Roddie`s Thesis - University of Stirling

Transcription

Factors affecting the abundance and distribution
of estuarine zooplankton, with special reference
to the copepod Eurytemora affinis (Poppe)
Thesis submitted for the degree of PhD
Brian D. Roddie
September 1988
.'
.
"
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Abstract
Factors affecting
the abundance and distribution
estuarine zooplankton
and laboratory
were investigated
of littoral
by means of both field
studies.
In the field, a 12-month survey was conducted at 6 stations
spanning a wide salinity range in the Forth estuary, to
investigate
the influence of geographical,
physical environmental
seasonal,
tidal and
variables on community structure.
Pump
samples, in two net fractions (69 urn and 250 um) were collected
on spring and neap tides, at high and low water over 9 complete
or partial lunar cycles.
The use of two concentric nets of differing mesh size extended
the size range of specimens caught, and permitted the observation
and enumeration
of small plankters such as rotifers, cope pod nauplii
and early polychaete
larvae. In early 1982, a clear temporal
succession
of rotifers> freshwater crustacea>
Eurytemora
was observed.
The data acquired on field distribution
Haranzelleria
larvae>
and abundance were analysed
in a variety of ways. The most effective approach was found to be a
combination
of polythetic,
divisive classfication
sepcies data, followed by Multiple Discriminant
the classification
using geographical,
data as the discriminating
seasonal,
(Twinspan) of
Analysis
(MDA) of
tidal and environmental
variables. The effects of geographical,
seasonal and tidal variation were removed by analysing subsets of the
data restricted
to one state of a variable at a time; comparison
of these restricted analyses with those performed on larger
data sets revealed, however,
that the relative influence of variables
on community structure could be readily discerned even when all
variables were considered
position of station
together. Salinity and geographical
were clearly the dominant factors in explaining
the species associations
suspended particulate
defined by classification
analysis; organic
material was closely associated
with these, and
temperature also but to a lesser degree. The influences of season and
primary production were linked, and were orthogonal
to the influence
of the dominant variables.
Classi~ication
analysis identified three main assemblages:
water community;
a low-salinity
group comprising Eurytemora
the freshaffinis
and Neomysis integer; A neritic assemblage dominated by Acartia spp.,
Pseudocalanus
meroplanktonic
and Oithona but also including Temora, Centropages
larvae. Pseudocalanus
and
and Oithona were more persistent
than the other neritic taxa, and were more often found in samples of
lower salinity and in the autumn and winter.
Predation and development
rate are two biological
directly influence the abundance and distribution
In the laboratory,
of individual
taxa.
studies were conducted a) on the rate of predation
of Neomysis on Eurytemora
availability
factors which
and b) the effects of temperature
on the development
and food
rate of Eurytemora.
Predation rates of adult mysids on adult Eurytemora were estimated
to
range up to 170 prey/day at 500 prey/litre, and the functional response
was adequately modelled by a Type II curve. It was experimentally
, demonstrated
that predation rates were not reduced in the dark or in the
presence of detritus, and it is inferred from this that Neomysis
relies on random foraging rather than on visual predation. Estimated
predation rates were sufficiently
high to suggest that Neomysis
predation may, at some times of the year, have a significant effect on
Eurytemora population size.
Development
rates in Eurytemora were not affected by food level, but
were qu~ntitatively
related to temperature. Development was approximately
isochronal, but the duration of the second naupliar ins tar was consistently
longer than that of other instars, especially at lower temperatures.
Total estimated development
times ranged from 39 days at 8 deg.C to
15.25 days at 20 deg.C, with the effect of temperature beini more marked
at low temperatures
than at high temperatures.
The results of the development study were applied to field observations
of instar body lengths, in order to estimate daily length increment for
9 dates in 1982. Field observations had indicated that, in contrast to
many other studies, body size did not bear a simple inverse relationship
to water temperature; whilst the smallest animals were observed
during the spring bloom .and midsummer,
the largest specimens were collected
in September when water temperatures were still high.
Highest growth rates were estimated for August (small animals) and
September (large animals) ; winter animals, although similar in size to
September specimens, had low estimated growth rates. The large size
of specimens encountered
in September suggests, when considered
in
conjunction with the low abundance at that time, that a switch may
have occurred from investment in reproduction
somatic growth.
to an investment in
GLOSSARY OF ABBREVIATED
TERMS USED IN TEXT
Authority
Taxon
% occurrence
Identifier
(Poppe)
Eurytemora affinis
Adult
copepodite
Nauplii
81.1
89.4
85.0
EAFF ADUL
EAFF COPE
EAFF JUVE
Harpacticid copepods
Rotifera
(mainly Synchaeta sp.)
Copepod nauplii (unidentified)
85.0
73.1
BARP SPEC
ROTI SPEC
73.6
COPE NAUP
Neomysis integer
Diaptomus gracilis
Daphnia sp.
Bosmina sp.
Chydorid spp.'
Cyclopoid spp.
20.7
24.2
13.2
18.5
4.0
10.6
NEOM
DIAP
DAPB
BOSM
CBYD
CYCL
INTE
GMC
SPEC
SPEC
SPEC
SPEC
.20.3
15.4
15.9
8.8
46.2
39.6
29.5
31.3
11.0
9.7
6.2
4.9
ACAR
ACAR
ACAR
ACAR
ACAR
OITB
PSEU
TEMO
CENT
SAGI
OIKO
CALA
INER
TONS
LONG
CLAU
COPE
SIMI
ELON
LONG
HAMA
ELEG
PLEU
BELG
25.5
11.9
51.5
20.3
10.6
29.5
4.8
LITT
LITT
CIRR
CIRR
TERE
POLY
ETEO
CAPS
VELI
NAUP
CYPR
LARV
CILI
LARV
22.5
MARA
MARA
MARA
BIVA
lIIRE
EGGS
TROC
LARV
Acartia·bifilosa inermis
Acartia tonsa
Acartia longiremis
Acartia clausi
Acartia sp. copepodites
Oi thona similis
Pseudocalanus elongatus
Temora longicornis
Centropages hamatus
Sagitta elegans
Oikopleura sp.
Calanus helgolandicus
Littorina littorea
egg capsules
veligers
Cirripede nauplii
cypdds
Terebellid larvae
Polydora larvae
Eteone sp. larvae
Maranzelleria wireni
larvae
eggs
trochophores
Bivalve larvae
Aphroditid larvae
(Leach)
Sars
Rose
Dana
Lilljeborg
Giesbrecht
Claus
(Boeck)
(O.F.Muller)
Lilljeborg
Verril
~(L.)
8.4
7.0
8.4
APHR LARV
GLOSSARY OF ABBREVIATED TERMS USED IN TEXT
Variable
Distance from upper
Shorthand identifier
Transformation
Dist
LoglO
Tide
Dummy 4-state
tidal limit
Tidal state (H,N etc)
variable
Date
Date
None, represented
as decimal month
Inorganic component
Inor
LoglO
Org
LoglO
%org
arcsin/-loglO
of suspended solids
Organi~ component of
.,'
suspended solids
Organic percentage of
(x
suspended solids
Salini ty
Sal
None
Temperature
Temp
None
Chlorophyll a
ChI a
LogiO
Acid Ratio
AR
LoglO
Phaeop igmen ts
Phae
LoglO
Freshwater discharge
Disch
None
at upper tidal limit
X+1
as proportion)
Contents
Introduction
Chapter
1 Field survey of littoral estuarine zooplankton
and associated environmental variables
1
Materials and Methods
Sampling equipment
Calibration and testing
Sampling programme
3 .
Sampling
.'
1
2
9
Estimation of small-scale
temporal and spatial
variability
10
Zooplankton sample processing
13
Numerical analysis of survey
data
17
Multivariate analyses
17
Relationship between community
structure and environmental,
spatial and temporal variables 19
Vater samples
21
23
Results
Environmental variables
23
Temperature and salinity
24
Suspended solids and
photosynthetic pigments
30
Comparative trends
38
Zooplankton
41
Taxa recorded and frequency
of occurrence
41
Distribution of taxa in
samples
43
Spatial and temporal patterns
of abundance in individual
taxa
48
Patchiness
54
Spatial variation
54
Temporal variation
57
Eurytemora affinis
Effects of
preservation
58
58
Variation in length 58
Abundance
65
Age structure
77
Multivariate analysis of field survey data
91
Environmental variables
91
Environmental data
93
Taxa included
94
Samples categorised by station 101
Samples categorised by sampling
cycle
135
Samples categorised by tidal
state
Chapter 2
162
Measurements of development rate in E.affinis
Material and Methods
181
-------------------Experiment 1: development rates in mass
culture at 15 and 20 deg.C
181
Experiment 2: development rates at four
temperature and two food levels
184
Results
189
Experiment 1: development rates in mass
culture
189
Experiment 2: development rates at four
temperatures and two food levels
191
.-
Chapter 3
Primary production
191
Comparison of development and
survival between replicates and
food levels
191
Estimation of egg development
times
193
Estimation of naupliar and
copepodite development rates
194
8 deg.C
194
12 deg.C
199
16 deg.C
205
20 deg.C
210
Average stage duration
210
Development rate as a function
of temperature
218
Estimation of daily length
increment from field and lab.
observations
220
Predation by Neomysis integer on Eurytemora affinis
227
Materials and Methods
--------------------Collection of experimental
material
228
Pretreatment and exposure of
mysids
229
a) effects of size on
predation rate
229
b) estimation of functional
response
231
c) predation in the dark
and in the presence of
detritus
232
Sorting and preparation of
cope pods
233
Results
235
Relationship between size and
predation rate
238
Functional response
242
Predation in presence of detritus
253
Foraging in the absence of light
255
Discussion
257
Field survey
Equipment performance
257
Taxa identified
257
Community structure and environmental
variables
260
Chi-squared test of association
between taxa and samples
261
Classification
262
Assessment of methods
266
Patchiness
Dunmore
Skinflats
Eurytemora affinis
Summary
268
268
268
269
Predation
269
Development rates
271
Field observations
274
277
Introduction
The zooplankton of estuaries has been widely studied (e.g.
Hiller 1983, Collins and Villiams 1982, Taylor 1987) and their
characteristics
described. The estuarine environment
particular problems for zooplankton,
poses
both physiological
behavioural. 'Lance(1965) has demonstrated
and
that osmoregulatory
ability may influence species distribution,
while Burkill et al
(1982) ~nd dePauw (1973) inter alia have suggested that
behavioural explanations
maintenance
need to be invoked to explain the
of endemic populations
in the face of high flushing
rates. For species that can successfully
the low-salinity
overcome these problems,
regions of estuaries may provide a rich resource
(Horris et al 1978, Heinle et al 1977) and lead to high secondary
production.
A feature of estuaries is often the high proportion
total area accounted for by
Roddie (1980) and dePauw(1973)
of the
the intertidal zone and shallows.
have demonstrated
of the littoral zone as a refuge from flushing,
the importance
but more
information on this habitat is needed in order to understand
the
extent to which its inhabitants constitute a true community.
The marked gradients present in estuaries (both spatial and
temporal) should constitute strong influences on the structure of
zooplankton
communities.
The estuarine habitat should therefore
be a suitable environment
and interpreting
A difficulty
assemblages
community structure.
in understanding
the structure of species
lies in objectively
relating a complex of
environmental
variables
of classification
uncovering
in which to test methods for resolving
to a complex of species.
A wide variety
and ordination methods are available for
species association.
to apply combinations
One aim of the present study was
of some of these methods to establish
to
what degree the littoral estuarine plankton of the Forth possess
discernible
structure,
quantitatively
and to what degree this structure could be
related to physical, spatial and temporal factors.
To help achieve this, a sampling programme was designed to cover
the major semidiurnal.and
semilunar cycles on a seasonal basis,
and thus to permit variation on a variety of levels to be removed
by the analysis of categorical
data subsets. The degree to which
a method succeeded could then be tested by examining
the extent
to which samples were sorted by logical category in the analysis.
Biological
factors, in addition to environmental
to structure communities.
and Heinle and Flemer(1976)
Eurytemora
factors, act
Roddie (1980), Burkill et al (1982),
have shown that the cope pod genus
can achieve high abundance at the upper end of
estuaries, and Roddie et al (1984) have demonstrated
of this species to regulate its haemolymph
conditions of low and fluctuating
characterised
the ability
concentration
under
salinity. Taylor (1987) has
the upper estuarine zooplankton
of the Forth
estuary as comprising
the mysid Neomysis integer and Eurytemora;
Heinle and Flemer (1976), Burkill and Kendal (1982) and others
have suggested
that Neomysis may prey on Eurytemora and this may
be an important route of energy transfer from the detrital to the
pelagic food web. A second aim in this work was, therefore, to
establish if predation did occur, and if so to quantify the
process and gain some insight into the potential impact of
Neomysis predation on the Eurytemora population.
The maintenance
of an indigenous population requires that net
population growth rate exceeds the flushing rate of the habitat.
One factor influencing population dynamics is the rate of
physiol~gical
development and maturation
(generation time) of a
species. Evidence indicates that, in copepods, development
is both temperature- and food-dependent
rate
(Corkett and McLaren
1970; Klein Breteler et al 1984). T~e third aim of this study was
to estimate the effect of temperature on the development of
Eurytemora
fed on environmentally-realistic
concentrations
of
natural suspended particulate material. Heinle et al (1977) have
shownth~t
copepods can grow and reproduce when fed
microbially-rich
that Eurytemora
detritus, and Boak and Goulder (1984) have shown
in the Humber estuary can graze on bacteria
attached to suspended particulate material. By applying
laboratory-derived
observations
estimates of development rate to field
of size distribution at different times of year, it
was intended to estimate the daily length increment of each
developmental
stage at a range of temperatures.
Chapter 1
Field survey of littoral estuarine
zooplankton and associated environmental
variables
_,'
Sampling equipment
The sampling equipment was required to effectively
small
«
10 mm) zooplankton
(Roddie,1980)
positioned
in shallow water. A previous study
had used concentric
suspended vertically
sample
250 um and 69 um mesh nets
in a vooden frame,the whole assembly being
in shallow water so that the net openings vere about
20 cm above the vater surface. 100 I of water collected with a
10 I bucket was filtered through these nets.
In the present study, it was considered necessary
to develop
a method which could rapidly sample a larger volume vith less
disturbance
of adjacent vater, and which was free of the depth
restriction
imposed by a wooden frame of fixed height.
Sampling requirements
centrifugal
were met by a l2v Teleflex-Morse
pump with a nominal delivery rate of lOOI/min,
modified as illustrated
in Fig.l.1 • Rigid PVC pipe was connected
to the pump outlet to act as a handle and permit sampling in
depths of up to 1m. A cone attached to the pump inlet reduced
the intake diameter
intake velocity
to 2cm • This had the effect of increasing
(and thus the velocity gradient close to the
intake), reducing the likelihood
that zooplankton
escape
responses would be effective in avoiding capture. 2 m of
flexible 2.5 cm diameter polythene tubing connected
to the rigid
pipe delivered vater to concentric 250 um (inner) and 69 um
(outer) mesh nets suspended from a frame attached to expanded
polystyrene
floats which held the net mouths 15 em above the
water surface. Each net terminated in a 2.5 cm diameter cod end
7.5 em in length, to the bottom end of which was welded a screen
of nylon net of the appropriate
mesh size.
Power was supplied to the pump via 10 m of cable by a l2v motorcycle
1
connection to
'1 m flexible hose
leading to net
support frame
1 m 1" PVC
piPe
.,.
•2 _
2
Cll
diameter intake
attached,__to_pum_p__,.in_le_t--lt-
Ilet support
f_ram--r"e
pump body
...,__s~~tyren.
2.50. WI net
----69
UIl
and cOl!l-end
net. and cod-end
Figure 1.1 Centrifugal puap configuration and concentric net design
battery mounted in a backpack. It was thus possible to operate
the sampling equipment in any location which was physically
accessible. The pump was activated by a toggle switch mounted on
the rigid PVC pipe.
Calibration and testing
The pump was tested to determine whether zooplankton could be
sampled without damage, and to establish the most convenient
method of calibration.
Preliminary sampling was carried out on
30.11.81 at S.Alloa(Fig.1.2).
prese~t.Microscopic
Eurytemora affinis was the only species
examination of the samples revealed that none
of the individuals collected had been physically damaged.
Calibration was achieved by recording the time taken to fill a
45 1 polythene bucket. Initial measurements
indicated that delivery
rate varied between 70 and 85 l/min at between 0 and 25 cm head.
25 cm was chosen as the standard elevation above sea surface for·
the highest point in the delivery system, and every effort was
subsequently
made to maintain this height as closely as possible
during sampling.
Calibration was carried out prior to sampling on every occasion.
Assuming a possible error of +/- 1 1 in volume measurement,
1s in time measurement,
sample volume was thus determined,
independently
for each sample, with an error(range)
approximately
+1- 5%.
2
of
and +/-
Sampling programme
Between December 1981 and December 1982, zooplankton
and water samples
were collected over a series of lunar cycles at 6 shore sites on the
Forth estuary (Fig.1.2,Table
at quarterly
1.1). Sampling was conducted less intensively,
intervals, in 1983. Four sites (Fallin, S.Alloa, Dunmore
and Kincardine)
were located in the upper estuary and were characterised
by low and variable salinity. Two sites (Culross and Skinflats) were
located in the middle estuary and were characterised
and more stable salinity(Roddie,1980).
as a ~ilot survey in determining
represented
The study of Roddie(1980)
higher
was used
the choice of stations. Those chosen
the full range of estuarine conditions;
to reflect considerable
by relatively
Fallin had been found
influence of freshwater planktonic
communities,
while Culross had been found to reflect some influence from coastal
communities.
All sites were readily accessible
was accomplished
was estimated
by road, and travel between sites
by road. Displacement
of high and low water at each site
from the tidal simulation models of Frazer et al (1972),and
trial runs conducted
to establish travel times between sites (Table 1.1).
These trials demonstrated
the feasibility of sampling all stations on the
same tide; in practice, it was possible to follow the tidal pulse up and
down the estuary, sampling within 30 min. of high and low water at each
of the upper estuary sites and within 30 min. of high water only at the
middle estuary sites. The extent of the intertidal zone at the middle
estuary sites made low water sampling impracticable.
Variation
in the timing of high and low water throughout
the year
prevented sampling under similar light conditions both within and
f
•..
Skinf lats
Skrn
Figure
1.2
The Forth estuary and location
sampling stations
4
of
Table 1.1 Tide time corrections for, and travel ti.es between, stations
TIDE TIME CCRRECTIC:iS
(:rtel:,tive to Rosyth )
Ba !Oedon model simulations
High tide
LOl'1
for spring tides
tic1e
Skin:'l!'lts
=
-1 hour
CuIrass
=
-1 hour
lrb.cardine
+10 min.
-1 hour
Dunmore
+20 min.
-145 min.
Allo~
+30 min.
+1 hour
Fallin
+40 min.
+1 hour
StirlinB'
+1 hour
+2 hour
...
TRAVEL TIit.£ BEI'~JEEN STATIOxs
Time in minutes
Journey
Stirling-Skinflats
Stirling-_Culross
35
.
Skinfla.ts-Klncardine
35
10
Kincardine-S •Allo~.
10
S.Alloa-Fallln
Fallin-Stirling
5
15
Kincardin-Dunmore
10
Dunmore-S.Alloa
5
5
cy F'l'a!:"e!'et al (1972)
Table
1.2
Summary of zooplankton samples collected between December 1981
and December 1982
Fallin S.Alloa Dunmore Kincardine CuI ross Skinflats
Date
Tide
,11.12.81
BTS
LTS
BTN
LTN
+
+
+
+
+
+
+
+
+
+
+
+
HTS
LTS
BTN
LTN
HTS
LTS
BTN
LTN
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
HTS
LTS
BTN
LTN
HTS
LTS
BTN
LTN
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
BTS
LTS
HTN
LTN
BTS
LTS
HTN
LTN
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
8.7.82
BTS
+
+
+
+
6.9.82
HTS
LTS
BTN
LTN
BTS
LTS
BTN
LTN
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
HTS
LTS
HTN
LTN
HTS
LTS
HTN
LTN
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
18.12.81
12.1.82
19.1.82
27.1.82
3.2.82
11.3.82
17.3.82
26.3.82
2.4.82
10.5.82
17.5.82
24.5.82
31.5.82
13.9.82
20.9.82
27.9.82
3.11.82
9.11.82
16.11.82
23.11.82
15.12.82
23.12.82
BTS
LTS
BTN
LTN
+
+
+
+
+
+
+
+
+
6
+
+
+
+
+
+
+
+
+
+
+
+'
+
+
+
:.""
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
.
Table 1.,
Recognition
characters for stages of calanoid copepods
Stage
Nauplius
1
3 4 5 6
2
.
Appendage
Copepodite
1 2
..
.
+ + + + + +
+ + + + + +
+ + + + +.+
maxi1lu1e
maxilla
0
0
•
0
0
0
maxi11ipeds
0
0
0
1st leg
2nd leg
0
0
0
0
0
3rd leg
0
4th leg
0
+
+
+
+
+
+
+
+
+ +
+ +
+ +
+ +
+ +
+ +
+ +
+ +
+.+
+ +
0
+
+ + + +
--- 0
0
0
0
0
0
0
0
-
0
0
0
0
0
0
0
0
0
0
0
0
•
•
0
0
0
0
•
•
•
•
3 4 5 Adult
..
'antennu1e
antenna
mandible
5th leg
caudal rami
.
-
+
+
+
+
+
+
+
+
+
---
+
+
+
+
+
+
+
+
+
+
+
+ ..
+fI)
q
+0
J
+~
....
.¥<
+~
+ip.,
'-'
+
+
+
+
I
relatively well developed,but not fully formed
- present as lobed structure, or otherwise undeveloped
• present as group of setae
o structure absent
+
7
between sites. Sampling on a given date commenced with either high or
low tide samples depending on the times of the tides. Sampling always
commenced at the seaward end of the study area.
Sampling in 1981-1982 was conducted during eight lunar cycles(Table
1.2);
within each cycle, sampling commenced on a spring tide and was normally
repeated at
weekly intervals until the following spring tide. At each upper
estuary site, a full lunar sampling cycle thus comprised a total of 8 samples.
Four samples per cycle were normally taken per cycle at the lower estuary
sites. On occasion, sampling was reduced to a single spring tide or a single
spring- neap pair. It was not always feasible to collect all the samples
scheduled for a given date. In 1983, samples were taken at the upper estuary
sites only, at quarterly intervals on high spring tides. On one of these
quarterly visits, a series
of 8 consecutive samples was taken at Dunmore in
order to evaluate small-scale
temporal variability.
On a separate occasion, a
series of samples was taken by boat on a grid superimposed
on the intertidal
mudflats at Skinflats to evaluate the magnitude of small-scale spatial
variability.
8
Sampling
A total of 248 visits to sampling stations were made, during which
528 zooplankton samples were collected.
500 1 volume were taken at each site on each
Samples of approximately
visit, with the sampling equipment described above. The pump was
calibrated immediately before each sample; normally,
the pump was
operated for about 6 min. to filter the required volume.
On most occasions,
there was a residual water flow at the sampling
locations of at least 10-15 cm/s, and the nets were positioned downcurrent of the pump to ensure that previously-filtered
water was not
,
re-sa~pled.Sampling
was conducted near the shore, in water of 50-100 cm
depth.The pump head was moved slowly and continuously
between the surface
and about 10cm from.-the bottom sediment. The bottom consisted of fine
I
mud at all sites except Culross, where sand and gravel predominated.
Yhen no residual water movement was apparent( most frequently at
Skinflats), a transect of approximately
100m was traversed slowly,with
the pump head held as far in advance of any disturbance as possible
(c. 2m).
Occasionally,
high turbidity led to net clogging. Vhen this occurred,
the pump and stopwatch were stopped, the nets agitated gently to clear
the mesh, and pumping and timing resumed. By following this procedure,
it was possible to ensure that 100% of the water pumped was filtered.
Sample volumes were subsequently calculated from the recorded sample
time (+/- O.ls) and the calibrated pump delivery rate(volume measured
+/-
0.5 1) for the appropriate sample.
9
Prior to the collection of each sample, 40% borax-buffered
Steedman's
solution (Steedman 1976) was made up to 4% using 20 um-filtered
ambient water. The use of local water minimised osmotic damage to the
sampled organisms, which were washed gently with a minimum of filtered
local water from the cod-ends of the nets into polythene jars. The
contents of the 250 um and 69 um cod-ends were preserved separately.
In conjunction with each sample,air temperature and local weather
conditions were noted; wind speed was estimated on the Beaufort scale.
Two 1-litre water samples were collected in acid-washed
polythene
bottles. Each bottle was first filled and rinsed with local water, then
squeezed flat,held 20-25 cm below the surface, and allowed to fill
rapidly.Vater
~
temperature was recorded to
+/-
0.5 deg.C using a direct-
reading total immersion mercury thermometer. Salinity was determined
+/-
0.5 ppt using an HCS conductivity
Estimation of small-scale
to
bridge.
temporal and spatial variability
The scales over which these sample series were taken was chosen to be
relevant to the question of the confidence with which a single sample
might be considered representative
considered
of local abundance;
this was
to be of the order of tens to hundreds of metres and of tens
of minutes.
On
23.6.82,
a series of 20 samples was taken over the intertidal
region at Skinflats(Fig.1.2)
to investigate spatial variation within a
short time period. Sampling was conducted over a 3S-minute period
from a small boat in a depth of between 1m and 2m, during slack high
water on a spring tide.
The pump and nets described above were mounted on the gunwale of the
boat; the nets amidships and the pump deployed laterally from the bow
with the intake at a depth of about 65 cm. Boat engine speed was regulated,
using a simple log and knotted line,to give a velocity of 0.5 m/s; at this
speed, the pump intake was effectively
free from any disturbance
propagated by the boat. The net cod-ends were replaced with small polythene
bottles containing 40% Steedman's solution. These bottles could be removed
and replaced with fresh bottles in a few seconds, permitting samples to be
taken virtually contiguously.
Samples were taken in a series of 5 parallel North-South
transects
(Fig.l.2), intended to provide a rectangular grid of 20 samples. Although
this took place at slack high water and in the absence of wind, buoys were
dropped at the beginning and end of the first and last transects to permit
subsequent detection of any displacement which might have occurred.
Each transect was divided into 4 60-second samples on a bearing of 0 or
180 degrees, and with a pump delivery of 76 llmin at
Samples were thus of approximately
a velocity of 0.5m/s.
801 volume, but the duration of sampling
and thus the volume was determined independently
end bottles were changed in approximately
for each sample. Cod-
lOs between samples, during
which time the pump output was diverted away from the nets.
Temperature
and salinity measurements were made at a depth of 50 cm
with an HCS temperature-conductivity
At the end of each transect
bridge.
the boat was turned on a course of 90
or 270 degrees for 60s before turning onto the next transect. The area
11
covered by the sample grid was thus approximately
120m by 120m. Pump
calibration was carried out before and after sampling.
The survey site was visited approximately
18 h later at low water.
The marker buoys were located and bearings taken on visible landmarks
to establish their position. These bearings were transferred to an
Admiralty chart in order to establish the actual shape of the sampling
grid.
On 3.3.83, a series of samples was taken on a high spring tide at Dunmore.
On this occasion, 8 consecutive samples were taken between high
water minus 30 min. and high water plus 30 min. to investigate short-term
tempo~al variability at a fixed point. The sampling equipment was
operated and calibrated as described previously for shore-based samples.
Note was taken of the direction of water movement, and the nets
positioned accordingly to avoid re-sampling water.
Zooplankton sample processing
Upon return to the laboratory, zooplankton samples were rinsed in
filtered seawater diluted with distilled water to the appropriate
salinity, and curated in fresh Steedman's solution made up to 4% with
water of the same salinity.
Samples frequently contained a considerable quantity of fine silt,
especially
in the 69 urn fraction.This
was often sufficient
to have
interfered with subsequent counting. Vhere this occurred, organisms
were extracted from the silt prior to final curation by a density
separ~tion
technique adapted from deJorge and Bouwman(1977).
This
method involves the use of a colloidal silica,Ludox TH, which has
density of 1.3 g/cc.
The sample was concentrated
of 50% Ludox(diluted
to a volume of 50 ml and added to 100ml
with distilled water). The mixture was stirred
thoroughly with a glass rod, a magnetic follower added, and a 1 cm
deep layer of distilled water floated on the surface of the mixture.
The addition of distilled water prevented dessication of the Ludox,
which causes a change of state from liquid to gel. The mixture was
placed on a magnetic stirrer, and the speed adjusted until sediment
was re-suspended
to about half the depth of the vessel. Stirring
continued for one hour, after which organisms were gently aspirated
from the Ludox-distilled
water interface into a l-litre Buchner
flask containing fresh 4% Steedman's solution. A fresh layer of
distilled water was floated on the surface of the Ludox and a
second extraction performed.
This method of extraction proved to be effective in separating all
soft-bodied
organisms, as well as foraminifera, small bivalve larvae
and juvenile gastropods(principally
effectiveness
Hydrobia). Attempts to assess the
of the method for soft-bodied organisms revealed that an
exhaustive examination of the remaining sediment was required to detect
the few remaining individuals. deJorge and Bouwman(1977)
formalin preservation
siltparticles;this
reported that
appeared to prevent adherence of specimens to
proved to be the case in the present study.
Following Ludox separation the remaining silt was scanned rapidly at
low maginification
to detect any larger, denser organisms, which were
removed and added to those already separated.
All zooplankton
counts were performed using either a Bausch and Lomb
or Vild M5A dissection microscope,
dark field illumination.
the latter equipped with light and
Subsamples or aliquots were placed in a modified
Bogarov tray with a volume of 2ml. Vhere practicable,
successive
aliquots were counted until the entire sample had been examined. This
minimised abundance estimation error and ensured that relatively rare
taxa were registered.
Vhere total numbers were sufficiently
large to make subsampling
necessary, the sample was diluted to an appropriate volume(determined
a preliminary
count),agitated
from
thoroughly, and subsampled with a 2ml
Stempel pipette. Successive subsamples were withdrawn
(without replacement)
until at least 100 of the more abundant categories had been counted
(Frolander,1968),
following which the entire sample was scanned with the
aid of a fine dissecting needle for less abundant taxa.
Identification
ICES Zooplankton
of taxa followed the authorities below:
sheets
Acartia
Centropages
Chaetognatha
Polychaete larvae
Rotifera
Appendicularia
Sars, G 0 (1895-1928)
An account of the Crustacea
of Norway
Copepoda
Rose, M 1933
Copepodes Pelagiques
Faune Fr. 26 Paris
Copepoda
Hartmann-Schroder 1971
Die Tierwelt Deutschlands
Annelida 58 Teil
Polychaeta
(esp. Maranzelleria)
.'
Newell G E and Newell R C 1977
Marine Plankton, Hutchinson
Additional information on a
variety of taxa
lS
Formal identification
organisms
was accomplished
by dissecting(where
in lactic acid, mounting the diagnostic
lactophenol,
appropriate)
structures
in polyvinyl
and examining at 400x under a compound microscope.
The commoner copepods were identified
to species, and, where practicable,
recorded as adult, copepodite or nauplius. A large category of cope pod
nauplii remained unassigned
All developmental
using Table 1.3
to species,however.
stages of Eurytemora affinis were enumerated
and decriptions
in Katona(197l).For
separately,
9 sampling dates
(Table 1.7) the mean length of up to 50 preserved specimens of each stage
was determined
using a calibrated eyepiece graticule.Nauplii
at 200x,to the nearest 10 um, while copepodites
and adults were measured at
120x to the nearest 16.6 um. For the purposes of examining
distribution
were measured
the size-frequency
of adults, copepods were grouped into 0.02mm size classes. Hale
and female copepodites
of stages 4,5 and 6 were counted and measured as
separate groups.
The length of nauplii was measured excluding
the caudal armature. Copepodite
length was measured as the length of the cephalothorax,
excluding
the 'wings'
on the final thoracic segment in the case of stage 5 and 6 females.
On 27.4.82, samples were taken to determine
the effect of preservation
on
adult body length. Copepods were first killed or rendered comatose by
raising the water temperature
to 30deg.C over a period of 30 min.
100
.
adult females were randomly removed from the sample and their cephalothorax
lengths measured as described above. They were then transferred
Steedman's
solution and stored for two weeks.Vater
of capture and 1.2 urn-filtered was used throughout,
changes which might have affected body dimensions.
copepods was re-measured
preservation
~ollected
to 4%
at the time
to minimise osmotic
The sample of 100
after two weeks and the mean pre- and post-
lengths compared by t-test.
16
Numerical analysis of zooplankton
survey data
The frequency of occurrence of taxa in all samples and by sample
category was computed. Samples were classfied by logical category
(high/low, spring/neap,
association
by station, etc.), and a chi-squared
test of
between frequency of occurrence and logical category
carried out.
Multivariate
analyses
The sur~ey data were analysed by logical category using classification
and ordination
techniques. The primary methods were routines in the
Cornell Ecology Series of programs:
Twinspan (classification:
Two-Yay
Indicator Species Analysis, Hill 1973) and Decorana (ordination: DEtrended
COrrespondence
ANAlysis, Hill 1979) •.
Twinspan (TS) is a polythetic divisive method of classification.
purposes of comparison, a synthetic agglomerative
For
method was also tested;
for each sample category a similarity matrix was generated using the BrayCurtis index (log x+1 transformed data) and a dendrogram constructed by
group-average
sorting (CLUSTAN, Yishart 1970).
All TS and Decorana (DCA) analyses were conducted on data subjected to
quasi-logarithmic
1000,1001-10000,
transformation;
abundance classes of 0-30, 31-100, 101-
10001~100000 and >100000 were used.
These intervals
were used as the cut levels for defining TS pseudospecies;
in effect, and
for the purposes of computation, each abundance class is regarded as a
'species', although indicator species are still defined by their presence or
17
absence. An upper bound of 30 was used for the smallest abundance class to
avoid undue influence from taxa occurring in low numbers. Cassie (1962) amongst
others has reported a high degree of uncertainty
in estimating population
abundance when the number of individuals in a sample is small, and it was
considered
that abundances between 10 and 30 per cubic metre could not be
reliably distinguished.
In both T5 and DCA analyses, the supplied option to downweight
rare species
was invoked.
For the purposes of T5 classification,
a-character
identifiers.
samples and taxa were each assigned
For samples, the identifiers were constructed as
follows:
Character
Purpose
1
Letter identifying station (F,5,D,K,G,C)
2-6
Digits identifying date
7-8
Letter identifying
tidal state (H5,L5,HN,LN)
Taxa were allocated 4 characters for generic name and four characters for
specific name, e.g.
were not identified
ACAR LONG represents Acartia longiremis. Vhere taxa
to species, a suitably obvious alternative was used, e.g.
rotifers were identified as ROT! SPEC.
A suite of Fortran programs was developed to re-format data structures
for the various analyses,produce
raw data tabulation and generate similarity
matrices (Roddie 1986).
18
Subdivision
of the entire data set into logical categories
removal in turn of seasonal,spatial
permitted
the
and temporal effects on community
structure.
Relationships
between community structure and environmental,
spatial and temporal variables.
a) DCA
For each analysis,
rank correlation
the first two DCA axes were plotted, and Spearman
coefficents
between major variables and the sample scores
on these axes computed.
b)
TS
For each analysis, a TS two-way ordered table and a sample dendrogram
were produced.
Multiple Discriminant
Analysis
used to relate environmental
(Green and Vascotto
Following
(MDA, • canonical variate analysis) was
variables
to classification
1978, Vright et a1 1984, Furse et al 1984).
the recommendations
(log x+l transformed)
which were implemented
of Green and Vascotto
were entered simultaneously
of sample taxonomic composition)
into the the analyses,
( in this case on the basis
and derives a linear combination
the ratio of between-group
those variables which had a significant
particular
(1978), all variables
by an SPSSx routine (Nie 1975). MDA uses the grouping
assigned to samples by a previous classification
which maximises
group membership
to within-group
univariate
of variables
variance. Only
F-ratio (p<O.05) for a
sample category were used in the analysis for that category.
19
Following derivation
percentage
successful
of the rule, the samples are re-classified
and the
re-classification
factors are
calculated.
Discriminant
derived which permit the samples to be located in 2-dimensional
discriminant
space and enable a visual assessment 'of the power of the analysis in
discriminating
A convenient
group structure on the basis of related variables.
method of displaying
the influence and relative importance
of variables
is to plot the classification
group centroids
in multiple
discriminant
space, and to project onto the plot vectors which indicate
the direction and relative magnitude of influence of the variables
discriminating
between groups (Overall and Klett, 1972). All vectors
originate at the overall sample group centroid in discriminant
endpoints are defined by co-ordinates
products of the univariate
space; their
which, for each axis, are the
F-ratio (between- to within-group
for a variable and the correlation
discriminant
in
variance ratio)
between group variable mean and group
function score mean. Thus, the direction of the vector is
a function of the degree of association
between each discriminant
and a variable, and the length of the vector a reflection of the
degree to which a variable is related to group structure.
20
function
Vater samples
Prior to 3.2.82, only salinity and temperature were measured, using an MCs
temperature-conductivity
bridge calibrated at regular intervals against a
silver nitrate titration. Salinity was recorded to 0.1 ppt, and temperature
0.5 deg.C. From 13.2.82, measurements
a and chlorophyll
made following
to
were made of suspended solids, chlorophyll
breakdown products (phaeopigments).
All determinations
were
the methods of Strickland and Parsons(1968).
Suspended solids were measured by filtering 250-1000 ml of sample water
onto pre-ashed,tared
(+1- 0.01 mg), Vhatman GF/C glass-fibre
a nominal retention of 1.2 um.At least two 1.2 um-filtered
of comparable
volume were carried through in conjunction
seawater blanks
with each set of
samples. A single sample only was processed in association
zooplankton
filters with
with each
sample. After filtration of each sample the filter was rinsed
three times with 10 ml distilled water to remove salts,placed
on clean
aluminium foil and dried at 60 deg.C for at least 12 h. Dried filters were
cooled in a dessicator
for 1h, weighed to the nearest 0.01 mg, placed on clean
aluminium foil, and ashed for 2 h at 450 deg.C. Filters were again cooled in
a dessicator
before re-weighing.
Dry and ash weights were corrected for
average blank changes, and the results expressed as corrected dry and
ash-free dry weight.
Analysis for chlorophyll
a and phaeopigments
was carried out on 200ml
samples filtered onto Vhatman GF/C filters. Filters, with retained
material, were folded twice and placed in polythene centrifuge
containing
tubes
12 ml 95% acetone (made up with Analar acetone and distilled
water, and neutralised
with sodium bicarbonate).
21
Centrifuge
tubes were
incubated overnight in a darkened refrigerator
extraction of photosynthetic
at 7 deg.C to allow
pigments to take place. After at least
ISh extraction, the tube contents were centrifuged at 3000 rpm for 10
minutes. The supernatant was decanted into a quartz spectrophotometer
cuvette with a 10 cm path length.
Optical density was measured on a Cecil UV spectrophotometer
95% acetone blank at 663 nm and 750nm. The latter wavelength
against a
was used
to correct for general sample turbidity; no known photosynthetic
pigments absorb at this wavelength. Following initial measurement,
the
sample and blank cells contents were acidified by the addition of 5
drops o~ 10% Hel and allowed to stand for 5 min. to allow any transient
turbidity to clear. Optical density was re-measured at the same wavelengths as before. Chlorophyll a concentration,
phaeopigment
concentration,
and the acid ratio(a measure of the health of the phytoplankton
population)
were calculated using the equations of Strickland and Parsons (1972).
22
Results
of
Environmental
field
survey
variables
Seasonal variation in measured environmental
on a station-by-station
variables will be examined
basis. The variables considered are:
salinity
temperature
chlorophyll a
phaeopigments
total and organic suspended solids
Additionally,
for the purposes
community structure,
of investigating
influences on zooplankton
tidal state, time, and freshwater discharge
to the
upper estuary will in a subsequent section be treated as environmental
variables.
All observations
in Appendix A.
on measured variables are tabulated by station and tide
Temperature
and salinity
Fallin
Low tide salinity at Fallin ranged from 0%0 to 0.8%0, whilst high tide
salinity encompassed
a range of 0%0 to 12.2%0 (Table Al ,Fig.1.3a). On only
two occasions did salinity exceed 5.7%0.
Temperature
symmetrical
corresponded
varied annually between 0 and 19 deg.C, following an approximately
curve (Table A1,Fig.1.3b).
High and low tide temperatures
closely throughout the year. A temperature
range of up to 6 deg.C
during a single sampling cycle (11-17 deg.C, May-June 1982) was observed.
S.Alloa
There was a wide discrepancy
at S.Alloa (Table A2,Fig.l.3c).
summer and least pronounced
between the high and low tide salinity curves
The difference
being most pronounced
in autumn and late winter/spring.
range was 0-5.5%0, and high tide salinity range was 2.0-22.6%0.
in
Low tide salinity
Low tide
salinity varied little within any sampling cycle, but high tide salinity
displayed a wide range (up to 13.9 %0) within sampling cycles.
Temperature
displayed a closely similar range and pattern to that at Fallin
(Table A2, Fig.1.3d).
~-
b)
~
Fallin
tempreature
0
v
~
?P
I ,0
,.;
~
\~
~
.'
~)
S.A1loa salinity
~)
S.Alloa temperature
.. _.o.. ....
-e:
-"'-...
~~-
--- -_
.................
.-~
------~ ~'
F\
M
Figure 1.3
o
N
1>
Annual variation in salinity and temperature at Fallin
(a,b) and S.Alloa (c,d). Open symbols .high tide
Closed symbols • low tide
25
Dunmore
The pattern of salinity variation at Dunmore was similar to that at
S.Alloa (Table A3, Fig.l.4a), with maximum salinities occurring in the
summer and during equinoctial
tides. Low tide range was wider (0.6-14.0%0)
than at S.Alloa, as was the high tide range(3.8-28.5%0).
Temperature
ranged between 0 deg.C and 17.0 deg.C, and the pattern again
matched that observed in stations further up-estuary(Table
A3, Fig.1.4b).
Kincardine
Minimum low tide salinity at Kincardine was 3.4 %0, and the maximum
observed was 23.0 %0 (Table A4,Fig.1.4c).The
high tide range was 10.4%0
to 31.5%0. Compared to stations further up the estuary, the difference
on any date between high and low tide salinities was relatively small.
Temperature
occupied the range 1.5-16.5 deg.C, and was thus slightly
reduced with respect to up-estuary stations (Table A4,Fig.l.4d).
The
annual pattern of variation in temperature was similar to that at upestuary stations.
26
et)
Dunmore salinity
10
'2S
\~~
____
----
'lD
~
..........
...0.. ...
.r
I!,
,0
"-,
~
b)
..... ~..
'"
.........
''
/
-1 \
,I
~:,,~,,~
Dunmore temperature
r=>
-.-'- ........_
",
~::--
r-----4
..
__ --~
'
c."
,/
)0
G.o. ..,
'2~
o
1.0
o..!2
IS
Kipcardine saiinity
,p............
,..
.......
..
..
:~~-_
<,
-..
---'{
I
/V~
,
I
It>
...........
S"
/
/\
Kincardine temperature
10
Figure 1.4
Annual variation in salinity and temperature at Dunmore
(a,b) and Kincardine(c,d). Open symbols .high tide
Closed symbols • low tide
27
Skinflats
Salinity varied between a maximum of 31.0%0 and a minimum of 15.8%0
(Table AS,Fig.1.5c),
and temperature between 0 and 21.0 deg.C (Fig.1.5d).
Neither the pattern of salinity variation nor that of temperature corresponded
closely to those at other stations.
Culross
Salinity at Culross varies relatively little throughout the year, showing
only on~ major reduction (to 10.6%0) in late September (Table A6, Fig.1.5a).
Salinity was normally in excess of 20%0, and reached a maximum of 32.6%0.
The reduction in salinity in late September closely matched similarly
abrupt reductions at Kincardine,Dunmore,
Temperature
and S.Alloa.
ranged from 5.5 deg.C to 27.5 deg.C; both minumum and maximum
values were higher than those of other stations, but the seasonal pattern
was otherwise similar.
28
Culross salinity
)0
,l''\,/
':lS
,
,,
,%0
0
~
,,
(~
1'0
Ij
Culross temperature
Po_
.,,,"
~)
0..........
Skinflats
\., V - ",/
~
" ......
"
...
salinity
'\
~
V - ----
pr ---- _--------~...
.
...
""".1\/'
It)
tN
Skinflats temperature
1.t'"
'Zo
v
0
\S'
fb
'5'
~ ..
"
r>
Figure 1.5
,,
A
s:
S
Annual variation in salinity and temperature at Culross
(a,b) and Skinflats(c,d). High tide samples only.
29
?
,
Suspended solids and photosynthetic
pigments
These variables will not be reported in detail here; the data will be
analysed more comprehensively
in conjunction with the analysis of
zooplankton survey data.
Fallin
Suspended particulate material exceeded 200 mg/l on 3 occasions on high
tide, but on 5 occasions on low tides (Fig.l.6a,b; Table Al). The organic
proportion lay between 10 and 20% for both tidal states. On low tides, there
was a t~ndency
towards higher suspended particulate loads on low spring tides.
,
Chlorophyll a and phaeopigments showed less variation between tidal states;
the latter pigment class consists of chlorophyll breakdown products and
predominated over chlorophyll a on all occasions except one (Fig.l.6 c,d).
The acid ratio, an indicator of phytoplankton health (1. senescent, 1.7.
healthy) reached a maximum on the same date, but did not otherwise exceed
1.52 and predominantly
took values indicating a low ratio of production
to breakdown.
S.Alloa
Both suspended particulates and pigments at S.Alloa showed less variability
and lower values than at Fallin on high tides, but greater variability and
magnitude than at Fallin on low tides (Table A2,Fig.1.7).
Acid ratio reached a maximum of 1.67 on the same date in July as the Fallin
, I
""""'_'QI
2
<:Ciiiii
o
~
,~
5
o
J
c:::::J
id
t
c
~
.'
~
....,
u
-e
-.1 _-.1
C -c u
as
bel v
.. .,
-e ..
... ,....,. ...
O-.t:l
I
WQlas
-"0
te c
CIl.l:_
"OCil
-
~asc
0
CIl~
"0
QI
...
.1:-
-CCJ
QI_CIl
as
c CIl
QI"'''O
Q.CQI
CIlQl ..."O
"0
I~
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-~
=' e .....
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maximum, but otherwise generally indicated a preponderance
products over viable photosynthetic
of breakdown
products (TableA2, Fig.l.7).
On low tides, there was a tendency towards higher particulate
loads on
spring tides. Particulate loads exceeded 200 mgll on eleven low tides but
on only two high tides. The organic percentage did not exceed 18% on low tides
but exceeded 25% on four high tides (Table A2,Fig.l.7).
Dunmore
Suspended particulate concentrations
exceeded 200 mgll on eight low tides,
but exceeded 100 mgll on only one high tide (Table A3, Fig.l.8); this
high tide value was, at 853.3 mgll, the highest value recorded during the
survey. The organic content of the suspended particulate material lay in
the range 13-22% for low tides but was 13-31% on high tides (Fig.l.8).
The seasonal trend in primary production was more clearly evident on high
tides than on low tides, a feature also of the data for S.Alloa and Fallin.
Pigment levels were higher and more variable on low tides than on high tides
and appeared to be to a large degree associated with suspended particulate
levels.
~IC===========~----~~
.....
-
.,.
,
.........
a
\
~
c
_ ..-
...
I
IV
e
\0-
j
0
=
.:!.
I
=
Ic:::::J
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.
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3.5
Kincardine
The variability
of suspended solids at Kincardine appeared considerably
reduced with respect to the upper-estuary
stations; on only four dates did
levels exceed 200 mg/l on low tides, and on only one date did levels exceed
100 mg/l on high tides. The organic proportion
ranged between 13-22% on
low tides and 15-26% on high tides.
Pigment concentration
(Table A4,Fig.1.9).
variability vas also reduced, especially on high tides
Again, the seasonal pattern in chlorophyll a vas most
evident on high tides. High levels of chlorophyll a on low tides vere
associated with high concentrations
of both total pigments and of suspended
solids; .this is likely to have been a consequence of the resuspension
~
benthic diatoms into already-turbid
of
water.
Culross and Skinflats
The range of values observed for variables at these two lower-estuary
stations indicated considerable damping compared to the upper estuary.
Organic content of suspended solids at Culross between 14 and 30% and at
Skinflats between 13 and 32% (Tables A5,6; Fig.1.10).
v
i.-
..
e-
~
~
P
~
0
.f'
et
h
~
J
I'"
11.
~
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et
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t
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eG
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ell ell
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(I
p
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ell rn
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rn "0 ...
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ell .... 0
c::~0.
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CIS"O~
=,QlCIlO
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;s
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~CIS"O'"
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rn ell w
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0/
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r
~
'J7
Comparative trends
To facilitate comparison of seasonal trends between stations, total
pigment concentrations
(Fig.1.11) and total suspended particulate material
(Fig.1.12) were plotted on a logarithmic scale versus time.
Total suspended particulate material
A
degree of coupling was apparent on high tides between the following
pairs of stations: S.Alloa/Fallin,
Dunmore/Kincardine,
Skinflats/Culross.
On low tides, Fallin, S.Alloa and Dunmore exhibited a strong degree of
coupling.
~
Total Pigments
On high tides, Fallin and S.Alloa appeared to be coupled to some extent.
Neither Dunmore nor Skinflats matched the patterns at any other stations, but
Kincardine and Culross showed similar patterns of variation.
On low tides, Fallin, S.Alloa and Dunmore again appeared strongly coupled,
whilst Kincardine had some features in common with these stations.
Trend Summary
Figures 1.11 and 1.12 demonstrate that, especially within the upper estuary,
patterns of temporal variation environmental variables may extend over a
scale of tens of kilometres.
Total Pigments - High Tide
100
10
rot
Fallin
~
1
::s
S.Alloa
Dunmore
Kincardine
---~-------------~/~
Skinfiats
Culross
Total Pigments - Low Tide
Fallin
S.Alloa
Dunmore
Kincardine
Figure 1.11
Seasonal trends in total photosynthetic pigments at all
stations on high and low tides. All values plotted on same
logarithmic scale
39
Total Suspended Particulate
Material - High Tide
Total Suspended Particulate
Material - Low Tide
1000
100
,....la
Fallin
f
S.Alloa
Dunmore
Kincardine
Skinflats
Culross
Fallin
S.Al1oa
Dunmore
Kincardine
Figure 1.12
Seasonal trends in total suspended particulate material at all
stations on high and low tides. All values plotted on same
logarithmic scale
40
Taxa recorded and frequency of occurrence
The following taxa were regularly recorded from samples collected in the
course of the survey; in association with each taxon, the authority (where
appropriate),
frequency of occurrence in samples, and the shorthand identifier
used in subsequent analyses are recorded (Table 1.4).
In addition to the above, the following taxa were encountered
on rare'
occasions;
..
-
Copepoda
Annelida
Eurytemora velox
Syllid sp.
Aphroditid larvae
Corycaeus anglicus
Acartia bifilosa
Acartia discaudata
Pseudocalanus
Mollusca
Nudibranch juveniles
Hydrozoa
Steenstrupia sp.
parvus
Sarsia sp.
Amphipoda Gammarus zaddachi
Hyale sp.
Orchestia sp ,
Chaetogammarus
Evadne sp.
Teleost
Pomatoschistus
sp.
Corophium sp.
Oligochaeta
Acaridae
Turbellaria
Nematoda
Foraminifera
Cladocera
41
sp.
Table ,.~ Taxa recorded, frequency of occurrence in samples, and
condensed taxon identifier
Taxon
Eurytemora affinis
Adult
copepodite
Nauplii
% occurrence
Authority
(Poppe)
Harpacticid copepods
Rotifera
(mainly Synchaeta sp.)
Copepod nauplii (unidentified) ,_
Neomysis integer
Diaptomus gracilis
Daphnia sp.
Bosmina sp.
Chydorid spp.
Cyclopoid spp.
Acartia bifilosa inermis
Acartia tonsa
Acartia longiremis
Acart~a clausi
Acartia sp. copepodites
Oithona similis
Pseudocalanus elongatus
Temora longicornis
Centropages hamatus
Sagitta elegans
Oikopleura sp.
Calanus helgolandicus
Littorina littorea
egg capsules
veligers
Cirri pede nauplii
cyprids
Terebellid larvae
Polydora larvae
Eteone sp. larvae
Haranzelleria wireni
larvae
eggs "
trochophores
Bivalve larvae
Identifier
(Leach)
Sars
Rose
Dana
Lilljeborg
Giesbrecht
Claus
(Boeck)
(O.F.Muller)
Lilljeborg
Verril
81.1
89.4
85.0
EAFF ADUL
EAFF COPE
EAFF JUVE
85.0
73.1
HARP SPEC
ROTl SPEC
73.6
COPE NAUP
20.7
24.2
13.2
18.5
4.0
10.6
NEOH
DlAP
DAPH
BOSH
CHro
CYCL
lNTE
GRAC
SPEC
SPEC
SPEC
SPEC
20.3
15.4
15.9
8.8
46.2
39.6
29.5
31.3
11.0
9.7
6.2
4.9
ACAR
ACAR
ACAR
ACAR
ACAR
OITH
PSEU
TEMO
CENT
SAGl
OlKO
CALA
lNER
TONS
LONG
CLAU
COPE
SIMI
ELON
LONG
HAMA
ELEG
PLEU
HELG
25.5
11.9
51.5
20.3
10.6
29.5
4.8
LlTT
LITT
CIRR
CIRR
TERE
POLY
ETEO
CAPS
VELI
NAUP
CYPR
LARV
CILl
LARV
(L. )
,22.5
8.4
7.0
8.4
42
MARA \lIRE
HARA EGGS
HARA TROC
. BIVA LARV
In terms of frequency of occurrence,
unidentified
dominated
E.affinis,
cope pod nauplii (predominantly
harpacticid)
copepods,
and rotifers
the survey; all these categories were found in excess of 70%
of samples. The next most frequently occurring
copepods Oithona similis, Temora longicornis
accounting
harpacticid
taxa were the neritic
and Pseudocalanus
e10ngatus,
for 30-40% of samples. Compound taxa such as cirripede naup1ii
(predominantly
Balanus and E1minius but not differentiated)
and the nauplii
of the Acartia genus were also present in a large proportion of samples.
The Acartia species complex was well-represented
in these littoral samples
(6 species in all); Taylor (1987) found a similar range of species in
his con~emporary
study of the plankton of the main channel of the Forth.
Distribution
The distribution
of taxa in samples
of species with respect to sample category was initially
assessed by tabulating
the number of samples within each category in which
a species occurred (Table 1.5) and then testing (chi-squared
the distribution
of occurrences
overall distribution
is relatively
between related categories
of samples between categories
insensitive
to hierarchical
test) whether
deviated from the
(Table 1.6). This approach
structure; variation between one
set of categories within another is likely to be obscured.
43
Table18 The distribution of species between tide and station
categories
Taxon
DAPH SPEC
BOSH SPEC
DIAP GRAC
CYCL SPEC
CHYD SPEC
NEOH INTE
EURY AFFI
ACAR CLAU
ACAR LONG
ACAR TONS
ACAR BIFI
ACAR INER
ACAR DISC
PSEU ELON
OITH SIHI
CENT HAHA
CALA HELG
TEHO LONG
PARA PARV
SAGI ELEG
OIKO''DIOI
ROTI SPEC
HARP SPEC
POLY CILI
MARA VlRE
MARA TROC
MARA EGGS
LITT CAPS
LITT VELI
APHR LARV
SYLL SPEC
ETEO LARV
TERE LARV
NERE LARV
CIRR CYPR
CIRR NAUP
BIVA LARV
Category
High Low Spring Neap HS LS HN
13
14
21
8
1
14
134
18
30
26
1
43
4
51
76
24
11
63
3
22
11
98
113
56
28
12
12
41
24
2
3
5
19
15
36
93
11
17
27
33
16
9
33
86
2
6
8
0
3
0
6
14
2
0
8
1
0
1
68
79
11
23
4
7
17
4
4
0
6
5
9
10
24
8
13
19
22
14
7
22
111
9
20
22
1
30
2
29
55
17
7
42
2
17
11
82
96
38
29
12
15
33
17
6
2
8
13
18
30
64
9
17
22
32
10
3
24
109
11
16
12
0
16
0
28
35
9
4
29
2
5
1
84
96
29
22
4
4
25
11
0
1
3
11
6
16
53
10
LN
4
9 9 8
6 13 8 14
8 14 13 19
5
9 3 7
1 6 0 3
8 14 5 19
71 40 63 46
9 0 9 2
18 2 12 4
19 3 7 5
1 0 0 0
28 2 15 1
2 0 2 0
28 1 23 5
51 4 25 10
16 1 8
1
7 0 4 0
40 2 23 6
1 1
2 0
17 0 5 0
10 1 1 0
49 33 49 35
57 39 56 40
32 6 24 5
.16 13 13 9
9 3 3 1
9 6 3 1
26 7 15 10
15 2 9
2
2 4 0 0
2 0
1 0
4 4 1 2
2 8 3
11
11 7 4 2
26 4 10 6
55 9 38 15
5 4 6 4
44
FA SA DU KI
SK CU
16
19
21
9
6
8
41
0
2
1
0
2
'0
2
1
0
0
0
0
0
0
35
36
5
7
0
2
8
1
0
0
0
0
3
2
3
0
0
1
0
1
0
1
20
6
3
6
0
14
2
15
15
4
3
15
1
12
4
13
19
11
4
1
1
8
5
0
2
1
3
3
6
16
4
7
9
20
7
4
15
49
1
3
5
0
4
0
7
11
0
1
5
0
0
0
44
44
11
11
2
4
7
4
0
0
2
4
4
8
14
0
5
8
7
5
0
16
47
1
10
9
0
10
0
17
20
7
2
15
1
4
2
45
36
10
13
3
4
12
5
1
0
3
3
7
9
26
2
2
4
6
2
0
7
45
9
12
7
1
10
1
16
28
8
2
24
2
3
3
31
40
18
12
8
6
10
6
5
1
5
9
5
13
39
12
0
0
0
0
0
0
18
3
6
6
0
6
1
10
15
4
3
12
0
3
3
7
20
12
4
2
2
13
6
0
0
0
5
2
8
19
1
•
Table 1.6 Association of taxa with tidal and geographical sample
categories. +,- indicate a significant chi-squared value:
the sign indicates whether a taxon could most easily be
considered biased towards or away from a particular
category
Taxon
}{
L
S
N
j·1S
LS
BOSM SPEC
+
+
DrAP GRAC
t
+
cyeL SPEC
+
+
+
MEOM IN'I'E.
",CAR CLAU
+.
ACAR lONG
+
ACAR TeNS
AeAR
I.CM
AeM
PSW
BIFI
INER
DISC
ElON
OrTf{ SIMI
,COO
HAMA
IN f"A
+
+
+
+
...
...
CALA HUG
+
TENO lOOG
+
FARA 'P/-.RV
SAGI ElEG
011(0 DIO!
+
1-
+
+
SA
DU
K
SJ(
Cu
+
+
+
+
+
+
CHYD SPEC
EURY AfT!
}IN
+
DAPH SPEC
.'
Station
Tides
+
+
+
+
+
t-
+
+
+
+
+
+ +
+
+
+
...
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
ROr.I SPE'C
~:A~..PSl'!':'
POLY
cm
+
MARA \4TRE
+
+
~ARA "['~
MARA EGGS
LITT CA:F5
lIT!' YEU
+
+
APHR WV
+
+
+
SYLL SfEC
ETEO lARV
TER8 LA.T<V
+
cm
+
+
+
nERe lAAY
CIP.:t
C!:R:t NAUP
+
+
+
+
+
ElVA LAAV
+
4S
+
+
+
High/low tides
Comparison of high and low tide occurrences revealed two readily-distinguished
groups; those which occurred significantly more frequently than expected on
high tides and those which showed a similar bias towards low tides.
The former group comprised mainly neritic calanoid copepods - Acartia spp.,
Pseudocalanus,Oithona,
Centropages,Calanus
and Temora- together with Oikopleura,
Sagitta and some lower-estuary meroplanktonic
forms (Littorina egg capsules,
cirripede larvae and Polydora larvae). The second group comprised predominantly
freshwater holoplankton
including Daphnia,Bosmina,
and the upper-estuarine
mysid Neomysis.
cyclopid copepods, chydorids
Spring/neap tides
The majority of taxa had occurrences more or less evenly divided between
spring and neap tide samples (Table 1.5). Only four groups showed a dev~ation
from the relative frequency of occurrence of spring and neap samples. The marine
genera Sagitta and Oikopleura both occurred significantly more frequently
than expected on spring tides, as did the larvae of aphroditid and nereid
polychaetes.
High and low spring/high and low neap tides
This comparison refines the evidence from the two previous comparisons.
The neritic copepods oc~urred significantly more frequently than expected on
high spring tides, as did Sagitta and Oikopleura.
significantly
Aphroditid
larvae were
associated with low spring tides, as were the freshwater
holoplankton.
Littorina egg capsules and veligers, and cirripede nauplii and cyprid larvae
were both significantly
associated with high spring tides.
Comparison
Twenty-eight
between stations
out of forty-two taxa showed a significant
degree of association
with station (Table 1.5, Table 1.6). Neritic copepods were most strongly associa
with Dunmore and Kincardine
(Fig.l.2), whilst freshwater
cladocera and cope pods
were st~ongly associated with Fallin and S.Alloa. Neomysis appeared
~
significantly
more often in S.Alloa and Dunmore samples than in collections
from other stations. Heroplanktonic
frequently at the up-estuary
larvae occurred significantly
less
stations Fallin and S.Alloa than the relative
number of samples taken at each station would predict. Haranzellaria
larvae
were most strongly associated with Dunmore and Kincardine,
while littorinid
and cirripede larvae were biased towards the lower-estuary
sites Culross
and Skinflats.
Sagitta and Oikopleura were also biased towards the lower-
estuary stations.
47
Spatial and temporal patterns of abundance of individual
species
Raw abundance data (individuals/cubic
metre) were tabulated by station and
tide (Appendix B), and abundances of selected taxa plotted on logarithmic
or linear scales as appropriate
to the abundance levels recorded
(Appendix
C, Figs.Cl to Cl9).
The contents of Appendices
will not be interpreted
Band
C are intended primarily
in detail. Six taxa have been omitted from tabulation
in Appendix B; Syllid larvae, Paracalanus
discaudata.
for reference, and
sp., Acartia bifilosa. Acartia
nudibranch juveniles and aphroditid
larvae all occurred in less
than four percent of samples.
Seasonal plots of abundance were constructed
following five categories
(holoplanktonic
station-by
taxa chosen on the basis of abundance
and/or their relative importance within a particular
E.affinis
Nauplii
Copepodites
Adults
Freshwater
Diaptomus
Daphnia
Bo~mina
Chydoridae
station for the
planktonic
association):
Mixed taxa
Unassigned copepod nauplii
Neomysis
Barpacticid copepods
Rotifera
Acartia genus
A.clausi
A.longiremis
A. tonsa
A.bifilosa inermis
r('~(,
)
~
S
Neritic, copepods
Temora longicornis
.,'
Centropages hamatus
Oi thona similis
Psudocalanus elongatus
Meroplankton
Several meroplanktonic
taxa,showed strongly seasonal patterns of abundance.
This was especially marked for the eggs and larvae of Littorina ,cirripedes,
and the spionids Maranzelleria and Polydora. Littorinid egg capsules were
largely confined to Culross between March and July 1982 (upto 4343/m-3)
although 692 m-3 were present on 15.12.82 at Culross(Table B6). Cirripede
nauplii displayed a similar pattern of abundance but a longer period of
~ccurrence
at Culross (Table B6), and were als0
abundant over the same
period (March-September)
at Kincardine and Skinflats. At each station at
which they occurred regularly, both Littorina capsules and cirripede nauplii
displayed a pattern of relatively higher abundance on high spring tides than
on high neap tides.
Larvae of Maranzelleria
were confined to a period
in March-April
1982, and
were most abundant (up tp 160000 m-3) at Skinflats although occurring at
all stations including Fallin. Larvae of terebellid polychaetes,
Eteone and bivalves attained
Polydora,
both maximum frequency of occurrence and maximum
abundance in May 1982 (Tables B 4 to 6). The spionids Maranzelleria
and Polydora displayed a succession of larval production.
E.affinis
The pattern of abundance of E.affinis will be described below in conjunction
with consideration
of the population size and age structure.
Freshwater taxa
The influence of the freshwater community extended, with diminishing
magnitude, as far down-estuary
as Kincardine
(Figs.C16 to C19). Abundances
were generally low for all taxa, rarely exceeding 200m-3. There was a
trend, becoming more marked down-estuary,
for abundances
to be greater on
low tides than on·high tides. At Fallin, S.Alloa (except Diaptomus on
low tides) and Dunmore. Diaptomus,Daphnia
so
and Bnsmina were most abundant
during the period January-April,
the period September-November.
whilst chydorids were most abundant during
Chydorids appeared at Dunmore in only one
sample (low tide), and were absent from all Kincardine,
Culross and
Skinflats samples. The pattern of abundance of Diaptomus, Daphnia and
Bosmina at Kincardine on low tides was similar to that further up-estuary.
Acartia genus
Members of the Acartia genus occurred on only five occasions at Fallin
(Table B1). The abundances of A.clausi, A.longiremis,
A.bifilosa
inerm1s
and A.tonsa have been plotted for high and low tides for S.Alloa, Dunmore
and Kincardine,
and for high tides at Culross and Skinflats(Figs.
A.clausi occurred infrequently,
in low numbers and predominantly
C11-1S).
on high
tides at all the above stations, exceeding 300 m-3 only once (Skinflats,
March 1982). A similar spring maximum occurred at Kincardine;
at the
remaining stations, maximum abundance occurred towards the end of the
year (Fig. C13).
A.longiremis
exceeded 100 m-3 only once, at Kincardine
remaining occurrences were patchily distributed
at Culross
and Kincardine.
in July 1982. The
in time and most frequent
Occurrences were predominantly
At Culross and Skinflats, A.longiremis
on high tides.
was most common in the latter half
of the year (Figs. C14,15), but at Dunmore and S.Alloa occurred most
frequently between May and July 1982. Occurrence was distributed
over a
longer period of time at Kincardine than at other stations, with a number
of appearances
in lo~ tide samples.
A.bifilosa
inermis occurred at all stations in generally low abundances
and in no obvious pattern. Numbers of over 1100 m-3 were recorded in a
single May sample at Skinflats, and reached 225 m-3 at Kincardine on the same
occasion (Table B4,5;Fig.C13,14);
abundance did not otherwise exceed
100m-3 at any station. Low tide occurrences were rare.
A.tonsa appeared at all stations except Fallin from September onwards. This
species showed considerable overlap in occurrence with A. longiremis.
Neritic copepods
~
Temora, Centropages,
Oithona and Pseudocalanus
(Table Bl-6;Figs C6-10)
showed 'no clear seasonal pattern of abundance, and there was little
correspondence
in occurrences between stations. At Dunmore and S.Alloa,
Centropages had peak abundance in May. At Culross and Skinflats, Centropages
showed a similar peak, but at both stations the maximum Pseudocalanus
numbers occurred in November. At Kincardine, Centropages and Pseudocalanus
were abundant only in November and December 1982.
Temora and Oithona showed intermittent peaks of abundance throughout the
year.
Mixed Group
Of the four taxa in tl.is group, only Neomvsis and the rotifers will
be considered here. Harpacticid copepods are predominantly benthic in habit,
and were present in samples mainly as a consequence of resuspension.
by definition
identified,
Although,
in this study, the mixed cop~pod nauplii were not fully
the majority were of harpacticid
species.
Neomysis occurrence was most frequent, and abundance highest, at S.Alloa.
The mysid occurred only in low tide samples at Kincardine,
was rare at
Skinflats and absent from Culross samples. Only at S.Alloa did Neomysis
occur during the early part of the year (Table B3;Fig C2a,b). At
Fallin and all other sites, Neomysis appeared only from May-July onwards, with
maximum abundances occurring at most sites in September.
Rotifera (predominantly
Synchaeta sp. , but including freshwater species
such as Keratella and Polyarthra
) occurred in high numbers at all stations
(excep~ Culross) between December 1981 and February 1982. Highest abundances
.
were recorded at S.Alloa during this period, when numbers exceeded
300000 m-3 (Table B3). Rotifera occurred almost exclusively
69 um net fraction.
in the
Patchiness:
spatial and temporal variation in abundance
and distribution
on the scale of sampling
Spatial variation
The position of the buoys used to mark the limits of the sampling
grid was checked, using a hand-bearing
compass, during low tide on the day
following sampling. Vithin the limits of resolution of the method, the grid
was found to be approximately
120m square and oriented
N-S E-V.
E.affinis, Polydora larvae, and Acartia clausi were found to be sufficiently
numerous in the approximately
modelled;
80 1 samples for their distribution
to be
these groups, together with total number of taxa, salinity and
temperature, were contoured using computer routines supplied by the
GINO SURF package.
The resulting plots are rather idealised, since each sample is treated as
a single point rather than as a segment of a transect. They nonetheless
serve to
give some indication of the presence and dimensions of plankton
patches in the littoral zone. In interpreting
these contour plots, it should
be borne in mind that the adjacent shoreline runs N-S , and that therefore
the horizontal axis of the plots is parallel to the shore.
Both salinity and temperature display a distinct front
shore (Fig.1.13e,f)
parallel' to the
Beyond 80 m from the inshore edge of the grid, both
variables were relatively constant at >26%0 and 11.5 deg.C respectively.
Moving inshore, salinity decreased rapidly to 1~%o over a distance of 20m,
and temperature increased to 14.0 deg.C. Vithin 40m of the westerly edge of
the grid, salinity had fallen to 16%0, but with some suggestion
low-salinity
that the
water was bounded to the south by a wedge of higher-salinity
water. Since sampling was conducted in about 1.5 m of water, and sample
bottles were immersed to a depth of 50 cm, it is unlikely that this pattern
could be attributed simply to a shallow layer of low-salinity
water
overlying more saline water.
The number of taxa present (Fig.1.13d) was reduced within the front, increasing
both inshore and, to a greater extent, offshore.
The three taxa represented
have been contoured in numbers/100l,
differing patterns of distribution.
range of abundance
(20-60/1001);
and showed
Polydora larvae showed a relatively small
the contours indicated patch sizes of between
30 and 100m radius (Fig.1.13c), with parts of up to four patches present within
the grid. One patch appeared to be located within the front itself.
In contrast, E.affinis ranged in abundance from less than 5 to more than 55
per 100 1, and the main patch (Fig.1.13a) did not appear to cross the front.
This patch had dimensions of 60-100m, and was oblate in a N-S direction.
Although E.affinis is generally associated with lower-salinity
instance the main concentration
water, in this
of numbers was on the high salinity side of
the front.
A.claus!
displayed yet another pattern distribution.
Numbers were at a
minimum of about 50/1001 within the front. To either side of the front, patches
of 40-100m diameter, with up to five times this minimum had developed.
Substantial numbers were therefore associated with both cooler, high-salinity
water and with warmer, low-salinity water.
E.affinis
A.clausi
No. of tau
]:
..
•...
..
iii
w
c
iii
C
Distance South(m)
Figure 1.13
Small-scale spatial variation in zooplankton distribution,
salinity, and temperature within a 120m square grid at
Skinflats. Zooplankton abundance contoured in nos./1001.
Temporal variation
In a series of 8 consecutive 5001 samples taken at Dunmore on 3.3.83,
numbers of E.affinis ranged from 137 m-3 to 909 m-3, with an arithmetic
mean of 496 m-3. The maximum abundance occurred in the middle of the sample
series, with numbers rising and falling more or less monotonically
on either
side. The 95% confidence interval for a single sample (expressed as a
percentage of the mean was calculated using the approximation:
C.l ••
[antilog (y+l- 2s)]
x 100
antilog y
(Gagnon and Lacroix, 1982)
where s - standard deviation of the log-transformed
values
y - geometric mean
This gave limits of 29-347% of the mena, and indicated that abundances in
samples taken over longer time intervals would need to differ by a factor
of 3.5 or more to be considered significantly different.
Eurytemora affinis: field observations
Effects of preservation
on length
A comparison between the length distributions
and unpreserved
female E.affinis
of subsamples of preserved
indicated that Steedman's
solution had no measurable effect on cephalothorax
length. Consequently,
no correction was made to the lengths recorded from preserved specimens
from the field survey.
Variation in length between tides and stations
In order to simplify the process of obtaining stage-specific
for E.affinis
length data
throughout the year, it was necessary to establish whether
samples from a single station and tidal state could be considered
representative
distribution
of the upper estuary as a whole. To this end, the size
of male and female adults was plotted for Fallin, S.Alloa
and Dunmore, on high and low tides, on consecutive spring and neap tides
(17.5.82,24.5.82,31.5.82).
On 17.5.82 there was no verall difference in size distribution
between
stations or tides (Fig.1.14), although the modal length of males in the
low tide Dunmore sample was greater than at the up-estuary stations. The
modal length of males ~~s consistently
less thall females.
On 24.5.82 and 31.5.82 , there was again no obvious pattern of difference
ht
It .
Lt
ht
Fa
S.A.
17.5.82
o
..
'
I
I.
'r,
.
LI
_.LII'L.___
!.
M
1"
2t..5. B2
31.5.82
Figure 1.14
Size-frequency histograms of male and female adult E.affinis
collected on high and low tides at Fallin, S.Alloa and Dunmore
on 17, 24 and 31 Kay 1982
Table "7(
Mean cephalothorax length of c6 female E. affinis
collected at HT S.Alloa on 18 dates
Length (mm)
standard error
mean
n
0.010
17
0.007
0.015
0.011
0.010
0.010
0.008
30
20
0.008
0.010
,50
20
0.693
0.007
30
0.9510.95'
0.905
0.01.0
20
0.011
9
0.009
zo
3.11. 8?
16.11.82
0.807
0.009
20
0.793
0.014
10
22.11.82
1,5.12.82
0.769
0.853
0.e48
0.008
0.006
30
:30
)0
Date
'<3.2.82
0.836
0.841
0.898
11.3.82
2.4.82
0.882
0.842
10.5.82
17 •.5.82
24.5.82
0.736
0.719
0.712
0.694
11.12.81
12.1.82
.: :31.5. 8?
8.7.82
6.9.82
13·9.22
2').9.82
23.12.82
20
30
30
,50
0.018
60
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lengths
A
S
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1981-2
Figure 1.15b Mean cephalothorax lengths of adult female E.affinis collected
on 18 dates at S.Alloa
62
1
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Figure 1.16
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Nt
Hean cephalothorax lengths of all developmental stages of
E.affinis collected on 9 dates at S.Alloa
6)
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64
•
in female length between tides and stations, but again some indication that
the modal length of males was greater at Dunmore.
On the basis of the above observations,
S.Alloa high tide samples were
selected as the source of specimens for biometric and age-structure
evaluation. Mean female c6 cephalothorax
(Table 1.7;Fig.1.1Sa,b),
length was determined
for 18 dates
and mean length of all stages determined
for
9 dates (Table 1.9;Fig.1.16a,b).
Female length distribution
(Fig.1.1Sa) was too irregular to permit the
discernment of multiple generations.
Both length frequency and mean length
(Fig.1.1Sb) displayed a bimodal temporal distribution,
with peaks of 0.898mm
on 3.2.82 and 0.9S2mm on 13.9.82. Mean length fell to 0.769mm on 22.11.82
before .returning in December to levels comparable with the same period in
1981. Minimum mean length was observed in May and July samples; these were,
at 0.693mm and 0.694mm respectively,
some 27% less than the maximum.
The annual variation in lengths of other stages followed a similar pattern,
which was most apparent in cS and c6 and increasingly damped in earlier
stages as variations
fell within the limit of precision of measurement
(Table 1.7;Fig.1.16a,b).
A summer minimum was still evident for n6 , however.
C6 females were consistently
larger than other stages. For both cS and c6,
females were always larger than males. There was a degree of overlap between
the lengths of c6 males and cS females.
Abundance:upper
estuary
The highest recorded abundances of E.affinis were at Fallin (exceeding
250000 m-3 on 24.5.82 and 31.5.82) and Dunmore (179796 m-3 on 2.4.82)
(Table 1.8). Maximum abundance at S.Alloa was 96280 m-3 on 24.5.82 and at
HIGH TIDE
• • • • • • • • • • • •• • • • • • • • • ••
cd...n-':TJN...JW (o~<D""'.I'I ~'8'_n,:;:J'~(_~'~.f)::l""'lI
•••••
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m
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a
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Fig. 1.17 a Abundance of
1:. affinis
I
66
Fallin high tide
tJ
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Fig. 1.17 b Abundance of ~
affinis
67
I
Fallin low tide
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1-4
HIGH TIDE
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Fig. 1.18 a Abundance of
!:. affiniscSouth
68
Alloa high tide
1~
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(J,.)
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!:. affinis
,South Alloa low tide
Cl
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HIGH TIDE
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Fig. 1.19 a Abundance of !.:. affiriislDunllore high tide
70
0
:z
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11
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ID
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Fig. 1.20 a Abundanceof
!:. affinislKinoardine
72
high tide
In
,
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, , ,~ (.1'1
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Fig. 1.20 b
Abundance of ~ affinislKincardine low tide
13
0
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Fig. 1.21
Abundance or
!:. aff1nis.Sk1nf1ats
74
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,....l
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Fig. 1.22
Abundanceof
E.!. aff1nislCulrols
75
high tide
Table 1.8 Annual variation in mean abundance of E.affinis (all stages,
nos. per cubic metre) at Fallin, S.Alloa and Dunmore on
high and low tides, 1982
Oate.
Tide.
.n. :t.2.81
S
HT
14-67l
15.12..81 N
.'
12.1.82
s
1.9.1.. e2
l'
-'
27.1.82
3.2.82
11.3.82
18.3.82
26.3.82
2.4.82
10.5·82
17 •.5.82
24.5.82
31.5.82
8.7.82
6.9.82
13.9.82
20.9.82
S
27.9.82
N
3.11.82
~.11.e2
16.11.82
22.11.82
1.5.12.82
23.12.82
S
N
s
N
S
N
S
N
s
N
s
....
Q
N
S
N
s
6822
122
186
727
LT
HI
'291
007)
495)
114
36
0
26
32
59
88
.82
307
2720.5
76
24279
95+
139,503
59888
98519
266032
5499
250741 132.561
25921
6
71
11
17
69
42
23
499
5+
8
71
176
N
S
N
51
0
14517
2223
4178
14145
3976'
355+
4981
14108
17682
63385
49388
72280
4.5348
201
128
1642
333
3296
682
708.5
3193
1,541
2206
76
l<incdrdine
Dunmore
S .Al1oa
fall in
L!
673
1397
90
260
599
51
465
5+.536
6868
18609
16629
96280
70390
LT
HI'
1235
261
2709
6530
1975
1007
251
1174
771
1550
399
2615
1793
6103
1658
12.53 12013
12258 179796
8880
358
26976
43620
10999 121389
6462 9.5063
9:fl
65
45
128
38
32
31
261
14
1594
51
0
39
18
19
10,56
6815
2440
1350
998
1470
48
312
358
376
~rr
45
3.51
LT
.518
597
1674-
32
492
57
133
114
382
418
4153
513
1921
361
390
.5302
1129
908
7r::A
48e.9
336
1263
1843
19922
7499
42
0
2
:fo
10
1)41
3977
1008e
:,>0
2116
422
97
26
182
419
288
31
·40C
30
269
639
280
169
1670
Kincardine
was 19922 m-3 on the same date.
At Fallin, high tide abundances
exceeded low tide abundances
the year, while at S.Alloa, Dunmore and Kincardine
generally
throughout
low tide abundances
exceeded high tide abundances.
At all stations, abundance declined dramatically
(Figs.1.17-1.22),
recovering
The spring population
between July and September
(least notably at Fallin) in November.
maximum coincided with the period of minimum size noted
above, and the late summer decline with the period of maximum size.
Naupliar,
facilitate
copepodite and adult abundances were plotted on log scales to
comparisons
(Figs.l.17-1.22).
of trends between these age groups and between stations
The same seasonal trend was evident at all stations, but more
striking at each station was the extent to which the age groups reflected the
~
same fluctuations.
The fluctuations
were, however, only comparable
in the
broadest sense between stations. The figures suggest that sampling variability
and/or small-scale
spatial variabilty
contribute an important element of noise
to abundance estimates.
Age structure
The percentage distribution
of stages within samples was calculated
Fallin, Dunmore and S.Alloa (Figs.1.23-1.25).
between S.Alloa and Dunmore age structures
for
Vhilst there was some consistency
prior to April 1982 (Figs.1.24,1.25),
there was no evidence of either consistent age structure over short time
periods or of systematic
change in age structure on a seasonal basis. It is
thus unlikely that structure was determined
by
overall synchrony of
reproduction
in the population (which might have led to the development of
identifiable cohorts) or that there was a strong influence of age- or densitydependent mortality in a continously-reproducing
population (which might
have led to a constant decline in relative abundance with age). It is likely
that reproductive processes are patch-dependent
and that the sampling method
integrates the information from several patches in an uncontrolled
The stage-frequency
manner.
histograms did, however, emphasise the presence of nauplii
in at least some samples on all sampling dates, showing that E.affinis breeds
throughout the year.
78
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83
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86
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0
t-
t-
N
.....
.....
U)
:J:
(.!)
.....
~
:I:
U)
~
e
~
oJ
N
N
c;)<S>c;)c;)CS)C!»~C!»C!»CS)CS)
(!;)tSl(!;)CS)c;)CS)~c;)c;)c;)CS)
(s)CT.lO)~(DIn"C"lN
(S)cr;.Cl.I~WIn"C"lN'"
....
N
M
N
\.
('oj
10
to
to
::>
..r-
..r-
o
UJ
Cl
N
.....
,_
UJ
N
D
W
:I:
(.!)
.....
..r-
:J:
.....
t:>
e
oJ
a:l~cs)IS)IS)CS)CS)1S)1S)1S)1S)
IS) cS) CS) IS) (!.) C!» CS) IS) IS) IS)'IS)
~O)CD .....W In V" C'?N-
~(T.IO'J""'WIf')TC'?N-
UJ
U)
(D
~
~
N
t-
U)
:J:
(.!)
.....
~
:I:
..rN
N
0
.....
W
UJ
0
.....
t-
~
e
oJ
N
U)
~
N
N
CS)(S)C!»CS)CS)CS)~CS)CS)CS)CS)
C!»<S>C!»CS)C!»CS)~C!»C!»C!»CS)
(s)CT.lQ)~(DIn"C"lN'"
(S)cr;.o)~u)In"C"lN""
'N
....
CO
\.
N
....
to
f.D
:;)
.....
..r-
e
UJ
N
0
.....
UJ
...J
"-
Q.
t-
W
:I:
(.!)
.....
..r-
:J:
z:
-c
u:
e
...
VI
Z
N
1S)cS)1S)1S)CS)1S)(!,)1S)1S)1S)1S)
r!;HTJCl) r- W In T C'?N -
"IS)(S)IS)IS)IS)IS)~IS)IS)IS)(S)
(S)(T.IOO.....
WIf')TC'?N-
Fig. 1.25 d Stage:-:frequeney diatrih1tionlDunllore
90
Multivariate
analysis of field survey data
Sample identifiers
In order to avoid confusion with S.Alloa, Skinflats has been
assigned the identifier G (for Grangemouth,
Environmental
an adjacent town).
variables
The following variables were used in analyses:
Variable
Distance from upper
Shorthand identifier
Transformation
Dist
LoglO
Tide
Dummy 4-state
tidal limi t .
.
'
Tidal state (H,N etc)
variable
Date
None, represented
Date
as decimal month
Inorganic component
Inor
LoglO
Org
LoglO
of suspended solids
Organic component of
suspended solids
Organic percentage of
%org
(x as proportion)
suspended solids
Salinity
Sal
None
Temperature
Temp
None
Chlorophyll a
ChI a
LoglO
Acid Ratio
AR
Phaeopigments
Phae
LoglO
Freshwater discharge
Disch
None
at upper tidal limit
91
Data on freshwater discharge was obtained from the Forth River
Purification
Board (FRPB 1983).
Suspended solids values were recorded in mgll, and Chla in ug/l.
The entire 1981/1982 data set was subdivided into logical
categories
for analysis. Subsets were extracted corresponding
station, date(sampling
to
cycle) and tidal state.
Station and date categories were subjected to DCA,TS and MDA
analysis. Similar analyses were conducted for tidal state, but
for tidal groups a further step was taken by projecting
significant environmental
the
variables onto the ordination space
defined by the first two discriminant
functions in order to
illustrate their relative magnitude and direction of influence in
discriminating
between groups. Accordingly,
less emphasis will be
placed on comparisons of DCA and TS analyses for tidal groups.
Twinspan
Analyses have been taken to the second (four groups) or third
(eight groups) level of division. The convention has been adopted
of referring to the groups thus defined in numerical sequence
from left to right on two-way ordered tables and top to bottom on
the corresponding
dendrograms. No correspondence
between TS groups
of similar number in different categories is implied.
On each dendrogram,
tJI~
indicator species defined by Twinspan at
each division have been shown using the shorthand identifiers
given in Table 1.4.
92
Environmental
data
A certain number of samples have been excluded from DCA
correlations with environmental data, and from HDA analysis, due
to the incompleteness
of the environmental
data set.
Correlations
between DCA axis 1 and axis 2 sample scores and
environmental
variables are summarised for the various sample
categories in Tables 1.12-1.14.
For MDA, variables with a significant univariate between-groups
0'
F-ratio for each sample category are shown in Table 1.10.
Variables with the strongest pooled between-groups
with scores on the first two discriminant
correlations
functions are given in
Table 1.11a, with the sign of the correlation indicated. The
percentage of variance accounted for by the first two functions,
and the percentage of samples correctly reclassified by the
linear discriminant solution, are given in Table 1.11b. For all HDA
analyses, samples were classified on the basis of prior
probability;
that is ,they were assigned a probability of
occurring in a given group equal to the relative size of the
group. In addition to the transformations
described above, all
variables were standardised before being entered into the
analysis, to remove any possible undue influence arising from a
difference in scale between variables.
9:3
Taxa included in analysis
Preliminary' trials indicated that the degree of association
between the three original Eurytemora categories was such that
combining them into one category produced identical results.
Ambiguous categories such as unidentified
copepod nauplii and
Acartia juveniles were excluded from the analysis and some rarer
categories such as aphroditid larvae and Acartia discaudata were
included when it became apparent that the downweighting
option
invoked in both TS and MDA analyses operated effectively on these
taxa.
Table
'·re
Vari abIes wi th a si gni f icant un ivari ate wi thi ngroup to between-group F-ratio (p( 0.05) : groups
derived by Twinspan classification
of samples within
logical categories
Variable
Category
Dist Tide Date Disch Inorg
GDl
Gp2
GD3
Gp4
Gp7
Gp8
Gp9
FA
SA
DU
KI
SK
CU
High
Low
Sprinq
Neap
HS
LS
HN
LN
*
*
*
*
~*
*
*
*
*
-
*
*
*
*
*
*
*
*
*
-
-
--
-
*
*
*
*
*
*
*
*
*
*
*
*
-
*
-
*
-
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
Drg ~org ChI a Phaeo
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
95
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
AR
*
*
*
*
*
*
*
*
*
*
*
*
Sal Temp
*
*
*
*
*
*
*
*
* *
** *
*
*
* *
* *
* *
* *
* *
* *
* *
* *
* *
Table I-ttl.
Summary of structure ma t r ices generated by Mul tipIe
Discriminant Analysis: Variables with highest poored
within-group correlations with discriminant function
scores and the sign of the correlation
Func 1
Func 2
H
Sal +
Dist +
Date +
L
Dist +
Sal +
Date +
ChI a-
Ne Sal +
Dist +
Date +
HS
Disch+
Date +
Temp-
LS ChI aAR Temp
Dist +
Date +
Sal +
~or9-
HN Sal +
Dist+
Date+
Disch+
LN Sal +
Dist+
Temp+
ChI a+
Disch+
Date+
-
Func 1
Func 2
FA
Temp +
Phae +
Disch+
~or9 +
Date -
SA
Sal +
Disch~org -
Date CHI aAR -
DU
Temp +
AR +
ChI a+
Disch-
Sal +
Tide +
Date -
KI
Temp+
AR +
Disch+
Sal +
Func 1 Func 2
GP2
Sp Sal +
Stat i oris
Dates
Tides
GP3
Dist + Temp +
Sal +
Inor +
Sal + AR Dist+
Tide+
~or9+
GP4
Sal + Dist +
Tide -
GP7
Sal +
Dist+
Inor
Phae
AR +
-
Tide+
GP8
Sal +
Dist+
Date +
Disch+
SK
Phae +
Inor +
-
CU
Date +
TempDisch+
Sal
-
Table "llb Multiple Discriminant Analysis: Percent variance accounted
for by first two discriminant functions, and percentage
successful reclassification
of samples
Variance
explained
Reclassification
tunc 1
tunc 2
Total
All samples
High tide
Low tide
Spring tide
Neap tide
High neap
Low neap
High spring
Low spring
51.4
50.6
43.2
46.2
54.8
56.0
75.0
83.8
69.0
31.3
35.1
28.5
39.0
25.0
82.7
85.7
71.7
85.2
79.8
76.2
93.6
91.6
87.8
70.3
71.1
75.0
77.3
72.7
79.6
88.6
85.5
90.9
Fallin
S.Alloa
Dunmore
Kincardine
Skinflats
Culross
63.4
58.3
50.1
49.9
77.1
26.5
35.6
34.6
41.2
16.6
5.8
89.9
93.9
84.7
90.3
85.5
81.1
91.2
100.0
100.0
Gp
Gp
Gp
Gp
Gp
Gp
3
64.1
59.3
4
7
96.8
89.4
8
89.9
83.9
2
'9
90.0
20.2
18.6
7.8
18.8
34.0
25.6
1.7
8.1
6.8
13.6
91.1
93.7
95.8
98.1
84.9
98.5
97.5
96.7
97.5
97
100.0
85.0
86.2
91.7
97.2
100.0
Table 1·12
Correlation between DCA first
and second axis sample
scores and log-transformed
environmental variables:
Samples categorised by tidal state
(Significance Levels: *= 0.05, **=0.01, ***=0.001)
Variable
Dist
Tide
Date
Disch
Xorg
Sal
Temp
Inor
Org
ChI a
AR
Phaeo
Tidal
state
High
Low
Spring
Neap
Ai A2
Ai A2
A1 A2
A1 A2
*** *
--- --*
*
*** *
***
*
*
..
-
*
*** *
***
---- ------------- ---**
**
**
**
**
*
*
**
*
**
***
**
*** ** ***
***
**
***
**
***
**
**
HS
Ai A2
LS
Ai A2
*
---- t---- ----
**
*
*
** **
**
*
*
**
***
**
HN
LN
Ai A2
Ai A~
***
**
--- ---- --- --- ---
***
**
**
***
**
***
***
*
**
**
**
Table
I"~
Correlation between DCA first and.second
scores and log-transformed environmental
Samples categorised by Station
Significance levels as Table '.1'-.
Variable
Station
Fallin
Al
Dist
Tide
Date
--*
Disch
~org
Sal
Temp **
Inor *
Drg
*
Chl a **
...
AR
Phaeo **
axis sample
variables:
A2
1----
**
S.Alloa
Dunmore
Kincardine
Skinflats
Culross
Al
Al
Al
Al
Al
A2
A2
--------- ~---~--------**
*
**
*
**
** *
**
** *
**
*
**
**
*
**
**
**
**
99
A2
------ r-----
**
*
**
A2
A2
r----- ---- ..--*
* *
**
**
**
*
*
Table 1·ILf
Correlation between DCA first and second axis sample
scores and log-transformed
environmental variables:
Samples cate90rised by Date
Si9nificance levels as Table ,'12
Sample
Variable
Gpi
Al
A2
Gp2
Ai
A2
group
Gp3
Ai
A2
Gp4
Al
A2
(chronolo9ical
Gp5
Ai
order)
Gp8
Gp7
A2
Al
A2
Ai
Gp9
A2
Al
A2
..
Dist
Tide
Date
Di !O.eh
~OT'g
Sal
Temp
InoT'
Or~
Chl a
***
**
**
--- ---- fo---- ---- ~---- ---
**
**
*
*
***
*
""**
**
**
AR
Phaeo
.'
*
**
*
*
**
**
----
***
---- r-----
**
***
*
*
*
100
---
**
***
*
*
*
*
***
*
***
**
----
**
***
*
**
*
**
---
***
**
----
**
--- ---- ---
***
**
1. Samples categorised
by station
Culross
There were a total of 20 samples in this group.
DCA anlysis indicated
that axes 1 and 2 were of equal length, a value
of 250 implying an approximately
(Gauch and Vhittaker,
correlated
75% turnover in species along each axis
1981)(Fig.1.26b).
Date was significantly
with both axes (p<0.05), while Temp and Disch were
highly correlated
«p<O.Ol)
with axis 1. AR was correlated
(p<0.05) with axis 2 onlY(Table
three levels of division,
(Fig.l.26a,
1.13). TS analysis was carried to
but only 5 groups could be derived
Table 1.15). The primary division was characterised
the indicators
Pseudocalanus
(winter/spring)
by
and Rotifera on one side
and Polvdora larvae and cirri pede larvae on the
other (summer/autumn).
At the second level, Hay samples were indicated by
terebellid,
nereid and cirripede larvae, while winter samples were
characterised
by a tight Haranzelleria
groups dominated
Eurytemora
group and two further
by Acartia spp. and Pseudocalanus.
were present in most samples.
Date, Disch, Sal, Temp had significant
univariate
Analysis
revealed
Function
1 (Table 1.11c"~ while the remaining
negatively
Oithona and
that Date was positively
associated
F-ratios
associated
(Table 1.10).
with
variables were
with Function 2. The groups were very well
101
Table
1.15
Twinspan
classification
of CUlross
(U
20 ROll
~Pf.C
OO~
2q j,1AHA
30 ;'IiAfiA
31 ~ARA
~'.IRE
'ROC
000
e.
ACAk
13 ACAj;
10 PSElJ
23 SAGl
9 ACAR
24 DIKO
5 fURY
12 ACAR
19 CALA
20 fMO
coo
000
EGGS
CLAU
001
001
CISe
ELON
E.LEG
LOi\G
--1
0
010
0101
AFfI
INfR
HELG
LONG
0110
2-11--
0110
0110
0110
0111
0111
10
10
17 OlTH SIMI
27 HARP SPE.C
10 ACAR TONS
32 LIlT CAPS
40 CIRR NAlJP
18 CENT hAMA
28 POLY elLI
33 LIlT VELI
37 lERE.· LARV
38 NERE LARV
39 CIRR CYPR
42 NUOI JUVE
1.11SIVA LARV
001
001
10
---
110
110
-12
110
---------
1110
1110
1110
III 0
III 1
1-
1--
000
onu
102
saaples
-
Cl1.03HS
C17.03HN
C2b.03HS
C02.01lHN
COb.OqHS
COl.llHS
COQ.l1HN
Clb.llHS
C27.0QHN
C23.11HN
C15.12HS
C2l.12HN .
Cl0.05HS
C17.0SHN
C2Q.OSHS
C31.0SHN
C08.07HS
C2b.08HS
C13.0QHN
C20.0QHS
MAlA
ACAR
ACA.
ACAIl
PSEU
11.0"
IlOTI SPEC
IREIl
LORG
'l'OlU
-
POLY
WIlE
PSEU
BloOM
TEltE
LAllV
CILI
-
-
ACAll
CAl
LONG
CLAU
POLY
CILI
CIlllt CYPR
CIllt RAUP
..
'
Flgure 1.26 a
i Dendrogram of Culross samples classified by Tvinspan. Indicator
: species for each division shown.
103
AXIS 2
-
.....
c;t
fI...)
UI
U1
ISl
co
~
(..11
J\.)
N
(..11
~
~
U1
UI
ISl
t'SJ
~
N
Iv
UI
N
ut
CO
c:!:J
G"'
0
\A
.9-
(.)
cri
.>
X·
.......
t'-
0
.....
.
Cj1_.
0
m
j
n
,
._.
if' •.,-,I
r~l
0
;::u
'_IfoI
,
."
J
!
0
0
t·,)
>
3
-u
0
;:;
CIt,.
e
0
v
-e-
Cl
5
r
m
Cl
;::u
_i ..,
C,
(.f)
=
;;,
0
I
-
•
-,
N
.-.
en
~
-
n
c
0
~
",0
0
..l>
0
~
0
~
CO
~
o
G"
Figure 1.26 b Decorana ordination of Culross samples on first two axes.
04
, .t,
.;-
.
.
t
..
r»
+
,..,
.,' r:
L
r»
..
,...
:
+
,
,..
~
:
p
t
!"I
Cl •
I
r-.
".
A
tl
1""".
T
,..
F
-. +
U
c"'
f""I
T
,..
C
,..,
J
-e: +
N
."
-c 2.
t
•
",
,..
..
-12
•
-'"
-4
lOS
+
4.
t-
Il.
+
GUT
separated in discriminant
space (Fig.l.26c). Date appeared to
account for most of the separation. Tide was not an influential
variable at Culross. The first two discriminant
functions
accounted for 95.8% of the variance, and 100% of samples were
correctly reclassified.
Skinflats
There were 20 samples in this group. DCA axis 1 was considerably
longer .than axis 2, indicating a greater turnover of species
.,'
along this axis(Fig.l.27b).
As with Culross, Date was correlated
with both axes. Disch was highly correlated with axis 2 (Table 1.13),
while Chla and Ar were correlated with axis 1.
TS division was carried to level 3, deriving 6 groups one of
which was a singleton sample. The primary division placed the
indicators Pseudocalanus,
Sagitta and Acartia clausi on one side
and Polydora larvae on the other(Table
1.16, Fig.l.27a).
As at
Culross, further division showed clustering by date, with group 1
characterised
exclusively
by Haranzelleria,
which was confined almost
to this group. In contrast, other polychaete larvae
were confined mostly to group 5. Samples in groups 1,2 and 3
contained,
in general. mostly neri tic/marine
Pseudocalanus
t axa
. Oi thona and
were present in most samples at Skinflats.
106
,Table 1,16
'l'winspanclassification
of Sklnflats
SK
10 ACAR
21 PARA
'"
TOI'~S
PARV
lCI\G
9 ACAR
2~ OIKO 0101
42 NUDl JUVE
q NEOioA INTE
19 CALA HElG
8 ACAR ClAlI
13 ACAR DISC
R A TROe
30 r·1A
~iARA
31
EGGS
16 PSEU ElON
23 SAGl ELEG
29 MARA ~ RE
17 OITH SI~I
39 CIRR CYPR
5 fURY AFFI
20 TEMO LONG
3Z LIlT CAPS
40 '·CIRR NAUP
12 ACAR INER
27 HARP SPEC
33 LIlT,VELI
35 SYLl" SPEC
26 o I SPEC
28 POLY elL
2 80S~ SPEC
18 CENT HAMA
37 TERE LARV
22 Cyel SPEC
30 E1EO lARV
3e. M:.F<E LARV.
ql rIVA L:lRV
0000 0
00000 o
00000 1
000001
000001
00001
00001
0001
0001
0001
0001
001
001
00
010
010
011
011
011
011
10
10
10
10
110
0
1110
110
1111
11111
11111
11111
11111
107
BaIlp1ea
J
Gll.03HS
G17.03HN
GZb.O)HS
GOb.09HS
G13.09HN 1
G20.09HS J
Gll.12HS
G03.11HS
G09.11HN
Glb.l1HS
G23.11HN
G15,12HS
G12,OlHS
G27.01HS
G03.02HN
G02.04HN G2'1.05HS
G19.01HN
GtO.05HS
G31.0SHN
POLY
CILI
pstU
SAGI
ACAR
ELON
ELEG
CLAU
POLY
HARP
CILI
SPEC
-
CIRR
EURT
pstU
CTpR
A"I
ELON
--
..
01
EURY
OITH
APrI
SIMI
S'1'LL s,eE"
IIVA
LAJlV
.'
Figure 1.27 a
Dendrogram of Skinflats samples classified by Twinspan. Indicator
;species for each division shown.
AXIS 2
")
a:.
-
~
:;,
o
.l-
0
:>X
(n
0
0
l)
_0
0
oQ
~
0
(.TI
~
':-)
';:
~I
AI
-
0
(t.1
::2.
_,.
\II
to
~
.P
,"SI
'-'
r
m
0
.'
t'"
-"J
c.n
.-.
,_,
-:c::
.....
-
0
•~
"tV
-Q
(5)
-
'.:I
In
~~
0
-
(/)
A
V\
(S)
~
CO
Cl)
tV
0
0
0
'"
CSI
0
tV
P
.".
Ci)
Figure 1.27 b Decorana ordination of Skinflats samples on first two axes.
109
o.
r.
"
IT
.
,.
•
J1&ure
-:!
-4
OUT
o
1.21 c
•
110
4
OIIT
MDA was carried out on TS groups defined at the second level of
division (four groups). Date, Disch and Temp all had significant
between-group
F-ratios (Table 1.10). Date and Disch were positively
related to Function 2, while on Function 1 all three appeared to
have equal influence (Table 1.11a). Discrimination
was again very
effective with 93.7% of variance explained by the first two
functions and 100% correctly reclassified,
but in contrast to the
DCA analysis, no influence of Chla or AR was observed.
Skinflats, like Culross, appears to be subject to seasonal rather
than semilunar influences, despite the entrainment of abundance
of eggs and larvae of some taxa already noted.
Kincardine
A total of 46 samples were taken at Kincardine.
Axis 1 represented a greater species turnover than axis 2 (Figure 1.2Bb).
Date,Sal, Temp, Chla and AR were all significantly
while Disch was signifcantly
TS classification
correlate~ with axis 1,
correlated with axis 2 (Table 1.13).
was taken to three levels (eight groups). The
main division was characterised
by Maranzelleria.
which was
confined to groups 7 and 8 (March and April), while the other
side of the division was dominated by the neritic copeopds and
meroplanktonic
larvae €,:cluding Maranzel1eria
(Table 1.17, Fig.l.28a).
Oithona and cirripede II~uplii were present in the majority of
samples, but cirripede cyprids were largely confined to group 1.
Eurytemora, Rotifera, and harpacticid
copepods occurred in almost
III
Table 1.17 Twinspan classification of Kincardine saap1es
KI
uOOO
4 tJ E 0 r· H;TE
16 PSEIJ I:LON
0000
0001
0001 !
0001
0001
0001
0010
0010
0010 0
0010 0
0010 0
0010 1
0010 1
0010 1
0010 1
·0010 1
0011
0011
10 ACAK TOl1S
1 1 ACAR 61ft
19 CALA HELG
2Ll 01KO DIOI
q
18
2~
36
38
41
13
21
33
3S
42
34
37
;S9
12
17
23
40
5
20
27
32
1
2
3
26
8
22
29
ACAR LOr-..G
CENf HAMA
POLY CILI
E1EO LARV
NERE LARV
alVA LARV
ACAR DISC
PARA PARV
LIlT VELI
SYLL SPEC
NUDI JUVE
APHR LARV
TERE LARV
IRR CYPR
ACAR INER
OITt1 SIMI
SAG.I· ELEG
CIRR NAUP
EURY AFfI
TEMO LONG
HARP SPEC
ITT CAPS
DAPH SPEC
BOS,"" SPEC
OIAP GRAC
ROTI SPEC
AC Arc CL ~lJ
eyeL SPtc
MARA ,"IRE
MARA TRee
00
01
01
01
01
-333332
10
10
10
10
110
43214i~3
212-122
4322121
110
110
110
111
1111
1111
111 1
111 1
30
31 r>1ARA EGGS
Cl111U
000111
. ".
112
K31.0SLN ·
KIO.OSHS
KO~.07HS
K13.0qHN
K17.0SHN
K21J.OSHS
K31.0SHN
K12.01HS
KI0.0SLS
KI7.0SLN
K21J.OSLS·
K06.0qHS
K03.11HS
K16.11HS
K23.11HN ·
K26.08HS ·
K27.0qlN
K13.0qlN
K20.0qlS
KOq.llLN
K23,11lN
K27.0QHN
KOQ.l1HN
K23.12HN
K27,_01lSl
Klo.11LS
K1Q.01HN!
K27.01H5'
K1S.12lS
Kll.12HS
K23.12LN
KIQ.01LN
K03.02LN
K1S.12HS
K18.12Hr~
K16.12LN
K12.01lS
K03.02HN
Kll.03HS
K17.03HN
K17.03lN
K2b.03HS
Kl1.0llS
K2b.03LS
K02.0QHN
K02.0QlN ·
TERE
CIRR
l.ARV
CiPR
-
alVA
CIRa
LARV
cypa
0
J
ROTI
SPEC
.
ACAR
'OLY
l.ONG
CIl.I
0\$0 !lIO\
AeAR Cl.AU
OITH
ACAR
IEOK
SIMI
l.onG
InTE
i
I
TEMO
·
TIMO
OITH
ACAR
LONG
MARA EGGS
MAltA WIRE
KARl. TROC
LOnG
SIMI
CLAU
MARA EGGS
KARl. WIRE
KARl. TROC
KAltA EGGS
Figure 1.28 a
Dendrogram of Kincardine samples classified
species for each division shown.
113
by Twinspan.
Indicator
AXIS 2
lSI
-
- -
-
,,_,
(.JI
"I
(D
U1
(!l
UI
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IS.)
l.n
Ut
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C)
~~
tv
N
N
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CS)
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'"to
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:>
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ID
-
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-
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t
t,lO
olI
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CJ1
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21
0
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N
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'SI
Figure 1.28 b
Decorana ordination of Kincardine samples on first two
<::$("-5,
114
•
".
..
..
..
..
..
..
·OUl
-6
X.
.
-4
-2
+
+
, ,
0
2
+
+. . . ..
4
. .+
x
b
.
•
",.
6 +
C
A
N
0
'I"
4
+
o
+
-;!
+
+
•. ';
+
+
C
A
L
,.
0
,.
,.
,.
,.
,..
t
S
C
~
I
M
J
,.
N
n
r
."
F
I.'
Il
,.
t;
,.
'.'
,.
,.
,.
•
+.
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,.
,..
-4
.+
-2
,." .. +
0
+.
DEC VAX-11/780
VMS VJ h
2
:·t·.. Kt .'
.• '.f
.:
•.. W
SlIflL.1NG
~
Figure 1.28 e
115
..
'I
'""0
all samples; Rotifera showed some bias toward the right-hand side
of the major division (Table 1.18) and thus had restricted overlap
with the neritic/lower
estuarine community.
At level 3, there was some tendency for high tides to be sorted
into group 1 and low tides into group 2. The same relationship
applied to groups 3 and 4. Groups 1 and 2 were mainly
Hay/June/September,
while groups 3 and 4 were mainly
September/November.
Groups 5 and 6 contained predominantly
samples from January and December and groups 7 and 8 from Harch
and April.
HDA was carried on level 2 TS classification.
Date, Disch, Chla
AR, Sal. and Temp all had significant between-groups
F-ratios
.'
(Table 1.10). The groups were well-discriminated
by HDA (Fig.1.28c),
with 91.1% of the variance explained by the first two
discriminant
functions and 91.2% of samples successfully
reclassified
using the linear discriminant
rule (Table 1.11b). Temp
and AR were the most strongly related variables on function 1,
and Disch and Salon
function 2. Thus, groups 3 and 4 (5,6,7,8 as
described above) were associated with lower temperature and lower
primary production
than 1 and 2 (1,2,3,4 as described above)
while groups 2 (3,4) and 3 (5,6) were associated with higher
freshwater discharge and lower salinity than groups 1
(1,2) and
4 (7,8). TS/HDA predicts that the neritic/meroplanktonic
assemblage
should be associated with higher temperatures and
higher primary production;
the division (Fig.1.28d
I,
this is the case on the left-hand side of
and MDA further cor rec t Ly summarises
116
the
distinction
between 1 (1,2) and 2 (3,4) in respect of higher
discharge and lower salinity in the latter. The biological
grouping shows
a considerable
reduction in diversity between
these sample classes, with most of the larval forms not present,
copepods such as Centropages and Temora reduced in frequency
of occurrence and species such as Pseudocalanus
and Acartia
becoming more common.
Dunmore
There were 48 samples in the Dunmore data set.
Axis 1 of the DCA ordination
(Fig.l.29b). was some 35% longer than
axis 2. There was no obvious clustering of samples, but sample 29
(D26.08HS) was an obvious outlier; only 3 taxa occurred in this
sample. Sal, Temp, Chla, AR, Tide, and Date were all correlated
with axis 1, and Sal was also significantly
correlated with axis
2 (Table 113.)3.
The major (level 1 ) division in TS was uneven (Table 1.18, Fig.
1.29a), and was defined mainly by the fidelity of A.clausi,
A.longiremis,
Polydora larvae, cirripede cyprids , Centropages,
terebellid larvae and A. bifilosa inermis to the right side
Groups 1 and 2 were pr~~ominantly
low tide samples. and mainly
117
collected in November/December/January.
characterised
association
These groups were
by high Rotifera abundance and distinguished
by the
of Diaptomus and Bosmina in group 1. Groups 3 and 4
differed from 1 and 2 by virtue of increased presence of
cirripede nauplii; group 3 was a cluster of samples containing
Maranzelleria
larvae and Littorina egg capsules, while group 4
contained an association
of Pseudocalanus,
Oithona and Temora.
Groups 3 and 4 were mostly high tide samples, and from the March
and December collections. Groups 5 and 6 displayed the highest
diversity, with a high frequency of occurrence of A.longiremis,
Polydora larvae, ~irripede cyprids, Centropages,
larvae" nereid larvae and A.bifilosa
terebellid
inermis. These two groups
~
differed little, and primarily in respect of the occurrence of
terebellid larvae. Groups 5 and 6 were, like 7 and 8 ,
predominantly
high tide samples collected in May-September.
MDA was carried out on level 2 TS classification;
therefore equivalent
to TS groups 1 and 2, and so on. Tide, Date,
Chla, AR, Disch, Sal and Temp
between-groups
MDA group 1 is
all had a significant
F-ratio; from Culross to Dunmore the number of
significant variables increased from 4 to 7. Temp, AR, Chia were
positively associated with
discriminant
function 1, and Disch
was negatively associated with this function. Sal, Tide, and Date
were negatively associated with function 2 (Table 1.11a). The
first two discriminant
functions accounted for 84.1% of the total
variance, and samples were reclassified with 81.1% success (Table
1.111.11b).
118
Fig. 1.18 Twinspan classification
of Dun.ore samples
DU
11U 12244 233:S44~
1
19139 3S3Sb7b1914735802
5 fUI-n
AFFI
27 HARjo' SPtc
32 LIlT CAPS
3e> ElEO LARV
29 MARA ~'iIKE
30 MARA. TROC·
1-----------12--1
0
0
0
0
31 ",lARA EGGS.
-----------------------------------1-------------·-1---··--------1-1---------1---
35 LIlT
VELI
SPEC
GRAC
SPEC
SPEC
SPEC
35522 '-SQ4Q4-1-4254S555
SPEC
12211
4 ~EOfl.\ INlE
8 ACAf< CLAU
10 ACAR TOf'.4S
24 OIKO 0101
9 ACAR LONG
23 SAG I ELEG
213 POLY tIll
39 CIRR CYfR
18 CENT .'t1AMA
Iq CALA HELG
21 PARA PARV
34 APHR LARV
_- __
31 TERE LAR'J
36 ~ERE LARV
42 ~..Unl JUVr::
1 DAPh
3 OIAP
15 CHYC
22 CYCL
2b ROll
2 80SM
12 ACAR
J blVA
lJ
:J:
-.-------------------------------.------- ..._----.--------- .._------------1------.--------------12-------------·-------.---------.-----.-._--_-_-.--------------------_.---------_-.- .. ...-.-
H~ER
LAR'J
C I !~I, I\AUP
r.SEl: EU;N
li
11 (lITh
~{,
Tt:·'(
1
011
00 00
100 00
100 00
100 01
100 01
100 01
100 01
100 100
100 101
100 101
10 101
10 101
10 101
10 101
10 11
10 11
--1--------------~--11---·--·-1----
101
, 1
SI jo, 1
11
11
L01>o;G
119
Ot9.0ILN
027.01LS
D03.02LN
Otl,03LS
023,t2LN 1002,04LN
Ol7,OSLN
024,OSLS
016,llLS
023,llLN
019,OlHN
031,OSLN
027.09HN
D27,09LN
D03,IILS
009.11LN
023.11HN
015.12LS D18,12LN
Ol2,OlLS
027.01HS
D03.02HN
018.12HN
D12,OlHS
OIO.05LS
Dil,03HS
017.03HN
026.C3HSJ
02~.03LS
009,11HN
D16,11HS
015,12HS
D23.12HN
Dl1,12HS
017,03LN
002.04HN
031,OSHN
008,07HS
D03.11HS
DI0,Q5HS ·
Ot7,OSHN
024.05HS
026.08HS ·
C06,09LS
D13,09LN
D20.09LS ·
D06,09HS
013,09HN
020.09HS
105M
SPEC
-
REO"
UITE
I
REOM
In'E
ROTI
..
1
SPEC
"
:..:-:a: CAPS
Taoc
. ~A'.A
I
I-
C IItR
'PSEU
ELOR
RAUP
PSEU
ELOR
MARA
TEMO
LITT
WIRE
LOIlG
VELI
..
·
ACAR
IllER
'1'ERELARV
10SM
CIR. JlAU'
POLY CILI
CERT KAMA
loCAl '1'ORS
SPEC
loCAl 'lOllS
ACAR
POLY
LOIlG
CILI
Figure 1.29 a
Dendrogram of Dunmore samples classified by Twinspan. Indicator
species for each division shown •.
120
I
\
AXIS
CSl
~
,.,J
(.JI
U1
0)
- - - -
re
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(JJ
(S)
0
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N
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t-.J
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Figure 1.29 b
Decorana ordination of Dunmore samples on first two axes.
121
".
".
ALL-GROUPS
CANONICAL
".
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A
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0
N
4
I
C
A
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...
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2
S
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til
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+ •••••••••••••••••••••••••••••
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t
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re
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DISCRIMINANT
A GROUP CENTROID
•
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c-
INDICATES
•
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t:.'
*
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-0
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0
X ••••••••• + •••••••••••••••••••••••••••••••••••••••
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Figure 1.29 e
M1>A!
122
0
2
4
+ •••••••••
•
X
OUT
The groups ordinated by MDA were less clearly defined than in
the stations considered previously, and analysis at TS level 3,
with eight groups might have been more successful. Nevertheless,
considering
the structure matrix summary (Table 1.11b), MDA predicts
that group 3 (TS 5,6) should have earlier date and higher salinity
than group 4 (TS 7,8), and this is in fact correct.
Dunmore showed an increased degree of tidally- and primary
production-associated
structure in comparison with stations
further down the estuary.
South Alloa
A total of 49 samples were taken at S.Alloa.
THe DCA axes were approximately
same amount of species turnover
equal in length, indicating the
along each axis(Fig.1.30b).
Tide,
Sal, Temp, Chla, Date and AR were strongly correlated with axis 1
and Sal also showed a degree of association with axis 2; this
pattern was similar to that at Dunmore (Table 1.13)3.
The major TS division was indicated by Neomysis and Diaptomus
versus Eurytemora,
Haranzelleria,
cirripede nauplii,Polydora,
Oithona and
but sample diversity was generally low (Table 1.19).
Group 1 was defined at level 2, and was characterised
presence of Acartia ~,
by the
whilst group 2 was characterised
by
cyclopid copepods. Group 3 contained the majority of occurrences
of Diaptomus, Daphnia al,d Bosmina. Group 4 ha~ few freshwater
taxa, but contained some Oithona, Polydora larvae and cirripede nauplii.
Group 5 was distinguished
Oithona,Polydora,
~ropages,
by the joint occurrence of
A.longiremis,
Pseudocalanus,
Temora,
and terebellid larvae, while groups 6 and 7 were
similar to each other and differed primarily in respect of higher
Eurytemora abundance in the former and the presence of Littorina
egg capsules in the latter. Groups 6 and 7 contained the majority
of Maranzelleria
summer/autumn
occurrences. Group 1 contained late
samples and mainly high tide (Fig.1.30a). Groups 2 and
3 were predominantly winter samples with no significant bias to
either high or low tides. Groups 4 and 5 were mainly May high
tide sa~ples, and were associated with the highest Eurytemora
abundances as well as the majority of neritic copepods and
meroplanktoniclarvae.
Groups 6 and 7 were exclusively March and
April samples and mainly high tide; the only samples from March
and April not included in this group were low tide.
MDA was carried out at TS level two, and the previous comments
regarding group numbering apply. Date, Disch, %org, Ar, Sal, and
Temp had significant
F-ratios (Table 1.10). Sal was positively and
Disch and %org negatively associated with function 1; Date, Chla and
AR were negatively associated with function 2 (Table 1.11a).
The first two functions accounted for 93.9% of variance, and 85.5%
of samples were correctly reclassified using the linear discriminant
rule derived by the analysis. The sample groups were clearly distiguished
U4
in
Table 1.19
Twispan claasification
sA
of South Alloa aaaples
_ 134~3q~
14 13~11?2 22
)3geb7~131568q027Sb~1
2 130sr'
LI NEO"
SPEC
rurc
6 ACAR CLAU
15 CHYU SPE.C
1 OAPH SPEC
3 DIAP GRAC
10 ACAR,lOi\S
22 Cl'eL SPEC
5 fURY AFfI
26 ROTl SPEC
27 HARP SPEC
16 PSEU ELON
39 CIRR CyPR
17 OITH SIMI
26POLY
eILI
o
o
o
9 AeAR LONG
36 ETEO LARV
38 NERE LARv
42 NUDI JUVE
16 CENT HA~A
19 CALA HELG
37 TERE LARV
o
1
1·
1
20 t€f-40 LONG
40 CIRR
30 MARA
31 MARA
33 LIlT
12 AeAR
NAUP
29
~.IRe:
MARA
TROC
EGGS
VELI
INER
32 LITT CAPS
us
)
S2b.OBHS ·
SOb.OQHS
S13.0QLN
S20.09HS ·
)SOQ.IIHtJ ·
S23,11LN
CYCL SPEC
S06.0qLS
S20.09LS
S27,09LN
S23,12LN
517,05LN ·
~18.12LN
S02.04LN
S03,11HS
S15.12HS
S15.12LS
~03.11LS
~09.11LN
Slb,llLS
Sl1,12HS
S12,OlLS
S19.01HN
DAPH SPEC
BAltP SPEC
S27,OlHS
EUltY ArrI
d27,OlLS
S03,02HN
S16,11HS
S19,OlLN
S11·~03LN !
~27, OQHN !
S23.11HN!
S03,02LN I
Sl1,03LS.J
S10,OSLS
S2Q,OSLS
~12,OlHS
S17.0SHN
S31,05LN 1------...,
S18,12HN
S06.07HS
S13,09HN
S23,12HN
m:m~}---_'CEIIT
'tORS
IIEOK lITE
DIAP GltAC
DAPH SPEC
,OSM SPEC
DIAP GRAC
ROTI SPEC
POLY CILI
BAKA
S17.03HN
S2b.03LS 1------...,
S02.04HN
.;11.03H5 ~
--'
S26.03HS
KARl. 'fROC
)
ACAR
EURY APn
CIIIR RAU.
POLY CILI
OITH SIMI
KARl. WIRE
KAltA WIRE
Figure 1.;0 a
Dendrogram of S.Alloa samples classified by Twinspan. Indicator
J
•
I ~
AXIS 2
f>_'
~
Ul
UI
.... ,3
(i'I
U1
et'
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m
UI
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(..I,
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01
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Figure 1.30 b
Decorana ordination of S.Alloa samples on first two axes.
127
ALL-GROUPS
aCATTERPLOT
CANONICAL
.,.
OUT
x
x •••••••••
-~
-a
- • INDICATES
DISCRIMINANT
-z
A 'ROUP
FUNCTION
0
CENTROID
1
2
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-
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+ ••••••••• • ••••••••• + •••••
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teDA SA TB
JI'1gure 1.)0 c
S.ALLoa..
128
0
••••••••
Z
••••••••••••••••••••••••••
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•
OUT
discriminant
space, with 2 (TS 2 and 3) and 3 (TS 4 and 5)
separated in terms of salinity and discharge
both of these distinguished
(cf Fig.l.30c), and
from group 4 (TS 6 and 7) by virtue
of the latter's earlier date and lower primary production. MDA
function 2 therefore reflects faithfully the effect of the
early-spring
peak of Maranzelleria
larval production, while
function 1 reflects other seasonal and spatial factors.
Fallin
There were 42 samples in the Fallin group.
Date,Temp,
Inor, Chla, Phaeo, AR were correlated with DCA axis
1, and Disch with axis 2 (Table 1.13). Axis length was comparable
to previous stations and their lengths implied that samples at opposite
ends of the axes would have about 25% of species in common(Fig.l.31b).
Sal was not an important variable at Fallin, but inorganic
suspended solids and phaeopigments
assumed an importance not
observed at stations further down the estuary.
TS classification
was carried to level three(Table 1.20). At
level one, the main division indicates high abundance of
Eurytemora on one side and the predominance
of freshwater species
on the other.
are almost evenly
Occurrences of Maranzelleria
divided between groups J and 8 at level three. Group 2 was
distinguished
by the presence of Daphnia, Bosmina and Diaptomus
in the relative absence of all other taxa except the common
Eurytemora, harpacticids,
and Rotifera. Groups 3 and 4 are mainly
~9
distinguished
by the clustering of most occurrences
of Neomysis
and chydorids, while groups 5 and 6 are characterised
abundances of Rotifera and Eurytemora respectively.
by high
Group 7
consisted of 4 samples containing Littorina egg capsules, and as
noted above group 8 was defined by occurrence of Maranzelleria~
High and low tides did not appear to be segregated
in any
particular manner (Fig.1.31a).
Disch, Org, %org, ChIa, Phaeo, Temp all had significant
F-ratios(Table
1.10). Temp, Phaeo were positively associated with
the first discriminant
function (Table 1.11a), Disch and % org
positively with function 2, and Date negatively with function 2.
89.9% of variance was accounted for by the first two functions
(Table 1.11b) and 90.3% were correctly reclassified.
MDA was carried
out on level two TS groups; when plotted in discriminant
space,
groups 1 (TS 1,2) and 4 (TS 7,8) were clearly separated from each
other and from the less well-separated
groups 2 (3,4) and 3
(5,6).
130
Table 1.20
Twispan classification
of Fallin saaples
FA
4 r~lOf\'
io ACAR
15 CI1YD
17 OITH
22 C~CL
1 DAPti
2 OOSft.
3 DIAP
31 ~ARA
F(
40
26 ROTI
12 ACAk
16
2B
33
38
39
~
PSEU
POLY
LITT
N£RE
RR
EUR¥
27 HARP
9 ACAk
2~ ~~ARA
32 LITT
..
'
0000000000000000
0 1111111111111111111
00000000000011111110000000000001111111
OOvOl1111tl1111110000
11000001111111
000111
000000000011100001
011110000111
oooe
0000001111
('00011
131
Fll!03HS
F17.03H N
F17.03l N
FZ6.03L SF19.01H W
F19.01l N
F27.0tL S
F03.0Zl N
FZ7.01H S
F03.02H N
Fl1.03l S
F09.1tLN
F23.1ZHN
F23.12LN
F27.09HN
F03,11HS
F09,11HN 1FJ3,09HN
F13,09LN
F20,09lS
F16.11LS
F06.09lS
F06.09HS 1
F27.09lN ~
F03,1llS
r·11.12HS I
FI8.12lN~. ---__
F12.C1HSJI
Ft2,01LS
FIO,OSHS
F17,05lN
F24,05HS
I F31.05HN
~-----'
F24.0SLS
F31, OSLN
F26,OBHS
F18.12HN
F02, OllLN
F17,OSHN
FOB.07HS
F26,03HS
F02.0~HN 1-------'
FIO.05LS
MARA
WIRE
DAPB
DIAP
BOSH
SPEC
ORAC
SPE
SPEC
ORAC
SPEC
SPEC
IUOM
IIfTE
BOSM
DIAP
DAPK
eYCL
,.
,
t-------
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CH'fD SPEC
CYCL
ACAa
e r ••
SPEC
LONG
CT'I
1-------
lOTI SPEC
,OLY
CILI
CILI
NE-let- \..I\tV
'POLY
EURr
LITT
MAlA
AF'I
CAPS
WIRE
1.12": CAPS
DAPK SPEC
MAaA WlaE
Dendrogram of Fallin samples classified by Twinspan. Indicator
species for each division shown •
•
132
AXIS 2
(1.1
cr,
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o
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...... tTJ
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-
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Decorana ordination of Fallin samples on first two axes.
lJJ
,..
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4-DEC-84
JU3U
J4
MOl. fA lS
UNIVERSITY
CLASSIFICATION
+ •••••••••••••••••••
-4
-2
Of SlI~LING
• •••••••••
0
• •••••••••••••••••••
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RESULTS·
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1)4
+ ••••••••••
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Sampling cycle (spring-neap-spring-neap
Date groups 1,5,6 and 9
successfully.
sequences)
contained too few samples to analyse
These correspond to December 1981,Ju1y 1982, August
1982, and December 1982. Analysis was completed for groups
consisting of samples from January/February
1982,Harch/April
1982, Hay 1982, September 1982 and November 1982. The summer
period was not therefore represented
in this category.
Date group 2 (Jan-Feb 1982)
This group contained 36 samples
DCA axis 1 was dominant (Fig.l.32b), with some evidence of sample
clustering. Dist, Sal, Temp, Inor Org, Phaeo were significantly
correlated with axis 1 (Table 1.14) and Phaeo was also correlated
with axis 2.
The main division at level 1 in TS was defined by the indicators
Diaptomus on one side and cirripede nauplii an the other.
Diaptomus was associated closely with Bosmina (Table 1.21) and, to
a lesser extent, with Daphnia, while the cirripede nauplii
co-occurred
to a large extent with the neritic copepods Oithona
135
Table 1.21 Twinspan classification
ot Date Croup 2 sa.plea
GP 2
110
110
10
01
01
01
o
001
001
001
000
000
..
1)6
•
r
G12. 01HS
I'OLY elLI
K12. OlHS
G27. OlHS
D12.01HS .
K19.01LN
K03.02LN
K12. OlLS
K03.02HN
S12.01HS
D12.01LS
K27.01HS
G03. 02HN G19.01HN
K19.01HN
D19.01HN
D27.01HS
D03.02HN
S12.01LS
F12.01HS
F12.01LS
S19.01HN
S27.01HS
S03.02HN
D03.02LN
S27.01LS
F03. 02HN
503. 02LN I
F03. 02LN I
D27.0ILS.J
D19.01LN
F19.01HN
F19.01LN
F27. 01HS I-K27. OILS
F27. OILS
S19.01LN
TEMO
MARA
WIRE
EURY
AFFl
....
SPEC
IIAU'
CAPS
LARV
C I R R CY PR.
EURY A,.'l
CIRR CYPR
OITH SIMI
DAPH SPEC
IIEOK.IIITE
ACAR INER
-
1
l
HARP
CIRR
LITT
alVA
LOIIG
DIAP
ACAR
GJtAC
lifER
I
i
~
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DIAP
G RAC
I).lP!! sPtC
CYCL
SPEC
...J
~~~~ :::~
Figure 1. 32 a.
Dendrogram of Group2 samples classified by Twinspan. Indicator
117
I )
AXIS 2
I\J
(S)
- - - -
UI
cs;)
U1
~
en
CD
tv
~
~
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N
N
UI
UI
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ffJ
(J)
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ro
vo~
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n
Cl
0°<..1
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V\
0
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co
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0
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l.I
r
m
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Cl
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(S)
0
0
0
-f:"
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Cl
"'0
N
0
-
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0
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(X)
0
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_0
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.J)
0
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0
...
0
c-
0
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.J)
...
N
(S)
t :
".'.'
Figure 1.:32b Decorana ordination
two axes.
of Group 2 samples on first
,i:
•
1:38
I
'
and Temora and were also associated with higher abundances of
Rotifera. Rotifera, harpacticid
abundantly
copepods and Eurytemora occurred
in all samples. At level 3, groups 1 to 4 were
predominantly
Skinflats, Dunmore and Kincardine
(Culross was not
sampled until March 1982), while 5 to 8 were biased towards
Dunmore, S.Alloa and Fallin (Fig.1.32a). Groups 5 and 6 were
mainly high tide samples, while 7 and 8 were mainly low tide
samples.
F-ratios were significant
for Dist, Inorg, Sal, and Temp(Table
1.10)
Dist was associated with function 1 and the remaining variables
with function 2 (Table 1.11a). MDA accounted for 98.1% of variance,
and 100% of samples were correctly reclassified
environmental
interpretation
data set was not sufficiently
(Table 1.11b).
The
complete to permit
in detail for this sample group.
Date Group 3 (March/April 1982)
This group consisted of 39 samples.
DCA axis 1 indicated considerably greater species turnover than
axis 2 (Fig.l.33b). Dist, Tide, %org, Sal and Temp were correlated
with axis 1, and Inorg and Org with axis 2 (Table 1.14).
Dispersion on axis 2 decreased with increasing value of axis 1
scores.
139
Eurytemora, harpacticids,Rotifera
and Maranzelleria
occurred in
almost all samples, and Littorina egg capsules and cirripede
nauplii in the majority (Table 1.22). The major level 1 TS division
was very uneven, and 7 groups were defined at level three.
Maranzelleria was the indicator for one side of the division, and
Diaptomus and Bosmina the indicators for the other. Group 1 at
level three was characterised by Sagitta, Oithona, Calanus,
Temora, Pseudocalanus
and Littorina veligers. Maranzelleria
trochophores and eggs were diagnostic of group 2, the majority of
samples in which also contained Oithona and Temora. Group 3 was
rather indifferent, containing samples with relatively few taxa
but particularly with a reduced frequency
..
and abundance of
'
Maranze11eria
eggs. Group 3 was also characterised by the highest
abundances of Eurytemora.
contained Oithona,Temora,
Pseudocalanus,
groups
The majority of samples in group 4
Acartia bifilosa inermis,
and Littorina veligers. The remaining three
contained samples devoid of neritic cope pods , with a
much reduced frequency of Maranzelleria,
cirripede nauplii and
Littorina egg capsules and with the majority of occurrences of
Daphnia, Bosmina and Diaptomus.
The dendrogram displayed considerable sorting of samples by
tide. Group 1 consisted of high spring tide samples and group 2
predominantly so. Group 3 contained mixed neap tide samples,
group 4 high neap samples, groups 5 and 6 mostly low tide samples
and group 7 low spring tide samples. As with date group 2, there
was also a clear bias ,..wards lower-estuary stations in groups
140
1-4, and to upper estuary stations in groups 5-7.
Dist, Tide, %org, Phaeo, AR and Sal all had significant
F-ratios
(Table 1.10). Sal, Dist, Tide and %org were associated with HDA
function 1 and AR with function 2 (Table 1.11a). 84.9% of the
variance was explained by the first two discriminant
and 85% of samples were correctly reclassified
functions,
(Table 1.11b). The
sorting of samples described above was well-represented
clustering in discriminant
by the
space of groups defined by level 2 TS
analysis (Fig.1.33c3 c.)
~l
Table 1.22 Twispan·classification
of Date Group :3 suples
GP 3
~122
22113331122323
33445757171374869688
..
__ ..-.--..-.....-..
•••••••••••
............
•1
••••••••••
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1·····-·····
....
11••••••• -•• 21·2
142
00000 •
00000
00000
00000
000001
000001
00001
00001
00001
;
C 11. 03H 5
G1l. 03H S
C26.03H S
G26. 03H S·
Kl1. 03H S·
K17.03HN
K26.03H 5
026.03H S
011.03H 5
Sll.03H 5
Kll. 03L S
S26.03H S
K26. 03L 5
C17.03H N
K17.03L N
C02. 04HN
,K02.04HN
K02.04LN
017.03HN
017.03LN
F26.03HS
S26.03LS
IF02.04HN
D26.031.5
D02.041.N
S02.04LN .I
S17.03HN
002. 04HN
:502.04HN
G17.03HN
G02.04HN
011. 03LS
F17.03HN
S17.03lN
,F17.03LN .
F11. 03HS.~
F26.03l5
F02. 04LN
S 11.03LS t-------'
F11. 03LS
JlSEU
CALI.
TEMO
SAGI
ELON
BELG
LONG
ELEG
TEMO
MARA
MARA
OITH
MARA
LONG
TROC
EGGS
sun
TROC
I
1
J
MAR A WIRE
EURY
APFI
CYCL
IUOM
SPEC
INTE
SPEC
SPEC
I
!
AeA .. !.!..
aD'll
BARP
DAPH
BOSM
BOSM
TEMO
S PEC
PEC
5
SPEC
LONG
Figlu'e 1.33 a
\
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AXIS 2
CS)
cD
tu
UI
U1
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- -
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0
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Figure 1.:3:3 b
Decorana ordination of Group3 samples on first tvo axes.
144
CANONICAL
OUT
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w •••••••••••••••••••
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OISCRI~INANT
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UNIVERSITY or STIRLING
+ •••••••••••••••••••••••••••••••••••••••
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DEC VAX·11/7eo
Figure 1.:)3 e
145
VMS Vl.6
•
x
X
OuT
Date group 4 (May 1982)
Group 4 contained 39 samples.
DCA axis 1 represented greater taxonomic diversity
than axis 2
(Fig.1.34b). Dist, Tide, % org, Sal, Temp, Inor, Ar and Phaeo were
correlated with axis 1; no variable was significantly
correlated
with axis 2 (Table 1.14)4.
The major TS division (Table 1.23) was characterised
Eurytemora abundance on one side and Centropages,
by high
Polydora and
terebe11id larvae and cirripede nauplii on the other.
A large
number of marine and neritic copepod species and additional
merop1anktonic
Freshwater
larvae also occurred in this latter group.
taxa were barely represented
in May samples. Samples
were classified at TS level three primarily into lower
estuary/high'tide
samples
(groups 1-4) versus upper estuary/low
tide samples (groups 5-8) (Fig.1.34a). There was a tendency for
the frequency of neap tide samples to increase from group 1 to 8.
Dist, Tide, %org, Phaeo, AR and Sal all had significant F-ratios
(Table 1.10). MDA was carried out
on TS level 2 classification
(four groups). Sal was associated with function 1, and Dist and
Tide with function 2 (Table 1.11a). 98.5% of variance was accounted
for by the first two discriminant
were correctly reclassified
functions, and 86.2% of samples
(Table 1.11b). Discrimination
on
function 2 was not good, but the correct order of groups was
recovered on function 1 (Fig.l.34c)34c.
146
Table 1.2)
TwinspaD classification
GP4
147
of Date Croup
4 eaap1es
.....
_0'1'% SPEC
K17.05HN ,
1\10.05HS
C17.05HN
C24. 05HS
C31. 05HN
CIO.05HS ;.
G24. 05HS "
K24.05HS
K31. 05HN
024.0SHS
010.0SHS
510. 05HS
S24. 05HS
031. OSHN
GI0. 05HS
G31. 05HN
I
K31.0SLN
017.0SHN
SI7.0SHN
S31. 05HN
KI0.0SLS
KI7.0SLN
K24.0SLS
024.05LS
S24.0SLS
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Decorana ordination of Group4 samples on first two axes.
149
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UNIVERSITY OF STIRLING
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CLASSIFICATION
ACTUAL
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RESULTS
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GROUP
Figure
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1.34 e
PREDICTED GROUP MEMBERSHIP
123
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OUT
Date Group 7 (September 1982)
Date group 7 contained 39 samples.
DCA ordination showed that axis 1 was considerably
stronger than
axis 2 (Fig.1.35b), with some indication of a cluster at low axis
1 scores. All variables except Chla , AR and Disch were
significantly
correlated with axis 1, and none with axis 2.
The major level 1 division in TS classification
the DCA ordination
corresponds
to
(Table 1.24), with level three groups 1,2,3 and
4 matching the cluster noted above. The pattern of tidal sorting
of samples (Fig.1.35a) corresponded
closely to that observed in Date
group 4; The frequency of marine/neritic
taxa and meroplanktonic
larvae declined from group 1 to 4, with only A.tonsa, Polydora
larvae, and cirripede nauplii being represented in groups 5~8.
Groups 5-8 were characterised
by increased frequency of
occurrence of Rotifera and the presence of freshwater taxa absent
from groups 1-4.
MDA was conducted on TS level 2 classification
(four groups).
All variables except Chla had significant F-ratios (Table 1.10).
Sal, Dist, %org and tide were associated with function 1, and
Inor, Phaeo, and AR were associated with function 2 (Table
).
97.5% of the variance was explained by the first two discriminant
functions (Table 1.11b). and 91.7% of samples were correctly
reclassifi~d
(Table 1.11b). Groups 1-4
1~
(TS 1/Z.3/4,5/6,7/8)
were
Table 1.24 Twinspan classification of Date Group 7 samples
GP7
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Dendrogram of Group7 samples classified
15) .
by Twinspan.
Indicator
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Decorana ordination of Group7 samples on first two axes.
l~
••••••••••••••••••••••••••••••
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DEC VAx-II/JaO VMS Vl.6
CLASSIFICATION RESULTS •
MDA·
Group 7
ISS
4
6
OUT
correctly ordered on the first discriminant
1/2
axis (Fig.1.35c). TS
(lower estuary, high spring tides) were separated from tS
3/4 on axis 2
by virtue of lower inorganic suspended solids,
lower phaeopigments,
and a more viable phytoplankton
population.
Date Group 8 (November 1982)
This group contained 36 samples.
DCA sample scores were more dispersed on axis 1 than on axis 2
(Fig.1.36b), with a tight cluster at an axis 1 score of about 150.
Only Dist, Tide and Sal were significantly
1, and no variables were significantly
correlated with axis
correlated with axis 2
(Table 1.14).
TS classification
was carried to level 3. Temora was the
indicator for one side of the major division, and Diaptomus and
Neomysis for the other. Neomysis was the only frequently-occurring
species in groups 4-6; freshwater taxa were only sparsely represented
in this sample set. Group 1 was notable for the co-occurrence
of five
of the 6 Acartia species present in the Forth, and also contained samples
with Oikopleura and Sagitta. Temora, Pseudocalanus,
Oithona and cirri pede
nauplii were present in almost all samples in groups 1-3. Group 2 was
distinguished
from groups 1 and 3 primarily by the absence of
1~
A.tonsa and A.bifilosa
inermis. 5amlples were again clearly
sorted by tide in the classification
high tide and Kincardine,
(Fig.l.36a);
1-3 were mainly
Culross and 5kinflats, while 4-6 were
mainly low tides and Fallin, 5.Alloa and Dunmore.
MDA was carried out on four groups defined by T5 at level 2.
Dist, Tide, Inorg, Org, Phaeo and Sal had significant
(Table 1.10). Sal and Dist were
F-ratios
associated with function 1 (Table
1.11a) and Phaeo and Inor with function 2. 96.7% of the variance
was explained by the first two discriminant
of samples were successfully
reclassified
groups were clearly distinguished
functions, and 97.2%
(Table 1.11b). The sample
in discriminant
space (Fig.l.36c)
on function 1 in terms of salinity and distance from upper
tidal limit. Little intelligible discrimination
function 2.
l~
was apparent on
Table 1.25
Twinspan classification
GPB
13 ACAR
158
of Date Group 8 aaaples
C03.. llHS
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species for each division shown •
•
15)
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Indicator
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Decorana ordination of GroupS samples on first two axes.
160
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CLASSIFICATION
RESULTS
Figure 1.:36 e
0
DEC VAX-Il/JIO
-
MD!: Group 8
161
2
VMS VJ ••
•
•
•
OUT
Tidal state (High, Low, Spring, Neap etc.)
Correlations
between DCA axis scores
and environmental
variables are summarised in Table 1.12. As with other sample
categories, axis 1 was most strongly related to most variables,
especially
to Dist, Sal, and Temp. Suspended particulate
characteristics
were most strongly related to axis scores in the
high spring tide category. On axis 2, Date was the most
consistently
correlated variable. Chla was most strongly
correlated with axis scores in the low Spring tide category.
Between 71.5% and 93.6% of variance was accounted for by the
first two discriminant
functions (Table 1.11b), and between 70.3%
and 90.9% of samples were correctly reclassified
(Table 1.11b).
Fucntion 1 invariably accounted for more variance than function
2; the difference was least in the Spring tide category and
greatest in the Low spring tide category. Of the variables
considered, only Inorg, Org, And Phaeo did not have significant
F-ratios for the majority of categories
(Table 1.10). Vith the
exception of the High spring and Low spring categories, Sal and
Dist were the most influential variables on
1.11a). Date and Disch ~ere most consistently
function 2.
MDA'
function 1 (Table
a~sociated with
Variables were projected onto discriminant space for
all-sample, high, low, spring, and neap categories; further
subdivision of tidal state did not recover additional useful
information. For the purposes of vector projection, variables
have been labelled as follows:
a
Distance from upper tidal limit
b
Date (decimal)
c
Discharge of freshwater into the estuary
d
Inorganic suspended solids
e
Organic content of suspended solids
f
~hlorophyll a
g
Phaeopigments
h
Acid ratio
i
Salinity
j
Temperature
k
Tidal state
16)
High tide
THe TS indicators for the major level 1 division were Rotifera on
one side versus Polydora, cirri pede nauplii,Temora,
Pseudocalanus
Oithona and
on the other (Table 1.26). Maranzelleria
forms
created a distinct cluster of samples which formed groups 5 and 6
at level three, as did the freshwater taxa Daphnia, Bosmina and
Diaptomus in group 8. Group 1 was a singleton with very few taxa
present except Neomysis in high abundance. Indicators
were terebel1id larvae, Centropages,
for group 3
cirripede cyprids,nereid
larvae and bivalve larvae (Fig.l.37); this group also contained
many neritic and marine taxa which occurred in other groups.
Group 4 was distinguished
from 3 primarily by the absence of
terebel1id larvae, Centropages and cirripede cyprids. Littorina
capsules and veligers were also less common in group 4 than 3;
while both groups had a high frequency of high spring tides,
group 3 was identifiable as comprising samples collected in the
spring, and group 4 was predominantly
composed of samples taken
in November and December.
MDA (Fig.l.41a, Table 1.11a) shows Date, Dist and Sal vectors most
influential, with TS groups 1, 2, 3 and 4 (at level three)
separated from 5, 6, 7 and 8 on the basis of Dist and Sal, and 3/4
separated from 5/6 on the basis of date.
1,2 and 4 showed some small positive influence of Chla, AR and
Temp, while 3 was associated with %org to a gr€~t€r extent than
l~
other groups. Group 3 indicated the association
and Temora with lower-estuary,
higher-salinity
group 4, with a strong Pseudocalanus/Oithona
of larval forms
conditions, while
association was
linked to later dates and rather lower salinity.
In contrast, Groups 5-8 were associated with a range of dates,
tending to later dates from 5 to 8 but also to lower salinities
and increased dominance of freshwater taxa. Group 8 showed little
date preference,
Groups 2 and 3
but a very strong tendency to low salinity.
were more strongly influenced
than other groups
by all variables except Date; 5 and 6 included almost all samples
from the third date group and none from any other date group and
were , as noted above, characterised
by Maranzelleria.
~
The variables of minor influence (e,f,h,j) all tended in the
same direction as Sal and Dist.
165
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167
1.37 nend rogram of High
t.Lde
samples classif ied by Twinspan.
Indicator speci es for each
division shown.
Low tide
TS and MDA analyses were carried out to division level three (eight groups).
Indicators
for the major division were cirripede nauplii and Oithona versus
freshwater chydorids and cladocera. Group 1 was composed of samples with joint
occurrence
of Maranzelleria
stages. Group 2 was rather sparse of species
and was dominated by high abundance of Eurytemora
contained a considerable
(Table 1.27). Group 3
number of larval orms (Eteone,Nereis,
bivalves,
Polydora) but differed from group 2 by virtue of high rotifer abundance.
Group 4 included the majority of occurrences
and A.tonsa, but only isolated occurrences
of cirripede nauplii, Oithona
of the larval forms common in group
3.
Groups.5-B
~
were characterised
by the almost complete absence of the neritic
cope pods and benthic larval forms, and by increasing
occurrence of freshwater
(from 5 to 8) frequency of
taxa. Chydorids and cyclopids were, however, largely
confined to group 6.
The sample sequence (Fig.1.38) showed evidence of strong selection
for date
in groups 1 and 2. Groups 3 and 4 consisted almost entirely of Kincardine
Dunmore samples and differed considerably
and
in date. Group 5 covered the same
time period as 4, but included more S.Alloa samples, while 6 again covered
the same general time period as 4 and 5 but included more Fallin samples.
Groups 7 and 8 were mainly winter samples, and 8 had the highest relative
frequency of Fallin samples.
The HDA plot showed, as with high tide samples, that 1,2,3 and 4 were clearly
separated from 5,6,7 and 8 on the basis of Dist. Disch, and Sal; the former
set of groups had higll€! salinity and the latl@1 were associated with
.
higher freshwater discharge. The effects of discharge and distance in
1~
distinguishing
4 and 5 can be seen by comparing their taxonomic composition;
group 4 (with the majority of September samples) with lower discharge and
greater distance from upper tidal limit contained more meroplanktonic
larvae,
while group 5 contained samples dominated by the more persistent upper-estuary
forms Eurytemora,Rotifera
and harpacticids.
Group 4 was also associated with
higher Chla values.
Temp and %org were influential in the same direction as Sal and Dist, but
Phaeo was more closely linked to Date and Disch. The MDA plot correctly
indicated that groups 1,2,3,7 and 8 were distiguished
A considerable
from 4,5 and 6 by Date.
amount of community structure was revealed, despite the fact
that 20 out of 88 samples were excluded from the analysis due to incompleteness
of the environmental
data set.
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Spring tide
The major division was characterised
Oithona,Temora
by Pseudocalanus,
cirripede nauplii,
and Polydora on one side and freshwater taxa on the other
(Table 1.28). TS and MDA analyses were carried out to level three (Eight groups
). Group 1 was most clearly defined by the presence of Centropages,
contained a large variety of neritic copeopds and meroplanktonic
Littorinid eggs and veligers were most faithful to groups land
2 contained a cluster of A.tonsa occurrences
but
larvae.
3, while group
but no terebellid larvae and few
Centropages
(Table 1.28). Group 3 was defined by a cluster of Haranzelleria
occurrences
and the absence of A.tonsa. Group 4 differed little from group 3.
Group 5 contained, as did group 1, a cluster of samples containing Nereis
larvae, but, with the exception of Polydora , lacked frequent occurrences
of
other larvae and contained virtually no neritic copepods.
Group 6was
defined by another cluster of Maranzelleria,
Littorina egg capsules. Group 7 was characterised
associated with
by another cluster of
samples containing A.tonsa, together with the majority of occurrences of
Neomysis; while group 8 was identified by the concentration
of occurrences
of Daphnia, Bosmina, Diaptomus and chydorids.
The dendrogam
(Fig.1.39) indicates that groups 1-4 were mainly high tide
samples, and almost exclusively
Culross. Group lwas
from Kincardine, Dunmore, Skinflats and
defined by date (May 1982), while group 2 was predominantly
composed of samples collected
in September and November-January.
The
association of these temporal groups is a feature common to all the tidal
,
categories considered
thus far.
Group 5 consisted of ~ mixture of predominantl~
included representatives
of all upper-estuary
172
autumn and winter samples,
stations but excluded Culross
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1.)9 Dendrogram
Indicator
of Spring tide samples
species
classified
for each division
174
shown.
by Twinspan.
and Skinflats. Group 6 is confined to March 1982 and mainly low tides.
Group 7 is a mixture of predominantly
autumn and winter samples, mainly low
tide and with a bias towards the upper estuary stations, and group 8
continues this trend with an increased representation
of Fallin an S.Alloa
samples.
The MDA plot (Fig.1.42a) reflects the above observations
clearly; the noted
simialrity between groups 1 and 2, and also between 3 and 4, is welldefined. The vectors correctly indicate that these two pairs should differ
most in
respect of AR, Chla, Temp and Date, but little in respect of
Sal.
Dist, %org and Sal are clearly aligned, are virtually orthogonal
to
Date an~ Disch (which are opposite to each other), and act in the opposite
.,.
direction to Inor and Phaeo. Chla is aligned with Date, and helps to
distinguish groups 5 and 7 from 3,4 and 6. Groups 3,4 and 6 are in fact all
samples taken in March 1982.
Neap tide
The pattern for neap tides was similar to that for spring tides (Table 1.29,
Fig.1.40) but with a more restricted occurrence and abundance of most
larval taxa. TS and MDA were carried out to level three division, producing
eight groups.
The major division
\.IW.'
characterised
by c i rr i nede nauplii on one side and
175
and Neomysis, Diaptomus, Bosmina, Daphnia on the other.
At level three, groups 1 and 2 contained most occurrences
of polychaete larvae.
Centropages was confined to group 2. Groups 3 and 4 contained few larvae other
than Polydora and Haranzelleria,
but Pseudocalanus,
cirripede nauplii were moderately well-represented.
Temora, Oithona and
The highest Eurytemora
and rotifer abundances occurred in group 4. Groups 5 and 6 were distinguished
by Daphnia, Bosmina and Diaptomus, with some occurrences
of Neomysis. Few
taxa were present in samples in groups 7 and 8; Neomysis was the species most
diagnostic of group 7.
The pattern of sample clustering by attribute was similar to that noted
already for other categories; groups 1-4 tended to be associated with seaward
stations, groups 5-8 with landward stations, with a similar trend for tidal
state. Sorting by date was most marked in groups 1-4.
HDA illustrated
the contrast between spring and neap tides (Fig.l.42b). The
relative magnitude and direction of influence of variables was similar
(although Date and Disch were antagonistic
on spring tides and similar on neap
tides), but neap tide group centroids were less widely dispersed; 5,6 and 7
especially differed little in location. Disch was the variable which best
distinguished
8 from 5,6, and 7, and accounts for the concentration
of
freshwater taxa in the latter.
On neap tides, the importance of distance relative to salinity increases.
%org was still aligned with these two variables, but Temp was also more aligned
with them than on spring tides. Date and Disch were, as in previous categories,
almost orthogonal
to th~ major spatial variables. Groups 1 and 3 differ
from 2 and 4 in these l~P€cts, and this separation seems to relate to Oithona
in the former pair (autumn/winter)
and high abundance of Eurytemora and Rotifera
in the latter (winter to spring).
176
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Dendrogram of Neap tide samples classified by Twinspan. Indicator
178
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Figure 1.41
Vector plots for a) high tide samples and b) low tide' samples
showing the location of the centroids of 8 Twinspan groups
ordinated in Discriminant space on the basis of sample
'environmental' attributes. Vectors indicate positive direction
and relative magnitude of influence of variables in discriminating
between groups. Vector labels indicated in text.
179
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Vector plots for a) spring tide samples and b) neap tide samples
showing the location of the centroids of 8 Twinspan groups
ordinated in Discriminant space on the basis of sample
'environmental' attributes. Vectors indicate positive direction
and relative magnitude of influence of variables in discriminating
between groups. Vector labels indicated in text.
180
8
10
Chapter 2
-
Me~ure.ents of developMent rate in E. affinis
.'
Experiments
were carried out in June and July 1983 and Harch-Hay
to measure the development
of temperatures
representative
20 deg.C, Figs.1.3-1.5).
food at approximately
of conditions
in estuary water at a range
in the Forth estuary( 8 deg.C-
Attempts were made to maintain lighting,salinity
and
natural levels.
Experiment
A preliminary
~
rates of E.affinis
1984
1 : Development
rates in mass culture
study was conducted in June-July
1983 to establish
feasibility
of culturing E.affinis
particulate
material. A variety of methods have been described
following the development
the
in estuary water on natural suspended
of copepods through successive
for
instars; these
range from the regular sacrifice of replicate cultures (Heinle 1966) to
the repeated subsampling
of a continuous
culture (Vidal, 1980a). The
latter method was adopted in the first instance in this study as offering
the expectation
Egg-sac-bearing
of a more exact relationship
between successive
samples.
female E.affinis were obtained by hand-net at S.Alloa(Fig.1.2)
Animals collected in the net were washed gently into the cod-end and decanted
into a 1 litre container. half-filled
with habitat water.Salinity
temperature during sampling were continuously
temperature-salinity
and
monitored using an MC5
bridge. A 25 1 water sample was collected when the
salinity was between 14%0 and 16%0. Temperature
deg.C during sampling.
181
was between 15 deg.C and 17.5
The copepod container was placed in a cool-box, and transferred
to the
laboratory within 1 hour of collection. Animals were sorted immediately on
arrival,in a CT romm at 15 deg.C, using a Perspex sorting device (Nival and
Nival 1970). Copepods were drawn into the observation
siphon from a reservoir;
chamber by
the chamber acted as a tap by means of which
individuals could be directed either to a collecting vessel or to waste.
200 egg-sac-bearing
female E.affinis were sorted in this manner without
sustaining visible damage.
A 1 litre sample of habitat water was filtered onto a pre-ashed,
tared,
Vhatman GF/C filter. The filter was rinsed 3 times with 10 ml distilled
water, dried at 60 deg.C for 24h and weighed after cooling in a dessicator.
Following this,the filter was ashed for 6h at 450 deg.C and re-weighed.
~
A blank filter (filtered seawater only) was subjected to the same treatment.
Dry and ash weights were corrected for blank values, and ash-free dry weight
calculated as mg/l.
The remaining 24 litres of water sample was screened through 69 um nylon
mesh to remove microzooplankton,
and held in dim light at 15 deg.C until
required.
Vidal (1980) followed development of Calanus and Pseudocalanus
in 10 1
vessels from which 10% of the population was subsampled at varying intervals.
In the present experiment, 2 litre vessels were established at 15 deg.C and
20 deg.C, using well-mixed habitat water (15 +/- 1 %0) screened and held
as described above. 15 deg.C water was allowed to acclimate to 20 deg.C for
24h before use. Initial suspended solids were estimated to be 75 mg/l.
Suspended solids levels in the culture vessels were not monitored during the
experiment.
182
On 26.6.83, 100 egg-bearing
females were added to a vessel at each temperature.
The 15 deg.C vessel was maintained
cycle adjusted to contemporary
under ambient lab. conditions
in a cooled incubator with the lighting
conditions.
The 20 deg.C vessel was maintained
and was therefore subject to less control.
Both vessels were covered to minimise salinity changes, and were topped up to
where necessary
to a marked level with distilled water before sampling took
place. Vessel contents were not mixed between sampling events, and particulate
material eventually
settled out on the bottom of the vessels.
Each vessel was sampled at intervals of approximately
at mid-afternoon.
Vessel contents were well-mixed,
one and three days,
and S% of the volume
sampled by rapidly immersing and filling a 100ml beaker. The contents of the
beaker w~re preserved by the addition of Steedman's solution to a final
concentration
by decanting
of 4%
and were subsequently
through a 100 ml polystyrene
concentrated
to a volume of 5 ml
vessel with 69 um mesh-covered
holes near the base. The sample was then counted, and specimens identified
to developmental
stage. The descriptions
by Katona (1971) were followed carefully,
of ins tar characteristics
given
to minimise the possibility
of
including ins tars of other species which may have been included as eggs in
the initial inoculum.
Following sampling,
the 24 I reservoir was thoroughly agitated and a 100 ml
sample withdrawn and added as replacement
to each culture vessel.
The experiment was terminated in each vessel when either a cohort appeared
to have reached adulthood,
or no remaining animals could be detected •.
Experiment
2
development
rates at four temperatures
and two food levels
Between 26.3.84 and 21.5.84, development
rate in E.affinis was investigated
in greater detail and under more carefully controlled
Throughout
this series of tests, regular collections
and realistic conditions.
of habitat water were
made in order to maintain a degree of coupling between the laboratory and
estuarine systems.
Development
rate was measured at four temperatures
(8,12,16 and 20 deg.C)
and at two food levels (10 and 50 mgll natural suspended particulate
material). The range of temperatures
spanned, within practical limits, the
greater.
part of the range experiences
~
annually in the Forth. Temperatures
5,10,15 and 20 deg.C would have been preferable
and permitting
of
<as spanning a wider range
a more direct comparison with the literature), ·but it was not
feasible to achieve a temperature
lower than 8 deg.C with the available
facilities.
The food levels were chosen to reflect the range typical of Forth nearshore waters under calm, slackwater
conditions
(see results of field survey).
Neither food level was expected to be limiting in terms of available energy and
nutrients;
particulate
measurements
in 1982 indicated that the organic content of suspended
material never fell below 9.7% and was commonly 20-25%. If half of
the organic content is assumed to be carbon, then a minimum of approximately
ugll would have been available initially in any treatment. This matches the
highest feeding level used by, for instance, Klein Breteler et al (1982) in
their study of the growth and development
calanoids. The nutritional
of four species of North Sea
quality of t~e suspended particulate
not known, but field observations
had shown previously
l~
material was
that the estuary could
500
support locally substantial
productivity.
populations during periods of low primary
At each temperature, cultures were replicated four times at the
higher food level and twice at the lower food level.
Adult female E.affinis with eggs in their body cavities and bearing
spermatophores
were obtained and sorted as described above, from S. Alloa
on 23.3.84. A further sample of males and females were held as stock for
re-mating. Salinity was 12.6 %0 and the temperature 8.5 deg.C.
Females were placed individually
in 5 ml of habitat water in tissue culture
plates, held in a cooled incubator at 8 deg.C, and observed daily until egg
sacs were produced. When sufficient eggs had been produced to initiate a
test at,one temperature,
the egg-bearing
females were transferred
vessels in an incubator or C.T. room at the appropriate
and the vessels allowed to equilibrate.
to 20 ml
experimental
temperature
These vessels were examined daily
at 1000 h and all nauplii present collected with a Pasteur pipette with a tip
drawn out to a diameter of approximately
24h age were used per experimental
100 um. 600 naupliiof
less than
temperature.
Each of the six vessels used to investigate naupliar development at each
temperature
diametrically
consisted of a 120 ml ploystyrene jar with 2 1cm holes cut
opposite each other 5 mm from the base. The holes were covered
with 69 urn mesh nylon net, and permitted the vessels to be gently drained and
the nauplii concentrated
observation,
into a small volume of water. This facilitated
sampling, and the changing of water.
Vhen development
to the first copepodite stage had been completed, animals
were transferred to 69 urn mesh net cylinders of approximately
18S
500 ml volume.
These cylinders were equipped with a clear polystyrene
dish) into which the copepodites
could be concentrated.
base (90 mm Petri
The cylinders were
stiffened with glass rods to enable them to stand upright in the culture
medium.
The development vessels were placed in polythene baths of 101 capacity,
which were filled to a depth of 1 cm less than the vessels' height with
with 69-um screened habitat water.
This water had been adjusted with 1.2 um
filtered river water (0 %0) to a salinity of 15 %0. Further adjustments were
made with either a filtered volume of this 15 %0 mixture or with resuspended
filter contents to produce a nominal suspended particulate concentration
of 10 or
50 mg/l. Suspended solids determinations
~
were carried out on the
source water as described previously, but the accuracy of the suspended
solids adjustment was not routinely verified.
As the vessels were placed in the tanks, they rapidly filled with water
and suspended particles. To maintain the paticulate material in suspension
and ensure circulation
a
through the vessels, each tank was equipped with
small centrifugal pump (Eheim aquarium pump, 41/min. flow rate). The
pump abstracted water from one end of the tank, and returned it via a simple
manifold to the other end. Preliminary
testing with Rose Bengal dye
introduced via a Pasteur pipette indicated that this was sufficient
to
generate gentle water flow through the development vessels and to ensure
even temperature distribution.
Expanded polystyrene
and attenuated
approximately
sheets were arranged over the vessels, and these diffused
the overhead lighting (natural light/dark cycle) to
1.2 Lux (Leakey, 1983).
The water in the baths was replaced every 48h, from stock water collected
weekly at high tide at S.Alloa and Stirling. This water was adjusted
approximately
experimental
the appropriate
day. Chlorophyll
to establish
successive
temperature.
salinity and particulate
content, and stored at
Stock water was mixed vigourously
a concentrations
to
twice per
were measured for each batch of stock water
if any increase in primary production
had occurred between
collections.
Tests were initiated by the addition of 100 nauplii to each of the 6 vessels
at a given temperature.
subjected
Two of the 50 mgll replicates
to detailed monitoring.
at each temperature were
This took place at 2-day intervals at 8 deg.C,
one-day. intervals at 12 and 16 deg.C, and 12h intervals at 20 deg.C. Test
~
vessels were gently removed from their tanks, concentrating
the animals within
into a small volume of water. In the copepodite development
vessels, a wash
bottle filled with tank water was used to gently rinse animals down the mesh
into the base dish. Animals were aspirated gently into a Pasteur pipette with
a 90 degree bend 2 cm from the tip. The pipette was connected by a length of
silicone rubber tubing to a rubber bulb which could be compressed
by a screw
clamp (Boleyn 1967). The Pasteur pipette tip was placed flat on the base of a
Petri dish containing
tank water, and the animals expressed slowly, one or two
at a time, into the dish.
Copepods in the last 2 cm of the pipette were observed at 100x magnification
and their developmental
.stage recorded with the aid of a bank of tally counters
labelled for each stage. A minimum of 50 individuals were staged on each
occasion.
The two remaining high-food replicates,
maintained
and the two low-food replicates, were
as controls for handling effects on development
1~
rate and mortality.
These vessels were only drained gently and checked at low power for the first
appearance of copepodite stages 1 and 6.
The test continued at each temperature until all specimens in a sample were
adult. C.T. room temperature and tank water temperature were recorded daily.
Egg development
rates were estimated separately. Thirty mated females were
held individually
in 2ml tissue culture well plates, and observed at 12 h
(16 and 20 deg.C) or 24h (8 deg.C) intervals. The approximate
sac production and naupliar hatching were recorded.
1M
times of egg
Results
Preliminary
experiment:
development
rates
in mass culture at
15 and 20 deg.C
The preliminary
experiment was of limited success. The 20 deg.C culture
collapsed after 14 days, following the first appearance of c4 copepodites.
Naupliar development
rates appeared relatively constant between
ins tars; the first cl individuals appeared after 5 days. Subsequently,
development proceeded to c3 in a further 4 days before stagnating during
the remaining 6 days.
Development was more complete at 15 deg.C (Fig.2.1). The culture was terminated
after 30 days, following the first appearance of adults of both sexes on
day 27. A small number of adult females were recorded on days 20 and 23,
earlier appearances of adult females were attributed
to the initial inoculum.
Individuals were widely distributed amongst stages on most days. The first
copepodites were present in samples between 6 and 9 days, with c3 copepodites
appearing on the 12th day. Total copepodite development
be approximately
twice total naupliar development
The persistence of early developmental
time.
stages throughout the experiment may
indicate that a male or males were inadvertently
included in the initial
inoculum, and that some ~f the females were subsequently
l~
time thus appeared to
re-mated.
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190
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Experiment 2 : Development
of cohorts at two food levels and
four temperatures
Primary production
Chlorophyll
a concentrations
in sample water did not exceed 4.7 ug/l
during the experiment.
Comparison of development
and survival between replicates and
food levels
The date of first appearance of cl and c6 stages was recorded for each of the
6 vessels at each of 4 temperatures
(Table 2.1). The comparisons of most
interest are a) between high-food replicates subjected to detailed monitoring
and those followed with minimal disturbance and b) between either high-food
category and the low-food replicates. A full comparison was not possible
for b) at 16 and 20 deg.C as the appearance of cl individuals was missed. It
is nevertheless
discernible
apparent that neither handling nor food level had
systematic effect on development
handling or lower food availability
rates; specifically,
any
repeated
did not impede development with
respect to animals subjected to minimal handling or fed ad lib.
Overall, survival ranged from 59% (16 deg.C, high food) to 86% (12 deg.C,
high food)(Table 2.2). Females constituted between 45% and 58% of the final
population. There was no trend in survivorship between high and low-food
treatments or between temperatures.
191
Table
1..
0
\
Day of first appearance of copepodite 1 and copepodite 6 stages
at 4 temperatures and two food levels. earliest day for
combined replicate counts underlined.
TEMPERATURE (oC)
soon
LEVEL (mg/l)
10
50
8
o'
cl
c6
cl
c6
15
33
17
35
17
33
17
35
12 .l2
12
9 21
..
10
22
22
-8 16
20
5
14
6
15
-6
14
9 23
10
-
23
14
15
5.5 13.0
- 13.0
5.5 13.5
- 14.0
!h..2
14.0
192
Table 2.2
Percent survival and percent adult female 1n replicate
trials conducted at four temperatures and two food levels
Food level
High
Low
Percent survival (% female in brackets)
8
12
16
1
76(52)
86(58)
73(54)
80(49)
2
74(48)
64(50)
63(56)
68(53)
3
60(51)
71(47)
83(53)
67(46)
4
76(55)
65(53)
59(48)
64(52)
1
66(49)
77(47)
71(52)
72(57)
2
79(45)
69(49)
75(55)
66(50)
20
.,'
In the light of the above observations it was considered justifiable
to
proceed to estimate stage durations from the combined counts of the two
more intensively-monitored
high-food replicates at each temperature.
Estimation of egg development times
,
Successful development of eggs occurred only at three temperatures
(Table 2.3); 8,16 and 20 deg.C. Mean development
8 deg.C to 0.5d at 20 deg.C. Development
time ranged from 4.7d at
time at 16 deg.C was much closer
to that at 8 deg.C than ,at 20 deg.C.
Table 2.3
Mean egg development
t,imes at 3 temperatures
Temperature
8
Mean (Days)
Range
12
4.7
(3.5-6.0)
-
16
20
4.0
0.5
(0)
(0)
193
Estimation of naupliar and copepodite development
The process of cohort development
rates
through ins tars can be considered
to
fall into the category of time-percent solutions derived in toxicoloiical
studies. In such studies, cumulative percent reaction versus time follows
a characteristically
S-shaped curve. Figure 2.3
emphasises
the applicability
of this approach to development studies. The passage of individuals
through
each instar is spread over time, with a few entering the next stage at first,
followed by the majority at an approximately
constant rate, and with a small
number completing development after a longer time interval.
Stage duration in the work reported here viII be defined as the interval
between the median time at vhich animals enter a stage and the median time
at which they enter the next stage. To estimate median development
(DT50), the cumulative percent having moulted beyond
vas plotted against time on log-probability
a particular stage
paper and a straight line
fitted to the points following Litchfield(1948).
graphically
time
The DT50 vas estimated
from this line.
Development at 8 deg. C
Development
to c6 vas complete after 45 days (Tables 2.5a-c, Fig.2.2 ), at
vhich time percent survival ranged from 60 to 79%. Females constituted between
45% and 55% of the survivors.
Naupliar stages appeared generally equal in duration, with the exception of
a protracted n2 stage. The distribution
1~
of individuals in stages broadened,
Table 2.5a Distribution in stage on each sampling
high-food replicate 1
occasion
8°C High food level, replicate 1
1
3
5
7
0
50
50
0
0
1lJ.
0
0
11
35
6
37
5
')
0
2
6
40
0
lJ.
9
15
17
.'
nl n2 nJ n4 nS no cl c2 cJ c4 cS co
50 0
4 46
,~
"-7
21
0
8
J9
0
6
0
:;
,t:
... 25
0 43
8 39
0
"
~
..,
v
"-
'
2;
2)
7
25
25
12
;6
2
27
29
31
J3
5
20
25
0
37
J9
41.
2
6 4~
J
0
35
0
0
1
16
~
'".......'"
0
0
40
10
0
Jb. 16
3
0
42
8
3~ 20
... 47
)
43
0
195
50
at 8 deg.C;
Table 2.5b Distribution in stage on each sampling occasion at 8 deg.C;
BOe Hie;h food level,
nl
1
3
.5
7
9
--
",,,,
.'
n2
re:plic;o:te 2
nJ n4 n5 n6 cl c2 cJ c4 cS c6
50 0
9 41
0
.50
0
0
27 17
7
2
.50
0
44
0
0
",-:
9
0
.50
0
0
-I
2
41
.,,...
.5
1"\
-.-'
8
15
0
,-
.~~
21
.
2'"
..J.J
'"
'"
.,.,
.,..,
~
(
.J~
0
3B
?'?
.... i
_,
"
.,,,
..:..'-
0
,~
_,r:
25
9
j-
-,
10
27
29
0
27
2;
0
0
44
6
24 26
0 44-
31
J3
0
6
0
35
37
39
41
27 1t.;. 9
J 38 9
0 42
8
13 37
43
8 42
45
0
196
.50
Table 2.Sc Distribution in stage on each sampling occasion at 8 deg.C;
percentage values from combined replicate counts
eOc
1.
J
5
7
9
11
..
n2
100
0
nJ
n4
0
0
62 )0
8
n5
n6
cl
c2
100
c6
6
8
0
19
0 ;Z
.58
10
22
0
25
27
29
c5
0
15
17
23
C) c4
14 86
o 100
82 8 4
0 10 74 16
0
0
4 90 6
0 10 82
13
_
nl
High :food level, combined counts (percentages)
0
80 20 0
50 44- 6
22 66 124 48 48
0
6 88
6
0
31.
0
".11
40
0
-'.-
:35
37
39
41
60 o
B4 16 0
61 26 12
2 80 18
0 72 28
43
18 82
8 92
45
100
197
:;
g.
Cl
g.
-..
'15
~ SO
u
1
i
....
ai
..
~
....
I
!
i
!
:;
e
~
t
I
I
g",
I
i
,;
0/
/'
L.I
5
10
15
20
25
30
35
40
Age of culture (days)
Figure 2.3 Cumulative percent of population beyond a given stage on each
sampling date at 8 deg.C, high food level. Curves fitted
by eye
and stage duration increased, in the later copepodite stages.
Naupliar stage durations, derived from the DT50 v~lues (Table 2.4), were, with
the exception noted above, of about 2.5 days. The n2 stage lasted eonsiderably
longer at 6.75 days.
Development
development
to e1 was estimated to take 18.20 days (Table 2.4), while full
( 50% moulted to c6) was estimated at slightly more than twice
this at 39.0 days.
Development
at 12 deg.C
All surviving copepods had reached adulthood after 29 days. As at
the distribution
8 deg.C,
of individuals in stages broadened with increasing age
(Fig.2.4), but the development
curves were considerably
8 deg. and the stage intervals less variable.Total
straighter
than at
median development
time for
all naupliar stages was 10.40 days, and full development was estimated to
take 26.0 days (Table 2.4). The n2 stage was again slightly longer at 2.55
days than the other naupliar stages, whieh ranged from 1.20 to 1.80 days.
Copepodite development
took longer than naupliar development
(15.6 days
versus 10.4 days). Individual stage durations ranged from 3.00 to 3.50
days.
Vith the exception noted above (n2 stage), there was no systematic variation
in naupliar instar duration or in copep~dite instar duration.
199
Table 2.4 Stage duration and median stage development time at four
temperature, high-food treatment
8°C
12°C
Stage
Duration
DT50
NI
N2
N3
N4
N5
N6
Cl
C2
C3
c4
C5
c6
2.45
6.75
1.95
2.10
2.55
2.40
4.05
4.25
4.00
4.00
4.50
0
2.45
9.20
11.15
13.25
15.80
18.20
22.!}J
26.50
30.50
Y+• .50
Duration
1.tu
2.55
1.e5
1.30
1.20
1.80
3.10
3.25
3•.50
2.75
3.00
39.00
D·r50
0
1.7J
4.25
6.10
7.40
8.60
10.4{)
13•.9J
16.75
20.25
23.00
26.00
..
..
_~o_
,,(0,....
... '"
Stage
NI
N2
N3
N4
N5
N6
Cl
C2
C3
c4
C5
c6
Duration
1.UO
1.90
0.95
0.75
0.95
1.10
1.35
2.10
1.90
2.10
2.50
cv ;;
DT(;n
_,;-
0
1.40
3.30
4.25
5.00
5·95
7.05
8.40
10.50
12.40
14.50
17.00
200
Duration
0.95
1.40
1.le
1.05
1.05
1.45
1.55
1.95
1.75
1.65
1.35
IJ!50
0
0.95
2.35
3.45
4.50
5·55
7.00
8.55
10.50
12.25
13.90
13.25
Table 2.6a Distribution in stage on each sampling occasion at 12 deg.C;
1
2
3
4
5
6
7
8
_
•.
9
10
11
12
13
14
15
'.6
,.,
-,
1M
19
20
.21
22
23
24
n1 n2 .n3 n4 nS n6 cl
'.50 0
0
7 43
0 43 7
34 16
17 33 0
2 28 20 0
0 14 31 5
0 7 43 0
0 30 20 0
0 36 14
31 16
10 30
0 37
28
17
0
c2 c3 c4 cS
0
3
10
13
22
33
ac;...
0
5
22 ,R
... 5 :39
0
25
26
27
28
29
0
6
44
6
29 21 0
19 28 3
10 31 9 0
0 26 23
1
12 3.5 .J
0 40 10
31
25
21
0
201
c6
19
2.5
29
50
1
2
3
4
5
6
7
R
9
10
11
12
13
14
..
··15
16
17
18
19
20
21
22
23
24
25
26
27
28
n1 n2 n3
:fJ 0
12 38 0
0 49 1
39 11
24 26
6 20
0 8
0
n4 n5 n6 cl c2 c3 c4 cS c6
0
24
0
J6 6 0
13 31 6
0 20 30 0
0 41 9
29 21 0
13 35 2
0 ;4 16
16 J4
8 42
,.,
v
48
0
2
18 32 C
0 40 10
38 12
23 27 0
9 )2 9
0 )) 17 0
8 37 5
3· 30' 17
0 31 19
24 26
9 41
0 50
202
Table 2.6c Distribution in stage on each sampling occasion at 12 deg.C;
1
2
3
4
5
6
7
8
9
10
11
12
13
14
.' 15
.,~
4'_·
17
lR
19
20
21
22
23
24
25
26
n1 n2 n3
100 0
19 81 0
0 92 8
73 27
41 59
8 48
0 22
0
n4 n5 n6 cl e2 c3 c4 c5 e6
0
0
67 11 0
20 74 6
0 50 .50 0
0 77 23 0
60 37 3
23 65 12
0 71 29
44
44
56
25 75
0
Q1
,.,
0
.,,
40 60 0
5 79 16
0 82 18
.52 48
28 60
10 63
0 J4
15
0
0
12
27 0
60 6
65 20
71 29
55 45
J4 66
21 79
o 100
27
28
29
20)
Lon
m
o
m
Percent survival ranged from 65 to 86, and females constituted
and 58% of the population(Table
Development
Development
2.2).
at 16 deg. C
to c6 was complete after 20 days. The development
almost linear (Fig.2.5), although, as at the lower temperatures,
duration broadened as develpoment
Median naupliar development
between 47%
progressed
(Fig.2.5,Table
curves were
stage
2.10a-c).
time was 7.05 days, and median stage duration
ranged ~from 0.75 to 1.90 days. The n2 stage lasted longer than any other
naupliar stage. There was no trend in the remaining stage duration variation.
Total median development
time was 17 days, and median copepodite development
time, at 9.95 days was again rather longer than naupliar development
time.
Median copepodite stage durations ranged from 1.35 to 2.50 days, and did not
change systematically
with increasing age.
Survival ranged between 59% and 83%, and females numbers between 48% and 56%
of the population
(Table 2.2)
205
Table 2.10a Distribution in stage on each sampling occasion at 16 deg.C;
nl n2 n3 n4 n5 n6
1
2
50
0
0 50
0
3
35 15
4
15 35
5
6
7
8
0
.,.
10
."0°.,.,
.....
,~
.....
cl c2 c3 c4 c5 c6
0
0
0
5
16
.. "
Cl
0
?C
I)
2J 30
0
0
11 39
0 50
"v
°7 43
0
0 35 15
0
.~I
.,~
.,- .,.,
., ,::;
,.. _ ... ... ...
-..;
,~
':'l
.:.;
-..;
~o
0
'1 ",0,
1.4
0 39 1.1
0
15
26 23
1
16
15 29
6
17
0 30 20
18
24 26
19
13 37
0
20
206
50
n'. n? n3 n4 n5 n6 cl
1
5J
0
2
0
50
3
4
5
6
7
8
9
10
..
-
11
.....
..l~
c2 c3 c4 c5 c6
0
;0 20
0
5 27 lE 0
0
0 7 4~
0
~
0 0 11 39 0
0
0
8 42 0
0
0
0 41
4 46 ,...
0 31 19
,...,
v
le 25
;
1.1
0
20
""
,c.
14
10
15
0
1~
16
17
lP.
19
-'_'
0
7
.,)'w'
1'\'"
C:
26
6
....,.,_ 16
0
2
32 8
0 29 21
17 ;3
8 42
0 50
10
Table 2.10c Distribution in stage on each sampling occasion at 16 deg.C;
1
2
3
4
5
6
?
8
9
10
11
12
13
·-14-
n1 n2. nJ n4 n5 n6 cl c2
100 0
o 100 0
65 35 0
20 62 18 0
0 5 23 72 0
0 0 31 69 0
0 19 81 0
0 90 10
11 89
0 66
30
0
15
16
17
18
19
20
208
c3 c4 c5 c6
0
J4
0
50 20
47 5.3
29 65
10 :fj
0 36
15
0
0
6
30
2
.,.,
0
,
""
0
.58 27
47 53
J2 68
13 87
o 100
r---------------~----------------~
~
.c:
CJ
ell
CU
"0
c::
CU
...
cu ....
0'"
bO~
CIS
.,.
-17)
m cu
>
c ...
CU ::I
....bO>u•
....
ell CU
>
"OCU
c:: .....
o
~"O
CU 0
.00
~
o.c:
c::
.......
....bO
.....::ICIS.c:..
a.U
o •
Q.bO
CU
~"O
o
........
c::
\0
CU'"
...CUCJCISIV
Q. ...
CIS
IV"O
>
....bO
... c::
CIS .... CU
.........
~
::I Cl. Cl
S S
::IeII~
Urn.o
.
an
o
o
.-
o
LIl
Development
at 20 deg. C
All surviving specimens had completed development
to c6 after 15.5 days.
Stage duration was much reduced compared to lower temperatures,
was less obvious variation
in the distribution
(Fig.2.9, Table 2.7). Median development
that at 16 deg. C. The median development
difference
of individuals
and there
within stages
time to n6 was 7.00 days, similar to
time to c6 was 15.25 days. The
between naupliar and copepodite development
times at 20 deg. C was
therefore small compared to that at lower temperatures.
Median naupliar instar duration ranged from 0.95 to 1.40 days; the n2 stage
again occupied a longer period than other stages.
0'
Median copepodite development
periods were 1.35 to 1.95, with some indication
of a trend to reducing period with increasing age.
Survivorship
was between 64% and 80%, with females constituting
between
49% and 57% (Table 2.2).
Average stage duration
Vhen median development
slope of copepodite
times were plotted against stage, the shallower
development
apparent, as is the approximately
at all temperatures
is more readily
linear nature of the naupliar and
copepodite segments (Fig.2.6). In order. to estimate mean instar durations,
210
Table 2.7 a
nl
n2 n3
c • .5
50
1.0
30 20
2.5
20°C High food level, replicate 1
n4 n5 n6
l.j,2
8
2 •.5
3.5
15
0
3.00
27
'8
5
9
.,,..,
l,f-
20
0
5.0
5.5
')
6.0
"
C •.,J"
()
Cl
5
22 28
.,....~ ~,
..1'-..I
0
1.;.
J
':,'.'7
1;)
."1
..II
",..!.
20
2,8
1
to\
v
..I
9 i.:1 C
7 26 17
0
21 29
15 35
11
39
0
0
28
22
0
20
27
3
10
"A
t:.~
0
32
12
18
'2.0
36
1Ii-
'2.5
1.7 33
;3.0
15 33
10.0
10.5
11.0
1,1. 5
,..,",.J~
c6
.,
8 1,.C
r.
o 38 20
1~
-' 30
4.5
cl} c5
s:
2. Cl
L~.O
c3
0
..I
:3.5
c2
0
45
0
cl
0
12 28 10
o
13 32
6 39
3
2
0
14.0
14.5
0
15.0
23 27
0
15·5
211
5
.50
0.,5
1.0
1.5
?O
2.5
3.0
3.,
4. o
4.5
5.0
.'
5.. s
6.0
6.5
7.0
7.5
8.0
8.5
9.0
9·5
10.0
10•.5
11.0
'1. 5
12.0
'2.5
13.0
13.5
14.0
14.5
15.0
15.5
n1 n2 n3
:fJ 0
42 8
0 5) 0
47 "':I~'
36 4
21 24
0 1.7
n4 n5 n6 cl c2 c-::
.., c4 c5 c6
0
.5
33 0
0
2 ?9 '0
-,
')
_,- 1?
7 ~,
.5 33 _t::.
,
...
0
,
...
.(
...'"
))
10 40
0
:."""
....
v
:;
7
3 31 16 0
0 14 29
7
6 31 1:;
c 28 22
26 24
0
9 41
0 17 33 0
11 32 7
4 34 12
0 40 10
36 14
le 32 0
11 36 3
7 27 16 0
0 3 24 23
0 11 39
23 27
0 50
212
Table 2.7c Distribution in stage on each sampling occasion at 20 deg.
0.5
1.0
1.5
2.0
2.5
3·0
3·5
ls.......r.
4.,5
5.C
5.5
6.0
~ ~
.'"
7.0
7.5
.
p, ~-,
- '"
o =
"_' •
.II'
9.0
9.5
10.0
10.5
11.0
11.5
12.0
12.5
13.0
13.5
14.0
14.5
'5. 0
15.5
nl n2
100 0
72 28
0 95
89
71
48
n3 n4 n5 n6 cl c2 c3 c4 c5 c6
0
5
11
29 0
42 10
9 29 62 0
0 10 6e 21
0
"
0 '37 51 12
20 63 17
0 39 61
0
2.5 71
4
10 77 13
.4 51 440 7.3 70
0
1)
~9
0
t:;7
.....
1
7
C0
49 _.-
41 59
20
0
eo
0
45 55 0
31 59 10
14 62 2l;.
0 72 28
67
41
26
19
2
0
213
33
.59 0
71 3
55 26 0
16 ~ 26
6 50 44
0 23 77
o 100
r----------------.------------------~
~
o
m
•..
o
N
'"•
N
o
o
o
U"'I
214
soC
C6
CS
Cl.
C3
C2
0
0
0
0
0
0
0
0
Cl
0
0
H6
HS
NI.
N3
N2
ell
01
tU
12°C
0
•
•
H1
•
•
0
•
•
•
•
•
•
Vl
..
'
;;
16°C
c t5
t5
[ Cl.
B Cl
ell
0
0
0
0
0
0
0
0
0
•
•
•
MS
NI.
H3
N1
0
0
~ C2
~ Cl
N6
N2
20°C
0
•
•
•
•
•
•
5
10 15 20
25 30
35
5
10
15 20 25
30
Median development time(days)
Figure 2.6 Median development times for all developmental
temperatures,high food level treatment
215
stages at four
35
the regression of stage on time was calculated separately for nauplii and
copepodites at each temperature (Table
Table 2.8
).
Regression parameters: developmental stage versus time
Nauplii
Temp.
Constant
Slope
8
-0.326
0.328
12
-0.311
16
.,'
20
r
Copepodites
Constant
Slope
r
92.8%
1.64
0.241
99.9%
0.595
97.9%
2.64
0.319
99.8%
-0.656
0.925
97.1%
3.09
0.471
99.7%
0.116
0.853
99.6%
1.83
0.593
99.4%
The slope of each regression has units of stages/day. For nauplii, there
is an increase in slope from 8-16 deg.C, with a slight decrease at 20 deg.C.
For copepodites, there is a monotonically increasing trend in slope with
increasing temperature. Comparison of slopes fornauplii
and copepodites
at any temperature emphasises that copepodites develop more slowly than
nauplii •
o
N
~
N
...-i
-
~
U
.......
0
,_,
(U
...-i
;J
.....
,_ttl
c:.
"-D
t"
'V
~
1-1
0
c.::t-
~
QI
E
..-t
<t.I
N
.'
0
\C
,...
).::t
'""
.
<t.I"O
c: QI
QlC:E..-tll"l
._.
0..0 ,....
N
OEo\
r-40P"4
QlU;>
..
QI (l)r-4
"0 41..-4
bOQI
"Onll-lCo
C:<t.I<t.I..-4
nI{I)<t.I.c:
o III
QI r-4 P:) c:
1-1..-4
0
='nI~..-4
~
0"nll-IC:nI
1-100..-4
QI~..-t
CLI
<t.I1-I
Co
S "0 nI
CLI c: =' 1-1
~nlC'CLI
41 :-
C\I
r-'
QlCLlQlCo
QI<t.I.c:
:- ..-t tJ
<t.I"O
QlObObO
0
C\i
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'-0
r-'
~
c:
,......
C
u
..._,
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r-I
CO
Q)
~
;:
r:s
rCl
;;J_
F.
'-0
.::T
'6
<II
~
.c
0.
0
'-0
( S!.'i:p) alJH
217
g
C\I
,+Uol.!JaC·L CiJI.:-O
nI
c: c:
QI..-t..-t
..-to='='
.c:U
"0"0
III
.. CLI Cl)
c:
o ..-t tJ tJ
..-t..-ttJtJ
tJ ..-4 ..-t ..-t
nI o.~ ~
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C\I
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r-I
(I)
CIS.o
Development rate as a function of temperature
Following Bottrell (1975), equations
InD.
a - b(lnT)**2
of the form
where D • development time in days
T • temperature in deg. C
a,b
are constants
were fitted to the development times for nauplii and copepodites and
for total development
(excluding egg development)
(Table 2.9,Fig.2.7a).
For purposes of comparison, power functions were also fitted (Table 2.9,
Fig.2.7b). The power function equations gave a marginally better fit for
nauplii and copepodites (Table 2.9), but
when extrapolated
appeared biologically less realistic
outside the range of experimental
temperatures. Bottrell's
equation was therefore preferred as a basis on which to predict daily growth
rates from field observations of water temperature and mean instar lengths.
218
Table 2.9
DEVELOH-.El:rI' RATE E~UA1'IONS
A) Based on Eottrell's e~uation:
1) Nau:plii lnD = J.76-0.2l5(lnT)2 1'2 = 9J.::.%
2) Cope:po~ites lnD = 3.90-0.207(lnT)2 r2 = 94.6%
J) Total
l~ = 4.53-0.2ll(lnT)2 r2 = 98.4%
B) Power function&
1) Nauplii
logl0D
=
2.23-1.10 loglO T
2
r - 94.~
2
2) Copepodites loglOD - 2.26-1.05 loglO T r - 97.l¥/o
J) Total
loglOD ..2.55-1.07 log10 T r2 ..98.1%
.,.
219
Estimation
of daily length increment from field and laboratory
observations
Using the equations
fitted above, naupliar and copepodite
were predicted at the ambient S.Alloa temperatures
development
times
on the 9 dates in 1981-2
for which detailed field length data were obtained (Table 1.7). A simplification
was introduced
by ignoring the fact that, at some times of the year, animals
would experience
a changing temperature regime.
The coefficients
of the regressions
Estimated field naupliar development
42.6d on. 12.1.82; corresponding
~
stages/day
times ranged from 7.13d on 7.7.82 to
copepodite development
were 8.76d and 49.4d respectively.
0.139-0.842
of stage on time were obtained (Table 2.8).
times for these dates
These values translated
for nauplii and 0.101-0.571
into rates of
stages/day
for copepodites
(Table 2.11).
Field observations
relating cephalothorax
showed a slightly curvilinear
be adequately
relationship,
modelled by linear regression
yielded coefficients
length to stage (Figs.2.8a-c)
but one which in all cases could
(Table 2.12). The regressions
(mm/stage) of between 0.034 (31.5.82,16.4
deg.C) and 0.049
(2.4.82,7.6 deg.C) for nauplii and between 0.076 (31.5.82) and 0.101 (11.12.81)
for copepodites.
In general, growth per .stage, and stage duration, were low at high temperatures
and high at low temperatures.
An exception to this was the sample for 20.9.82,
in which animals of all stages were unexpectedly
220
large (Fig.1.6).
Table 2.11
DEVELOPMENT TIMES AND RATES OF LENGTH INCRE!'lENT PER STAGE AND PER DAY
FOR 9 DATES IN 1982
D\~
?em;-
DEV
" .12
12.1
3.2
2.4
31.5
7.7
20.9
16.11
7.0()
10. ('3
n
o
L~2.95
4.06 28.16
7.56 17.82
16.4:1 7.99
18.00
7.13
14.25
9.42
6.08 21.32
23·12 1.83 39.71
Deve
:L~ten
'1atee
Inc:1
22.:/49.4
32.90
21..18
9.78
8.76
11.46
25.17
45.81
0.3' 5
0.139
0.213
0.337
0.751
0.842
0.637
0.281
0.151
0.222
O.CL~
C.l01
0.152
0.236
0.511
0•.57l
0.436
0.199
0.109
.'
221
lnce
DEily r,
C. '.0"1 0.0137
0.045 0.091 0.00(.3
o. oL~6 0.096 0.0097
0.049
o.oy~,
0.035
0.046
0.043
0.044
0.0166
0.076 0.026
0.079 0.029
0.100 0.0289
0.087 0.0119
0.097 0.0067
0.099
DE.i1~r
e
0.0224
0.0092
o,01~-5
0.0233
0.0388
0.0451
0.0436
0.0172
0.0106
Table 2.12
llDEC
EQUATIONS RELATING LENGTH TO STAGE
Copepodites
Nauplii
L = 0.0446 + 0.043.55 L ...-0.395 + 0.101S
2
r = 95.4 %
r2."98.2 %
12JM;
L = 0.04'.9+ 0.04538
2
r=97.8%
L = -0.261 + 0.09158
2
'r = 93.2 %
3FEB
L = 0.0448 + 0.0458S
L - -0.296 + 0.09568
2
r =97.5 %
2
r = 97.3
2APR
3ll1AY
.,' 7JUL
ut
10
L =0.025 + 0.04928
r2= 96.6 %
L = -0.36 + 0.09878
2
r = 96.7 %
L = 0.0,588+ 0.03435
r 2 - 98.8 %
L = -0.251 + 0.07615
r 2...97.0 %
L - 0.0608 + 0.0)485
r2...99.4 %
L ...-0.268 + 0.079S
r 2...98.6 %
L - -0.346 + O.lS
2
r ...91.9 %
20SEP
L ..0.0399 + 0.04558
2
r ..98.1 %
l6NOV'
L" 0.059-1+ 0.04258
2
r - 99.5 %
L ..-0.259 + 0.08658
r2_ 98.3 %
23DEC
L ..0.0.56+ 0.0444S
L ..-0.339 + 0.09735
2
r ..97.5 %
2
r - 98.8 %
222
11
DEC
.913
•
•
•
•
•
•
••
•
06
c6
1.ea
--
12 JAN
.913
i .sa
~ .re
g .Ge
r-I
~ .~a
~
o ... 13
.'
:So .3a
r-I
lIS
•
•
•
•
~ .20
Q)
t)
.re
•
•
•
•
•
•
•
•
r~
c6
n6
1.ea
3 FEB
.913
•
•
•
••
••
n6
•
•+
c6
Figure 2.8 a
Relationship
collected
between cephalothorax
length and stage in E.affinis
at S.Alloa on 11.12.81, 12.1.82 and 3.2.82
223
2
.se
I\PI<
,.....
~ .se
.._"
~ .70
g .se
r-4
a .se
.<4e
o
:So .3a
M
I'd
is. .ze
Cl)
u
.ie
.ea
•
•
••
•••
•
•
•
• f'
r
(6
n6
1.0a
31 MI\T
.se
...-.. .se
-=
.70
-'=
.se
.se
~Cl)
rot
."
a
.40
-'=
~
.3a
0
0
r-I
•2a
~Cl)
u
.re
.ee
•
•
•
•
•
•
•
•
••
•
~
~
(6
n6
1.ea
7 JUL
.se
...-..
~
~
s::
Cl)
r-I
.se
.70
.se
•~a
E
... 13
~
.ae
i
• 28
0
0
Cl)
u. .10
.ee
••
•
•
•
•
•
n6
•
•
•
~~
(6
Figure 2.8 b
Relationship between eephalothorax length and stage in E.affinis
collected at S.Alloa on 2.4.82, 31.5.82 and 7.7.82
224
I .M
2e SEP
.ge
'i
••
.ae
••
•
•
•••
•
•
•
n6
c6
1.0e
16 NOV
.se
'"'
~ .se
.....,
~
_70
53 .se
r-i
.
'
2
.~a
.,o
.c
~
.3a
•28
1 .re
Cl)
tJ
••
•
•
•
•
•
•
••
n6
c6
1.0a
23 DEC
.ge
•
•
•••
••
n6
•
•
•
c6
Figure 2.8 e
Relationship between cephalothorax length and stage in E.affinis
collected at S.Alloa on 28.9.82, 16.11.82 and 23.12.82
22.5
Daily length increment (mm/day) was estimated from the product of development
rate (stages/day) and growth rate (mm/stage) (Table 2.11, Fig.2.8). This reveale
that, although copepods achieved a greater final length in each stage at low
temperatures
(with the exception of 20.9.82), growth rates were considerably
higher in spring and summer as a consequence of the temperature-enhanced
development
rates. Highest growth rates were predicted' for 20.9.82 for
nauplii ( 0.029 mm/day), and 7.7.82 for copepodites
(0.045 mm/day). Lowest
growth rates were predicted for 12.1.82, and were, for nauplii and copepodites
respectively,
only 22% and 21% of the highest predicted rates.
226
Chapter :3
.'.
Predation b.Y Neomysis integer on Euryteaora
affinis
Predation of Neomysis on Eurytemora affinis
Between 29.9.83 and 31.10.83, a series of experiments were
conducted to investigate
could prey
the effectiveness
with which Neomysis
on Euryternora. These comprised;
a) the investigation
of the
relationship between mysid length
and predation rate
b) the generation
of a functional response for adult mysids by
determining predation rate at a range of prey densities
representative
of those observed in the Forth estuary
c) the investigation
of the effect of the absence of light and
the pre~ence of natural suspended particulate material on
predation rates in adult mysids.
All experiments were carried out at 15+1- 1 deg.c and at 10 +1-1
ppt. Vater for each experiment was made up from
S.Alloa water
collected on high tides, filtered to 1.2 um, and diluted as
necessary with fresh(O%o) Forth river water similarly filtered.
exception was the investigation
An
of the effect of suspended
particulate material on predation rate, when the suspended solids
content of the medium was adjusted to approximately
65 mg/l. All
experiments were carried out over 24h in a 12:12 LD lighting
cycle, with the exception of the test of predation rate in the
dark.Illumination,
when present, was diffused and attenuated by
expanded polystyrene
Experiments
sheets to approximately
1.2 Lux.
in b) and c) used adult mysids of between 14 and
227
17mm in length, which had been held in filtered 10 ppt estuary
water for 24 h prior to introduction
to the test vessels.Hysids
used in a) had been similarly treated, but ranged in size from
5.04 to to 19.0 mm in length. Only mysids which had not moulted
in the 24h prior to testing were used, and only one mysid was
introduced
to each test vessel.
All trials were conducted in
acid-washed glass beakers, loosely covered to minimise
evaporation
losses.
Preliminary
tests were conducted under less well-controlled
circumstances
to establish whether or not Neomysis
could capture
Eurytemora under laboratory conditions
Collection of experimental material
Copepods, mysids and water collected from S.Alloa on high tides
between 26.9 and 27.10. Additional water was collected on each
occasion from Stirling bridge. Animals were collected using a
hand-net equipped with a transparent cod-end. Following each net
sweep, the contents were washed gently into the cod-end and the
cod-end detached and decanted through a 1mm mesh (through which copepods
passed) into a polythene aspirator containing 5 litres of ambient water.
Juvenile and adult mysids retained on the mesh were washed gently into a
separate and similar container.
Inspection of the contents of
the cod-end permitted the size of the catch to be approximately
estimated.
The contents of the aspirator were placed in a cool-box and
transported to the laboratory within 1 hour of collection.
228
Pre-treatment
and exposure of mysids
a) effects of size on predation rate: 29.9.83
The contents of the aspirator were mixed thoroughly, and a
l-litre subsample rapidly taken to obtain a sample of
approximately
30 mysids of varying sizes.
Half of these animals
were gently transferred
individually with a wide-bore pipette
into beakers containing
100 ml filtered habitat water, and the
time noted. The remaining mysids were placed in 51 of unfiltered
habitat water for 24h. During this period, sufficient adult
E.affinis
for the experiment were sorted. Damaged individuals
were rejected, and
1500 intact and active specimens were placed
in groups of 100 into 15
200ml polystyrene
beakers containing
filtered 10ppt estuary water. These beakers were placed in a C.T.
room at 15 deg. C. A further 1500 adult E.affinis vere placed in
10 1 unfiltered
estuary vater (10 ppt) and held in the same room.
After 24h, mysids held in filtered vater were examined, and
active,unmou1ted
intervals,
animals were transferred
individually, at 10min.
to 21 beakers each containing 100 adult E.affinis.
The mysids held in unfiltered water were transferred
into 100 ml of filtered water.Staggering
predation
trials allowed sufficient
without affecting
individually
the initiation of the
time to terminate each trial
the duration of any other, although since all
trials vere conducted in the same C.T. room there was a
consequent
variation in the lighting history of successive
trials. During the first set of trials, the remaining 1500
E.affinis were sorted into groups of 100 and held in filtered 10
ppt estuary water.
229
Each trial was terminated 24h after initiation, by
decanting
the contents of the beaker through 1mm mesh into a
lOamI container with 69 um mesh- covered holes near the base.
Neutral red stain ( 1:200000,Dressel
the material retained, which
et a1,1972)
was added to
was stained for.30 min. and then
preserved in acidified 4% Steedman's solution and transferred
a Beatson jar for subsequent examination.
to
Staining served to
assess the degree of mortality in the prey sample, and thus to
indicate the extent to which results may have been biased by
"uncontested"
unsuccessful
capture.It was expected that mortality due to
attacks would be evident from physical damage
sustained
by such copepods during the attack.
~
Following the termination of the first set of trials, the 21
experimental
beakers were rinsed with filtered test water (10
ppt) and refilled. 100 adult E.affinis were added to each beaker,
and the second batch of mysids exposed to prey in the same manner
as were the first batch.Two- controls with 50 prey but without
predator were run in conjunction with each batch.
b) Estimation of functional response
During the period 2.10.83 to 22.10.83, trials were conducted at
densities of 10,25,50,100,200,300
and 500 prey/I. Each treatment
was replicated five times, with two controls, ,and each treatment
tested on 5 separate occasions.
Pre-treatment
of mysids was as
described above, except that individuals were briefly restrained
in a small trough and measured to the nearest millimetre.
For
each treatment, five mysids between 14 and 17mm in length were
selected and held in filtered water for 24h. Adult E.affinis were
sorted during the starvation period and added to the experimental
beaker$ just before the mysids.
~
Beaker volume differed between treatments.At
10 prey/l
5 litre
beakers were used( 50 prey per mysid). At 25 prey/litre, tests
were replicated
prey/litre
in both 5 and 2 litre beakers, and at 50
in both 2 and 1 litre beakers. The remaining prey
densities were tested in 1 litre beakers. All beakers were filled
to their nominal capacity. Hysids were exposed to prey for 24h,
and replicates were initiated at 10-minute intervals to eliminate
variation
in exposure period due to the time taken to terminate
each one.
At the end of the exposure period, the vessel contents were
decanted as described above and the contents preserved.
c) Predation
in the dark and in the presence of suspended
sediments
l4-l7mm mysids were obtained on 28.10 for the investigation
the effects of darkness and on 31.10 for the investigation
effects of the presence of suspended particulate
were measured, held in filtered water for 24h
of
of the
material. Hysids
and the tests
initiated as described above, with 5 replicates and two controls
per treatment.
Predation
in the dark was measured at 50 and 100 prey/litre
in 2
litre beakers filled to nominal capacity. Mysids were exposed at
these .concentrations
'
for 24h in a light-tight et room at 15 deg •.
C.
Predation in the presence of detritus was tested at 25,50 and
100 prey/litre,
also in 2 litre beakers. Habitat water, adjusted
to 10 ppt, was screened through a 69 um mesh, and 3 250 ml
subsamples
filtered onto tared Vhatman GF/C filters and dried for
24h at 60 deg. C.
The mean suspended solids content was 65.08
mgll, which was considered sufficiently
estuarine conditions
representative
to require no adjustment.Hysids
of
were exposed
for 24h. The beakers were not mixed during the exposure period,
and the particulate
material gradually settled out on the bottom.
Each replicate was obser.ved at intervals during the exposure
period to obtain an impression of the effect of this sedimented
layer on the behaviour on both mysids and copepods.
In both experiments,
the pre-exposure
described above.
adult E.affinis w~re,as above sorted during
period. Both sets of trials were terminated as
Sorting and preparation
The experiments
E.affinis.
of copepods
described above required a total of 13875 adult
The constraints
of time imposed insetting
up each
treatment demanded a rapid, efficient and gentle method of
sorting.
Folowing each collection,
cumulatively
concentrated
69 um-mesh-screened
subsamples
from the collection were
in a 100mI polystyrene
container with
holes near the base. Vhen several
hundred had been concentrated
into about 15 ml of estuary water,
they !ere aspirated gently into a sorting device consisting of a
Pasteur pipette with the final 2 cm bent to 120 degrees, attached
via 20cm of 3mm internal diameter polythene tubing to a 20ml
thumbwheel-controlled
pipette filler. This device permitted
precise control of the movement of water into and out of the
pipette tip.
The pipette tip was transferred
to a 90mm Petri dish which had a
divider glued in place acros the diameter. The final two 2cm of
the tip was placed flat on the bottom of the Petri dish, below
the surface of filtered estuary water, and the position of the
dish adjusted so that the
2cm section was within the field of
view of a Vild M5 stereo dissection microscope.
The refractive
index of the water and of the pipette glass were sufficiently
close that the contents of the pipette tip could be resolved
clearly enough to identify individual cppepods to species and to
stage. The contents of the device were expressed slowly into one
half of the dish.Vhen an adult male or female Eurytemora
approached
manipulated
the end of the pipette, the pipette filler was
until the individual was at the pipette end. The
pipette was raised and the individual ejected. into a drop at
the pipette tip, which was then dipped gently and briefly into
the water in the other half of the Petri dish.The pipette was
transferred back to the original side of the division, and the
sorting process continued; unwanted copepods were expressed
directly without manipulation.Given
the largely monospecific
nature of the samples from S.Alloa, it proved possible to sort in
excess, .of 1000 adult E.affinis per hour
Cope pods were transferred in groups of 25,50 or 100 into 100ml
beakers of filtered 10 ppt estuary water and held in dim light at
15 deg. C until required. For each set of trials, an additional
25 copepods were held in filtered water; these animals were used
to replace
any which died or became moribund during the holding
period and ensured that only active, apparently healthy specimens
were used as prey.
Evaluation
of results
The preserved specimens from each trial replicate were
concentrated
into 2ml of fresh 4% Steedman's solution made up
with 10 ppt filtered estuary water. This volume was pipetted into
a Bogarov counting tray, and the copepods counted. Each was
examined for signs of damage, and any which had not taken up the
Neutral red stain (and were therefore dead at the end of tbe
trial) were noted.
Small mysids were placed in tbe Bogarov tray and tbe length from
rostrum tip to telson end measured with a calibrated eyepiece
graticule
.'
to O.lmm. Larger mysids were placed on a piece of graph
paper and their length similarly measured to 0.5mm.
Faecal pellets produced by mysids during trials were examined and
their contents qualitatively
assessed.
The effect of mysid size on predation rate was evaluated
regression
of rate in prey/mysid/day
For the remaining experiments,
on mysid length in mm.
feeding rate was calculated as
clearance rate (Fulton,1982a;Frost,1972,
1975; Cooper and
Goldman, 1980)
(equation 1)
Cr -( -In(Nt/No)V)/(NpT).
Vbere
(lIh)
Cr - clearance rate
No - ini tial(control)
number of prey
Nt _ number of prey at time T
V
from tbe
- volume of experimental
(b)
vessel
Np - number of predators
23.5
(1)
The relationship
between clearance rate and prey density was
tested by analysis of covariance
rates were normalised
by log(x+l) transformation,
were found to be heterogeneous
partitioned
(ANCOVA) in which clearance
by Bartlett's
as the variances
test. The model
variance between prey density and.blocks(days),
with
mysid length as a covariate.
Predation rate was calculated as:
(equation 2)
P • (No-Nt)/TNp
Although Frost(1972),Landry(198l)
Butler(1986),inter
and Villiamson
alia, have suggested the use of mean, rather
than initial, prey density in the representation
relationships,Harin
of predation or clearance rate. It is
to determine a functional response based on mean
density.Further,the
replication
of predation
et al (1986) have pointed out that mean prey
density is not independent
thus incorrect
and
use of mean prey density invalidates
the
of treatments and means that there is no independent
estimate of error in clearance rate for any treatment.
Clearance rate was also calculated following Harin
et al(1986) as
Cr.
V«Co-Ct)/Co)
l/h
(equation 3)
where V • vessel volume .
Co,Ct • initial(control)
and final prey
densities respectively
This equation is appropriate when clearance rate is not constant
but declines with increasing food concentration.
Ingestion rate may be calculated from equation 3 simply as
I • CrCo
231
Results
Relationship
between mysid size and predation rate
All copepods recovered from experimental vessels in this experiment
stained strongly with Neutral red, and were presumed to have been alive
and active throughout the experimental
period~ Two mysids moulted during
the exposure period, and these results were discarded.
,A
linear relationship
adequately described (r-0.93, p<O.Ol) the observed
increase in predation rate (copepods/mysid/day)
with mysid length (Fig.3.1,
Table 2.1). Since neither variable was measured without error, and the
size distribution
of mysids must be regarded as truncated, a functional
or geometric mean regression was calculated
(Ricker, 1973):
y- -11.59 + 2.60x
where y
x
is predation rate (copepods/mysid/day)
is mysid length (mm)
The regression line intercepted the x-axis at a mysid length of 4.4Smm.
Hysids below this length did not appear to be able to feed effectively on
adult Eurytemora.
Predation rates for mysids in the 14-17mm
experiments
size range used in subsequent
clustered between 25 and 33'copepods/day
at a prey density of
50/litre. The highest rates were observed in the two largest mysids, which
captured 39 and 40 prey respectively during the 24h exposure period. This
\
.......
,
so
r c 0.934
, Y - -11.59 + 2.60x
......
't1
...... •
t
't1
• .-4
III
•
40
.
>.
e
III
'8Pt
Cl>
Pt
30
0
.....,
(J
..
'
Q)
~
tl 20
.0..
-""
~
re
't:i
Cl>
t;
10
•
4
8
Mysid
12
16
20
length (mm)
Functional regression of mysid length on predation rate;
correlation coefficient for least-squares included as an
approximate indication of the strength of the relationship
Table3.' The relationship between mysid length and predation
rate at a prey density of 50 prey per litre
No. prey
taken
No. prey
moribund
No. prey
recovered
Control
recovery
5.04
3.82
4.60
4.39
6.60
9.00
7.92
7.48
10.51
9.07
6.77
8.64
46
48
51
49
46
40
36
41
37
36
45
35
(51,50)
4
2
0
1
4
10
14
9
13
14
5
15
0
0
0
0
0
0
0
0
0
0
0
0
14.50
12.50
16.50
13.00
19.00
17.00
13.00
14.00
13.00
13.50··.
28
32
27
37
21
10
29
22
26
28
(50,50)
22
18
23
13
29
40
21
28
24
22
0
0
0
0
0
0
0
0
0
0
Hysid
length(mm)
241
was equivalent
Comparison
to a clearance rate of 1.6 l/mysid/day.
of the mysid size range used here with the data of Astthorsson
and Ralph (1984) indicate that the greater part of the normal size spectrum
of Scottish Neomysis
certainly
(-3mm to >16mm) was sampled, but that the sample almost
contained representatives
of two generations.
are likely to have belonged to the summer generation,
«10mm)
The larger mysids
while the smaller
mysids will have been members of the autumn-born
overwintering
generation.
Functional
response: the relationship
between prey density and
predation rate
In all replicates,
uneaten copepods stained strongly with Neutral red,
and, as above, it has been assumed that all constituted
normally-available
prey
up to the end of the" exposure period. None of the 35 mysids used moulted
during exposure; Assthorsson
and Ralph (1984) reported that the intermoult
period for mature mysids is between 12 and 18 days, so moulting would not
be expected
to be a frequent occurrence.
Mysid faecal pellets recovered from experimental
exception
loosely-packed
vessels were without
and contained identifiable
copepod remains.
Mean raw predation rate ranged from 24.2 copepods/day
127.4 copepods/day
monotonically
at 500 prey/l (Table" 3.2,Fig.3.2).
at 10 prey/l to
Predation rate increased
with prey density up to 11S/day at 200 prey/I, declined slightly
at 300 prey/I, and increased to 127/day at a density of 500/1.
242
,
'"
r-t
I
tU
.
Functional response, uncorrected for
differences in vessel volume
150
r-I
I
tU
.'
.~
en
>,
El
•
en
tU
0
100
Pt
Q)
Pt
0
0
'-'"
Q)
+'
ro
1-1
50
r:::
.~
0
+'
ro
reQ)
~
100
200
400
JOO
.
500
1
Prey density (copepods.l- )
Figure 3.2
Table
~.2.
Prey
density
10
Mysid predation on E. affinis at a range of prey
densities; number of prey recovered,number of
prey taken, and predator length.
Mysid
Vessel
volume(l) length(mm)
16.0
16.5
5
16.0
15.5
16.0
2
25
5
25
1
50
No. prey
recovered
30
27
25
22
25
16.5
15.0
15.5
17.0
16.0
31
30
33
27
34
16.5
15.5
17.0
16.5
15.0
93
89
87
92
79
16.5
14.0
14.5
14.5
15.0
8
30
21
22
26
Control
recovery
(52,49)
No. prey
taken
20
23
25
28
25
No.
dead
0
0
0
0
0
19
20
17
23
16
0
0
0
0
0
31
35
37
32
45
0
0
0
0
0
42
20
29
28
24
0
0
0
0
0
55
55
76
53
51
45
(100,100) 45
24
47
49
0
0
0
0
0
54
57
54
37
48
46
(100,100) 43
46
63
52
0
0
0
0
0
120
112
118
110
115
0
0
0
0
'0
112
106
115
114
110
0
0
0
0
0
150
124
113
132
118
0
0
0
0
0
(50,50)
(123,124)
(50,51)
,,-
50
2
15.0
15.5
15.5
16.0
15.0
100
1
14.0
14.5
15.0
15.0
14.5
200
1
15.5
16.5
16.0
16.5
16.0
80
88
82
90
85
15.5
14.0
16.0
14.5
14.5
189
195
186
187
191
14.5
14.0
14.0
14.0
13.5
349
375
386
367
381
300
500
1
1
r
(201,199)
(300,302)
(497,501)
Table;·3 Clearance rate of mysids feeding at a range of
prey densities; rates calculated using models
which assume a) constant clearance rate with
declining prey density and b) decreasing clearance
rate with increasing prey density.Values are mean
llh ,95% confidence intervals in brackets
10
25
Prey density
50
100
200
300
500
a)
5
3.38
(0.65)
Vessel volume
5
2
2
'1
1
1
1
1
1.70
0.96
1.11
0.95
0.70
0.86
0.46
0.30
(0.37) (0.26) (0.36) (0.59) (0.21) (0.05) (0.03) (0.05)
b)
2.42
(0.33)
1.48
0.76
0.84
0.57
0.50
0.54
0.37
0.25
(0.22) (0.13) (0.23) (0.20) (0.09) (0.08) (0.01) (0.02)
Table
Mean predation rates of mysids feeding at a range of prey
densities(copepods/mysid/day);
a) actual numbers taken in 24h
b) numbers corrected to unit vessel volume
10
..
100
50
25
a)
5
24.2
(3.4)
5
36.0
(6.4)
'Vessel volume
1
2
2
28.6
42.0
19.0
(3.1) (11.7) (9.5)
b)
4.8
(0.7)
7.2
(1.3)
9.5
(1.6)
'
21.0
(5.9)
"
245
200
300
500
1
1
1
1
50.0 115.0 111.0 127.4
(.92) (4.7) (4.1) (16.6)
"
"
"
"
4
OJ./
-.c:
3
~
0
5-~
•
'2..-
~
~~
\-I!a:..
0
G>
+'
~
2
G>
0
f
S
~
1
tJ
..
t
1
1
Q
.,
1025 SO
100
200
Prey
300
400
500
density (copepoda/litre)
.'
Figure ,.,
The relationship between clearance
prey density (no./l) in vessels of
a) clearance rate calculated using
b) clearance rate calculated using
I(1986)
rate (l/mysid/day) and
different volumes
the equation of Frost (1972)
the equation of Marin et al
o
o
o
..-0
.-4
t:IO
0
r-4
'"sRI::
...41as
.0'
r"
0
0
..--
r
w
"
01
41
U
V
=RwI
t
RI
41
U 41
".
r-4
~
041
>.
.-4
t:IO>.
0.0
~
r-4
=41'"'"41
0
..--
...:. ~....
41'"
41
.041
=
......c::0. ....
~
.
(I)
s::>.
.::t-
n•
or--------~---;,,-------J.. ....-
..--
..
o
..--
i
~
o ...
........
"'(1)
RI
s::
(II'"
w
r-4(11
G1~
..c::w
~o.
,
247
Clearance rates were calculated using both the conventional
expression
exponential
(e.g. Frost, 1972) and that of Marin et al (1987) (Table 3.3, Figs.
3.3a,b). Rates based on the latter equation were consistently
lower at
any given prey density than those based on the former; on both bases, however,
rates exhibited a declining curvilinear
characteristic
relationship
(Fulton 1982) of a Type II functional
The curvilinear
nature of the relationship
prey density is emphasised
with prey density,
response (Holling, 1965).
between clearance rate and
if 10g10 clearance rate is plotted against prey
density (Fig.3.4).
Because the experiments
were carried out on a number of days, and because
mysid length differed between treatments, analysis of covariance was carried
out to a) determine whether either of these covariates had a significant
~
effect on the results and b) to correct the estimates of clearance for any
effects that were detected. The analysis was conducted on log(x+l)-transfromed
data because of the obvious dependence of the variance on the mean (Fig.3.3).
Neither day nor mysid length had a significant
only the treatment effect was significant
effect on the results, and
(Table 3.4). In the light of this
result, clearance rates were not adjusted.
At 25 prey/I, clearance rates were significantly
higher (p<O.Ol, t-test)
in 5 litre vessels than in 2-litre vessels. At 50 prey/litre,
differences
statistically
independent
however,
between clearance rate in 2-litre and 1-litre vessels were not
significant.
( p>0.05). Clearance rates thus appeared to be
of vessel volume at prey densities
above 50 prey/litre,
mysid foraging activity appeared at lower densities
while
to be greater in larger
vessels than in smaller vessels.
Under circumstances
Fulton(1982)
where clearance rate is independent
of vessel volume,
has pointed out that prey density will decline faster in smaller
vessels; consequently,
predation rate will be higher, for a given clearance
rate, in larger vessels. This observation has been used, following Fulton (1982)
to generate a functional response standardised
The results of standardisation
to unit volume (Figs.3.5,3.6).
( computing predation rate by holding v • 1
in the equations of Frost (1972) and Marin et al 1987) are indicated in
Table 3.3: values for larger vessels are substantially
between different-sized
reduced, and differences
vessels at 25 and 50 prey/litre also much reduced.
The functional response was modelled in two ways. Firstly, a rectilinear
model, frequently applied to invertebrate predators (Cooper and Goldman 1979),
fitted by eye (Fig.3.5). A chi-squared goodness-of-fit
test yielded a value
of 8.64 (p<O.Ol). Secondly, an attempt was made to fit Bolling's
(1965) disc
equation (Fig.3.6):
~
Na
•
aNT
1+ aNTh
where
Na
• no. of prey eaten
a
• instantaneous
N
• no. of prey available
Th
T
coefficient of attack
• handling time (d)
• duration of experiment (d)
This yielded an estimate of 0.873 for a and of 6.97 minutes for Th. The
curve corresponding
to this solution was generated by a computer plotting
routine (GINOGRAF) (Fig.3.6). The chi-squared value was 8.06, indicating an
adequate fit. The 'disc' model suggests that predation rate will increase
at a decreasing rate up to prey densities in excess of 1000/litre; this
..
.,w
8
•
.~
LIl
~
.~
Ct-t
.
8
J::::l
~
Q
b.!l
.~
-g
0
•
Cl)
•
<:>
rn
a.>
0
C>
•.-1
N
~ §
U)
Q
0
0
'-'
~
~
..-c
~ ~S>
a.>
0
CIl
Q
<l>
e:
0
U)
..-c
,.-I
r-4
0
Ct-!
I
r-i
P4
<l>
Pi
0
Ct-t
-.
.-I
"cd
~
a.>
f-I
Pot
0
~
..-c
Cl)
0
0
~
(J)
8.
•.-1
~
~
ro
(J)
r;;l
s
(J)
•.-1
~
CIl
~
~
0
'"
•
.-, " n
§
~
Cl)
0
LIl
.-
0
C>
,
0
LIl
.-
(I_P.I_P1SAw.spoaaaoo) a~~~ UOl~~pa~
i
.:
,
2.50
2.51
Table;·5
Mean clearance rates ( llh and 95% confidence intervals)
of mysids feeding at 50 and 100 prey/litre in the dark
and at 25,50 and 100 prey/litre in the presence of detritus
Clearance rates calculated using exponential equation.
Prey
density
Clearance
rate(l/h)
95%
C.l.
.50
1.28
0.39
100
1.41
0.20
25
0.98
0.17
50
0.76
0.18
100
0.67
0.10
Dark
Detritus
Table3·~
Analysis of covariance of log(x + 1) -transformed
. clearance rates measured at a range of prey
densities; variance partitioned by prey level
and blocks(days) with predator length as a covariate.
Source of variation
df
Mean square
F
Residual
Regression
(length on rate)
Constant
Prey density
Blocks(days)
23
1
0.00163
0.00668
4.09
1
6
0.02310
0.10822
0.00187
14.16
66.37
1.15
.~
4
2.52
Significance
NS
0.01
0.001
NS
is approximately
the maximum abundance observed in UK estuaries
(Burki11,
1982) and is almost double the highest abundance recorded for Eurytemora
in the Forth (Roddie, 1980). At 1000 prey/litre,
to be 167 copepods/day,and
this may be considered
predation rate is predicted
the effective maximum
rate likely to occur in the upper Forth estuary. It should be noted that in
fitting the above curves, predation rates at 25 and 50 prey/litre vere those
derived from experiments
in the larger of the two vessel sizes used.
Predation in the presence of detrital material
No mortality vas observed amongst unpredated
cope pods in this experiment.
Kysid faecal material recovered from the experimental
colour, and contained considerable
vessels was dark in
amounts of compacted material. By virtue
of the contrast between these faeces and those produced in the previous
experiment,
it vas inferred that the compacted material was of detrital
origin. The faeces also contained identifiable
Since the vessels were unstirred
copepod remains.
throughout the exposure period, the mysids
had clearly spent some of their time foraging on the vessel bottoms. This
inference vas supported by direct observation;
was not constructed,
apportioned
although a formal time budget
intermittent viewing indicated that time vas
between bottom-feeding
and active swimming. It was also observed
that a variable proportion of copepods vere located at the sediment surface
whenever viewing took place, so it is not clear whether or not ingestion of
detritus vas simply a consequence
of foraging for these animals.
Predation rates (Table 3.7) and clearance rate (Table 3.6) were similar to
those at the same prey densities in the absence of detritus. Predation ranged
from a minimum of 15/day at 25/litre to a maximum of 54/day at 100 prey/litre.
)
)
'"'
M
''d
M
•
.-.-
''d
...t
co
t-•
M
....,
O-
2
CD
~
CD
c
1
"'-1
S litre
vessel
2 litre vessel
1 litre vessel
I
.J
,!
25
SO
~
CD
2
.-
0-
b)
24h clark
12.12 LD
.-
2
0-
f
1
I
1
I
M
0
c..)
6S mg.1 -1 detritus
1.2 um filtered
!
.'
SO.
. 100
0
SO
Prey density(eopepods.1-1)
)
Figure
'J
)
'.7
The effects of vessel size (a), absence of light (b) and
presence of detritus (c) on the clearance rate of N.integer
feeding on E.affinis
100
Mean clearance rate in the presence of detritus was compared statistically
at each prey density with clearance rate in filtered seawater. F-ratios
for each pair indicated that variances were homogeneous,
performed on untransformed
so t-tests were
values. At no density were significant differences
apparent (p> 0.05, Fig.3.7c). Foraging on copepods did not therefore seem to
be affected by the presence of a potential al~ernative
food source.
Predation in the absence of light
As with the experiments
reported above, no copepod mortality
other than
directly due to predation was observed. Predation rates at 50 prey/litre
ranged form 21-41/day, and at 100 prey/litre from 70 to 83 prey/day (Table 3.6).
Clearance rates (Table 3.5, based on equation (3»
had a mean value of
1.28 ltd at 50 prey/litre and 1.41 ltd at 100 prey/litre;
was not statistically
significant
(p>0.05, Fig.3.7b). Clearance rate at each
density was compared with clearance rate
lighting conditions
at 50 prey/litre
therefore apppeared
at the same density under normal
(Fig.3.7b). Vhilst there was no significant difference
(p>0.05), a statistically
at 100 prey/litre
this difference
significant difference did exist
(Fig.3.7b, p>O.OOl). At higher prey density, foraging
to be more effective in the dark.
255
Table
Prey
Density
!.~Mysid
predation on E.affinis in the absence of light
Mysid
Vessel
volume{l) length{mm)
1
50
1
100
No. prey
recovered
Control
recovery
No. prey No.
taken
dead
14.0
15.0
13.5
13.0
13.5
9
13
29
14
(SO,SO)
41
37
21
36
39
0
0
0
0
0
14.5
15.0
13.5
13.0
14.5
30
26
17
23
27
(98,99)
70
74
83
77
73
0
0
0
0
0
11
Table;'rMysid predation on E.affinis in the presence of
< 69 um naturally-ocurring detrital material
No. prey Control No. prey
Mysid
Vessel
Prey
Density volume{l) length(mm) recovered recovery taken
No.
dead
2
17.0
16.5
15.5
15.0
16.0
28
32
35
28
31
(50,50)
22
18
15
22
19
0
0
0
0
0
50
1
13.5
15.5
13.0
14.5
16.0
23
17
29
26
24
(50,50)
27
33
21
24
26
0
0
0
0
0
100
1·
16.0
14.5
15.0
16.5
15.5
46
59
46
54
51
(100,100) 54
41
54
46
49
0
0
0
0
0
25
.,.
...
Discussion
Discussion
Field
survey
Equipment performance
The sampling equipment performed reliably throughout the survey
period, and proved to be an effective method of sampling
zooplankton
in shallow and turbid waters. No physical damage was
incurred by animals present in samples, but passage through the
pump did detach eggs sacs from species such as.Eurytemora.
A
field assessment of fecundity in this species was therefore not
possible.
Taxa identified
A total of 55 taxa were found in the samples collected, although
some of these were identified only to family or higher level.
This contrasts with the 135 taxa identified in a contemporary
study of the main channel Forth zooplankton by Taylor (1984,
1987). Taylor encountered euphausids,
considerably
ctenophores, mysids and a
greater variety of meroplanktonic
found in the littoral/intertidal
larvae than were
zone. His samples were, however,
numerically dominated by the same restricted group of neritic
copepods (Acartia, Oithona, Centropages, Temora, Pseudocalanus)
as were the samples in the present survey. A feature of the
present study was the persistent presence
and numerical
dominance of Eurytemora at all stations throughout much of the
year. This species, although showing a seasonal and spatial
pattern similar to that reported by Taylor (1984,1987) appears,
as observed by Roddie (1980) and dePauw (1973), to concentrate
in
the margins of estuaries.
The use of a 69 um mesh net in the present study permitted the
observation
and enumeration of groups such as the small cyclopoid
Oithona, rotifers, copepod nauplii and cirripede nauplii which
are frequently missed in zooplankton surveys. Amongst the
meroplanktonic
.'
larvae, cirripede nauplii were abundant throughout
much of the estuary and most of the year.
holoplanktonic,
The other,
groups, were also widespread and abundant, and
would clearly contribute much to
seasonal estimates of biomass
and production.
Rotifers are rarely reported in zooplankton surveys by virtue of
their small size; $ynchaeta.
the genus dominant in the upper
Forth, would not be retained by a mesh of greater than 100 um.
Hulsizer (1976) and Allan et al (1976) noted high rotifer
abundance during February and March in Chesapeake Bay estuaries,
which corresponds
to the present findings. These authors also
found, however, that this peak coincided with the maximum
abundance of Eurytemora;
in the present study, the Eurytemora
peak succeeded the rotifer peak •
.The extension of the survey area upstream of the range covered
by Taylor (1984) enables the interface between freshwater and
estuarine communities
seasonal,
to be more clearly reflected,
tidal and spatial variation
and the
in freshwater
influence was
readily apparent from the survey results. Peak abundance of
Daphnia, Bosmina, and Diaptomus at all upper-estuary
stations
occuured in January and February 1982, just after the rotifer
peak and just before a peak in abundance of Maranzelleria
larvae.
There was thus a succession of abundance maxima;
Rotifera
>
freshwater
sp.
>
Maranzelleria
>
Eurytemora
at Fallin, S.Alloa, Dunmore and Kincardine.
Hulsizer (1976)
reported
a similar succession of rotifers and cladocera in
~
Chesapeake
Bay. In contrast, an additional
chydorid cladocera,
occurrence
freshwater
taxon,
achieved maximum abundance and most frequent
in late summer and autumn. This pattern appears to be
typical of chydorids
(Vhiteside,
1974).
The winter peak of rotifers coincided with a winter minimum of
Eurytemora.
Synchaeta has similar linear dimensions
nauplii, but can, by reproducing
to Eurytemora
parthenogenetically
and at a
small size, maintain a higher intrinsic population growth rate at
low temperatures
than can Eurytemora.
Competition
nauplii and rotifers for resources may exacerbate
low temperatures
on Eurytemora
abundances,
the effects of
population growth. Conversely,
may be that rotifers can only compete successfully
at low Eurytemora
between
it
for resources
although this might imply a second
peak in. rotifer numbers during the summer decline in Eurytemora.
Villamson and Butler (1986) have demonstrated
that Diaptomus can
feed effectively
on small rotifers, and it is further possible
that the succession of rotifers and Eurytemora is a consequence
of predatory interaction.
Amongst the meroplankton,
Littorina egg capsules were, as
previously shown by Alifierakis
(1980), clearly associated
Although not numerically
and Berry(1980) and Roddie
in abundance with high spring tides.
reported, the presence on occasion of
large numbers of turbellaria
in S.Allloa samples is worthy of
note, as these animals may exert significant predation pressure
on zooplankton
(Maly et al 1980).
Community structure and environmental
variables
The zooplankton data were rather sparse, with low abundances
and infrequent occurrences of the majority of taxa. Vith the
exception of Eurytemora and the rotifers, Taylor (1984) found
higher abundances of most taxa common to both studies. This
contrasts with the pilot intertidal study of Roddie (1980) which
found generally higher abundances of most taxa than in the
present study. Further, since not all samples planned could be
taken, the distribution
of samples among attributes
station, date) was unbalanced.
260
(tide,
Despite these shortcomings,
reasonably
successful
relationship
the methods applied in analysis were
in describing
with environmental
Chi-squared
community structure and its
variables.
test of association
between occurrence
and
sample type
This approach proved reasonably effective in discriminating
between species and sample categories. A strong distinction
was
apparent between the neritic copepod community and marine
~
incursors, and the freshwater community. The relationship
between
these groups and tidal and spatial variation was also clear.
The larvae of benthic invertebrates
spring tides (where any association
were associated with high
existed), and their
occurrence was biased toward lower-estuary
stations. This is a
reflection of both the location of the parent populations
the advantages
and of
in larval dispersion derived from release during
periods of maximum tidal excursion. Greater tidal penetration
spring tides is also likely to account for the higher relative
frequency of occurrence
of neritic copepods and marine species
during this phase of the semilunar cycle.
261
on
Classification
Classification
analysis indicated that Centropages was confined
to spring/summer
samples, and was closely allied with the
occurrence of terebellid larvae. Oithona and Pseudocalanus
were
most abundant in spring, but occurred frequently throughout the
summer and autumn, and Temora was associated with these species
although occurring less frequently. Taylor (1984,1987) found
these copepod species generally most abundant in winter.
The seasonal pattern of the Acartia spp. matched that found by
Taylor (1984,1987); A.bifilosa
inermis and A.longiremis
were most
abundant in spring and autumn, while A.tonsa had a late summer.
peak. Polydora and cirripede cyprids occurred primarily in spring
but, as noted above, cirripede nauplii we/present
in many,
samples throughout the year. The nauplii and cyprids were not
distinguished
in the samples, but are likely to have been
predominantly
Balanus in the spring and Elminius during the
summer.
In general, the neritic and meroplanktonic
assemblages were
seen to be associated with higher temperatures and primary
production. TS/HDA analysis indicated that Pseudocalanus
Oithona were more persistent than Temora and Centropages;
and
the
former pair were common .in samples associated with lower salinity
and higher freshwater discharge ( e.g. in the Skinflats data
set). It was evident that primary production increased as a
discriminating
factor with decreasing distanc~ from the upper
tidal limit. This was related to the joint advection of lower
estuary phytoplankton
and zooplankton
also revealed the increasing
community
assemblages.
Classification
importance of the freshwater
in the same direction. The species associations
were
preserved, however, wherever overlap between stations occurred.
Analysis by date group emphasised
the importance of seasonal
factors, with groups in the latter part of the year reflecting
greater positive influence of primary production
and temperature.
The impact of benthic larval production was exagerrated
within
date groups 4 and 7.
Analysis of samples grouped by tide indicated that information
on
~
structure was preserved in larger, more diverse sample sets. The
Oithona/cirripede
nauplii/Pseudocalanus
a robust diagnostic
association
proved to be
feature of sample division. Salinity was
relatively more important for the spring and neap tide
categories
than for high and low tide categories due to the
inclusion of semidiurnal
For completeness,
tidal variation in the latter.
TS/MDA was conducted on the entire data set.
Vhilst it was impracticable
table and dendrogram,
illustrate
to represent the results as a two-way
the vector plot was constructed
the overall relationship
The plot emphasises
between variables
to
(Fig.4.1).
the overall dominance of salinity, and shows
the strong association
between salinity and distance from upper
tidal limit. Percent organic and tidal ~tate are also clearly
linked to salinity, although purely for reasons of definition
26)
in
....
N
o
r::
0
or4
...
co
.-4
CIS
...
CIS
.'
(,)
0
r-4
CO
~
N
m
r-4
r-4
as
e
•
...,.~
E-t
,
N
r-4
•
.::t
~
~
~
N
0
264
N
I
'"
I
r::
QI
r::
e
GIr::
..c
0
... 'CI-I
Gl0r4
bO'" >
r:: CIS r::
or4r::QI
>O'C
QI
..c1-l.-4
UlOCo
S
.. Ul CIS
'C Co Ul
QI ::s
r::0~
or41-10
.abO
e
Ul
0r::..-4
(,) CIS Ul
Co CIS
Ulrn,.Q
GI r::
r-40r4Q1
o.>..c
E~'"
CIS
Ul co r::
0
r-4 ~
r-40Ql
CIS
(,)
U2n1
1-1'C
0..-4
Co
Ul
~O
1-1'"
c::
o r:: CIS Ul
r-4G1r::QI
0.(,)..-4'"
e ::s
I-IGI_.a
o..c 1-1or4
...... (,)1-1
(,)
Ul'"
......
GI~-
>o'CnI
...
the latter case.
Date (association with sample cycle) is, as noted previously,
approximately
orthogonal
to salinity and is the second most
important variable. Inorganic suspended solids and phaeopigments
tended to be higher at low salinities and up-estuary, a
reflection of the existence of a turbidity maximum. Chlorophyll a
and acid ratio were closely linked, and were also orthogonal
to
salinity, while temperature tended in the same direction as
salinity but was less influential. Temperature
showed some link
with chlorophyll a. Discharge acted in the opposite direction
to
salinity and temperature and was also associated with low primary
productivity.
~
The species associations
identified in the analysis .of data
subsets were preserved in the all-sample analysis. MDA correctly
reclassified
70.3% of samples. Vright et al (1984), using similar
methods, found approximately
the same success in reclassification
for a larger sample set. In the present study success in
reclassification
decreased.
generally improved as the size of the sample set
Assessment
Vhilst DCA ordination
environmental
of methods
followed by correlation with
variables demonstrated
it was difficult
to quantitatively
influence as well as TS/HDA,
relate the results to
community structure since the information
accessible
in the final ordination •• This constraint also applies
to other ordination
methods such as non-metric
scaling as well as to to classification
similarity
about species is not
indices. The Bray-Curtis
multidimensional
methods based on
classifications
in the present study were much less intelligible
classifications
carried out
than the TS
and have therefore not been reported. A major
advantage of TS is that the end-product
is a two-way ordered
table in which both species and samples are classified.
only can association
identified,
Thus, not
between species and sample clusters be
but the degree of order in the table can be
interpreted
as an indicaton of the success of the analysis (or
of the degree of underlying order).
In the present work, it was possible to independently
sensitivity
of the analysis since the association
assess the
of samples
belonging to the same logical categories was readily apparent in
the dendrogram
derived from the table; i.e. within each TS group
it was relatively easy to see if samples were grouped on the basis
of station, tide or date. Further analysis was,strictly,
necessary where a heterogeneous
group was observed.
association
only
of samples within a
The utility of MDA has long been recognised in disciplines
such
as numerical taxonomy and sociology, and Vright et al (1984) and
Green and Vascotto (1978) inter alia have demonstrated
application
its
in ecology. By adopting a simple method of variable
visualisation,
the present study has achieved. two things.
Firstly, the relative influence of environmental,
temporal variables
in distinguishing
can readily be demonstrated.
spatial and
between biological groups
Secondly, the overall relationships
between variables can be concisely summarised.
The method appears to be robust, in that useful and intelligible
information was recovered from an unbalanced and rather 'noisy'
data set. In fact, most of the information seemed to be preserved
~
if the data were treated
in terms of presence and absence.
Beyond this, semi-quantitative
data was sufficient
influence of seasonal fluctuations
Eurytemora,
to reflect the
in abundance of taxa such as
rotifers and meroplanktonic
larvae.
Although Green (1974) described a more rigorous approach to the
use of MDA in analysing survey data with temporally-varying
environmental
data, the present methods were reasonably effective
in simultaneously
distinguishing
the effects of spatial and
temporal variables.
267
Patchiness
1. Dunmore samples: temporal variation at a single site
The back-transformed
95% confidence
limits for a single sample
were 29-347% ,within the range reported by Gagnon and Lacroix
(1981) for a variety of taxa, and are similar to those reported
by Roddie et al (1984) from a previous study on the Forth. The
uncertainty
of an estimate appears to increase with abundance
(Gagnon and Lacroix 1981), so abundance estimates based on a
single sample need to be treated with caution.
2. Skinflats samples: spatial variation
The patch sizes defined in this work syggest that the routine
practice of traversing approximately
should have
100m in sampling still water
ensured a reasonably representative
sample. The
maximum range of variation for a single taxon ( about a factor of
11 for Eurytemora)
was similar to that found at Dunmore. The
contrast in distribution
about the salinity-temperature
front between
the cope pods and the Polydora larvae is probably a consequence
distribution
of the parent polychaete population;
be unrelated
to the development
this would, of course,
of ephemeral high tide water patterns.
The existence of the front is interesting
indication of the degree of small-scale
physical characteristics
of the
of an estuary.
in itself, as an
hetereogeneity
in
Eurytemora
affinis
The results of field and laboratory
will be considered
investigations
on Eurytemora
together in this section.
1. Predation
Vhilst clearance rates appeared to be independent
of vessel
volume at 50 prey/I, ther was evidence at 25 prey/l that foraging
activity might be relatively
constrained
in smaller vessels.
Thus, mysids in 5 litre vessels appeared to search a larger
.'
volume in 24h than those in 2 litre vessels. This contrasts with
the findings of Fulton (1982) who reported, in a study of
Neomysis and Hysidopsis
predation on Acartia and Centropages
that
clearance rate was independent of vessel volume at all prey
densities.
In the present study, at 25 prey/I, final prey density
was rather similar in both 2 and 5 litre vessels at about
16-18/1. Prey density declined at roughly the same rate in both
sizes of vessels; the measured clearance rates may reflect a
steady acclimation
of effort to declining prey density, and thus
point at low prey densities
to some inhibition in smaller
vessels.
Irrespective
prey densities,
of volume, clearance rates were higher at lower
declining approximately
logarithmically
as prey
density increased. Clearance rates show~d evidence of decline at
densities above saturation;
Frost 1975) characteristic
this is considered
(Fulton 1982,
of animals displaying a Type II or
curvilinear
functional response (Holling 1959), although the
chi-squared
test did not distinguish effectively
between the
goodness of fit of this model or of a rectilinear model. The
curvilinear model yielded an estimate of handling time
and
maximum daily predation rate of 6.97 min. and about 170/d
respectively,
while the rectilinear model sugested values of 12
min. and 120/day. Predation rates at up to 100 prey/l were
similar to those estimated by Fulton (1982) for Mysidopsis
feeding on Acartia. Predator and prey in Fulton's experiments
were very similar in size to Neomysis and Eurytemora
~
respectively.
Cooper and Goldman (1980), however, reported
predation rates of 15mm Mysis relicta which were approximately
50% higher than those reported here.
Visual predation did not appear to be important in NeomysiSI
indeed, predation rates at higher prey densities were higher in
the dark than in the light, which might suggest that the level of
iilumination
interfered with mysid behaviour or perception •.
Cooper and Goldman (1982) found no difference between foraging
rates of Mysis relicta in the light and dark, while Ramcharan and
Sprules (1986) found that Mysis relicta had significantly
higher
predation rates on Daphnia in the light. The high turbidity
characteristic
especially
aggregrate,
of the Neomysis/Eurytemora
habitat would,
in the shallow regions where both species tend to
render visual predation ineffective much of the time.
The omnivorous, nonselective
nature of Neomysis foraging was
270
illustrated
by the lack of effect of detritus on predation rate,
despite the evidence of ingestion of detritus. A rectilinear
model of foraging may, in the light of this, be more appropriate,
as it implies a random search pattern and the ingestion of food
items in proportion
to their abundance.
Neomysis could not be quantitatively
sampled in the present
study but could, even at a density of 5/m-3, account for a
considerable
proportion
of the Eurytemora
population during the
late summer. The potential impact on.naupliar
copepodite
stages was not investigated,
additional
pressure on recruitment
2. Development
Development
and early
but could constitute
an
success (eg Fig. 4.2).
rates
rate in Eurytemora was not affected by differences
in food level, but was quantitatively
related to temperature.
Final body size was not measured, so unfortunately
no information
was obtained on growth rate. Corkett and McLaren (1970) noted an
inverse relationship
Development
between size and development
was approximately
line could successfully
rate.
isochronal, .in that a straight
·be fitted to the plot of stage against
time. Kelin Breteler et al (1982) described isochronal
development
in four species of North Sea calanoids, and the data
of Vuorinen
(1982) for E.hirundoides
similarly suggest isochronal
211
5
D
J
F
M
A
M
J
1981-1982
.,'
filled circ1es- E.affinis
open circ1es- N.integer
Figure 4~2 'Meanupper-estuary
272
al:wldances
J
A
5
o
N
development.
It should be noted, however, that in the present
study the duration of the n2 stage was consistently
longer than
that of the other naupliar stages. Landry (1983) reported similar
observations
for Acartia and Pseudocalanus.
The longer n2 stage
may be a consequence of the need to replace energy expended
during the nonfeeding nl stage.
Development
times were similar to those determined
for
Eurytemora using a variety of techniques. Vuorinen estimated
total development
time at 10 deg.C to be between 27 and 33 days
(cf 39 days at 8 deg.C), while Heinle and Flemer (1976) estimated
a shorter period of 26 days at the same temperature.
At 20 deg.C, agreement was closer, with estimates of 15.5 days
~
and 13.5 days respectively
Vijverberg
(cf 15.25 days). Estimates of
(1980) at 15 deg.C were 8.3 days for nauplii and 9 to
12 days for copepodites,
giving a total of 17.3 to 20.3 days (cf
17.0 at 16 deg.C).
The shape of the curve relating temperature to development
rate
indicated that the effects of temperature were most marked at
lower temperatures.
C
Little reduction was apparent from 16 deg.
to 20 deg.C, and it can be concluded that generation
time is
relatively independent of temperature above 14 deg.C and is
highly dependent below this temperature.
273
3. Field observations
The annual variation
in length followed an unusual pattern,
most marked in adult females. Conflicting
relationship
des~riptions
between body length and temperature have been
reported in the literature. A negative relationship
was described
of the
for cope pods
by Deevey (1960) and Harsha11 and Orr (1955), but Evans
(1977) described a positive relationship
for Pseudoca1anus.
Evans (1977)
also reported a bimodal pattern for Temora, similar to that found
for Eurytemora
in the present work. Be subsequently
length variation
~
Variation
in Temora to variations
in length in Eurytemora
in food quality.
in the Forth led to largest
body size in September, at water temperatures
which the annual length minimum occurred;
be the controlling
related
similar to that at
temperature
cannot, therefore,
factor. The length variation was less evident
in earlier stages, which were measured only for 9 dates; the more
frequent measurements
for adult females confirmed that the peak
was real. The subsequent decline in body size was followed by a
return to normal winter size. The dramatic increase in length (c.
~O%)between August and September argues that generation
time was
short, as no size overlap was apparent. The clear synchrony of
change in size between all stages similarly suggests a short adult
life span at all seasons.
This study has demonstrated
temperature-dependent.
with field observations
that development
Combining
rate is strongly
the findings on development
of the relationship
214
rate
between length and
stage led to the prediction of highest daily length increments
August and September and lowest daily length increments
midwinter.
in
in
Thus, on the one hand, growth rates are similar
between samples of radically different body size growing at
similar temperatures
and on the other growth rates differ widely
between specimens of much more similar size growing at very
different
temperatures.
controlling
Clearly, temperature
cannot be the factor
body size and, as noted above, the lack of overlap
between samples suggests rapid development
so food should not be
limiting.
Eurytemora
displayed a clear peak of abundance in May, reaching
numbers .in excess of 250000 m-3. Abundances
~
were considerably
lower
than found by Roddie (1980) somewhat later in the season, and
very much lower than maximum numbers reported for this species
elsewhere
(Burkill et al 1982, Heinle and Flemer 1976). Taylor
(1984,1987) has demonstrated,
however,
that variations
in zooplankton
abundance between consecutive years can exceed one order of
magnitude.
Following
the spring peak, abundance declined to a minimum at
the same time as body size reached a maximum. A reduction
in fecundity might leave more energy available for somatic
growth, if the reduction were not itself a consequence
of food limitation.
The rapidity with which .body size increased makes food limitation
an unlikely possibility.
Alternatively,
increased predation
pressure from mysids could lead to selection for increased body
size as a refuge from attack, although ~arly stages would still
275
be vulnerable were they present.
One further, simpler , hypothesis might be that the sudden
change in size was due to the replacement of the local population
by a larger adjacent population. Measurements
size distributions
were not made of
at other sites to test this hypothesis,
but
evidence can be adduced that in high spring tide samples in May
mean female lengths did not differ significantly
between Fallin,
S.Alloa and Dunmore. It would be likely that, if morphologically
distinct populations existed, evidence of advection woud have
been apparent at that time. In either case, it is possible that
seasonal genetic selection could account for the change in size;
Bradley (1978) demonstrated
~
tolerance characteristics
seasonal selection for thermal
in an American.Eurytemora
population.
Summary
Application
of classification
and discriminant
year's field survey data demonstrated
objectively
analysis to one
the practicability
of
relating zooplankton community stfucture to physical,
spatial, and temporal variables. Littoral zooplankton have been
shown to have a clear structure which agreed well with that
descibed by other studies on the Forth, and which could
successfully
be defined in terms of the above variables.
Field and laboratory studies relating to Eurytemora affinis
demonstrated:
a) seasonal variation in'abundance and size
b) a well-defined
relationship
beteen temperature and development
rate
c) the capacity for predatory interaction between this species
and Neomysis, one of the co-dominants of the low salinity region
of the estuary.
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."
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28.5
Appendix A
Environmental
variables
Variables are identified in tables as follows:
Tide
Date
FV disch
Susp.solids
%organic
Chla
Phaeo
Sal
Temp
..
'
tidal state
decimal mon th
mean monthly freshwater discharge (Cumecs)
total suspended particulate material (mg/l)
percent of susp.solids lost on ignition
Chlorophyll a (ug/l)
Phaeopigments (ug/l)
salini ty (%0)
temperature (deg.C)
Table Al
Tide
Date
Environmental variables : Culross
FY'
Disch.
S
N
S
N
S
N
S
N
S
S
S
N
S
N
S
N
S
N
S
N
3.37
3.60
3.86
4.07
5.32
,5.55
5.77
6.00
7.25
8.84
9.20
9.43
9.67
9.90
11.10
11.30
11.53
11.73
12.48
12.74
113.1
113.1
113.1
25.1
22.9
22.9
22.9
22.9
12.1
24.1
77 .9
77.9
77 .9
77.9
135.2
135.2
135.2
135.2
114.3
114.3
Susp.
solids
% org
Chla
Phaeo
Sal
Temp
50.80
40.30
74.18
39.78
17.80
33.20
122.18
8.30
7.45
21.18
8.18
5.75
7.80
22.50
0.00
0.32
3.95
1.18
1.18
9.35
8.61
6.73
3.04
4.35
4.36
1.49
4.02
12.69
*
*
3.11*
25.8
28.6
26.8
23.6
27.5
27.5
28.7
26.3
32.6
27.1
26.6
27.0
29.9
10.6
22.9
23.5
26.6
18.6
22.9
26.1
5.5
9.5
10.5
11.5
15.5
14.5
16.5
17.5
27.5
19.0
14.5
14.0
16.5
13.0
11.0
11.0
8.0
6.5
13.5
6.5
*
*
48.76
77.56
40.60
257.90
30.35
33.15
40.08
185.77
61.20
238.50
51.52
*
*
13.60
24.32
2.54
8.75
45.00
6.90
6.30
7.85
35.70
15.40
32.13
11.50
4.33*
2.72
7.36
2.48
2.00
2.20
9.88
1.34
6.68
0.00
2.26
1.03
8.49
4.06
0.33
1.89
12.70
2.40
16.55
2.90
Table A2 Environmental variables : Skinflats
Tide Date
S 0.35
S 1.39
N 1.61
S 1.87
N 2.11
S 3.37
N 3.60
S 3.86
N 4.07
S 5.32
S 5.77
N 6.00
S 9.20
N 9.43
N 9.90
S 11.10
N 11.30
S·11.53
N 11.73
S 12.48
FY
Susp. %org
Disch solids
100.0
97.2
97.2
97.2
79.4
113.1
113.1
113.1
25.1
22.9
22.9
22.9
77.9
77.9
77.9
135.2
135.2
135.2
135.2
114.3
*
*
*
*
Chla Phaeo Sal
*
*
*
*
154.60 25.85
56.70 9.40
.32.20 5.45
·38.33 11.73
29.68 7.63
33.80·8.75
35.08 7.80
*
*
*
*
0.00
0.00
0.11
0.75
1.18
1.39
4.41
* 31.0
* 27.0
* 15.0
* 25.0
3.45
2.84
4.08
3.81
2.64
4.08
3.14
21.5
27.4
18.6
27.4
23.6
23.4
29.6
*
*
*
* 26.2
57.36 14.85 7.68 3.66 27.7
34.18 6.23 1.34 3.00 27.8
·24.55 5.65 3.00 1.79 10.4
43.10 6.60 2.97 2.99 18.0
53.98 8.00 1.67 2.77 20.3
205.27 33.13 7.81 12.74 23.9
51.93 10.00 1.91 4.78 15.8
170.23 27.37 1.00 20.95 27.6
Temp
7.0
0.0
3.5
3.5
1.5
5.0
8.5
9.5
7.5
21.0
14.5
25.0
12.5
15.0
12.5
10.0
10.0
7.0
5.5
7.5
Table A3
Tide
Environmental
variables : Kincardine
Date FV
Susp. %org
Disch solids
Chla
Phaeo Sal
0.35 100.0
* 31.5
*
*
*
0.58 100.0
*
30.5
*
*
*
LN 0.58 100.0
*
23.0
*
*
*
HS 1.39 97.2
* 25.0
*
*
*
LS 1.39 97.2
*
18.5
*
*
*
HN ·1.61 97.2
*
11.5
*
*
*
LN 1.61 97.2
* 12.0
*
*
*
HS 1.87 97.2
*
23.0
*
*
*
LS 1.87 97.2
*
15.0
*
*
*
HN 2.11 79.4 26.80 5.60 0.00 1.79 17.5
LN 2.11 79.4 28.65 5.85 0.00 2.04 11.0
HS 3.37 113.1 63.90 10.00 0.00 1.65 18.4
LS 3.37 113.1 494.40 71.80 0.00 18.70 4.6
HN 3.60 113.1 12.50 2.90 0.00 1.27 20.2
LN 3.60 113.1 25.55 5.10 0.53 1.48 15.4
BS ·3.86 113.1 52.23 13.78 0.85 3.78 24.4
LS 3.86 113.1 140.15 29.95 0.67 11.67 10.4
HN 4.07 25.1 37.77 8.13 1.18 4.06 15.4
··w 4.07 25.1 156.65 23.45 2.00 8.93 14.0
HS 5.32 22.9 22.50 5.15 4.57 1.45 26.7
LS 5.32 22.9 75.10 14.55 2.63 7.97 9.7
HN 5.55 22.9 17.10 4.20 2.24 3.18 25.0
LN 5.55 22.9 394.66 60.33 8.19 21.59 16.3
HS 5.77 22.9 31.73 6.05 4.74 5.21 29.8
LS 5.77 22.9 60.83 10.85 2.40 6.31 15.6
BN 6.00 22.9
* 22.9
*
*
*
LN 6.00 22.9
* 21.5
BS 7.25 12.1'104.28* 23.08* 8.33* 5.52 30.5
LS 8.84 24.1
* 24.9
*
*
*
HS 9.20 77 .9 33.91 8.90 7.01 2.80 29.1
HN 9.43 77.9 23.20 4.25 2.16 1.58 23.1
LN 9.43 77.9 70.40 15.30 3.20 4.46 14.4
HS 9.67 77.9 259.90 48.50 6.94 8.03 30.4
LS 9.67 77.9 61.20 12.30 3.47 3.25 9.6
LN 9.90 77.9 34.02 7.68 2.34 0.82 8.2
HS 11.10 135.2 61.40 11.80 1.74 2.54 24.3
HN 11.30 135.2 50.03 8.60 1.04 1.82 14.4
LN 11.30 135.2 92.60 14.07 1.47 8.19 3.4
HS 11.53 135.2 53.32 9.18 2.80 2.87 24.5
LS 11.53 135.2 637.97 96.63 21.22 24.19 8.2
HN 11.73 135.2 37.35 8.43 0.00 1.98 23.6
LN 11.73 135.2 55.60 12.40 0.00 5.69 10.0
HS 12.48 114.3 106.77 16.70 0.45 10.46 22.8
LS 12.48 114.3 462.82 62.29 15.14 21.27 13.6
HN 12.74 114.3 32.37 7.75 0.80 1.53 23.5
LN 12.74 114.3 49.12 10.55 1.17 3.62 9.1
HS
HN
Temp
7.0
2.0
2.5
2.5
1.5
3.0
2.5
4.0
3.0
5.0
5.0
4.5
3.5
5.0
6.5
9.0
6.5
8.0
9.0
13.5
11.0
13.5
16.0
13.5
14.0
15.0
16.0
16.5
16.5
13.5
13.5
13.0
14.0
15.0
12.0
10.5
9.5
9.5
8.5
7.0
7.0
5.5
7.0
6.0
5.5
4.0
Table A4 Environmental variables : Dunmore
Tide
HS
HN
LN
HS
LS
HN
LN
HS
LS
RN
LN
HS
LS
HN
LN
HS
LS
RN
LN
HS
LS
HN
LN
HS
LS
HN
LN
HS
HS
HS
LS
RN
LN
HS
LS
RN
LN
as
LS
RN
LN
HS
LN
RN
LN
as
LS
RN
LN
Date FV
Susp. %org
disch solids
Chla Phaeo
Sal
Temp
0.35 100.0
*
*
*
* 28.0 7.0
0.58 100.0
*
*
*
* 24.0 1.0
0.58 100.0
*
*
*
* 14.0 0.5
1.39 97.2
*
*
*
* 22.0 0.0
1.39 97.2
*
*
*
* 10.0 0.0
1.61 97.2
*
*
**
4.0 2.0
1.61 97.2
*
*
*
* 2.5 1.5
1.87 97.2
*
*
*
* 14.0 3.5
1.87 97.2
*
*
*
* 2.5 4.0
2.11 -79.4 22.60 5.20 0.00 2.44 9.0 4.0
2.11 79.4 38.40 5.80 0.00 0.43 1.5 4.0
3.37 113.1 49.20 7.20 0.00 2.64 11.4 5.0
3.37 113.1 304.20 41.00 0.00 10.20 0.6 4.5
3.60 113.1 26.65 4.40 0.00 1.72 6.8 4.5
3.60 113.1 33.20 5.45 0.21 1.81 6.0 5.5
3.86 113.1 35.53 11.031.07 3.27 20.8 8.5
3.86 113.1 319.15 55.55 3.87 17.92 2.0 7.0
4.07 25.1 31.83 7.53 0.85 3.03 20.0 7.5
4.07 25.1 100.15 20.95 1.74 7.89 6.4 8.0
5.32 22.9 27.70 7.95 3.50 3.15 19.2 8.5
5.32 22.9 200.20 36.65 2.31 20.43 4.5 12.0
5.55 22.9 44.13 8.13 2.72 4.06 12.8 15.0
5.55 22.9 20.26 4.46 0.85 2.75 9.5 15.5
5.77 22.9 46.63 9.50 4.21 2.49 28.5 14.5
5.77 22.9 104.33 17.70 4.37 19.96 7.2 14.5
6.00 22.9
*
*
*
* 24.6 15.0
6.00 22.9
*
*
*
* 11.1 16.5
7.25 12.1 853.30 142.40 23.96 41.97 25.1 17.0
8.84 24.1
*
*
*
* 21.7 16.5
9.20 77.9 29.76 7.10 18.69 2.57 17.3 13.0
9.20 77.9 225.71 39.25 15.42 23.69 1.5 12.0
9.43 77.9 36.80 7.55 3.74 2.01 21.2 14.0
9.43 77.9 79.45 14.10 4.24 7.05 3.6 13.5
9.67 77.9 43.65 8.30 3.11 2.90 24.2 14.0
9.67 77.9 143.90 28.50 .5.61 6.95 4.3 15.0
9.90 77.9 20.40 3.95 3.24 2.56 4.2 11.5
9.90 77.9 284.75 41.95 7.08 19.18 5.4 11.0
11.10 135.2 53.35 8.05 1.17 5.02 11.5 9.5
11.10 135.2 190.95 30.45 1.67 16.55 0.5 9.0
11.30 135.2 56.987.75
2.94 6.57 14.5 10.0
11.30 135.2 96.13 13.00 5.52 4.69 2.6 9.0
11.53 135.2 57.03 12.37 3.60 3.31 15.2 7.0
11.53 135.2 383.97 56.43 17.49 17.27 1.8 6.0
11.73 135.2 60.60 13.53 0.98 4.00 3.8 6.5
11.73 135.2 588.80 80.53 12.03 16.72 1.4 5.5
12.48 114.3 86.96 15.10 0.66 7.32 20.6 7.5
12.48 114.3 503.93 83.60 17.81 18.33 4.0 5.0
12.74 114.3 21.36 4.80 0.33 1.54 8.0 2.0
12.74 114.3 10.86 1.95 0.50 0.90 1.9 1.5
Table AS Environmental variables : S.Alloa
Tide
Date FV
Susp. %org
disch solids
Chla
Phaeo Sal
Temp
HS 0.35 100.0
*
*
*
* 18.0 7.0
HN 0.58 100.0
*
*
*
* 22.5 0.5
LN 0.58 100.0
*
*
*
* 5.5 0.0
HS 1.39 97.2
*
*
*
* 16.5 0.0
LS 1.39 97.2
*
*
*
* 3.5 0.0
HN 1.61 97.2
*
*
*
* 3.0 2.0
LN 1.61 97.2
*
*
*
* 0.5 1.0
HS 1.87 97.2
*
*
*
* 6.0 3.0
LS 1.87 97.2
*
*
*
* 1.5 3.0
HN 2.11 79.4 15.40 3.95 0.10 0.87 3.5 4.5
LN 2.11 79.4 73.45 10.35 0.00 1.51 0.5 4.0
HS 3.37 113.1 314.70 43.50 0.26 3.04 4.4 5.0
LS 3.37 113.1 267.80 28.70 0.80 7.04 0.0 4.0
HN 3.60 113.1 34.45 6.30 0.00 3.06 2.6 4.5
LN 3.60 113.1 38.15 6.05 0.42 2.64 0.6' 5.5
HS 3.86 113.1 37.23 8.78 1.18 3.76 12.2 8.0
LS 3.86 113.1 282.35 49.45 2.14 21.04 0.8 7.0
HN 4.07 25.1 88.95 16.45 1.34 6.78 11.6 7.5
LN 4.07 25.1 237.75 35.45 4.41 9.71 1.0 8.0
HS 5.32 22.9 36.10 9.05 4.27 4.42 12.1 12.5
LS 5.32 22.9 202.70 35.45 2.31 23.95 2.4 11.0
HN 5.55 22.9 16.66 4.26 2.06 2.15 12.6 14.5
LN 5.55 22.9 51.73 9.13 2.19 3.75 1.6 15.0
HS 5.77 22.9 85.93 16.60 2.94 7.18 22.6 14.0
LS 5.77 22.9 266.35 40.20 7.78 18.16 3.5 14.5
RN 6.00 22.9
*
*
*
* 20.0 16.5'
LN 6.00 22.9
*
*
*
* 3.2 17.0
HS 7.25 12.1 42.72 11.08 10.95 0.45 19.0 18.0
HS 8.84 24.1
*
*
*
* 17.2 15.5
HS 9.20 77.9 92.51 19.90 8.51 9.95 9.8 11.0
LS 9.20 77.9 465.21 77.95 13.82 42.77 1.0 11.0
HN 9.43 77.9 25.50 4~15 4.67 2.34 10.4 13.5'
LN 9.43 77.9 122.65 19~10 4.76 5.08 0.9 13.5
HS 9.67 77.9 77.23 14.40 2.55 2.12 15.9 14.0
LS 9.67 77.9 461.50 70.10 14.77 21.38 1.1 14.5
HN 9.90 77.9 30.40 5.39 4.20 2.15 2.0 11.0
LN 9.90 77.9 338.15 48.35 2.60 13.42 0.8 10.5
HS 11.10 135.2 50.30 7.59 3.10 5.00 6.4 9.5
LS 11.10 135.2 278.35 41.45 8.48 13.81 0.0 9.0
HN 11.30 135.2 59.63 8.50 4.17 3.66 2.6 '9.5
LN 11.30 135.2 113.40 17.13 4.32 6.96 0.0 8.5
HS 11.53 135.2 63.50 10.03 4.67 7.94 4.3 6.0
LS 11.53 135.2 349.57 51.63 14.02 18.04 0.2 5.5
HN 11.73 135.2 33.67 7.33 1.02 3.65 2.5 6.0
LN 11.73 135.2 118.00 19.13 4.14 5.74 0.0 5.0
HS 12.48 114.3 217.16 31.30 6.23 6.23 7.7 7.5
LS 12.48 114.3 372.64 52.80 13.19 12.78 0.5 5.0
HN 12.74 114.3 20.12 4.95 0.33 1.89 6.1 2.0
LN 12.74 114.3 26.27 4.80 1.84 1.08 0.0 1.5
Table A6 Environmental variables : Fallin
Tide
HS
BN
LN
BS
LS
BN
LN
BS
LS
BN
LN
BS
LS
BN
LN
BS
LS
BN
LN
BS
LS
HN
LN
BS
LS
HN
LN
BS
HS
BS
LS
BN
LN
LS
BN
LN
HS
LS
BN
LN
BS
HN
LN
Susp. %org
Date FV
disch solids
0.35
0.58
0.58
1.39
1.39
1.61
1.61
1.87
1.87
2.11
2.11
3.37
3.37
3.60
3.60
3.86
3.86
4.07
4.07
5.32
5.32
5.55
5.55
5.71
5.77
6.00
6.00
7.25
8.84
9.20
9.20
9.43
9.43
9.67
9.90
9.90
11.10
11.10
11.30
11.30
11.53
12.74
12.74
100.0
100.0
100.0
97.2
97.2
97.2
97.2
97.2
97.2
79.4
79.4
113.1
113.1
113.1
113.1
113.1
113.1
25.1
25.1
22.9
22.9
22.9
22.9
22.9
22.9
22.9
22.9
12.1
24.1
77.9
77.9
77 .9
77.9
77.9
77.9
77.9
135.2
135.2
135.2
135.2
135.2
114.3
114.3
Chla
Phaeo
*
*
*
*
*
*
*
*
*
*
*
*
*
*
*
15.300
*
*
*
*
*
*
*
*
*
*
*
*
2.55 0.00
92.450
215.500
95.800
9.000
14.600
167.500
264.250
83.250
4.065
74.700
249.500
50.900
22.460
238.950
173.950
10.20 0.00
22.60 0.00
10.80 0.00
1.45 1.28
2.45 0.96
30.75 3.38
41.75 1.87
13.85 1.82
7.55 0.69
16.20 2.90
47.75 5.34
8.46 3.31
4.46 4.67
32.90 2.49
28.30 11.39
0.90
2.32
3.74
3.74
0.81
0.31
12.94
14.02
6.48
3.44
22.39
18.02
3.49
2.70
15.08
16.92
*
*
33.51
*
15.35
*
*
1.70
*
38.38
*
*
*
*
*
*
*
*
62.480* 13.88
109.710* 29.55*
*
458.110
77 .050
86.850
375.200
42.200
648.950
368.950
90.450
31.330
25.270
136.970
19.410
16.400
*
*
*
66.55 8.89 24.06
8.90 2.14 4.41
12.83 2.50 3.45
61.10 11.99 16.07
5.75 3.00 2.49
91.75 17.11 44.22
49.45 9.35 12.38
9.68 2.89 4.89
3.05 1.57 2.15
3.80 2.32 0.80
17.30 4.14 6.14
2.60 0.33 1.89
3.00 1.34 0.65
Sal
Temp
11.0
5.0
0.5
5.0
0.5
0.5
0.0
0.0
0.0
0.0
0.0
0.2
0.0
0.2
0.0
2.2
0.2
1.4
0.1
3.5
0.8
1.3
0.2
12.2
0.0
6.5
0.8
5.7
0.2
2.6
0.2
1.5
0.2
0.0
0.2
0.1
0.0
0.0
0.0
0.0
0.0
0.4
0.0
7.0
0.0
0.0
0.0
0.0
1.0
1.0
2.5
2.5
4.5
3.5
4.5
4.0
5.0
5.0
7.5
6.5
7.0
7.5
11.0
11.0
14.5
14.5
14.5
14.5
16.5
17.0
19.0
15.0
10.0
10.5
13.0
13.0
14.0
12.0
11.0
9.5
9.0
9.0
8.5
5.5
2.0
2.0
_)
.,'
Appendix El Zooplankton abundances
Table Bl al Fallin
STATION ETEOLARV NERElARV MARAwIRE MARAEGGS MARATROC BIVAlARV
Fll.1ZHS
F18.1Zl-tN
F18.1ZL~
F12.01HS
F12.01l5
F19.01HN
F19.01LN
FZ7.0U"5
F27.01L5
F03.021'ofN
F03.02L~
Fll.0JH5
Fll.03LS
FI7.0.3HN
FI7.03L~
FZ6.03HS
F26.03LS
F02.04HN
F02.04L~
F10.05HS
FI0.05LS
F17.051'1N
F17.05lN
F24.051-1S
F24.05lS
F31.05HN
F31.05lN
F08.0nofS
F26.08t'!S
F06.09HS
F06.09l5
F13.09HN
F13.09lN
F20.09LS
F27.09HN
F27.09L~
F03.11HS
FOJ.lllS
F09.11HN
F09.11lN
F16.11lS
F2J.12HN
F23.12LN
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
104
2·
0
0
9
0
0
0
0
0
0
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
141
4
142
2
321
0
0
0
0
0
0
0
17
0
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
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
lZ
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Table Bl b. Fallin
STATION LITTCAPS LITTVELI CIRRNAUP CIRRCYPR TEREL.ARV POLYCILI
·Fll.12HS
F18.121-1N
F18.12L.N
F12.01t~S
FI2.OILS
F19.01HN
F19.01LN
F27.01HS
F27.01LS
F03.02HN
F03.02LN
Fll.031-1S
Fll.03LS
F17.03HN
F17.03LN
F26.03HS
F26.03LS
F02.04HN
F02.014LN
FIO.05~S
FIO.05LS
F17.051-1N
F17.05LN
F24.0SHS
F24.05LS
F31.0SHN
F31.0SLN
F08.071-1S
F26.08HS
F06.0qI-lS
F06.0QLS
F13.0QI-lN
F13.09LN
F20.09LS
F27.09HN
F27.0QLN
F03.11HS
F03.1ILS
F09.111-1N
F09.1ILN
FU,.11LS
F23.121-1N
F23.12LN
0
0
0
0
0
0
0
0
0
58
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
5
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
27
0
2
0
5
'0
0
1
0
0
2
0
0
2
0
0
0
0
0
0
0
0
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
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
2
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
0
0
0
0
0
2
0
.
0
9
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
17
4
0
2
0
2
4
0
0
o .
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Table Bl c. Fallin
PSEUELON TEMOLONG CEt-4THAMACALAHELG SAGIELEG 011<00101
STATION
,
Fll.12HS
F18.12HN
F18.12LN
F12.01HS
F12.01LS
Flq.01MN
Flq.01LN
F27.01HS
F27.01lS
F03.02HN
FOl.02lN
Flt.03MS
Fll.03lS
F17.03HN
F17.G3l~
F26.03HS
F26.03lS
F02.04HN
F02.04LN
FI0,05HS
F1O,05lS
F17.05HN
F17.05LN
F24.05HS
F24.05LS
F31.05HN
F31.05LN
F08,07MS
FZ6,08HS
F06.0qHS
F06,OqLS
fll,OqH~
FI3,OQlN
FZO.OQlS
F27.0QHN
F27,OqU~
F03.1lHS
FOl.IIlS
FOQ,l1HN
FOQ.l1lN
F16.11lS
F2l.12HN
F2l.1ZlN
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Z
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
.
'0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
38
0
0
0
0
Table Bl d. Fallin
..
•
-
_._
•
-
0
••
-
•
STATION ACARINER ACARTONS ACARlONG ACARCLAU ACARCOPE OITHSIMI
Fll.12MS
Fle.tzHN
F18.12lN
F12.01HS
F12.01lS
F19.01HN
F19.01lN
F27.01MS
F27.01lS
F03.02HN
F03.02lN
Fl1.0lMS
Fll.0llS
F17.Q.lMN
F17.0llN
F2b.OlMS
F2b.OllS
F02.04HN
F02.04lN
Fl0.0SH5
Fl0.0SlS
F17.05HN
F17.05lN
F24.05H5
F24,OSl5
Flt.05MN
Fll,05lN
Foe,07HS
F2b.OBMS
FOb.09HS
FOb,09lS
F13.09HN
Fll.09lN
F20.09lS
F27.09HN
F27.09l~
F03.11HS
F03.lllS
F09.IIHN
F09.1IlN
FU,.lllS
F23.121'fN
F23,12lN
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
'0
0
0
0
0
'0
0
0
0
,0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
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
0
0
0
17
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
·0
24
0
9
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
B
0
0
0
2
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
0
0
0
0
0
0
o·
0
0
0
0
0
0
0
0
0
0
3
0
0
0
0
4
0
0
0
0
0
0
0
0
·0
0
0
0
0
0
0
0
Table ai £1 Fallin
STATION EAFFAOUL EAFFCOPE EAFFJUVE HARP SPEC ROTISPEC COPENAUP
Fll.12"'S
FI8.12t-N
F18.12LN
F12.01"S
F12.01LS
F19.01"'N
F19.01LN
F27.01HS
F27.01LS
F03.02MN
F03.02lN
Fll.03HS
Fll.03lS
F17.03 ..N
F17.03lN
F2&.03HS
F2b.03lS
F02.04MN
F02.04LN
F10.05HS
Fl0.05LS
F17.05HN
F17.05LN
F24.05HS
F24.05LS
F31.05HN
F31.0SLN
F08.07~S
F20.08HS
F06.09HS
F06.09LS
F13.09t'!N
F13.09LN
F20.09LS
F27.09HN
F27.09LN
F03.11HS
F03.11lS
F09.11HN
F09.11lN
F16.11lS
F23.12t-1N
F23.12LN
770
328
8
552
0
0
10
09
0
35
8
11
0
4
2
1450
24
1348
38
14927
309
30139
38540
14521
128
3168
81275
9818
2
32
3771
245
51
597
22
44
8
113
0
lb3
0
25
0
43
15
&173
74
2538
28
36214
109
22750
20b36
98390
262
53950
70317
4573
0
5
0
0
7
2
0
0
26
2
136
12
39
2
84
28
0
0
0
19
68
25
0
4
0
18
0
o .
159740
57
14487
126b33
437
530
53
10117
0
232
6144
92
78
12
4
0
528
18
23
32
34
71
19582
209
20392
12
883&2
475
39&30
712
153120
5108
193621
309b8
11565
4
. 34
6
8
600
298
05
1179
177
25
41
17
17
5
11
11
0
41
69
16
2
298
17
32
2
92
6
0
22
14
0
7
17
21
13
0
8
0
9
4
18
29
0
4
14
277
31
56
18
2
90
12
0
56
19
6
4
43
15
7 .
6
0
1044
105
IOb47
109
57
30
39
87
0
14
10
4
2192
77
0
0
82560
0
91
0
.59
0
0
0
0
0
0
37
0
0
0
0
0
43
30
96
48
0
29
112
9
0
0
1232
0
0
369
0
0
0
0
9
oJ
0
8
10
20
11
7
9
28
0
32
39
7
166
25
64
66
46
15
- 0
0
4
0
Table B2 &.South Alloa
..
-. -
'.-
-.
-
-
___
._
_
__
_
...
__
_
••
••
4
_.
•
STATION ETEOLARV NERELARV MARAWIRE MARAEGGS MARATROC'BIVALARV
SII.12~S
S18.12tiN
S18.12lN
S 12.0 HIS
S12.01LS
S19.01tiN
S19.01lN
S21.01~S
~27.0IlS
S03.0ZHN
S03.02lN
Slt.03HS
Slt.03lS
S17.03HN
SI7.0~lN
SZb.OltiS
SZb.03lS
S02.04tiN
S02.04lN
SIO.OS"'S
SIO.05lS
S17.05HN
S17.05lN
S24.0SHS
SZ4.05LS
S31.05HN
S31.05lN
S08.07HS
S2b.OeHS
SOb.09HS
SOb.09LS
S13.09HN
S13.09LN
S20.09HS
S20.09LS
S27.09HN
SZ7.09lN
S03.11MS
SOl.IILS
S09.1IHN
S09.11L~
Slb.l1t-tS
Slb.tllS
S23.l1HN
S23.11LN
S15.12HS
SIS.l2lS
S23.12HN
S2l.12lN
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
27
b
0
0
4
11
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1436
9
574
13
3t45
t6
57055
83
0
0
0
0
0
0
0
0
0
0
0
700
2
0
0
390
5
0
0
0
0
0
0
0
0
0
0
0
0
0
150
0
0
0
440
0
0
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
12
0
0
0
0
0
0
0
0
0
0
0
0
2
0
. 0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
.. .- -.
Table B2 bl South Alloa
STATION LITTCAPS LITTVELI CIRRNAUP CIRRCYPR TERELARV POLYCILI
SII.12HS
SI8.12t1N
S18.12LN
S12.01tiS
S12.01LS
S19.01HN
S19.01LN
S27.01HS
S27.01LS
S03.02HN
S03.02LN
S11.03t-1S
Sl1.03LS
S17.03HN
SI7.0_3LN
S2.6.0lHS
S26.03LS
S02.04HN
S02.04LN
SI0.0SHS
SI0.0SLS
S17.0St-lN
S17.0SLN
S24.0SMS
S24.0SLS
S31.0SHN
S31.0SLN
S08.07HS
S26.08HS
S06.09HS
S06.09LS
S13.09HN
S13.09LN
S20.09HS
S20.09lS
S27.09HN
S27.09lN
S03.11HS
S03.tllS
S09.tlHN
S09.11lN
S16.11HS
S16.11lS
S23.11HN
S23.11lN
S15.12HS
SlS.12lS
S23.12HN
S23.12LN
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
9
0
4
0
0
0
0
0
0
0
2S
0
6
0
2
7
0
0
0
0
0
7
0
8
0
2
0
0
2
2
0
0
4
8
0
0
6
8
0
16
0
0
0
0
0
0
0
0
27
18
7
0
0
25
0
0
0
0
0
0
0
"
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
11
0
'4
0
0
0
0
0
0
0
0
0
0
0
0
5
0
0
0
0
0
0
0
0
0
,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
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
"0
2
0
0
0
0
0
0
0
2
0
0
0
0
0
S
10
0
0
0
0
0
0
0
22
0
0
0
9
0
0
0
0
0
0
0
0
0
4
2
0
0
0
8
0
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
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
21
2
3&
0
113
86
12
0
9
5
34
0
9
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
Table B2 C I South Alloa
...
-
..... -
~
._
.... .-.....
-
-
,_
~....... - ...
-
.
.. ~
STATION PSEUELON TE""OlONG CENTHAMA CAlAHElG SAGIElEG 011<00101
Sl1.12t-1S
SI8.1ZI1N
SI8.IZLN
S12.01HS
SIZ.0IlS
S19.01t'lN
S19.01lN
S27.01t-1S
S27.01lS
SOJ.02I1N
SOJ.OZlN
Sl1.0JHS
SII.0llS
S17.03HN
S17.0llN
SZb.03HS
SZb.03lS
S02.04HN
S02.04LN
S10.0S115
SIO.OSlS
517.0511N
S17.05LN
S24.05HS
S24.0SlS
S31.05HN
S31.05lN
S08.07t~S
SZb.08t-1S
SOb.09t-1S
SOb.09lS
S13.09HN
S13.09LN
S20.09HS
S20.09lS
S27.0911N
SZ7.09LN
S03.11~S
SOl.l1lS
S09.11~N
S09.11lN
SI6.11HS
SI6.11LS
SZ3.11HN
SZ3.11lN
SIS.12MS
S15.12lS
S23.12t4N
S23.1ZLN
0
0
0
.0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
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
2
II
0
0
0
0
0
0
0
b
8
0
0
0
25
0
4
0
0
0
0
0
30
0
0
0
0
17
0
0
0
100
0
0
0
0
0
0
0
0
0
0
0
0
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
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
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
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
4
0
0
2
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
2
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
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0;
0
,
Table B2 d. South AlIos.
OITHSIMI
STATION ACARINER ACARTONS ACARLONG ACARCLAU ACARCOPE
13
0
0
0
0
0
S11,12HS
30
0
0
0
0
0
S18,12HN
0
0
0
0
0
0
S18,12LN
0
0
0
0
0
0
S12.01HS
0
0
0
0
0
0
SI2,OlLS
4
0
0
0
0
0
S1Q,OIHN
0
0
0
0
0
0
SlQ.OILN
0
0
0
0
0
4
S27.01MS
0
0
0
0
0
0
S27.01lS
0
0
0
0
0
0
S03,02HN
0
0
0
0
0
0
S03,02lN
4
2
0
0
0
0
S11,03MS
0
0
0
0
0
0
Slt.03lS
4
2
0
0
0
2
SI7,03HN
0
0
0
0
0
0
S17.03LN
0
0
0
0
0
0
S2b.03HS
0
0
0
0
0
0
S2b.03LS
2
0
0
0
0
2
S02,04HN
0
0
0
0
0
0
S02,04LN
4
2
0
0
0
0
SlO,05~S
0
0
0
0
0
0
S10,OSLS
0
0
0
0
0
0
S17,OSMN
0
0
0
0
0
0
S17,OSLN
6
82
0
43
0
8
S24.05t-tS
0
0
0
0
0
0
S24,OSLS
0
0
0
6
0
0
S]1.05HN
0
0
0
0
0
0
S3t.OSLN
S
2q
. b8
0
0
0
SOe,07HS
0
12
0
0
41
0
S2b,06HS
0
45
0
0
llb
0
SOb,OqHS
0
0
0
0
0
0
SOb,OqlS
2
11
0
0
11
0
S13,OQHN
0
0
0
0
5
0
S13.0qlN
0
bO
2
0
eb
0
S20.0QHS
0
5
0
0
0
0
S20.0QLS
0
0
0
0
0
0
S27.0QHN
0
0
0
0
0
0
S27,OQLN
0
8
0
0
0
0
S03.11HS
0
0
0
0
0
0
S03,IILS
0
0
0
0
0
0
SOQ.IIHN
0
0
0
o .
0
0
SOQ,11LN
0
2
0
0
0
0
Slb,11MS
0
0
0
0
0
0
Slb,IILS
0
2
0
0
0
0
S23,IIHN
0
0
0
0
0
0
S23.11LN
0
0
0
0
0
0
SIS.12MS
0
0
0
0
0
0
SIS,12LS
4
0
0
0
0
S23,12MN
0
0
0
0
0
0
S23.12LN
"
I
Table B2 ea South Alloa
STATION NEOMINTE OIAPGRAC DAPH5PEC BOSMSPEC CHYDSPEC CYCLSPEC
SII.12MS
S18.12tiN
S18.12LN
512.011'''5
S12.01LS
519.0ltm
S19.01LN
S27.01HS
S27.01LS
S03.02HN
S03.02LN
SI1.0lHS
Stt.03LS
517.0lHN
517.0·3LN
S26.03HS
52b.03LS
S02.04HN
S02.04lN
S10.05H5
. S10.0SLS
. S17.05MN
SI7.0SLN
S24.05HS
52".05LS
5l1.05HN
Slt.05LN
S08.07H5
526.08H3
S06.09H5
S06.09LS
S13.09HN
S13.09LN
S20.09t-1S
520.09L3
S27.09HN
S27.09LN
S03.11t-15
S03.IILS
S09.11HN
S09.IILN
S16.11HS
Slb.IILS
S23.11t-4N
S23.11LN
S15.12H3
S15.12LS
S23.12HN
S23.12LN
0
0
0
0
0
12
12
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
10
0
0
0
0
"
0
0
0
0
883
2
5
0
9
0
"0
11
0
14
0
0
0
0
0
0
0
0
0
8
12
4
12
53
' 35
15
9
27
2
11
22
17
0
0
0
2
0
0
0
0
0
0
0
0
"
0
0
215
2
0
0
0
2
0
0
0
0
6
0
Z
0
0
0
0
0
2
13
4
0
0
0
0
0
0
0
0
0
35
29
41
27
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
6
0
0
0
0
0
7
0
2
5
0
0
0
0
0
0
0
0
0
2
12
0
0
3
12
0
0
0
0
0
0
0
0
0
0
0
0
0
0
16
0
50
0
0
0
0
0
0
0
21
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
b
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
53
0
0
. 0
10
0
0
Z
2
0
0
0
2
0
0
0
0
0
0
0
0
0
10
0
9
0
0
0
0
0
11
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
7
0
0
0
0
1
Table 132fa South Alloa
STATION E.FFAOUl EAFFCQPE E.FFJUVE HARPSPEC ROTISPEC COPENAUP
Sll.12HS
SI8.12t-1N
S18.12lN
S12.01HS
S12.01lS
S19.01HN
S19.01lN
S27.01HS
S27.01lS
S03.02t-1N
S03.02lN
S11.03MS
S11.03lS
S17.03MN
S17.03lN
S2b.OlMS
S2b.OllS
S02.04MN
S02.04lN
SIO.OSHS
S10.0slS
S17.05HN
S17.05lN
S24.05HS
S24.05lS
S31.05HN
S31.05lN
S08.07HS
S2b.08HS
SOb.09HS
S06.09lS
S13.09HN
S13.09lN
S20.09HS
S20.09lS
S27.09HN
S27.09l~
S03.11HS
S03.lllS
S09.11HN
S09.11l~
Slb.l1HS
Slb.l1lS
S23.11HN
S23.11lN
SIS.t2HS
S1S.t2lS
S23.12HN
S23.12lN
3428
758
92
141
53
78
lb
382
15
191
254
178
Ib
91
53
150
20654
307
495
121
2540
1971
7228
7913
2739
3209
15895
770
17
57
6
15
l5
129
0
0
19
382
8
84
4
348
lb5
166
269
71
1183
306
17
3298
3674
365
1005
48
362
0
1552
22
3010
27
55&
27
125&
124
1&94
18702
3348
600
7333
4043
34852
1491
33999
28759
40168
19824
15215
7
9b
50
9
0
285
14
b3
5
1782
8
183
12
1781
30
458
4
411
115
986
b
2143
523
416
13371
1296
1785
74
2244
224
10944
318
3242
8
2205
286
l141
15180
10449
5775
10229
12025
265&4
7889
7476
64782
28909
34670
29364
0
. 46
9
104
0
1128
114
270
' 14
1132
20
415
15
4950
bb
2575
8
1059
2b9
914
24
118
790
4
833
b29
50
45
47
22
95
41
554
43
13
31
3&
45
51
182
21
132
b
1154
78
115
18
47
23
20
0
68
7
0
23
53
7
5
8
71
0
137
82
88
0
2
183
854
12
0
13550
338495
241
94102
37212
23662
107
52435
8952
341250
4545
1415
32
21
114
47
0
94
224
73
0
ll85
0
19217
2304
5685
0
8370
0
37
62
42
25
144
19
12
14
53
30
567
21
13439
&2
2016
32
6449
228
7510
58
0
0
0
0
0
0
0
0
0
0
0
502
27
87
164
293
0
539
0
1587
56
1774
430
739
1086
0
0
81b3
131
572
13b
721
16
1822
110
80
9
31
30
21
38
10
19
60
0
10
142
14
b
Table B.3 al Durutore
STATION ETEOLARY NERELARY MARAWIRE MARAEGGS MARATROC BIYALARY
011.12HS
018.12~N
018.12l.N
012.01HS
012.01LS
019,01HN
019,01l.N
027. 01t~S
027~01LS
003.02HN
003,02LN
011.03HS
011.03LS
017.03HN
017.0·3LN
026.031'15
026.03LS
002.04HN
002.04L~
010.051'15
010.05L5
017.05HN
017,O5l.~
024.051'15
024.05lS
031.0SHN
031.05lN
008,071'15
026.081'15
006,091'15
006.09LS
013.09HN
013,09lN
020.091'15
020.09l5
027.09HN
027.09LN
003.111'15
003,11lS
009.11HN
009.11lN
016.111'15
016.11lS
023.11HN
023.11lN
015.121'15
015.12l5
023.12HN
023.12l.N
;
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
6
0
0
0
3706
0
0
0
0
0
0
0
225
5653
973
16022
2258
0
1060
350
0
b8
0
0
0
0
0
9
9
0
0
0
0
0
0
0
0
1120
73
0
0
0
2bl
0
0
la
43
6
30
0
0
0
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
860
117
0
0
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
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
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Q20
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
20
0
0
0
0
0
3383
78
0
0
4
0
0
27
12
13
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Table BJ ba Dunmore
STATION LITTCAP5 LITTVELI CIRRNAUP CIRRCYPR TERELARV POLYCILI
Dll.12~S
DI8.12~N
D18.12lN
OI2.01~S
D12.01lS
O19.01MN
D19.01lN
027. 01t~S
027.01lS
003.02MN
003.02U04
011.03MS
011.03lS
017.03HN
C17.03lN
026.03MS
026.03LS
002.0"MN
002.04lN
010.05,;S
010.05lS
D17.05HN
017.05LN
024.05HS
D24.05LS
031.05MN
031.05LN
008.07MS
02b.08';5
00b.09';S
006.09LS
013.0QHN
O13.09LN
020.09MS
020.09L5
D27.09HN
027.09LN
003.11MS
D03.lllS
009.11';N
D09.11LN
Olo.I1HS
Olb.llLS
D23.11HN
023.11lN
015.12"'S
C15.12LS
023.12HN
D23.12LN
0
4
0
18
0
0
0
0
0
0
0
0
0
0
0
0
0
0
50
7
2
0
0
2
0
8
0
0
2
2
0
2
0
0
4
0
22
0
0
0
4:
0
0
0
"
0
0
4
0
2
2
0
0
237
0
14
0
0
0
0
0
0
0
0
0
a
0
199
0
12
0
0
0
0
0
0
0
0
0
0
0
38
0
0
0
0
0
0
0
125
2
55
2
295
0
0
0
0
0
0
0
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
4
0
0
0
0
0
0
0
011
0
14
0
b
00
0
0
20
0
700
20
125
0
0
0
0
0
0
2
0
0
0
0
0
0
1.1
0
5
0
0
8
4
23
18
. 2
0
2
0
0
0
19
1.13
0
0
3
0
2
0
31
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
75
0
0
0
0
0
0
"00
0
7
4
22
0
0
0
0
0
0
0
0
0
0
0
0
0
52
0
0
0
0
0
2
2
0
0
0
0
0
0
0
0
0
0
0
0
0
"2"0
0
0
0
0
0
0
0
0
0
0
30
24
0
0
0
0
0
0
0
0
0
0
0
'0
0
0
Table BJ c. Dumlore
STATION PSEUELON TEMOLONG CENTHAMA CALAMELG SAGIELEG OIt<OOIOI
011.12HS
Ole.12M~
018.l2LN
O12.01MS
012.01lS
019.01HN
019.01lN
027.01HS
D27.01lS
D03.02HN
003.02lN
OII.OlHS
OI1.03lS
D17.03MN
D17.o.3lN
D2b.03HS
D2b.OllS
002.04HN
002.04LN
010.0SHS
OIO.OSlS
017.05H~
017.05lN
024.05HS
024.0SLS
D31.05MN
OJ1.0SlN
D08.07HS
OZ,b.OSHS
006.09MS
006.09LS
013.09HN
O13.09lN
Oz'O.09HS
OZ,O.09LS
027.09HN
027.09LN
DOl.llMS
D03.1IlS
009.IIMN
009.11lN
Dlb.llMS
Olb.lllS
Oz'3.1IHN
D23.11LN
D1S.1Z,MS
D1S.t2lS
Dz'3.t2HN
D23.12lN
144
48
33
0
0
0
0
0
0
0
2b
0
0
0
0
0
0
0
0
0
0
0
0
0
0
"
0
0
4
170
8
0
0
0
0
0
0
0
0
0
0
0
0
0
0
b
0
0
0
0
2
2
0
.0
223
0
163
0
0
317
0
16
0
0
0
0
0
0
4
174
0
b
178
0
10
0
0
0
27
0
2.
0
2
4
0
"00
0
22b
0
0
0
0
0
0
81 .
0
0
0
0
0
<7
0
0
0
6
0
45
q
0
0
Z
34
0
0
0
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
4
0
0
0
,4
0
0
0
2
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2.
2
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
13
0
0
0
0
0
0
0
0
0
2.
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
10
5
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Table
B3 dl Dunmore
STATION ACARINER ACARTONS ACARLONG ACARCLAU ACARCOPE OITHSIMI
Dl1.12HS
C18.12~N
018.12LN
OI2.01~S
OI2.0ILS
OI9.01~N
O19.01lN
D27.01~S
027.01LS
003.02~N
003.02lN
Dll.0l~S
Oll.OllS
017.0l~N
O17.0lLN
02b.03MS
02b.OllS
002.04t'fN
002.04lN
010.0S"'S
010.0SLS
OI7.0S~~
OI7.0SLN
024.0S"'S
024.05LS
Oll.OSI-IN
D31.05LN
D08.071-1S
026.08"'5
006.091-1S
OOb.09lS
013.09"'N
013.09lN
020.09H5
020.09LS
027.091-1N
027.09L~
003.11HS
D03.IIlS
009.11HN
009.11lN
Olb.llr1S
Dll>.l1lS
023.11HN
c23.l1lN
cIS.1ZHS
cIS.12lS
023.12t-tN
023.12lN
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
0
0
0
0
6
0
0
0
0
0
0
4
0
2
0
30
0
95
0
4
0
"0
0
0
0
0
0
0
0
0
8
0
0
·0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
19
207
18
Ib
11
542
31
0
0
17
0
0
0
0
0
0
4
0
0
0
0
0
0
0
2
0
41
0
665
0
0
0
. 0
0
0
3
81
0
12
0
0
0
3
0
9
0
0
·0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
27
0
0
0
0
0
0
0
0
20
0
0
"00
0
0
0
"0
0
0
0
0
9
0
11
0
0
0
0
0
0
0
0
0
0
0
0
0
0
"
2
0
0
0
2
0
0
0
0
0
0
0
0
0
22
0
4
0
0
0
0
186
0
10
0
0
0
0
108
0
456
0
2
0
0
24
197
12
99
0
1962
17
0
0
9
0
0
0
88
0
0
0
0
0
0
0
0
0
10
0
0
0
0
0
0
0
4
0
0
0
0
137
75
0
31
0
198
0
7
0
5
0
31
0
0
0
47
0
15
2
62
Z
0
0
I>
8
98
0
0
10
0
Table B3 e I DunIlore
STATION NEOf-lINTEOIAPGRAC OAPHSPEC BOSMSPEC CHYOSPEC CYCLSPEC
Ott.12MS
018.12tofN
018.12LN
O12.0ltofS
D12.01LS
O19.0ltm
D19.01LN
027.01tofS
027.01LS
D03.02MN
D03.02lN
Oll.03HS
Dll.q.3LS
O17.03MN
017.03LN
C2b.a3MS
02b.03LS
002.04MN
C02.04LN
CIO.05~S
C1O.05lS
017.05HN
D17.05LN
024.05HS
OZ4.0SLS
C31.05HN
031.05LN
008,07HS
D2b,08HS
006,09HS
DOb.09LS
D13.09MN
013.09lN
020.091'15
020.09l5
OZ7.09HN
027.09LN
C03.UMS
003.11lS
OOq.llHN
Da9.11LN
Olb.llHS
Olb.llLS
023.11tofN
023.11LN
C15.12H5
OIS.t2lS
023.12HN
023.12LN
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
6
0
0
0
0
0
0
0
0
0
0
0
4
0
0
4
0
17
0
0
a
2
4
a
0
0
0
0
0
0
0
0
0
4
0
2
0
0
0
0
0
0
0
4
53
0
141
0
22
0
16
0
0
0
0
0
0
0
0
159
0
27
0
7
0
0
0
0
0
0
0
a
a
a
a
0
a
a
a
a
0
a
0
.a
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
a
a
0
2
4
324
0
0
0
0
0
0
0
0
a
0
0
0
0
0
0
0
0
0
0
b
0
4
22
12
0
0
0
0
0
a
0
0
0
0
0
0
0
a
0
0
2
0
0
a
2
0
0
0
0
0
0
0
0
0
a
0
0
0
0
30
0
a
2
a
0
0
34
0
0
0
a
0
0
a
a
0
0
a
a
2
2
8
4
0
·24
16.
44
a
4
0
a
0
a
a
0
0
0
0
0
a
a
0
0
0
0
0
0
0
a
2
a
0
a
a
0
0
0
0
"a
58
a
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
a
a
a
4
0
a
0
0
Table
B3 fa Dunmore
STATION EAFFAOUL EAFFCQPE EAFFJUVE HARp SPEC ROTISPEC COPENAUP
Cll.12~S
DI8.12~N
D16.12lN
C12.01HS
D12.01lS
D19.01~~
O19.01LN
027.01HS
D27.01LS
D03.02HN
D03.02lN
011.0·3HS
Dll.03lS
D17.03HN
D17.03lN
026.03HS
D26.03LS
002,04~N
002.04lN
OI0.05HS
010,05lS
017.0SI'lN
D17.05lN
024.05HS
D24.0SlS
D31.05HN
D31.0SLN
D08.07HS
026.08HS
C06.09HS
C06.09lS
C13.091'1N
D13.09lN
020.09HS
C20,09lS
027.09HN
027.09LN
003.11HS
D03.IIlS
C09.11HN
D09.11LN
O16.11HS
D16.11LS
023.11HN
D23.11LN
CI5.12t1S
C15.12LS
C23.12HN
C23.12LN
4b
2b
0
358
159
20
57
4
17
23
850
77
194
2
858
42
194
308
495
522
12
b
5184·
1739
3152
205
5812
77
0
0
15
0
7
4
26
2
5
202
0
178
21
235
189
240
12
154
253
120
155
209
879
38
408
612
584
32
89
50
176
165
773
230
1248
2585
194
2134
5634
16538
6292
158
25990
21560
7521
33673
2780
52830
362
0
0
6
9
27
14
48
6
137
783
4
6405
379
2119
910
762
46
812
456
772
64
980
1804
223
1209
5784
403
168
678
1107
200
S36
1766
1367
414
2665
1015
8686
6314
162762
2066
187
17628
232
1738
84564
3476
3b420
SIS
. 0
0
27
30
278
0
284
13
235
71
50
406
3577
92
8985
344
30
30
1407
578
183
183
293
204
496
1511
28
37
26
75
13
22
25
336
8
73
220
119
68
4981
b8
2127
69
1695
18
11382
500
0
143
0
9
15
34
653
0
2
2
17
36
0
6
66
115
4938
82
142
20
88
21
145
379
10b74
5269
74867
48180
1145
752
47019
12912
47881
49522
130
96
0
0
52144
309
204
2921
68
0
63
0
783
11956
12
2572
0
0
0
0
0
0
0
50
0
7
45
0
182
3062
111
7157
64
1597
2245
438
0
0
589
111
0
0
0
0
0
0
1484
427
46
491
7336
648
1393
2022
354
22
717
0
1500
0
741
0
180
342
1023
68
1354
162
1657
391
9
~O
81
0
208
471
376
4254
10
5b
261
10490
6
4034
32
105
153
Table B4 al kincardine
STATION ETEOLARV NERELARV MARA~iIRE MARAEGG5 MARATROC BIVALARV
Kl1.12HS
K18.1211N
K18.12LN
K12.0111S
K12.01LS
K19.01HN
K19.01LN
1<27.01t~S
K27.01LS
KOl.O.2H~
K03.02LN
Kl1.03HS
Kl1.03LS
KI7.03H~
K17.03LN
K26.0311S
1<26.03LS
K02.0411N
K02.04LN
Kl0.05HS
KI0.05LS
K17.05HN
K17.05LN
K24.0SHS
K24.05LS
K31.0SHN
K31.05LN
K08.07HS
K26.08H5
K06.09HS
K13.09HN
K13.09LN
K20.09LS
K27.09HN
K27.09LN
K03.11HS
K09.11HN
K09.l1LN
KI6.11I1S
K16.l1LS
K23.11HN
K23.11LN
1<15.12HS
K15.12LS
K23.1211N
K23.12LN
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
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
6
0
0
0
250
0
0
36
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
.0
0
0
0
0
0
0
0
0
14
0
q48
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
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
2379
502
15
0
1219
903
3
l
0
14
181
0
0
0
0
0
0
0
0
0
0
32
4
0
0
0
0
0
4063
l
0
361
0
26
0
0
0
0
0
0
0
0
0
0
18
0
0
0
0
5
0
9
0
26
0
0
0
20
2
0
0
0
t0604
4414
35806
198
3535
1929
t08
119
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
7
0
59
0
b8
b
16
2
248
14
0
0
"3
0
0
0
0
'0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Table B4 b, Kincardine
Kll.12HS
K18.12HN
K18.12LN
K12.01HS
K12.01LS
K19.01HN
K19.01LN
K27.01~S
K27.01LS
K03.Q-2HN
K03.02LN
Kl1.03HS
Kll.03LS
K17.03~N
K17.03LN
K26.03~S
1<26.03LS
t<02.04~N
Ka2.04LN
Kla.OSHS
Kl0.05LS
1<17.0S~N
K17.05LN
K24.0S~S
K24.05LS
K31.05HN
1<31.05LN
1<08.07~5
K26.08H5
K06.09~S
K13.09HN
K13.09LN
K20.09LS
K27.09HN
1<27.09LN
K03.11t-1S
K09.11HN
K09.11LN
K16.11HS
K16.11LS
K23.11t-lN
K23.11LN
K15.12HS
K1S.12LS
K23.12HN
K23.12LN
0
0
0
a
4
0
0
a
0
0
0
0
0
0
4
5
6
4
2
10
0
0
2
0
o·
2
0
14
0
0
0
0
0
0
0
a
0
0
a
0
0
0
0
0
a
0
0
0
0
a
0
0
a
a
0
0
0
0
a
a
a
a
0
a
a
a
0
22
0
0
2
38
78
"
a
0
0
32
26
22
40
27
0
4
9
0
54
5
34
a
38
31
113
11
47
16
406
3
104
8
695
. 2
16
0
3bb
127
1502
630
0
0
0
2
0
57
26
31
17
6
6
0
0
0
a
a
0
0
0
a
0
4
0
18
0
0
0
0
0
0
a
0
0
0
0
2100
0
13
0
a
a
a
0
0
a
0
0
0
0
a
a
0
a
a
a
a
0
a .
34
0
10
4
60
0
20
a
a
a
0
0
a
4
0
32
34
16
0
2b
.147
16
0
141
29
7
2
0
0
0
0
4b
0
0
0
0
0
0
0
a
0
a
199
0
4887
48
615
107
14480
189
61
0
1556
20
3
0
12
0
9
11
a
a
8
8
8
0
0
0
0
0
a
a
0
2
7
15
11
0
0
0
0
7
2
0
a
0
0
a
0
0
0
a
0
0
a
0
0
0
0
0
0
0
2
0
0
0
0
a
2
0
Table B4 c I Kincardine
Kl1.121-13
K18.1ZHN
K18.1ZL~
KI2.011-13
K12.01LS
K19.011-lN
K19.01LN
K27. 0 1t~3.
K27.01LS
KOJ.02MN
K03.02U~
Kll.03MS
KII.03LS
K17.03HN
KI7.03Lt-4
K26.0311S
K26.03L3
K02.04HN
K02.04LN
KIO.05HS
Kl0.05L3
K17.05HN
K17.05LN
K24.05H3
K24.05LS
K31.05HN
K31.05LN
1<08.071-13
1<26.08t-43
K06.091-15
1<13.09t-4N
K13.09LN
1<20,09LS
K27.09HN
1<27.09LN
K03.11113
K09.11HN
K09,11LN
K1I>.IIH5
'<l6,lllS
K23.11HN
K23,11l~
KI5,lZHS
1<15.12LS
K23.121-1N
K23.12LN
CALAHELG
TEMOLONG CENTHAMA
STATION PSEUELON
0
0
0
4
0
0
0
0
0
0
14
"0
0
0
5
0
0
0
0
0
0
0
0
0
172
25
1&
2
0
218
2
2
0
42
0
12
0
10
32
22
0
2
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
47
4
2
0
0
0
2
0
0
4
0
1"
0
6
0
6
0
0
0
0
0
0
54
e
0
0
0
53
0
16
0
0
18
136
0
0
0
0
53
0
0
0
0
0
0
2
0
2
0
4
.
40
6
32
b5
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
65
2
0
0
0
o .
2
0
10
0
2
0
2
0
011<00101
0
0
0
0
0
0
0
0
0
lJ
48
SAGIELEG
5790
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
277
0
0
0
0
0
0
0
17
0
0
0
.0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
Table B4 d I Kincardine
STATION ACARINER ACARTONS ACARLONG ACARCLAU ACARCOPE OITHSIMI
Kll.t2t-4S
K18.t21-1N
K18.12LN
K12.0ltofS
K12.01LS
K19.0ltiN
K19.01LN
1(27.01t~S
1<27.0.1
LS
K03.02HN
K03.02LN
K11.03HS
Kll.03LS
K17.03HN
K17.03L~
K26.03HS
K26.03LS
K02.04HN
K02.04LN K10.05HS
K10.05LS
K17.05H~
K17.05LN
K2'1.05to1S
K2'1.05LS
K31.05HN
K31.05LN
K08.07HS
K26.08HS
K06.09HS
K13.09HN
Kll.09LN
K20.09LS
K27.09H~
K27.09LN
K03.11HS
K09.1IHN
K09.IILN
K16.11HS
K16.11LS
K23.11HN
K23.11LN
t<IS.12HS
K1S.12LS
K23.12HN
K23.12LN
0
0
0
0
e
0
0
4
0
31
0
0
0
404
189
0
18
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
4
0
0
8
0
0
163
4
226
0
135
0
0
0
0
0
0
0
0
0
0
118
0
0
0
0
32
17
0
0
0
0
0
5
255
0
0
0
0
0
0
0
0
0
0
0
0
2'1
0
0
0
0
18
4
86
0
0
0
0
0
0
0
10
0
7
0
0
0
0
0
0
0
203
6
228
0
0
0
22
0
0
0
0
0
-0
0
93
0
0
0
0
26
0
0
0
0
0
39
0
0
0
0
0
0
0
0
0
0
0
0
0
40
28 .
0
7
10
0
7
104
0
O'
·0
0
48
0
0
0
0
0
0
2
0
0
31
4
'I
0
0
22
4
0
0
2
0
0
0
59
0
15
0
0
0
0
0
3
0
0
2
0
6
0
0
0
0
2
0
0
95
0
3173
24'
26b8
0
55
32
178
102
66
36
13
17
2
2
699
264
2
lSI
2
505
2
6
7
265
0
0
0
18
17
0
0
0
4
5
0
12
2
"00
0
2
0
0
0
0
0
0
16
'14
29
118
7
3
0
Z
0
410
2
II
121
6
256
18
41
7
38
l
Table B4 el Kincardine
STATION
NEOMINTE
Kll.121-1S
1<18.12HN
1<18.12lN
1<12.01HS
K12.01lS
t<19.01HN
K19.01LN
1<27~OIHS
K27.01LS
KOl.02totN
KOl.O.zl~
Kll.03~S
Kll.OlLS
1<17.03totN
K17.03L~
K2b.03totS
K26.03lS
I<02.04~N
I<02.04LN
KI0.05HS
KIO.05lS
K17.0StotN
KI7.05Lt~
KZ4.05HS
K2l4.05LS
Kll.OS"'~
t\ll.0SLN
K08.07t~S
t<26.08HS
K06.0q ...
S
Kll.09HN
K13.09lN
K20.0qLS
K27.09totN
K27.09lN
K03.11 totS
K09.11HN
KOq.l1LN
Klb.l1MS
K16.11LS
K2l.11totN
t<2l.I1LN
K15.12HS
K15.12LS
K23.12"'N
KZ3.12LN
OIAPGRAC
DAPHSPEC
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
44
0
0
22
0
0
0
0
BOSMSPEC
CHYOSPEC
0
0
CYCLSPEC
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
l4
0
0
0
0
0
0
0
0
0
0
0
2
6
1
0
9
0
0
0
0
8
0
b
0
0
0
0
0
0
0
0
0
0
0
0
0
b
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
. 0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
. 0
0
0
'0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
·0
0
0
0
0
0
0
2
0
0
0
94
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
3
0
0
0
0
0
11
0
0
0
0
0
4
0
0
0
0
0
0
0
0
0
2
10
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0'
0
0
Table Elf. fa Kincardine
STATION EAFFAOUL EAFFCOPE EAFFJUVE HARPSPEC ROTISPEC COPENAUP
Kll.12HS
K18.1211N
K18.12LN
K12.01115
K12.01LS
K19.0111N
K19.01LN
K27.01HS
K27.01LS
K03.0_2MN
K03.02LN
1<11.031'15
Kll.03LS
K17.03HN
K17.03LN
K2b.03HS
K26.03LS
K02.041'1N
K02.04LN
Kl0.051'15
KI0.05L5
K17.05HN
K17.05LN
K24.05H5
K24.05LS
K31.05HN
K31.05LN
K08.071'1S
K26.081'1S
1<06.091'15
Kl].09I'1N
K13.09LN
K20.09L5
K27.091'1N
K27.09L~
K03.11H5
K09.111'1N
K09.11LN
1<16.111'15
K16.11LS
K23.l1HN
K2].11LN
K15.121'15
K15.12L5
KZ3.12HN
1<23.12LN
0
13
17
13
119
0
12
0
0
0
0
9
32
2
122
14
126
Z
44
12
23
12
34
0
847
24
57
8
0
0
0
7
24
5
2
121
.0
73
8
8
100
00
349
7
63
756
0
49
176
b2
518
12
217
13
0
32
32
6b2
80
100
366
79
2112
20
58
126
877
34
167
347
10032
61
4223
34
4
0
2
7
146
0
9
1183
77
187
18
84
82
178
35
37
ZOb
876
45
189
404
443
1037
20
2b3
144
133
386
82
3482
204
401
21b
1828
2651
339
234
252
363
5256
1642
782
9043
823
3209
0
0
0
0
17
·230
5
19
37
20
9
0
547
0
42
35
125
19
38
45
0
61
2bl
2897
78
56
13
22
18
23
0
82
31
26
0
50
0
200
135
2161
2
383
10
123
2
1106
92
78
316
19775
4376
80900
54]
b4197
535
3444
30504
177133
2283
107
5b
109
609
0
99
0
0
56
94
3203
0
30782
0
0
0
8
0
5b
21
44
14
0
0
0
216
0
0
5
b2
0
9
34
201
14
10
2
676
6
200
0
0
8
51
0
829
"
4b
47
1853
27
137
3601
1708
2b3
1439
341
0
24
44
0
1412
41
12899
0
2227
334
3544
0
b79
454
555
62
12073
903
3652
0
2181
823
721
Sib
1502
567
544
3501
83
298
0
4775
77
1988
141
6820
20
200
235
3824
111
Table B5
STATION
Skinnats
NERElARV
ETEOLARV
Gl1.1211S
G12.01H5
G19.01HN
G27. 0 1I'~S
G03.02tm
Gl1.03115
G17.01HN
G26.03115
G02.04t1N
G 1O. Q.5I1S
G24.05115
G31.0511N
GOb.09HS
G13.09HN
G20.09"'S
G03.11HS
G09. tlHN
GI6.11115
G21.11MN
G15.12HS
&1
0
MARAWIRE
MARAEGG5
MARATROC
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
98bO
0
1231
0
0
0
15
0
4
8
621
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1la83
159277
730
0
0
0
0
0
0
0
0
0
319
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
eIVAlARV
0
0
20
0
0
0
0
0
7
29
0
82
0
0
0
0
0
0
0
0
Table BS b. Skinflate
STATION
LITTCAP5
Gll.12HS
G12. 0 1I"IS
G19.0um
G27.01HS
G03.02HN
Gll.0311S
G17.03HN
G2b.03HS
G02.04HN
Gl0.QS~S
G24.0SMS
G31.05H~
GOb.09HS
G13.09HN
G20.09~S
G03.111-15
G09.11I-1N
G16.111-1S
G23.ll~N
G15.121-1S
-;
LITTVELI
0
0
0
0
0
0
4
0
0
O.
107
91
9
4
4
4
61
CIRRNAUP
78
16
0
17
0
104
9
274
9
bS
138
0
16
23
CIRRCYPR
0
0
0
0
0
TERELARV
0
0
0
0
0
POLYCILI
0
q
12
22
19
0
0
0
0
0
0
IS
0
0
2
54
2
0
0
3771
68
4837
0
9
20
4
49
0
2b
.6
0
8
0
340
0
0
0
12
0
50
14
0
0
2
0
0
0
149
.0
0
2
0
0
0
11
4
0
0
0
0
0
0
0
0
0
0
3
2
0
0
0
0
6
0
0
0
0
0
0
Table B5 ea Skinflats
STATION PSEUELON TEP-10LQNGCENTHAMA CALAHELG SAGIELEG OIKOOIOI
GH.12HS
GI2.0H"S
G19.01HN
G27.01HS
G03.02HN
Gll.03HS
GI7.03t1N
G26.03t-1S
G02.04HN
GI0.05HS
G24.05HS
G31.05HN
G06.09HS
G13.09HN
G20.09HS
GOl.llt1S
G09.l1HN
G16.11tiS
G23.11HN
G1S.12HS
99
31
0
0
0
210
2
88
la
0
83
0
2
2
26
121
1559
1692
74
241
6
Itl
0
0
0
0
0
0
0
0
4
0
4
0
160
2
7
0
0
0
0
0
0
0
11
0
0
0
0
8
0
0
0
0
0
0
0
0
0
0
0
0
0
0
67
6
10
8
8
2
2
0
0
0
8
0
0
0
65
0
473
20
65
0
0
140
29
13
0
0
0
0
0
13
143
78
5
0
0
21
6S
310
0
0
38
8
"
7
0
0
0
0
0
0
0
0
2
0
2
6
Table B,5d. Sklnflats
STATION ACARINER ACARTONS ACARLONG ACARCLAU ACARCOPE OITH9IMI
Gll.12HS
GI2.0U'IS
G19.0um
G27.01H9
G03.02HN
Gll.03H9
G17.03HN
G26.03HS
G02.04HN
GI0.O-5HS
G24.0SH9
G31.05HN
G06.09HS
G13.09HN
G20.09HS
G03.11HS
G09.11HN
GI6.11HS
G23.11HN
G15.1211S
20
8
0
96
0
0
201
102
11
6
1152
16
0
0
0
11
24
6
4
b
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
\)
0
0
0
0
12
41
460
34
51
0
"0
185
0
0
0
0
. 0
0
321
31
0
0
0
0
0
0
36
0
0
242
13
0
0
0
2
0
"
37
229
27
0
87
"
16
454
113
34
"
2391
0
54
243
302
S9
157
68
14
58
144
53
0
200
0
89
0
264
4
4
44
0
49
18
175
748
214
7b
0
7b
Table B~ el Sldnfiats
STATION
Gll.12~S
G12.01HS
G19.01HN
G27.0HiS
G03.02HN
Gl1.03HS
G17.0ltiN
G26.0lHS
G02.0I&tiN
Gl O. O.5HS
G24.05HS
G31.05HN
G06.09MS
G13.09HN
G20.09MS
G03.11HS
G09.11HN
G16.11HS
G23.11HN
G1S.12HS
OAPHSPEC
OIA~GRAC
NEOMINTE
0
0
0
0
0
0
0
0
0
WO
0
0
0
BQSMSPEC
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
0
0
0
0
0
0
0
0
0
0
0
0
0
.0
' 0
0
0
2
0
0
·0
0
0
0
0
CYCLSPEC
0
0
0
0
0
0
0
0
0
0
CHYDSPEC
0
0
0
0
0
0
0
0
0
0
0
0
'0
0
0
0
0
0
0
0
0
0
-0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
Table BS f. Sldnflats
STATION EAFFADUL EAFFCOPE EAfFJUVE HARPSPEC ROTISPEC COPENAUP
Gll.12HS
G12.01HS
G19.01HN
G27.01HS
G03.02~N
Gll.03~S
G17.03HN
G2b.03t1S
G02.04HN
GI0.05HS
G24.0sHS
G31.0SHN
GOb.09I1S
GI3.09~N
G20.09t1S
G03.l1HS
G09.11 t1~
Glb.llt-tS
G23.11HN
G15.12"'S
27
44
0
13
0
187
4
25
11
15
201
21
0
2
2
14
0
1312
4b
60
85
278
4
b9
101
2007
0
13294
0
180
507
4110
2271
3b
40b3
4bOb
580
154
1239
12
5
0
10
911
50
·2483
94
139
3b32
194
0
0
0
0
0
b741
338
227
4b
0
~4s
1288
219b
72b
240
99
1349
859b
3502
'5517
1685
39910
0
b6
355
Ib3
b7
227950
811
1801
Ib28
4b9
bb73
18b4
971bO
5b75
1589
1712
5000
0
1834
0
1;?t;0
55b2
b292
35
1217
0
91
0
0
bOb7
3340
1457
2b09
0
cH9
1831
715
4045
92
17(1
10
0
33b
185
2(1b
33
0
53
Table B6 a. Culroes
STATION
CI1.03"'S
C17.03HN
C26.03HS
C02.04HN
CIO.OSHS
CI7.0SHN
C24.05HS
C3t.OSHN
C08.07MS
C26.lt8HS
C06.091-1S
C13.09HN
C20.09MS
C27.09HN
C03.l1HS
COQ.tlI-fN
CI6.11l-lS
C23.11J04N
C15.121-fS
CZ3.12HN
NERELARV
ETEOLARV
0
0
0
0
MARA~IRE
876
1884
399
0
55
0
0
120
to
1
0
0
0
85
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
'4
0
8
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
BIVAL.ARV
0
0
0
0
0
fo'ARAEGGS MARAT~OC
0
0
0
0
0
0
. 0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
Table B6b.
CUlross
Cll.03MS
C17.03MN
C2b.03~S
C02.04M~
CIO.OSHS
C17.0SHN
C24.0SMS
C31,OSHN
C08,07MS
C26,(}8HS
C06,09HS
C13,09HN
C20.09HS
C21,09HN
C03.11H5
COq.llHN
C16.11HS
C23.11~N
C1S.12MS
C23.12HN
1645
55
2'530
40
1083
23
4343
2349
27q
0
7
0
82
0
e
0
0
0
692
0
0
2
20
0
10
2S3
9
0
5
0
0
0
0
0
0
0
0
0
0
0
134
28
63
79
4860
967
196
0
0
0
2
0
0
0
0
0
0
192
31
4
0
0
8232
271
204
10
S9
477
e
2290
1652
976
399
2244
7
45
13
5
la
12
222
0
4
28
126
2
24
14
0
0
0
0
0
0
0
29
0
0
0
0
23
34
7
6
0
0
0
0
0
0
0
0
0
0
0
Q
0
b
0
2
Table B6
Cl
Culross
STATION PSEUELON TEMOLONG CENTHAMA CAlAHElG SAGIELEG OIKOOIOI
Cll.03HS
C17.03HN
C26.0lHS
C02.04HN
CIO.05HS
C17.05"'~
C24.051-!S
C31.0SHN
C08.07H5
C26.o.e·HS
C06.0911S
C13.09HN
C20.09HS
C27.091-1N
C03.11HS
C09.11HN
C16.11MS
C23.11"'N
C15.12H5
C23.12HN
0
0
7
0
29
0
0
0
0
0
2
0
5
0
674
bb
t03
10
2
2
23
2
6
0
44
0
4
2
0
0
2
2
0
0
56
4
6
2
0
0
2
0
0
0
35
0
0
2
2
0
.0
2
0
0
·0
8
0
0
0
0
2
0
0
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
0
232
0
0
0
10
0
0
0
0
0
6
14
2
0
0
0
0
0
0
0
0
0
0
0
Table B6 dl Culross
STATION ACARINER
Cll.03HS
C17.03HN
C26.03t1S
C02.04t1N
Cl0.05t-4S
CI7.0St-4N
C24.05HS
C31.05HN
C08.07HS
Ccb.08HS
COb.0"9HS
CI3.0Qt-4N
CZO.I)9HS
Cc7.09H~
C03.11HS
C09.11HN
C16.11t-4S
C23.11HN
C15.12HS
C23.12HN
ACARTONS ACARLONG ACARCLAU
0
0
0
0
0
0
0
0
0
0
0
0
0
5
0
23
5
5
0
82
0
0
0
0
0
0
35
0
20
0
0
0
0
0
39
0
2
6
0
0
0
0
0
0
0
0
0
38
0
0
0
0
0
25
0
8
2
2
0
14
00
6
0
0
0
0
0
0
0
0
6
11
0
0
0
0
4
0
0
0
ACARCOPE OITHSI~I
12
20
32
0
286
2
129
2
29
4
b3
34
110
2
0
312
16
22
0
32
0
226
0
11
0
5
39
28
2
28
0
432
8b
44
30
12
10
24
2
8
Table B6 e I CUlross
STATION NEOMINTE DIAPGRAC OAPHSPEC BOSMSPEC CHY05PEC CYCL5PEC
Cl1.0]H5
CI7.03Ht-.
CZ&.03HS
C02.04HN
Cl0.05H5
C17.0SHN
CZ4.05HS
C31.05HN
C08.07H5
C2b.0~H5
COb.OqHS
C13.0QHN
C20.0QHS
C27.0QHN
C03.11H5
COc;.11HN
Clb.l1H$
C23.11HN
CI5.1ZHS
C23.12HN
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
.0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
Table B6 f.
Culro8S
STATION EAFFAOUL EAFFCQPE EAFFJUVE HARPSPEC ROTISPEC COPENAUP
Cll.OlHS
CI7.0]HN
C26.03HS
C02.04t-1N
CIO.05HS
C17.05HN
CZ4.05HS
C31.05HN
C08.07t~S
C26.0~HS
C06.09"'S
CI3.09t-1N
C20.09HS
C27.091'"N
C03.11'-'S
COq.ll~N
C16.11t1S
C23.11HN
C15.12HS
C23.12HN
287
0
0
27
0
0
]9
0
0
0
2
0
5
0
3
2
bOb
b
47
0
100
16
9
7
338
10
1b5
14
5
0
2
9
0
0
Z05
0
404
0
53
0
18
240
14
112
0
14
126
0
0
0
7
0
0
0
62
9
0
0
18
15
34
140
43
54
159
225
48
119
20
4
178
18
167
2
76
101
46
130
167
48
to]
24
0
0
0
0
0
0
0
0
0
0
0
0
404
13
0
776
94
17
Itt.
1769
84
2b3
169
70
302
74
408
176
0
180
895
229
118
113
36
156
404
48
Appendix CI~ooplankton abundance IFigures
Cl
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p..J
w
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en
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CD
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n
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>
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m
r.7'
m
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(,(1
er:1
r~1
j.!:1
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Figure Cl
al
Annual ab.mdance of'mixed group taxa at Fa11in,high tide.
Logar! thmic scales.
'f)B/m'
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~
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rn
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Figure Cl bIAnnual abmdance of mixed group taxa. at Fallln,low tide.
Logarithmic 8cales,Nos/m3
:r.
~l
~-
0
At
""Cl
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..,m
):
coo:.
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m
G,)
m
m
::tI
~.
;.:0
cc·
Ct·
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Figure C2 al Annual abmda.nce of mixed group taxa at South A110a at high tide.
Logarithmic 8cales.Nos/m3
c:.
....;,
r.:,
-fA
..-
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r
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::.:-
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Cr
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--r
1-1
m
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m
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JJ
:>
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Figure C2 bl Annualabundanceof wed
Logarithllic Bcles,Nos/.'
group taxa at South Alloa at low tide.
r::1
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(,-,
r
~.
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At
'"IJ
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m
"
m
C)
m
A1
~.
A:I
01
(11
~l
Figure CJ al Annual abundance ot aixed group taxa at DunIloreat high tide.
Logarithmic 8cale.Nos/.3
Figure C) bl Annual &bmdance of aixed group taxa. at Dunmore at' .low tide.
Logan thaic
Bcales ,108/1l:3
J:
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Figure c4 al Annual abundanceof wed
Logarithmic scales,Nos/m3
group taxa at Kincardine at high tide.
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rr.
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r.
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).
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Figure c4 bl Annual ahmda.nce of mixed group taxa at Kincardine at low t1de.
Logarithm1c ecalee.Noe/.3
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Figure CS a. Annual abundance ot
wed
group taxa at Sk1nnats
at high tide,
Logarithmic scales.Nos/m3
.'
t·.)
':' UI
to ...J
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•
-t
m
(j)
m
Al
Figure CS bl Annual abmdanee of wed group taxa at CUlross at low tide.
Logarithmic
scales.Nos/.3
Gl
Q
~
U1
Q
Q
..r-f
0
-~
..r...
r.:n
.1;.""')
-_
0
::::0
.c..
n
o
:-rJ
m
1---1
(/)
""()
o
o
in
(}"J
~
I
C)
:J'':
-1
1-4
f::::J
ITl
~
I
~
en
I
-I
1-1
CJ
m
Figure c61 Annualabundanceof neritic copepods at South Alloa.high and low
tides.
Linear scales.Nos/mJ
t··.) Jl. ITJ CO
oQ
lSI C5:1I!;)
-" -" -" _. _,. ".J
(SI I\.) ~
~~ cs;)
0")
lSI C5:l
CD lSI
(SJ
Q
n
Cl
"1J
m
T.I
Cl
o
(.f)
Cl
C
z:
_. ......-" r·J r··.)
('~i(J)
f~)
t·)
+.
(D
f\_)
Ul CO -
..j::.
~S:I '::1 tl:I lSi t£, lSi 1St lSi
~.J( ...
)
~•.:: ~I'
_~
~S) lSI
(SI ~.~,
cc'
((I
r,_,
ISJ
lSI ~,
f~.±-~~~
t··._)
::r
(oJ (..:, ~
(T) r':';;:l ~
((I"'._)
r..:r_,IS)
~:S:II:;:) I:;:) I~I
I~
__~~~~~~~
r.~:;)
.
r
-~
::J:3:
::p.
G.i
.:=
(J"J
:r:
m
Cl
I:;)
:k-t
,=
in
-l
al
t>
.q
r
r.l)
1:;:1
0
:?.:
r._,
1-'
jj
m
tJ
,......,
"1J
z
,T
-"' -"""r'J ".J
,....!
--I
1--1
0
IT1
I
1-1
(1.1
I
-!
t-1
0
m
Figure C? I Annual abundanceof nerl tic eopepods at Dunmore,high and low tides.
Linear scales,.os/.J
-I
a
r
I::J
Z
.r.J:)
1-1
r
r.""')
1-1
(:::.
'''fJ
::T.)
"7
~
r,:O
r-1
(J)
......
Ci:l
1',.1
....
__.
....:. .....:.. ...... ....:. t·, 'I
r- '1 t, .,
r.) Id' .•..1 I~I ".,)(.11 ~-.1 I~! t,::!di
I:::'
~_n1..::.,1en
I"
(n 13) ':." I:;:J I~n
~s:.'
(.J1
I~
r:5l
(!:I
r~1
I~
I::;)
~:S:I
z
n
~.
AJ
o
z
m
(""J
:.:r:
:;:p.
:-=
::;p.
-f
,
'-
Cr)
.'-
r::.q
r
c)
:7"':
-1
1-4
0
m
Figure
cB.
Annual abundanceof neritic
Linear BcaleB.NoB/.3~
"I
~
(j)
I
~
1-1
Cl
ITI
copepoda at Kincardine. high and low tides.
~
t'..'1
a
~l
r
er
Z
en
,_
(TJ
(")
0
...:::.
AI
(.(1
I'-t
(()
\.I
I,:)
"1J
m
-"
I.)
r.::!::1
I~
a
(r')
t-<J
(r")
A
--- _..... -- t··.J
-iL (T') (Cl 0 r··.J -+ Ij) m ~
r~ 1::1 J'S:) I::;:' I~'_' I~
(S:I (:::' I:::' r,.D
~::;:j 1:::1 I~ I~I I~' I:D I::) :':;:1 1'::1 (S)
--..:. --:. ~
r-..)
'S)
1',) .../:.
lTI
t:1 lSI lSI lSI
0) r.:!:) 1'-,) .../:.
IS'
r.:1
1:sJ
~I
__:. _..:..t-.J
en oJ
I~
lSI (~.' ~
("",)
1'--:1
::r:
::""
-'"
.:. ..
(I')
OJ
:.3
_:;-......f
I:":'
r.J::I
~.
T
'-I
r,S)
(i)
T
-l
t<1
~
0
m
Figure C91 Annual abundaneeof ner.ttie eopepcds at Sk1nflats at high and
low tides. Linear ecales,Nos/m3
-
~.j
en ....
-
(JJ .J:,. U1
J CO (!) I:s:l
0 0 lSI ':S) 'SI CS' IS)
crI '~I lSI 'SI
~~~~~~~~~~_.~~
~
II
r
f;:)
Z
(j')
D
C
r
_.
.....
fr.
t··.)
I..•.)
1:=.
~
en
Al
';T!
Cl
-;.•J
({J
rn
•
:::c
-.J>
(f)
l:=-
I
~
(j)
::I.
-I
t-1
o
m
Figure C10, Annual abundance of neritic
copepods at Culross at high and
3
low tides. Linear scales,Nos/.
r-.j
IS:)
'S,
';S:J
w
'SI
~
0
c:J
):..
-
1",)
•
11
-A
IS)
Q
r
,=,
:>
~
r
en
t-1
1-1
(II
W
~S)
"0
m
+
~")
GJ
lSi
tl.~1
Q
:::J:-
l=
((:1
'J1
:"..
(I:!
((I
Z
m
c::;
A.1
=
.........
(f)
~.~:
.~>-
fo·..::
CJ
+
r··...,
t···)
~
~
1-l-<
r.=:t
..:.....
--:..._
1)-'
I::J)
-::'"
:~.
!..i)
--"
C!:)
en
:.'0
III
~--:,.
_..
I~!:
r..:O
.~
~r-~
I
~
::s::: CD
Cl
_.......
(
r~1
--f
t··.·,
r-,
I
~ -i
Cl 1-1
m
0
m
Figure OIl, Annual abmdanee of Aeartia spp. at South Alloa. high and
low tides.
Linear . 808.les .NoS/1l3
__.t·) (,J -l.I..
::) ~~: 'S' lSI tSJ
en eT)
'-..,J
t~'ISJ 'Si
,:0
.....
(D
__. ~,) (,J ~ (.0 Cl') ...~ ((I (D ~I
Q 'SJ Q lSI 0 (~lSI ,3) 'SI (:i:.l Q
c:;)
(1:1 '3:1 ~~
•
'Il
t-1
T1
t-1
r
I~
.n
((I
~:F-
).~
C)
)
.....
z
::0
--I
1""'1
:J:t··:
_..
r~1
IS'
I~' (S)
t°.,J
(....)
~
IS)
IS:'
m
is)
lSI
lSI
13)
(::'
,3)
(1":1
"'-~
I':::'
rIl
'SI
(5:.'
__.
IS)
(~I
:l:-
(::' ':.:'
t··J
"7
U)
(:::1
(_"J
~
IS)
':SI
lSI
13)
(.r1
'.:n
.::::"1
1':::1
1::.1
(S)
'-' •..1
'::,3)
-:.P'
-
r-
-1
i:::'
t··.J
I::'
-+,
"_:1
-,
.~r:;)
,-<
:.J::-
:t'J
!"n
(0
'-::1
-'I',
r
'«:
r'::l
~:.
I
.Li
-r-
.J..
-f
r'-.1
~
,~
-!
.-1
in Cl
111
Figure C121 Annual ahmdance of Acartia spp. at Dun.ore, high and
low tides. Linear scales,Nos/.3
t·,)
.n -;-J
~s:;i:~!1 r:)
(.11
__...-.:.-"
1::1
13)
t·..)
(J1
!::I
I~n
_. ~'J r',J !',J
'-.J lSI r,j .n
I;_n (:~l 'Jl (.5)
rs)
-
T'J
(,j .j).. (..J1
lSi
I:::)
I~) ~:;)
'SI
m
.......
J
'<,
m ill LW
G) 1;:1 I:.:;:) ~:::) 13)
>,
...
( )
):-
:D
--I
-
1'-1
):-
(,1)
z:
rn
t··..::
.
_J:. •
r....p.
':=J
-:::"
,(7.,"1
r-,
-' ,
i
i":'(
I
::n
m
-:;:
'--=1
'p~~-:)
-
T
,;::)
1-1
-c:
G'J
--i
r·.:,
~".
......
CJ
I
--I
1"""1
iT1 :.::J
III
Figure Cl). Annual abundance of Acartia spp. at Kincardine, high and
low tide.
Linear sca1es,Noe/m)
t·.,) ,
):-
•m
1-1
-+--
11
>..
1'<1
()
r
4<
>-
'-
t-I
r
I"r"·
_I .'
I~
(f)
CD
1"""1
(()
:;F-
co
1::'(1
1-1
Z
--:,
m
__,
N
Ci:l
--.....-
'=1
(J')
~.J
r...J
.....
".J
....;. ....;.
~.J
l ..p. +<,,":1 r.....
r.:..n Cl U1 I:;' U1 Cl (Jl I!:I (.11
IS
'S) rSl 'SI
I:;:! 1;s:J I~
1;s:J
I::::, 'SI
IS)
Z
11
(-',
r~,
( •.11
r~'
rS)
lSI
__"
-"
I::;:'
en
lSI
r0
I:;)
~!I G
t-.J
lrJ
r
I~I
I::;:)
IS)
-I
en
if)
1::5)
>.
I',_)
I~
~
r_:::)
:.z
CJ
0)'')
o-
>
.
r
-I
Z
):-
(I')
:::D
rn
.......
0:1
......
~
(J")
I:=I
1-1
J::!:I
G1
:c
-I
" •..'1
t-I
0
111
Figure 014, Annualabmdance of Acartia epp. at Skinflate. high and
low tides. Linear sca1ee.Nos/m3
........
-:..
..... kl I~IJ~ 01 0'1 .....
J (0 ({J rS:1
G) (:;:1Cl (:;:1
IS) IS.1:-:;:"J Cl
_.
~::rISJ
G) Cl cS)
):-
•
1'-.."1
CO
~
II
,=,
(oj ..A (J1 CT) '-J if.J
1;9 (SI >SI ISJ lSi
uJ
a: !SI
G'
13)
>a
0
~,
t-f
r
1'<1
13:!
(n
r
:;.:.
r.=
oJ)
I'"""'!
(()
::J:.-
~
Z
01
m
_..
7J
-,.
_.
r.:!;)
.......
in
::t:-
1'<.)
!'-),
~
Cl
z
u·)
):-
~
-r::J:Cl
Z
-...
r_::_'
en
o~
In
CO
:-:I
I_
r.:.I')
i>
I
1-1
.:J.l
r~'1
I
to,,'!
t-t
-i
Cl
m
Figure CIS. Annualabmdance of Aca.rtl& spp. at Culross, high and
low tides. Linear acalestNos/m)
0
t,,)
u
G)
:::D
~
::;,:.
1-.
r
t-"
G,I
......
rj)
(Cl
r..!.1
k)
.......
0")
-
;=' en
(.,)
'SI
...J:;.. 1:r:1 ~.::J I:.LJ 1:;::1 1',,) (I.)
Ijl ,:;;)
I~n 1:5:) I:;:, '~n
en
CS) +-'---A.--II..-"'__'_-'-......L--J_.L.-I
r.:..rl
r~'
en
t:!l
I~,~_~
__
L- ___'__~
lSI
__~
.....:1
en
I
-f
t-1
C)
t>.;;,
'lJ
t",~1
Cl
::z
.......
o
m
.~:
Cl
((j
+
I
-P
Z
):.
i:~-.
I::.:J
:lJ
~
0
(()
'.:CI
((.
(::,
r'::::',
'-'
'1J
t·,,J
Figure c16 al ~ual
abundanceof freshwater taxa at Fallin at high tide.
Linear scales,Nos/m3
to,)
lSi
to.o)
0'1
0
r-oJ
a
r.:J.;!
:::0
-4
:):>
-(""')
r
GI
(J')
co
-"
11
r~1
"J>
rr
.
~.)
......
z
-
(I)
~
1:r:1 <I (J:I IS) 1',) (I_-I
IS) en IS! Ul IS) (n IS) I~n tSI IJI
en
r5:.'
t1)
-~
IS)
en
lSI -j.'--I.-L-..L-...L--L.-I..-J......I~J.....I
Cl
r~..)
co
..
l:tJ
lJ
~o.)
1::1
..,.-,
iT,
==
....p.
...,..
..._
>-4
):-
m
I
..-::~
0
I~
':T.l
I-<
0)
0
.:j?
"0
(C!
0'
::'::t
r.:i'
t._)
~'o.'l
OJ
Figure c16 be Annualabunda.nceof freshwater taxa at Fallin at low tide.
Linear scales.Nos/m3
...:..
.
~.)
r.~
lSI
r::S:1
t·..:;
_. _. _. t-.J f'>.) f'>J
'.. J & r<:r U1
~,(_" Q1I1 C~
'Si en 'SI
t·,.) Ul '''"~ IS) f'>.)
en
'SI
r..n
Cl
..G)
Cl
:)!.
IJ
:r:
-J:;..
2
AI
...j..
:;»
C)
>-1
:~.
tT'
.......
r
0.1
......
(.1')
(1:~
(0
Tl
~)
~:
__;,
-"
I:::
r.ZI
((I
•
>r
r',J
r
Cl
i:,
t8
".J
1:::;:,
(_,j
lSi
-~
lSi
r.:.n
t-.) (I,) ~r':'l1
~~,
r.SI lSi
1:2:1
1::::1
m
Cl
+
-:,,-
IS;!
r.='
en ~.~0:1 U,)
G:I
-...:.
1--1
I~
G1
lSi lSi (~) lSI
1",)
::c
-;
C")
I
-:::
,'f)
::r
......
~
- Gl 'S)
>:c
......
0
m
0
4(II
I':::>
::T.J
~
0
,
..
I.-,)
0::
rn
I::'
:"~I
1
OJ
'-0
t,J
Figure C17 a. Annual abundanceof freshwater taxa at South A110a a.t high
tide.
Linear sca1eS.Kos/.3
):-
r
r
I:)
>....lo
en
-
r:S:_,
r:::iJ ,:;:)
en
t-,)
(,j ~
r':'l1
I:;:)
ISJ
~:s;J lSI Q ':SI
I~
'<,
.l (() r.j) IS!
c:S:'
IS)
--f
t-I
(""J
m
t-<I
I:J
o
m
-(,
Cl
'J")
_p-
z
:>
I
O
AJ
1--1
rn
Cl
((J
'"lJ
0)
-1J
~l)
r-.J
Figure cl? bl Annual abundance of freshwater taxa at South Alloa at low
tide. Linear scales,Nos/.3
~~......::..~
-
~ L"
0
y.
'"0
J:
?"
.-.
iJ..:1 .A
IS)
(J) '--~
Ul Cl
UJ ~
h) 6.)
en IS) Ul m en
t··,J
Cl
D
(j)
A:I
...j...
">-
~
:l:-
1fI
IS)
C")
~
-
tJ:i
r
U)
rj)
T1
::0
.._;,
'::'
~
-"
C!.)
,=-~
0
.:::..
,....
-,
i
-l.
Cl
m
"
J:
~
rSJ
I'-,j
Ul
r~1
(.~
13)
Gl
:c
-t
C")
I
-:::
0
m
0
r.::J
.~
I~
CJ
((j
"-0
""D
Figure Cl8 a. Annualabmdance of freehlfater taxa at DunlIoreat high tide.
Linear ecales,Noe/.3
--:.
~
..
j
-
'-::1
en
iJj ~
is! 01
11)
IS)
....:a __:.. ~
···.1 UJ lSI r,j w Ul
(n G) C~ lSI (11 lSI
o
:):-
0
"1J
"(J.l
z
:):.
::r:
:::r.J
-C)
r
(I')
T1
':<J
~::
Cl
1=
Z
r·.J
:::3:
r·J
Cl
~J
m
~!I
I:"'~
rSI
hJ
()l
r
Ct
:s:
I:;:)
-i
t-1
tJ
:r:
~:
0
m
C)
r--'
:.:r,:.1
1-01
CJ
({)
''1;1
v
Figure C18 b. Annual abundance of freshwater taxa at DwlIlore at low tide.
Linear scales,Nos/.3
y ...)
,!:"
Cl
'::'
U
~:,
(~f
'SI
h:.J
'S'
0
0
:):-
a
(j)
iJ
:r:
:::u
:::J:-
z
n
......
r
......
t-!
:l:(r:J
(J)
(X,
"T1
::n
...........
......:.
'
...
1;::::1
1
A
t-1
Z
r,")
n
..
):
AJ
o
I'-'-i
Z
i::l
':...n
'S)
( ..n
t··..:"
r:,
:os.'
U1
r.,s)
,-,.)
I:;)
m
-.:r:
UJ
n
CO
,=,
I
({J
0
-:;.
::z
_.,
z
.1'::;'
AJ
,-
~~.
:r:
--I
I-~
Cl
m
Q
«)
IJ
'iJ
.~I
r,.:s:1
t'~1
Figure 019 al Annual abundance of freshwater taxa atKincardine at high
tide.
Linear sca1es.Nos/.3
t,J
is}
',SI
IS}
l.,_i
'31
s:
fSJ
en
r.:s..,
r.5J
0
:~.
r:'SI
Cil
lJ
tJ
~
IS)
.A
U1
13)
I,S)
t··.)
0
a
'iJ
I
Z
~
=~
(j)
AJ
...j.L
:>
C)
.......
r
.......
•:J)
(f')
(0
Tl
::OJ
-.::........
•--!'
cr,
7\:
......
Z
Cl
r'J
>:lJ
CJ
......
Z.
m
-~~
l~
en
,~,
I···.)
(..11
r.~,
m
Cl
t'·)
(SI
(..~
I:;:)
tJ
1"'.1
40-
~
:,..
(SI
lSI
,:.0
=
.....
.....
(.~
I
.-.<::
0
c.,
r
Cl
~..:
-I
.....
CJ
m
AJ
>--4
IT!
C.J
(()
OJ
1:.1
IJ
r!)
Figure C19 b. Annual abundanoe of freshwater taxa at Kinoardine at low
tide. Linear soales.Nos/m3

Roddie`s Thesis - University of Stirling

Transcription

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