Journal of Plankton Research Volume 4 Number 3 1982

Composition and differential distribution of zooplankton in

Arcachon Bay
Jacques Castel and Claude Courties
Institut de Biologie Marine, University de Bordeaux I, 33120 Arcachon, France
(Received July 1979; accepted March 1982)

Abstract. Specific composition, distribution and seasonally of zooplankton in Arcachon Bay was
studied along a salinity-temperature gradient from coastal neritic to estuarine conditions. Zooplank-
ton numbers ranged from 23 m" 3 to 127 000 m~3 and biomass from 0.08 mg dry wt m~3 to 525.5 mg
dry wt m " 3 . The maximum density and biomass of zooplankton occurred between April and June in
the three stations investigated. Copepods were the dominant taxon comprising 71.4% in number and
91.7% in biomass of the total zooplankton. The three different water masses of the bay were

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characterized by copepod associations of species. The most common meroplankters were Cirripedia
nauplii. Other important groups were polychaete, gastropod, bivalve larvae and the larvacean
Oikopleura dioica. Biomass increased significantly from the oceanic water to the inner neritic water
mass while the species diversity was decreasing. As mean densities and biomass increased, fluctuations
were more important. The community structure has been measured as a mathematical description of
the species-abundance relations. Total zooplankton and pelagic copepods fitted both a geometric
series and a log-normal model. In general, a closer fit to a log-normal distribution was due to the addi-
tion of several assemblages of species. The fit to both models was affected by the salinity-temperature
gradient. The community structure was especially unbalanced in the inner bay, directly subjected to
influx of fresh water.

Introduction
Very few workers have studied the zooplankton of Arcachon Bay. The study
from Lubet (1953) mainly considers the qualitative aspect. Only the specific dis-
tribution and seasonality of polychaete larvae is well-known since Cazaux (1973).
This author found an important endogenous annelid meroplankton occurring in
the bay. It is characterized by its abundance and short periods of production.
In a recent study (Castel and Courties, 1979) we made a comparison between
the open bay and lagoonal impoundments, considering only the copepods and
polychaete larvae. These impoundments, situated on the East coast of the bay
('Certes' in Figure 1), receive allochthonous zooplankton populations when they
are flooded with sea water. These planktonic immigrants cannot colonize the
ponds for a long time.
In the present work we describe the composition, differential distribution and
seasonal pattern of zooplankton living in the bay (particularly copepods). Distri-
bution laws of specific abundance are adjusted to the data (numbers and
biomass) to detect variations of the community structure from oceanic to
estuarine conditions. The interest and validity of the most common used models
of species abundance patterns are discussed.
Description of area
The Bay of Arcachon (44°40'N, l°10'W) forms a triangular embayment on

© IRL Press Limited, Oxford, England. 0142-7873/82/0403-0417S2.00/0. 417

J. Castel and C. Courties

the South-West Atlantic coast. The channels and intertidal areas cover 155 km2,
but only 40 km2 of the bay are subtidal. The 'pass' connecting the bay and the
ocean is a narrow channel through which the oceanic water enters at each tide.
The Arcachon Bay receives direct fresh water input: the Leyre river to the
South-East, Lege canal to the North and Cazaux canal to the South. About a
dozen small streams drain into the eastern side of the bay. The volume of water
introduced during mean tides is - 370 million m3. The mean discharge through
the inlet is - 1 7 000 m3 s~l. Hydrographic studies of the Arcachon area
(Bouchet, 1968) have shown that the bay is not homogeneous since the waters
undergo temperature and salinity variations directly proportional to the distance
'upstream' from the inlet. The bay is flooded with neritic coastal waters. In the
inner bay the waters mixed with fresh water are inner neritic waters. The mixed
waters of the intermediate part of the bay constitute the intermediate neritic
waters.
According to this classification of the water masses, three zooplankton sampl-

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ing locations were selected (Figure 1). Station A was located near the inlet (water
depth, 15 mat high tide). Station B was situated in the main channel in 4 - 5 mof
water. Station C, in the inner bay, was located near Spartina marshes (1 m depth
at low tide).

Material and Methods

Field procedures
Sampling was performed monthly from February 1976 to January 1977. Two
tows were made at each station: just before and after high tide at station A, dur-
ing ebb-tide and during the flow at station B, at the end of ebb-tide and at low
tide at station C. This was done to have the greatest chance to sample each water
mass.
Zooplankton was collected with a WP2 nylon net (UNESCO, 1968). The aper-
ture of the mesh was 200 /im. Each tow was ~ 2 min long. Samples were obtained
against the current (0.30-1 m s~ *). The volume of water filtered through the net
was estimated with a T.S.K. flowmeter. Filtration coefficient was calculated in
comparison with a flowmeter placed outside the net mouth. The filtered volumes
and the filtration coefficients were usually 4 - 2 5 m3 and 75 - 9 5 % , respectively.
The catch was preserved in buffered 4% seawater formalin.
Surface temperatures and salinities (Electronic Switchgear bridge) were
measured every 2 h during the tide cycle.
Laboratory procedures
Samples often contained a large amount of detritus which was separated from
zooplankton by filtration through 100 |im mesh sieve. Densities of zooplankton
were determined by volumetric subsampling with replacement. One to four sub-
samples were counted in total. Copepoda, generally the most important group,
were identified to the species level from aliquots of at least 100 individuals.
Biomass was obtained by weighing three to twenty animals representative of each
taxon on a Mettler ME 22 microbalance (sensitivity: 0.1 /ig). Before weighing, the

418
Zooplankton in Arcachon Bay

LEGE

5 km*

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Fig. 1. Map of Arcachon Bay showing the three sampling sites (A, B and C).

animals were rinsed three times in distilled water and dried at 80°C for 24 h. Each
weighing was replicated three to six times.
Data treatment
Species diversity was calculated using the Shannon-Wiener information func-
tion H' and evenness J (Pielou, 1969) with natural logs.
Two distribution laws of specific abundance have been computed for each sta-
tion averaging monthly estimates of densities and bio mass. The frequency distri-
bution of species has been tested against the most commonly used models: the
log-linear (Motomura, 1947) and the log-normal (Preston, 1948, 1962) distribu-
tions. For further information on the mathematical treatment and the ecological

419
J. Castel and C. Courties

interpretation, see May (1975) and Daget (1976). In the log-linear distribution or
'niche preemption model', there is a linear relationship between the logs of densi-
ties (or biomass) and the rank of the species. The antilog of the slope of the distri-
bution, m, depends on diversity.
Preston (1948, 1962) found that the frequency distribution of living organisms
was better described by a log-normal distribution where the logarithms of densi-
ties (or biomass) are randomly distributed around their mean with a standard
deviation s = V l / m ' , m' being a constant related to the diversity of the com-
munity (Edden, 1971). The diversity is maximum when s = 0. The lower the value
of s, the higher is the diversity. The quality of the adjustment between observed
data and calculated values has been tested by the Bravais-Pearson correlation co-
efficient r (according to Inagaki, 1967) and the chi-square evaluation (for distri-
butions of densities only). This latter was not used as a test for goodness-of-fit
but as an index of distance allowing comparison between the two models. The
values of the calculated chi-square are very high; the table of probability cannot
be used directly in this case. However, since the total number of species is nearly

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the same at each station (when frequencies lower than 5 are pooled), the degrees
of freedom are about the same and the chi-square values can be compared.
Results
Station characteristics
Considering the temperature-salinity diagram (Figure 2) of the surface water
during the year investigated, the three water masses of the Arcachon bay are
clearly separated. There is an increasing gradient of salinity from the inner neritic
water to the inlet. Salinity was most variable at station C (8 — 30°/oo) compared
with station B (22-34°/ o o ) and station A (30-35°/oo).
Surface water temperatures were 6-25°C in the inner bay, 6-24°C in the in-
termediate neritic water mass and 7 — 20°C in the entrance of the bay. Tempera-
tures were homogeneous in the bay in February-March and September-October.
In spring and summer the temperatures were notably higher in the inner bay than
in the inlet; the contrary was observed in autumn and winter.
Standing stocks of total zooplankton
Replicate tows of zooplankton showed no significant differences (p >0.05) in
total numbers or biomass. We assumed with Youngbluth (1976) and Castel and
Lasserre (1977) that 50% difference of densities are significant when comparing
two samples in this type of environment.
The list of the principal taxa collected, their mean density and biomass at the
three sampling stations and the mean values for all three sites combined are given
in Table I. Copepods constitute the dominant group in density (71.4% of the
average number of individuals) and biomass (91.7% of the averaged biomass).
The proportion of copepods was higher at station C compared with station B and
station A. The mean percentages of individuals, based on monthly sampling were
respectively 87.8 ± 6.0%, 58.5 ± 7.9% and 49.3 ± 8.8%. The mean biomass of
the total zooplankton increased significantly from the coastal neritic waters (14.2
mg dry wt m~ 3 in station A) to the intermediate neritic waters (21.2 mg dry wt

420
Zooplankton in A reaction Bay

o
1,25
0 O st A
st B
dl
Q. st C
Q)
~ 20"

15-

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10-

Jan

20 25 30 35
salinity (%,)

Fig. 2. Monthly temperature and salinity of surface water in the three sampling locations,
1976-1977.

m - 3 in station B) and to the inner neritic waters (90.5 mg dry wt m - 3 in station

C).
Monthly variations of zooplankton in density and biomass in each site are pre-
sented in Figure 3. Kendall's tau coefficient of rank correlation indicates that bio-
mass and total numbers are directly related in the three stations (station A, p =
0.035; station B, p = 0.051; station C, p = 0.014). The maximum of density was
recorded between April and June at the three stations. Considering the biomass,
two peaks were recorded at station A, one in April and one in September. Sta-
tions B and C did not show the autumnal maximum. The largest peak occurred in
April 1976 at station C with 127 000 ind.m" 3 and a corresponding biomass of
525.5 mg dry wt m~ 3 . Minimum numbers of individuals were 23 m~ 3 with a cor-
responding biomass of 0.08 mg dry wt m~ 3 .
In terms of abundance, the standing stock was more variable at stations A and
C than at station B. However, the fluctuations of biomass did not have the same
pattern. It appeared far more stable at station A (variance -s- mean ratio = 9.44)
than at station B (s2 -s- x = 31.17) and station C (s2 -s- x = 226.96).
In station A, the strong increase in density recorded in June-July was due to the
dinoflagellate Noctiluca scintillans (see Figure 5) whose weight (0.19 /tg dry wt)

421
to

Table I.
J. Castel an

a.
Taxa (tg dry wt/ Station A Station B Station C X
individual Number Biomass Number Biomass Number Biomass Number Biomass
p

HOLOPLANKTON:
Cou •ties

Noctiluca scintillans 0.19 ± 0.04 7670 1457 60 11 171 33 2634 500

Cladocera 1.68 ± 0.08 30 50 + + 10 17
Ostracoda 3.89 ± 0.59 + + + + + +
Copepoda 4.45 ± 0.44 2193 9760 3997 17 786 19 866 88 405 8685 38 650
Copepod nauplii 0.80 ± 0.10 + + + + + + + +
Sagitta spp. 6.5S ± 0.81 35 230 14 94 8 54 19 125
Oikopleura dioica 5.30 ± 0.42 291 Downloaded from http://plankt.oxfordjournals.org/ at University of Sussex on October 3, 2012
1543 243 1289 327 1735 287 1521
MEROPLANKTON:
Coelenterata 7.35 ± 2.05 22 162 1 12 5 39 9 69
Polychaete larvae 2.78 ± 0.13 27 75 31 87 97 270 52 144
Cyphonaut larvae 8.62 ± 1.65 50 433 9 74 + + 20 170
Echinoderm pluteus larvae 2.00 ± 0.11 2 4 + + 1 2
Gastropod larvae 1.62 ± 0.40 38 61 172 278 198 321 136 220
Bivalve larvae 2.07 ± 0.32 113 234 30 62 10 21 51 106
Cirripedia nauplii 1.91 ± 0.10 14 27 570 1089 53 101 212 406
Cirripedia cypris larvae 4.93 ± 0.67 4 18 27 134 11 53 14 69
Crab zoea 5.30 ± 0.51 20 104 57 302 1 5 26 138
Mysid larvae 10.00 ± 1.41 3 30 1 10
Fish larvae 18.00 ± 1.00 + + + + + +
TOTAL ZOOPLANKTON 10 509 14 158 5211 21 218 20 751 91067 12 157 42 147

Average based on 12 months sampling of numbers m~ 3 and biomass (jig dry wt m~ 3 ) at the three stations in Arcachon Bay, 1976-1977.
Zooplankton in Areachon Bay

100,000 1,000
station A
10,000 100

1,000 10
....o

100
E
It
m I I I I
£ 100,000 station B ~ 1,000
CD
6
c E
-~ 10,000 100

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U)
C 1,000 10 I
o
..a
100 •(->••'

o
I I I I I I I I
1,000 -
station C -

100

10

j I J I J I J I
F M A M J J A S O N D J
Fig. 3. Fluctuations of the numbers and biomass of zooplankton at stations A, B and C express-
ed as logt0 per m3.

yields a low biomass. This shows that the variability of densities in station A is
not correlated with a variability in biomass.
During cold months (January, February, March) the density and biomass were
higher in the coastal neritic water than in the inner bay where the temperatures
were very low. After the spring homothermie, the increase of standing stock is far
more important in the inner neritic water mass than in the oceanic water. The
mean biomass increased significantly from the coastal neritic waters (14.2 mg dry
wt m ~ 3 in station A) to the intermediate neritic water (21.2 mg dry wt m ~3 in sta-
tion B) and to the inner neritic water (91.1 mg dry wt m~ 3 in station C).

423
J. Castel and C. Courties

Table II.

Species Station A Station B Station C

I II 1 II I II

Calanus helgolandicus 1
Clausocalanus arcuicornis 1 0.4
Oithona plumifera 2
Temora stylifera 3 0.2 2 0.2 1 0.5
Isias clavipes 3 0.1 2 0.7 2 0.1
Pseudocalanus elongatus 5 1.5 2 0.5 1
Centropages typicus 6 0.4 4 0.1 1 0.1
Acartia clausii 8 2.5 2 0.3
Corycaeus anglicus 18 2.8 5 0.5 2 0.6
Oithona helgolandica 18 9.5 6 0.9 2 0.3
Oncaea spp. 21 7.3 8 0.4
Temora longicornis 21 13.5 11 4.6 7 0.9
Euterpina acutifrons 24 15.6 21 3.2 7 0.8

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Paracalanus pan/us 24 42.1 21 8.1 9 0.7
Oithona nana 18 3.7 14 1.3 7 0.4
Acartia discaudata 21 19.0 2 0.1
Parapontella brevicornis 3 0.2 23 26.0 9 3.4
Acartia (Paracartia) grani 1 11 34.3 9 11.6
Acartia bifilosa 2 0.1 24 80.6

Frequency of occurrence (I) and relative abundance in per cent (II) of the copepod species based on
monthly duplicate samples during 1976-1977.

Differential distribution and seasonally of individual taxa

Copepoda. Nineteen species of copepods were identified during this study. Their
frequency of occurrence and relative abundance are indicated in Table II.
Calanus helgolandicus, Clausocalanus arcuicornis and Oithona plumifera were
collected only at station A. These oceanic species (Beaudouin, 1975) are im-
migrants from the Bay of Biscay. Temora stylifera, Isias clavipes, Pseudocalanus
elongatus and Centropages typicus are typical coastal neritic species but were
sporadic and did not really develop in the Arcachon Bay for a long time. Among
the characteristic species of this coastal neritic contingent, Acartia clausii, Cory-
caeus anglicus, Oithona helgolandica and Oncaea sp. are more abundant. A.
clausii and O. helgolandica were at a maximum in spring while C. anglicus and
Oncaea sp. did not show a clear seasonal pattern (Figure 4).
Temora longicornis, Euterpina acutifrons, Paracalanus parvus and Oithona
nana, comprising 74.9% of the copepods collected at station A, were the domi-
nant species of this group. They were also frequent in the intermediate neritic
water mass (Table II). Their possibility of colonizing the bay seems to be more
important (Figure 4).
Three species, Acartia discaudata, Parapontella brevicornis and Acartia
(Paracartia) grani characterized station B (79.3% of the copepod population in
this station). These authochtonous species are rarely found at station A and are
far less frequent and dominant at station C than at station B. Their maximum
abundance occurred in summer (Figure 4). The seasonality was the most marked

424
Zooplankton in Arcachon Bay

station A station B station C

A. clausii

A,
C. anglicus
IX...A/w,1

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A. discaudato

P. brevicornis

A A
A. grani

A
A. bifilosa

F'M'A'M'J'J'A'5'O'N'D'J' 'F'M'A'M'J'J'AVO'NVJ' 'F'M'A M J J A 5 O N D J

Fig. 4. Monthly fluctuations of the most abundant pelagic copepods from the Arcachon Bay,
1976-1977.

for A. grani which was restricted in time from April to September. This latter
species can also colonize the inner neritic water mass during the warmest months.
At station C, Acartia bifilosa was the most common and dominant (80.6%)
species exclusively living in the inner part of the bay. A maximum of abundance
was observed in spring (April-June) and a second maximum in October-Novem-
ber (Figure 4).

425
J. Castel and C. Courties

Other holoplanktonic taxa. Other components displayed a differential distribu-

tion (Table I and Figure 5). Among the holoplanktonic forms, Noctiluca scintil-
lans, Cladocera, Sagitta sp. and Oikopleura dioica clearly came from the coastal
oceanic waters. Their abundance decreased from the inlet to the inner bay. The
peridinien Noctiluca scintillans had a maximum of abundance in June-July when
it comprised up to 56.6% of the total number but only 5.5% of the total biomass.
Among the Cladocera, Podon spp. (mainly P. polyphemoides) and Evadne spp.
(mainly E. normanni and E. spinifera) appeared during the spring season while
Penilia avirostris was characteristic of the summer (not shown in Figure 5).
Chaetognath (Sagitta sp.) comprised 0.15% of the total number and 0.29% of the
total biomass and was the most abundant in summer (Figure 5). The pelagic
tunicate Oikopleura dioica was a non negligible component of the zooplankton
(2.4% in number and 3.6% in biomass).

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station A station B

Sogitta

]...^..O,l

F'MVMVJ'A'S'O'N'D'J' 'F'M'A'M'J'J'A'S'O'N'D'J' 'F'M'A'M'J'J'A'S'O'N'D'J

Fig. 5. Monthly fluctuations of the most abundant zooplanktcrs from the Arcachon Bay,
1976-1977.

426
Zooplankton in Arcachon Bay

Meroplankton. Meroplanktonic larvae were more abundant at station B

(average 897 ind. nr 3 ) than at station A and C (averages 290 ind. nr 3 and 378 ind.
m 3 respectively). Coelenterata, cyphonaut larvae and echinoderm larvae mainly
colonized the coastal neritic waters, but were never abundant (Table I).
Cyphonaut larvae of Bryozoa appeared in October, reaching a maximum in
March and disappeared in late spring. Coelenterate medusae did not have a clear
seasonal pattern. They were not collected in winter from November to January.
Bivalve larvae were found mainly in station A. In this part of the bay, sandy
sediment prevails allowing the development of important bivalve populations.
Bivalve larvae reached a maximum in March-April (Figure 5). Dominant species
were Mytilus edulis, Cerastoderma glaucum, Tellina tenuis. Oyster larvae
(Crassostrea gigas) were rarely collected, probably because of the mesh size of the
plankton net which was too large to retain these small larvae (maximum 300 /tin
when pelagic).
Crab zoea (mainly Carcinus maenas, Portunuspuber, P. arcuatus), collected at

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station A and mostly at station B had a maximum in summer (Figure 5). Cir-
ripedia larvae had a clear maximum of abundance in the intermediate neritic
waters. Barnacles are associated with oyster cultivation in this zone. Cirripedia
nauplii were the most abundant meroplankter (1.8% in density and 1.1% in
biomass) with a maximum in April-May and a peak in September (Figure 5). Two
species were dominant: Chtamalus stellatus and Balanus perforates.
Gastropod larvae were more abundant in the intermediate and inner neritic
waters where adults live among seagrasses {Zostera marina, Z. nana). They reach-
ed maximum abundance in summer but were also abundant in spring (Figure 5).
The most common larvae were: Nassa reticulata, Gibbula spp., Ocinebra sp.,
Hydrobia ulvae).
Polychaet larvae were slightly more abundant in the inner part of the bay where
a few species produce blooms of larvae. The differential distribution and seasonal
cycles of polychaet larvae have been studied by Cazaux (1973). The seasonal vari-
ations were the same in both studies.
Diversity and community structure
Copepoda. The species diversity H' and evenness J indices were computed for
the copepods collected in each station over the sampling period. Average values
for diversity indices were quite similar for station A (H' = 1.51 ± 0.10; J = 0.68
± 0.04) and station B (H' = 1.57 ± 0.07; J = 0.73 ± 0.09). They were signifi-
cantly higher than values computed for station C (H' = 0.58 ± 0.19; J = 0.31 ±
0.09). Averages for all the stations combined were H' = 1.26 ± 0.10 and J =
0.59 ± 0.04. H' diversity has also been computed for polychaet larvae collected
at the same stations during a one-year period (after the work of Cazaux, 1973).
The average value obtained was 1.23 ± 0.10, quite the same value calculated for
copepods. Seasonal variations of diversity were not simple. Low values were rela-
ted to blooms of species (e.g., Paracalanusparvus from August to September at
station A, Acartia grani in August at station B and Acartia bifilosa in spring and
autumn at station C).
Species-abundance relations have been determined for copepods considering

427
J. Castel and C. Courties

the annual relative abundance of the species collected in each station (Table II).
The parameters of the distributions are given in Table III. Whatever the distribu-
tions, the quality of the adjustment is better at station A than at station B and
much better than at station C. The log-linear distribution fits better with the data
at station A, only slightly better at station B, whereas at station C the best adjust-
ment is obtained with the log-normal model. Considering the frequency distribu-
tions of the copepods for all the three stations, the log-normal model is the best
fit (Table III).

Table III.

Stations Rank of Log-normal model Log-linear model

the species m' r m r

1-17 0.866 0.986b 884 0.648 0.994c 465

1-9 3.244 0.987" 280 0.683 0.984" 544

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10-17 3.991 0.968" 8.5 0.681 0.979" 5.5
1-16 1.119 0.985b 1121 0.655 0.987b 403
1-3 27.431 0.999c 0.7 0.743 0.999c 0.7
4-8 3.850 0.984b 19.6 0.572 0.989" 14.9
9-16 6.121 0.983" 5.2 0.737 0.980" 3.9
1-14 0.940 0.963a 3600 0.619 0942n.s. 7790
1-3 0.964 0.992° 130 0.206 O.992c 130
4-10 17.561 0.974a 3.7 0.818 0.971" 3.2
10-14 7.924 0.983" 0.2 0.635 0.982" 0.2
A + B +C 1-19 0.633 0.990° 750 0.635 0.984 b 4260

Parameters of the log-normal and log-linear distributions for the pelagic copepods based on average
over the 12 months sampling period. Goodness-of-fit according to Inagaki (1967): n.s. = not signifi-
cant, (") = approximative, (b) = good, (°) = very good.

However, from Figure 6 it is apparent that the frequency distributions can be

decomposed in two or three segments closed to a log-linear distribution. Each
structure corresponds to an assemblage of species which is not randomly formed.
The parameters of each distribution are given in Table III. At station A, the first
group (species 1 —9) constitutes the dominant species of the coastal neritic con-
tingent: P.parvus, E. acutifrons, T. longicornis, O. helgolandica, Oncaea sp., O.
nana, C. anglicus, A. clausii, and P. elongatus. The other species (species 10-17)
are immigrant species from the Bay of Biscay or from the intermediate neritic
water mass. At station B, one group (species 1 - 3 ) corresponds to the three domi-
nant and authochtonous species: A. grani, P. brevicornis, and A. discaudata.
The species 4—8 are the most abundant species living in the coastal neritic waters
{P. parvus, T. longicomis, E. acutifrons, O. nana, and O. helgolandica). The
others (species 9 — 16) are rare and sporadic species in this site. At station C, A.
bifilosa, A. grani and P. brevicornis constitute the first segment of the distribu-
tion. Species 4 —10 are members of the coastal neritic contingent (T. longicornis,
E. acutifrons, P. parvus, C. anglicus, T. stylifera, O. nana, and O. helgolan-
dica).

428
Zooplankton in Arcachon Bay

- 4n

o
sr A

2-

1-

r i l i i i I i i i i i

1 5 10 15 20
rank i

* * • / sr B

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O •
2-
• •
1- •
1 I 1 T 1 I I

10 15 20
rank i
or

2-

1-

10 15 20
rank i
Fig. 6. Annual species-abundance relations for the pelagic copepods collected in Arcachon Bay,
1976-1977.

Total zooplankton. The distribution laws of specific abundance have been com-
puted for each station averaging monthly estimates for all the species. Calcula-
tions were made both on numbers and biomass (Table IV). The best adjustment is
found for the log-normal model as indicated by the tests for goodness-of-fit. One
exception occurs at station A where the distribution of biomass is slightly better

429
J. Castel and C. Courties

Table IV.

Stations Log-normal model Log-linear model

m' r x2 m r x2

(number) 1.109 0.990° 5031 0.794 0.973" 15 911

(biomass) 1.256 0.990c 0.796 0.992c
(number) 1.199 0.990° 325 0.788 0.978a 289
(biomass) 1.115 0.984" 0.782 0.964a
(number) 0.867 0.981b 9496 0.742 0.960" 29 139
(biomass) 0.847 0.975a 0.750 0.945 n !
(number) 0.980 0.992° 840 0.776 0.981" 5416
Tbiomass) 0.988 0.989"

Parameters of the log-normal and log-linear distributions for the zooplankton based on average over
the 12 months sampling period. Goodness-of-fit according to Inagaki (1967): n.s. = not significant,
(a) = approximative, (") = good, (°) = very good.

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adjusted to a log-linear model. However, the difference is probably not signifi-
cant. For the models applied to the distribution of densities, the best fit is found
for station B and then for station A, the worse fitting being for station C. How-
ever, considering the distribution of biomass, the order of goodness-of-fit
becomes: station A, station B and station C. Most of the difference can be ex-
plained by the abundance of the peridinien Noctiluca scintillans. The community
structure is unbalanced by the dominance of this species. When considering the
biomass, the dominance of this species decreases drastically and the community
appears more equilibrated.
The values of the constants m and m' indicate that the diversity is not strongly
different between the stations A and B, but decreases significantly in the inner
bay (station C). These constants have about the same values when comparing the
distribution of densities and biomass for each station.
Discussion
There is a salinity-temperature gradient in the Arcachon Bay from oceanic con-
ditions at the entrance of the bay to estuarine conditions in the inner part. From
the hydrographic studies of Bouchet (1968) it can be assumed that, whatever the
season, the proportion of the three water masses does not vary significantly, even
in conditions of temperature homothermie (i.e., in spring and autumn). Further-
more, the renewal of the water is very slow. The intermediate neritic waters are
not evacuated seaward but tend to become inner neritic waters. These latter are
eliminated as sheet flow or nappes at the surface.
Similar conclusions can be drawn from the present study. We found a differen-
tial distribution of zooplankton from coastal neritic community to estuarine
species. Each water mass is characterized by a type of zooplankton which is main-
tained seasonally. A more detailed study of the copepods shows that the specific
composition and the community structure is different between the three investiga-
ted sites.
Quantitatively, the biomass increases from the inlet to the inner bay, whereas

430
Zooplanklon in Arcachon Bay

the diversity decreases, especially at station C. As mean densities and biomass in-
creases, fluctuations are more important. Only station A exhibits a spring and
autumnal bloom as many coastal neritic waters in the temperate zone. In the
other sites, the maximum of abundance and biomass occurred between April and
June.
As stated by Lonsdale and Coull (1977) the number of individuals alone cannot
be used as an indicator of a species contribution to the total standing crop. In the
inlet, blooms of Noctiluca scintillans unbalanced the plankton community in
terms of density. But the weight of this species (0.19 jig) is low compared with the
other species and its contribution to total biomass is not so pronounced. How-
ever, this species may have an important role in the pelagic ecosystem since it is a
predator of copepod eggs (Sekiguchi and Kato, 1976).
The community structure of copepods and total zooplankton is measured as a
mathematical description of the relationship of individuals to species. The
MacArthur's broken-stick model has been tested against our data but the latter

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did not fit the distribution which is not, in fact, ecologically relevant. The log-
normal model fits better than the log-linear model when the total populations of
copepods are considered as an average for all the stations over the year and in the
case of complex multispecies assemblages. This seems to be consistent with the
conclusions of Gray (1978) who found that the log-normal is generally a closer fit
to the data in the marine environment. Such a distribution might define a com-
munity in a equilibrium state (MacArthur and Wilson, 1967). However, it has
been shown that a log-normal distribution arises from the interplay of many in-
dependent factors (Whittaker, 1970; May, 1975). The addition of several com-
munities always yields a log-normal distribution (MacNaughton and Wolf, 1970).
In our study, considering the overall distribution of copepod species, it cannot be
assumed that a log-normal distribution might define an homogeneous and stable
community since several assemblages are involved, each of them having generally
a better fit with a log-linear model. This becomes more obvious when considering
complex communities such as total zooplankton which is composed of very dif-
ferent taxonomic groups: crustaceans, mollusks, annelids, tunicates, etc.
In one case the fit to a log-normal might have an ecological sense in this study.
The distribution of the most abundant copepods living at station A (species 1 —9)
fits a log-normal model slightly better than a geometric series (see Table HI). This
community lives in a relatively stable environment and may be considered as
more stable and homogeneous than the other assemblages of copepods living in
the Arcachon Bay. A log-normal distribution may be, in this case, an index of
stability.
As stated by Castel (1980) many factors are involved in determining the species
abundance patterns of marine invertebrates and each figure must be carefully ex-
amined before any ecological conclusion can be made. All the parameters used in-
dicate that the community structure is affected by the salinity-temperature gra-
dient and the variability of hydrological factors, specially in the inner bay.
The diversity, expressed as the slope of species-abundance relations or as a
single index, is not very different between the stations A and B although the
specific composition is not the same. However, the fit to the models is generally

431
J. Castel and C. Courties

less good in station B than in station A indicating a more pronounced heterogen-

eity. In station C, the community structure is clearly unbalanced, the diversity is
low and the log-normal and log-linear distributions are not a good fit to the data.
Comparing diversity of copepods and polychaete larvae, we found that the
values were the same for the two communities (H' = 1.26 and 1.23, respectively)
although the number of species was not the same. Furthermore, in a similar bio-
tope, and for copepods, Lonsdale and Coull (1977) computed an average value
for H' of 1.32 ± 0.32. The copepod diversity of the North Inlet system was very
much like other systems of the United States East coast although the specific
composition was different. It seems that Copepod populations in the Arcachon
Bay reach the same 'standard range' of diversity occurring in estuarine bays. Fur-
thermore, the species-abundance pattern of the total zooplankton community is
nearly the same when comparing the North Inlet and the Arcachon Bay. Using
the average number m' 3 and biomass of each species for the entire year from the
work of Londsale and Coull (1977) we computed m = 0.804, r = 0.988

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(numbers) and m = 0.774, r = 0.980 (biomass) for the log-linear model and m'
= 1.203, r = 0.988 (numbers) and m' = 0.837, r = 0.976 (biomass) for the log-
normal model, which are similar to values obtained for the Arcachon Bay (Table
IV).
In conclusion, the zooplankton of Arcachon Bay is characterized by a rela-
tively diverse authochtonous community in the intermediate neritic water mass
whose structure is comparable to other similar environments. In contrast, the
diversity and community structure of zooplankton is clearly unbalanced in the in-
ner bay or in true estuaries where the river flow is significantly more important.
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