Hydrobiologia 481: 137–145, 2002.

© 2002 Kluwer Academic Publishers. Printed in the Netherlands.

137

Spatial distribution patterns of Littoraria species in Hong Kong mangroves

Olive H. K. Lee & Gray A. Williams

Department of Ecology & Biodiversity, The University of Hong Kong, Pokfulam Road, Hong Kong
E-mail: olivelee@graduate.hku.hk

Received 13 September 2001; in revised form 3 April 2002; accepted 6 May 2002

Key words: littorinids, mangroves, spatial distribution, temporal variation, vertical distribution, Hong Kong

Abstract
Variation in the abundance and distribution of two species of mangrove littorinid was investigated using a nested
sampling design at different spatial scales, in the two dominant seasons, in Hong Kong. The abundance of Littoraria
melanostoma, which was less abundant than L. ardouiniana, showed large-scale spatial variation whilst abundance
of L. ardouiniana varied on both large and smaller spatial scales and interacted with seasons, indicating that the
abundance of this species varied between mangroves in summer and winter. Small-scale variation suggested a
patchy distribution of littorinids within a mangrove, whilst the large-scale variation might reflect changes in phys-
ical factors possibly associated with habitat fragmentation. Investigation of the vertical distribution of the littorinids
on the mangrove trees revealed that L. melanostoma were located at a lower overall level than L. ardouiniana. This
zonation pattern may be a result of the morphological differences of the two species, as L. melanostoma has a
thick shell, whilst L. ardouiniana has a thinner, but colour polymorphic shell. Both species of mangrove littorinid
showed patchy distributions at a variety of scales and these patterns highlight the importance of using a hierarchical
sampling approach when investigating spatially fragmented habitats.

Introduction
Mangroves are functionally important systems (Lean
et al., 1990) influencing both marine and terrestrial
ecosystems and offering a diversity of microhabitats
supplying food and refuges for a variety of biota (Por
& Dor, 1984; Lean et al., 1990; Hyde & Lee, 1995;
Tam & Wong, 1997). On a global scale, mangroves
cover about 181 000 km2 of the world’s surface area
(Spalding et al., 1997). Millions of hectares have,
however, been lost due to reclamation, pollution and
many other anthropogenic disturbances (Lean et al.,
1990; Spalding et al., 1997). Due to this rapid frag-
mentation, accurate assessments of the status of the
natural resources in mangroves are difficult to make
and a repeatable method to investigate the distribu-
tion of their associated organisms would benefit such
assessments.
Forty-three distinct mangrove forests have been
identified in Hong Kong, most of which are found in
the relatively undeveloped western and eastern New
Territories (Tam & Wong, 1997). The fauna and flora
of these mangrove forests have been described in a
number of qualitative (e.g. Chang et al., 1985; Sadaba
et al., 1995) and semi-quantitative surveys (e.g. Yipp
et al., 1995; Tam et al., 1997). Most of the general,
nonspecific surveys were, however, conducted in the
early 1990s and are unlikely to represent the current
status of mangroves in Hong Kong since the environ-
ment has been changing rapidly and many mangroves
have been destroyed for reclamation and development
(Yipp et al., 1995; Li & Lee, 1997; Tam & Wong,
1997).
The study of spatial patterns is essential for as-
sessing the status and ecology of a natural com-
munity (Brown et al., 1995; Underwood & Chapman,
1996) and, therefore, its conservation value. Hong
Kong mangroves are becoming increasingly fragmen-
ted, resulting in the need to assess different spatial
scales (within and between mangrove forests) when
investigating populations within these habitats. To ac-
commodate this, nested, hierarchical sampling designs
138
Figure 1. Locations of the six mangrove forests in Hong Kong. CK:
Chek Keng; SPN: Sheung Pak Nai; TC: Tung Chung; TFC: Three
Fathoms Cove; TK: Ting Kok; TT: Tai Tan.
should be applied to study the spatial and/or tem-
poral variation in the abundance of organisms (e.g.
Morrisey et al., 1992a, b; Underwood & Chapman,
1996). Within the mangrove canopy, snails of the fam-
ily Littorinidae are common, being found on the roots,
trunks and leaves (Reid, 1985, 1986, 1992a). The
present survey investigates the vertical and spatial dis-
tribution patterns of the two most common mangrove
littorinids, Littoraria melanostoma (Gray) and L. ar-
douiniana (Heude) on Kandelia candel and Aegiceras
corniculatum in Hong Kong mangroves.
Description of study sites
Surveys were conducted within the mixed Kandelia
candel and Aegiceras corniculatum zone (20–50 m
width) at six representative mangrove forests in Hong
Kong: Chek Keng (CK), Sheung Pak Nai (SPN), Tung
Chung (TC), Three Fathoms Cove (TFC), Ting Kok
(TK) and Tai Tan (TT, Fig. 1). All of the mangrove
forests are relatively large (∼0.7–8.8 ha, Tam & Wong,
1997) and affected by different water sources (CK,
TFC, TK and TT are influenced by oceanic currents,
salinity ranges between 31–35‰ in winter and 17–
26‰ in summer, while SPN and TC are influenced
by freshwater from the Pearl River, with salinity of
26–35‰ in winter and 14–17‰ in summer; Morton
& Morton, 1983; Lee, 2001).
Materials and methods
Surveys were conducted in the winter (February 1999)
and summer (September 1999, see Kaehler & Willi-
ams, 1996 for definitions of Hong Kong summer and
winter). All sites were sampled once within a period of
3–4 weeks in each season and as no replicate sampling
was conducted within each season, this design does
not, therefore, address temporal, small scale (within
season) changes in spatial patterns (see Underwood &
Chapman, 1996). The abundance of littorinids on the
two most abundant tree species, Kandelia candel and
Aegiceras corniculatum, were compared within, and
between, the six mangrove forests in the two seasons
using a nested, hierarchical sampling design.
Littorinid abundance
The Kandelia candel and Aegiceras corniculatum belt
within each mangrove forest was approximately di-
vided, along the coastline, into three equally sized
areas, ranging from ∼0.2 ha (Chek Keng) to ∼3 ha
(Ting Kok) (mean±s.d.=1.3±1.0 ha; Tam et al., 1997).
In each area, three 10×10 m plots (plots 1–3) were
designated using random number tables. The plots ex-
amined were scattered around the mangrove forests,
so no bias might be caused by zonation patterns, if
any, within the mangrove forest. To estimate littorinid
density, trees inside each plot were systematically
searched for 5 min and all littorinids found counted.
A three-factor, mixed model ANOVA, with Season
(Se) as a fixed, orthogonal factor, Mangrove forest
(Ma) and Area (Ar) as random factors, with Mangrove
forest as an orthogonal factor and Area nested within
Mangrove forest (
n=2 seasons×6 mangroves×3
areas×3 counts (plots)=108) was used to investigate
variation in abundance of Littoraria ardouiniana and
L. melanostoma in summer and winter, separately. Ho-
mogeneity of variances was tested using Cochran’s
C test (Underwood, 1997) and where data failed to
meet this assumption, square root transformation was
applied.
Vertical distribution of the littorinids
Five Kandelia candel and five Aegiceras corniculatum
trees were haphazardly selected from the three areas
of the mangrove forest at Sheung Pak Nai, where both
Littoraria species were abundant, in June 2000. The
tree height, canopy height (defined as the height of
the lowest leaf from the ground) and the height of the
high water mark (HWM, determined by observation
of the water mark from the previous high tide period),
of each tree were measured (±1 cm) using a telescop-
ing measuring rod and/or a meter ruler (Fig. 2). The
trees were searched by eye from the bottom to the top
 139
Table 1. Three-factor ANOVA to investigate variation in abundance of Littoraria ardouiniana in areas (Ar) within different mangrove forests
(Ma) in the summer and the winter (Season, Se) (n=3). Significant p-values (p<0.05) are indicated in bold. Variances were homogenous
(Cochran’s C=0.18, p>0.05)

Source of variation SS DF MS F p F versus

Se 200.08 1 200.08 2.51 0.174 Se×Ma

Ma 1140.71 5 228.14 7.56 0.002 Ar (Ma)
Ar (Ma) 362.00 12 30.17 2.61 0.006 RES
Se×Ma 398.19 5 79.63 4.81 0.012 Se×Ar (Ma)
Se×Ar (Ma) 198.89 12 16.57 1.43 0.171 RES
RES 832.67 72 11.56
TOT 3132.55 107

To investigate the variation in the height of the

Figure 2. Illustration of the variables measured in the mangrove
forest.
with the aid of a ladder and all L. ardouiniana and L.
melanostoma were scored according to their position
(Fig. 2), i.e. height above the ground (and therefore
height above the HWM) and substratum they were at-
tached to (trunk=diameter ≥30 mm, branch=diameter
<30 mm, or leaf; Ohgaki, 1992).
The number of littorinids found on the different
substrata, leaf, branch or trunk, the number of the
two littorinid species located on the leaves of the two
mangrove species and variation in the mean number
of each species of littorinid located above the HWM
on the two mangrove tree species were compared with
Chi-square tests (using a 2×2 χ 2 contingency table
in the case of tree×littorinid species test). Differences
in the number of littorinids located on the upper and
lower leaf surfaces in the three areas were investigated
with a t-test.
littorinids above the ground, a two-factor ANOVA
was used with mangrove species (Tree) and littorinid
species (Snail) as fixed, orthogonal factors. Forty-
eight Littoraria ardouiniana and 48 L. melanostoma
on Kandelia candel and Aegiceras corniculatum were
randomly chosen in order to maintain a balanced
experimental design (
n=2 tree species×2 snail
species×24 individuals=96). Variation in the height of
the two littorinid species (Snail) above the high water
mark (HWM) was investigated on different mangrove
species (Tree), from 20 randomly selected L. ardouini-
ana and 20 L. melanostoma located above the HWM,
again on K. candel and A. corniculatum, respectively,
using a two-factor ANOVA (
n=2 tree species×2
snail species×20 individuals=80). The ratios of the
height of the littorinids above ground to tree height
were also calculated; the greater the ratio the higher,
relatively, the littorinid was located on a tree. Twenty-
four L. ardouiniana and 24 L. melanostoma on K.
candel and A. corniculatum, respectively, were ran-
domly chosen. Differences in the ratio (height of the
littorinid/tree height) between the two littorinid spe-
cies (Snail) on the two mangrove species (Tree), was
investigated using a two-factor ANOVA (
n=2 tree
species×2 snail species×24 individuals=96). Trees
without littorinids, or shorter than the height of HWM,
were not sampled.
Results
Abundance of littorinids
Chek Keng (CK) and Tung Chung (TC) had the
highest abundance of Littoraria ardouiniana, whilst
Sheung Pak Nai (SPN), Three Fathoms Cove (TFC)
and Ting Kok (TK) had the lowest densities (Fig. 3).
140

Figure 3. Mean abundance of littorinids from 5-min searches (n=3, +s.e.) in three areas (1–3) of six mangrove forests (CK: Chek Keng; TT:
Tai Tan; SPN: Sheung Pak Nai; TK: Ting Kok; TFC: Three Fathoms Cove; TC: Tung Chung) in winter and summer. Grey bars: Littoraria
ardouiniana; white bars: L. melanostoma.

The number of L. ardouiniana varied from 0 to 24 term, Table 1), suggesting the distribution of this
in a 5-min count, and abundance varied both within species was patchy within mangroves (note differ-
and between the mangrove forests (significant Ar (Ma) ences between mean abundance for the three areas
 141

Figure 4. Mean percentage (+s.e.) of littorinids on different Figure 5. Mean ratio (+s.e.) of the position of the littorinids or
substrata of mangrove trees. Black bars=Littoraria ardouiniana high water mark (HWM) to tree height. LA=Littoraria ardouiniana;
(n=157); white bars=L. melanostoma (n=68). LM=L. melanostoma.

at CK and SPN, Fig. 3). There was a significant Table 2. Two-factor ANOVA to investigate variation in the
height of the two littorinid species (Snail) above the ground on
Season×Mangrove interaction, which indicated that different mangrove species (Tree) (n=24). Significant p-values
the seasonal difference in abundance of L. ardouini- (p<0.05) are indicated in bold. The variances were homogen-
ana varied between mangrove forests. As the factor ous (C=0.40, p>0.05). SNK multiple comparison tests were
Mangrove (Ma) was random it is not appropriate to conducted where appropriate
further investigate this term statistically. Inspection of Source of variation DF SS MS F p
the mean values, however (Fig. 3), suggested that the
abundance of L. ardouiniana was consistent between Tree 1 0.52 0.52 6.47 0.013
the two seasons for most mangrove forests, but dens- Snail 1 0.65 0.65 8.10 0.006
ities were higher in the winter than the summer at Tree×Snail 1 0.05 0.05 0.64 0.427
TC. RES 92 7.37 0.08
TOT 95 8.58
Both Sheung Pak Nai (SPN) and Tung Chung
(TC) had similar, high abundances of Littoraria
SNK tests
melanostoma, whilst abundance in the other four man-
Tree species
grove forests was similar (Fig. 3). The numbers of L.
Kandelia candel>Aegiceras corniculatum
melanostoma varied from 0 to 10 in a 5-min count, and
the density varied between mangrove forests (Three- Snail species
factor ANOVA, DF=5,12, F=36.5, p<0.001). The Littoraria ardouiniana>Littoraria melanostoma
density, however, did not vary within mangrove forests
(Three-factor ANOVA, DF=12,72, F=0.7, p>0.05) as
the distribution of L. melanostoma was similar in areas
within the different mangrove forests (Fig. 3) and on the leaves, more being found on branches (14%)
with season (Three-factor ANOVA, DF=1,5, F=1.9, and on the trunks (18%; Fig. 4). The number of
p>0.05). littorinids located on the three substrata differed signi-
ficantly (χ 2 =8.11, DF=2, p<0.05), more being found
Vertical distribution of littorinids on the leaves of A. corniculatum than K. candel (2×2
contingency table, χ 2 =11.3, DF=1, p<0.05). More
The mean heights (±s.e.) of Kandelia candel and Ae- littorinids were also found on the upper than the lower
giceras corniculatum were 1.83 (±0.29) m and 1.24 leaf surfaces (t=3.80, DF=4, p=0.02).
(±0.13) m, respectively. None of the canopies of K. Littoraria ardouiniana ranged from 0.41 to 2.45
candel extended below the high water mark (HWM), m (mean±s.e=.0.88±0.02 m, n=187) above ground,
whilst 47% of A. corniculatum did. More than 80% whilst L. melanostoma was found between 0.14 and
of Littoraria ardouiniana were located on the leaves 1.34 m (mean±s.e.=0.65±0.03 m, n=93). Littorin-
of the mangroves, whilst ∼8% and 11% were loc- ids were significantly higher above ground on Kan-
ated on the branches and trunks of the mangroves delia candel (mean±s.e.=0.86±0.05 m, n=24) than
(Fig. 4). Fewer (70%) L. melanostoma were found Aegiceras corniculatum (mean±s.e.=0.71±0.03 m,
142
n=24) and L. ardouiniana (mean±s.e.=0.86±0.05 m,
n=24) was, in general, higher than L. melanostoma
(mean±s.e.=0.70±0.04 m, n=24; Table 2).
The number of littorinids above the high water
mark (HWM) varied significantly between the lit-
torinid species and the mangrove species (χ 2 =5.58,
DF=1, p<0.02). The height occupied by littorin-
ids above the HWM was not significantly different
between the two mangrove species (Two-factor AN-
OVA, DF=1,76, F=0.03, p>0.05), but Littoraria ar-
douiniana (mean±s.e.=0.63±0.02 m) was, however,
found significantly higher above the HWM than L.
melanostoma (mean±s.e.=0.39±0.03 m; Two-factor
ANOVA, DF=1,76, F=11.7, p=0.001).
Littoraria ardouiniana was located significantly
higher on the trees (mean ratio±s.e.=0.64±0.01) than
L. melanostoma (0.57±0.02), regardless of the man-
grove tree species (Two-factor ANOVA, DF=1,92,
F=11.5, p=0.001). The mean ratio (±s.e.) of high
water mark (HWM) to tree height was 0.19 (±0.02),
which was significantly smaller than the ratio for the
littorinids, showing that the littorinids were usually
located above the HWM, regardless of the height of
the trees (Fig. 5).
Discussion
Littorinid abundance varied over different spatial
scales. The abundance of Littoraria melanostoma var-
ied over a large scale, i.e. between different man-
grove forests. The distribution of L. ardouiniana var-
ied, however, over a smaller scale, within mangrove
forests. The small-scale variation in abundance in-
dicated that the distribution of littorinids was patchy,
probably behaviourally determined which is known to
cause individuals to aggregate in favourable locations
(Brown et al., 1995; Underwood & Chapman, 1996).
This is most likely mediated through patterns of set-
tlement or mortality as horizontal movement within
the mangrove canopy is probably limited. Littorinids
were more abundant in mangrove forests on the west
coast of Hong Kong, and least abundant within Tolo
Harbour (Three Fathoms Cove, TFC and Ting Kok,
TK), perhaps as a consequence of the extensive pol-
lution and reclamation in Tolo Harbour (Sin & Chau,
1992; Tam et al., 1997). This spatial variation may
be related to physical factors such as salinity (Reid,
1986), as well as tidal magnitude, nutrient levels and
environmental stresses (Cintron & Schaefer-Novelli,
1984). The western coast of Hong Kong experiences
lower salinities than the east, which may affect the
occurrence of more oceanic species, as has been sug-
gested for L. pallescens (Reid, 1992a). Water currents
may also affect the recruitment of these littorinids as
both have a planktonic larval stage (see discussion in
Scheltema, 1971; Johannesson, 1988). Littoraria ar-
douiniana is ovoviviparous, whilst L. melanostoma is
oviparous (Reid, 1989) and the different reproduct-
ive strategies of the two species may contribute to
differences in their abundance.
The composition of the littorinid fauna in Hong
Kong mangroves appears to have changed in recent
years. Littoraria ardouiniana is the most abundant lit-
torinid in Hong Kong mangroves except in Sheung Pak
Nai, where L. melanostoma is more abundant. In con-
trast, L. ardouiniana was rarely recorded in a survey
conducted in the mid 1990s (Tam & Wong, 1997).
Littoraria articulata was previously recorded to be
as abundant as L. ardouiniana and L. melanostoma
(Walthew, 1995), but was seldom found in the present
survey. This difference may be due to changes in the
natural environment, and/or differences in the sur-
vey methods used. Previous surveys, for example,
may also have included areas of cobbles and mudflat
on the fringes of mangroves, where most L. artic-
ulata occur (Reid, 1992a). Since different sampling
techniques may give different survey results, a stand-
ardized technique, for example, counting numbers in
a fixed time period, should therefore be used to record
faunal abundance in 3-D environments.
No temporal variation in littorinid abundance was
found between winter and summer. When the abund-
ance of organisms varies among sites, but is consistent
over time, the major influence can be assumed to be
associated with the sites themselves (Thrush et al.,
1994). Temporal variation in dispersal of the gast-
ropod, Bembicium auratum, in Australian mangrove
forests was also unrelated to season (Crowe, 1999).
The abundance of littorinids might, however, fluctuate
within shorter time periods since aseasonal temporal
variation in dispersal might occur.
Most littorinids occurred on the leaves as opposed
to trunks or branches of the trees and exhibited ver-
tical zonation within the mangrove trees, Littoraria
melanostoma being found lower on the trees than L.
ardouiniana. The location of the littorinids ranged
from nearly ground level to >2 m, although it has
previously been stated that these littorinids were rarely
found >1 m above the ground (McMahon & Brit-
ton, 1985). This distribution appeared to be, as in
some other littorinid species (e.g. Littoraria irrorata;

Bingham, 1972), unrelated to food supply, as these
mangrove littorinids appear to be generalist grazers
ingesting any material on the surface of the substrata
(Lee et al., 2001).
The vertical distribution of the littorinids is likely
to be influenced by physical factors (see Evans, 1965;
Hawkins & Hartnoll, 1983; Henry et al., 1993),
particularly the upper limit which is thought to be
set by desiccation or temperature stress during peri-
ods of emersion (Reid, 1985). Littoraria ardouini-
ana is located higher in the mangrove trees than L.
melanostoma, which may reflect the morphological
differences between these two species, as L. ardouini-
ana is highly colour polymorphic and has a thinner
shell than L. melanostoma (Reid, 1992b; Lee & Dav-
ies, 2000; Lee, 2001). Littoraria ardouiniana might,
therefore, be better suited to live in a higher position
within the mangrove canopy, whilst thicker shells may
protect L. melanostoma from predators lower on the
mangrove trees (Reid, 1986, 1992a, b; see Duncan &
Szelistowski, 1998). The ovoviviparous development
of L. ardouiniana, which permits rapid spawning dur-
ing a brief contact with the high tide (Reid, 1991),
may also be an evolutionary adaptation to its relat-
ively high location in the mangrove canopy. Littorinids
were always located higher on Kandelia candel than
on Aegiceras corniculatum trees. This may simply
be a function of the fact that K. candel was always
taller than A. corniculatum. The ratio of the position
of the littorinids to tree height did not, however, dif-
fer between the two tree species. Since the littorinids
are more abundant on the leaves than on the trunk,
the bushy morphology of A. corniculatum may also
contribute to the different heights occupied on the two
trees. Regardless of tree height, the littorinids were al-
most always located above the high water mark, which
reduces the risk of being dislodged by waves and/or
being attacked by marine predators when inactive.
Unlike in other areas where crabs, conchs and fish
are known to be important predators of many littorinid
species (Warren, 1985; Reid, 1992b), there were no
visually obvious predators, including predatory ar-
boreal crabs, in Hong Kong mangrove forests. Aquatic
predators may have entered the mangrove during high
water, but evidence (crushed shells, etc.) is seldom
seen for this. Since many mangrove flora and fauna
are located near their northern limit in Hong Kong,
the conditions of Hong Kong mangrove forest may
not be typical. The vertical zonation pattern seen may,
however, be due to evolutionary constraints on the
littorinids, as the larvae of these littorinids are plank-
143
totrophic (Reid, 1986, 1999) and so may experience
selective predation pressures in other regions where
predators are present.
Most littorinids were found on the leaves, on which
they feed (Lee, 2001). It has been reported that lit-
torinids mostly attach themselves to the lower sides
of the leaves (Kohlmeyer & Bebout, 1986), probably
to reduce thermal stress (Plaziat, 1984). In contrast,
more littorinids were found on the upper leaf sur-
faces in this study. The percentage of littorinids found
on leaves was higher on Aegiceras corniculatum than
Kandelia candel. This might simply be due to the fact
that A. corniculatum was more bushy, and therefore
had relatively more leaves and a lower canopy level
than K. candel (see Boneka, 1994). Approximately
one-third of Littoraria melanostoma and L. ardouini-
ana were found on the branches and trunks (also see
Reid, 1992a). Littoraria melanostoma in Malaysia, as
in Hong Kong, has been recorded to be confined to
the leaves, stems and aerial roots of mangroves, ran-
ging from ground level to >2 m above ground (Berry
& Chew, 1973), whilst juveniles occurred mainly on
leaves (Reid, 1986).
Nested sampling designs provide a means of as-
sessing the distribution of organisms at different
scales, and quantifying the relative importance of dif-
ferent scales to the overall variability (Morrisey et
al., 1992a; Underwood & Chapman, 1996) and are,
therefore, particularly useful for assessing distribution
patterns of flora and fauna in mangrove ecosystems.
Mangroves are currently threatened habitats and their
destruction and fragmentation increases the distance
between the remaining habitat islands (Yipp et al.,
1995; Li & Lee, 1997). Large-scale reclamation in
Hong Kong has already impacted mangrove com-
munities, and may also affect the dispersal of the
planktonic larvae of the littorinids. Pollutants from the
Pearl River and new towns of the north-west New Ter-
ritories, plus construction in Deep Bay will also affect
the abundance and species diversity of the mangrove
forests. Assessment of the abundance and species
richness of littorinids as well as other mangrove or-
ganisms, using a hierarchical sampling approach is,
therefore, needed when investigating the biodiversity
of mangrove habitats, and conservation value of these
threatened environments.
144
Acknowledgements
We thank Dr Lisandro Benedetti-Cecchi for advice on
experimental design, Dr Benny K. K. Chan, Stephen
Y. L. Tai and Dennis Tong for assisting with the field
work and Ms Kelly Lau and Helen Leung for tech-
nical support. Colleagues of the Hard Rock Ecology
Group and Dr Kevin Hyde, HKU are thanked for com-
ments on the experimental design and the manuscript.
Olive H. K. Lee acknowledges a HKU Postgraduate
studentship.
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