~
ZOOPLANKTON
ALONG THE
CONTINENTAL
SHELF OFF
NEWPORT,
OREGON,
1969-1972:
distribution,
abundance, season~
cycle, and year-to-year
variations
William Peterson
Charles Miller
OREGON STATE UNIVERSITY
SEA GRANT COLLEGE PROGRAM
Publicationno. ORESU-T-76-002
OREGON STATE UNIVERSITY
SCHOOL OF OCEANOGRAPHY
JUNE 1976
authors
WILLIAM T. PETERSON
is a Resea~ch Assistant
Oregon State University.
CHARLESB. MILLER is Associate
University.
Professor
(Unclassified)
of Biological
in Biological
Oceanography
Oceanography at
at Oregon
State
The Oregon State University
Sea Grant College Program is supported
cooperatively
by the National
Oceanic and Atmospheric
Administration,
U.S. Department
of Commerce, by the State of Oregon, and by participating
local governments
and private industry.
The OSU Sea Grant College Program attempts
to foster
discussion
of important
marine issues
A balanced presentation
by publishing
reports,
sometimes dealing with controversial
material.
is always attempted.
When specific
views are presented,
they are those of the authors,
not
of the Sea Grant College Program, which does not take stands on issues.
related
publications
OREGON'S ESTUARIES: DESCRIPTIONS AND INFORMATIONSOURCES, by K.L. Percy,
D.A. Bella and P.C. Klingeman.
Publication
No. H-74-001.
294 pp.
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Summarizes much of the known information
about Oregon's
13 major estuaries,
excluding
only the Columbia River estuary.
Numerous citations
of literature
and agencies
from which
further
information
can be obtained
make this an excellent
starting
point for assembling
physical,
chemical and biological
data about Oregon's
estuaries.
THE FUTURE MANAGEMENTOF THE OREGONCOAST: PROCEEDINGS OF A SYMPOSIUM HELD AT THE LAW
CENTER, UNIVERSITY OF OREGON, OCTOBER 27, 1972.
Publication
No. W-74-001.
167 pp.
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for the Oregon coast.
Panels discussed
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contents
Introduction
Part
Part
II:
I: general
remarks
S
9
taxa
37
Bibliography
SS
Part III: appendices
S9
remarks
on important
introduction
This technical report summarizes the
results of a three year study of the zooplankton living in the nearshore zone off
Newport, Oregon.
The sampling program was
part of the Early Life History (ELH) project
of the Oregon State University Sea Grant Program and was designed to study the ecology
of the larval forms of all fishes, shrimps
and. crabs, as well as the zooplankton.
These samples are valuable because they are
the first from Oregon to have been collected
and analyzed from stations located very near
to shore.
In the past, zooplankton studies
have been based on samples collected along
transects beginning at stations 5 or 15
miles from the beach, or on samples collected well offshore over the continental slope.
Our samples are also the first collected
within the zone of intense coastal upwelling. Huyer (1974), among others, has
identified this zone as extending from the
beach out to about 10 km from shore.
Other available Oregon zooplankton data
are from one of three sources:
(1) Oregon
State University Master's and Doctoral
Theses of Cross (1964), Lee (1971), Hebard
(1966) and Laurs (1967), and (2) University
of Washington .samples collected at an irregular set of stations off the Washington and
Oregon coasts during 1961 and 1962 (Peterson
and Anderson, 1966; Peterson, 1972).
In
addition (3), a limited amount of data were
collected in August-October 1972 by L. Smith
(Holton and Elliot, 1973).
This report is divided into two parts.
Part I contains general remarks on zooplankton distribution, abundance and frequency
of occurrence in samples collected during
several summer upwelling and winter downwelling seasons. Differences between seasons
are stressed.
Part II contains detailed discussions of the ecology of important taxa.
Population dynamics, sex ratios and miscellaneous observations are presented where possible. Some of the results of this study
have been presented elsewhere (Miller and
Peterson, 1974; Peterson and Miller, 1975).
Part III, the appendix contains the zoo~
plankton enumeration data.
5
MATERIALS
AND METHODS
TSKflowmeters were mounted off-center
Zooplankton samples were collected along
the Newport Hydrographic (NH) line (on 440
40'N). This line was chosen because an extensive set of hydrographic data were available from the year 1960 to present, as
Oregon State University, Department of
Oceanography Data Reports.
Our samples were
gathered at stations NH 1, NH 3, NH 5, and
NH 10 during the period 22 June '1969 to
5 August 1972. After 3 February 1971 samples were collected as far seaward as 60
nautical miles (NH 60), weather permitting.
Figure 1 is a map of the study area.
45'
00'
45'
00'
in
each net mouth.
Tows were wade obliquely
over the entire water column (or from 0 to
140 m offshore of NH 20) using either a Vfin or Kite-Otter depressor.
A time-depth
recorder was used to record the towing track.
Samples were preserved with buffered formaldehyde solution.
Surface temperature was measured and a
surface salinity sample was taken at nearly
all stations.
Bottom salinity samples were
taken on many cruises.
Bathythermograph
traces were usually taken.
In the laboratory, samples were poured
into 500 ml pharmaceutical graduates and
allowed to settle. Several hours later the
settled volume was read. Water was then decanted off or added to make a diluted volume
of five times the settled volume.
Aliquots
for counting were removed from this volume
with a 1 ml piston pipette, after the animals were suspended by agitation.
w
~
40
30
20
10
44'
30'
44'
30'
'25'
00'
fig.
1.
Animals were enumerated with the aid of a
'24'
30'
Map of the study area showing the
Newport hydrographic line (solid
dark line) and the sampling positions.
Inset map shows detail of
the first ten miles of the coastline
with bathymetry.
Samples were collected with 20 cm
"Bongo" nets without an opening~.closing
mechanism.
To construct the net frame, two
20 cm diameter x 30 cm PVC plastic cylinders
were bolted to either side of a pivoting
wire clamp. When mounted on the towing
cable, the net frame was free to move in the
vertical plane about 1200 of arc, and it
could rotate horizontally about the towing
cable.
The two plankton nets were 145 cm long
and were constructed of NITEX nylon with
mesh apertures of 505 ~m and 240 ~m, The
ratio of net mesh area/net mouth area was
about 11.5:1 for the smaller mesh net. The
cod ends were 9 cm dia. x 16 cm PVC plastic
buckets with stainless steel mesh screens
cemented to laterally positioned
windows.
filtering
The two cod ends were fastened
approximately 15 cm apart with a stainless
steel strap, which kept the nets from wrapping about the cable while being towed.
6
binocular dissecting microscope at 25x magnification.
Usually five aliquots were
drawn from each sample. Taxa were counted
in the first and successive aliquots until
50 in a category were enumerated.
Approximately 400 animals were present in each aliquot. The counts were multiplied by appropriate factors to arrive at a number of individuals in a category per five aliquots.
A computer was used to convert these raw
data into number of individuals per cubic
meter and to carry out much of the summary
data analysis.
Two morphs of the genus Metridia were seen
and were separated on the basis of shape of
the prosome in lateral view.
The Metridia
pacifica type is more robust and has a steeply sloping forehead while the M. lucens type
has a much less sloping forehead.
Detailed
morphological study of the two types has not
been done. All adult copepods were divided
by sex and identified to species.
Copepodite
stages were identified for all species except ,those of Clausocalanus, Gaidius,
Gaetanus, and Heterorhabdus.
Only the nauplii of Calanus were distinguished.
Both
euphausiids and cladocerans were identified
to species: Euphausia pacifica, Thysanoessa
spinifera, and Evadne nordmanni and Podon
leukartii. The larvaceans rOikopleuraJ were
not identified,to species.
The chaetognaths
were not counted by species but the following were identified:
Sagitta elegans,
Eukrohnia hamata, Sagitta euneritica, S.
scrippsii
S. bierii, and S. minima.
The
holoplanktonic mollusks were represented by
Limacina helicina and Clione limacina.
'J
All other holoplanktonic taxa were counted
as general groups, e.g., amphipods, ostracods, scyphomedusae, ctenophores, siphonophores, etc. Meroplankton also were counted
by general categories such as barnacle
nauplii, bivalve veliger, and gastropod
veligers.
The crab larvae were not counted
because they were studied .separately from
these samples by R.G. Lough (1974).
THE COASTAL ENVIRONMENT
Aspects of the descriptive physical
oceanography of Oregon coastal waters can be
found in Burt and Wyatt, 1964;
Pattullo and
Denner, 1964; Cross and Small, 1967;
Collins, et al., 1968; Bourke, 1972; Pillsbury, 1972; Huyer, 1974; and Smith, 1974.
Water Circulation Patterns
The California Current is a slow
southward flowing current situated within a
broad band 300 to 800 km off the Oregon
coast.
Inshore of this permanent feature
the circulation changes with season. During
spring and summer months, nearshore flow is
southward, driven by northerly winds.
During autumn and winter months, after cessation of upwelling, nearshore flow becomes
northward.
This flow is called the Davidson Current.
Summer Hydrography
Northerly winds predominate from April
to September.
They produce a generally
southward transport of surface water with a
component to the right of the wind, away
from the coastline.
This offshore, nearsurface transport is balanced by northward
and onshore transport at depth of cold,
high salinity, nutrient rich water. When
northerly winds are strong and/or persistent,
this cold, high salinity water appears on
the beach and throughout the entire water
column out to about 10 km from shore. This
condition is called active or intense upwelling.
When north wind stress is weak or
nonexistent, the onshore deep transport is
weakened and nearshore waters are warmed by
.solar radiation.
deeper waters, upwelling is indicated in
hydrographic sections by upward sloping isolines of temperature, salinity and density.
The upward slope develops in April or May
and remains until cessation of upwelling in
September.
Offshore waters are warm (as
high as 170C) and a mixed layer is well developed.
Surface salinities are reduced as
a result of the Columbia River plume.
During any given summer, intense
southwesterly storms may approach the Oregon
coast. The result is an onshore transport
of warm surface water.
If the winds are of
sufficient intensity and duration, water will
pile up nearshore.
Nearshore hydrography is
modified such that isotherms, isohalines and
isopycnals either lie parallel to the sea
surface or slope downward toward the shoreline. This condition is called downwelling,
and is a special case of the overall April
to September upwelling condition.
Offshore
of approximately 10 km, isolines of temperature, salinity and density maintain their
upward slope, although the slope may be decreased somewhat.
See Huyer (1974)
for representative hydrographic sections.
Winter Hydrography
The autumn and winter months are
characterized by a high frequency of southwesterly winds.
The effect is a transport
of offshore surface waters onshore.
These
onshore winds may also drive the Davidson
Current, although this is not certain.
The
offshore waters show the characteristics of
any North Pacific Ocean station in winter:
the top 100 meters or so are isothermal and
the seasonal pycnocline disappears.
Salinities are reduced by rainfall, and at the
inshore stations they are reduced by runoff
as well.
Sea surface temperature over the
continental shelf are generally around 8 to
100C. The nearshore ocean can be warmer in
winter than it is in the active upwelling
period in summer. This is because warm
water is advected from the south by the
Davidson Current.
Intense upwelling is not continuous.
If
northerly winds blow for a day or two,
active upwelling will develop.
If the wind
ceases or changes direction, this upwelling
ceases.
During July and August, northerly
winds are often persistent enough that intense upwelling may continue uninterrupted
for several weeks.
Waters offshore of 10 to 20 km do not
experience these rapid changes.
In these
7
part I: general remarks
SUMMERHYDROGRAPHYAND WINDS
We have compared our surface
temperature
and salinity
observations
to Pillsbury's
summary (1972) of all
surface-to-lOm
temperature
(T) and salinity
(S) observations
collected
at NH 3 and NH 5 between 1960 and
1970.
The modal group (28 per cent of the
data)
in a bivariate
plot
of these observations
was bounded by temperatures
of 8.1oC
to 9.0oC and salinities
of 33.1 to 34.0
°ioo.
This is called
the modal cell.
The T-S data are shown in Figure
2.
The
bounds of the modal cell
are indicated.
The
upwelling
season of 1971 is shown to be
quite
different
from 1969 or 1970.
About
26 per cent of our data from 1969 and 1970
fell
within
Pillsbury's
modal cell,
Only
6 per cent of the 1971 data fell
within
the
limits.
Surface
temperatures
were higher
and surface
salinities
were lower in 1971,
indicating
the presence
of mixed Columbia
River plume water and surface
oceanic
water.
Upwelling
was weak in 1971.
Wind data support
the conclusion
that
the
1971 upwelling
season was weak.
We have con~tructed
Progressive
Vector
Diagrams
(PVD's),
of winds measured at the south jetty
off
Newport,
Oregon.
The PVD's for the years
1969, 1970, 1971 and 1972 are shown in
Figures
3,4 and 5.
Winds during
the upwelling season (1 May-30 September)
are quite
different
between these years.
The 1969
upwelling
season had relatively
weak winds
with only about two and one-half
months of
persistent
northerlies,
while
the 1970 season had about four months of northerlies.
A higher
total
northerly
component was
achieved
in 1970 than in 1969.
The 1971
upwelling
season was strikingly
different.
North wind miles
are low and west wind miles
are exceptionally
high.
As a result,
upwelling should have been weak in 1971.
This explains
the low frequency
of temperature
and
salinity
observations
in the modal cell
in
1971.
Winds during
the 1972 upwelling
season
had both a strong
northerly
component
(like
1970) and a strong
westerly
component
(like
1971).
All four years have the same pattern'
in September and October:
winds are light
and variable.
9
II
','
.1
-i
,.,
,',
,
""
'.'"
I. "..
1
.,
"
"::'
. . . . 0 .
"
..
".
"i!"
"
o.
II
IO[
,.
,
1 ,
II
,
.
,
.
..
.''.
,
.
"
"
o.
. '.,
'r
:..
..,;..->
"
o.
50
to
I.
'.'"
..
"71
.
1970-71
"
..
.\
."
-i
c I.
::0
,.,
..'". "4
;..,.
,
".
".'-.7.
.
"
,., to
::0
J>
, .,.
0.
..
-i
,.,
."
"
,.,
::0
J> ,.
1870
-i
C
::0 .1
,.,
,. t-
II
II
t
,
SALINITY
,
,
II
(L)
..
.
1971-n
,
i
'. ".,
.
o.
SALINITY
"
(%.)
. Fig. 2. Temperature-salinity diagrams for NH1, NH3, NH5, and NH10 for the summersof
1969,.1970 and 1971 between April and September. and winters of 1959~70. 1970-71,
and 1971.;.72from October through Harth
Winter HydX'ography
and Winds
Winter T-S scatter diagrams are also
shown in Figure 2. The winter of 1969~70 is
seen to have been warmer than the other winters. This may indicate that the transport
of the Davidson Current was greater during
the winter of 1969-70.
The winds during the winter of 1969-70
were--quite-dIfferent compared to 1970~71 and
1971-72. During the fall and winter months
of 1969, there were three intervals with
winds from the east: most of October, 23
November-S December, and 1-12 January.
The
entire six-month period of winter winds
lacked the southwesterly storms that are
characteristic of most winters.
T~e other
two winter wind patterns shown in Figures
3-5 are probably typical.
Patterns of Surface Salinity
As a result of rainfall and runoff,
salinities less than 30 0100 occur at stations near the beach (NH 1, NH 3 and occasionally NH 5) during the months of January
and February.
Beyond NH 5, salinities were always
greater than 32 0/00 during these two
months.
March and April are months of
transition.
In May and June salinities are
again lowered, but this time by the Columbia
River plume.
Surface salinities of 26-30
0/00 are co.-only encountered at varying
distances fro. shore, for exa8p~e during
10,
middle and late June of 1969 and 1971,
salinities of 30-31 0/00 were found as close
as 1 mile from shore.
In 1970, water of reduced salinity was never found closer to
shore than 10 miles on any cruise.
Surface salinities in excess of 33.5 %0
in the coastal zone are generated only by upwelling.
Water of this salinity or greater
first appeared in Mayor
June in the years
of our study (13 May 1969, 4 June 1970 and
29 May 1971). The autumn months (OctoberDecember) are months of transition to
lowered salinities as a result of rainfall
and runoff.
Bakun's Upwelling Indices
Another parameter useful in studying
differences between seasons is the upwelling
index.
Bakun (1973) has calculated indices
which express mean monthly amounts of upwelling or downwelling at points along the
west coast of the United States ranging from
Baja California (Lat. 1SoN) to the Gulf of
Alaska (60oN). Monthly index values were
calculated for the years 1946-1971.
The indices are still being calculated but on a
daily instead of monthly basis (Bakun, personal communication).
To calculate the indices, Bakun estimates the monthly mean wind
stress at selected geographic points from
the known atmospheric pressure field. From
this he computes the Ekman transport and
finally resolves the component of this transport perpendicular to the coast. The units
of the index are cubic meters of water
/970
EAST
---
.mn
-
6
/969
-,._.~--~---
.
EAST
n
IM
t
,
I-
31 DEC
I JUNE
6
'\
I
6\
I
EC
[
r
r14
I
SOUTH
1
I
SOUTH
I/o
1
11"-1
SEPT
Fig. 3. Progressive vector diagrams for the wind at Newport for the years 1969
from 16 April--31 Decemberand all months of 1970.
3 I DEe J
/97/
c
A
EAST
20j
)
en
z
c
en
I
0
I
r
"'TI
NORTH
:E
Z
/0
C
I
"-
r
0
3:
'ITI
-I
ITI
::u
en
d
4 JAN
/0
20
THOUSANDS
OF
30
WIND
-
40
KILOMETERS
Fig. 4. Progressivevector diagramfor the wind at Newportduring 1971.
11
30
-I MAY
/972
EAST
~
20
--I
::I:
0
C
(J)
1>
Z
0
(J)
10
0
."
10
:E
z
THOUSANDS
0
A
r
0
20
OF
WIND - KILOMETERS
1
3:
rrI
--I
rrI
::c 10
NORTH
(J)
/'
L DEC
I NOV
20
Fig. 5. Progressive vector. diagram for the wind at Newport during 1972.
12
I~
upwelled
(or downwelled)
100m of coastline.
per
second
per
The mean monthly values of the index and
the anomalies
(i.e"
the deviations
from the
20-year mean) are shown below.
The anomalies show that the summers of 1969 and 1970
were near normal, and that the summer of
1971 was unusually
low indicating
weak upwelling.
This supports
our observations.
20 year
mean
index
1969
1970
1971
34
-22
48
74
51
17
+13
+32
- 5
-11
-- 12
+32
-36
- 3
+23
- 9
-27
+7
+12
-48
for month
May
June
July
August
September
Tota1s
- 5
- 8
The winter upwelling
index data are
shown below.
The winter of 1969-70 is
quite different
from the other two winters.
Onshore transport
was anomalously
high in
1969-70 and anomalously
low in 1970-71 and
1971-72.
This does not agree entirely
with
our Newport wind data.
Both Bakun's and
our data agree that the winter of 1969-70
was different
but we disagree
as to the direction
of surface
water transport.
20 year
index
19691970
19701971
19711972
-74
-93
-94
21
-64
-47
-24
+19
-12
+62
+32
+34
+66
+75
+56
-71
+101
+119
mean
for month
November
December
January
February
Totals
-4
ZOOPLANKTONDATA COLLECTION AND GENERAL
OBSERVATIONS
Zooplankton Data And Their Limitations
A total of 523 Bongo net samples were
collected
at stations
NH 1 through NH 60.
At stations
NH 1, 3, 5, and 10 only, we collected
329 samples and have counted 213 of
them.
In addition,
40 samples have been
counted from the set collected
'offshore
of
NH 10. The total
number of samples counted
is 253.
Within the extensive
set of 523
samples are many replicate
samples and daynight comparisons.
None of these have been
counted.
This report
will deal primarily
with the samples collected
within 10 miles
of shore since that data set is much more
complete.
Appendix I is a list
of the
sampling dates and indicates
the types of
samples collected
(day-night,
replicate,
day only) and which samples were counted.
Appendix II contains
zooplankton
enumeration
data listed
by taxa for stations
NH 1, 3, 5,
and 10. Appendix III lists
the data collected offshore
of NH 10.
In this report,
summer is defined
as the
upwelling
season (April-September)
and winter
is defined as the other months (OctoberMarch) .
There are important
limitations
on these
zooplankton
data.
We chose to express nu~
merical
abundance as numbers of individuals
per cubic meter "(no. 1m3).
Since our nets
were towed obliquely
through the entire
water column, quantitative
abundance estimates are actually
abundances
averaged over
the water column.
If an animal is equally
abundant at all depths then oblique
tows
will adequately
estimate
its abundance.
If
an animal is restricted
to a narrow layer
then its abundance will be underestimated.
We could have used number of individuals
beneath a square meter water column by multiplying no. 1m3 by the water depth, but the
effect
of tow depth can be handled without
this conversion.
Recent work by ourselves
and Myers (1975)
has shown that highest
zooplankton
abundances
are found within the top 20 to 30m of the
water column.
Therefore,
our oblique
tows
from depths greater
than about 30m do underestimate
zooplankton
abundances.
This becomes a problem in tows taken at stations
farther
from shore as the water depth increase~ because an increasing
fraction
of
the water column sampled contains
few animals.
Therefore,
one is not justified
in
discussing
abundance gradients
between stations NH 1 (water depth = 20m) and NH 10
(water depth = 80m) unless
abundance differences are greater
than a factor
of four.
Abundances are also underestimated
for
many copepod taxa because the small copepodite ;stages could easily
pass through our
0.24 mm mesh net.
Copepodites
of Pseudocalanus and Acartia
species
younger than
stage III were seldom seen in our samples.
Probably only stages IV and V were sampled
quantitatively.
This data set gains its value from being
a three-year
time series
of samples collected
in exactly
the same manner at the same stations.
As such, these are. good baseline
data
to which future work can be compared.
The
data clearly
show
(1)
seasonal
and (2) year-
to-year
variations
in zooplankton
species
composition
and abundance,
and (3) spatial
13
(i.e.,
inshore~offshore) differences in
species
composition.
Point estimates
of
abundance have little
meaning, but comparisons of abundances between seasons and years
at a set of stations
are valid and meaningful.
Fpequency of Occuppence of ZoopZankton Taxa
Copepods were the most frequ~ntly
occurring
and the most abundant members of
the zooplankton
community in the nearshore
region off Newport, Oregon.
A total
of 58
species
were seen in our samples (Table 1).
Thirty-eight
species
were found in the summer samples and 51 species
in the winter
samples.
~ring
our study, species
from the
Subarctic,
Transition
and Central
Pacific
water masses were taken.
The copepods in Table 1 can be grouped on
the basis of patterns
of occurrence.
Eight
species
occur commonly during both winter
and summer months: CaZanus mapshallae3 PapacaZanus par'Vus3 Pseudocalanus
sp., Metpidia Zucens3 ACaPtia clausii3
ACaPtia longi~
pemis3 Oithona simiUs3
and Oithona spinipostpis.
Six species were found onlyduring the
summer months and so probably have northern
affini ties:
Aetideus paci.ficUB. Gaidius
immatures , Gaetanus 'iminatures, Racovitzanus
antaPcticas
s .1., Metpidia pacifica3 and
Oncaea media hymena.
Only one species
was common during
the
summer and uncommon during
the winter:
Centpopages
abdominaUs.
This species
has
northern
affinities.
A group of six species
had the opposite
characteristic,
i.e.,
common during
the winter
but uncommon or rare
during the summer:
CaZanus tenuicoPnis3
CZausocaZanus apcuicoPnis3 CZausocaZanus
pepgens3 CtenocaZanus vanus s.l., ACaPtia
tonsa and Copycaeus angZicus.
All of these
species are common in warmer water south o£
Oregon.
The majority of the copepod species (43)
were always uncommon in our samples and
probably have unimportant roles in the community.
However, taxonomic study of these
rare or uncommon species is important because
in many cases their presence indicates
the presence of a particular water type or
mixture of types. Most of the species that
are found off Newport only during winter
months have southern affinities (Central
Pacific water mass).
They are transported
north along the continental shelf by the
very near
Davidson current and are probably
the extreme northerly limit of their range.
14
Someof these species were Mecynocera
CaZocaZanus styZipemis3
CaZocaZanus
tenuis3 CaZocaZanussp., CZausocaZanusmas-
cZausii3
tigophopus3 CZausocalanus fupcatus3 ClausocaZanus jobei3 ClausocaZanus papapepgens3
CZausocalanus pauZuZus3 ACaPtia danae3 CoPycaeus amazonicus3 Oncaea dentipes and Oncaea
subtUis.
Other species that were found only during
winter months have northern affinites and are
usually found in deep water over the continental slope. They were probably transported
shoreward as a result of onshore winds.
These species are CaZanus cpistatus3 Gaidius
bpevispinus3
Gaetanus
simplex3
Candacia
co-
Zumbiae3 Hetepophabdus immatures. PZeupomamma bopeaZis3 and PleupomammaabdominaZis.
Eight of the remaining 14 species in
Table 1 were uncommon in both summer and winter samples: EucaZanusbungii3
Micpocalanus
pusillus3 ScoZecithpiceZZa minoP3 Lucicutia
fZavicoPnis3
amphitpites),
EpiZabidocepa Zogipedata (E.
Toptanus discaudatus3 Micpo-
setella spp., and Sapphippina spp. Six species were rare during both seasons: RhincaZanus nasutus3 PaPaeuchaeta japonica3 Oncaea teneZla3 Oncaea bopeaZis3 Oncaea conifepa3 and Oncaea medittepanea.
Abundance of ZoopZankton Taxa
Tables 2 and 3 list the average relative
density and frequency of occurrence of the
copepod species, totaled over the four stations, for three individual years, during
the summer and winter respectively.
Relative
density is an animal's mean abundance in
those samples in which it occurred.
The most
abundant copepods found during the summer, in
order of decreasing total average abundance
werePseudocalanus sp., Acaptia cZausii3
Acaptia Zongipemis3 CaZanus mapshaZZae3 Centpopages abdominaZis3 and Oithona simiZis.
These six species almost certainly control
the grazing dynamics of the zooplankton community.
No other copepod species ever approach numbers consistently reached by these
six species.
During the winter months, the zooplankton
assemblage is quite different and abundances
of all species are much lower. Four species
are prqbablY dominant: PseudocaZanus sp.,
Oithona simiZis3
PapacaZanus papvus3 and
ACaPtia Zongipemis.
Other species that may
play important roles at certain times between October and March are CtenocaZanus
vanus3 CZausocaZanus immatures, ACaPtia cZausii3 Centpopages abdominaZis3 and CaZanus
mapshaUae.
~
11
C = occurrence in greater than 50% of the samples taken,
U = occurrence in less than 50% but more than 5 samples taken,
R = occurrence in less than 5 samples.
COPEPOD SPECIES
Calanus marshallae
C. tenuieoPnis
C. plumehY'us
.C. eY'istatus
Rhinealanus nasutus
Euealanus bungii
MeeynoeeY'a elausii
Paraealanus paY'Vus
Caloealanu8 styliY'emis
C. tenuis
C. sp.
Pseudoealanus sp.
MieY'oealanus pusillus
Clausoealanus mastigophoY'Us
C. fUY'eatus
C. areuieoPnis
C. jobei
C. pegens
C. paY'apegens
C. paululus
Ctenoealanus vanus
Aetideus paeifieus
Gaidius immatures
Gaidius bY'evispinus
Gaetanus immatures
Gaetanus simplex
Paraeuehaeta japoniea
Raeovitzanus antaretieas
SeoleeithY'ieella
minoY'
Table 1.
S
W
.C C
U C
R U
R
R R
U U
U
C C
U
U
R
C C
U U
U
R
U C
R
U C
U
.R
U C
U
U
R
R
R
R R
U
U U
COPEPOD
SPECIES
MetPidia lueens
M. paeifiea
Lucieutia flavieoPnis
Candaeia eo lumbiae
C. bipinnata
HeteY'oY'habdus immatures
PleUY'omammaborealis
P. abdomina lis
Centopages memuY'Y'iehi
Epilabidoeea
amphitY'ites
Aeartia elausii
A. longiemis
A. tonsa
A. danae
EUY'ytemoY'asp.
TOY'tanus diseaudatus
MieY'oseteUa sp.
SapphiY'ina sp.
Oithona similis
o. BpiniY'ostis
Oneaea tenella
o. bOY'ealis
o. eonifeY'a
o. medi tteY'anea
o. dentipes
O. subtilis
o. media hymena
Coyeaeus anglieus
C. amazonieus
S W
C C
U
U U
R
R R
R
R
R
C U
U U
C C
C C
U C
R
R
U U
U U
U U
C C
C C
R R
R R
R R
R R
R
R
R
R C
R
A checklist of copepod species
taken off Newport, Oregon in summer (S)
and winter (W) months during the period of our study.
.
15
COPEPODSPECIES
.
TOTAL RELATIVE DENSITY
Calanus marshallae
C. tenuieornis
Euealanus bungii
Paraealanus parvus
Pseudoealanus sp.
Mieroealanus pusillus
Clausoealanus areuieornis
C. pergens
C. immatures
Ctenoealanus vanus
Aetideus paeifieus
Gaidius immatures
Raeovitzanus antaretieas
Seoleeithrieellaminor
Metridia lueens
M. paeifiea
Lueieutia flavieornis
Centropages abdominalis
Epilabidoeera longipedata
Aeartia elausi
A. longiremis
A. tonsa
Tortanus diseaudatus
Oithona simi lis
o. spinirostris
Coryeaeus anglieus
FREQUENCY
1969
1970
1971
69
70
71
1475.2
1.3
21.1
80.7
23776.3
2.2
0
20.2
0
4.8
1.5
2.5
0.6
9.3
21.4
6.7
0
371.8
2.9
1178.1
1078.2
130.7
17.5
482.5
5.1
3.1
147.3
6682.4
18.5
1.4
5.9
0.5
31.0
2.3
3.4
2. 1
4.6
16.3
436.1
7.9
9.0
16.8
3994.5
1.8
4.0
6.6
2.1
11.0
2.7
3.7
1.3
16.0
48.6
2.9
3.7
0.4
686.2
10.8
6045.1
1509.2
27.5
12.3
2.0
110.7
0
414.9
1331.1
0
0
32
1
13
29
33
4
0
5
0
5
4
3
1
7
18
2
0
29
5
31
33
31
3
44
4
10
21
44
17
3
5
5
7
4
3
2
14
29
5
1
42
6
37
44
19
10
40
14
15
20
40
2
7
9
2
16
1
2
2
16
26
6
9
23
0
29
38
0
0
369.9
19.9
2.8
275.0
16.0
8.0
416.2
55.5
3.6
32
19
2
42
28
10
40
31
3
The following species were found in less than five samples: Calanus plumehrus,
Gaetanus immatures, Paraeuehaeta japoniea immatures, Candaeia bipinnata,
Eurytemora
thompsoni,
Rhinaalanus
nasutus,
Oneaea borealis,
Oneaea tenella,
Oneaeamedia hymena, Oneaeamediterranea, Sapphirina sp. and MieroseteUa
Table 2. Total relative
density and frequency of occurrence of the copepod species
taken within 18 km of the Oregon coast, during the upwelling seasons of
1969, 1970, and 1971. Realative density is average number per m3 of
individuals in samples in which the species occurred. Table entries are
sums of relative densities at each of four stations.
33 samples were
collected in 1969, 44 in 1970 and 40 in 1971.
16
sp.
TOTALRELATIVEDENSITY
COPEPOD
SPECIES
FREQUENCY
1969-70 1970-71 1971-72 69-70
62.7
15.1
359.6a
3.4
0.8
570.6
7.4
3.3
445.6
3.6
2.3
38.9
41.1
5.8
156. 6a
189.0
***
17.6
0.5
48.0
1.6
186.4
332.8
121. 9
8.5
6.8
189.0
7.9
1.9
***
0.4
277.7
0.8
0.3
2011 .1
1.1
***
36.5
22.8
3.7
38.2
98.8
0.9
42.9
0.4
33.3
0.8
78.8b
84.9
8.5
6.3
4.2a
46.6
14.3
2.3
3.5
***
351. 4
1.0
0.5
1260. 1
1.3
***
31.5"
30.3
***
32.4
65.9
1.1
40.8
0.8
127.5
1.2
95.3
25.0
7.4
4.4
***
22
11
4
6
5
22
10
7
22
3
6
13
14
8
17
16
0
16
6
8
5
14
18
19
13
9
20
14
1
0
2
18
3
1
20
1
0
16
15
3
16
16
4
14
4
5
2
8
20
8
4
3
15
12
3
8
0
16
1
1
16
2
0
10
12
0
9
10
1
10
3
7
1
12
15
4
4
0
1064. 0
15.3
15.2a
397.1
0.9
246.9
7.1
0.3
20.1
1.1
625.4
22
21
2
21
10
20
8
3
11
1
16
8
0
0
0
CaZanus marshaZZae
C. tenuieornis
C. PZumehrus
EueaZanus bungii
Meeynoeera eZausii
ParaeaZanus parvus
CaZoeaZanus styZiremis
C. tenuis
PseudoeaZanus sp.
MieroeaZanus pusiZZus
CZausoeaZanus mastigophorus
C. areuieornis
C. pergens
C. parapergens
C. immatures
CtenoeaZanus vanus
SeoZeeithrieeZZa minor
Metridia Zueens
Lueieutia fZavieornis
Centropages memurriehi
EpiZabidoeera amphitrites
Aeartia eZausii
A. Zongiremis
A. tonsa
A. danae
Tortanus diseaudatus
Oithona simiZis
o. spinirostris
Oneaea teneZZa
Coryeaeus angZieus
C. amazonieus
70-71 71-72
3.5
***
***
***
= high value the reult of one station or sampling date
b = a value of 4500/m" on 20 October 1970 was omitted from the summation
a
CaZanus erisThe following species were found in less than five samples:
tatus, RhineaZanus nasutus,CaZoeaZanus
sp., CZausoeaZanus jobei, C. pauZuZus, Gaidius brevispinus,
Gaetanus simpZex, Paraeuehaeta japoniea, PZeuromamma boreaZis,
P.
abdominaZis,
Heterorhabdus
immatures,
Candaeia
eoZum-
biae, C. bipinnata,
Oneaea boreaZis, o. eonifera,
o. dentipes,
o. subtiZis,
o. meditteranea,
MieroseteZZa sp., Sapphirina sp., Harpacticoid copepods.
Table 3.
Total relative
density and frequency of occurrence of the copepod species
taken within 18 km of the coast during three winters.
Table entries are
sums of relative
densities
at each of four stations.
22 samples were
collected during the winter of 1969-70, 20 during 1970-71 and 16 during
1971-72.
A rough estimate of the magnitude of
differences in abundance between summer
dances (No./m3) in each yea~ have been
summed from Tables 2 and 3 to get an over-
and winter for the most important copepods
all
can be seen in the following
table.
total
abundance.
Abun-
17
CopepodSpecies
Acartia
SUMMER
7,638
WINTER RATIO
BETWEEN
SEASONS
360
- 21.2
clausii
34,453
3,717
- 9.2
2,394
299
- 8.0
Centropages
1, 169
209
- 5.8
Acartia
3,918
1,143
- 3.4
1,061
1,936
+
0.6
1,200
+
4.9
354' +
7.5
Pseudocalanus
sp.
Calanus
marshaUae
abdomina lis
longiremis
Oi thona simi lis
Paracalanus
parvus
245
Ctenoca lanus
47
vanus
Abundances of other
zooplankton
(the
holoplankton
and meroplankton)
indicate
that
they were generally
not important.
Overall
abundances
and frequency
of occurrence
of
these taxa are given in Tables 4 and 5 for
summer and winter
respectively.
The plankton that were sometimes abundant during
the
summer were Calanus nauplii, Calanus eggs,
Other taxa that were
and euphausiid eggs.
occasionally
abundant
include
decapod shrimp
mysis,
barnacle
nauplii
and bivalve
veligers.
During the winter
months,
Oikopleura spp.
became abundant, and barnacle nauplii and
bivalve veligers remained somewhat important.
would be likely to have considerable impact
on community
structure
if it made up at least
1 percent
of the total catch in five or more
samples.
This rather generous filter removed all but 15 copepod species and 19
other
zooplankton taxa from the total species
list. Many of the taxa excluded were important to the results of this study only as
indicator species.
The remainder of this
report will deal primarily with the 34 dominant taxa.
Table 6 lists these animals.
A comparison
of the first two columns reveals that only
a few taxa were important in all summer
samples. The only taxa that occurred often
and frequently made up greater
than 1 per
cent of the total
catch were copepods.
With
few exceptions, all other taxa made more
than 1 per cent of the catch in less than
25 per cent of the summer samples.
The preceding portions of this report
have centered
on general statements about
distribution and abundance of taxa that make
up the different zooplankton assemblages
found off Newport, Oregon during summer and
winter.
For simplicity, we disregarded temporal variations and treated
the 10-mile
wide study area as a homogeneous
area is not homogeneous however.
the
animals
are
not
equally
Dominant Taxa
22
18
23
28
18
of
over
distributed
ABUNDANCE PATTERNS
Spatial Abundance Patterns
UpUJeUing Season
our data which objectively categorizes importance.
It was assumed that an animal
The
Most
the study area and are not equally abundant
during all months of a particular sea~on.
Both spatial
and temporal
abundance gradients
are present and they are quite pronounced
for some of the most important copepod
species.
The reader can make the comparisons
between summer and winter abundances from
Tables 4 and 5.
Some taxa have similar
abundances
during
both seasons:
Calanus
nauplii, other copepod nauplii and euphausiid nauplii, chaetognaths, scyphomedusae,
Podon leukartii, pteropods, barnacle nauplii,
bivalve veligers and gastropod veligers.
are seen for
Marked differences in abundance
Thysanoessa spinifera, Evadne nordmanni,
decapod mysis, Calanus eggs and euphausiid
eggs. These were all more abundanti during
the summer. Oikopleura spp. were more
abundant during the winter.
Further discussions of abundance patterns
will be limited
to the more important animals.
We have applied a simple filter to
zone.
During the
Three of the six most abundant copepod
species almost always have their
greatest
summer abundance at the station
shore (NH 1).
These three
are
sp.,
Acartia
inalis.
clausii,
The abundance
nearest
and centropages
gradients
to
Pseudocalanus
abdom-
can be
quite steep, for example,pseudocalanus
sp.
had the following abundances {no. 1m3) at'
the four
stations:
DATE.
NH 1
NH 3
NH 5
NH 10
1969
1969
1970
1971
13,97,5
'43,053
23,773
5,259
2,322
2,706
487
178
2,000
8,960
19
78
461
351
104
***
21 July 1971
5 Aug. 1972
7,330
1,524
155
337
132
140
290
June
July
June
June
***
I:
"
I
I
TOTAL RELATIVE DENSITY
TAXA
FREQUENCY
I
1969
1970
1971
69
70
71
119.5
43.1
8.5
46.3
13.3
30.2
35.4
73.7
2.8
10.2
89.4
69.2
6.0
22.9
695.5
68.1
18.5
85.9
14.5
13.6
4.0
58.9
1153
246
50.3
85.7
2.5
70.9
172.7
52.3
15.7
84.0
17.2
17.7
87.3
9.8
5.2
60.6
30.8
66.5
34.9
22.8
21
10
5
5
4
14
2
17
2
11
25
11
7
13
40
20
15
26
17
20
7
26
12
22
33
15
5
28
28
20
14
18
11
10
11
2
1
35
34
21
19
22
decapod shrimp mysis
barnacle nauplii
barnacle cypris
polychaete posttrochophores
bivalve veligers
gastropod veligers
hydromedusae
unidentified
annelid
without parapodia
plu1;eus
142.7
59.3
4.4
52.6
168.3
64.0
45.3
231.4
8.3
16
8
2
24
32
19
22
28
10
16.2
170.5
28.9
6. 1
20.1
258.9
79.2
3.2
21.4
68.3
42.2
10.3
5
20
16
2
23
40
33
2
15
27
23
11
8.2
0.0
23.1
16.0
35.8
117.6
3
0
3
5
16
11
large round eggs (fish)
36.8a
870.1
55.0
70.0
19.1
25.0
168.7
686.1
57.5
35.1
17.8
226.1
449.6
39.6
34.3
11
10
11
2
12
13
28
29
16
18
12
25
24
14
18
Cal-anus naupl i i
Other Copepod nauplii
Amphipods
Euphausiid nauplii
Euphausiid calyptopis
Euphausiid furcilia
Thysanoessa spinifera
Evadne nordmanni
Podon l-eukarti
Pteropods
Chaetognaths
Oikopl-eura
Ctenophores
Scyphomedusae
I
II
I
II
I:
I'
I
I
Cal-anus eggs
I
euphausiid eggs, early
euphausiid eggs, late
other fish eggs
I',
.
a = biased by a single observation of 760 individuals/m3.
I
The following taxa were found in less than five samples: radiolarians,
foraminifera, siphonophores, planula larva, trochophores, Tomopteris3
heteropods, Cl-ione3 phoronid larva, ascidian larva, salps, auricularia
larva, immstarfish, decapod protozoeas, unusual barnacle nauplii, Styl-ocheiron abbreviatum3 anchovy eggs, and four miscellaneous unidentified
meroplanktonic taxa.
I
Il
I
Table 4.
I
Total relative density and frequency of occurrence of other holoplanktonic
taxa and meroplankton taken within 18 kmof the coast during 1969. 1970
and 1971 upwelli ng seasons. Table entri es are sums of average abundances
at each of four stations.
I
I
'
I
19
TAXA
CaZanus naup1ii
Other Copepodnaup1ii
Amphipods
Euphausiid naup1ii
Euphausiid ca1yptopis
Euphausiid furci1ia
Evadne nordma:nni
Podon Zeukarti
Pteropods (Limacina)
Chaetognaths
OikopZeura spp.
Ctenophores
Scyphomedusae
Sa1ps
Isopods
Mysids
decapod shrimp mysis
barnacle naup1ii
barnacle cypris
polychaete post-trochophores
bivalve ve1igers
gastropod ve1igers, assorted
gastropod A
hydromedusae
annelids lacking parapodia
echinoderm p1uteus
large round eggs (fish)
CaZanus eggs
euphausiid eggs
TOTALRELATIVEDENSITY
FREQUENCY
1969-70 1970-71 1971-72 69-70 70-71
71-72
10
11
12
4
13
7
2
4
17
20
22
8
5
9
2
2
15
13
4
5
4
2
2
2
15
19
16
8
6
0
3
1
15
12
10
4
8
5
4
4
13
13
15
9
10
0
0
2
5.6
77.9
16.8
70.8
118.4
37.2
***
3.3
21. 9
22.1
7
11
4
12
20
19
0
4
5
5'
10
6
4
8
18
18
6
2
4
2
11
12
12
11
15
15
0
3
11
4
4.9
4.7
2.8
6
10
0
11
11
6
8
4
3
1188.7a
29.1
5.9
2.8
6.4
3.1
5.8
126.3a
66.0
62.9
551.9
7.0
10.0
0.9b
0.5
0.2
3.1
309.1
8.7
41. 5
87.8
31.3
***
165.9
122.5a
4.8
108.4a
56.1a
0.4
24.1
27.3
88.0
47.4
101.2
6.2
94.3
***
0.7
3.3
21.4
192.7
188.1 a
13.5
98.2
27.6
1.0
9.2
1.8
40.0
41.7
74.9
0.8
9.0
36.5
***
5.5
36.7
274.7a
35.1
20.2
5.0
3.4
14.5
7.6
4.8
116.4a
14.2
22.4
75.6
10.3
16.6
***
***
2.1
value
the re3u1t
of October
one station
sampling
date
= ahigh
value
of 34.3/m
an 29
'1969orwas
ommitted
from the summation
ba =
The euphausiids
The following taxa were found in less than five samples:
Thysanoessa spinifera
and Euphausia pacifica,
amphipod larvae and eggs,
ostracods,
cumaceans, siphonophores, Sagitta scrippsii,
S.
bierii,
S.
minima, Lepas nauplii,
other unidentified
barnacle nauplii,
echinoderm
bipinnaria,
imm starfish,
imm sea urchins, planula larvae, trochophores,
foraminifera,
radiolarians,
Tomopteris, cyphonautes larvae, other fish
eggs, and six miscellaneous unidentified
merop1anktonic taxa.
Table 5.
20
Total relative density and frequency of occurrence of other ho1oplanktonic
and meroplanktonic taxa taken within 18 km.of the .coast during three
winters. Table entries are sums of relative densities at each of four
stations.
I
I
}
~
\
!
I
\
r
I
J
~
TAXA
Pt
Ii
,
t
,(
't
l
It
I
II
vI
,I
I
,
II
(
{
PseudoaaZanus sp.
CaZanus marshaUae
Oithona simiZis
Aaartia Zongiremis
Aaartia aZausii
ParaaaZanus'parvus
GaZanus naupl i i
OikopZeura sp.
Gentropages abdominaZis
bivalve veligers
Limaaina heZiaina
Chaetognaths
barnacle nauplii
GaZanus eggs
euphausiid eggs
GtenoaaZanus vanus
Metridia Zuaens
GZausoaaZanus immatures
Oithona spinirostris
CZausoaaZanus pergens
C.
arauiaornis
Aaartia tonsa
euphausiid nauplii
gastropod veligers
worms lacking parapodia
Goryaaeus angZiaus
echinoderm pluteus
decapod shrimp mysis
Scyphomedusae
cope pod nauplii
Ctenophores
GaZanus tenuiaornis
Podon Zeukartii
misc. fish eggs
SUMMER
WINTER
BOTH
ALL
FI LTERED
SAMPLES SAMPLES
FILTERED
SAMPLES
TOTAL
FILTERED
141
140
138
139
'115
76
109
53
110
107
88
107
83
82
48
39
81
8
97
26
12
53
53
88
30
15
19
74
78
57
42
23
19
54
141
125
107
114
73
16
44
17
,49
28
28
14
26
29
33
4
21
1
19
3
1
13
16
7
7
0
6
11
9
4
7
0
1
6
53
33
48
40
16
49
20
40
6
-
23
15
27
13
7
2
30
10
24
5
19
21
5
2
9
9
14
7
2
2
6
2
9
6
0
196
158
155
154
89
65
64
57
55
51
43
41
39
36
35
34
31
25
24
22
22
18
18
16
16
14
13
13
11
10
9
9
7
6
(
I
t
I
.'
II
Table 6. A list of the species
and
taxa whoserelative
abundance was
greater
than 1%of the total catch in at least five samples. This filter
removed about two-thirds of the taxa collected during the period of
this study. Differences between frequency of occurrence in all samples
and frequency in only those samples in which relative abundance was
greater th~n 1%are compared in the table for the summerdata. The list
is compiled from all data collected from 1969-1972, a total of
141 summerand 58 winter samples.
<i
21
~
-l
'\j
Someof the steepest
gradients observed for
Centropages abdominalis were:
DATE
18
25
23
2
28
July
July
June
July
June
1969
1969
1970
1970
1971
NH 1
NH 3 . NH 5
NH 10
14
55
83
128
6
7
0
4
107
0
NH 1
NH 3
NH 5
249
655
1,370
1,655
102
0
0
3
80
0
and for Acartia clausii:
DATE
NH 10
18 July
23 June
1969
1970
528
25,661
8
58
35
19
2
12
21
5
1970
1971
1971
1972
19,807
2,640
762
297
100
127
0
***
47
16
0
8
-b
3
3
0
0
July
June
June
Aug.
o
The other three important copepod species
(Acartia longiremis3 Calanus marshallae3 and
Oithona similis) did not show such regular
patterns.
Acartia longiremis was always
more abundant at NH 1 in 1971 than at sta~
tions farther offshore, but in 1969 and 1970
it was usually more abundant offshore of
NH 1. Calanus occasionally had their greatest abundance at NH 1, but in all years it
usually did not. Oithona abundances fluctuated widely at all stations.
The other zooplankton taxa listed in
Table 6 had the following patterns of
abundance:
1. Highest abundance at NH 1, decreasing
offshore
decapod shrimp mysis
barnacle nauplii
and
2.
gastropod veligers
bivalve veligers
Usually abundant only
at NH 1, NH3
NH 5
Acartia tonsa
euphausiid nauplii
copepod nauplii
worms lacking parapodia
3.
NH1
Scyphomedusae
Chaetognaths
other fish eggs
echinoderm pluteus
Usually more abundant offshore of
Ctenophores
Pteropods
Oithona spinirostris
4. Abundance increases as distance
offshore increases
Metridia
22
lucens
5. Noapparent pattern
~
can be abundant
anywhere
Paracalanus parvus
Calanus nauplii
Calanus eggs
Oikopleura spp.
euphausiid eggs
The abundance patterns of these individual
taxa are discussed in greater detail in Part
II of this report.
These gradients deduced from absolute
abundance data give information about a population's response to the environment.
Nothing is learned about the complete zooplankton assemblages.
Since different gradient' patterns are exhibited by populations,
the zooplankton. assemblage changes between
stations.
Another way to look at gradient patterns
is with ranks of abundance.
Ranking results
in some understanding of how a population's
abundance changes relative to the abundance
of the total zooplankton assemblage..
Table 7a lists the average rank of
abundance of the most important taxa at each
station. We ranked only the filtered data,
i.e., an animal received a rank in a sample
only if it made up greater than one per cent
of the catch in that sample. Also given is
the mean rank over the four stations.
We
have listed only those taxa that occurred in
ten or more filtered samples summed over the
four stations.
A value was entered under a
station label only if the taxon occurred in
five or more filtered samples collected from
that station.
1
1
j
~
1
.,
,
f
J
1
~
1
.~
Among those copepods which had their
highest abundance
at NH 1, only Acartia
clausii had low ranks only at that station.
Centropages abdomina lis had a mean rank of
approximately 5 at NH 1, 3, and 5 and Pseudocalanus sp. had a mean rank of 1.5 at all
four stations.
For those copepods that were
usually more abundant offshore, Oithona
similis is seen to have its lowest ranks at
NH 1, 3, and 5. Acartia longiremis shows
evidence of having lowest ranks offshore.
There is little doubt that Calanus marshallae
prefers water offshore of NH 1.
Table 7b shows a similar result but is
here expressed 'in terms of an average relative abundance. Pseudocalanus sp. and
Acartia longiremis are clearly the two most
important copepods, on a numerical basis,
making up large percentages of the catch.
Acartia clausii is important only at NH 1 and
Calanus marshallae is again seen to be
t
~
j
)
1
!
important only offshore of NH 1.
Spatial
.
'1
Abu:ndanae Patterns
in WinteY'
The winter data are very different (Tables
8a and 8b). Immediately obvious from a
comparison to the summer data is the difference in rank order between the two seasons
for the endemic zooplankton species. Pseudoaalanus s~. was rank 1 during both seasons
as averaged over the four stations. During
the summer, AaaY'tia longiY'emis~ Calanus .
maY'shallae~ AaaY'tia alausii
and Oithona
simiUs
occupied
ranks 2 through 5. During
the winter, Oithona similis occupied rank 2,
and ranks 3, 4, and 6 were occupied by copepods with southern affinities: PaY'aaa"lanus
paY'vus~ Ctenoaalanus
ang Uaus.
.
vanus and COY'yaaeus
Few definite winter patterns are seen in
Tables 8a and 8b. Pseudoaalanus
and Oithona
similis have greater relative abundance offshore in contrast to the summer. Ctenoaalanus vanus and COY'yaaeus angliaus
had their
lowest average ranks nearshore at NH 1 and
NH 3.
Relative abundances of the important
summer copepod species change during winter
months. Pseudoaalanus sp., AaaY'tia longiY'emis and Calanus maY'shaUae make up a
smaller percentage of the total catch.
Oithona similis percentages increase significantly during the winter.
Abu:ndanae DiffeY'enaes
were more abundant in 1970. Maximum abundances for the three species in Figure 6
(Pseudoaalanussp., AaaY'tia alausii
and
CentY'opages abdominaUs)
occurred
in June
and July of all years. In addition, AaaY'tia
alausii
can have a second peak in September.
Differences between stations will be discussed later in this report, and are well il~
lustrated in Figure
-
6.
Figure 7 shows the abundances of Calanus
maY'shallae~ AaaY'tia longiY'emis
and Oithona
These are the copepods that were
similis.
usually more abundant offshore of NH 1.
Calanus maY'shallae was clearly most abundant
in 1969. Maxima occurred in August and
September
after .the Pseudoaalanus
maximum.
No peaks were seen in 1970. Peak heights
are greatly reduced in 1971 and are seen
only in June and July. AaaY'tia
longiY'emis
was abundant only in September of 1969 whereas in 1970 it was abundant from July through
September.
Oithona simiUs
was most abundant in June and July of 1969, September 1970
and sporadically in 1971.
The major differences between the two
years 1969 and 1970 are perhaps best illustrated by percentage of Pseudoaalanus
sp. in
the total catch. Average percentages are
shown below for the comparable periods during the two upwelling seasons:
Between SummeY's
One of the most intriguing problems in
the data set is the differences between
zooplankton assemblages sampled during 1969,
1970 and 1971.
Abundances
were so very
different between the seasons that one wonders whether any but the most general patterns can be described.
Elsewhere (Peterson
and Miller, 1975) we discussed some peculiarities in species composition and abundance associated with the 1971 data and related
our observations to anomalous winds
occurring during that summer.
Differences
between 1969 and 1970 were noted, but not
discussed.
Standing stocks were high during both of these upwelling seasons, but
abundances of certain copepod species were
very different between the two years.
Different species dominated the catch. Many
of the differences between the three years
can be seen in Tables 2 and 4 of this report.
Figure 6 shows the abundance data for
three copepod species which had their greatest abundance at the nearshore station.
Abundances of Pseudoaalanus
sp. and AaaY'tia
alausii
look as though they may be negatively correlated at station NH 1, both within
and between the upwelling seasons of 1969
and 1970. Pseudoaalanus sp. were clearly
more abundant in 1969.. Acar+ia clausii
22 June
23 June
- Sept.
1969
Sept.
1970
-
NHl
NH3
NH5
NH 10
68%
44%
67%
43%
58%
33%
54%
29%
Differences in relative abundance for the
other copepod speciescan be seen in Figures
6 and 7. As previously pointed out, AaaY'tia
alausii
and AaaY'tia longiY'emis were more
important during the 1970 upwelling
season.
Another way to illustrate differences
between the upwelling seasons is with an
analysis of the frequency with which various
species held different ranks of abundance.
Table 9 presents this analysis.
We show
only the data for the most important copepods. Abundances were ranked over comparable time intervals in 1969, 1970 and 1971:
from 22 June 1969, 23 June 1970 and 12 June
1971 through September of the three years.
During 1969, Pseudoaalanus
sp. were rank 1
(i.e., most abundant) in 7 of 9 samples from
NH 1, 8 of 9 samples from NH 3, 8 of 10
samples at NH 5 and 5 of 7 samples from
NH 10. During the same period in 1970,
Pseudoaalanus
ranks were quite different.
They were rank 1 in only 3 of 7 samples at
NH 1, 5 of 7 samples at NH 3, 3 of 8 samples
at NH 5 and 2 of 8 samples at NH 10.
By
contrast, AaaY'tia alausii
was rank 1 in 4 of
23
COPEPODSPECIES
PseudoaaZa:nus
Aaartia Zongipemis
GaZa:nusmarshaZZae
Aaartia aZausii
Oithona simiUs
GentPopages abdominaZis
Aaartia tonsa
ParaaaZanus pa1'VUS
MetPidia Zuaens
Oithona spini:r>ostpis
NH 1
NH 3
NH 5
NH10
1.69
3.48
5.41
3.03
4.55
4.80
1.42
3.71
3.68
4.67
4.71
5.57
5.80
1.46
2.81
3.43
3.81
4.70
5.50
1.55
2.80
3.37
4.83
5.63
6.40
7.29
SUM
MEAN
RANK
6.12
12.80
15.83
16.34
17.59
1. 53
3.20
3.96
4.09
4.40
20.88
22.86
5.22
5.72
6.28
4.79
6.88
OTHERTAXA
Limaaina heZiaina
OikopZeum sp.
Sagitta sp.
GaZa:nus eggs
GaZanus naup1i i
euphausiid eggs
euphausiid nauplii
barnacle nauplii
bivalve veligers
decapod mysis
Table 7a.
5.20
4.79
6.50
5.85
6.10
6.00
7.66
6.63
6.77
5.13
6.44
6.67
8.32
5.09
5.67
5.13
6.46
5.29
5.83
7.50
6.00
8.27
6.50
Average rank of abundanceat each stations sum of ranks and overall
rank for those taxa that were taken in five or more summer samples
at each station.
Rank 1 indicates the greatest abundance.
TAXA
NH 1
NH3
NH 5
NH10
Pseudoaa Za:nus s p.
Aaartia Zongipemis
GaZa:nusmarshaUae
Aaartia aZausii
Oithona simiUs
GentPopages abdomina Us
GaZa:nusnaupl i i
euphausiid eggs
51.6
11.3
3.3
19.5
6.3
4.1
7.5
2.6
46.8
18.3
10.7
9.6
6.4
3.4
7.1
9.1
45.0
18.7
11.3
12.0
7.0
3.2
4.2
8.8
43.4
34.6
10.5
4.5
7.4
3.0
3.8
8.6
Table 7b. Meanpercent-of-total-catch averagedover only those samples in which
the taxa made up greater than 1% of the catchs in five or more
samples at each station.
Only the more important taxa are listed.
24
I
COPEPODSPECIES
NH 1
NH 3
NH 5
Pseudoealanus
sp.
Oithona similis
Pa:raealanus pa:rvus
Ctenoealanus vanus
Aea:rtia longiremis
Coryeaeus anglieus
Calanus ma:rshallae
Clausoealanus areuieornis
Clausoealanus immatur
Clausoealanus pergens
Aea:rtia elausii
Aea:rtia tonsa
Centropages abdominalis
Calanus tenuieornis
Metridia
lueens
3.17
3.75
3.77
4.00
4.70
6.00
7.50
8.67
8.00
9.33
3.57
6.00
8.75
2.08
3.00
3.73
3.80
5.18
4.75
6.25
6.50
8.87
9.25
6.60
7.00
2.08
2.58
4.00
5.44
6.50
8.40
7.50
8.20
7.00
9.62
NH10
2.36
2.64
3.43
5.60
5.33
- 8.40
6.62
7.22
7.25
6.83
SUM
MEAN
RANK
9.69
11.97
14.92
18.84
21.71
27.55
27.87
30.59
31.12
35.03
2.42
2.99
3.73
4.71
5.42
6.88
6.96
7.64
7.78
8.75
25.21
6.30
9.66
5.66
.
OTHERTAXA
Oikopleura spp.
Chaetognaths
Pteropods
Calanus nauplii
barnacle nauplii
bivalve veligers
gastropod veligers
polychaetes
other worms
Table 8a.
5.88
7.42
6.33
8.66
5.33
6.60
5.85
7.66
6.33
5.20
6.00
8.25
10.33
8.66
5.90
9.40
10.00
8.33
5.66
6.60
7.10
7.70
8.75
8.66
10.80
Average rank of abundance at each station,
sum of ranks and overall
rank for those taxa that made up greater than 1% of the catch in
the winter samples.
TAXA
Pseudoealanus sp.
Oithona similis
Pa:raealanus pa:rvus
Ctenoealanus vanus
Aea:rtia longiremis
Coryeaeus anglieus
Calanus ma:rshaUae
Oikopleura
Calanus nauplii
Aea:rtia elausii
Table 8b. Meanpercent-of-total-catch
NH 1
NH 3
NH 5
NH10
23.9
13.9
16.0
3.4
10.8
12.6
4.0
5.1
8.8
21.8
29.7
18.0
12.2
10.2
9.2
18.5
2.7
8.1
6.0
15.0
31.0
13.9
14.9
10.3
5.0
6.3
9.1
5.5
3.3
29.8
18.3
13.5
10.5
8.8
3.0
3.4
3.9
***
***
***
averaged over only those samples which
the taxa made up greater than 1%of the catcht in five or more
samples at each station.
Only the more important taxa are listed.
25
N
0\
I5
w
5
L
Pseudocalanus
NH-IO
!ill:
.
10'0
1
L
!.
a
500
1'0
.
:.:
5
ci
LL
~ 1000
~
-
L..L
L..L.LJ.
1...& LI....
2.
~"
<t
~
LL
'"''
. r",'02
<t 15
5
WlLL
'0.
J
J1969AS.
NH-3
.
I
.
",,,n.
",
01.1
. J.
--
A
J
A
M
A
J1970
M
S
i
500
~
,
J
1971.
.
"
1S00
LL
,
lLl
Ul
500
,
LJ
'~III
'.,.,
"O~
1000
NH-I
L
clausii
NH- 10
L:s
z=> 1000
p
LL Lc
"
ISOO
NH-5
,.
:=
~
,
15
Acarlio
II
L'
I
r.
NH-
SOO
A
S
J
J
A
1969
S
LLJJ:I
M J J A
AMJJAS
1970
S
1971
I
NH-IO
200
100
L
mcmurrichi
fEr.
l
NH -5
UJ
~
300
<t200
c
2
::;) 100
III
<t
LR
".
ill
''to....
300
...
200 .
100
J
J A
1969
Fig. 6. Density of PseudoaaZanus sp.. AaaPtia aZausii~ and
S
A
M J
J
1970
A
S
M
J
J
A
1971
S
Cent:r>opagesmarrru:1':r>iahi
at NH1. NH3. NH5. and NH
10 during the summer upwelling seasons of 1969.
1970 and 1971.
'-
Acarlia
I 50 A
~
Calanu5
P.
I
51
I.
1000
NH-IO
P II.. P.
L"
500
LL.
l. II II
II
pP ,
p
Luhu LL-'
U
i
1500
II
~~
1000
oLJL~~
r:1
.
1
'"'
1
il~LL:
500
1500
::J
1500
1000
~
::1
~~~
J
300
200
100
J
A
1969
1
S
Oilhona
i
::11.
-
'
z 100~.
I
1111
I.,
.1
.00
0
200
lliJ
J
N
-..;j
J
A
1969
S
A
M
J
J
1970
A
S
J
A
I ,II J
S
AMJJAS
.1.
~
1
I
II
MJJAS
1970
,"-'
1971
NH-5
Lill
.
Uw
M
UL1 LL'
NH- 10
U
~;LL L
WJ
Z
300
J
1969
~
5imili5
L
J
1971
1970
lli
500
MJJAS
AMJJAS
P
~
'"'
I
U
Z 1000
::J
ro
<t 500
J
A
1971
S
N H-3
N H-I
Fig. 7. Density of Calanus marshallae,
Aaartia longiremis
and Oithona similis at NH 1, NH 3, NH5, and NH 10
during the summer upwelling seasons of 1969, 1970
and 1971.
7 samples at NH ~ as compared to a rank of 1
in only 1 of 8 samples in 1969. Ca~us
I1laX'shaZZae
is seen to have higl:J.est
ranks in
1969 at all stations. Aaartia Zongiremis
had its highest ranks in 1970.
Three other copepod species had different abundance patterns between the two years
of good upwelling, 1969 and 1970. ParaaaZanus parvus occurred commonly during the
months June-September 1969 but in 1970 it
was common only during the spring and fall
months.
During 1970, the population was
shifted offshore during late June, July and
early August.
The relative density of
Aaartia tonsa was greatest during the 1969
upwelling season (see Table 2). These two
species are the only coastal neritic species
with temperate-subtropical affinities that
occur off Oregon.
The third species,
Euaalanus
bungii
occurred in an equal number
of samples from the three upwelling seasons
but was much more abundant during the 1969
upwelling season. Mean density was 211m3 in
1969, 31m3 in 1970 and 91m3 in 1971. These
averages are based on 13, 10 and 15 samples
respectively.
EuaaZanus bungii is a subarctic oceanic species.
It always had its
greatest abundance at the offshore stations
and was taken at NH 1 only during the 1971
upwelling season.
The only other taxon that had a clear
pattern related to the 1969 upwelling season
was barnacle nauplii.
They occurred in only
a small fraction of the samples compared to
1970 or 1971.
The 1970 upwelling
season was the best
one for the copepods Centropages
abdomina lis
and Microcalanus
pusillus.
Although
average
density of Centropages
abdominaZis was about
the same in 1969 and 1970, the peak densities were much higher during the 1970 season
(see Figure 6 and the appendix data). Microcalanus pusillus
appeared much more frequently and was more abundant during the 1970
'upwelling season.
The strong dominance of Pseudocalanus
sp.' during
the 1969 upwelling season, the
persistence of ParaaaZanus parvus and Aaartia
tonsa through the summer, and the low abundance of the other two Acartia species in
1969 compared to 1970 when dominance was
shared by Pseudoca~us
sp. and the two
Acartia species, must somehow be related to
environmental ,factors. There were some
environmental differences between 1969 and
1970 but they were difficult to,evaluate.
The pattern of northerly winds is different
between the two years.
In 1969, northerly
winds did not begin to blow steadily until
the end of June. PseudocaZanus
sp. had
28
reached high numbers by this time.
In 1970,
northerly winds blew from late April through
August (see Figure 3). As a consequence of
this wind pattern, the Columbia River plume
was found nearshore at NH 1 through late June
1969 but was never found within 10 miles of
shore in 1970.
The extent to which this water of reduced salinity and high temperature may modify the planktonic environment is uncertain.
It is likely that different kinds of phyto~
plankton would be found in surface waters influenced by the plume compared to surface
waters influenced by active upwelling.
Different food resources may explain the occurrence of a different zooplankton assemblage
found during the late June-July period of
the 1969 upwelling season.
Perhaps Acartia
clausii could not utilize the food resource
resulting from phytoplankton growing in a
mixture of plume and coastal water in 1969
and as a result no population outbursts were
observed.
As the 1969 upwelling season progressed, Acartia clausii became important at
NH 1 in August. Acartia longiremis
became
important offshore, at NH 10, only after intense upwelling had developed.
The 1970 upwelling season began in April.
Intense upwelling was frequently present nearshore from
mid-May through September.
During this season, Acartia clausii was more abundant than
Pseudocalanus
sp. on most sampling dates
(see Figure 6), and was frequently dominant.
Beyond the data period, we have observed the
sp. dominance
1969 pattern of Pseudocalanus
with Acartia clausii in lesser abundance'
during early summer of 1973. The wind patterns in 1969 and 1973 were very similar.
Plume water was onshore throughout June, and
active upwelling did not begin until early
July. Acartia clausii was not abundant until
mid-August 1973.
Abundance Differences
Between Winters
Between winters, variability in species
composition was the most nqtable difference,
but there were differences in abundances as
well. These differences may be a function
of total northward transport of the Davidson
Current.
Unfortunately, estimates of the
transport are not available.
However, our zooplankton data, temperature-salinity diagrams, Newport wind data
and Bakun's upwelling index data all indicate that the winter of 1969-70 was an unusual one. Some of the southern zooplankton
species that had their greatest abundance
during this winter were Paracalanus parvus,
immature Clausocalanus,
Ctenoca~us
vanus,
Acartia tonsa and Corycaeus anglicus. Other
copepod species with southern affinities
NH 1 1969
2 3 4 5 6 7 8 9
23456
Calanus marshallae
Pseudocalanus
Acartia clausii
A. longiremis
1 1 1 2 1
7 1
1 3 1 1
1 1
NH 3
Calanus marshallae
Pseudocalanus
Acartia clausii
longiremis
A.
Calanus marshallae
Pseudocalanus
Acartia clausii
A. longiremis
*
#
1969
1
1 4 4
8 2
2
1 3
1 1 3 3
NH 10
Calanus marshallae
Pseudocalanus
Acartia c lausii
A. longiremis
1969
1 2 3 1
1
8
3 2
1 1 1
1
NH 5
NH1 1971
NH 1 1970
1969
2 2 3
5 2
1 1
1
2 2 1
1
3 3 1
4 2
1 1
NH3
123456789
1
1411
2
4
1 5
1
1 1 1
1
I
1 1 31 2
1
2
1
1
NH 5 1971
1 3
232
2
411
1
1
NH 10 1971
NH 10 1970
2 4 2
1
5 2 1
1
321
3
1
1
2 4
= Pseudocalanus was rank 11 in one sample and Acartia clausii was rank 10 in one sample.
= Acartia clausii was rank 12 in one sample.
Table 9.. Frequency distribution
of ranks of abundance of the four most abundant copepod species found in the
Oregon upwelling zone. Ranks 1~6 are shown for the 1969 upwelling season and Ranks 1~9 for 1970 and 1971.
Temporal variations are seen reading across the table and spatial variations down the table.
The table is
interpreted as follows. for example: Reading across the table. Pseudocalanus
was Rank1 in 7 samplesand
Rank 2 in 1 sample from NH 1 in 1969; Rank 1 in 3 samples, Rank 2 in 3 samples and Rank 3 in 1 sample from
NH 1 in 1970. etc.
N
\0
*
*
5
1
2 2
3 3 1
1
3 2 1 1
NH 5 1970
3 1 1 1 1 1
3 4 1
1
2 1
3 2 3
11
NH3 1971
1970
1
2 1 1 1
5 2
2
2 1 1 1
1
2 2 2
2
#
appeared in samples with
much
greater fre-
quencyduringthe winterof 1969-70. They
were: Mecynocera clausii, Calocalanu8 styliremis, Calocalanus tenuis, Clausocalanus..
mastigophorus, Clausocalanus parapergens,
Acartia danae and Corycaeus amazonicus.
In
addition, four other southern species were
seen only during this winter: Clausocalanus
jobei, Clausocalanus paululus,
Oncaea dentipes,
and Oncaea subtiUs.
All of the
above 16 copepods are indicators. of water
originating south of at least Cape Mendocino,
California (Olsen, 1949; Fleminger, 1967;
Frost and Fleminger, 1968).
Seasonal Cycle of Total Zooplankton Abundance
A diatom bloom was in progress at this time
as well. Our nets were clogged with the
diatom Thalassiosira at stations NH 1, 3,
and 5. The 16 February 1971 peak h~d high
numbers of Pseudocalanus
sp. ~680/m
= 12%
of the catch), Calanus (240/m
15%) and
Calanus nauplii (192/m3
12%). The Pseudocalanus
sp. population was 50% adult females
and 50% immatures.
The Calanus
population
was almost entirely stage I copepodites.
Female Calanus
were found farther offshore.
These facts again indicate an actively reproducing adult copepod population.
In both
of these years, abundances decreased after
the February peak to lower values in March
=
or
Apri
=
1.
Given the variability between individual
summers and winters, generalizations about
the annual cycle of zooplankton abundance
are difficult to make. The problem is compounded by our lack of understanding of the
timing of upwelling events and other environmental factors, and the response of zooplankton production to such events.
In 1972, no samples were collected in
Januaty of February. The 15 March sample
sp. (1844/
had high numbersof Pseudocalanus
m3 = 62%), Oithona simiUs
(69Ojm3 = 23%)
The annual cycles for stations NH 1, 3, 5,
and 10 are shown in Figure 8, 9 and 10 respectively.
The simplest temporal pattern
is the annual cycle. Abundances are high
during the upwelling season and low during
the winter months.
All four stations have
this basic pattern.
There are important
differences between the cycles at the four
stations. Variations on the basic pattern
seem to be related to distance from shore.
At the nearshore stations (NH I and NH 3),
the winter low extends from mid-November
through January, while at the offshore stations, the low extends at least through February.- Peak amplitudes are also a function
of distance from shore. Station NH 1 had
The months of April and May are periods
of transition in ~he field of the wind.
The
atmospheric high pressure cell begins to
form over the North Pacific Ocean and the
winds begin blowing from the north with
greater frequency.
These months are also
months of transition for the zooplankton.
Copepod species with southern affinities are
generally missing by Mayor
are taken in
greatly reduced numbers.
In all years of
this study, thick phytoplankton blooms were
seen during this period.
The blooms are
probably associated with the replenishment
of nutrients within the photic zone by upwelling. The blooms were found on 27 April
1970, 3 and 14 May 1971, 20 April 1972 and
22 May 1972.
Zooplankton abundances were
low at these times.
16 sampling3dates with abundances greater
than 5000/m.
Median abundances also show
the same pattern and are listed below:
STATION
NH
NH
NH
NH
1
3
5
10
SUMMER
4350/m3
2250
1550
1000
WINTER
850/m3
800
530
365
The annual cycle at NH 1 has the most
detail.
The winter low has a duration of
approximately two and one half months, from
mid-November through January of all years.
A peak in zooplankton was seen during all
three years in either February or March.
The 25 February 1970 sample had high numbers
of copepod
nauplii
(1840/m3
= 27% of the
catch).
This indicates the presence of an
actively reproducing adult copepodpopulation.
30
and Acartia
longiremis
(265/m3
= 9%). Half
of the total catch were immature Pseudocalanus sp~ and half of the Acartia longiremis
were immatures.
By
29 May
reached
through
late May and June (4 June 1970 and
1971) zooplankton abundances had
5000/m3.
Numbers remained high
June and July of all years, reaching
maximum numbers o£ 44,000/m3 on 18 July 196~
and 51,372/m3 on 23 June 1970.
The peak in
1971 occurred on 21 July and WaS only 9577/m3.
No large peak was seen in our 1972 samples.
Abundances were lower in August, as compared to June and July, during all years of
this study. Abundances continued to decline
through September and October of 1969 and of
1971.
In 1970,
a large peak was seen on 25
September, 9 October and 20 October. Acartia
clausii was the single dominant, making up
50% of the catch in September and 83% and
61% on the October dates.
It)
105
8-1969
8--1970
1971
:E
......
-
.-+- 1972
0
z
W
I
I
I
104
U
Z
I \
0
z
:::> 103
m
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\i""'::"
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(!)
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t
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I \'
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i'
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II
!\~-
'
\1
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102
J
M
F
A
M
J
J
s
A
0
N
D
M 0 N TH S
Fig. 8. Annua1 cyc1e of tota1ed zoop1ankton abundance at NH1. during the three
year study period.
It)
~
NH 3
......
0
-w
z
104~
.-+-I~(~
U
'\
Z
0
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m
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0102
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10'
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t .,.\.,;:;-.4
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;
F
J-
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.
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J
AV
V
M
A
M
J
J
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"'"
\"'1/'0,.
A
s
0
N
\.
,
D
M 0 N THS
Fig. 9. Annua1cyc1e of tota1ed zoop1anktonabundance
at NH3, during the three
year study period.
31
8-1969
.-1970
1971
8---1972
104
NH 5
~
!! i\!.
".........
~
~
"
,..,.."
103
z
/
'-
-....
7'
..
/
/
.
.
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'
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"x.
r
' ...or
...",,'JO.
0
/;:0
\.-",-~ \ .
.
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.../
w
U
Z
~
..(:Jt.1
.
-,
,
V!
":'
:..
.f
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'11
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f
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102
F
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A
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D
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8.--
m
«
8-+-
(!)
10
1969
1970
1971
1972
103
0
.-J
102
J
F
M
A
M
J
J
A
5
0
MONTHS
Fig.
32
10.
Annual cyc1e~ of totaled zooplankton
during the three year study period.
abundance at NH 5 and NH 10
N
D
Our data indicate that the transitional
period between an upwelling system and one
dominated by the northward flowing Davidson
Current extends at least through October.
By December of all years copepod species
with southern affinities were well represented and abundant off Newport.
The annual cycle at NH 3 is similar to
the cycle at NH 1, but amplitudes are lower.
The pattern of low abundance in December
and January is present.
The February peak
was seen only in 1970. The low abundances
extended through February in 1971 and 1972.
The late spring-early summer peak that began to develop in June at NH 1 did not develop at NH 3 until July of 1969 and 1970.
The 1971 peaks are strikingly reduced, and
are probably a result of reduced coastal upwelling during that summer. The September
and October peak seen in 1970 at NH 1 is
also seen at NH 3. On 25 September and 9
October, the NH 3 peak was associated with
Pseudocalanus
sp. dominance (53% and 45%
respectively) rather than dominance by
Acartia clausii as at NH 1 on the same date.
The 20 October peak at NH 3 was dominated
by Acartia clausii (60%).
At NH 5 the annual cycle is different.
Amplitudes are greatly reduced as compared
to NH 1 and NH 3, and winter-spring peaks
do not develop until April and May.
There
seems to be a recurrent summer abundance
pattern:
peaks are seen through July and
mid-August, and a low is present from midAugust through September of all years. An
autumn peak developed in September and
October of 1969 and 1970. The September
peak was dominated by Acartia clausii and
the October peak by Pseudocalanus sp. in
1970, and by Pseudocalanus sp. and Calanus
marshallae in early and mid-September 1969,
by Aeartia longiremis on 28 September 1969,
and by winter species in late October 1969
(Paraealanus parvus~ Oithona similis and
Oikopleura spp.).
The annual cycle at NH 10 is similar to
that at NH 5, in that the late winter peak
seen at the nearshore stations is not present and the spring peak does not begin to
develop until May. Two peaks are present
during the summer months, in July-August
and September-October.
August lows are
seen at NH 10. Abundances were never
greater than 5000/m3 at NH 10.
SEASONAL CYCLE OF RELATIVE SPECIES ABUNDANCE
The seasonal cycle of the relative abundance of the abundant (and therefore important) copepod species is shown in Figure 11
for NH 1, NH 3, NH 5 and NH 10. The figures
are cumulative percentage
of the total catch
sp., Aeartia
elausii~ Centropages abdominalis~ Aeartia
longiremis~ Calanus marshallae adults, immatures and nauplii pooled, Paraealanus
parvus and Oithona simiU8.
The most general result contained in these figures was
summarized in Tables 7b and 8b.
for the copepods Pseudoealanus
The figures are rather complex but deserve careful study because some interesting patterns are present.
The simplest pattern is the sinusoidal annual cycle. The
same pattern was seen in the plots of total
zooplankton abundance.
One can conclude
from a comparison of the zooplankton abundance plots (Figures 8, 9, and 10) and the
relative species abundance plots (Figure 11)
that low abundances during winter months are
coincident with a decrease in relative abundance of the endemic copepod species, and an
increase in importance of non-copepod taxa.
Oikopleura
spp, chaetognaths and warm-water
copepod species become important during the
winter.
There is considerable seasonality in the
relative abundance of each taxa. At NH 1,
Pseudoealanus sp. were important through
September 1969, April-September 1970, February-July 1971 and late September-November
1971. Aeartia elausii and A. longiremis
were always important during the autumn
after the cessation of upwelling.
Centropages abdominalis was never important after
August, with the exception of November 1971.
Calanus mar8hallae copepodites and nauplii
had their greatest importance during the
spring. Paraealanus parvus and Oithona
similis appear most importantly during the
winter.
There are large differences between years
at NH 1 as previously noted. Pseudoealanus
sp. had a much higher relative abundance
during the 1969 and 1971 upwelling seasons.
During the 1970 season, Aeartia elausii and
Pseudoealanus sp. shared dominance in many
samples. Centropages abdominalis were less
important during the 1971 upwelling season.
Aeartia longiremis were about equally important at various times during all three
upwelling seasons. Oithona similis was
more important during the 1969 and 1971 upwelling seasons. Paraealanus parvus were
important over broader time intervals in
1969 and 1970 as compared to 1971.
At NH 3, the most striking aspect of the
annual cycle compared to NH 1 is the greatly
decreased importance of Acartia elausii and
increased importance of Aeartia longiremis
and Calanus marshallae.
Acartia elausii
made up a large fraction of the catch only
33
during October 1970.
Acartia longiremiB
were important over
Calanus marshallae
and
broader intervals in 1970 and 1971 at NH 3.
The annual cycle of PseudocalanuB
sp.
relative abundance at NH 3 was about the
same as for NH 1 except for two periods:
July of 1970 and 1971. During both times
Pseudocawnus
sp. was important at NH 1,
whereas Acartia longiremis
was important at
NH 3.
The NH 5 plot is similar to that for NH 3,
particuarly between November 1969 and May
1970, and between January and July 1971.
Similarly to NH 3, the importance of Acartia
clauBii is greatly reduced and the importance
Calanus marshallae~
of Acartia longiremis~
and Oithona similis
are increased, relative
to the stations closer to shore.
The NH 10 plot follows the NH 5 plot
closely during 1970 and 1971 with one exception:
in September 1970 Acartia clausii was
important at NH 5 but not NH 10.
SUMMARY
OF PART
I
The zooplankton community is different
between summer and winter partly as a result
of major changes in nearshore circulation
patterns.
In summer when the net water
transport is to the south, species with
northern affinities dominate.
In winter
when transport is northward, species with
southern affinities are mixed with the endemic fauna. Abundances are about an order
of magnitude higher in summer than winter
presumably because of coastal upwelling.
We found year-to-year variations during
both summer and winter in zooplankton abundance and species composition.
During the
summer upwelling season, the observed "differences can be explained by differences in
upwelling strength resulting from differences in the pattern of the wind stress during each of the three summers. During the
winter months, the differences in species
composition were probably a result of either
differences in transport of the Davidson
Current or else in the position of the current relative to the nearshore zone. Both
cases are a function of the pattern of the
winds.
On a spatial scale, zooplankton were
consistently most abundant at the station
one mile from shore. Persons interested in
coastal zone management should be interested
in this result because man's activities that
could have an impact on zooplankton are most
likely to be most intense very close to
shore.
35
part II: remarks on
important taxa
In this section taxa will be discussed
separately.
Seasonal cycles of abundance
will be presented and distribution patterns
will be related to environmental variables
wherever possible.
Life cycle information
and sex ratios will be discu~sed for the
copepod species.
Emphasis will be given to
the 34 taxa listed in Table 6. Also included
in this section will be observations on
phytoplankton blooms, large numbers of
medusae, and other miscellaneous occurrences.
COPEPODSPECIES
Pseudocalanus
sp.
The importance of Pseudocalanus
sp. in the
zooplankton assemblages occurring off Newport
was shown in Tables 2, 7, and 9 above.
It is
the one copepod that frequently had a rank of
abundance of 1 or 2 in samples collected from
all four stations (see Table 9). . When
Pseudocalanus
sp. was not rank 1 it was
usually replaced by an Acartia species.
High
abundances of Pseudocalanus sp. frequently
occur on days of active upwelling, and it is
on those days that the dominance of the zooplankton by this species is most extreme.
The contingency of abundance and dominance
upon upwelling conditions was tested in the
following manner.
Abundance data were divided into high and low categories by ranking
the 34 samples collected at NH 1 between the
dates 1 May and 30 September.
Ranks 1-17
were called high abundance and ranks 18-34
were called low abundance.
Dominance data
were divided according to whether Pseudocalanus
sp. was more or less than SO per cent
of the total assemblage.
The criterion for
active upwelling on a particular date was
surface salinity 33.5% or greater.
If salinity data were not available for a date, then
the wind data (Fig. 4) were consulted.
If
northerly winds preceded the sampling date py
a few days, upwelling was assumed to have
been active.
37
The contingency tables are as follows:
Pseudoealanus
sp. abundance at NH 1
High
Active
Inactive
X2
= 5.79,
Pseudoealanus
4
5
13
P
= 0.02
sp. dominance at NH 1
>50%
<50%
11
5
6
13
term episodes of
the upwelling process.
At NH 3, no relationship between abundance
and upwelling was expected because, as noted
earlier in this report, abundances are fre~
quently low there during active upwelling,
when strong abundance gradients are seen between N1k1 and offshore stations.
Abundances
may be reduced due to upwelling circulation
patterns.
NH 3 is an area of divergence, so
low zooplankton abundances are not surpris~
ing. Note however, the relationship between
dominance and upwelling was checked.
The
contingency table was:
Low
12
short
Pseudoealanus
Active
Inactive
>50%
<50%
10
6
6
11
Active
X2
= 3.43, P = 0.07
The conclusion is that Pseudoealanus
Inactive
sp.
abundance tends to be high at NH 1 on days of
active upwelling, and that on those days it
tends to be more strongly dominant. The six
sp. was strongly
dates on which Pseudoealanus
dominant despite weak upwelling were primarily in May and June:
22 June 1969, 14 May
1971, 28 June 1971, 22 May 1972, 28 June
1972, and 30 August 1969. It is possible
that dominance by this species during the
spring period is related to the timing of
the spring phytoplankton bloom, and not to
upwelling.
Hints like these from the animal
data make it clear that better information is
needed on the annual cycle of plant abundance in the Oregon coastal zone. The five
dates on which Pseudoealanus
sp. was not
dominant, but when active upwelling was in
progress, were 4 June, 23 June, 16 July
and 11 September 1970 and 21 July 1972.
Aeartia elausii was rank 1 and Pseudoealanus
sp. was rank 2 in three of the samples (4
June, 23 June and 11 September 1970).
Pseudoealanus were rank 1, but not quite
50 per cent of the plankton in the other
two samples:
46 per cent on 16 July 1970
and 44 per cent on 21 July 1972. Rank 2 was
occupied by Aeartia
longiremis
on both of
these dates.
= 1.47,
X2
p>0.10
There is no strong relationship between
sp. dominance and upwelling at
NH 3. There is, however, some suggestion of
a pattern. Pseudoealanus
sp. was dominant
in spite of no upwelling on 22 June and 29
June 1969, 3 May 1971 and 22 May 1972, and
26 August 1969 and 25 September 1970. The
first four dates are in spring months.
The
opposite relationship of no dominance during
upwelling had the following dates:
23 June,
2 July, 16 July 1970, 6 July and 21 July
1971, and 21 July 1972. These are all early
summer months. Pseudoealanus
sp. was rank 1
but not dominant in two of the samples (23
Pseudoealanus
June
1970
=
44 per
cent
37 per cent).
Aeartia
1 in the
four
other
and
2 July
longiremis
1970
=
was rank
samples.
At NH5, there were only 10 sampling dates
with active upwelling, as compared to 16
dates at NH 3 and 15 dates at NH 1. No relationship between Pseudoealanus sp. dominance
and active upwelling is seen in the contingency table.
But once again, a pattern can
be recognized in the opposite cells.
Pseudoaalanus
The analysis used here is based on the
untrue assumption that Pseudoealanus
abundance and dominance are independently determined on the different dates of the study.
Thus significance can arise if Pseudoealanus
is dominant and abundant
thToughout one year
of strong upwelling and less abundant
in
other years.
In fact, it is certain that
the observed relationship is at least partly
a result of differences between the years.
We also think, however, that Pseudoealanus
is probably concentrated near shore by the
38
dominance
>50%
dominance
<50%
Active
3
7
Inactive
9
12
X2
= 0.086,
p>0.75
Dates when Pseudoealanus
sp. were
dominant when upwelling was weak or nonexistent were from April through July (22 June,
10 July, 18 July and 6 August 1969, 22 May,
4 June and 2 July
1970, 14 May 1971, and 20
April and 21 July 1972). Dates when upwell~
ing was strong but Pseudocalanus sp. were
rank 1 in four of the samples but were not
dominant:
26 August 1969
31 per cent;
3 September 1969 = 40 per cent; 11 September
1970 = 47 per cent; and 5 August 1972 = 42
per cent. The other three dates were 14
September
1969, Calanus
= 39 per cent; 13
August 1970, Acartia longiremis = 32 per
cent; and 21 July 1971, Acartia longiremis
-
38 peT
NH 5
NH 10
2
4
3
2
3
5
5
10
8
6
5
7
4
2
2
4
3
2
1
1
2
1
3
2
5
1
9-10
5
2
5
1
2
2
3
>10
5
3
5
5
<1
i
3-4
4-5
LL
-....; 5-6
.
6-7
7-8
IX
8-9
i
r;
cent.
NH 3
";;;- 1-2
;!!. 2-3
=
=
NH 1
en
CLI
.....
VI
Totals
4
20
24
17
14
16
5
5
5
4
1
18
Active upwelling as indicated by surface
salinities in excess of 33.5% was never
seen at NH 10. The upwelling front always
intersected the sea surface inshore of this
station.
Virtually
no information on the life
sp. can be gleaned
from the data of the Early Life History
Program.
The mesh size of the plankton nets
used was too large to capture young copepod~
ite stages, nauplii, or eggs. Copepodite
stages I and IT were captured only occasion~
ally. Stage III were often seen, but were
probably not sampled quantitatively.
There~
fore, it was not possible to follow peaks of
eggs and nauplii through successive stages
to adult in order to estimate generation
history of Pseudocalanus
time.
Sex ratiodata for adultPseudocalanus
sp.
indicate a preponderance of females. Uneven
sex ratios favoring females are characteris~
tic of members of the copepod section Amphascandria.
Adult males do not feed, so
they suffer mortality soon after molting.
However, there were four samples in which
males were more abundant than females.
All
four samples were from 1969, the year of
greatest Pseudocalanus sp. abundance: 6
August and 30 August at NH 1 and 30 August
~nd 3 September at NH 3. The frequency
distribution of sex ratios is shown below.
The dispersion of observed sex ratio is
greatest at NH 1 and least at NH 10. We
have no explanation for this slight trend.
012345
m
ffi
SEX RATIO (FEMALES/MALES)
m
~~f~
~
~
Sex ratios of immature Pseudocalanus sp.
were also determined.
This was possible because immature males have fifth legs whereas
females do not. The data shown below are
pooled copepodite stages IV and V. The range
of ratios is much less compared to the data
on adultPseudocalanus
sp. About 80 per cent
of the data were sex ratios between 1:1 and
1:2 males: females.
Since most of data fell
within this interval, we have expanded it to
intervals of 0.25 over the ratio range of
1:1.00 -- 1:1.99.
In all cases N>40.
Ratio
NH 1
NH3
<1. 0
1. 00-1. 24
1. 25-1. 49
1. 50-1. 74
1. 75-1. 99
2-2.99
33.99
4-4.99
4
11
5
5
2
5
6
5
7
2
1
NH 5 NH 10
5
5
9
3
3
2
1
3
5
10
1
0
5
L
17
27
29
16
7
7
1
1
Acartia clausii
Acartia clausi has its greatest abundance
at NH 1 and was most abundant during the
swnmer of 1970. Highest numbers were seen
in June of 1970, 1971 and 1972.
It is possible that the beginning of our sampling program on 22 June 1969 missed a 1969 June peak.
A September peak developed in 1969, 1970,
and, according to data of L. Smith (in
Holton and Elliott, 1973), in 1972 as well.
It is only at this time that Acartia clausi
can be found in large numbers offshore of
NH 1. This autumn period corresponds to a
time when the winds are light and variable,
fluctuating between northerly and southerly
directions.
No peak was seen in September
1971. August and September 1971 were both
months of anomalously low numbers of all zooplankton (see Figures 8-10). These low numbers probably resulted from an effect on the
nearshore environment of onshore transport
of warm, low salinity water.
39
There is no statistical
relationship
between Acartia cZausi abundance
and the
short term intensity of upwelling at NH 1,
The contingency table (see PseudocaZanus sp,
discussion above for the method) was:
abundance
Acartia cZausi
High
Low
7
'9
Active upwelling
8
Inactive upwelling
10
= 0,12
X2
~,~
~
0
z
-- ILIIO'
';;
<)
. z
;; «
C>
.. Z
::>
I
2
~
1D'0'
'", «
The precipitous
drop in A, clausi
abundances between 2 July and 16 July 1970
is not yet understood. The wind had been
steady from the north from 2 through 14 July,
On 15 July the wind switched around to the
southwest.
Our sampling followed that switch
by only one day,
It is doubtful that this
brief shift in wind direction could have been
responsible for the fifteenfold reduction in
A, cZausi abundance,
Abundances of CaZanus,
PseudocaZanus, and Oithona were unchanged between the two dates. Acartia Zongiremis increased threefold and Centropages abdominaZis
decreased threefold over the 2 July to 16
July interval.
The extreme reduction in A,
cZausi may have been caused by some highly
specific predation or other biotic process,
A preliminary study of stomach contents of
juevenile smelt (AZZosmerus eZongat~) ~eing
carried out by one of us (WTP) is showing
that these fish selectively eat Acartia
cZausii when that copepod is abundant.
These
fish are of 5.5 cm fork length. They were
collected 22 August 1974. We will pursue
this preliminary observation further.
The sex ratio data are shown below.
'0
DAYS
Fig, 12,
.0
'0
AFTER
120
100
27 APRI L
are
1970
Abundance of Acartia dZausii at
NH 1 during the 1970 upwelling
season over the six month period
beginning on 27 April,
Abundance
data are plotted against time
in Figure 12 for the 1970 upwelling season.
Two peaksare seen: in June and September.
Peaks of males, females, and immatures are
synchronous.
It seems likely that eggs,
which were responsible for these peaks,
were laid over very short intervals, probably less than one week. Generation times
might be estimated to be about 60-70 days.
The time interval between 27 April 1970,
when 45 individuals/m3 were present to 23
June 1970 when about 25,000/m3 were present
is about 60 days,
40
~
in
sex
1;'atios, between
There
upweU.,..,
ing seasons, During the 1969 season, males
were more abundant than females in 12 of 19
samples.
During the 1970 season, males were
more abundant in only 5 of 28 samples.
This
agrees with the PseudocaZanus
data where we
found more males than females only in 1969.
Heinle (1970) found that sex ratio in
Acartia was a function of population density,
At low densities, females were more abundant.
We find here the opposite relationship:
In
the year that Acartia cZausi was a dominant
organism, sex ratios were usually between
1:1 and 2:1::females:males.
V)
UJ
<
::IE:
"V)
UJ
.;..J
There are no published data on fecundity
of Acartia cZausi. Acartia tonsa has been
shown to produce 1200-1300 eggs in a 40 day
adult life span (Johnson, 1974; Wilson and
Parrish, 1971). Fecundity of A, cZausi is
likely to be of the same general order,
If
there were no mortality this would be sufficient to account for the entire increase
within one generation,
However. any more
probable mortality rate requires that much
of the population increase must have come
from overlap df several generations,
It is
likely, therefore, that the generation time
is much less than 60 days.
differences
<1
1-2
2-3
3-4
<
4-5
UJ
I.L
5-6
6-7
::IE:
'-'
-
0
....
.
><
UJ
V)
1969
1970
1971
1972
12
1
1
1
5
13
8
2
2
3
3
4
1
1
5
2
7-8
8-9
9:.10
10-1
11-1
,
1
1
Centropages abdominaZis
There was an onshore-offshore gradient in
standing stocks of Centropages abdominaZis,
Abundances
were always highest at NH 1.
over rather narrow limits compared to
sp. or Acartia clausii.
The
highest abundances and median abundances
listed below are remarkably similar at each
station:
There was also a gradient in frequency of
occurrence in the samples:
over all samples
collected 1969 through 1972, this copepod
occurred in 34 of 35 samples at NH 1, decreasing to 27 of 34 at NH 3, to 27 of 35
at NH 5, and to a low of 22 of 33 at NH 10.
Pseudocalanus
STATION
NH 1
NH 3
NH 5
NH 10
In our data (listed in the appendix), the
population peaked on 25 July 1969, 2 July
1970, and 12 June 1971. There was an autumn
peak in two of the three years:
29 October
1969 and 6 November 1971. Typically, no
C. abdominalis
are found during the winter
months frqm December through March.
Notable
exceptions were seen at NH 1 on 25 February
1970 and 16 February 1971. Both of these
dates have been shown to be unusual in that
zooplankton production was seen. Pseudocalanus sp. and Calanus marshallae both
showed peaks on these two dates. This matter was discussed briefly on page 30 of this
report.
)
I
Absolute abundances were different on a
year-to-year basis.
The upwelling season of
1970 had highest abundances, followed by
1969 and 1971. Relative abundances in 1969
and 1970 however, were similar. At NH 1,
from 22 June through September 1969, C. abdomina lis made up 3.4 per cent of the catch.
From 23 June through September 1970, it made
up 4.2 per cent of the catch.
In 1969,
C. abdominalis had an average rank of 3.0,
but in 1970, its mean rank dropped to 4.2.
It was displaced one rank in 1970 because
Acartia clausii was more abundant during
that year.
MAXIMUM
MEDIAN
1477/m
1618/m3
1776/m3
1142/m
108/m
159/m3
163/m3
262/m
Also, some of our data suggest that
Acartia longiremis abundance changes very
little between different upwelling seasons.
Total relative.densities (i.e., relative
density summed from each station) listed in
Table 2 varied little between the three
upwelling seasons.
Table entries for 1969
were 1078/m3, for 1970 were l509/m3, and
for 1971 were l33l/m3.
There is no direct relationship between
active upwelling and abundance of Acartia
longiremis.
Contingency tables were set up
in the same manner as described for Pseudocalanus
sp., and are shown below for stations NH 1, 3 and 5:
Acartia longiremis abundance
<.!J
z:
......
NH 1
NH 3
NH 5
High Low
High Low
High Low
8
;-H
::::
Active
I..LI
3
Sex ratios were variable.
Males were
more abundant than females in one half of
the 1969 samples. Males were more abundant
in 73 per cent of the 1970 samples, but only
33 per cent of the 1971 samples. The data
are:
RATIO FEMALES:MALES
1969
1970
1971
<1
1: 1
12
24
6
4
4
4
Inactive
9
8
13
14
None of the tables is significant.
We
conclude that the abundances of this species
are not affected by short term variations
in coastal upwelling.
Further support for a hypothesis that
upwelling differences have little effect on
>1
numbers of A. longiremis is ~iven below.
The average abundance (no./m ) at each station during each upwelling season is not
very different between stations,and seasons.
8
5
8
NH 1
Acartia
9
NH 3
NH 5
294
491
238
208
380
385
281
134
NH 10
longiremis
Two general abundance patterns have
already been noted in this report for this
species.
First, Acartia longiremis has its
highest ranks at the offshore stations, and
secondly, this species was abundant longer
and more extensively during the 1970 upwelling season than in other years.
Abundance of Acartia
longiremis
varies
1969
1970
1971
1972
97/m3
273
593
237
307
441
218
284
On a date-by-date
basis, there was no
consistent pattern in the"abundance of" Acartia
longiremis.
In 1970, numbers were generally high between May and December.
In
both 1971 and 1972, high abundances were
41
seen only from June through mid-August.
near shore and the older copepodites
There was no September peak in either 1971
or 1972.
offshore,
The trend is clearly shown in
Table 10. This table lists the abundance of
eggs, nauplii, and copepodite stages at the
four study stations, averaged over 33 dates
when data were available for all stations on
a date. The abundance data have also been
converted to percentages of the total abundance of each category over the four stations.
The importance of Acartia longiremis
within the zooplankton assemblage can only be
seen by examining its relative abundance.
Table 7b, page 24, shows that the average
relative abundance of this copepod changes
from 11 per cent of the catch at NH 1, to
18 per cent at NH 3 and NH 5, to 34 per cent
of the catch at NH 10. Clearly, this species is increasingly important offshore.
Sex ratio data for Acartia longiremis are
similar to data for other copepods: ratios
are variable from year to year. The data
are listed below:
-
V>
LLI
...J
<1
.......
1:1
<
V>
LLI
1-2
...J
2-3
3-4
"'-"
0
.....
.....
4-5
<
LLI
I.J...
1969
1970
1971
1972
14
3
8
3
23
1
15
1
7
4
5
3
2
1
3
2
7
10
5
1
41
30
20
3
5-6
6-7
>7
><
Lo.I
V>
TOTALS 33
During 1970 when A. longiremis abundances
were highest, sex ratios were about equal.
Males dominated in 56 per cent of the samples and females dominated in 42 per cent.
There were no ratios in excess of four females per male. The range in female:male
ratios was narrower in 1972, but females
were more abundant than males in all but 20
per cent of the samples.
In 1969 and 1971
the range of ratios was broad. There was
one ratio in 1969 of ten females per male.
In 1971, the seven ratios in excess of six
to one were 11, 12, 13, 13, 19, 21, and
28:1 females per male.
We do not yet have
-any ideas about what relationship sex ratio
has to environmental factors in this species.
.Calanus marshallae
Calanus
marshallae
is the only copepod
for which we have adequate life history
data. Since it is a much larger copepod
than either Pseudocalanus
sp. or the Acartia
species, all copepodite stages were retained by our plankton nets.
The onshore-offshore distribution of eggs,
nauplii and copepodites was interesting because the younger stages tended to be found
42
Eggs, nauplii and stage I and II
copepodites were most abundant at stations
NH 1 and NH 3. Stage III copepodites were
abundant at NH 1, 3, and 5. Stage IV copepodites were most abundant at NH 3 and 5
and Stage 1['s at NH 3, 5, and 10. Adults
were most abundant at NH 3 and 5 only. This
observed distribution of stages is probably
linked to coastal water circulation patterns.
The vertical distribution of the stages must
be known before we can understand these observations.
Another problem apparent in Table 10 is
the difference in abundance of each stage.
Natural and predation mortality should lead
to decreased abundance of each successive
copepodite stage with time, if the time spent
in each stage is the same and if predation
is not age specific.
However, stages IV and
V are much more abundant than stages I-III.
This cannot be explained by differential
escapement through net meshes because our
.24mm mesh net was small enough to retain
even the youngest copepodite stage. A likely
explanation is that stages IV and V are
longer lived than stages I-III.
Because of the onshore-offshore gradient
in distribution of life cycle stages, it is
difficult to objectively follow development
through time utilizing data from one or
several stations.
But we have learned something about generation times simply by looking at abundances of eggs or nauplii with
time because they appear in pulses.
The
beginning of a generation is indicated by
the presence of large numbers of eggs, nauplii or stage I copepodites.
This occurs at
regular intervals and several generations
per year are indicated.
In both 1970 and
1971, generations began in mid-February,
late April, in June, and in August.
A fifth
generation began in October 1971. Less than
12 months of data are available for 1969 and
1972. In 1969, generations appear to have
begun in late May, mid-July and late October.
In 1972, the first generation began in February, the second in April but the third
generation did not appear until August.
These observations
are in agreement with
those of Marshall and Orr (1955)
from the
NH1
EGGS
NAUPUI
STAGEI
STAGEII
STAGEII I
STAGEIV
STAGEV
43.4
14.0
8.8
14.6
33.9
24.6
1.7
17.6
9.7
2.7
CJ 0.6
Q 3.2
cJ
31.8
74.5
51.5
33.1
25.5
10.1
3.3
44.9
16.0
29.8
28.6
30.3
35.3
29.5
29.4
40.0
10.0
<;(
7.2
Abundances (no.1m3) of CaZanus~ averaged
from February to October for 1970 and 1971
are shown below:
NH 1
NH3
NH 5
NH 10
1970
1971
75.1
151.6
131.3
74.8
49.3
133.1
94.9
121.2
Little difference in abundance is seen
between these two years. CaZanus marshaZZae
and Acartia Zongiremis have similar abundance patterns.
Both are usually more abundant offshore of NH 1 and neither shows much
year-to-year variation in numbers.
Sex ratios of adult CaZanus are shown
below.
Females are usually more abundant
than males.
Males can be more abundant than
females at times preceding spermatophore
transfer because stage V CaZanus males molt
the
HB 10
8.6
15.7
5.2
5.8
13.7
36.6
30.1
2.4
16.6
3.7
10.1
3.6
6.0
7.5
15.8
26.1
1.1
6.6
16.2
5.8
11. 1
18.9
28.6
38.1
36.0
40.9
37.7
7.1
3.7
7.6
19.5
15.6
16.5
31. 3
19.7
15.0
Abundance of Calanus marshalZae eggs, nauplii and copepodite stages
at the four sampling stations.
Data were averaged over 33 dates when
samples were successful~y obtained from all stations on a date. The
upper table is numberlm and the lower table is percentage of a
category at each station.
Eggs, nauplii, and younger copepodites are
found nearer to shore than the older copepodites.
English Channel and Clyde Sea area.
Generation times there are about two months.
The first generation begins in mid-February,
the second in April-May, and the third in
July. Usually no more than three generations are seen. There are four off Oregon
in most years.
before
NH 5
23.7
16.8
202.2
24.1
10.2
12.3
EGGS
NAUPUI
STAGEI
STAGEII
STAGEII I
STAGEIV
STAGEV
Table 10.
NH 3
females
(Marshall
and
Orr,
-
V1
I.LI
...J
1969
1970
V1 <1
1971
1972
2
2
1-2
1
2
I.LI
...J
2
2-3
I.LI 3-4
t::. 4-5
3
1
6
4
1
1
2
3
2
11
4
0
5-6
1
4
1
22
8
7
6
......
I- >6: 1
c::c
0::
X
I.LI
V1
Males have a regular pattern of
occurrence.
They usually were found only
between about February and July in 1970,
1971, and 1972. In 1969, males persisted
between July and September.
When males are present in the samples,
females are typically present also. There
is a slight tendency for males to be found
without females at NH 1. It is not statistically significant.
Occurrence between the
months of February and October at each of
the four stations is shown below:
.
,
1955).
43
-
---
--
-
-
--
Station
NH 1
Number
of
NH 3
NH 5
NH 10
48
49
51
47
Number
containing
females
19
38
45
36
Number
containing
males
8
20
27
21
Number
of
joint
occurrences
4
Dates -"
Sampled
19
26
20
Of the zooplankton
occurring
off Oregon,
Oithona simiZis
was one of the few taxa
important
in both summer and winter samples.
Its mean rank and mean percentage
of the
total
catch during summer and winter are
shown below.
It was relatively
more important during the winter.
AVERAGE RANK OF ABUNDANCE
NH 1
NH3
NH5
4.6
3.8
4.7
3.0
4.7
2.6
AVERAGE PERCENT-OF-TOTAL
Summer
Winter
NH 10
5.6
CATCH
NH 1
NH 3
NH5
NH 10
6.3
13.9
6.4
18.0
7.0
13.9
7.4
18.3
NH 1
114.3
53.6
138.0
NH 3
128.2
106.2
107.9
NH 5
NH10
78.0
48.3
91.2
75.4
66.9
79.1
We tried to determine if O. simiZis
abundance was related to sea surface temperature, or if abundance had some temporal
pattern.
All of the quantitative
estimates
of the abundance of O. simiZis (including
44
the
This copepod is a broadly neritic
temperate-subtropical
form that occurs at
least
as far south as Baja California
and as
far north as the Queen Charlotte
Islands
(Fleminger,
1964; Cameron, 1957).
Off
Oregon it is most abundant during the winter,
but frequently
occurs during the summer upwelling
season.
Winter and summer densities
(no. 1m3) are shown below:
Winter
NH 1
NH3
NH 5
1969-70
1970-71
1971-72
123.3
139.2
144.4
117.0
67.3
74.4
Summer
NH1
NH3
NH5
NH 10
1969
1970
1971
23.9
31.1
4.4
26.4
55. 1
6.3
13.9
37.7
3.4
13.7
22.7
2.7
170.2
33.6
84.8
NH10
160.4
37.6
47.8
2.6
In addition, b. simiZis is another taxon
which was about equally
abundant during all
three upwelling
seasons.
No large differences are seen in the table of mean relative
density
(no. 1m3) shown below:
1969
1970
1971
were plotted against
surface
temperature
at the station.
(fig.
13).
The only clear conclusion
is that the higher
abundances
(greater
than 200/m3) mostly occur
at surface
temperatures
greater
than 10oC.
The best explanation
is that this species
is
not abundant in the very deep layers
which
are moved up and ashore during the strongest
upwelling
events.
These events also result
in the lowest temperatures
observed
in the
sampling area.
There was no temporal
pattern
in the data.
Dates from all months of the
year, except April,
were represented
in the
20 dates with highest
abundance.
:ParacaZanus parvus
Pi thona simi lis
Summer
Winter
zero estimates)
ParacaZanus was abundant over the entire
study area during the winter of 1969-70, but
during the other two winters,
it had its
greatest
average abundance nearshore.
ParacaZanus was not abundant
(and occurred
infrequently)
during the 1971 upwelling
season.
There were also abundance differences
between the 1969 and 1970 seasons that are not
apparent
in the above averaged densities.
ParacaZanus was common throughout
the June
to September period in 1969.
In 1970, it
was common only in the spring and fall,
and
the population
was shifted
offshore
during
late June, July and early August.
600
t689 (8) t2309(8)
.
500
"
'~
Q)
+'Q)
E
400
()
..........
.
300
.
.
.
.
(f)
E
(f)
. .
200
0
C
0
-'=
+-
0
"
.
...0
::J
U
.
.
.
100
.
.
.
.
.
"
.
,
"
.
.
.
." .
" ..
.
.
"
.
.
.
.
10
Temperature
Fig. 13.
"
.. .
.
". "" " "
.
.
...II'
.
.
. .
8
.
. ." " . ..
. . .. ...
..
..
.
.
- °C
Scatter diagram of Oithona similis abundance vs. sea surface temperature.
Caption
Ctenoaalanus
symbols are
8 = NH 1.
. = NH 3. 'If' = NH 5. and"
vanus
Ctenoaalanus vanus abundance patterns are
similar to Paraaalanu8:
Highest abundances
are seen during the winter months.
Cteno-
= NH 10.
aalanus, like Paraaalanus, were most abundant during the winter of 1969-70. particularly between 29 December 1969 and 29
January 1970. Average densities
(no./m3)
are shown below for the three winters:
, 45
1969-70
1970-71
1971-72
NH1
NH3
NH 5
NH 10
37.3
21. 9
15.8
42.5
20.3
13.2
61.3
22.5
19.5
47.9
24.1
17.4
Abundance and occurrence
are usually
low
during the summers.
During the summer of
1969, Ctenoaalanus
occurred
in only 5 of 33
samples and averaged
1.6/m3.
During the 1970
upwelling
season,
it was taken ln 7 of 44
samples.
Five of the seven samples were in
the months of April and May before the onset
of active upwelling.
Average abundance was
7.8/m3.
During the weak upwelling
season of
1971, Ctenoaalanus
occurred
in all months
from May through September in 16 of 40
samples with an average density
of 2.8/m3.
In 1972, it also occurred
in a large number
of samples (10 of 21 samples) and averaged
2.9/m3.
Of the 38 summer occurrences,
Ctenoaalanus
occurred
at NH 1 in only four
samples.
abundant over that summeras a whole.
OithDna spinirostris
There were no large differences between
summer and winter average abundances (no.1m3)
of Oithona spinirostris, nor were there
any obvious differences between statlons.
Abundances were highest during the 1971 upwelling season, like those of Metridia
luaens.
Summer
NH 1
NH3
NH5
NH 10
1969
1970
1971
1972
5. 1
3.8
19.6
1.4
7.0
4.8
6.1
4.2
4.8
4.4
15.6
2.8
3.0
3.0
14.2
5.2
4.7
0.4
6.8
6.6
0.0
1.9
1.9
2.4
7.3
2.1
4.3
2.6
Winter
1969-70
1970-71
1971-72
Metridia luaens
Metridia luaens was one of the small
number of taxa that increased
in abundance
as distance
offshore
increased,
at least
to 10 miles offshore.
The abundance gradient was evident during both summer and
winter as shown by the table of average
densities
(no. 1m3) below:
Summer
1969
1970
1971
1972
Mean
NH 1
NH3
NH 5
NH 10
3.3
3.4
2.1
0.0
2.9
4.2
4.3
6.3
9.9
6.2
4.7
2.6
11.8
9.9
7.3
9.2
5.9
28.4
29.6
18.3
Frequency of occurrence was also similar
to Metridia luaens. In all summer samples,
Oithona spinirostris occurred in only seven
samples at NH 1, but in 22 from NH 3, in 25
from NH 5, and in 24 from NH 10.
These similarities in occurrence between
Metridia and Oithona led us to look at joint
occurrences of these two species. Using an
index of affinity (Fager & McGowan, 1963), we
found that the two are related and that the
degree to which they occur together increases
offshore.
The data are shown below. Values
of the index greater than .500 were taken to
be indicative of significant association.
NH 1
NH 3
NH 5
NH 10
8
20
19
26
7
22
25
24
5
15
17
23
.479
.608
.680
.819
Winter
Occurrences
1969-70
1970-71
1971-72
Mean
2.3
1.3
2.4
2.0
4.4
1.0
3.2
2.9
1.1
4.1
19.3
8.2
9.8
36.5
15.9
20.7
There
was a similar gradient in frequency
of occurrence.
It occurred in only eight
summer samples at NH 1, but in 21 at NH 3,
in 24 at NH 5, and in 31 at NH 10. In the
winter samples, the gradient was 6, 11, 11,
and 12 at NH 1, NH 3, NH 5, NH 10 respectively.
Metridia luaens was much,more abundant
during 1971. This fact lends support to the
hypothesis that water which usually lies
farther offshore was advected onshore during
this weak upwelling season. High 'average
abundances in 1972 were a result of high
numbers in April and June only; it was not
46
of Metridia
luaens
Occurrences
of Oithona
spinirostris
Frequency of
Joint
Occurrences
Index of
Affinity
This result is expected since both are increasingly frequent and abundant at the offshore stations.
I
1
9lausoealanus
areuieornis
This copepod occurs most frequently and is
most abundant during the winter.
It first
appeared in the nearshore zone in the month
of December in all years of this study.
In
1970 and 1972 it persisted until April.
In
1969 it persisted until May.
In 1971, the
year of weak summer upwelling, it was found
in May, June, August and September.
qlausoealnus
and (2) as occurring off the coasts of
California and Baja California (Acartia
tonsa, Paracalanus
parvus,
Temora discaudata,
Clausocalanus
farrani
Type 1, and Candacia
curta), only Acartia tonsa and Paracalanus
parvus
are found off Oregon.
In our summer
data, both of these copepods were abundant
only in 1969 and 1970. During the winter,
both species were abundant at all stations
in 1969-70, and abundances were reduced during the other two winters.
pergens
Calanus tenuicornis
The abundance pattern of Clausocalanus
pergens was similar to that of C. areuieornis.
It was important only during the winter
months, being advected to the Oregon coast
by the Davidson Current.
It did not appear
in the plankton until December in 1969 and in
1970.
It was taken through mid-May of 1970,
1971 and 1972. After the month of May, it
either did not occur (1970) or else was found
only at NH 10 (1971 and 1972). In 1969,
Clausocalanus pergens was found nearshore in
June and July.
This observation appears to
be exceptional.
I!-eartia
This copepod never made up more than 1 per
cent of the catch in any summer sample.
It
only occurred in one sample in 1969, and four
samples in 1970. By contrast, it was taken
frequently during the 1971 upwelling season,
occurring in 14 of 40 samples.
The copepod
Ctenocalanus
vanus had the same pattern of
greater occurrence in 1971. Abundance of
C. tenuicornis was not great in winter either
but it did make up more than 1 per cent of
the catch in nine of 37 samples in which it
occurred.
The densities (no.1m3) are shown
below:
tonsa
The distribution and abundance of Aeartia
tons a were peculiar because its abundance declined and it disappeared from the study
area during the three-year sam~ling period.
Mean relative densities (no.1m ) are tabulated below for summer and winter:
Summer
NH 1
NH3
NH 5
NH 10
1969
1970
1971
1972
44.8
10.5
0.0
0.0
36.0
7.7
0.0
0.0
41. 0
3.3
0.0
0.0
8.9
6. 1
0.0
2.0
Winter
NH 1
NH3
NH5
NH10
69-70
70-71
71-72
38.4
2..7
2.7
46.6
2.1
1.0
22.7
1.6
3.0
14.2
2.1
0.7
Abundances were about constant between the
summer of 1969 and the following winter, and
between the summer of 1970 and the following
winter.
Abundances decreased between these
two periods.
No Aeartia tonsa were taken
during the summer of 1971 and none occurred
after April 1972. We conclude that the decline is not clearly related to a seasonal
cycle.
There are similarities between the
distribution and abundance of Acartia tonsa
and Paracalanus parvus.
Of the five copepods listedby Fleminger(1967) as (1)
coastal-neritic temperate-subtropical species
69-70
70-71
71-72
NH 1
NH3
NH 5
NH 10
0.6
1.2
2.3
8.5
0.8
3.7
4.5
1.2
5.2
1.5
4.7
3.1
Corycaeus anglieus
This cyclopoid copepod was abundant only
during the winter of 1969-70.
It appeared
first in September, was most abundant in
December and January, and disappeared by June
in two years.
It did not appear in any
samples collected during the summer of 1971
or 1972, nor during the winter of 1971-72.
Other copepods that occurred during the first
two winters but not during the winter of
1971- 72 were Meeynoeera
clausii,
Clausocala-
nus parapergens,
Tortanus discaudatus,
tenella,
and Coryeaeus amazonicus.
OTHER HOLOPLANKTONIC
Euphausiid
Oneaea
TAXA AND MEROPLANKTON
Developmental
Stages
Euphausiid eggs and nauplii were counted
in our samples, but we cannot be certain if
they are those of Euphausia paeifica or
Thysanoessa
spinifera.
The latter species is
a coastal-neritic form (Brinton, 1963), and
it is likely that most of the eggs and nauplii are of that species.
Furcilia and juveniles in our samples were Thysaneoessa
spinifera.
In samples collected on the ex-
tended transects
pacifica
of 1971 and 1972, Euphausia
adults or juveniles did not occur
47
1969
1970
1971
APRIL
JUNE
JULY
AUGUST
SEPTEMBER
OCTOBER
JULY
V1
<!J
<!J
I..LJ
MAY
JUNE
JULY
AUGUST
APRIL
1972
APRIL
MAY
JUNE
/
AUGUST
APRIL
MAY
......
......
...J
0...
JULY
JULY
AUGUST
SEPTEMBER
OCTOBER
:::>
~
z:
AUGUST
SEPTEMBER
APRIL
V1
......
0...
0
JUNE
JULY
JULY
I0...
>...J
SEPTEMBER
OCTOBER
~
U
~
......
...J
......
U
c:::
:::>
LL
JULY
AUGUST
SEPTEMBER
JULY
JUNE
JULY
MAY
JUNE
SEPTEMBER
NOVEMBER
V1
I..LJV1
...J I...J
z: :::>
1..LJe::!
>~,
:::>
"J+
JUNE
JULY
JUNE
SEPTEMBER
,
Table 11.
48
The months of the year when euphausiid developmental stages were
present in densities greater than 101m3. '
less than 15 miles from shore. The months
in which eggs, nauplii, calyptopis, furcilia
and juveniles were abundant are shown in
Table 11. There are temporal patterns in
the table, but they are different in different years.
This makes the patterns hard to
evaluate.
Two spawning periods are seen in
1970, 1971 and 1972. These are in April-May
and in July.
Either Thysanoessa
spinifera
spawns twice yearly, or we are seeing the
eggs and nauplii of T. spinifera at one.
time and E. pacifica at the other.
We could
find no published data on the spawning times
of T. spinifera~ but since it is'a coastalneritic species adapted to an upwelling environment, it is reasonable that it would
spawn in the spring so that actively feeding
furcilia and juveniles are present in July
during the most active upwelling.
Spring or
late winter spawning is the rule for most
crabs (Lough, 1974), for pink shrimp
PandaZus jordani Rathbun (Rothlisberg,
1975), and for most fish (Sally Richardson,
personal communication).
E. pacifica
is
known to spawn off Oregon in July (Smiles
and Pearcy, 1971). Even though adult E.
pacifica are found only offshore of 15
.
miles, their eggs could be spawned at depth
there and be carried into the nearshore zone
by the shoreward-moving deep water.
This
hypothesis can be checked if nauplii andlor
calyptopis can be identified.
We did not
try to do this.
The frequency of abundance estimates for
Thysanoessa spinifera within various intervals, and the average abundance of each developmental stage are tabulated below.
abundant very close to shore than E.
pacifica is in the adjacent zone to seaward.
CZadocerans
Evadne and Podon were both seen
frequently, but they seldom occurred together.
No taxonomic study was done on
these animals, and it was assumed that the
specific names in common use at OSU are correct: Evadne nordmanni
and Podon Zeukartii.
Evadne
PeZagic MoZZusks
The pelagic mollusks are represented off
Oregon by Limacina heUcina
(Type B,
McGowan, 1963) and CZione Zimacina.
C.
Zimacina appeared in only three samples
from the nearshore, on 29 May 1971 at NH 10
and 23 September 1971 at NH 5 and NH 10.
They were found offshore on 12 June 1971 at
NH 30, 35, 40 and 50, on 21 July 1971 at
NH 40. Limacina
heZicina
occurredin over
half of our samples from the nearshore zone.
It was one of several zooplankton species
that was most abundant and occurred most
frequently during the weak upwelling season
of 1971 (Peterson and Miller, 1975).
Mean relative density
percentage
This table is provided as a summary of
the mean abundance and ranges of abundances
because the data for each stage are listed
in the appendix on separate pages and are
not easily compared.
These data can be
compared to previous data on Euphausia
pacifica off Oregon.
Smiles and Pearcy
(1971) found about 1 individual/m3 with maximum values of 27 and 141m3. They sampled
at stations from 15 to 65 miles from shore
off Newport.
Hebard (1966) found 21m3 at
NH 15 and 91m3 at NH 25 with a maximum of
501m3. Thus
spinifera appears from our
data to be about an order of magnitude more
were abundant in July and late
August 1969, and June, July and SeptemberOctober of 1970. They occurred in only two
samples in 1971 in June and did not occur
in 1972. Podon were abundant only in the
autumn on 29 October 1969, September-October
1970 and November 1971.
frequency
(no./m3) and
of occurrence
or-L.
heZicina are listed on the following page
for summer and winter:
T.
ABUNDANCE
INTERVALS- NO./m3
>1000
Eggs
Nauplii
Calyptopis
Furcil i a
Juveniles
Adults
}
500-1000
100-500
50-100
3
3
12
3
13
5
2
10-50
<10
20
17
7
10
8
30
37
55
57
93.7
8.4
4.1
5.4
14
15.4
MEANDENSITV
49
Sununer
1969
NH 1
NH3
NH5
NH10
Frequency
1970
7.1
5.8
8.3
3.4
59%
7.1
2.1
1.0
33%
1971
26.5
.10.5
5.8
17.8
88%
1972
40.0
261.3
139.8
108.5
95%
Winter
1969-70
1970-71
1971-72
NH 1
NH3
NH5
NH10
30.1
16.5
10.5
8.9
32.3
20.8
31.9
3.0
3.0
1.6
4.9
4.7
There is a suggestion in the data that
chaetognaths were more abundant during the.
two summers of good upwelling, 1969 and
1970~ than durin~ 1971. The mean relative
densities (no.1m ) at the four stations are
shown for summer and winter.
There is very
little difference between average summer
and winter abundances.
SUMMERAVERAGED ABUNDANCE
1969
1970
1971
1972
During the summers of 1969, 1970, and 1971
heZiaina occurred most frequently offshore of NH 3. During the summer of 1972
they occurred frequently at all stations.
and
June
were
the
months
when
L.
heZiaina were most abundant during all four
years of our study. . In 1971, they were
abundant in August as well. Phenomenally
large numbers were found on 28 June 1972 at
NH 3 and NH 5. Abundances were l85/m3 and
l12/m3 respectively.
On 30 May 1972 several
samples were collected that qualitatively
appeared to be almost entirely Limaaina
heZiaina, at a station ten miles off Cascade
Head (latitude 45° 16' N).
Large numbers
were also encountered once outside the
May-June period on 20 October 1970.
Chaetognaths
A
number
of species
of chaetognaths
occur off Oregon, but they were not differentiated in our counts. During the summer,
Sagitta eZegans predominates, but Eukrohnia
hamata also occurs. During the winter
months, both Sagitta eZegans and Sagitta
euneritiaa are important.
Other species
found close to shore off Oregon during the
winter and offshore during the summer are
Sagitta
sarippsii,
S. bierii and S. minima.
Chaetognaths occurred in 10'7of 141
summer samples and 52 of 58 winter samples.
They made up more than 1 per cent of the
total catch in only 14 summer samples, but
in nearly half (27 of 58) of the winter
samples.
There was no relationship between
chaetognath abundance and upwelling.
Of the
". 34 samples with abundances greater than
,101m3 during the upwelling season, there
w~re 17 from days of relaxed upwelling and
IT,from days of active upwelling.
Seven of
the":;?4samples were from NH 1, 17 from NH 3,
8 frim;l.oNH
5 and only 2 from NH 10.
50
NH3
NH 5
NH 10
43.5
6.5
8.3
6.9
25.6
30.8
13.3
2.5
15.3
8.6
7.5
5.0
4.4
1.7
2.5
5.3
WINTERAVERAGEDABUNDANCE
L.
May
NH 1
1969-70
1970-71
1971-72
NH 1
NH 3
NH 5
NH 10
17.3
1.9
7.1
15.8
25.5
5.6
20.5
11. 0
7.1
9.3
9.0
2.6
OikopZeura
OikopZeura were more abundant during the
fall, winter and spring months and were most
abundant during the winter of 1969-70.
Mean
relative density (no.1m3) at the four stations during the summer and winter sampling
periods are shown below:
Sununer
NH 1
NH 3
NH 5
NH 10
1969
1970
1971 .
1972
18 1
10.1
18.7
10.6
35.1
55.3
18.0
0.0
12.5
15.0
14.2
7.0
3.5
5.3
15.6
8.3
Winter
NH 1
NH 3
NH 5
NH 10
226.8
40.4
21. 1
85.8
33.8
15.9
39.6
12.3
11.9
1969-70
1970-71
1971-72
199.7
14.7
26.7
Presence of OikopZeura in samples
collected during the upwelling season was
strongly correlated with. surface salinity.
OikopZeura occurred in 53 of 130 summer
samples from 1969-1972.
We have surface
salinity data from 47 of the 53 samples.
Presence and absence in the 47 samples were
compared to surface salinities greater than
and less than 33.0% and 33.5% with contingency tables (2 x 2). The tables and respective chi-square C 2) values are shown
below.
Both tables are highly significant,
indicating an association between presence
of OikopZeura and lowered surface salinity.
*
PRESENT
~:?
<33.0
<as::
4- .,....
5';j >33.0
V) V)
ABSENT
PRESENT
8
20
<33.5
9
63
>33.5
x2 = 36.8
p<O.Ol
ABSENT
~
3f34
x2 = 16.0
p<O.Ol
These data indicate that Oikopleura is an
offshore form found only in waters of reduced salinity and suggest that it may be
particularly abundant in the Columbia
River plume.
Oikopleura
was seldom present
in active upwelling areas (salinities
>33.5 0/00).
Ctenophores
Ctenophores occurred infrequently during
the 1969 and 1970 upwelling seasons. The
average abundance was 21m3 in seven samples
in 1969 and 0.8/m3 in five samples in 1970.
Ctenophores were never taken at NH 1 during
1969 and 1970. For the winter of 1969-70,
mean abundance was 1.81m3 in eight samples;
for 1970-71, 1.61m3 in eight samples, and
for the winter of 1971-72, 2.6/m3 in nine
samples.
Greatest abundances occurred during the 1971 and 1972 upwelling seasons.
Average abundance was 8.7/m3 in 19 samples
from 1971 and 7.6/m3 in 11 samples from
1972. Ctenophores were abundant only during
May and June. This agrees with observations made by Nielsen (1937) in the North
Sea and off Iceland. Ctenophores were not
taken in the autumn months, except during
1971. Hirota (1974), in work off La Jolla,
California, found ctenophores to be most
abundant in the autumn months.
Medusae
In some of our samples, medusae were
quite numerous.
The dates when they made up
a large fraction of the total sample settled
volume are listed in Table 12. We have no
estimates of absolute abundance of large
medusae because all medusae with diameters
greater than about 2 cm were removed from
the samples before subsampling.
The conclusions that can be drawn from Table 12
are that medusae were never abundant in
1969, and were most frequently abundant in
1970. They were important only in June of
1971 and 1972.
Decapod Shrimp Mysis
i
j;
These meroplankton appeared in 74 samples
collected during the summer months.
The
frequency distribution of abundance is shown
below:
ABUNDANCE INTERVALS (NO./MJ)
>u
z:
L.1.J
>40
30-40
20-30
10-20
1-10
<1
10
1
3
14
35
11
CT
L.1.J
c:::
u..
The highest abundance was 620/m3. This
category made up 1 per cent of the total
catch in only 11 summer samples.
During the winter months, abundances are
greatly reduced.
Decapod mysis were seen in
28 samples but they made up less than 1 per
cent of the total catch in all but two
samples.
The two estimates were 471m3 on
16 February 1971 at NH 1 and 71m3 on 30
March 1971 at NH 1.
Barnacle Nauplii
Barnacle nauplii appeared in 76 summer
samples and 29 winter samples.
Abundances
were usually low. More than half of the
abundance estimates were numbers less than
10 nauplii/m3.
Frequency of abundances
within seven abundance intervals are shown
below:
> 1000 1m3
>100
50-100
20-50
10-20
1-10
<1
Summer
Winter
1
9
8
9
8
32
9
4
2
5
2
10
6
The maximum abundance was 1105/m3 at NH 1
on 12 June 1971. Barnacle nauplii were virtually absent in all samples collected in
December and January.
They were seen on all
but two summer dates at either NH 1 or NH 3
during the upwelling seasons 1970-72. Those
dates were 23 August 1970 and 21 July 1971.
By contrast, there were many dates during
the 1969 upwelling season when nauplii were
not taken at any station:
25 July, 6 August,
26 August, 30 August, 14 September, and 28
September 1969.
There is a suggestion in the data that
highest abundance occurs during relaxed upwelling.
In 1969 northerly winds blew nearly
continuously from 29 June to 16 August, were
calm for six days, then blew steadily from
the north until mid-September.
Barnacle
nauplii were not found in any samples during
51
Table 12.
DATE
STATION(NH)
23
16
13
9
20
4
5, 10
10
1, 3
3
10
3, 5, 10
JUNE 1970
JULY
AUGUST
OCTOBER
OCTOBER
NOVEMBER
12 JUNE 1971
28 JUNE
6 JULY
5
3
1, 3
28 JUNE 1972
5
Dates when medusaecomprised at least 100 ml of the sample settled
this extended event. In 1970 both dates
having zero barnacle nauplii were at the end
of extended upwelling events. High numbers
on 25 February 1970, 25 September and 20
October 1970, and the three 1971 dates were
associated with calm or southerly winds.
Of the other five dates, 4 June, 2 July and
11 September 1970 were preceded by about a
week of calm or southerly winds. Nearshore
waters could have easily been warmed during
these intervals. High abundances on 16
July 1970 and 5 August 1972 were not in
agreement with this hypothesis.
Both were
during active upwelling and were preceded
by many days of active upwelling.
Bivalve
Veligers
No attempt was made to differentiate this
taxon into species.
It represents the lar~
vae of both clams and mussels.
They were
found in 107 of 138 summer samples and 54 of
58 winter samples.
Peak numbers were seen
once in the summer and once in the autumn.
No dramatic peaks were seen during the
winter. More than half of the abundance
estimates were numbers less than 10/m3. The
frequency distribution of abundances is
shown below:
Summer
>100/m3
40-100
30-40
20-30
10-20
1-10
<1
52
8
10
7
7
8
51
14
Winter
3
2
3
2
7
28
9
volume.
The highest
summer abundance was 769/m3 on
11 September 1970 at NH 3, and the highest
winter abundance was 2ll/m3 on 6 November
1971 at NH 1. Bivalve veligers had high
ranks of abundance in five samples.
They
were rank 4 on 30 August 1969 and 11 September 1970 at NH 1, rank 3 on 11 September
1970 at NH 3, and 23 September 1971 at NH 1,
and rank 1 on 11 October 1971.
Gastropod Veligers
This is another broad category.
It
represents a large number of gastropod mollusk species which occurred in 88 summer
samples and nearly all winter samples (52
of 58).
Abundances were greater than
60/m3 on only two dates:
(227/m3)
25 Se~tember 1970
and 2 July 1970 (173/m).
Usually
abundances were less than 101m3. The frequency distribution of abundances is shown
below:
Summer
2
0
8
10
53
15
>100/ m3
60-100
30-60
10-30
1-10
<1
There
is
little
Winter
3
7
33
9
evidence of pattern in the
abundance data. This is no doubt a result of
lumping the veligers of a large number of
gastropod species into this category. Highest
abundances in 1969 were seen in July and
September; in 1970 in February, June and
September; in 1971 in May, June, October and
November, and in 1972 in August.
Echinoderm
Larvae
Given the great abundance of echinoderms
along the Oregon coast (sea urchins, star~
fish~ sea cucumbers, crinoids and brittle
stars), we were surprised that their larvae
seldom appeared in our plankton samples.
Auricularia occurred in three summer samples
and no winter samples.
Bipinnaria occurred
in two winter samples but no summer samples.
Plutei occurred in 11 of 58 winter samples
and 16 of 117 summer samples. They were
most abundant during 1971 on 12 June and 19
August.
Their rare occurrence indicates
that elther the pelagic larvae spend little
time in the plankton, or the larvae are not
dispersed more than one mile from the shore.
53
100
ACART/A CLAUS//
IICENTROPAGES
50
IHHHA.
LONG/REM/S
NH-I
0
CALANUS.
NAUPL//
100
III
PARACALANUS
50
O/THONA
-~
L
~
~
\.>
(;j
q:
NH-3
0
100
50
NH-5
0
100
50
Fig. 11. Seasonal cycle of relaNH-IO
0
J
A
/969
S
0
N
D
M
A
M
J
J
/970
A
0
N
D
M
A
M
J
J
/97/
A
0
N
D
F
M
A
M
J
/972
J
A
S
0
N
tive abundance (% of total catch) of the most
abundant copepod species
at NH 1. NH 3. NH 5. and
NH 10 over the three year
study period.
bibliography
Bakun, A. 1973. Coastal upwelling indices
west coast of North America, 1946-71.
U.S. Dept. Commer., NOAA Tech. Rep.
NMFS SSRF-671, 103p.
Bourke, R.H. 1972. A study of the seasonal
variation in temperature and salinity
along the Oregon-Northern California
coast. Ph.D. Thesis, Oregon State Univ.,
Corvallis, 107p.
Brinton, E. 1962. Variable factors
affecting the apparent range and estimated
concentration of euphausiids in the North
Pacific.
Pac. Sci. 16:374-408.
Burt, W.V., and B. Wyatt.
1964. Drift
bottle observations of the Davidson
Current off Oregon.
In K. Yoshida
(editor), Studies on Oceanography, p.
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Cameron, F.E.
1957. Some factors
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165-202.
Collins, C.A., C.N.K. Mooers, M.R. Stevenson,
R.C. Smith, and J.G. Pattullo.
1968.
Direct current measurements in the frontal
zone of a coastal upwelling region.
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Cross, F.A.
1964. Seasonal and
geographical distribution of pelagic
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Fager, E.W., and J.A. McGowan.
1963.
Zooplankton species groups in the North
Pacific.
Science 140:453~460.
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Ocean. Fish. Invest., Atlas no. 7,213p.
55
Frost, B. and A. F1eminger. 1968. A
McGowan, J.A.
1963. Geographical variation
in Limacina helicina in the north Pacific.
Speciation in the Sea. Systematics Assn.
Pub. No.5,
Brit. Mus. Natl. Hist.,
London, 109-128.
revision of the genus Clausocalanus
(Copepoda:Calanoida) with remarks on distributional patterns in diagnostic characters.
Bull. Scripps Inst. Oceanogr.
12:1-235.
Miller, C.B. and W.T. Peterson.
1974. Yearto-year variations in coastal zooplankton
community structure in the upwelling zone
off Oregon. Invited paper, AAAS meeting,
1 March 1974, San Francisco, California.
Hebard, J.F.
1966. Distribution of
Euphausiacea and Copepoda off Oregon in
relation to oceanographic conditions.
Ph.D. Thesis, Oregon State Univ.,
Corvallis, 85p.
Heinle, D.R. 1970. Population dynamics of
calanoid copepods.
Helgolander wiss.
Meeresunters 20:360-372.
Myers, A.H. 1975. Vertical distribution
of zooplankton in the Oregon coastal zone
during an upwelling event. M.S. Thesis,
Oregon State Univ., Corvallis, 60p.
Hirota, J.
1974. Quantitative natural
history of Pleurobpachia bachei in La
Jolla Bight. Fish. Bull. U.S. 72:295335.
Nielsen, E. Steeman.
1937. On the relation
between the quantities of phytoplankton
and zooplankton in the sea. J. Cons.
into Explor. Mer. 12:147-155.
Holton, R.L. and W.P. Elliott.
1973. The
development of methods for studying physical and biological processes in the nearshore zone on the Pacific coast of the
United States. Progress Report.
School
of Oceanography, Oregon State Univ.,
Corvallis, Ref 73-3, 18 num pages and
appendices.
Olsen, J.B.
1949. The pelagic cyc1opoid
copepods of the coastal waters of Oregon,
California and lower California.
Ph.D.
Thesis, University of California, Los
Angeles, 208p. and XLI Plates.
Huyer, A. 1974. Observations of the
coastal upwelling region off Oregon during 1972. Ph.D. Thesis, Oregon State
Univ., Corvallis, l49p.
Johnson, J.K. 1974. The dynamics of an
isolated population of ACaPtia tonsa
Dana (copepoda) in Yaquina Bay, Oregon.
M.S. Thesis, Oregon State Univ.,
Corvallis, 97p.
Laurs, R.M. 1967. Coastal upwelling and
the ecology of lower trophic levels.
Ph.D. Thesis, Oregon State Univ.,
Corvallis, 2l2p.
Pattullo, J. and W. Denner.
1965.
Processes affecting seawater characteristics along the Oregon coast.
Limnol.
Oceanogr. 10:443-450.
~
Peterson, W.K. and G.C. Anderson.
1966.
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Pacific Ocean: Columbia River effluent
area, 1961, 1962. Univ. Wash. Tech.
Rep. 160, 225p.
~eterson,
W.K.
1972. Distribution of
pelagic Copepoda off the coasts of
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(editors), The Columbia River Estuary and
adjacent ocean waters, p. 313-343.
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Wash.
Press,
Seattle.
Lee, W. 1971. The copepods in a collection
from the southern Oregon coast, 1963.
M.S. Thesis, Oregon State Univ., Corvallis, 62p.
Peterson, W.T. and C.B. Miller.
1975.
Year-to-year variations in the p1ankto1ogy
of the Oregon upwelling zone. Fish.
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Lough, R.G. 1974. Dynamics of crab larvae
Anomura,
Brachyura) off the central
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Ph.D. Thesis,
Oregon State Univ., Corvallis, 299p.
Pillsbury, R.D. 1972: A description of
~ydrography, winds, and currents during
the upwelling season near Newport, Oregon.
Ph.D. Thesis, Oregon State Univ.,
Corvallis, l63p.
Marshall, S.M. and A.P. Orr. 1955. The
biology of a marine copepod Calanus
finmaPchicus
(Gunnerus).
Oliver and
Boyd, Edinburgh, vii and l88p.
56
Rothlisberg, P.C.
1975. Larval ecology of
Rathbun. Ph.D. Thesis,
Oregon State Univ., Corvallis, 117p.
Pandalus jopdani
Smiles, M.C. and W.G. Pearcy.
1971. Size
structure and growth rate of
Eupahusia pacifica off the Oregon
coast.
Fish. Bull. U.S. 69:79-86.
Smith, R.L.
1974. A description of
current, wind, and sea level variations
during coastal upwelling off the Oregon
coast, July-August 1972. J. Geophys.
Res.
79:435-443.
Wilson, D.R. and K.K. Parish.
1971.
Remating in a planktonic marine ca1anoid
copepod.
Marine Biology 9:202-204.
57
part III: appendices
DATE
22 June 1969
"
29 "
"
10 July
"
"
18
25 "
30 "
6 Aug
26 "
30
"
3 Sept
"
14
28
8
23
29
11
18
2
9
29
"
"
"
"
"
"
"
"
"
"
"
Oct
Nov
"
Dec
"
"
"
"
"
"
13 Jan 1970
"
29 "
13 Feb "
"
25 "
9 Mar "
"
16 Apr
27
"
1 May
6 II
22 "
4 June
II
23
2 July
16 II
II
29
13 Aug
II
27
.
11 Sept
25 II
9 Oct
20
II
II
II
II
II
II
II
II
II
II
II
II
II
II
"
"
II
21 Dec
6 Jan 1971
II
18 II
APPENDIX1.
NH5
NH10
X
X
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
X
0
R
X
0
X
X
X
0
X
0
R
X
0
X
X
X
0
X
0
0
X
0
A list
X
X
0
X
X
0
X
0
X
X
0
0
0
X
X
0
X
0
X
0
0
X-R
X
X-R
X
0
X
R
X
X
0
0
X
0
. X-R
X
R
X
0
X
X-R
X
X
X
X
X
X
X
0
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
0
X
II
II
4 Nov
4 Dec
NH3
X
"
"
"
"
NH1
X
X
X
X
X-R
X
X
X
0
0
0
0
X
0
X
0
X
0
X
0
X
0
X
X
a
X
X
0
X
0
X
0
X
a
0
0
of sampling dates and samples collected
X = Sample counted
0 = Sample collected but not counted
R = Replicate samples collected but neither counted
X-R = Replicate samples collected but only one counted
59
0\
a
NH 1
DATE
3 Feb 1971
II
16 II
20 Mar "
"
30 "
22 Apr
3 May.
14 May
29 May
1 June
12 "
28 "
N28 "
29 II
N29 "
6 July"
21 "
22 "
"
"
II
"
II
"
"
"
"
"
"
"
NH 10
NH 30
NH 35
NH 40
NH 50
NH 60
0
X
0
X
0
X
0
0
0
0
R
X
0
X
0
X
0
0
X
0
X
0
X
X
X
0
0
0
0
0
0
0
0
0
0
0
X
0
X
0
0
0
0
0
0
0
0
X
0
X
0
X
0
X
0
X
0
X
0
X
0
X
0
0
0
0
0
0
0
0
0
R
0
0
X
R
X-R
X
R
X
X
R
X
X
X
X
X
X
X
X
X
X
X
X
X
0
X-R
X-R
R
X
X
X-R
X-R
R
X
X
X-R
X-R
R
X-R
X-X
X-R
X-R
X-R
X-R
X-R
R
X-R
X-R
X-R
X
X-R
X-R
X
X
0
0
0
0
0
0
X
X
X
X
0
X
R
X
0
0
0
0
0
0
X
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
O.
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
X-R
X-R
R
X
R
0
X-R
X-R
R
0
0
0
R
R
R
0
0
0
0
R
R
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
X-R
"
"
R
X-R
0
X
X
X
X-R
0
X
X
X
X
0
X
0
R
R
R
X-R
X
X
X
X
X
X
X
X
X
X
X-R
0
X-R
X-R
0
X-R
0
X
X
X
X
0
X
0
R
X
X-R
X
X
X-R
0
X-R
.X
0
X
X-R
0
APPENDIXI (CONTINUED). A list
X
X
0
X-R
X-R
"
NH 25
0
R
22 May
11 June"
"
28 "
"
"
N28
21 July"
"
22 "
"
5 Aug
NH 20
0
R
X-R .
II
NH 15
X
X-R
23 Sept" "
11 Oct
6 Nov "
7 Dec "
3 Mar 1972
"
15 "
"
"
29
11 Apr
" "
20
NH 5
"
2 Aug "
N2 "
19 "
N19 "
NH 3
X
of samplinq dates and samples collected.
N = Night samples
r; U i1 ':3L "
p.."
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07/iO/6CJ
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06/06/69
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12/02l6CJ
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01/29/70
02/25170
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05/06/10
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06/04/70
06/2J170
07/02110
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08/13/70
06/23/70
09/11110
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12/04110
01/06/71
02/16/71
03130111
05/03111
05/14/71
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06/02171
06/12/71
06/26/71
07/06/71
07/21/71
08/0U71
08/19/71
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10/11171
11/06/71
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2.2
.«:
1.4
3.7
14.5
0
42.1
4E.6
4.0
1.3
13.4
41.9
0
24.2
0
5.5
3.3
42.2
1.0
25.3
2.2
5.e
31.3
11.1
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1.2
10.E
35.1
0
1.7
...:t
NUMBER
N
PER CUBIC
06/22/69
06/29/69
07/10/69
07/18/69
07/25/69
08/06/69
08/26169
08/30/69
09/03/69
09/14/69
09128/69
10/08/6CJ
MALES
0
0
0
0
0
0
0
0
0
0
0
0
10/29169
12/02169
12/29/69
01129/70
02/25/70
04/27170
05/06110
OS/22110
06/04/70
06/23/10
01/02/10
01/16/70
07/29/70
08/13/70
08123/70
09/11110
09/25/70
10/20110
.1
02/16/71
03/30/11
05/03/71
05/14171
05129/71
06/02171
06/12171
06/28/11
07/06/71
07/21/71
08/02171
08/19171
OCJ/23171
10/11/71
11/06171
12/01/71
03/15/72
04/20/72
OS/22172
06/11/72
06/28/72
07/21172
08/05/12
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
12/04/70
01/06/11
a
0
0
0
0
0
0
0
0
0
a
0
0
0
0
0
0
0
0
OF
O. SPINIRCSTRIS
FEMALES
0
0
0
0
0
0
0
0
5.1
a
0
0
2.1
2.2
0
.4
17.1
a
1.8
0
2.9
0
0
0
0
0
1.8
0
0
0
0
.5
0
0
0
0
0
0
0
39.3
0
0
7.8
0
0
0
0
3.9
0
0
2.6
1.2
0
.6
0
IMPS
0
0
0
0
0
0
0
0
0
0
0
0
.1
0
.2
0
0
0
3.5
0
1.5
0
{)
0
0
0
0
0
0
0
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0
0
0
0
0
0
0
0
5.4
1.6
0
4.7
0
0
1)
0
2.g
0
0
0
0
1.3
0
0
MALES
FEMALES
0
0
0
0
3.1
0
2.1
0
2.1
0
0
0
13.1
0
0
7.11
14.6
0
0
0
0
0
0
IJ
0
0
0
0
0
0
0
0
a
0
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0
0
0
0
0
0
0
0
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0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
AT
EARLV
LIFE HISTQRY STATIONS
NH05
N03
NH01
DATE
METER
0
11.0
.7
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2.0
a
2.8
0
1.G
0
1.0
4.2
7.2
0
0
16.0
1.8
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0
0
0
0
0
0
2.1
3.5
3.2
15.5
6.7
1..8
14.8
0
2.7
0
1.2
0
0
0
0
8.7
2.0
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0
IMPS
0
0
0
2.1
MAL E"S FEMALES
0
0
0
0
0
0
0
0
0
0
0
a
0
4.'=1
0
0
0
0
0
0
4.0
0
0
0
0
0
0
0
0
0
1.4
2.0
0
0
0
0
0
0
0
0
0
0
0
0
1.1
0
1.g
1.6
0
0
0
2.7
0
0
0
1.2
0
0
0
NHI0
0
0
0
0
0
0
0
C
0
0
0
0
0
0
0
0
0
0
C
0
0
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0
0
0
0
0
0
0
0
0
0
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0
0
0
0
0
0
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0
0
0
0
0
0
0
0
0
2.7
6.6
1.0
6.7
0
4.1
0
2.7
5.5
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0
3.1
0
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0
0
0
3.3
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0
3.8
1.2
3.7.
6.0
2.0
6.8
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0
0
0
0
0
0
0
4.1
0
0
0
0
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0
0
0
0
0
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0
0
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0
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28.3
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0
0
0
0
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4.5
11.6
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0
0
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0
0
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1.0
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3.7
0
4.1
18.7
4.1
0
0
0
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0
4.0
3.6
0
0
0
0
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0
0
0
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0
0
0
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0
0
0
0
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0
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0
0
a
0
0
0
0
0
0
0
0
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0
0
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0
0
0
0
0
0
0
0
0
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0
0
0
0
0
0
0
0
0
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n
0
0
0
0
0
0
0
0
0
0
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0
1.3
0
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0
0
0
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0
0
0
2.0
0
3.4
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1.3
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1.3
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0
2.0
2.2
0
0
1.2
4.2
2.15
10.2
2.6
4.7
3.5
0
0
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14.1
20.6
0
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0
25.2
1+.9
17.7
7.4
1;.2
4.4
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2.1
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11.3
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1.1
4.8
5.6
2.Q
0
1.7
0
0
0
0
0
0
0
0
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0
0
0
0
1.5
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0
0
0
1.7
4.1
0
0
0
2.2
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0
G.4
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0
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1.3
0
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2.4
0
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P£R CUBIC
06/22169
06/29/69
07/10/69
07/18/69
07/25/09
0/06/69
0B/26/69
08/30/69
09/03/69
OQ/14/69
09/23/69
10/08/69
10/29/69
12/02/69
12/29/69
01129/70
02125/70
04/27/70
05/06170
OS/22170
06/04/70
0&/23/70
07/02/70
01116170
07/29/70
08/13/70
08/23170
09/11170
/5/70
/ 0/70
12/04/10
01/06/71
02116/71
03/30/71
05/03/71
05/1'+/71
OS/29/71
06/02/71
06112/71
06/28/71
07/06171
07/21/71
08/D2171
08/19/71
09/23/71
'"
(A
10/11/ 71
11/06/71
12/07/71
03/15/72
04/20/72
OS/22/72
06/11/72
06/28/72
07/21/72
08/05/72
ALi:.S Fc./'tALlS
0
0
0
0
0
0
0
0
0
0
0
0
0
0
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0
0
0
0
0
0
0
0
0
0
0
0
0
0
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2.2
0
0
0
0
0
0
0
0
0
0
0
0
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0
1.5
0
0
0
0
0
0
0
C. ARCUICCR"'IS
0
0
0
0
0
0
0
0
0
0
0
0
0
0
7.0
5.2
0
0
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0
0
0
0
0
0
0
0
0
0
0
0
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22.3
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0
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0
2.6
0
0
0
0
0
0
0
0
0
1.0
1.8
0
0
0
0
0
0
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.
0
0
0
0
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0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
n
11
0
0
0
8
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
II"PS
H,ALc:.S
FEMALES
0
0
0
0
0
0
0
0
0
0
0
0
0
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10.0
0
0
0
0
0
0
0
0
0
0
0
0
0
1.8
0
2.9
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1.1
5.1
0
a
0
0
0
0
T ERLY
LIFE
H1S1C
SlTIC"S
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NH05.
NH03
I'<HOI
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0
0
0
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0
0
0
0
0
0
0
0
0
12.1
5.6
2.0
0
0
0
0
0
c
0
0
0
0
0
3
5.5
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IS.0
1.0
0
0
0
0
0
0
0
0
0
0
0
0
0
1.1
20.3
0
0
0
0
0
0
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0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
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0
0
IJ
0
0
0
0
0
0
0
0
0
0
0
1.0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
C
0
0
0
0
0
0
0
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14.e
15.2
11.2
8
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1.1
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1.0
0
c
0
0
0
0
0
0
0
0
0
0
1.0
2.2
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1.2
0
0
0
0
0
c
c
c
c
c
c
c
0
0
0
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IHFS
C
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
c
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1.1
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0
0
0
0
0
0
0
0
0
0
0
0
0
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0
0
0
c
0
0
0
0
0
0
0
c
1.3
3.1
0
0
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0
0
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0
0
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c
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3.0
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2.4
0
0
0
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0
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0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
C
0
0
0
0
0
0
C
0
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0
0
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0
0
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0
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0
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0
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0
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14.1\
3.5
16.3
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0
0
0
0
0
0
0
0
0
0
0
22.5
5.1
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4.9
2.7
1.7
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0
0
0
0
0
0
0
0
0
0
4.1
S.C;
18.8
0
0
0
0
0
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0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
8
0
0
0
0
0
0
0
0
0
0
0
0
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0
0
0
0
0
0
0
0
0
0
0
0
--.J
NUMBER
PER CUSIC
06/221613
06/29/69
07/10/69
07/18/69
07/25/69
\J8/06/6C1
08/26/613
0 /3D/6CJ
09103/613
09/14/69
09/28/69
lIJ/08109
10/29/69
12102169
12129/69
01129/70
02/25/70
04/27170
05/06170
05/22170
06/04/10
06/23110
01102110
07/16/70
07129/70
08/13/70
08/23170
09/11/70
09/25170
10120/70
12104110
t1ALt.S FMAL[S
0
0
0
0
0
0
0
0
0
0
0
0
0
0
03/15/72
04/20/72
05/22/72
06/11/72
06/28/72
07/21/72
08/05/72
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2.6
0
0
0
0
0
0
0
0
0
0
0
0
0
0
12/07/71
0
0
0
0
0
0
0
0
0
0
17.1
11/06/71
10/11/71
0
0
0
0
5.7
05/03/71
0'5/14/11
0 '5129/71
06/02/71
06/12171
06/28/71
07/06/71
07/21/71
08/02/71
08/19171
09123/71
03/30111
0
0
0
0
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C
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
5.6
10.9
0
1.4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
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0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
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0
0
0
0
0
0
0
0
0
0
0
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0
0
0
0
0
0
0
0
0
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3.6
0
0
0
0
0
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0
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0
0
0
0
0
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0
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0
0
0
0
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0
0
0
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0
0
0
0
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0
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3.6
24.0
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0
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0
0
0
0
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0
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0
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0
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0
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0
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0
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0
0
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0
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0
0
0
0
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0
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0
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0
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0
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0
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0
0
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0
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0
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1.9
0
7.6
0
0
0
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0
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0
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0
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Nt-05
t-1ALES FEMALES
0
0
0
3.1
0
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0
0
0
0
0
0
0
0
0
0
0
0
01/06/71
02116171
14.7
If'!PS
0
0
FERGENS
NH03
NH01
DATE
METER CF CLAUSO.
0
0
0
0
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0
0
0
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0
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0
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0
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0
0
0
0
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0
0
0
0
c
0
c
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2.5
0
0
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0
0
0
0
0
0
0
0
0
0
0
0
0
3.1
0
28.5
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0
0
0
0
0
0
0
0
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3.0
5.5
0
0
0
0
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0
0
0
0
0
0
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0
0
0
0
0
0
0
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0
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10.5
0
0
0
0
0
0
0
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0
0
0
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0
0
0
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20.3
0
0
0
0
0
0
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0
8.2
15.6
18.8
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07l10/6Q
07/16/69
07125/69
08/06/69
08/26/69
D8/3D/59
09/03/69
09/1"/69
OQ128/69
ID/O/6
10/2 /6
12/02/69
12/29/69
01/29/70
02/25/70
04/27/70
05/06/10
OS/22170
06/04110
06/23/10
07/02/70
07/16110
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0
08/13/70
08123/70
09/!1/10
09/ 5/70
ID/20/70
12104/10
01/06/71
02116/71
03/30/71
05/03/71
05/14/71
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06/02171
06/12/71
06/26/71
07106/11
07/21/71
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09/23171
10/11/11
11/06/71
12/07171
03/15/72
04/20172
OS/22172
06/p172
06/
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07/21/12
08/05/72
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LIFE
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DATE
06/22/69
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I
NUMBER
PER CUalC METER OF EUPHAUSIIC EGGS AT EARLY LIFE HISTORY STATIONS
DATE
06/22/69
06/29/69
07/10/69
07/16/69
07/25/69
£18/06/69
08/26/69
08/30/69
09/03/69
09/14/69
09/26/69
10/06/69
10129/69
12/02169
12/29/69
01/29110
02/25170
04/21110
05/06/10
OS/22/10
06/04/10
06/23/70
07/02/10
07/16/10
07/29/10
08/13/10
08/23/10
09/11/10
09/25/10
10/20110
12/04/10
01/06/11
02/16/11
03/30/11
05/03/11
05/14/11
OS/29/11
06/02/11
06/12171
06/28/71
07/06/11
07/21/11
08/02111
08/11Ul1
09/23/71
10/11/11
11/06/11
12/01/11
03/15/12
04120/12
05122172
06/11/12
06/28/12
01/21/12
06/05/12
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08/26/69
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09/14/69
09/26/69
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12/29/69
01/29/70
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06/23/70
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DATE
06/2U6C)
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07/10/69
07/16/6c)
07/25/69
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08/26/69
08130/6c)
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0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
FEMAlES
0
0
OS/22170
06/04/70
06/23/70
01/02170
I) 1/16/70
07/29/70
08/13/70
08/23/70
09/11/70
09/25/70
10/20170
12/04/70
01/06/71
02/16/71
03/30/71
05/03/71
05/14/71
0
0
0
1 C).2
0
0
0
0
0
0
12.7
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
06/02/71
06/12/71
06/28/71
07/06/71
07/21/71
08/02171
08/19/71
09/23/11
0
'5.2
It
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
10/11171
1.1
0
12/01/71
03/115/72
0,./20/72
OS/22/72
0
0
15.3
0
0
0
11/06/71
0&/11/72
06128/72
01/21/72
08/05/72
.9
0
2.1
8.2
0
0
0
0
0
0
0
0
3.0
05129/71
0
0
AT EARLY
CAlYPTCPIS
NH03
01./29170
02/25/70
0,./27170
05/06170
00'
....
MALES
OF
METER
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
{)
0
0
I)
0
0
0
0
0
0
0
0
0
0
0
0
0
0
It
0
0
!)
0
0
0
0
0
0
1.6
1.0
.4
0
2.0
.8
0
0
0
0
0
0
2.4
0
0
0
3.E
26.5
It
0
0
0
1.8
3.6
0
0
31.8
0
0
J
0
.0
0
3.5
0
0
Q
0
0
0
0
a
lIFE HISTOY
5TATIONS
NH10
NH05
IMPS
FEALES
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
{I
0
0
0
0
0
:J
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
(I
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
U
0
0
0
0
0
0
0
0
a
0
0
0
0
a
0
0
0
0
0
0
o.
0
0
0
0
0
0
MALES
G
0
0
0
0
0
0
2.0
0
0
0
.0
2.5
0
0
0
0
3.3
0
2.5
1.2
0
0
0
0
. 0
0
0
0
0
(I
0
0
C
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
I)
0
0
2.7
2.5
I'1PS
Q
0
0
0
0
:J
0
0
0
0
0
0
0
0
0
0
0
0
.7
16.4
FEMALES
0
a
!)
0
0
0
0
0
0
0
0
,0
0
0
C
n
0
0
0
0
0
0
0
0
0
0
0
0
0
MALES
0
0
0
0
0
0
0
C
0
0
0
0
0
0
0
..,
u
0
0
15.2
0
0
.2
0
1.0
2.2
0
0
..2
2.1
7.4
10.2
.5
1.3
1.1
0
0
0
0
1.1
a
2.9
0
1.2
0
0
0
.'5
0
15.0
0
0
0
0
0
0
0
0
0
0
0
a
0
0
0
a
6.0
.8
a
0
a
0
a
0
0
0
a
0
0
0
.3
.7
.Cj
0
0
0
0
.
0
0
0
0
0
Q
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
G
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
C
0
0
0
0
0
0
0
0
0
0
0
0
0
a
0
0
.6
0
0
(j
0
0
0
0
0
0
.B
0
0
0
0
0
0
0
..1...,
0 .
0
0
0
I)
0
0
0
0
0
0
a
(j
1.3
0
0
0
IMPS
0
0
0
0
0
0
0
0
0
0
!J
0
0
1)
a
0
0
FEMALES
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
00'
NUMBER
N'
PER
CUBIC
06/2216CJ
06/29/69
07/10/69
07/18/69
07/25/69
08/06/69
08/2 6/6CJ
08/30/69
DCJ/03/6CJ
09/14/69
OCJ/28/6CJ
10/08/69
10/29/69
12/02169
HALES FEMALES
IMPS
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
11.8
0
0
0
0
0
0
0
1.3
0
0
0
0
0
.0
12/29/69
01/29/70
02/25i10
0
0
0
0
0
0
05/06/70
05122110
06/04/70
06123110
0
0
0
0
0
0
0
0
04/21/70
0
0
0
01/06/71
02/16111
03/30/11
05/03/11
05/14/11
05129111
0
0
0
0
0
0
0
0
0
0
0
0
I)
0
0
0
0
0
06/12/71
06/28111
07/06/11
07/21/71
08/02171
08/19111
09/23/11
10/11/71
11/06/71
12/07111
03/15/72
04/20/12
OS/22/72
06/11/12
06/28/72
07/21112
08/05/12
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
6.3
0
AT
0
0
0
0
{)
0
.2
0
0
0
23.7
0
0
0
9.1
0
0
0
0
0
0
0
0
0
2.8
26.7
0
0
0
0
0
0
0
0
0
.7
0
0
0
0
0
0
0
0
lIFE HISTOPY
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
'
0
0
0
0
0
0
0
0
0
0
0
0
c
1.6
0
0
0
.6
0
.5
.1
0
0
0
0
0
0
Ii
0
0
0
0
0
0
0
0
2.0
20.3
21.9
2.7
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2.2
2.6
12.3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1.3
0
0
0
0
0
0
0
2.5
0
18.2
1.7
IMPS
0
0
0
0
0
0
0
0
0
0
0
0
{)
C
.3
0
0
0
0
MALES FEMALES
4.2
0
0
0
0
0
0
9.0
.7
0
0
7.7
0
2.3
0
0
STATIONS
NH1D
0
0
0
0
0
0
0
0
0
0
0
0
.4
0
0
0
0
0
0
0
0
0
0
0
0
a
4.1
1."
52.1
0
0
0
0
0
Q
0
0
0
0
0
0
0
IJ
0
0
0
0
2.4
0
0
UPS
0
0
0
0
0
2.1
2.1
27.6
EARLY
NH05
0
0
2.<3
0
0
0
2.8
2.1!
.8
0
0
0
0
0
0
0
0
0
0
A
0
0
3.1
0
0
0
0
0
0
0
0
0
0
0
0
0
06/02/11
LI
0
0
0
0
0
0
0
0
0
2.0
0
0
0
0
0
0
0
0
0
FU ReI
HALES FEMALES
0
I)
0
07/02170
07/16/10
07/29/10
08/13/10
08/23110
09/11170
09/25/10
10/20/70
12/04/10
OF
NHC3
NHD1
DATE
METER
0
0
0
0
0
0
0
0
.0
0
0
0
0
0
0
0
0
0
0
0
(j
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
9
0
0
0
0
0
()
0
0
0
0
0
n
0
IMPS
MALES FE"'AlES
0
2.5
0
0
0
0
0
0
0
0
0
(J
0
0
0
0
0
0
0
0
0
0
0
0
.q
0
.2
.4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
5.4
1.8
1.1
0
0
0
0
0
0
0
.7
0
4.Q
0
0
10.2
0
.4
.5
1.7
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
()
()
3.8
0
0
0
0
0
0
I'!
3.3
.7
0
D
0
0
Q
0
0
.6
1.6
0
2.3
1.7
0
0
U
I)
0
I}
0
G
0
0
0
0
.0
0
0
0
1.
"'.;
.
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
iJ
0
0
Q
0
0
U
0
0
l1
0
iJ
0
0
NUMBER PER CUBIC
06/22/69
06/29/69
07/10/69
07/18/69
[) 7/25/69
[) 8/06/69
[) 8/26/69
HAlES
75.4
58.6
11.8
0
0
0
2.6
0
08/30/69
09/03/69
OQ/14/69
0
0
OQ/28/69
10/08/69
10/29/69
12/02169
1212(U69
01/29/70
02/25170
04/21170
05/06/70
OS/22/70
06/04/70
1.5
16.0
[)7/02/70
[)7/16110
19.2
18.3
06/23/70
07/29/70
08/13/70
00;
I
0
0
08/23/10
oQ/11170
OQ/25170
10/20/10
12/04170
01/06171
02/16/11
03/30/71
05/03/11
05/14/11
OS/29/71
06/02/71
06/12/11
06/28/71
07/06/11
07/21/11
08/02111
08/19/71
09/23/71
10/11/11
11/06171
12/01/11
03/15/12
04/20112
OS/22/12
06/11/72
06/28/72
01/21/12
08/05/12
4.2
0
0
0
0
0
0
27.8
0
.9
81.9
17.5
.
7.8
1.2
IMPS
FEALES
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
IJ
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
IJ
0
0
0
0
(I
0
0
. 0
I)
0
0
0
0
0
0
0
0
OF
AT EARLV LIFE HISTORV STTIONS
EVADNE
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
MALES FEMALES
IMPS
0
0
0
.0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
!J
0
0
I)
0
0
0
0
0
0
0
0
0
0
0
0
2.1
0
0
0
50.7
0
1.6
1.E
IJ
0
0
0
0
(J
0
0
n
.8
51.5
4.0
53.0
33.7
0
1.4
0
0
0
0
0
0
0
.0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
23.2
0
0
0
0
0
0
0
0
0
a
0
0
0
0
f)
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
4.0
(!
6.E
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
o.
0
0
0
0
0
0
0
0
0
0
0
o.
0
0
0
0
0
0
1.2
NHI0
NH05
NH03
HMOl
DATE
ETER
0
0
0
0
MALES FEMALES
40.4
23.1
0
0
4.6
1.3
.6
36.8
8.2
0
0
0
C
0
0
0
0
0
0
0
0
0
7.0
2.5
3.5
23.2
1.3
3.5
1.4
2.0
0
D
0
0
0
0
0
0
0
0
0
0
0
IJ
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
. a
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0 .
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1.0
IMPS
0
0
J
0
0
Q
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
MALES FEMALES
0
3.11
0
0
1.3
0
0
IJ
.q
0
0
0
0
0
0
c
0
0
0
0
0
1.2
0
0
0
.7
0
0
0
0
0
0
0
0
0
C
0
0
0
0
0
0
0
0
0
I]
0
0
(J
0
0
0
0
0
0
0
0
0
0
0
0
0
G
a
0
0
0
0
0
0
0
0
0
0
0
0
9.1\
1&.0
0
()
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1.4
IMPS
0
0
0
0
Q
0
0
a
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
00
oj:>.
NUMBER PER curc
09/28/69
10/08/69
10/29/69
12/02169
12129/69
01/29170
NH01
HAlES FEMALES
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
.0
0
0
0
0
0
0
0
0
63.9
0
0
0
0
0
0
05/06/10
OS/22/70
06/04/70
06/23170
07/02170
01/16/10
01/29/70
08/13/10
08/23/70
09/11/10
09/25/70
10120/70
0
0
0
0
0
0
0
0
0
1.8
181.1
20.1
DATE
06/22169
06/29/69
07/10/6'=
07/18/69
07/25/69
08/06/6<3
08/26/69
08/30/69
09/03/69
09/14/69
02/25/70
04/27/70
12/0'+/10
01/06/71
02/16/71
03/30/11
05/03/71
05/14/11
OS/29/11
06/02/71
06/12/11
06/28/71
01/06/11
07/21/11
08/02/11
08/19/11
09/23/71
10/11/11
11/06/11
12/01/7.1
03/15/12
04120172
OS/22/12
06/11/72
06/28/72
071Z1/12
08/05/72
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NUMBER
DATE
06122169
06129/69
07/10/69
07/18/69
07125/69
08/06/69
08/26/69
08/30/69
09/03/69
09/14/69
09/28/69
10/08/69
10129/69
12/02169
12/29/69
01/29170
03/30/71
05/03/71
05/14111
OS/29/71
06/02171
06/12/11
06/26171
01106/11
07/21/71
08/02171
08/19171
09/23/71
10/11/11
11/06/11
12101111
03/15/72
04/20/72
OS/22112
06/11/12
06126/12
07/21172
08/05172
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0
0
0
0
0
0
1)
0
0
0
0
0
0
46.9
2.0
.3
0
0
0
0
.5
0
6.8
0
0
0
0
0
0
I)
0
0
0
0
3.2
.3
6.3
72.5
0
2.6
0
0
IMPS
ALES FEMALES
121. 1
5.0
C
18.0
0
11.4
1.8
0
0
0
{J
0
0
a
0
0
0
0
C
0
0
0
0
0
(}
0
0
0
0
0
0
0
0
n
v
0
0
C
0
0
0
0
0
0
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0
0
0
0
0
0
0
0
0
0
a
0
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0
0
0
0
1.5
1.2
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1.0
2.7
[]
0
!)
0
(J
0
a
0
0
0
0
0
0
0
0
0
0
0
0
0
1.2
a
STATIONS
HISTOQY
tlH11J
0
a
0
0
0
0
C
0
0
0
0
0
0
0
0
0
0
0
0
C
0
0
0
f)
0
0
0
0
0
0
0
0
0
0
0
0
n
6.1
0
It. 1
0
0
0
0
0
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1.1
0
1.5
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0
0
0
0
0
0
0
0
0
0
0
0
Q
0
0
(!
0
0
D
0
0
0
IJ
0
0
0
0
0
0
0
0
0
a
0
0
0
0
0
0
0
0
0
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0
1)
!j
0
o.
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0
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0
0
0
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2.8
2.1
2.1
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0
O
0
Q
0
0
0
0
0
0
0
0
0
0
0
0
0
I)
0
0
{)
0
MALES
0
0
0
0
0
0
0
0
0
6.0
IMPS
0
0
0
0
0
0
0
Q
2.0
FEMALES
0
0
0
0
0
0
0
0
0
0
0
2.7
0
0
0
0
.11
I}
LIFE
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Q
5.5
2.R
0
.1
0
0
0
MALES
0
31. II
3.9
.4
AT EARLY
NHOS
NH03
0
0
0
0
I]
0
0
0
10.2
50.1
192.5
16.4
02/16/71
OF
0
0
0
0
0
0
0
0
48.4
06/04/70
061Z3/10
01/02170
07/16/70
01/29170
08/13110
08/23/10
09/11/10
091Z5/10
12/04110
01/06/71
ETER
IMPS
02/25/70
04/27170
10/20/10
CUBIC
NHOl
MALES FEMALES
S.7
05/06170
OS/22/70
PER
0
0
0
.2
0
0
2.2
G
0
.2
.
4.q
0
1.5
FEMALES
C
0
0
0
0
0
0
0
0
0
0
0
C
. 0
()
0
0
a
0
0
0
0
0
0
0
0
c
0
0
D
0
0
0
0
0
0
.5
.4
1.3
.6
0
0
0
0
0
0
fJ
n
D
0
0
0
0
0
0
0
Ij
0
0
0
0
0
[)
0
1.1
c
.5
.3
0
0
0
0
0
0
0
0
0
0
0
0
0
HPS
iJ
0
0
G
0
0
v
0
0
a
0
0
0
0
0
{)
0
0
0
0
0
0
0
0
0
0
a
0
0
0
0
0
{)
0
0
0
0
0
(j
0
0
:]
0
0
0
0
0
0
0
a
0
0
C
0
0
I
NUMBER PER CUBIC
06/22169
06129/69
07/10/69
07/18/69
07/25/69
08/06/69
08/26/69
08/30/69
09/03169
09/14/6Q
09/28/69
10/08/69
10/29/69
.12102/69
12129/69
01/2917 [
02/25170
04/27170
()5/06110
()5/22/70
(J6/04/7D
10/11/71
11/06/71
12/01/71
0 3/1
5/72
0'+/20/72
OS/22/72
06/11112
06/28/72
01/21/72
08/05/12
0
\)
f}
f}
0
2.0
0
()
0
0
0
0
0
'0
0
.4
2.6
0
130.7
0
0
0
0
0
0
0
0
S.e:
.
0
3.1
0
5.6
0
14.6
11.1
0
0
1105.3
150.0
3.3
0
106.2
154.2
21.8
31.8
29.4
0
1.8
60.0
46.1
10.7
0
38.3
146.7
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
()
0
0
0
0
0
0
0
0
0
0
0
0
0
o.
0
0
0
0
0
\)
0
0
0
0
45.2
4.0
166.!!
4.8
0
296.1
14.3
0
0
0
0
0
0
0
0
0
0
0
0
216.6
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
IMPS
32.5
0
0
0
0
0
0
0
0
0
0
0
II
0
140.4
0
013/23/71
f}
0
0
0
0
0
0
0
0
0
0
0
0
10/20/70
12/04170
06/02/71
()6/12/71
06/28171
07/06/71
07/21/71
08/02171
08/19/71
0
0
0
0
a
0
0
1.0
5.4
8.0
5.2
1.6
120.Q
0
0
0
1.8
47.e;
8.0
14.1
6.E
.15
1.5
0
95.q
0
9.0
.6
0
AT
EARLY
LIFE HISTOY
STATION$
NH10
NH05.
MALES FEMALE'S
0
0
0
0
0
0
0
5.8
4.7
8.0
0
II
257.1
22.3
269.5
51.1
0
55.3
0
62.8
226.7
OS/2C)/71
IMPS
0
70.0
14.7
0
0
0
0
1.3
0
0
0
20.4
.2
0
0
592.5
06/23170
07/02/70
07/16/70
07129/70
08113/70
08/23/10
09/11110
013/25170
01/06/71
02116171
03/30/71
05/03/71
05/14/71
<D
.....
MALES FEMALES
0
BARNACLE NAUP
NHOJ
NHOl
DATE
METER OF
0
0
0
0
0 I
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
I}
MALES FEMALES
IMPS
0
0
2.8
0
18.8
0
0
0
0
0
0
C
0
.1
17.7
0
0
0
87.5
0
0
' 0
0
0
0
0
0
0
0
0
0
0
0
0
0
9.5
0
5.4
.5
0
0
0
0
0
0
0
0
0
0
0
0
fI.O
8.8
0
5.2
0
.4
10.9
0
0
9.4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
34.4
5.3
4.9
0
85.0
0
0
0
0
1.2
0
7.6
35.2
0
.4
0
5.4
0
6.9
0
4.6
(J
>
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
o.
0
0
0
0
0
0
0
0
0
0
0
C
0
0
0
0
0
0
0
MALES FE!o'ALES
'
2.
0
.
0
0
Q
0
0
0
0
0
C
0
0
0
0
a
0
0
J
0
0
(}
0
0
.
0
0
0
0
0
0
0
fj
0
0
0
n
0
0
.1\
13. R
f)
0
0
0
.4
4.4
1.8
0
.2
G
0
415.0
0
,)
,..
..
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
C
0
0
0
0
0
0
.5
0
0
2.7
0
0
0
0
0
0
0
0
.3
0
3.3
23.2
0
0
0
.3
0
1\.5
n
0
(I
0
0
C
0
0
G
0
0
0
0
0
0
0
0
0
iJ
0
0
0
0
0
0
0
0
0
0
0
0
IMPS
0
0
0
0
0
U
0
0
G
0
0
r
0
0
0
G
0
0
0
0
0
0
0
0
0
0
0
tJ
0
0
0
0
0
0
0
0
0
0
'
"
0
0
0
"
n
(]
0
0
0
0
0
0
C
0
0
C
0
0
G
0
0
0
n
0
f)
0
0
11
0
(1
0
..,
0
0
"
0
0
0
0
0
NUMBER
<D
N
PER
CUBIC
06129/6<J
07/10/&<J
07/18/&9
07/25/&9
08/0&/6Q
08/26/6Q
08/30/69
O<)/OJ/&<J
0<)/14/69
09/28/69
10/08/69
10/2<J/69
12/02169
12/29/&9
iJ1/29./ 10
02/25110
04/21/70
MALE'S FEMALES
0
&.0
80.8
13.3
0
0
0
0
362.6
27.0
0
0
0
1.4
.1
13.9
2.4
17.1
.4
0
0
0
0
0
0
0
0
0
0
0
0
0
08/13110
08/23/70
0<)/11110
0<)125/10
10/20110
12/04110
01/06/11
02/16/71
03130/11
27.1
3.6
86.1
49t.6
105.4
0
0
0
0
0
0
0
0
05/03/71
05/14/11
OS/29/11
06/02171
2CJ.6
34.6
3.3
4.2
.8
6.0
0
0
0
0
0
0
0
0
06/12/11
0&128171
07/06/71
07121/71
08/02171
08/1<J/71
254.CJ
39.3
0
0
9.1
0
0
0
0
0
0
10/11/11
11/06/71
12/07/71
03/15172
94.6
211.5
2.4
10.9
0
0
0
0
09/23111
04./20112
OS/22172
06/1111?
06/28/12
01121172
08/05172
6.2
73.1
3.5
33.1
4.7
29.4
2.2
99.0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1.8
0
0
0
0
0
0
0
0
0
0
0
0
0
32.4
4.1
&6.6
250.6
519.7
1.3
0
IMPS
0
0
0
05/0&110
OS/22110
0&/04110
0&/23170
07/02170
07/1&/10
01/29/70
OF
IVALVE
VELIGERS
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
;)
0
0
0
0
0
0
0
0
0
0
I)
0
0
0
0
n
0
0
0
0
0
0
0
0
MALES FHAlES
0
2.0
15.5
4.1
6.2
0
0
0
21.0
22.4
133. 3
0
0
14.2
4.0
8.13
.3
30.1
6.7
9.3
14.3
1.0
34.9
21'.2
53.0
33.7
6.9
7&8.7
65. 9
165.6
1.8
16.7
25.1
2.9
3.6
2.7
0
1.8
63.6
0
.4
0
0
4.1
10.7
39.3
63.3
1.6
.5
0
35. E
2.5
3&.0
3.7
0
UPS
LIFE
HISTOPY
STATIONS
NH10
0
MALES FEMALES
a
a
0
0
0
0
0
0
!J
0
0
0
0
0
Q
0
0
6.8
0
0
0
0
0
0
0
0
0
0
0
0
7.8
0
0
0
I]
0
0
0
0
0
0
0
0
0
0
0
0
0
n
0
0
0
AT EARLY
NH05
NH03
MHOI
DATE
06122169
METER
n
0
0
0
IJ
0
0
:J
0
0
a
0
0
u
0
0
0
0
0
I]
0
0
n
0
0
0
0
a
1)
0
0
0
0
0,
0
0
0
a
0
0
0
0
0
0
0
0
0
0
()
0
0
!1
0
"0
0
0
0
0
0
oJ
0
0
5.4
c
53.2
54.1
13.7
11.0
0
0
0
.0
5.1
.8
0
0
0
0
0
0
1.8
1.6
14.9
0
3.7
2.0
0
0
0
0
I)
.1
3.8
2.3
2.4
116.1
1.7
.3
23.7
30.4
0
4.2
6.6
0
18.2
0
3.4
0
0
4.6
1.8
.9
15.7
19.1
4.4
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
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0
0
0
3.3
0
1.1
0
3.9
6.9
21.9
a
0
0
0
0
0
0
0
0
0
0
0
0
0
0
()
0
0
0
0
0
0
0
0
Q
0
0
0
2.7
6.5
IMPS
0
0
0
0
Q
0
0
0
0
0
Q
0
0
0
0
0
0
0
0
0
0
0
0
0
I)
0
0
0
0
0
0
0
0
MALES FEMALES
a
G
t.3
0
0
0
0
G
0
0
0
0
0
. 0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
D
0
"
c
c
0
0
0
0
D
1.3
0.3
G
I)
0
1.7
0
3.
.1',
2.5
.2
.4"
.6
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4.
12.3
94.4
c
2.'3
3.0
4.0
.9
c
0
.7r,
"
0
0
rJ
1.6
.IS
1.0
D
0
G
3.3
0
0
0
.7
4.3
1.0
.7
.5
0
0
0
2.1
1.2
1.1
0
0
0
u
0
0
0
0
0
0
0
0
0
0
0
0
!t'PS
0
0
0
0
0
0
0
0
0
0
0
c
0
0
0
0
0
0
0
J
0
0
0
0
0
0
U
0
0
0
0
0
a
0
0
0
0
0
0
0
[1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
a
Q
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
r
NUMBER
PER CU8IC METER OF GASTQO VElIGERS AT EARLY LIFE HISTORY STATIONS
OATE
0612Zf&9
06129/69
07110/69
07118/&9
07125/&9
08/06/&9
08/26/&9
08/30/69
09/14/69
09/28/&9
10/08/&9
10/29/&9
12102169
12129/&9
01/29/10
02125/10
04/27/10
MALES FEMALES
0
0
0
0
32.3
0
0
0
0
0
0
0
0
0
0
9.7
0
1.1
0
0
0
0
0
0
.1
0
0
5.9
2.7
0
0
1.0
22.8
0
0
0
05/06/10
OS/22/10
06/04/7Q
06/23/70
07/02170
7.0
0
4.4
33.4
173.2
DCJ/03/&9
a 7/16/70
07/29/10
08/13/70
08/23/70
0«:1/11170
0«:1/25/70
10/20170
12/04/10
01/0&/71
02/16/71
03/30/71
05/03/71
05/14/71
OS/2'U71
06/02171
06/12/71
06/21\/71
\0
\101
J7/06/71
07/21/71
08/02/71
08/19/11
09123/71
10/11/71
11/06/71
12/07171
03/15/72
04/20/72
OS/22/72
06/11/72
06/28/72
07121172
08/05/72
0
0
7.5
0
21.0
227.5
17.5
.4
.9
8.CJ
3.3
30.2
5.6
42.3
0
52.0
30.4
0
0
3.1
2.0
2.7
31.3
5q.9
.5
.9
10.6
15.6
1.2
5.1
1.7
41.7
NH05.
NH03
NH01
0
0
0
0
0
0
0
0
0
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ON,08/19/71
1.9
CAL. MARSYALlAE
CAL. TENUICORNIS
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