GRADIENT ANALYSIS OF PHYTOPLANKTON
PRODUCTIVITY AND CHEMICAL PARAMETERS IN
POLLUTED AND OTHER NEARSHORE HABITATS

John Victor Rowney

Library

Naval Postgraduate School

Monterey, California 93940

NAVAL POSTGRADUATE SCHOOL

Monterey, California

THESIS

GRADIENT ANALYSIS OF PHYTOPLANKTON PRODUCTIVITY
AND CHEMICAL PARAMETERS IN POLLUTED AND OTHER

NEARSHORE HABITATS
by

John Victor Rowney

Thesis Advisor:

E. D. Traeanza

March 1973

kppKjovo.d &OA. pubtic A.e£cose; &li> tiibixtlcn ujitunLtad.

T153557

Gradient Analysis of Phytoplankton Productivity

and
Chemical Parameters in Polluted and Other Nearshore Habitats

by

John Victor Rowney
Lieutenant, United States Navy
B.S., United States Naval Academy, 1967

Submitted in partial fulfillment of the
requirements for the degree of

MASTER OF SCIENCE IN OCEANOGRAPHY

from the

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c,/

Libr

Naval Po
Mont.

ABSTRACT

Measurements of primary productivity, chlorophyll standing crop, and
nutrient concentrations were made along a gradient of five nearshore hab-
itats, a seaward transect, and a longshore transect to determine environ-
mental relationships. The effects of municipal sewer outfalls, type of
shoreline, and degree of exposure to high winter seas were found to be
dramatic. The behavior of nutrient ratios suggest their use as pollution
tracers in certain circumstances. The ratios of productivity to chloro-
phyll demonstrated physiological regimes among the phytoplankton in the
sampling area. Comparison of data with carbon monoxide and methane con-
centrations provided a possible correlation between phytoplankton pro-
ductivity and carbon monoxide production.

TABLE OF CONTENTS

LIST OF TABLES 5

LIST OF FIGURES 6

I. INTRODUCTION 8

II. MATERIALS AND METHODS 10

A. STATIONS 10

1. Nearshore Stations 10

2. Seaward and Longshore Transects 12

3. Diurnal and Depth Surveys; Deep Ocean Reference
Stations 14

B. SAMPLING TECHNIQUES 14

C. TOOLS USED IN GRADIENT ANALYSIS 16

1. Productivity Measurements 16

2. Nutrient Measurements 20

3. Chlorophyll and Acid Factor Measurements 20

4. Nitrogen/ Phosphorous Calculation 22

5. Productivity/ Chlorophyll Calculation 22

6. Carbon Monoxide, Methane, and Productivity
Comparison 22

III. RESULTS 23

A. DEPTH AND DIURNAL STUDIES 23

B. GRADIENT ANALYSIS 27

1. Productivity 27

2. Chlorophyll 27

3. Nutrients 33

4. Acid Factor 33

5. Nitrogen/ Phosphorous Ratios 33

6. Productivity/Chlorophyll Ratios 38

7. Carbon Monoxide and Methane 38

IV. DISCUSSION 42

A. PRODUCTIVITY, CHLOROPHYLL, AND NUTRIENTS -- 42

1. Nearshore Gradient: Del Monte to Point Joe 42

2. Seaward Transect: From Del Monte and Point

Cabrillo 43

3. Longshore Transect: Del Monte Beach 44

B. ACID FACTOR - 45

C. NITROGEN/ PHOSPHOROUS RATIOS «% 45

D. PRODUCTIVITY/ CHLOROPHYLL RATIOS 46

E. CARBON MONOXIDE, METHANE, AND PRODUCTIVITY RELATIONSHIPS 47

F. ENVIRONMENTAL EFFECTS 47

V. SUMMARY 50

APPENDIX A - CRUISE DATA 51

REFERENCES -- 61

INITIAL DISTRIBUTION LIST 63

FORM DD 1473 64

LIST OF TABLES

Table I. Productivity, Chlorophyll, and Nutrient Data on the

Nearshore Gradient 34

LIST OF FIGURES

Figure Page No,

1. Chart of the Monterey Peninsula showing nearshore stations-- 11

2. Environmental conditions during the sampling period 13

3. Del Monte Beach kelp bed 15

4. Incubation box (after Doty, 1959) 17

5. Diurnal Productivity survey at Monterey harbor 24

6. Productivity and chlorophyll relations at the deep water
station 25

7. Nutrient concentrations at the deep water station 26

8. Average productivity, chlorophyll, and nutrient data on the
nearshore gradient 28

9. Average productivity and chlorophyll relations on the sea-
ward transect 29

10. Average nutrient concentrations on the seaward transect 30

11. Productivity, chlorophyll, and nutrient data on the first
longshore transect 31

12. Productivity, chlorophyll, and nutrient data on the* second
longshore transect 32

13. Nitrogen/phosphorous ratios on the nearshore gradient 35

14. Productivity/chlorophyll (Pc) ratios on the nearshore
gradient 36

15. Productivity versus chlorophyll at Del Monte and Point
Cabrillo 37

16. Carbon monoxide versus methane at deep station B 39

17. Carbon monoxide and methane on the nearshore gradient 40

18. Carbon monoxide and methane on the longshore transect 41

19. Time study of productivity at the nearshore stations

during the sampling period 49

ACKNOWLEDGEMENTS

I wish to thank my advisor, Professor Eugene D. Traganza, Department
of Oceanography, for his assistance and advice during the preparation of
this thesis. His careful analysis, critical evaluation, and support
were much appreciated. I also thank Dr. Malvern Gilmartin and Mr. Tom
Bowhay of Hopkins Marine Station for their help in the productivity method-
ology and for use of their geiger counter. Appreciation goes to Professor
Stevens P. Tucker, Department of Oceanography, and to Mr. Kenneth Graham,
Department of Research Administration, for their assistance in the chloro-
phyll determinations. I am also indebted to LCDR Gaylord Paulson who
took the time to show me the intricacies of the Auto analyzer despite a
deadline to meet on his own thesis.

A special thanks must be expressed to the NPS boat crew, of the
First Lieutenant's Division. Their tireless help and cheerful support,
despite adverse weather conditions in potentially hazardous waters, was
instrumental in the completion of the sampling phase.

Finally, I would like to thank Professor Eugene C. Haderlie, Depart-
ment of Oceanography, for his critical and constructive evaluation of the
final manuscript.

I. INTRODUCTION

This study was designed to test the hypothesis that measureable rela-
tionships exist between primary productivity, standing crop, and the chem-
ical environment (nutrients , carbon monoxide, and methane) of sublittoral
phytoplankton communities. The data is presented as a gradient analysis
of a variety of nearshore habitats characterized by large stands of macro-
algae including Macrocystis and Nereocyst is . The gradient analysis is a
study of a series of biological and chemical parameters which vary along
an environmental gradient to determine which factors, if any, are limiting,
The study was carried out in late fall to midwinter, a poorly documented
season in the literature, but during which numerous and noticeable oceanic
and atmospheric effects on the phytoplankton community took place.

Part of the impetus for this thesis came from a desire (from the
Naval Research Laboratory) to examine relationships, if any, between pri-
mary productivity and carbon monoxide and methane concentrations in an
oceanic environment. Therefore, sampling was carried out simultaneously
with James T. Welch and the data analysed together (see Welch, 1973, and
this report) .

Although much work has been done in recent years measuring and sur-
veying the primary productivity of the world's oceans, most studies have
been confined to relatively deep water, especially in coastal areas, and
accomplished in the spring and late summer, the times of upwelling and
phytoplankton blooms. Little work has been done less than 1000 meters
from shore in the sublittoral environment. In Monterey Bay, Cowles (1972)

beyond the intertidal, but with waves and turbulence still in-
fluencing the entire vertical range of depth.

8

has studied the sublittoral nearshore, but only in a transect from one
nearshore location to deep water in the spring season.

II. MATERIALS AND METHODS

A. STATIONS

Sea water samples for the measurement of primary productivity, chloro-
phyll, and nutrients were taken at five nearshore stations representing
a variety of habitats (Figure 1). Data were taken during a ten week

period from 10 October to 15 December, 1972, in Monterey Bay, California.

2
The study began in the local "Oceanic Period", characterized by calm

seas and warm sea surface temperature. A sharp drop in temperature and

an influx of clear, oceanic surface water (Figure 2) marked the onset of

3
the "Davidson Current Period". Heavy rainfall and high seas accompanied

the passage of storms during the fifth, sixth, and ninth weeks.

Five nearshore stations represent a gradient from a protected (bay)
environment to an exposed (oceanic coast) environment. A seaward transect
from two of these stations illustrate a gradient to open water and the
effect of kelp beds on productivity. A longshore transect was designed
to examine a sewer outfall and the gradient of productivity in a large
kelp bed. Local depth and diurnal surveys were made in order to obtain
maximum results when sampling. A station in deep water was made in order
to provide an oceanic deep water index for this season.

1. Nearshore Stations

Station 1 was the center of an extensive kelp bed at Del Monte

2

a period of variable winds and calm seas when cool, saline surface

water (previously upwelled) sinks and is replaced by warm oceanic water

(Welch, 1967).

3

a period when Davidson Current develops, flowing north along the

California coast; net transport is on-shore due to the Coriolis effect
(Welch, 1967)

10

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Figure 1. Chart of the Monterey Peninsula showing nearshore stations

11

beach, adjacent to the Postgraduate School beach laboratory and about
200 yards offshore. Water depth was about eight meters over a sandy
bottom. In general, the area was relatively calm, less affected by the
stormy seas further along the coast. A sewer outfall which was pumping
effluent directly into the kelp bed was located near the eastern edge.
Station 2 was located in a kelp bed off Point Cabrillo, adjacent to the
Hopkins Marine Station. Here the bottom and shore was rocky. Depth was
about 10 meters 100 yards offshore. The area was subjected to some heavy
swell from the north and northwest during the study. Station 3 was lo-
cated in the kelp bed between Point Pinos and Lucas Point (denoted "Point
Pinos North" hereafter). Samples were taken from the approximate center
of the kelp bed about 300 yards from the rocky shoreline and in about
14 meters of water. This area received very heavy swell most of the
time, as did the following two stations. Station 4 was a kelp bed just
south of the main rock outcropping of Point Pinos. This station (denoted
"Point Pinos South" hereafter) was about 300 yards offshore in about 15
meters of water and was the first "oceanic influence" station. It had a
sewer outfall in the rocks about 200 yards from the kelp bed. Point Joe,
Station 5, was the furthest from the influence from the bay. Sampling
took place in the kelp bed center which was in about 15 meters of water
and 300 yards offshore. Besides receiving heavy swell, the turbulence
of the area was increased by large eddies formed between the kelp bed
and the shore.

2. Seaward and Longshore Transects

Transects were made from the kelp beds at Del Monte beach and
Point Cabrillo to the bell buoy adjacent to Point Cabrillo. The buoy is
in open water of about 40 meters depth. In each transect, samples were
taken in the center of the kelp, and at equidistant intervals from the

12

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Figure 2. Environmental conditions during the sampling period
Periods of rain and Davidson current intrusion are noted.

13

edge of the kelp to the buoy. Another transect was made across the ex-
panse of the Del Monte beach kelp bed (Figure 3). Six locations approx-
imately 400 yards apart were selected for sampling: at the east edge, at
the sewer outfall, adjacent to the Water Pollution Facility pumphouse,
adjacent to the NPS beach laboratory, adjacent to La Playa apartments,
and at the west (harbor) edge.

3. Diurnal and Depth Surveys; Deep Ocean Reference Stations

A diurnal time series survey was taken at Monterey harbor wharf
#2, near the west end of the Del Monte beach kelp bed. Samples were taken
every two hours for 24 hours. Two depth surveys each were taken at Del
Monte kelp bed and Point Cabrillo kelp bed. Two deep stations were taken
in the Monterey canyon at 36°44'N, 122°7'W and 36°44'N, 122°5'W, aboard
R/V Acania.

B. SAMPLING TECHNIQUES

A 40 foot boat was used for a total of 22 cruises to the inshore
stations. Although analyses could not be performed aboard, the boat
size made it easier to get close to shore. Heavy seas during the fifth,
sixth, and ninth weels did make it impossible to collect samples at Point
Pinos and Point Joe. Water samples were collected from each station in
a two liter, opaque, PVC Van Dorn bottle with hand lines for lowering and
tripping. Subsamples were then dispensed into three 125 ml Pyrex bottles
for productivity, one 500 ml Pyrex bottle for chlorophyll, and one five
ml plastic Technicon Autoanalyzer cup for nutrient analysis. All bottles
were put in a dark plastic bag of seawater for the remainder of the
cruise and transportation to the lab. Sea surface temperatures and a
Secchi disc reading were taken at most stations. All samples were taken
between 0800 and 1000 local time and within two meters of the surface.

14

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C. TOOLS USED IN THE GRADIENT ANALYSIS
1. Productivity Measurements

Food-chain ecology of the sea has attracted a great deal of
interest in recent years. Researchers have been striving to accurately
measure primary productivity, the basis of life in the sea. They have
tried to relate phytoplankton standing crop or biomass , nutrient concen-
trations, and chlorophyll concentrations to primary productivity but with
the possible exception of nutrients, have not had a great deal of success
due to interfering processes (Ryther, 1956). Initially, primary produc-
' tivity was measured by the relative production of oxygen in light versus
dark bottles, but incubation periods are long and the ratio of C09 (assim-
ilated) to 0„ (produced) while close to unity is variable (Ryther, 1956).
On the other hand, the uptake of C09 is equivalent, mole for mole, to
carbon production and represents the most direct approach to primary pro-
ductivity measurements. The method of radioactive carbon fixation was

first described by Steemann-Nielsen (1952). A known activity of radio-

14
active carbon (C ) is added to a phytoplankton culture and incubated at

14
appropriate light levels. The radioactivity of the C assimilated by the

14
plants is measured on a geiger counter. Steemann-Nielsen claimed the C

method measured gross production (total carbon produced) while later re-
searchers claimed the method to yield net production (carbon produced
less carbon used in respiration) (Ryther, 1956). The general consensus is
to regard the results as somewhere between gross and net production,
longer term experiments favoring a net value (Riley and Skirrow, 1965).
Artificial light may be used instead of in situ or natural illumination
to determine "relative productivity" (Jitts, 1961).

Other modifications to Steemann-Nielsen's (1952) carbon-14
method for measuring primary productivity have been used. The method

16

Figure 4. Incubation bo ifter Doty, 1959)

coolant enters the box Erom tubing above.

Sea water

17

employed here was based on that of Stickland and Parsons (1968) and used
a fluorescent light incubator (Doty and Oguri, 1959) as shown in Figure

4. Two light and one dark bottle from each station were inoculated with

14
one ml of NaC 0„ from glass ampoules containing about five microcuries

of radioactivity,, The bottles were incubated under fluorescent light

("cool white" bulbs, about 0.06 langley per minute) for three to four

hours. Taylor and Hughes (1967) have shown that carbon uptake is linear

over the first two to eight hours. Use of a short incubation period

14
also eliminated the loss of C by respiratory oxidation (Harvey, 1955)

and "bottle effects" such as bacterial growth on walls. The bottles
were cooled in the incubator by filtered sea water which was pumped from
the sea just off of Del Monte beach. The incubator cooling was always
within +2 C of sea surface temperature. In the depth analyses, neutral
density filters were used to reduce incubator light to approximately
that which was present _in situ (McAllister and Strickland, 1961).
After incubation the bottles were filtered through 0.45 micrometer Milli-
pore HA-type filters under 0.5 atmosphere vacuum. Bottles of 125 ml
capacity were used due to high filtering times encountered with 250 ml
bottles of the particulate laden nearshore water. The method is inde-
pendent of volume filtered. Filters holding the radioactive phytoplank-
ton were rinsed with a few ml of filtered sea water, sucked dry, and mounted
on copper planchettes with rubber cement. The above procedures were car-
ried out aboard ship for the deep water stations only. The filters
were dried in a dessicator for 24 hours and transferred to airtight jars
containing silica gel and soda lime to insure dry, carbon dioxide-free
storage. The radioactivity of each sample was counted within one week
of collection. Samples can be stored for several weeks in this manner
without physical loss of radioactivity (Strickland and Parsons, 1968).

18

The mounted filters were counted on a Nuclear Chicago Scalar
geiger counter (model 161A) equipped with a model D47 gas flow chamber
and a "micromill" window. Light bottle counts were held to 5000 counts
or three minutes, whichever came first. Dark bottles were counted two to

three minutes. It was also necessary to determine the total amount of

14
activity in the C ampoules. The first batch used was pre-calibrated

at the factory. The second batch was calibrated on a Nuclear Chicago

liquid scintillation counter in the method of Jitts and Scott (1961).

This method has proven to be much more accurate than the barium carbonate

precipitation method described in Strickland and Parsons (1968). Errors

inherent in extrapolating to "zero" thickness are eliminated. Primary

productivity is then given by the formula:

mg C m hr

(Rg-Rg) x W x 1.05

R x N

where: R = light bottle count in counts per minute (CFM),

R^ = dark bottle count in CPM,

R = total absolute activity of an ampoule in CPM,

W = weight of carbonate carbon in the water
in mg C/m ,

N = number of hours incubated, and

1.05 = an isotope discrimination factor.

3
In this study W was assumed to be constant at 24000 mg C/m . Periodic

checks in various locations showed this to be valid within 1%. The dis-

14

crimination factor accounts for the difference in behavior of the C

12
isotope from the C isotope found in nature (Strickland and Parsons,

1968). In the calculations, dark bottle counts were subtracted from

light bottle counts. The dark bottle acts as a control representing a

combination of non-photosynthetic fixation of carbon and absorption of

14
C by particulate matter and detritus (Taylor and Hughes, 1967).

19

2. Nutrient Measurements

In sea areas where nutrient cycles and phytoplankton have been
studied, it is clear that certain minor inorganic constituents of sea water
are required for growth and reproduction. Due to the low concentrations
of these nutrients in sea water, phytoplankton production may become
limited by their depletion (Raymont, 1963). Study of the nutrient environ-
ment, therefore, may give significant insight to variations in primary
productivity and the "fertility" of the seas.

Upon reaching the laboratory after collection, the five ml
nutrient samples were frozen in a dark freezer until enough samples
were collected to warrant automated analysis (Steven, Brooks and Moore,
1970, and Corcoran and Alexander, 1963). Samples were analyzed on a
Technicon Autoanalyzer II with concentrations reported in microgram-atoms
per liter ((i.g-at/1) of nitrate (N0„ ), nitrite (N0„ ), phosphate (princi-
pally HPO ~), and silicate (H.SiO,- H + H SiO ~). Overall, the sensi-
tivity, reproducibility, and accuracy of this system for sea water
nutrient analysis have been found to be very satisfactory. The last batch
of phosphate samples run (representing 9 November - 15 December collections)
are somewhat suspect due to erratic behavior of the phototube.

3. Chlorophyll and Acid Factor Measurements

A 500 ml water sample was used to make an estimate of standing
crop of phytoplankton by measurement of chlorophyll a. Grazing pressure
was estimated by calculation of the chlorophyll a to phaeophytin a fluor-
escence ratio. Using chlorophyll a as an index of standing crop is valid
as long as the species composition remains relatively stable .between
stations and over the time period of collection (Cowles, 1972). Chloro-
phyll degradation products, such as phaeophytin, are often found in areas
.of zooplankton activity. These compounds fluoresce strongly at the same

20

wavelength of maximum fluorescence of the chlorophylls. Undegraded
chlorophyll a can be determined by measuring the decrease in fluorescence
(occuring at 665 run) which takes place when the pigment extract is treated
with dilute acid to convert it to phaeoph>tin a. The ratio of the unacid-
ified fluorescence to the acidified fluorescence, or the "acid factor",
is a measure of zooplankton grazing pressure under ideal conditions. An
acid factor of close to one, indicative of a very high phaeopigment con-
centration, has been found below the photic zone and in the vicinity of
large populations of zooplankton. Ratios above 1.7 indicate high concen-
trations of undegraded chlorophyll. The exact value will depend on the
relative amounts of chlorophyll a, b, and £ present (Holm-Hansen, et. al.,
1965, Yentsch and Menzel, 1963).

In this study, the experimental method of Strickland and Parsons
(1968) was used to determine chlorophyll a and acid factor. Each 500 ml
water sample was filtered in the laboratory within an hour of collection
through a Whatman GF/C glass fiber filter under 1/3 atmosphere vacuum.
Samples from the deep water station were processed aboard ship. One ml
of magnesium carbonate suspension was added to the final small volume
of each sample passing through the filter. The filter containing the
sample was folded in half and frozen in the dark until fluorometric de-
termination. A time study, in which three replicate samples were stored
for five days, three weeks, and two months respectively, showed no de-
gradation of pigment with this procedure. The filters were removed from
the freezer, ground in a tissue grinder, and centrifuged. The fluorescence
of the supernatant liquid extract was measured on a Turner Model 111
fluorometer. Two drops of dilute HCl were then added, and, after five
minutes, the fluorescence was measured again. The acid factor was then
calculated .

21

The fluorometer was calibrated by measuring the chlorophyll
extinction of one sample on a Beckman spectrophotometer and calculating
concentration with the equation of Strickland and Parsons (1968):

_,. h „ , 3 U-6E6650 - L-31E6450 ' °-14E6300

mg Chlorophyll a/m =

V
where: E = extinction value at the specified wavelength
V = volume of sea water filtered in liters.

4. Nitrogen/Phosphorous Calculation

From the nutrient data, nitrate to phosphate ratios (N:P), or
phytoplankton "assimilation ratios", were calculated to estimate chemical
interrelationships in the sea water (Riley and Skirrow, 1965).

5. Productivity /Chlorophyll Calculation

Productivity to chlorophyll (Pc) ratios were calculated as an
index of physiological or population changes within the groups of phyto-
plankton and habitats studied (Riley and Skirrow, 1965).

6. Carbon monoxide/ Methane/ Productivity Comparison

Carbon monoxide and methane concentrations were compared to
productivity data to determine if any interrelationships exist. See
Welch (1973) for methods of measurement.

22

III. RESULTS

A. DEPTH AND DIURNAL STUDIES

Maximum daily production rates were desired for this study. It was
assumed that the bulk of the phytoplankton population would be in the up-
per two meters of water owing to the reduction of natural light in the
winter and the turbidity of inshore waters (Taylor and Hughes, 1967).
According to Riley and Skirrow (1965), photosynthesis in phytoplankton
occurs at its maximum rate in the late morning hours. Separate experi-
ments confirmed these assumptions. Depth studies taken on different days
at Del Monte beach and Point Cabrillo confirmed maximum production to
take place from one-half to two meters in depth (Appendix A, Cruises 4,
13, and 18). A twenty- four hour study (Figure 5) showed the greatest in-
crease in production to occur in late morning with a maximum at 1330 local
time. A similar phenomenon has been noted by other researchers in other
waters (Doty and Oguri, 1957, McAllister, 1963, and Lorenzen, 1963).

At the deep water stations in the Monterey submarine canyon (Figure
6), productivity and standing crop at the surface was comparable to the
values obtained at Point Pinos and Point Joe. The values decreased rapid-
ly with depth to a low at 46 meters, the depth at which only 1% of the
surface light was available as determined by Secchi disc. As expected
the acid factor indicated heavily degraded chlorophyll at this level.
Figure 7 shows "reactive" nitrate and silicate both increasing to deep
water maximums while high values of "reactive" phosphate and nitrite
remain in the surface layer. Acid factors were roughly on the same order
as the inshore stations.

23

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at 36° 44* N, 122° 05' W.

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26

B. GRADIENT ANALYSIS

1 . Productivity

The gradient of productivity between the five nearshore stations
for each week data was obtained, is presented in Table I. In addition,
Figure 8 shows average values of all productivity measurements taken
during the study. Productivity generally decreased toward the most ex-
posed station, i.e., from Del Monte to Point Joe. The seaward transect
shown in Figure 9, represents an average of two runs from Del Monte to
the buoy and two runs from Point Cabrillo to the buoy. Runs were made on
different days. Productivity increased from Point Cabrillo to the buoy,
but decreased from Del Monte to the buoy. In each case, however, pro-
ductivity was lower inside the kelp bed (mostly Macrocystis) than at the
edge just outside it. The longshore transects across the width of the
Del Monte kelp bed are presented in Figures 11 and 12. The first run
showed maximum productivity from the beach laboratory to the harbor. On
this day, the "boil" from the sewer outfall could not be found. The
second run also showed high productivity from the laboratory to the harbor,
The "boil" was sampled and found to be low in productivity, but increase
was rapid towards the west.

2. Chlorophyll

Chlorophyll, like productivity, generally decreased from Del
Monte to Point Joe along the nearshore gradient as shown in Figure 8 and
Table I. Little change was noticed in the seaward transect from either
Del Monte or Point Cabrillo to the buoy (Figure 9). Figure 11 shows
chlorophyll to be variable in the first run of the longshore transect;
when the "boil" was sampled (Figure 12), a very high chlorophyll standing
crop was evident.

27

? 2

n.

O

i

3.

1.5

L.

o

U

o

u 1.0

0

CO

E

2

X

E

U

U

1

o»

Ol

E

E

x H3Sio;

• NO"2

xhpo;

• pp

*Chl

Figure 8. Average productivity (PP) , chlorophyll (Chi) and nutrient
data on the nearshore gradient. Data plotted from Station 1 (Del
Monte) to Station 5 (Point Joe).

28

L.

o

u

o

1.5

S 1.0

CO

u

E 0

• Del Monfe-buoy

* Pt Cabrillo- buoy

U

w
E

kel

edg<

buoy

Figure 9. Average productivity & chlorophyll relations on the seaward
transect. Samples were taken at the respective kelp bed center, kelp
bed edge, and from there at equidistant intervals to the buoy.

29

10

H3SiO;

no:

O

I

no;

0

hpo;

• Del Monte -* buoy
X Pf Cabriilo ■* buoy

Figure 10. Average nutrient concentrations of the seaward transect.
Samples were taken at the respective kelp bed center, kelp bed edge,
and from there at equidistant intervals to the buoy.

30

0

I

O 1
i

en

3.

o 15

u

0

o

O

10

u u

en en
E E

• H3Si04
x HPOi

"8DT

•NO
*NO

8 12

yards x10~2

16

20

Figure 11. Productivity, chlorophyll, and nutrient data on the first
longshore transect. Zero is the eastern edge of the kelp bed at Del
Monte beach.

31

•H3SiO;
* hpo;

•^ 2
o
i
en

a o

u
O

U

D

'o

o

1 -

•41.6

•NO
*NO

Figure 12. Productivity, chlorophyll, and nutrient data on the sec-
ond longshore transect. Zero is at the eastern edge of the kelp bed
at Del Monte beach. 400 is the sewer outfall.

32

3. Nutrients

Nutrient concentrations for each week in the nearshore gradient
are presented in Table I. The average value plot (Figure 8) shows phos-
phates decreasing toward Point Joe, nitrates increasing toward Point Joe,
and silicate highs at both ends of the gradient. The seaward transect
(Figure 10) shows most nutrients decreasing towards open water at the
buoy. Concentrations generally increased slightly at the edge of the
kelp over those in the kelp bed center in both areas. The day the "boil"
at the sewer outfall could not be found in the longshore transect (Figure
11), high silicate and phosphate concentrations were noted west of its
supposed location. When the boil was sampled, very high concentrations
of silicate and phosphates were found (Figure 12); these decreased with
distance from the outfall. Nitrite, however, increased with distance from
the outfall.

4. Acid Factor

In the nearshore gradient (Figure 8 and Table I), acid factor
was not very definitive but generally was highest at Station 2 off Point
Cabrillo. Acid factors in the seaward transect generally increased once
out of the respective kelp beds (Figure 9). In the first run of the
longshore transect (Figure 11), acid factors indicated higher grazing
pressures towards the west (harbor) end. The second run showed a very
high acid factor at the sewer boil, with diminishing values again towards
the harbor (Figure 12).

5. Nitrogen/Phosphorous Ratios

In Figure 13, nitrate to phosphate ratios are plotted on the

nearshore station gradient for four of five weeks. In these diagrams,

the N:P ratio increases from Del Monte, Station 1, to Point Joe, Station

5. In the seventh week analysis, undetectable phosphate concentrations

were suspect and therefore not considered.

33

TABLE I

Productivity, chlorophyll and nutrient data on the nearshore grad-

-3 -1 3

ient. Productivity given in mg C in hr ; chlorophyll given in mg/m ; and

nutrients given in ug-at/1. Station 1 is Del Monte; Station 2 is Point

Cabrillo; Station 3 is Point Pinos North; Station 4 is Point Pinos South;

Station 5 is Point Joe. Symbols are: PP(productivity) , CHL(chlorophyll) ,

AF(acid factor), N3(nitrate), S(silicate), P(phosphate) , N2(nitrite).

Week

Station

PP

CHL

AF

N3

S

P

N2

3

1

11.4

.85

5.05

1.64

.28

2

3.08

0.0

2.75

0

.20

3

1.79

.38

2.60

.54

.39

5

.37

4.72

2.55

.32

.42

4

1

6.80

3.39

1.4

0

8.15

.86

.14

2

5.12

.72

1.49

0

2.15

.17

.22

3

3.05

.15

3.85

.55

.28

4

2.08

2.90

4.40

.65

.46

5

.37

8.42

7.00

.48

.27

7

1

1.89

.65

1.17

3.08

8.05

0

.20

2

.99

.56

1.46

3.48

4.95

0

.28

3

.95

.42

1.17

4.08

4.35

0

.27

4

.67

.32

1.13

4.78

3.90

.10

.25

5

. .12

.48

1.21

5.10

4.05

.22

.37

8

1

1.36

.56

1.15

6.31

12.4

1.37

.61

2

2.83

.58

1.20

5.22

7.80

1.08

.76

3

1.59

.51

1.23

7.10

7.45

.83

.72

4

1.02

.36

1.20

8.52

8.00

.80

.74

5

1.07

.39

1.20

7.47

7.50

.50

.59

10

1

3.02

5.75

9.55

1.19

1.21

2

1.70

5.80

7.40

.40

1.35

3

1.31

6.25

6.90

1.08

1.37

4

1.30

5.72

6.70

.07

.87

5

1.15

5.18

5.70

0

.75

34

Z

80

25|-
20
15
10
5

week 10

• oo

a.

18-
16 -
14
12
10
8

• week 3
xweek 4
oweek 8

2 3
stations

Figure 13. Nitrogen/phosphorous ratios on the nearshore gradient,
Data plotted from Station 1 (Del Monte) to Station 5 (Point Joe).

35

1 -

2 3
stations

Figure 14. Productivity/chlorophyll (Pc) ratios on the nearshore
gradient. Data plotted from Station 1 (Del Monte) to Station 5
(Point Joe) .

36

»- 4

U

E

Del Monte

mgChl/m3

U
E

Pf Cabrillo

.25

m

.5
gChl/i

.75

m

1.0

Figure 15. Productivity versus chlorophyll at Del Monte and Point
Cabrillo.

37

6. Productivity/Chlorophyll Ratios

In Figure 14, average productivity to chlorophyll (Pc) ratios are
plotted along the nearshore gradient for the two weeks that data was ob-
tained for all areas (weeks seven and eight). Del Monte and Point Cabrillo
(1 and 2) seem to represent similar conditions, as do the stations at
Point Pinos (3 and 4). Point Joe (5) is by itself with a very low Pc
ratio. When productivity was plotted against chlorophyll for Del Monte
and Point Cabrillo (the two stations with the most chlorophyll data),
definite but different relationships were obtained (Figure 15).

7 . Carbon Monoxide and Methane

Carbon monoxide and methane data are from Welch (1973). Carbon
monoxide, methane and depth relations are plotted in Figure 16. This
figure represents conditions at one of the deep water stations studied.
Both CO and CH, increased with depth to 15 meters, then CO abruptly de-
creased with depth to 100 meters with methane still increasing slightly.
Both decreased from there to 1000 meters. In the nearshore gradient
(Figure 17), average values of methane behaved very much like silicate
(Figure 8) with a source at Del Monte and Point Joe. Carbon monoxide
fluctuated but generally increased toward Point Joe. The sewer outfall
showed strikingly high values for both CO and CH, in the longshore tran-
sect (second run) as shown in Figure 18. Concentrations decreased with
distance from the boil, but both began a rise toward the harbor.

38

o

2

IE

1000

0 u
0

• depth in meters

100

\

\

200 <k

/

/

/ 0 o

700 ^

75
o

50

30

15

~X5>

CO (ML/L x 104)

Figure 16. Carbon monoxide versus methane at deep station B,

39

o CO

5r

4

O
v-

>:

Siafi

ons

Figure 17. Carbon monoxide and methane on the nearshore gradient
Data plotted from Station 1 (Del Monte) to Station 5 (Point Joe).

40

x CH

A

6

o

4

V

it

_1_

X

0

.4

.8 1.2

Kiloyards

1.6

2.0

Figure 18. Carbon monoxide and methane on the second longshore
transect. Zero is the eastern edge of the Del Monte beach kelp
bed . v

41

IV. DISCUSSION

A. PRODUCTIVITY, CHLOROPHYLL, AND NUTRIENTS

1. Nearshore Gradient: Del Monte to Point Joe

The high productivity and chlorophyll standing crop encountered
at Del Monte and the decrease seaward was due to a combination of factors.
Dunstan and Mengal (1971) have shown that "seawater diluted with secondary-
treated sewage effluent provides excellent enrichment" for the maintenance
of natural phytoplankton communities. At Del Monte Beach, municipal sew-
age effluent was present and this, combined with a relatively protected
and stable environment, allowed the phytoplankton to proliferate at rates
uncommon to the season. Out of the influence of the effluent and into
more turbulent waters, productivity and standing crop fell gradually to
the outermost station as shown in Figure 8 and Table I. Deep station pro-
ductivity was on the same order as that of Point Joe.

The high average inorganic nutrient values indicate that Del
Monte beach is a phosphate and silicate source (see section 3, below).
Nitrates and silicates are replenished at Point Pinos and Point Joe by
turbulent upwelling of bottom water in the shallow nearshore. The low
values of nitrate and silicate found at the surface of the deep station
will be replenished in an analogous manner with the spring upwelling of
water. High concentrations of nitrite, sometimes a product of phytoplank-
ton as well as from bacterial reduction of nitrate and nitrification of
ammonia, are often found in highly productive areas (Raymont, 1963, and
Riley and Skirrow, 1965). In Figure 8, the highest values a nitrite were
found inshore, at Del Monte and Point Cabrillo, the areas of highest pro-
duction.

42

In contrast to the effect of the sewer outfall at Del Monte, the
outfall at Point Pinos had little success stimulating productivity. In
fact, it was noticed that the Del Monte outfall is surrounded by healthy
kelp while the Point Pinos outfall is devoid of any visible life for about
200 yards. It was surmised that either the Point Pinos effluent was very
toxic or that any nutrients there were dispersed quickly by turbulence. It
was later found that the Point Pinos sewage is chlorinated to excess while
the Del Monte effluent is only lightly chlorinated every three to four
days.

2. Seaward Transect: From Del Monte and Point Cabrillo

As previously discussed, productivity increased from open water
to the Del Monte kelp bed and decreased from open water to the Point Cab-
rillo kelp bed. The latter result is probably more typical of uninfluenced
kelp beds (i.e., without a sewer outfall) and agrees with the findings of
Cowles (1972). In his study of the Point Cabrillo transect, he attributes
the lower level of productivity inshore to cell damage due to physical
contact with the rocky shore, high bacterial activity, and photodegrada-
tion of chlorophyll. The Del Monte phenomenon is apparently due to the
enrichening influence of the sewer outfall in a less turbulent or violent
environment (sandy beach) which more than compensates for other nearshore
hazards to phytoplankton. The fact that the plankton productivity was
higher just outside either kelp bed than inside them may be explained by
the presence of the kelp itself. The huge biomass of benthic algae is
probably in direct competition with the phytoplankton for available nu-
trients. Those phytoplankton in the bed may simply not have as many
nutrients available to them as those just outside. This is borne out to
a certain extent in Figure 10, which shows a decrease in silicates,
nitrates, and, in one case, phosphates, but more data needs to be taken.

43

Furthermore, the mass of benthic algae may have a shading effect in the
bed preventing phytoplankton there from utilizing available light, which
is low in the winter.

3. Longshore Transect: Del Monte Beach

Stevenson (1964) has noted in a current survey of Del Monte
Beach that a southwesterly longshore current may develop when a north or
northeasterly wind is blowing. It is surmised that these conditions were
in effect when the first Del Monte transect was made. Since the outfall
"boil" could not be found, it was probably being pumped under reduced
pressure. The discharge, indicated by diluted, but still high, phosphates
and silicates, was detected southwest of the outfall where the current
had moved it. The next time the transect was sampled, the phosphate
and silicate source at the Del Monte outfall was confirmed and concentra-
tion decreased with distance along the beach. The concentrations of
phosphate and silicate at the outfall boil were approximately ten and
eight times their respective average values at the most oceanic nearshore
station studied (Point Joe). Moreover, the outfall was fifteen and
thirteen times the surface concentrations at the deep station with respect
to phosphate and silicate.

Productivity was very low, near zero, at the outfall which may
have been due to the low salinity or toxic material in the effluent.
Dunstan and Mengal (1971) found in a laboratory experiment that sewage:
seawater mixtures in excess of 20:80 were inhibiting to marine diatoms
with normal seawater salinity requirements. Once out of the immediate
vicinity of the outfall, however, productivity increased greatly. High
production adjacent to the beach laboratory indicated this to be about
the optimum distance from the outfall. Nitrite increased with productiv-
ity hinting again that it might be a growth product. The high phytoplankton

44

standing crop at the outfall contained nearly pure chlorophyll as indi-
cated by the very high acid factor (see below).

B. ACID FACTOR

In the nearshore gradient, the high acid factors indicated the least
grazed chlorophyll at Del Monte and Point Cabrillo. However, the range
of values showed that a large herbivore population might have been present.
As Cowles (1972) points out, low acid factor in the nearshore may also
be the result of processes other than grazing, e.g., bleaching of chloro-
phyll after cell damage in waves or on rocky shores. In fact, in the
nearshore, grazing may be the least important factor in evaluating high
phaeophytin concentrations. The seaward transect indicated that the
lowest acid factors (highest degraded chlorophyll) were indeed the furthest
inshore, at or near the kelp beds. This again, could be due to heavy
grazing and/or physical cell damage. The longshore transect indicated
the Del Monte sewer outfall to be just as inhibiting to zooplankton as
to phytoplankton. Acid factor was very high suggesting that the chloro-
phyll present was not being grazed. Grazing pressure did increase, how-
ever, with the distance from the outfall boil.

C. NITROGEN/ PHOSPHOROUS RATIOS

In most oceanic waters of the world, the ratio of nitrogen to phos-
phorous (also called the phytoplankton "assimilation ratio") has been
reported to be on the order of 15 or 16 to 1 for both oceanic seawater
and uptake by phytoplankton (Riley and Skirrow, 1965). This ratio is
somewhat variable, especially in the coastal areas, and has been reported
from 5:1 to 24:1. The nearshore gradient, from Del Monte to Point Joe,
in four of five weeks showed a remarkably similar trend from low to high
N:P ratio. In three of five weeks, the values at Point Joe, from 14.5:1

45

to 17.5:1 were similar to Riley and Skirrow's figgure. It seems that
in this area, the assimilation ratio could be used as a "pollution tracer"
with respect to the sewer outfall at Del Monte. The deep station reflected
Riley and Skirrow's figure only at a depth of 100-200 meters due to upper
layer depletion of nitrate. It is suspected that during the spring up-
welling the ratios will become more typical of oceanic water.

D. PRODUCTIVITY/ CHLOROPHYLL RATIOS

Pc ratios, that is, the ratios of primary productivity to chlorophyll
concentration, have been shown to be a good index of comparative plant
physiology (e.g., the effect of nutrient deficiencies) and species compo-
sition changes (Riley and Skirrow, 1965). High Pc ratios indicate a
healthy or efficient state in a given population or characterize efficiency
differences in different species. In Figure 14, Del Monte and Point Cab-
rillo seem to be on the same physiological level as do the two Point Pinos
stations with a higher ratio. The much lower Pc ratio at Point Joe
(Station 5) may reflect the low phosphate concentrations or some other
physiological deficiency. There was a different genus of kelp at Point
Joe, Nereocystis, which may indicate some environmental change. On the
other hand, the different levels may be due to changes of the species
composition of phytoplankton populations living on bay coast as opposed
to oceanic coast. The deep water stations gave Pc values of 2.9 and 2.8
which indicate conditions or populations similar to those between Point
Pinos and Point Joe. Figure 15 shows a definite proportionality between
productivity and chlorophyll concentration suggesting most all the chloro-
phyll is contained in living plant cells (Steele and Baird, 1961).

46

E. CARBON MONOXIDE, METHANE, AND PRODUCTIVITY RELATIONSHIPS

Comparisons between carbon monoxide, methane, and productivity at
the deep stations and at the outer nearshore stations, Point Pinos and
Point Joe, are interesting. Comparing Figures 6 and 16, it is evident
that there was a CO source in the upper 15 meters were productivity was
productivity was highest. Below 15 meters, where productivity decreased,
CO decreased rapidly also. It is theorized that certain bacteria which
utilize CO reduced its concentration in the deeper water (Welch, 1973).
In the nearshore gradient (Figure 17), a rise in CO occured between Point
Pinos and Point Joe. In previous discussion, it was seen that the pro-
ductivity and standing crop between these last two stations were very close
to the deep station values. Moreover, Pc ratios showed like physiological
conditions or populations in the area between Point Pinos and Point Joe
and at the deep sations. Hence, the similar phytoplankton populations
in. the ares between Point Pinos and Point Joe and at the deep stations
both showed a strong correlation with carbon monoxide production. Much
more work needs to be done, however, to show a causal relationship.

Very little, if any, correlation was seen between primary production

and CO and CH, concentrations in the longshore transect (Figures 12 and 18)

4

This is probably due to large perturbations caused by the sewer outfall
which effectively masks any correlations.

F. ENVIRONMENTAL EFFECTS

The period of study was characterized by a series of miniature blooms
which are not uncommon in midwinter in shallow water (Doty, 1961). Small-
scale, short-term vertical turbulence caused by storms periodically add
nutrient-rich bottom water to surface layers in sublittoral areas, but
nutrients are quickly depleted in the new mixed layer and are not renewed

47

until the next storm. In addition, rain and land runoff may lower salin-
ities, prohibiting phytoplankton from frequenting the uppermost layers
where they can best utilize the limited light available.

Figure 19 shows a time study of productivity for the five nearshore
stations. Primary productivity was high during early weeks but was cut
down drastically during the first heavy rains (Figure 2). Periods follow-
ing the rain during" the fifth, sixth, and ninth weeks showed productivity
minimums. The reduction of light and the lowering of temperature with the
onset of winter also limited the maximum value of the observed "mini-
blooms" (Raymont, 1963). Nutrient renewal by turbulence was very evident
at the outer nearshore stations (Point Pinos and Point Joe), but there
also could have been nutrients added by land runoff. In fact, this factor
could have varied from station to station.

48

r io

Figure 19. Time study of the productivity at the nearshore stations
during the sampling period.

49

V. SUMMARY

As a result of this study, the following conclusions were made.

1. The value of the "gradient analysis" in revealing relations
between biological and chemical parameters is seen, and it is recognized
as a useful tool for monitoring and prognosticating the state of the
environment. In particular:

a. Primary productivity and standing crop in the sublittoral
zone decrease significantly from Del Monte beach around the coast to Point
Joe. A nutrient-rich and non-toxic sewer outfall at Del Monte beach and
the gradient from a calm, sandy, bay environment to an exposed, rocky,
oceanic coast are responsible for this trend.

b. The ideal growth conditions at Del Monte also reverse a
trend for productivity to increase from the nearshore to an open water
environment .

c. Nitrogen to phosphorous ratios are pollutions tracers for
certain sewer outfalls.

d. Productivity to chlorophyll ratios indicate unique phyto-
plankton habitats or populations in the area studied.

2. Two different municipal sewer outfalls have profoundly different
effects on the environment. In just meeting the "letter of the law", the
Del Monte outfall enhances an already productive area. The Point Pinos
outfall, by adding toxicants beyond that required, creates a local abiotic

area.

3. A correlation is seen between carbon monoxide production and
primary productivity at the deep water stations and the nearshore oceanic
coast stations.

50

APPENDIX A - CRUISE DATA

Symbol Moaning

CRU cruise number

Z depth in meters of sample

T sea surface temperature in C

LOC location

transect

Se Secchi depth in meters

-3 -1

PP productivity in mgC m hr

3

Cl chlorophyll in mg/m

C/P acid factor

N3 nitrate in ug-at/1

S silicate in ug-at/1

P phosphate, in ug-at/1

N2 nitrite in ug-at/1

DM Del Monte

PC Pt. Cabrillo

PN Pt. Pinos North

PS Pt. Pinos South

PJ Pt. Joe

CC buoy adjacent to Pt. Cabrillo

HAR Monterey harbor

51

Week of

St

udy

Cruise

number

' applicable

1

1,2

2

3,4

3

5,6

4

7,8

5

10

6

11,

12, 13

7

14

8

15,

16

9

17,

18, 19

10

20,

21

52

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60

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Welch, R.H., A Study of the Stratification of Phytoplankton at Selected
Locations in Monterey Bay, California, M.S. Thesis, Naval Post-
graduate School, 1967.

Yentsch, C.S. and Menzel, D.W., "A Method for the Determination of Phyto-
plankton Chlorophyll and Phaeophytin by Fluorescence," Deep-Sea
Research, v. 10, p. 221-231, 1963.

62

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Naval Postgraduate School

Monterey, California 93940

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Department of Oceanography

Naval Postgraduate School
Monterey, California 93940

4. LT John V. Rowney, USN 1
1208 Await Drive

Mountain View, California 94040

5. Department of Oceanography 3
Naval Postgraduate School

Monterey, California 93940

6. Oceanographer of the Navy 1
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63

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{Security classification ol title, body ol abstract and indexing annotation mu»l be entered whrn the overall rrf'" '< f'"" j

2b. CROUP

"ri&inatinG activity (Corporate author)

I Naval Postgraduate School
Monterey, California 93940
_ . —

":EPORT TITLE

Gradient Analysis of Phytoplankton Productivity and Chemical Parameters
in Polluted and Other Nearshore Habitats

ie. IKPONI »LCU»IT» CL»1JIFK<

Unci as si fi ed

4JESCRIPTIVE NOTES (Type ol report and,lncluaive dates)

Master's Thesis; March 1973

~KU THORI5I (First name, middle initial, last noma)

John Victor Rowney

e 3EPOR T DATE

March 1973

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PROJEC T NO

7«. TOTAL NO. OF PAGES

65

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2 5

Da. ORIGINATOR'S REPORT UUMStRHI

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Approved for public release; distribution unlimited.

SUPPLEMENTARY NOTES

2. SPONSORING MILITARY ACTIVITY

Naval Postgraduate School
Monterey, California 93940

ABSTRAC T

Measurements of primary productivity, chlorophyll standing crop,
and nutrient concentrations were made along a gradient of five nearshore
habitats, a seaward transect, and a longshore transect to determine en-
vironmental relationships. Hie effects of municipal sewer outfalls, type
of shoreline, and degree of exposure to high winter seas were found to be
dramatic. The behavior of nutrient ratios suggest their use as pollution
tracers in certain circumstances. The ratios of productivity to chloro-
phyll demonstrated physiological regimes among the phytoplankton in the
sampling area. Comparison of data with carbon monoxide and methane con-
centrations provided a possible correlation between phytoplankton pro-
ductivity and carbon monoxide production.

)D,fr,\.1473

/N 0101 -807-681 1

(PAGE 1)

64

Security Cl«»»ilic«

Uon

1- 3140S

UNCLASSIFIED

Security CUtsif iretion

key wo RDI

NOLI I '• "

primary productivity

plankton

pollution

nearshore

nutrients

chlorophyll

sewage

effluent

carbon monoxide

methane

sublittoral

zooplankton

grazing pressure

acid factor

DD,f.r..1473 <B"*>

UNCLASSIFIED

S/N 0101 -807-6821

65

Security Cl««»ific«tion

A • 3 1 409

i 4 M A Y 7 5

I'll T -*

2 3 8 7 9

Thesis H6ZM

R823 Rowney

c.l Gradient analysis of
phytoplankton produc-
tivity and chemical
parameters in polluted
and other nearshore hab-
itats.

m

I 4 MAY7S

2 3 8 7 9

Thesis 1^252

R823 Rowney

C.l Gradient analysis of
phytoplankton produc-
tivity and chemical
parameters in polluted
and other nearshore hab-
itats.

thesR823

Gradient analysis of phytoplankton produ

3 2768 001 97079 1

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