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J. Moll. Stud. (1986), 52, 83-90.

DISTRIBUTION OF MOLLUSCS ACROSS A PNEUMATOPHORE


BOUNDARY IN A SMALL BAY IN NORTHWESTERN AUSTRALIA
FRED E. WELLS
Western Australian Museum, Perth 6000, Australia
(Received 18 June 1985)
ABSTRACT
The number of species, total density and total biomass
of molluscs were examined for 50 m either side of an
Avicennia pneumatophore boundary in the Bay of
Rest, North West Cape, Western Australia (2218'S;
114 08'E). Two mollusc assemblages were found, sep-
arated almost exactly at the pneumatophore bound-
ary. The mudflat association had more species of
molluscs but a lower total density and total biomass
than the assemblage in the Avicennia zone. Stations
among pneumatophores on the seaward fringe of the
Auicennia zone were more diverse and had a greater
density and biomass than stations among the trees.
Possible reasons for the pneumatophore area having
a higher density and biomass of molluscs than the
adjacent mudflat and tree zones are discussed.
INTRODUCTION
Mangroves fringe the shoreline of coastal bays
and estuaries across northern Australia and in
many other tropical and subtropical areas of the
world. There has recently been an increasing
awareness of the role of mangrove trees in pro-
viding a major source of primary production not
only in the forest, but also in adjacent marine
areas. Few animals consume the plants directly;
instead they are broken down into detritus by a
combination of biological and physical factors
to form the basis of a detrital foodweb (Day,
1975; Evink, 1975). Invertebrates in the man-
groves are thought to be an important link in
energy flow to higher order predators such as
birds and fish (Odum and Heald, 1972), but
there are few quantitative data on the diversity,
density and biomass of invertebrates in man-
groves (Milward, 1982; Redfield, 1982).
As one step in documenting the importance
of invertebrates in mangrove systems I recently
(Wells, 1983; 1984) examined the marine invert-
ebrate fauna of a small bay, the Bay of Rest, at
North West Cape, Western Australia (22 18'S;
11408'E). The bay was divided into four
distinct intertidal habitats: seaward mudflat,
landward backflat, and the intermediate Avicen-
nia marina and Rhizophora stylosa tree zones.
There was a steady decline in diversity of species
and total invertebrate density with the mudflat
stations being richest and the Avicennia, Rhi-
zophora and backflat stations being progress-
ively less diverse and having lower invertebrate
densities. Dry, decalcified biomass followed
essentially the same progression except that it
was higher in Avicennia than on the mudflat,
due entirely to the gastropod Terebralia sulcata,
which constituted 55.4% of the biomass of all
marine invertebrates in the Avicennia zone.
Within each habitat stations to be sampled
were selected randomly; transects were not
used. Despite this there was an apparently
greater density and biomass of marine invert-
ebrates among the pneumatophores at the sea-
ward edge of the Avicennia zone than either
among the trees themselves or on the adjacent
mudflat. The present paper examines the num-
ber of species, density and biomass of molluscs
in the transitional zone between the open
mudflat and the centre of the mangrove forest.
Molluscs were selected for investigation because
they were the most important group in terms of
numbers of species and biomass in both the
mudflat and Avicennia stations sampled, and
were also important in terms of density (Wells,
1983; 1984).
MATERIALS AND METHODS
Eight transects 100 m long were established in May-
June 1983 to cover the major areas of the bay. Each
transect was centred on the junction of the mudflat
with the Avicennia pneumatophores, at a midtidal
level of about 1.4 m (Wells, 1980). When the stations
were sampled they were marked with surveyor's tape.
Shore heights relative to the pneumatophore bound-
ary were measured with a metre stick on the sub-
sequent high tide. At each station the following physi-

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84 F.E. WELLS
cal measurements were made: sediment grain size,
pH, salinity and sediment surface temperature. The
same techniques were used as described by Wells
(1983). Salinity measurements were incomplete due
to an equipment malfunction on an early transect.
More comprehensive information on physical con-
ditions in the Bay of Rest is contained in Wells (1980;
1983; 1984).
Molluscs were sampled by placing a 0.75 m diameter
steel ring on the sediment surface, and sieving the
upper 5 cm of sediment through a 2 mm mesh. The
deeper sediment was searched by hand for larger
species. Four samples were made at each station and
pooled. Molluscs were preserved in 10% formalin,
buffered with borax and transferred to 70% alcohol
on return to the laboratory, where specimens were
sorted, identified and counted. Voucher specimens of
each species were deposited in the Western Australian
Museum where they have catalogue numbers WAM
4310-83 to 4351-83. Dry weight determinations were
made as described previously (Wells, 1983).
Overlaps between stations were calculated using
Jaccard's coefficient, c/ a+b- c where a is the number
of species in sample A, b is the number of species in
sample B, and c the number of species in common at
stations A and B (Popham and Ellis, 1971). The
dendrogram was constructed using the weighted pair
group method with average linkage (Sokal and
Sneath, 1963). The primary species of each assem-
blage were determined using the ranking method out-
lined by Sanders (1960). A rarefraction technique
(Simberloff, 1978) was used to determine the number
of species which could be expected to occur in a sample
of a given size from each habitat.
RESULTS
A total of 38 species was collected: 17 occurred
only on the mudflat; 7 only in Avicennia; and
14 species in both habitats (Table 1). The total
density of molluscs in the mudflat stations was
half that in the A vicennia and the biomass was
only 14% as great. The mudflat was not domi-
nated by a single species or genus. Cerithidea
cingulata was the most numerous species, with
36% of the total, followed by Marginella sp.
with 8%. In terms of biomass the most important
molluscs on the mudflat were Saccostrea sp. with
38% of the total and Rhinoclavis vertagus with
19%. In Avicennia, Terebralia sulcata formed
50% of total numbers and 85% of total biomass.
T. palustris was relatively minor with 1% of total
numbers and 4% of total biomass.
In the Avicennia stations the mean number
of total mollusc species was greatest near the
pneumatophore boundary and declined after
10 m shoreward (Table 2). Total mollusc density
and biomass, largely T. sulcata, were greatest at
5 m upshore from the boundary. The Avicennia
habitat can be subdivided into two areas: sta-
tions in the first 10 m upshore from the pne-
umatophore boundary, which are among pne-
umatophores but seaward of the actual tree
trunks, and stations 15 m or more from the
boundary, which are among the trees (Table 2).
Considered in this manner the pneumatophore
stations were more diverse (mean number
of species per station is 3.29 versus 1.81),
had a greater total density of molluscs (115.8 vs
41.9/m
2
) and a greater total biomass (21.6 vs
8.4 g/m-) than stations among the tree zone.
All of these differences are statistically signifi-
cant (t-test, p < 0.05). In addition, all of the
species found in both mudflat and Avicennia
stations occurred in the pneumatophore
stations.
In the mudflat stations the total number of
molluscan species at each level was large, rang-
ing from 10 to 18. The mean number of species
per station, total density and total biomass, were
all greatest or second greatest at the station 1 m
downshore from the pneumatophore boundary.
The mean number of species per station declined
2 m downshore and remained low until the last
two stations. Total density declined to its lowest
level at the 5 m downshore from the pneu-
matophore boundary then increased consist-
ently. Total biomass was lowest at the 5 m down-
shore station then fluctuated with increasing
distance from the pneumatophore boundary.
Analysis of affinity levels between stations
shows two distinct clusters, with an affinity of
only 0.12 between them (Fig. 1). Mangrove
stations form one cluster at an affinity level of
0.36. The downshore pneumatophore station,
one metre from the pneumatophore boundary,
fits into the mudflat cluster. The remaining
Avicennia stations form a second cluster. Within
this grouping the pneumatophore stations (2 m,
5 m, 10 m upshore from the boundary) had the
greatest affinity, 0.43 or more. Stations among
the actual tree zone (15 m upshore) had pro-
gressively lower overlaps with the pneum-
atophore stations. The rarefraction analysis
(Simberloff, 1978) provided additional evidence
that there were two separable mollusc assem-
blages. If the sample size was 250 individuals
from each habitat there would be little dif-
ference in the number of species expected in the
Avicennia stations, with 12.2 species among the
trees and 13.5 species among the
pneumatophores. A combined sampling of 250
individuals in Avicennia would be expected to
yield 14.3 species, about half of the 25.1 which
would be expected in the mudflat. The Sanders
(1960) ranking method showed that the mudflat

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Table 1. Molluscs collected at stations sampled across a pneumatophore boundary in the Bay of Rest in northwestern Australia.
Species
lUlitA(\rtt ntnl
IVI
Total
Indiv-
iduals
2
192
12

39
38

43
33
36
25
24
2
18
14
12

6
6
6
2
5
4

4
3
3

2
1

1
1
1
1
1
1
Density
(no./
m
2
)
0.1
13.6
0.9
2.8
2.7

3.0
2.3
2.6
1.8
1.7
0.1
1.3
1.0
0.9
0.4
0.4
0.4
0.1
0.4
0.3

0.3
0.2
0.2
0.1
0.1
0.1
0.1
0.1
0.1
0.1
0.1
Biomass
(g/m
2
)
>0.0
0.3
>0.0
0.8
0.2

>0.0
0.1
>0.0
>0.0
0.2
>0.0
0.4
>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
Pneumatophore
Total
Indiv-
iduals
388
101
161
59
26
23
26
2
1
13
2

2
6
1
1
1

Density
(no./m
2
)
54.9
14.3
22.8
8.4
3.7
3.3
3.7
0.3
0.1
2.1
0.3

0.3
0.8
0.1
0.1
0.1

0.4
0.1

Zone
Biomass
(g/m
2
)
19.0
0.3
0.8
0.2
1.1
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

Avicennia stations
Total
Indiv-
iduals
169
6

73

2
20

1
5

11
2
1
4
1

Tree Zone
Density
(no./m
2
)
23.9
0.8

10.3

0.3
2.8
0.1
0.7

1.6
0.3
0.1
0.6
0.1

0.3

Biomass
(g/m
2
)
6.7
>0.0

0.3

>0.0
>0.0

>0.0
>0.0

1.3
>0.0
>0.0
0.1
>0.0

>0.0

Total
Indiv-
iduals
557
107
161
132
26
25
46

2
2
18
2

11
4
6
1
1
5
2

3
1
2

Overall
Density
(no./m
2
)
39.4
7.6
11.4
9.3
1.8
1.8
3.3

0.1
0.1
1.3
0.1

0.8
0.3
0.4
0.1
0.1
0.4
0.1

0.2
0.1
0.1

Biomass
(g/m
2
)
12.8
0.2
0.4
0.3
0.6
0.1
>0.0

>0.0

>0.0

>0.0
>0.0

0.6

>0.0
>0.0
>0.0
>0.0
0.1
>0.0

>0.0

>0.0
>0.0

z
o
o
w
2
o
r
( s
c
s
Terebralia sulcata (Born, 1778)
Cerithidea cingulata (Gmelin, 1791)
Clypeomorus sp.
Cerithium coralium Kiener, 1843
Saccostrea sp.
Cominella acutinodosa (Reeve, 1846)
Modiolus auriculatus Krauss, 1848
Marginalia sp.
Nassarius livescens (Philippi, 1849)
Dentallum sp.
Laternula creccina (Reeve, 1860)
Anomalocardia squamosa (Linnaeus, 1758)
Marinula sp.
Rhinoclavis vertagus (Linnaeus, 1767)
Atyid
Circe sulcata Gray, 1838
Terebralia palustris (Linnaeus, 1767)
Polinices conicus (Lamarck, 1822)
Placamen gravescens (Menke, 1843)
Salinatorsp.
Acanthopleura gemmata (Biainville, 1825)
Nerita chamaeleon (Linnaeus, 1758)
Ischnochiton sp.
Septiferbilocularis (Linnaeus, 1758)
Cerithidea largellierti (Philippi, 1836)
Onchidium sp.
Placamen callophylla (Philippi, 1836)
Morula margariticola (Broderip, 1832)
Patelloida mimula (Iredale, 1924)
Katelysia hiantina (Lamarck, 1818)
Thais sp.
Neritina crepidularia (Lesson, 1831)
Trochus hanleyanus (Reeve, 1843)
Vexillum vulpecula jukesii (A. Adams, 1851)
Venerid sp. (juvenile)
Nassarius dorsatus (Roding, 1798)
Area sp. (juvenile)
Isognomon sp. (juvenile)
TOTALS 538 38.3 2.1 817 115.8 21.6 297 41.9 8.4 1114 78.8 15.1

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Table 2. Biological characteristics for all mollusc species and physical parameters of stations sampled across a pneumatophore boundary in the
Bay of Rest in northwestern Australia.
Distance
from
pneumatophore
boundary
(m)
M1
M2
M5
M10
M15
M20
M30
M50
Means
A1
A2
A5
A10
A15
A20
A30
A50
Means
Total
Species
18
12
12
10
12
13
18
17
14.0 3.2
12
8
8
12
6
10
8
3
8.4 3.0
Biological
Mean spp./station
1S.D.
4.50 2.07
2.13 1.55
1.75 2.19
2.25 2.05
2.50 1.41
2.88 1.00
4.25 2.55
4.63 2.92
3.11 1.17
3.38 1.69
3.38 0.74
3.13 1.64
3.25 2.38
1.63 1.51
2.63 1.60
2.00 1.93
1.00 0.93
2.55 0.90
Mean density
(No./m
2
1S.D.)
Meanbiomass
(g/m
2
1S.D.)
Mudflat Stations
8.89 11.13
3.29 3.32
1.75 2.02
4.01 3.06
4.48+ 3.45
4.27+ 4.26
6.65+ 6.66
5.74 5.37
4.89+ 2.19
0.4 0.6
0.3 0.7
0.1 0.1
0.4 0.8
0.1 0.1
0.2 0.2
0.2 0.2
0.5 1.1
0.3 0.2
Avicennia Stations
10.50+11.31
13.02 9.60
22.82 24.24
10.92 11.30
4.62+ 5.72
7.14+ 3.81
6.58+ 8.37
2.52 3.33
9.76+ 6.31
1.9 1.7
2.1 2.1
5.0 3.8
2.4 2.1
1.5 1.8
0.9 0.8
1.3 1.3
0.7 1.3
2.0 + 1.3
Grain size
(01S.D.)
2.68 0.22
2.59 + 0.22
2.78 0.28
2.68 0.26
2.70 0.25
2.69 0.25
2.64 0.29
2.70 0.26
2.68 0.05
2.78 0.23
2.74 0.28
2.85 0.17
2.86 0.12
2.88 0.13
3.00 0.14
2.83 0.21
2.84 0.14
2.85 0.08
Shore
(cm
height
1S.D.)
-3.3 3.3
-6.6
-9.6
-12.1
-13.6
-16.3
-17.4
-24.5
-12.9
+4.0
+9.1
+ 18.6
+30.8
+37.6
+46.0
+ 50.8
+57.9
+31.8
5.4
8.1
9.5
9.8
+ 8.4
7.5
9.7
6.7
2.1
3.4
8.6
11.4
7.6
13.2
10.2
11.8
19.8
pH
(1S.D.)
7.0 0.5
7.0 0.4
7.1 0.4
6.9 0.4
7.0 0.5
7.1 0.5
7.2 0.5
7.4 0.6
7.1 0.1
6.9 0.4
7.0 0.3
7.0 0.3
6.9 0.4
7.1 0.5
7.2 0.5
7.2 0.5
7.1 0.6
7.1 +0.1
Physical
Salinity
(%o)
45
43
42
42
43
42
42
40
42
45
46
45
44
42
44
43
43
44
Temperature
(C 1S.D.)
22.3 3.3
22.6 3.0
23.0 3.1
22.8 3.5
22.5 3.1
23.0 3.5
22.9 3.1
22.8 + 3.7
22.7 0.3
21.6 3.6
21.4 3.6
21.3 4.0
21.2 3.0
21.6 3.6
21.2 3.5
21.0 3.9
21.7 3.7
21.4 0.2
m

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MANGROVE MOLLUSCS
STATION
87
Mudflat Avicennia
7T 0.5-
Fig. 1. Dendrogram of mollusc associations calculated by the weighted pair group method with average
linkage in stations sampled across a pneumatophore boundary in the Bay of Rest in northwestern
Australia.
mollusc assemblage can be characterised by Cer-
ithidea cingulata and Marginella sp., while the
Avicennia grouping can be characterised by Ter-
ebralia sulcala and Cerithium coralium.
Shore height increased consistently across the
mudflat and into the tree zone (Table 2). The
other physical factors fluctuated erratically with-
out demonstrating consistent patterns. Linear
correlations were performed on all possible
combinations of the biological factors of total
species, species per station, density, and bio-
mass using the raw data, not the derived aver-
ages on Table 2, and the physical factors of
sediment grain size, shore height, pH, salinity,
and sediment surface temperature. Significant
relationships (t-test; p < 0.05) were found only
between the number of species and depth and
sediment surface temperature. In both cases the
r
2
value was below 0.09. The same analyses were
performed for all combinations of biological and
physical factors separately in each habitat. For
the mudflat stations significant relationships (t-
test; p < 0.05) were obtained for the number of
species versus depth, distance from the pne-
umatophore boundary interface, and sediment
surface temperature. The greatest r
2
was 0.13
for number of species versus distance from the
pneumatophore boundary. For the Avicennia
stations significant relationships (t-test;
p < 0.05) were obtained for number of species,
density and biomass versus both depth and dis-
tance from the pneumatophore boundary. In
general these r
2
values were higher, but the
greatest was only 0.23 for the number of species
versus distance from the pneumatophore
boundary.
Epifaunal species dominated on the mudflat
in all parameters measured: number of species,
density and biomass (Table 3). Infaunal species
were an important component, with 32% of the
total number of species, 24% of the density and
29% of the biomass. Epifaunal species domi-
nated in Avicennia stations; there were only
two infaunal species which had an insignificant
portion of the total density and biomass. The
mudwhelk genus Terebralia, with two species
had a density of 40.7/m
2
and a biomass of
13.4 g/m
2
. The remaining eighteen epifaunal
species together had a density of 38.1/m
2
and a
biomass of 1.7 g/m
2
. Table 3 also shows that
epifaunal species were dominant in both the tree
zone and the pneumatophore zone; only isolated
infaunal species occurred in either zone.
Arboreal molluscs do occur in low numbers in

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88
F.E. WELLS
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the Bay of Rest (Wells, 1983; 1984), but none
was encountered in the present study.
DISCUSSION
Cantera, Arnaud and Thomassin (1983) ana-
lyzed gastropod species reported from man-
groves in geographical areas throughout the
world, and concluded that the gastropods found
among mangroves are primarily a soft bottom
fauna and only 20% of the species are restricted
to the tree area. Despite the overlap in species
with the mudflat, the mollusc assemblage in
Avicennia in Bay of Rest is substantially dif-
ferent from that which would occur in the
absence of mangroves. While arboreal species
of molluscs were not encountered in the present
study they do occur in low numbers among
Avicennia (Wells, 1984). Teredo are present in
an unknown, but presumably low, density in the
trees themselves. The dense root mat in the
sediment of the Avicennia zone prevents infau-
nal molluscs from living in the area (Wells,
1984). This is a general feature of mangroves
which has been reported by Berry (1963), Mac-
nae (1967), Brown (1971), Sasekumar (1975),
Wells and Slack-Smith (1981) and Wells (1984).
The association of molluscs among Avicennia
was dominated in terms of density and biomass
by two species of Terebralia, which occur only
among mangroves. Of the two species T. sulcata
lives lower down the shore and is particularly
abundant in the seaward pneumatophore zone
(Wells, 1980; present study). The dense popu-
lations of these snails would attract potential
predators, particularly fish. Rays are major
predators of invertebrates in the Bay of Rest
and the holes they make in digging invertebrates
out of the sediment are abundant on the mudflat.
The rays seem to be largely unable to feed
among the pneumatophores, and the population
of T. sulcata is thus protected by the presence
of pneumatophores.
The pneumatophore zone in the Bay of Rest
has been shown to have a greater density and
biomass of molluscs than either the adjacent
mudflat or the actual tree zone. The number of
species of molluscs in the pneumatophore area
is.intermediate between the two adjacent areas.
There is evidence in the literature that this is a
general feature of molluscs in mangroves. Pla-
zait (1975) working in Fiji and Murty and Rao
(1977) working in India both found the density
of molluscs to be greatest near the fringe of the
forest and to decrease progressively into the
trees. Richmond and Ackerman (1975) working

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MANGROVE MOLLUSCS
89
in Fiji found that oysters occurred only at the
seaward fringe of the forest. Unfortunately
these studies provide evidence in favour of the
decline in molluscs among the trees being a
general feature but they did not compare mol-
luscs at the seaward fringe with adjacent
mudflats. The question of whether the pneu-
matophore fringe has a greater density and bio-
mass of invertebrates other than molluscs has
not been answered. Kolehmainen and Hildner
(1975) showed that the greatest biomass of
invertebrates in a Puerto Rican mangrove
occurred on prop-roots of Rhizophora which
hung into the channels and did not touch
bottom; biomass declined in the tree zone.
While the pattern is the same as shown here
for molluscs the Puerto Rican situation can be
considered to be a fouling community, an
entirely different habitat.
The question of why the pneumatophore
fringe has a greater density and biomass of mol-
luscs than the adjacent mudflat or tree zone
must also be considered. Increased feeding time
could help to explain the greater density and
biomass of molluscs among the pneumatophores
than in the trees, but if this was the case density
and biomass on the mudflat should be greater
still, and they are not. If the basic source of
nutrition for marine invertebrates in the Bay of
Rest is the breakdown of mangroves into
detritus as proposed earlier (Wells, 1983), it
could be expected that density and biomass
would be greater at source in the Avicennia
zone than in the mudflat. The molluscs which
dominated in the Avicennia zone are in fact
dependent on the trees for their nutrition. Pla-
zait (1977) has shown that juvenile Terebralia
palustris feed on microflora on the mangrove
surface but adults consume fallen leaves. Yipp
(1980) found vascular plant tissue to be a major
component of gut contents of T. sulcata in a
small mangrove in Hong Kong. Vermeij (1973)
has suggested that the adverse physical
conditions, particularly acidity, make survival
in the tree zone difficult for molluscs with cal-
careous shells. If, as could be expected, the
adverse physical factors in the tree zone are
ameliorated at the seaward margin, an increased
density and biomass of molluscs could be
expected there.
ACKNOWLEDGEMENTS
C. Bryce provided considerable assistance in the
fieldwork, under difficult conditions, under-
taken for this study. Mr and Mrs J. Lefroy,
owners of Exmouth Gulf Station, were generous
in providing access to the Bay of Rest. M. Bez-
ant treated the sediment samples. Dr R. Black
of the University of Western Australia kindly
performed the rarefraction analysis. Dr R.
Humphreys and Ms D. Jones critically read a
draft of the manuscript.
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