carry a load of negatively
charged organic particles which are precipitated by the positively
charged metallic ions of salt water (Darnell, 1967). The great bulk
of this organic detritus in estuaries is derived from vegetation.
These include autochthonous sources as phytoplankton, marginal
submerged vegetation, mud-flat diatoms, and periphyton growing on
stems of emergent plants and other surfaces. The allochthonous
sources can include river borne phytoplankton and detritus, beach
and shore detritus washed in during storms or during high water,
windblown material including leaves and pollen grains, and phyto­
plankton and other material originating in the adjacent marine
environment.
Due to the morphology of Yaquina Bay the organic particles
carried by the river are deposited on the Sally's Bend tidal flat.
Besides the deposition of the allochthonous fluviatile material men­
tioned above, several sawmills and wood processing plants on the
shore of Yaquina Bay contribute very large amounts of sawdust to the
sediment (Morrison, 1966). Ku lm (1965) noted that the particles of
sawdust occur in greatest abundance in the finer sediment of the tidal
flats. He determined the percentage of organic matter in the Yaquina
Bay sediment and obtained values ranging from 0. 86 to 3. 22% at Sally's
Bend as determined by wet oxidation with hydrogen peroxide.
The ash-free dry weight determinations indicate a wide range in
143
the amount of organic matter, with the cores from the Sally's Bend
tidal flat having much higher values than Southbeach. This would be
expected since the Southbeach tidal flat is not under the fluviatile
realm and would not receive much allochthonous material from the
river. Also, the tidal current is weaker in the Sally's Bend area and
less detrital material is removed. It has been noted by ZoBell (1946)
that higher concentrations of organic material tend to produce an
anaerobic environment because of bacterial action. Finer sediment
also decreases the rate of diffusion of the dissolved oxygen into the
interstitial water. These conditions at the Sally's Bend study area
may prevent a large standing crop of microalgae from developing.
The configuration of the bay and its resultant hydrography
determines the physical conditions which effect the distribution of the
microalgae present. The two tidal flats studied were exposed to
different realms of sedimentation and subsequently each consisted of
a different assemblage of minerals as well as being of different
textures. Exposure of the Southbeach tidal flat to marine conditions
contrasts sharply with the Sally's Bend tidal flat which is character­
istic of fluviatile conditions. The intertidal flora which may be
physically attached to the sand grains or which may be free-living in
the sediment is largely determined by the conditions of the substrate.
Particle size and organic content of the substrate play an important
role in the distribution of littoral species. Hendey (1964) indicates
144
that mud and sand provide warmth for the diatom community during
the colder months of the year and shelter from direct sunlight.
The
diatoms of the intertidal zone of Yaquina Bay were found to persist
more or less throughout the entire year with individual species
showing a certain amount of periodicity. Hendey (1964) also found this
to be the case in his study of British littoral diatoms.
An interesting feature noted in the distribution of diatoms in the
intertidal zone was that most of the species that were found in the
lower intertidal zone also occurred in the mid and upper intertidal
zones as well. This situation occurred both at the Southbeach and
Sally's Bend study areas. Smyth (1955) also found a similar occur­
rence in his investigation of Loch Sween. It is possible that some of
the diatoms may have been passively carried to the upper part of the
intertidal zone by tidal current action but the frequency with which
these species occurred in this zone probably indicates that these taxa
are present throughout the entire intertidal zone. The major difference
between the lower, mid, and upper intertidal zones is not a difference
in species composition but a difference in abundance.
Taxa from
the Monoraphidineae and Biraphidineae were prominent in the
flora even though many Araphidineae were also present. In an
unstable envirnoment which is exposed to sedimentaticn and burrowing
invertebrates, taxa with true raphes may have an advantage over
rapheless forms in that they are capable of motility. However, many
145
taxa without raphes such as Biddulphia alternans were present in
great abundence throughout the year and the absence of motility does
not seem to hinder this species.
A large number of diatom species are considered to be
cosmopolitan in distribution.
The prevalence of this type of distribu­
tion implies genetic stability over a long period, broad physiological
tolerance and a means of transportation (Curl, 1959). Cleve (1894­
1895) states that although many species are cosmopolitan there are
many species which occur only in certain seas and climates.
Patrick (1948) does not agree that diatoms are generally cosmopolitan
and states that endemism may range from 10 to 30% in a local flora.
She indicates that so little is known about diatoms in certain regions
of the world that it is difficult to draw any general conclusions
concerning the geographic distribution of diatoms and in many cases
identification of diatoms have been made by people unfamiliar with
the literature, Patrick (1948) also states that there is a great
similarity between the floras of eastern North America and western
Europe but that there is considerable difference between the diatom
floras of eastern and western United States.
The results of this taxonomic study of benthic diatoms supports
the cosmopolitan theory. Almost all the taxa identified from the
Yaquina Bay estuary have been described elsewhere from throughout
the world indicating an extremely wide distribution. Castenholz
146
(1963) states that almost all species found in Oregon also occur in
Britain and that most littoral and sublittoral diatom species appear to
be world-wide in distribution in north temperate shores. Several
species in this study could not be identified and these may be new to
science or they may have been reported in some systematic treatment
not examined by the investigator. Hendey (1958) reports that of 194
species reported from the littoral of West Africa, 100 were also found
in the Caribbean. Many of the diatom species reported from West
Africa were also found in Yaquina Bay.
It has generally been considered that centric diatoms are usually
planktonic while pennate diatoms are benthic. Snyth (1955) found
pennate forms the most important in his study of benthic diatoms and
states that bottom-living centric forms as Melosira sp. were rare.
On the other hand, Hustedt and Aleem (1951) found, in the littoral
zone, such species as Melosira sulcata which thrives equally well both
in the plankton and in the littoral region. The authors point out that
the origin of such species is in the littoral region and that later on
they are recruited into the plankton where they still retain the capacity
to multiply. Many centric diatoms were identified from the estuarine
sediments from Yaquina Bay. It is possible that these taxa were
carried in by the tide and were filtered out and remained in the
sediment as the tide ebbed. However, the frequency of occurrence of
many of these species suggested that these centrics can thrive equally
147
well in the benthic as well as in the planktonic habitat. Wood (1963)
thinks the idea that benthic diatoms are pennate while the phyto­
plankton are centric is a fallacy since he has found Coscinodiscus,
Auliscus, and Actinoptychus as part of the benthic flora in sediment of
a Texas bay.
It is true however, that some phytoplankters do settle out on
the sediment surface. On one occasion in this study Chaetoceros
didymus was found in abundance. Also occasionally auxospores of
Chaetoceros have been found. Wood (1963) has found spores of
Skeltonema costatum in surface sediments and these germinate in situ
before the forming of plankton blooms. Smyth (1955) also reported
finding spores of algae in the littoral zone.
Many species which are considered fresh-water forms have been
found in the sediment of Yaquina Bay as Achnanthes lanceolata,
Cymbella ventricosa, Diatoma hiemale, Diatoma vulgare, and
Gomphonema angustatum.
Since identification of the diatoms is
dependent on the cleaned silicious frustule there is no way of deter­
mining whether these taxa are actually part of the benthic estuarine
community or are allochthonous material washed down by the river
and deposited in the sediment. Since the frequency of finding these
typical fresh-water forms is low the latter is the most plausible
explanation. Hustedt and Aleem (1951) found that the flora of mud
flats consist of taxa from three allochthonous sources: 1) attached
148
forms from rocky shores, 2) pelagic forms from the plankton, and
3) fresh-water forms from rivers and lakes.
The diatom taxa identified indicated that the study area is a
typical coastal brackish water environment (Hendey, 1964). Many of
the species identified have never been found in Oregon prior to this
study.
This study extends the distribution of many taxa which have
been described from other areas throughout the world.
149
SUMMARY
1.
In vertical distribution, the greatest biomass of microalgae was
found in the upper centimeter of sediment at the time collections were
made. This was attributed to the fact that light extinction in sediment
is very rapid and no light was found to penetrate more than a few
millimeters in depth. The motile epipelic algae migrate to the
surface of the sediment because only in this area is autotrophic
growth possible.
2.
The large number of microalgae found below the photic zone was
attributed to a vertical migration rhythm which is thought to be
sensitive to light and also involves a positive geotaxis. Viable di­
atoms were found at 18 cm beneath the sediment surface.
3.
The greatest biomass of microalgae was found at the lower inter­
tidal zone with the upper intertidal zone having the lowest biomass.
This distribution is thought to be the result of fluctuation in various
environmental factors as temperature, salinity, oxygen content, and
water content which vary with intertidal height. The upper intertidal
zone has a longer period of exposure and thus has greater fluctuations
in the various factors than the lower intertidal zone. The fact that the
microalgae at the lower intertidal zone would be exposed to nutrientcontaining seawater for a longer period of time may also explain a
greater biomass at the lower intertidal zone.
150
4.
The absence of seasonal fluctuations as exhibited by the phyto­
plankton is thought to be the result of a continuous supply of nutrients
being carried down by the river or regenerated in situ by bacterial
action in the sediment.
5.
The low standing crop present on the Sally's Bend tidal flat
relative to the Southbeach tidal flat has been attributed to the hydro­
graphy of the bay and the resultant realms of deposition. Fine silt
and organic debris on the Sally's Bend flat results in a decrease in
light penetration, an increase in bacterial activity and a resulting
decrease in oxygen content. These conditions would tend to keep the
standing crop of microalgae minimal. Also an extensive grazing by
invertebrates on the tidal flat may also keep the standing crop low.
6.
The low potential gross productivity of the algae in subsurface
sediment from the Sally's Bend tidal flat is most likely due to the
large biomass of bacteria which exist in this type of sediment. Res­
piration of a sample core often exceeded photosynthesis. Since
fewer bacteria are likely to be found in coarse sediment and the
standing crop of microalgae was higher, rates of oxygen production
exceeded respiration in sample cores from the Southbeach tidal flat.
7.
Pheopigments were abundant on both tidal flats but more so on
Sally's Bend. Since it has been shown that there is more organic
detritus on this tidal flat it is logical to assume that it is the result
of the degradation of chlorophyll of allochthonous material. The
151
greater amount of pheophytin at the lower depths sampled at Southbeach
is thought to be caused by the death of microalgal cells which have
been in the dark for long periods of time.
The greater amount of
pheophytin at the high tide level is thought to be due to detrital sea­
weed decomposition as well as autochthonous microalgae not capable
of withstanding fluctuations of various environmental factors.
8.
Most of the species that were found in the lower intertidal zone
also occurred in the mid and upper intertidal zones. The major
difference between these zones is not one of species composition but
one of biomass.
9.
The microflora is almost exclusively composed of diatoms.
Species from the Biraphidineae predominate but species from the
Monoraphidineae and Araphidineae are abundant as well. The taxa
identified from the sediments in the bay are typical of littoral areas
throughout the world and many of the species are considered cosmo­
politan in distribution. Many species identified have not yet been
reported from Oregon and this study constitutes new distribution
records for many of these species.
152
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APPENDICES
Appendix Table 1. Number of cells/cm 3 in the sediment layers 0. 0-1. 3 cm and 1. 3-2. 6 cm below
the surface at Southbeach. Each value is the mean of 2 or 3 samples.
.
Date
Lower
Intertidal
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
12-27-67
1-25-68
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
*
0. 0-1. 3_cm Layer
Mid
Upper
Intertidal
*
1.
Intertidal
*
3-2. 6 cm Layer
Lower
Mid
Intertidal
Upper
Intertidal
Intertidal
*
*
30, 160
23,760,
21, 093,
30, 520*
23, 973*
10, 240;
36, 560
41, 680
57, 200
40, 480
18, 693
15, 014
28, 959
30, 080
11, 960
27, 000
12, 800
43, 800
23, 280*
39, 200*
32, 640
24, 880
45, 560
35, 400
47,000
39,333
45,240
43,920
31,227
44, 160
53, 720
54, 560
44, 613
43, 786
42, 399
41, 386
34, 772
17, 280
41, 440
47, 200
41, 440
48, 360
46, 480
37,720
51,920
49, 520
32, 640
39, 560
39, 160
37, 413
35, 146
43, 893
43, 199
29, 413
51, 279
42,920
46, 160
52, 400
44, 640
0.,
..,,
Counts of only epipelic algae.
16, 960*
24, 000,
18, 213
12, 187
22, 427
23, 866
*
19, 800*
11, 680*
12, 440
9, 320
11,720
41, 600
29,200
18, 240
27, 080
39, 340
18, 060
40,720
37, 480
Appendix Table 2. Number of cells/cm3 in the sediment layers 5. 2-6. 5 cm and 7. 8-9. 1 cm beloW
the surface at Southbeach. Each value is the mean of 2 or 3 samples.
Date
5. 2-6. 5 cm Layer
Lower
Intertidal
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
12-27-67
1-25-68
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
*
10,960,0,
29, 440*
31, 280
26, 080
40, 160
44, 000
16, 880
28,720
52, 840
50,560
31, 640
49,720
53, 800
7. 8-9. 1 cm Layer
Mid
Upper
Mid
Upper
Lower
Intertidal
Intertidal
Intertidal
*
7,440,
*
4, 986,
11, 466;
9, 253
6, 320
13, 546
3,760
9, 520
23, 226
25, 467
32, 666
32, 026
21, 194
30, 373
Counts of only epipelic algae.
820 *
720
10,945
1, 680
4, 200
7, 640
6,720
14,720
30, 680
7, 040
5,960
6,920
4, 400
*
20,480
-...
Intertidal
:,'4
4, 400*
18,933- ,,
Intertidal
320
440
21, 240
23, 560
26, 680
6,213
10,960
5,786
5, 880
5, 440
7, 760
35,720
7,960
38,720
4, 693
10, 320
17, 019
36, 293
34, 559
30, 479
25, 573
42, 800
15,706
17, 040
13, 360
31, 160
47, 440
26, 000
5, 000
9, 360
3, 200
6,400
4,960
5, 160
3, 000
*
Appendix Table 3. Biomass expressed as concentration of chlorophyll a in pecm 3 in the sediment
layer 0.0-1.3 cm below the surface. Each value is the mean of 2 or 3 samples.
Date
Southbeach
Lower
Mid
Intertidal
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
19.5
21.5
19.5
12-27-67
1-25-68
17.5
18.0
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
23.5
26,0
17.0
16.5
27.5
21.5
41.0
33.0
Intertidal
27.5
6.0
15.0
17.0
15.5
12.0
17.0
35.0
24.5
27.5
23.0
53.0
37.5
Upper
Intertidal
Date
Sally's Bend
Lower
Mid
Upper
Intertidal
Intertidal
Intertidal
8-23-67
9-20-67
10-18-67
11-15-67
12-13-67
10.0
21.5
17.0
16.5
12.5
11.5
17.0
13.5
10.5
16.0
21.0
8.5
7.5
10.5
14.0
20.0
1-11-68
2- 9-68
3- 9-68
4- 9-68
5- 7-68
10.0
15.0
10.5
9.5
11.0
8.5
11.5
6-10-68
7-14-68
9.5
9.5
9. 5
9. 5
9. 5
10.0
12.0
9. 5
23.5
16.5
11.5
28.0
7.5
19.5
21.0
43.5
40.0
40.0
27.0
9.0
8.0
5.5
11.0
9. 5
8.0
12.5
Appendix Table 4. Biomass expressed as concentration of chlorophyll a in pg /cm3 in the sediment
layer 1.3-2. 6 cm below the surface. Each value is the mean of 2 or 3 samples.
Date
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
12-27-67
1-25-68
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
Southbe ach
Lower
Mid
Intertidal
20.5
23.5
21.5
25.5
21.0
15.0
18.5
23.5
23.5
27.0
24.5
34.5
35.0
* No date available.
Intertidal
21.0
17.0
12.5
15.5
16.0
11.5
12.0
21.0
18.0
21.0
30.5
33.0
24.5
Upper
Intertidal
10.0
7.5
10.0
11.0
12.0
12.5
8.0
13.0
11.5
35.5
25.5
23.5
15.0
Date
8-23-67
9-20-67
10-18-67
11-15-67
12-13-67
1-11-68
2- 9-68
3- 9-68
4- 9-68
5- 7-68
6-10-68
7-14-68
Sally's Bend
Lower
Mid
Upper
Intertidal
Intertidal
Intertidal
9.5
11.5
12.5
12.0
10.0
9. 0
10.0
11.5
11.0
8.5
10.5
7.5
9.0
8. 0
9. 0
11.0
6.5
10.5
8.5
10.0
11.0
10.5
4.5
9.0
14.5
12.0
11.0
12.5
9.0
7.5
10.5
8.5
10.0
9. 0
5. 5
Appendix Table 5. Biomass expressed as concentration of chlorophyll a in pg/ ,cm 3 in the sediment
layer 5.2-6.5 cm below the surface. Each value is the mean of 2 or 3 samples.
Date
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
12-27-67
1-25-68
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
Southbeach
Lower
Mid
Intertidal
10.5
19.5
19.0
20.0
19.0
17.5
21.0
20.5
17.5
28.5
26.0
29.5
25.5
Intertidal
Upper
Intertidal
10.0
7.5
1.0
4.5
5.0
11.0
3.0
7.5
8.5
1.0
9.5
15.5
8.0
13.0
15.0
13.0
14.0
2.0
Date
5.0
4.0
5.5
5.0
Intertidal
7.5
8-23-67
9-20-67
10-18-67
11-15-67
12-13-67
15.5
13.5
12.5
1-11-68
8.0
2.5
13.5
15.5
7.5
Sally's Bend
Lower
Mid
2- 9-68
3- 9-68
4- 9-68
5- 7-68
6-10-68
7-14-68
9.5
9.0
8.5
7.0
10.0
8.5
8.0
Intertidal
12.5
8.5
9.5
8.0
6.0
7.0
8.5
6.0
7.5
9.5
9.5
8.0
Upper
Intertidal
8.5
7.5
9.5
5.0
8.0
7.5
8.0
5.5
5.5
8.0
15.5
8.5
Appendix Table 6. Biomass expressed as concentration of chlorophyll a in ilg/, cm 3 in the sediment
layer 7. 8-9. 1 cm below the surface. Each value is the mean of 2 or 3 samples.
Date
Southbeach
Lower
Mid
Intertidal
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
11. 5
12. 0
11. 5
15. 5
14. 0
12-27-67
1-25-68
22.5
18.5
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
17. 5
15. 5
20.5
32.0
17. 5
22.5
Upper
Intertidal
Intertidal
Date
7. 5
13. 5
3. 5
8. 0
3. 5
2. 5
11. 0
15. 5
14. 5
15. 0
20. 0
15. 5
8. 5
1. 5
8-23-67
9-20-67
10-18-67
11-15-67
12-13-67
1.5
10. 0
9. 5
7. 0
5. 5
6. 0
11. 0
4. 0
7. 5
5. 5
6. 0
5. 5
1-11-68
2- 9-68
3- 9-68
4- 9-68
5- 7-68
6-10-68
7-14-68
Sally's Bend
Lower
Mid
Intertidal
8.5
Intertidal
7.5
9. 5
9. 5
8. 0
7. 5
6. 0
7.0
7.0
7.5
7.0
9.0
7.5
9.0
6. 0
6. 5
6. 0
6. 0
8. 0
8. 0
9. 0
9.5
15.0
9.0
Upper
Intertidal
7. 0
7.0
9. 0
4. 5
5. 5
7.5
6. 5
7. 5
6. 5
8. 5
11. 5
7.5
Appendix Table 7. Concentration of pheophytin, expressed as percentage of total chlorophyll
(chlorophyll a and pheophytin), in the sediment layer 0. 0-1. 3 cm below the
surface. Each value is the mean of 2 or 3 samples.
Southbeach
Lower
Mid
Date
Intertidal
Intertidal
Upper
Intertidal
Date
Sally's Bend
Lower
Mid
Intertidal Intertidal
Upper
Intertidal
J.
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
12-27-67
1-25-68
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
12, 0
2.5
3. 0
2. 5
7. 0
6. 5
3. 0
0. 0
8.5
14.5
30. 5
26. 5
J.
No data available.
13. 0
8. 0
23. 0
12. 0
13. 0
14. 0
10. 0
7. 0
19. 0
10. 0
36. 0
23. 0
31. 0
8. 0
39. 0
19. 0
3. 0
6. 0
9. 0
5.5
31. 5
19. 0
15.5
6. 0
8-23-67
9-20-67
10-18-67
11-15-67
12-13-67
61.5
73.5
62.0
58.5
60.0
57. 0
56. 0
60.6
68.6
51.5
1-11-68
59.5
60.5
61.5
63.0
56.0
53.0
61.0
56. 0
55. 0
60. 0
61. 0
55. 3
59. 6
61. 0
2- 9-68
3- 9-68
4- 9-68
5- 7-68
6-10-68
7-14-68
62. 0
67.6
66. 5
56. 0
55. 0
57. 0
67.0
77. 0
54. 5
61. 0
66. 0
57. 0
Appendix Table 8. Concentration of pheophytin, expressed as percentage of total chlorophyll
(chlorophyll a and pheophytin), in the sediment layer 1.3-2.6 cm below the
surface. Each value is the mean of 2 or 3 samples.
Date
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
Southbeach
Lower
Mid
Intertidal
1.5
4.0
6.5
3.0
0.0
12-27-67
1-25-68
12.5
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
3.0
0.0
9.0
3.0
22.0
24.0
No data available.
Intertidal
17.5
12.0
13.0
12.0
14.0
8.0
16.0
6.0
10.0
15.0
29.0
11.0
Upper
Intertidal
40.0
12.0
26.0
18.5
5.5
15.2
22.0
15.0
21.5
15.0
16.0
14.5
Date
Sally's Bend
Lower
Mid
Intertidal
Intertidal
8-23-67
9-20-67
10-18-67
11-15-67
12-13-67
60.0
68.0
65.5
55.0
51.5
63.0
52.0
1-11-68
58.0
56.0
61.0
62.3
55.6
53.3
64.6
56.0
58.6
2- 9-68
3- 9-68
4- 9-68
5- 7-68
6-10-68
7-14-68
62.5
59.0
57.0
59.0
58.3
55.0
58.3
*
Upper
Intertidal
62.0
64.0
60.5
59.0
58.0
60.5
70.5
80.0
60.0
60.0
71.0
87.0
Appendix Table 9. Concentration of pheophytin, expressed as percentage of total chlorophyll
(chlorophyll a and pheophytin), in the sediment layer 5. 2-6. 5 cm below the
surface. Each value is the mean of 2 or 3 samples.
Date
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
12-27-67
1-25-68
2-23-68
3-19-68
4-18-68
5-16-68
6 -28 -68
7-26-68
Southbeach
Lower
Mid
Intertidal
8. 0
6. 5
5. 0
3. 0
0. 0
1. 5
5. 0
1. 5
6. 5
16. 0
17. 0
6. 5
No data available.
Intertidal
33. 0
52. 0
51. 0
37. 0
66. 0
40. 0
33. 0
14. 0
24. 0
15. 0
43. 0
11. 0
Upper
Intertidal
73. 0
2 8. 0
58. 0
39. 0
46. 5
52. 0
17. 0
5 8. 5
53. 5
42. 0
43. 0
54. 5
Date
Sally's Bend
Lower
Mid
Intertidal Intertidal
63.0
61.0
8-23-67
9-20-67
10-18-67
11-15-67
12-13-67
5 8. 0
57.5
53.5
58.6
1-11-68
62.5
68.3
60.0
64.6
59.3
59.9
59.6
63.3
2- 9-68
3- 9-68
4- 9-68
5- 7-68
6-10-68
7-14-68
5 8. 5
56.5
61.0
64.0
63.0
63.0
54. 5
55. 0
61. 6
64. 0
Upper
Intertidal
68. 5
7 1. 5
62. 0
69. 5
56. 0
62. 5
62. 0
77. 0
71. 5
60. 0
45. 0
67. 5
Appendix Table 10. Concentration of pheophytin, expressed as percentage of total chlorophyll
(chlorophyll a and pheophytin), in the sediment layer 7. 8-9. 1 cm below the
surface. Each value is the mean of 2 or 3 samples.
Date
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
12-27-67
1-25-68
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
Southbeach
Lower
Mid
Intertidal
10. 0
31.0
12. 5
3.0
6.5
7.0
6.0
3.0
2.0
30.0
10. 0
2. 5
No data available.
Intertidal
27. 0
60. 0
54. 0
47. 0
45. 0
32. 0
23. 0
21. 0
11. 0
7. 0
17. 0
27. 0
Upper
Intertidal
87. 0
65. 0
45. 5
44. 0
60. 0
30. 0
20. 0
37. 5
16. 0
38. 5
39. 5
50. 0
Sally's Bend
Lower
Mid
Upper
Date
Intertidal
8-23-67
9-20-67
10-18-67
11-15-67
12-13-67
67.0
62.5
61.5
59.5
59.0
59. 0
58. 3
64. 3
54. 6
63. 3
72. 0
77. 0
1-11-68
61.0
59.5
61.0
63.0
68.5
55.6
64.3
64.3
65.0
63.0
65.0
64.3
53. 5
66. 0
65. 5
70. 0
69. 5
56. 0
66. 5
2- 9-68
3- 9-68
4- 9-68
5- 7-68
6-10-68
7-14-68
63. 5
62.5
Intertidal
Intertidal
65. 5
76. 0
75. 0
Appendix Table 11. Rates of gross production in pg 02/cm3/hr in the sediment layer 0.0-1.3 cm
below the surface. Each value is the mean of 2 or 3 samples.
Southbeach
Lower
Mid
Upper
Date
Intertidal
Intertidal
Intertidal
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
85.27
117.63
110.08
116.64
147.16
136.61
191.45
26.83
163.35
144.73
129.81
211.61
165.29
76.59
86.74
54.11
88.24
88.00
68.52
90.35
54.13
140.58
73.29
60.63
93.84
137.26
62.56
100.76
172.36
21.11
96.14
159.29
153.70
202.53
95.17
12-27-67
1-25-68
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
167.81
139.92
234.75
105.98
Date
8-23-67
9-20-67
10-18-67
11-15-67
12-13-67
1-11-68
2- 9-68
3- 9-68
4- 9-68
5- 7-68
6-10-68
7-14-68
Sally's Bend
Lower
Mid
Intertidal
19.19
22.21
13.97
- 30.16
- 7.05
0.81
17.97
- 47.55
-
-22.98
- 31.75
- 56.04
27.14
Intertidal
Upper
Intertidal
86.96
61.50
4.89
43.83
74.27
38.48
35.15
50.68
43.44
14.91
65.59
54.12
84.93
60.17
24.08
51.18
43.90
50.05
51.81
65. 66
61.88
58.34
55.33
40.37
A
Appendix Table 12. Rates of gross production in p.g 02/cm3/hr in the sediment layer 1. 3-2. 6 cm
below the surface. Each value is the mean of 2 or 3 samples.
Date
1
1
1
1
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
12-27-67
1-25-68
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
Southbeach
Lower
Mid
Intertidal
47. 66
Intertidal
124. 17
73. 60
100.31
176.37
163.93
170.04
116.93
178.50
48. 13
192.06
75.13
81.59
68.25
52.71
72.99
56.57
75.52
179. 34
140. 47
215.21
142.53
130.87
118. 50
117. 32
95.02
Upper
Intertidal
16.02
12.11
67.58
32.76
66.11
55.40
68.18
2.43
35.95
113.34
45.56
61.80
77.19
Date
Sally's Bend
Lower
Mid
Intertidal
Intertidal
Upper
Intertidal
8-23-67
9-20-67
10-18-67
11-15-67
12-13-67
3. 49
13. 83
- 16. 69
31. 34
- 25.51
36.78
74. 23
- 11. 80
- 15. 83
- 3. 36
16.76
41. 00
4.96
1-11-68
- 2. 62
5.82
41.20
41.20
- 36. 49
- 72. 64
- 39. 22
- 4. 32
- 51. 31
8. 98
3. 63
10. 62
21. 25
40. 83
2- 9-68
3- 9-68
4- 9-68
5- 7-68
6-10-68
7-14-68
40. 84
47. 80
9.18
-
29.69
22.85
17.76
3.13
5.13
23.22
Appendix Table 13. Rates of gross production in t_ig 02/cm3/hr in the sediment layer 5.2-6.5 cm
below the surface. Each value is the mean of 2 or 3 samples.
Southbeach
Lower
Mid
Date
Intertidal
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
23.68
78.09
12-27-67
1-25-68
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
82.07
106.21
136.56
133.90
88.00
71.13
136.35
136.14
80.34
110.29
131.85
Intertidal
30.67
27.64
16.67
25.08
59.47
14.31
34.91
54.95
35.12
53.22
66.61
33.25
68.88
Upper
Intertidal
5.78
10.90
57.61
27.18
40.10
21.94
33.68
2.89
79.21
9.54
7.98
3.16
- 6.15
Date
Sally's Bend
Lower
Mid
Intertidal
Intertidal
8-23-67
9-29-67
10-18-67
11-15-67
12-13-67
18.10
8.06
- 15.13
- 12.40
- 3.67
43.51
46.17
14.51
21.94
30.64
1-11-68
- 15.79
- 14.21
- 12.53
26.84
26.84
28.40
- 5.29
11.04
2- 9-68
3- 9-68
4- 9-68
5- 7-68
6-10-68
7-14-68
-20.98
-
8.92
3.62
27.61
15.27
23.85
Upper
Intertidal
6.65
14.24
30.99
10.91
- 0.30
-
4.80
8.34
30.72
2.04
-25.35
- 19.52
- 0.67
Appendix Table 14. Rates of gross production in p.g 02/cm3/hr in the sediment layer 7.8-9.1 cm
below the surface. Each value is the mean of 2 or 3 samples.
Southbeach
Lower
Mid
Date
Intertidal
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
26.54
58.37
37.55
1.34
93.25
99.31
12-27-67
1-25-68
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
114.33
68.70
147.12
149.08
80.35
84.53
90.58
Intertidal
5.92
33.83
31.64
- 0.21
20.46
42.01
91.91
80.18
35.12
73.90
96.04
52.57
38.18
Upper
Intertidal
3.32
7.98
48.49
0.53
18.30
27.73
40.63
67.75
33.07
12.93
17.99
7.35
2.90
Date
Sally's Bend
Lower
Mid
Intertidal Intertidal
8-23-67
9-29-67
10-18-67
11-15-67
12-13-67
- 11.58
9.88
0.66
- 8.49
0.76
26.27
39.11
1-11-68
- 12.73
1.67
0.60
- 14.83
- 17.13
- 18.69
- 4.03
10.47
10.47
2- 9-68
3- 9-68
4- 9-68
5- 7-68
6-10-68
7-14-68
Upper
Intertidal
-42.48
19.63
9. 02
2.26
16.62
14.79
37.75
28.63
23.72
- 8.92
12.53
23.71
31.86
-
10.34
7.61
11. 01
- 14.01
- 11.10
4.25
- 8.05
Appendix Table 15. Biomass expressed as ash-free dry weight in mg /cm3 in the sediment layer
0. 0-1. 3 cm below the surface. Each value is the mean of 2 or 3 samples.
Southbeach
Lower
Mid
Upper
Date
Intertidal
Intertidal
Intertidal
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
40.5
31.7
65.8
37.9
45.3
36.7
32. 1
29. 6
134. 1
28.9
244.9
35.2
54.3
26.8
33.6
32.5
34.7
29.6
36.0
36.5
149.8
105.8
37.4
18.5
37.7
42.8
12-27-67
1-25-68
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
Date
Sally's Bend
Lower
Mid
Upper
Intertidal
Intertidal
Intertidal
8-23-67
9-20-67
10-18-67
11-15-67
12-13-67
124.7
165.0
102. 9
505.6
254.2
79.5
78.8
91.8
100.5
97. 8
21.3
1-11-68
27. 1
2- 9-68
3- 9-68
4- 9-68
5- 7-68
90.2
72.6
79.1
87.7
115. 4
87. 4
97. 4
120. 6
143.3
22. 8
39. 4
39.7
43.2
38.0
42.5
36.9
34.8
23.2
51.9
6-10-68
7-14-68
234. 6
39. 5
83.7
90.4
96. 1
240. 2
72.7
82.2
121.1
104.7
109.4
78. 8
91. 1
83. 3
93. 2
83. 7
122. 7
86. 4
3
Appendix Table 16. Biomass expressed as ash-free dry weight in mg/cm in the sediment layer
1. 3-2. 6 cm below the surface. Each value is the mean of 2 or 3 samples.
Southbeach
Mid
Lower
Upper
Date
Intertidal
Intertidal
Intertidal
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
33. 9
31. 5
33. 8
23. 2
55. 8
46.6
37.0
37.7
37.6
12-27-67
1-25-68
33. 3
19. 1
30. 1
28.7
24.7
33. 5
28. 4
30. 3
24. 9
26. 0
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
29.7
31. 9
31.7
32. 6
21. 5
33. 9
23. 4
41. 7
26. 6
24. 5
26.4
45.5
36.4
31.8
29.4
35.7
43.4
46.8
31.2
35.9
Date
Sally's Bend
Mid
Lower
Intertidal
Intertidal
Intertidal
111. 1
104. 3
277. 5
114. 1
96. 0
93. 3
132.1
358.7
118.4
103.3
103.7
106. 5
102. 5
111. 0
112. 3
100. 0
117. 1
87. 9
142. 1
90. 2
8-23-67
9-20-67
10-18-67
11-15-67
12-13-67
290.4
79.5
92.2
97.9
1-11-68
102. 1
2- 9-68
3- 9-68
4- 9-68
5- 7-68
78.8
78.0
6-10-68
7-14-68
Upper
85.0
85.1
113. 5
92.6
104.8
57.8
103. 4
96. 3
89.8
Appendix Table 17. Biomass expressed as ash-free dry weight in mg/cm 3 in the sediment layer
5.2-6.5 cm below the surface. Each value is the mean of 2 or 3 samples.
Southbeach
Lower
Mid
Upper
Date
Intertidal
Intertidal
Intertidal
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
35.1
29.0
57.2
37.8
20.7
63.7
106.3
35.1
83.7
32.7
61.6
39.2
53.9
12-27-67
1-25-68
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
45.6
50.9
22.9
64.6
41.6
44.5
35.0
25.2
32.3
50.5
35.0
32.4
28.3
73.8
45.9
56.0
34.8
31.2
45.1
38.3
45.2
39.9
48.8
41.4
30.0
45.0
Date
Sally's Bend
Lower
Mid
Intertidal Intertidal
Upper
Intertidal
8-23-67
9-20-67
10-18-67
11-15-67
12-13-67
201,1
131.5
109.4
94.3
96.9
149.5
176.7
125.8
100.5
114.7
581.1
239.4
105.5
1-11-68
106.9
78.1
97.1
90.2
97.7
130.9
102.3
100.1
120.7
104.2
2- 9-68
3- 9-68
4- 9-68
5- 7-68
6-10-68
7-14-68
175.9
95.1
97.8
128.6
105.0
97.0
105.2
94.4
119.0
115.0
103.1
244.6
85.0
Appendix Table 18. Biomass expressed as ash-free dry weight in mg/cm 3 in the sediment layer
7. 8-9. 1 cm below the surface. Each value is the mean of 2 or 3 samples.
Southbeach
Lower
Mid
Date
Intertidal
8- 9-67
9- 4-67
10- 4-67
11- 3-67
12- 1-67
29.9
34.2
64.6
37.1
42.6
49.8
46.5
30.3
21.4
35.2
31.0
29.6
12-27-67
1-25-68
2-23-68
3-19-68
4-18-68
5-16-68
6-28-68
7-26-68
28. 0
Intertidal
33. 6
48. 6
213. 3
45.9
66.4
27. 2
47. 3
54. 2
25. 3
37. 0
33. 6
34. 2
27. 7
Upper
Intertidal
74.2
44.5
75.1
34.9
44.8
Date
Sally's Bend
Lower
Mid
Upper
Intertidal
Intertidal
Intertidal
8-23-67
9-20-67
10-18-67
11-15-67
12-13-67
156.1
134.3
224.7
93.4
214. 4
199. 0
135.7
80.1
104.6
86. 1
100.7
85. 3
101. 0
1-11-68
99. 5
106. 1
114. 1
83.4
92.7
97.4
99. 8
118.7
111. 7
112. 8
93. 9
105. 1
110. 2
127. 2
85. 8
119. 8
99.2
105.7
196. 2
127.7
55. 2
80.2
27.0
28.9
58.6
46.2
39.6
55.9
2- 9-68
3- 9-68
4- 9-68
5- 7-68
105. 1
6-10-68
7-14-68
122.0
96.3
Plate I
Figure 1.
Achnanthes brevipes Agardh
Figures 2, 3.
Achnanthes haukiana Grunow
Figures 4, 5.
Achnanthes lanceolata (de Brebisson) Grunow
Figures 6, 7.
Achnanthes sp.
Figures 8, 9.
Achnanthes sp. 2
Figure 10.
Actinoptychus senarius Ehrenberg
Figure 11.
Actinoptychus marmoreus Brun
1
co
atinlatitcoy.tia
Ui
Plate II
Figure 1. Amphiprora alata (Ehrenberg) Kiltzing
Figure 2. Amphora elegans Peragallo
Figure 3. Amphora granulata Gregory
Figure 4. Amphora lineolata Ehrenberg
Figure 5. Amphora ocellata Donkin
Figure 6. Amphora ostrearia de Brebisson
Figure 7. Amphora truncata Gregory
Figure 8. Amphora proteus Gregory
C1
Plate III
Figure 1.
Amphora angusta var typica Cleve
Figure 2.
Amphora sp.
Figure 3.
Anaulus balticus von Riemer Simonson
Figure 4.
Auliscus coelatus Bailey
Figure 5.
Aulocodiscus probabilis A. Schmidt
Figure 6.
Bacillaria paxillifer (0. F. Willer) Hendey
Figures 7, 8.
Biddulphia alternans (Bailey) Van Heurck
Figure 9.
Biddulphia aurita (Lyngbye) de Br'bisson
1
CO
(NI
I
!,1 AA
1
1
o.
tX1
Plate Ili
Figure 1.
Biddulphia dubia (Brightwell) Cleve
Figure 2.
Biddulphia longicruris Greville (600x)
Figures 3, 4.
Caloneis brevis (Gregory) Cleve
Figure 5.
Campylodiscus echeneis Ehrenberg (600x)
Figure 6.
Caloneis westii (Wm Smith) Hendey
Figure 7.
Campylosira cymbelliformis (A. Schmidt) Grunow
00
LO
Plate V
Figure 1.
Cerataulus turgidus (Ehrenberg) Ehrenberg
Figure 2.
Chaetoceros didymus Ehrenberg
Figures 3, 4.
Cocconeis costata var pacifica (Grunow) Cleve
Figures 5, 6.
Cocconeis sp. 2
Figure 7.
Cocconeis sp.
Figures 8.9.
Cocconeis clandestina A. Schmidt
Figure 10.
Cocconeis dirupta Gregory
Figures 11, 12.
Cocconeis placentula var euglypta (Ehrenberg) Cleve
Figures 13, 14.
Cocconeis decipiens Cleve
Figure 15.
Cocconies scutellum Ehrenberg
1
4.1
CNA
Plate VI
Figure 1.
Coscinodiscus curvatulus Grunow
Figure 2.
Coscinodiscus eccentricus Ehrenberg
Figures 3, 4.
Coscinodiscus lineatus Ehrenberg
Figure 5.
Coscinodiscus marginatus Ehrenberg
Figure 6.
Coscinodiscus sp. 1 (800x)
Figure 7.
Coscinodiscus nitidusl Gregory
ftop
qbes.
ibTes,Z,
.4,t
.,
.0
es
t
-
,(
; *fl0114.Ott
0111
te
..11114
185
Plate VII
Figure 1.
Coscinodiscus radiatus Ehrenberg
Figure 2.
Cymbella mexicana Ehrenberg
Figure 3.
Cymbella ventricosa Kiitzing
Figure 4.
Diatoma hiemale var.mesodon (Ehrenberg) Grunow
Figure 5.
Diatoma vulgare Bory
Figures 6, 7. Dimmerogramma minor (Gregory) Ralfs
Figures 8. 9.
Dimmergramma sp. 1
Figure 10.
Donkinia recta (Donkin) Grunow ex Van Heurck
Figure 11.
Diploneis bombus Ehrenberg
Figure 12.
Diploneis crabro Ehrenberg
Figure 13.
Diploneis interrupta (Kiitzing) Cleve
Figure 14.
Diploneis smithii (de Brebisson) Cleve
E
),
4
.11
1
II
A
eh
..111
1
b
1
1
!
..
.4
4,..
4
a
'
a
,
,
r
" °,
11,
0.
se
* 04 ov elhos
7.
01
II. '610' rev
II
ar
co
is is
":,
-­
aV.O.,
11.
. rte~
.P*
1O
be.
00
*
'
Plate VIII
Figure 1.
Epithemia sorex Hazing
Figure 2.
Epithemia turgida Hazing
Figure 3.
Eunotogramma laeve Grunow
Figure 4.
Eunotogramma marinum (Wm. Smith) Peragallo
Figure 5.
Fragillaria pinnata Ehrenberg
6.
Grammatophora
Grunow
Figure 7.
Grammatophora angulosa Ehrenberg
Figure 8.
Grammatophora marina (Lyngbye) Kiitzing
Figure 9.
Gyrosigma fasciola (Ehrenberg) Cleve
Figure 10.
Gyrosigma obliquum Grunow apud. Cleve and Grunow
Figure 11.
Gyrosigma wansbeckii (Donkin) Cleve
Figure 12.
Gomphoneis herculiana var. robusta Grunow
Figure 13.
Gomphonema angustatum var. subcapitata Hazing
t
000\1\\\!""1111111
quilitillibilill11
Plate IX
Figure 1.
Gyrosigma balticum (Ehrenberg) Cleve
Figure 2.
Gyrosigma sp.
Figures 3, 4.
Hantzschia virgata (Roper) Grunow
Figures 5, 6.
Hantzschia marina (Donkin) Grunow
1
40141111116iiiim1111111111111111111iiitt.
7
ilit er,
I 14
f.11414hit
.
.
r
47;
ino
Plate X
Figure 1.
Isthmia nervosa Kiitzing (600x)
Figure 2.
Licmophora abbreviata Agardh
Figure 3.
Licmophora ehrenbergii (Kiitzing) Grunow
Figure 4.
Licmophora gracilis (Ehrenberg) Grunow
Figure 5.
Licmophora tenuis (Kiitzing) Grunow
Figure 6.
Mastogloia exigua Lewis
Figure 7.
Melosira moniliformis (Muller) Agardh
Figure 8.
Melosira sulcata (Ehrenberg) Kiitzing
Figure 9.
Navicula clavata Gregory
Figure 10.
Navicula abscondita Hustedt
Figure 11.
Navicula arenaria Donkin
Figures 12,13. Navicula cancellata Donkin
681 Plate XI
Figure 1.
Navicula complanta Grunow
Figure 2.
Navicula comoides (Agardh) Peragallo
Figure 3.
Navicula digito-radiata (Gregory) Ralfs
Figure 4.
Navicula directa var. remota Grunow
Figures 5, 6.
Navicula diver sistriata Hustedt
Figure 7.
Navicula finmarchica Cleve
Figure 8.
Navicula forcipata var. densestriata A. Schmidt
Figure 9.
Navicula grevillei (Agardh) Heiberg
Figure 10.
Navicula granulata Bailey
Figure 11.
Navicula litoricola Hustedt
Figure 12.
Navicula jamalinenses Cleve
Figure 13.
Navicula humerosa (de Bre'bisson) ex Wm. Smith
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Plate XII
Figure 1.
Navicula lyra Ehrenberg
Figure 2.
Navicula, lyra var. elliptica A. Schmidt
_Figure 3.
Navicula lyra var. subelliptica Cleve
Figure 4.
Navicula normaloides Cholnoky
Figure 5.
Navicula sp. 3
Figure 6.
Navicula palpebralis (de Brebisson) ex Wm. Smith
Figure 7.
Navicula pseudony Hustedt
Figure 8.
Navicula pusilla Wm. Smith'
Figure 9.
Navicula sp. 2
Figure 10.
Navicula sp.
Figure 11.
Navicula rhynchocephala Kutzing
Figure 12.
Navicula peregrina (Ehrenberg) Kiitzing
Figure 13.
Navicula sp. 4
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ie
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Plate XIII
Figure 1.
Nitzschia acuminata Wm. Smith
Figure 2.
Nitzschia angularis Wm. Smith
Figure 3.
Nitzschia apiculata Gregory
Figure 4.
Nitzschia granulata Grunow
Figure 5.
Nitzschia marginulata Grunow
Figure 6.
Nitzschia navicularis (de Brgbisson) Grunow
Figure 7.
Nitzschia spathulata var. hyalina Gregory
Figure 8.
Nitzschia socialis Gregory
Figure 9.
Nitzschia punctata var. coarctata Grunow
Figure 10. Nitzschia punctata (Wm. Smith) Grunow
Figure 11. Nitzschia vivax Wm. Smith
Figure 12. Nitzschia sp.
1
Figure 13. Nitzschia sigma Wm. Smith
ch
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01101111
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03
Plate XIV
Figure 1.
Opephora schwartzii (Grunow) Petit in Pellatin
Figure 2.
Opephora marina (Gregory) Petit
Figure 3.
Pinnularia cruciformis Donkin
Figure 4.
Plagiogramma staurophorum (Gregory) Heiberg
Figure 5.
Plagiogramma brockmanii Hustedt
Figures 6, 9.
Pleurosigma decorum Wm. Smith
Figure 7.
Pleurosigma affine var. normanii Ralfs
Figure 8.
Pleurosigma angulatum var. guadrata. Wm. Smith
7 0 3
Plate XV
Figures 1, 2.
Raphoneis amphiceros (Ehrenberg) Ehrenberg
Figure 3.
Raphoneis sp.
Figure 4.
R'hopalodia musculus Katzing
Figures 5, 6.
Rhoicosphenia curvata (Kazing) Grunow
Figure 7.
Stauroneis salina Wm. Smith
Figure 8.
Skeltonema costatum (Greville) Cleve
Figure 9.
Rhabdonema arcuatum (Lyngbye) Katzing
1
,1$
1,141111111111."0116.
111411NORItlif
L.
Plate XVI
Figure 1..
Stephanopyxis nipponica Gran and Yendo
Figure 2.
Synedra fasiculata (Agardh) Ku"tzing
Figure 3.
Synedra fasciculata var. truncata (Greville) Patrick
Figure 4.
Surirella fastuosa var. cuneata Witt
Figure 5.
Tabellaria flocculosa (Roth) Katzing
Figure 6.
Surirella gemma (Ehrenberg) Kiitzing
Figure 7.
Surirella ovata Kiitzing
Figures 8. 9.
Terpsinoe americana Bailey
Plate XVII
Figures 1, 2.
Trachyneis aspera var. aspera (Ehrenberg) Cleve
Figure 3.
Trigonium arcticum (Brightwell) Cleve (800x)
Figure 4.
Thalassiosira aestivalis Gran and Angst
Figure 5.
Trachysphenia australis Petit
Figure 6.
Tli 'alas sionema nitzschioides Grunow
Figure 7.
Tropodeneis lepidoptera var. minor Cleve
Figures 8,9.
Tropodeneis vitria (Wm. Smith) Cleve
0
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