distribution of intertidal flat macrobenthos in buntal bay ... · 50 bako-buntal bay (n...

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Distribution of Intertidal Flat Macrobenthos in Buntal Bay,

Sarawak, Borneo.

Journal: Songklanakarin Journal of Science and Technology

Manuscript ID SJST-2017-0393.R1

Manuscript Type: Original Article

Date Submitted by the Author: 20-Mar-2018

Complete List of Authors: Zakirah, Taufek; Faculty of Resource Science and Technology, Universiti Malaysia Sarawak, 94300, Kota Samarahan, Sarawak, Malaysia., Department of Aquatic Science Shabdin, Mohd; Faculty of Resource Science and Technology, Universiti Malaysia Sarawak, 94300, Kota Samarahan, Sarawak, , Department of Aquatic Science

Fatimah-A'tirah, Mohamad; Faculty of Resource Science and Technology, Universiti Malaysia Sarawak, 94300, Kota Samarahan, Sarawak, , Department of Aquatic Science, A. Rahim, Khairul Adha ; Universiti Malaysia Sarawak, Faculty of Resource Science & Technology

Keyword: Marine Benthos, Benthos Ecology, Intertidal, Macrobenthos

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Songklanakarin Journal of Science and Technology SJST-2017-0393.R1 Zakirah

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Distribution of Intertidal Flat Macrobenthos in Buntal Bay, Sarawak, Borneo.

Original Article

Mohamad Taufek Zakirah1*, Mohd Long. Shabdin

1, Abd. Rahim Khairul-Adha

1,

and Mohamad Fatimah-A’tirah1)

1Department of Aquatic Science, Faculty of Resource Science and Technology, University

Malaysia Sarawak, 94300 Kota Samarahan, Sarawak, Malaysia.

* Corresponding author, Email address: [email protected]

List of authors:

1. Corresponding author: Mohamad Taufek Zakirah. Department of Aquatic Science,

Faculty of Resource Science and Technology, Universiti Malaysia Sarawak, 94300,

Kota Samarahan, Sarawak, Malaysia. Email: [email protected]

2. Mohd Long Shabdin. Department of Aquatic Science, Faculty of Resource Science

and Technology, Universiti Malaysia Sarawak, 94300, Kota Samarahan, Sarawak,

Malaysia. Email: [email protected]

3. Mohamad Fatimah-A’tirah. Department of Aquatic Science, Faculty of Resource

Science and Technology, Universiti Malaysia Sarawak, 94300, Kota Samarahan,

Sarawak, Malaysia. Email: [email protected]

4. Abd. Rahim Khairul-Adha. Department of Aquatic Science, Faculty of Resource

Science and Technology, University Malaysia Sarawak, 94300, Kota Samarahan,

Sarawak, Malaysia. Email: [email protected]

Acknowledgment

I wish to thank Faculty of Resource Science and Technology, Universiti Malaysia

Sarawak for providing the research facilities and Ministry of Higher Education for

financial support through Fundamental Research Grants Scheme no. FRGS/STWN 04

(01)/1062/2013(08).

Running head title: Macrobenthos of Buntal Bay

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Abstract 1

Macrobenthos distributions in intertidal of Buntal Bay, Sarawak was studied 2

based on systematic sampling conducted in 2014. The aims of this study to determine 3

the intertidal macrobenthos horizontal distribution and their relation with environmental 4

parameters. The analyses of intertidal flat marobenthos community suggested 5

Polychaete dominated the community in terms of the number of individuals and species 6

and followed by crustaceans and molluscs. Polychaetes of families Nephtyidae, 7

Spionidae, Capitellidae and Magelonidae contributed to high densities of macrobenthos 8

found. Multivariate analysis performed by BIO-ENV indicated that communities in 9

Transect 1 (T1) and Transect 2 (T2) were best correlated with food availability 10

(sediment chlorophyll a), and heterogeneity of sediment type (percentage of fine sand 11

and very fine sand). Heterogeneity of sediment characteristic and food availability has 12

been identified as potentially playing a key role in shaping of intertidal macrobenthos 13

distribution in Buntal Bay. 14

Keywords: macrobenthos, intertidal flat, Buntal Bay, horizontal distribution, Sarawak, 15

Borneo 16

1. Introduction 17

Intertidal macrobenthos consists of a highly diverse group, comprising mainly 18

polychaetes, crustaceans and molluscs (Peterson and Peterson, 1979; Whitlatch, 1982; 19

Nakao, Nomura, & Abd. Satar, 1989; Netto & Lana, 1997; Lastra et al., 2006; Morais, 20

Comargo, & Lana, 2016), as well as three lesser groups, namely echinoderms, 21

nemerteans and sipunculids (Whitlatch, 1982; Morais et al., 2016). Early studies of 22

intertidal macrobenthos were concerned mainly with macrobenthos zonation, 23

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classifying low, mid and high intertidal zone on the basis of dominance species (Rodil, 24

Lastra, & Sánchez-Mata, 2006; Blanchet et al., 2014). Recently, enormous progress has 25

been made towards comprehension of macrobenthos communities and ecosystem 26

functioning in many parts of the world (eg: Magni, Como, Montani, & Tsutsumi, 2006; 27

Shin, Lam, Wu, Qian, & Cheung, 2008; Gerwhoing, Drolet, Hamilton, & Barbeau, 28

2016). 29

Previous studies have reported the environmental factors coupled with variations 30

in tolerance of the macrobenthos organisms influenced on macrobenthos communities 31

(Peterson & Peterson, 1979; Lu, 2005; Magni et al., 2006). Community structure 32

embodies all the various ways of individual members of communities relate and interact 33

with one another (eg: spatial and temporal abundance of macrobenthos) and how the 34

community level properties arising from these relations with environment and 35

biological factors (Giller, 1984; Tokeshi, 1993). Alteration in the environmental 36

characteristic of the habitat can strongly affect in composition and abundance of species 37

among sites, thus influencing the species diversity (Seiderer & Newell, 1999; Faraz et 38

al., 2016). However, the macrobenthos inhabiting tropically and particularly in Sarawak 39

has been poorly studied. Buntal Bay is one of few intertidal flats in Sarawak and served 40

as an important fishery area of economic species of Razor clam Solen spp. (Rahim, 41

2011). To date, macrobenthic communities in this intertidal flat of Buntal Bay have 42

never been described. Therefore, the present study aims to determine the intertidal flat 43

macrobenthos abundance in Buntal Bay with particular emphasis on the context of 44

community structure and the relation with environmental factors. Finding of this study 45

provides useful baseline data for future ecological and systematic study of 46

macrobenthos in this area. 47

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2. Materials and Methods 48

2.1 Study site 49

Bako-Buntal Bay (N 1°41’52.03’’, E 110°22’28.10’’) is a semi-circular bay 50

bordered by Gunung Santubong to the west and Bako National Park to the east (Figure 51

1A). Mangrove forest stretches between the two promontories. During neap tides, 52

almost a third of the Bay is exposed sand-mudflats. The Bay is globally importance as a 53

migration site for waterbirds (Howes, 1986; Mizutani et al., 2006). 54

2.2 Field sampling 55

2.2.1 Macrobenthos sampling 56

Macrobenthos sampling was conducted during low tide in May 2014. The 57

approach taken was by performing line transects methods. Two transects were 58

performed perpendicular to shoreline starting from low water mark to high water mark 59

(Figure 1B). The distance between transect was 1.5 km. A total of 21 sampling stations 60

were established on these two transects. A distance between each station was 150 m. At 61

each stations three quadrate 0.25 m2 (= three replicates) was placed at 5 meter interval 62

on the right and left hand side of the transect. Sediment in the quadrate were scooped 63

with spade approximately 15 cm deep based on preliminary sampling conducted on 64

vertical distribution. In field, all sediment samples were sieved through a 500 µm mesh 65

sieve and fixed in 5 % buffered formalin before further analysis in laboratory. 66

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2.2.2 Environmental parameters 67

Water parameters for interstitial water such as salinity, temperature, dissolved 68

oxygen and pH were measured in situ. Interstitial water was obtained by using a 69

modified device followed Giere, Eleftheriou and Munson (1988). Three replicates of 70

sediment sample (15 cm) were collected using pespex core at each station within the 71

macrobenthos sampling quadrates for the determination of grain size distribution and 72

total organic matter (TOM). Two replicates of 1 cm of surface sediment was also taken 73

within the quadrates using pespex corer and placed in a plastic bag for determination of 74

chlorophyll a. 75

2.3 Laboratory analysis 76

2.3.1 Macrobenthos study 77

The first step of macrobenthos extracted was carried out after formalin was 78

removed and transferred to 70 % ethanol before sorting process. Fine sorting was 79

carried out in order to separate organisms belonging to different high taxa under the 80

stereomicroscope. For a detail taxonomic identification of macrobenthos specimens, the 81

use of compound microscope was needed. Species were identified to the lowest 82

practical taxon by referring identification keys such as Day (1967) for Polychaeta, 83

Brinkhurst (1982) for Oligochaeta, Valentich-Scott (2003) for Mollusca, Gibson and 84

Knight-Jones (1994) for Nemertinea, Cornelius, Manuel, and Ryland (1994) for 85

Cnidaria, Abele, and Kim (1986) for Decapoda, Barnard and Karaman (1991) for 86

Amphipoda. 87

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2.3.2 Sediment analysis 88

The method used for determination of grain size was based on standard method 89

by Bale and Kenny (2005). The sediment grain size analysis determined by using dry 90

and wet sieving technique in order to determine the fraction mixture of gravel, sand, silt 91

and clay. A simple estimate of the organic contents can be derived from the mass of loss 92

of ignition (LOI). This method involved drying the samples at low temperature (40 ºC) 93

for 24 hours, then combusting the organic content at high temperature (450 ºC) for 4 94

hours (Greiser & Faubel, 1988). The loss of weight indicated the amount of total 95

organic matter (TOM) in the samples. 96

The chlorophyll a in sediments was done by followed method by Lorenzen 97

(1967). The chlorophyll a in sediments was done by grinding the sediment inside the 98

mortar with 90% of acetone. Then, 10 ml of aliquot was transferred into centrifuge tube 99

and left over a night before centrifuge for 30 minutes at 4000 rpm. The supernatants 100

were then transferred into cuvette and measured in spectrophotometer (HACH, 101

DR2800) before and after acidification. One drop of 0.2 hydrochloric acid (HCl) was 102

added to 1.5 ml of extract volume and absorbance at 665 nm were made. Turbidity 103

blank was measured at 750 nm. 104

2.4 Data analysis 105

Determination of quantitative macrobenthos composition and density was based 106

on number of macrobenthos individuals per 0.25 m². One-way analysis of variance 107

(ANOVA) was used to test difference between the environmental variables between the 108

stations. The statistical significance of differences among sites was assessed using 109

analysis of similarities (ANOSIM), a nonmetric method based on randomization of 110

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rank-similarities among all samples and multiple pair-wise comparisons (Clarke, 1993). 111

A significance level of ρ<0.05 was used in all tests. The number of species (S) in each 112

sample was used as direct measure of species richness index (d). The Shannon-Wiener 113

diversity index (H’) is widely used as an absolute measure of diversity and species 114

equitability was determined by Pielou’s evenness index (J’). Cluster analysis were 115

carried out to delineate the macrobenthic communities of the sampling station into 116

different groups using a Bray-Curtis similarity measure based on presence/absence 117

transformed data and group-average linkage. The relevance of the station groups 118

obtained was evaluated by the similarity profile routine (SIMPROF) tests (Clarke & 119

Gorley, 2006). Differences in the composition of the macrobenthos assemblages among 120

stations were verified through non-metric multidimentional scaling (nMDS). 121

Subsequently, the contribution of species in each group similarity was assessed using 122

the SIMPER (similarity percentages) procedure (Clarke & Gorley, 2006). 123

Macrobenthos assemblages were characterized using univariate and multivariate 124

measures by using PRIMER v6 for determination of community structure (Clarke & 125

Gorley, 2006). Biotic and Environmental linking (BIO-ENV) and Spearman’s rank 126

coefficient analysis was performed to test which environmental variables were 127

correlated with the macrobenthic community. 128

3. Results 129

3.1 Environmental parameters 130

ANOVA analysis showed that physico-chemical parameters of water in both 131

Transects were significantly different (p=0.0001). Generally, the water temperature in 132

Transect 1 and Transect 2 ranged between 33.30 ºC to 36.10 ºC and 29.03 ºC to 33.57 133

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ºC respectively. Salinity tended to be much higher in Transect 1 than in Transect 2 with 134

salinity ranged between 30.3 to 33.3 PSU and 20.33 to 24.73 PSU respectively. 135

Dissolved oxygen (DO) concentration in Transect 1 ranged from 0.14 to 2.76 mg/ℓ and 136

between 0.17 to 0.47 mg/ℓ recorded in Transect 2. Transect 1 recorded pH value 137

ranging between 7.47 to 7.93 and for Transect 2 pH ranged between 6.6 to 7.73. 138

Summary of sediment characteristics were presented in Table 1. The sediment 139

grain size in both transects consisted of medium sand and moderately sorted sands. 140

ANOVA analysis showed that total organic matter (TOM) were significantly difference 141

among the stations for both Transect 1 (P=0.0001) and Transect 2 (P=0.0001). Total 142

chlorophyll a (Chl a) concentrations ranged from 2.98 to 53.49 mg/m3 in Transect 1 and 143

1.73 to 83.05 mg/m3 in Transect 2 (Table 1). ANOVA analysis showed that chlorophyll 144

a concentrations were significantly difference among the stations in Transect 1 (ρ<0.05) 145

and Transect 2 (ρ<0.05). 146

3.2 Species composition and density 147

A total of 97 macrobenthos species were identified in the intertidal of Buntal 148

Bay, which mainly composed of Polychaeta, Crustacea, Mollusca and Nemertinea. 149

Majority of macrobenthos species was contributed by Polychaete worms (Annelida). 150

Other groups with low species number were Oligochaeta and Echinodermata. 151

In total, mean density of macrobenthos collected in both transect was 3461.67 152

ind.m-². Mean density of total macrobenthos varied between 63.33 to 397.67 ind.m

-² in 153

Transect 1 and 26.67 to 450.67 ind.m-² in Transect 2. In Transect 1, the highest density 154

was recorded at ST8 (397.67 ind.m-²) and followed by ST6 (296.67 ind.m

-²). The lowest 155

density was recorded at ST11 (63.33 ind.m-²). High density in ST8 and ST6 corresponds 156

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to occurrence of high density of polychaete Prionospio sp. 1 and Nephtys 157

sphaerocirrata. Four species of polychaetes recorded greater density value, namely 158

Nephtys sphaerocirrata (217.3 ind.m-²), Prionospio sp. 1 (186.7 ind.m

-²), Nephtys sp. 1 159

(109.3 ind.m-²) and Spiophanes sp. 1 (80.0 ind.m

-²). Whereas for two species of mollusc 160

Umbonium elegans (150.7 ind.m-²) and Tellina sp. 1 (126.7 ind.m

-²) were also 161

contributed to high density of macrobenthos found. 162

In Transect 2, the highest macrobenthos density was recorded at ST7 (450.67 163

ind.m-²), followed by ST8 (302.67 ind.m-²). The lowest density was recorded at ST4 164

(26.67 ind.m-²). The highest density in ST7 contributes by the occurrence of high 165

density of Magelona sp. (61.33 ind.m-²) and Tellina sp. 1 (66.67 ind.m

-²). Similar to 166

Transect 1, polychaetes species were recorded dominance in this transect and followed 167

by mollusc. Three species of polychaetes recorded highest density value which were 168

Nephtys sphaerocirrata (122.7 ind.m-²), Barantolla sp. (161.33 ind.m

-²) and Magelona 169

sp. 1 (146.7 ind.m-²). The occurrence of Tellina sp.1 (137.33 ind.m

-²) also contributed to 170

high density of macrobenthos found. 171

3.3 Species number, richness, diversity and evenness 172

In general, the total number of species was found higher in Transect 1 than 173

Transect 2 (Figure 2A and Figure 2B). With regard to the ecological indices between 174

tide marks, number of species, (S) and species richness index (d) showed similar 175

patterns in T1 (Figure 2C) and T2 (Figure 2D). In both transects the number of species 176

(S) and species richness index (d) was significantly different between stations (p<0.05). 177

In Transect 1, the highest number of species (S) and species richness index (d) was 178

recorded at ST8 (S= 23.67 ± 5.51; d= 3.78 ± 0.87) and the lowest number of species (S) 179

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and species richness index (d) was recorded at ST2 (S=4.33 ± 2.3; d= 0.74± 0.45). In 180

Transect 2, the number of species (S) and species richness index value (d) recorded was 181

slightly lower compared to Transect 1. The highest value was recorded in ST7 182

(S=20±6.56; d=3.10±0.86) and the lowest value was recorded in ST9 (S=3.67±1.15; 183

d=0.51±0.03). 184

In both transects, the species diversity index (H’) and species evenness index (J’) were 185

fluctuated in both transect from low tide (LTL) to high-tide level (HTL). In Transect 1, 186

the highest species diversity was observed at mid-tide level (MTL) stations 2.79 ± 0.21 187

(ST8) (Figure 2E and Figure 2F). High species diversity in these stations corresponded 188

to greater total number of species recorded. The lowest H’ value was recorded in ST2 189

(LTL) with value 1.21 ± 0.51. For other stations, the H’ value ranged from 1.15 to 2.29. 190

In Transect 2, the highest species diversity was observed at ST7 (MTL- 2.58 ± 0.34). 191

The lowest H’ value was recorded in ST9 (LTL) with value 1.03 ± 0.06. 192

High evenness index (J’) indicated that macrobenthos species were evenly distributed 193

among the stations (on a scale of 0-1). The highest J’ in Transect 1 was observed at ST1 194

(0.91 ± 0.1) and lowest at ST11 and ST12 with values 0.83 ± 0.09 and 0.83 ± 0.032 195

respectively (Figure 3.9). Values for other stations were ranged from 0.86 to 0.90. In 196

T2, the values recorded was highest at ST4 (0.94 ± 0.037) and lowest at ST1 (0.75 ± 197

0.17). Evenness index value for other stations was ranging from 0.83 to 0.93 (Figure 2E 198

and Figure 2F). 199

3.4 Community structure 200

Results of one-way ANOSIM test indicated significant distribution of 201

macrobenthos community between the stations in Transect 1 (r = 0.558, p = 0.1 %). 202

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Cluster analysis and SIMPROF test revealed that the macrobenthos assemblages in 203

Transect 1 (T1) significantly differed between stations (p<0.005) with a Bray-Curtis 204

similarity of 17 % (p = 0.001) and 33 % (p = 0.02). Based on 33 % similarities, MDS 205

ordination (stress: 0.11) and cluster analysis suggested that T1 consisted of two groups 206

(Figure 3). Group 1 comprise of ST2 and ST11. Group 2 had the highest number of 207

stations and species, representing middle tide zone and low tide zone (excluding ST1), 208

and relatively homogenous with respect to species composition. In T1, species 209

homogeneity was greatest in ST2 with average similarities 68.28 %, followed by ST1 210

(61.92 %), ST6 (53.27 %) and least in ST5 (15.53 %) (Table 2). According to the 211

SIMPER analysis, this shift was mainly caused by a difference in densities of Tellina 212

sp. 1, Barantolla sp., Prionospio sp. 1 and Nephtys sp. 1. 213

One-way ANOSIM test in Transect 2 (T2) indicated that significant distributions 214

of macrobenthos community was occurred (r = 0.859, p = 0.1 %). Based on cluster 215

analysis and SIMPROF, the macrobenthos distribution in T2 significantly differed 216

between stations (p<0.005) with a Bray-Curtis similarity of 11 % (p = 0.001) and 24 % 217

(p = 0.003). Based on 24 % similarities, MDS (stress: 0.05) and cluster analysis 218

suggested that the macrobenthos communities in T2 were distinguished by three groups 219

(Figure 4). First group was separated from others which represented by macrobenthos 220

communities from ST1. Second group was contributed by macrobenthos communities 221

from ST4 and ST9. The third group consists of the combination of ST2, ST3, ST5, ST6, 222

ST7 and ST8 communities. According to the SIMPER analysis, species similarities was 223

recorded highest in ST4 (79.48 %), ST9 (74.57 %) and ST1 (71.02 %) and least in ST6 224

(34.42 %). Difference in densities of Tellina sp. 1, Magelona sp., Glycera sp. 1 and 225

Barantolla sp. accounted for distribution differences in this transect (Table 3). 226

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3.5 Macrobenthos relationship with environmental parameters 227

To determine whether the environmental parameters might influence the 228

variability in the macrobenthos communities distribution in both transects, multivariate 229

analysis BEST (BIO-ENV) and Spearman’s rank correlations were carried out. BIO-230

ENV analysis showed that the correlations between environmental variables and 231

macrobenthos density were modest in T1 and T2 as presented in Table 4. Results of 232

BIO-ENV reveal that macrobenthos density were all significant at T1 (rho = 0.45, p = 233

0.26) and T2 (rho = 0.487, p = 0.37). When all environmental parameters are included 234

in the analysis, the macrobenthos communities in T1 were best correlated with 235

dissolved oxygen, pH, salinity, sediment chlorophyll a, and heterogeneity of sediment 236

type (percentage of fine sand and very fine sand). To gain an insight on how the 237

mention environmental parameters effects on structural of intertidal macrobenthos 238

communities, Spearman’s rank correlation was performed (r) between community 239

indices (species density, species diversity (H’) and species evenness (J’)) and 240

environmental parameters (Table 5). In T1, the strongest Spearman’s rank correlation 241

showed statistically significant correlation (p<0.05) between species evenness and 242

dissolved oxygen (DO). Based on BIO-ENV analysis, the macrobenthos communities in 243

T2 were best correlated with pH, chlorophyll a, salinity and heterogeneity of sediment 244

type (percentage of fine sand and very fine sand). In T2, the strongest Spearman’s rank 245

correlation revealed a statistically significant correlation (p<0.05) between species 246

evenness and chlorophyll a (Table 6). Species diversity showed significant correlation 247

between sediment type (fine sand) and salinity. Species density showed strong 248

correlation between salinity and temperature however no significant level was detected). 249

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4. Discussion 250

In this study, up to 97 species of macrobenthos was collected. The macrobenthos 251

species identified in Buntal Beach was mostly contributed by Polychaeta, Mollusca, 252

Crustacea and Nemertinea. Among the macrobenthos, the most taxonomically diverse 253

group was the polychaete which accounted 56 % to 62 % of total macrobenthos 254

collected in TI and T2, respectively which is previously reported as important taxa 255

inhabiting intertidal habitat in terms of species number and species density (Ditmann, 256

2002; Lastra et al., 2006; Morais et al., 2016). 257

Finding of the present study showed that high number of species collected 258

compared to other regions of the same habitat such as in Guaratuba Bay, Brazil with 75 259

species (Morais et al., 2016), Paranagua Bay, Brazil with 52 species (Netto & Lana, 260

1997), Gulf of Mexico with 21 species (Coblentz, Henkel, Sigel, & Taylor, 2015), 261

North of Portugal with 22 species (Veiga, Rubal, Cacabelos, Maldonado, & Sousa-262

Pinto, 2014), North of Spain with 31 species (Lastra et al., 2006), Wenzhou Bay, China 263

with 38 species (Bao-Ming, Yi-Xin, & Hong-Yi, 2011) and Karnafuly, India with 33 264

species (Islam et al., 2013). In the regional scale, macrobenthos communities studies in 265

an intertidal of Buntal Bay have not been in detail before. The number of macrobenthos 266

species collected in Buntal Bay showed relatively higher compared to other study of 267

similar habitat such as in Teluk Aling, Pulau Pinang, Malaysia with 46 species (Ahmad, 268

Fang, & Yahya, 2011); Kuala Selangor, Malaysia with 44 species (Nakao et al., 1989) 269

and Barangay Tagpangahoy, Philippines with 39 species (Medrano, 2015). The 270

macrobenthos species obtained at intertidal of Buntal Bay resembled that of 271

communities in other intertidal habitat reported in Malaysia coastal waters (Broom, 272

1982; Nakao et al., 1989; Ahmad et al., 2011). 273

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In this study, macrobenthos community pattern were fluctuated in both transects 274

and did not followed or varied with tidal gradients. Occurrence of some dominant 275

species was apparent at both transects. This can be explained by the high dominance of 276

Barantolla sp., Glycera sp., Nephtys sphaerocirrata and Carinoma sp. The dominance 277

of one or a few taxa in intertidal habitats has been observed in similar studies and the 278

variability of species composition (Mclachlan & Jaramillo, 1995; Coblentz et al., 2015). 279

Giller (1984) suggested that communities can differ in species diversity due to the 280

variability of available food resource, niche width of species composition, and lastly the 281

degree of niche overlap. 282

The environmental data and macrobenthos data make it possible for us to test the 283

general hypothesis that environmental factors influence the macrobenthos community 284

structure in an intertidal of Buntal Bay. Dissolved oxygen (DO) in interstitial water was 285

surprisingly positively correlated to species evenness in T1. The lowest value of 286

species evenness and species density were recorded at ST11 and ST12 (T1) with low 287

dissolved oxygen concentration (0.1 to 1.05 mg/ℓ). Bottom sediments are the final sink 288

for many anthropogenic contaminants and they can accumulate great amounts of 289

organic matter affecting the oxygen content of the bottom water (Whitltach, 1982). 290

Hypoxia/anoxia causes reduced macrofauna abundance in turn reduces the amount of 291

bioturbation activities (Rosenberg, Hellman, & Johansson,1991; Gray, Wu, & Or, 292

2002). This study observed that interstitial water salinity has showed significant 293

correlation with species diversity in T2. Lower salinity means influence by freshwater 294

input. Freshwater input not only lower the salinity but also nutrient enrichment, 295

resulting in a macrobenthic community different from that under high salinity 296

conditions (Nishijima et al., 2013). Thus, freshwater input from adjacent river or rainfall 297

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is suggested may be influence the salinity and affect the macrobenthos communities in 298

T2. 299

Sediment characteristics were among the major factors that best explained the 300

pattern of macrobenthos in T2. Results from correlation analyses in T2 showed that 301

sediment properties (sediment mean (ø), coarse sand and fine sand) were significantly 302

correlated with species diversity (H’). The most reported feature associated with 303

macrobenthos community of particular species or assemblages is sediment type 304

(Peterson & Peterson, 1979; Coblentz et al., 2015). In T2, sediment distribution 305

exhibited variability among the stations. Based on statistical values (mean, sorting and 306

skewness), most of sediment distribution in Transect 1 and Transect 2 consist of coarse, 307

medium, fine and very fine sand with more prevalence of medium sand. Gooday et al. 308

(2010) stated that, sediment habitat heterogeneity can be generated by hydrodynamics 309

features such as bottom currents and biological effects such as bioturbation activity. In 310

turn, creates variability of species distribution, species composition, and species 311

diversity. Linear regression showed that the species diversity value increase with 312

decreasing of sediment mean (phi) value (towards coarser sand). In agreement with 313

Lastra et al. (2006), they reported that grain size significantly affects the community 314

characteristic which coarse sand sediment result in low number of species and species 315

density than finer sand. Suggesting that species diversity of intertidal macrobenthos is 316

affected by sediment properties in this transect. 317

The variability of benthic chlorophyll a in an intertidal sediments has been 318

investigated by several authors at various spatial and temporal scales (Magni & 319

Montani, 2006; Sin, Ryu, & Song, 2009). Sediment microalgae (chlorophyll a) play an 320

important role both as food for benthic organisms and source of nutrients for the 321

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overlying water column after decomposition (Fabiano & Danovaro, 1994; Magni, Abe, 322

& Montani, 2000; Josefson, Forbes, & Rosenberg, 2002). Biomass of sediment 323

chlorophyll a ranged from 1.73 to 83.05 mg.m3 in this transect and was relatively high 324

compared with other intertidal system such as Kwangyang Bay, Korea (4.4 to 81.2 325

mg.m3) and Seto Inland Sea, Japan (4 to 25 µmol.g

1). The comparison between 326

chlorophyll a concentrations in surface sediment in this study further provides general 327

information for evaluating the extent of environmental variability at ebb tide of tropical 328

intertidal flat of Malaysia particularly. Cook, Butler, & Eyre (2004) reported that low 329

primary productions were observed at coarse sand sediments (high water energy) than 330

fine grain sand. In contrast, Sin et al., (2009) reported that grain size was not major 331

factor controlling the biomass of benthic microalgae (chlorophyll a). In this study, sand 332

percentage were composed of medium sand, fine sand and very fine sand but 333

concentration chlorphlyll a value were varied. However, further studies including 334

mesocosm experiments are required for better understanding the direct effects of 335

sediment chlorophyll a on macrobenthos community in Buntal Bay. 336

5.0 Conclusions 337

Present study showed that sediment granulometry and interstitial water salinity 338

was a significant explanatory factor in the structuring macrobenthos community in T2. 339

However, in T1 the best correlation was observed influence the community structure 340

was dissolved oxygen. Finding of present study suggests that the environmental 341

variables related to macrobenthos examined in small spatial scale resulting in difference 342

community structure and environmental factor regulating such community pattern. 343

344

References 345

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Ahmad, O., Fang, T.P., & Yahya, K. (2011). Distribution of intertidal organisms in the 346

shores of Teluk Aling, Pulau Pinang, Malaysia. Publications of the Seto Marine 347

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of laser diffraction particle size analyzers for inference on infauna-sediment 366

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intertidal mudflats of a temperate Australian estuary. I. Benhtic metabolism. 369

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Fabiano, M., & Danovaro, R. (1994). Compositon of organic matter in sediments facing 373

a river estuary (Tyrrhenian Sea): relationship with bacteria and microphytobenthic 374

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the coastal environment. Marine Ecology Progress Series, 238, 249-279. 384

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Giere, O., Eleftheriou, A., & Munson, D.J. (1988). Abiotic factors. In Higgins, R. P. & 386

Thiel, H. (Eds.), Introduction to the study of meiofauna (pp. 61-78.). Washington 387

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Gooday, A.J., Bett, B.J., Escobar, E., Ingole, B. S., Levin, L.A., Neira, C., Raman, 389

A.V., & Sellanes, J. (2010). Habitat heterogeneity and its influence on benthic 390

biodiversity in oxygen minimum zones. Marine Ecology, 31(1), 125-147. 391

Greiser, N., & Faubel, A. (1988). Biotic Factors. In: Higgins, R.P. and Thiel, H. (Eds.), 392

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Howes, J.R. (1986). Evaluation of Sarawak Wetlands and their importance to 395

waterbirds. Report 3. Pulau Bruit. Kuala Lumpur, Interwader Publication ,No. 10. 396

Islam, M.S., Sikder, M.N.A., Al-Imran, M., Hossain, M.B., Mallick, D., & Morshed, 397

M.M. (2013). Intertidal macrobenthic fauna of the Karnafuli Estuary: relations 398

with environmental variables. World Applied Sciences Journal, 21(9), 1366-1373. 399

Josefson, A.B., Forbes, T.L., & Rosenberg, R. (2002). Fate of phytodetritus in marine 400

sediments: functional importance of macrofauna community. Marine Ecology 401

Progress Series, 230, 71-85. 402

Lastra, M., Huz, R. D.L., Sánchez-Mata, A.G., Rodil, I.F., Aerts, K., Beloso, J., & 403

López, J. 2006. Ecology of exposed sandy beaches in northern Spain: 404

Environmental factors controlling macrofauna communities. Journal of Sea 405

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Lorenzen, C.J. (1967). Determination of chlorophyll and pheopigments: 407

spectrophotometric equations. Limnology Oceanography, 12, 343–346. 408

Lu, L. (2005). The relationship between soft-bottom macrobenthic communities and 409

environmental variables in Singaporean waters. Marine Pollution Bulletin, 51, 410

1034-1040. 411

Magni, P., Abe, N., & Montani, S. (2000). Quantification of microphytobrnthos 412

biomass in intertidal sediments: layer-dependent variation of chlorophyll a content 413

determined by spectrophotometer and HPLC methods. Société franco-japonaise ď 414

océanographie La mer, 38, 57-63. 415

Magni, P., Como, S., Montani, S., & Tsutsumi, H. (2006). Interlinked temporal changes 416

in environmental conditions, chemical characteristics of sediments and 417

macrofaunal assemblages in an estuarine intertidal sandflat (Seto Inland Sea, 418

Japan). Marine Biology, 149, 1185–1197. 419

Magni, P., & Montani, S. (2006). Seasonal patterns of pore-water nutrients, benthic 420

chlorophyll a and sedimentary AVS in a macrobenthos-rich tidal flat. 421

Hydrobiologia, 571, 297–311. 422

Mclachlan, A., & Jaramillo, E. (1995). Zonation on sandy beach. Oceanography and 423

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benthic fauna. Marine Ecology Progress Series, 79, 127-131. 449

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161. 462

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Massachusetts. Biological Bulletin of Marine Biology Laboratory, 152, 275-294. 470

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Figure 1: A) Map of Sarawak showing the location of Buntal Bay. B) Illustrations of line

transects performed for horizontal distribution study in intertidal flat of Buntal

Beach. T1-Transect 1, T2- Transect 2, ST-Station, HTL-High tide level, LTL-Low

tide level.

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Figure 2: Means and SD for univariate measures of macrobenthos community indices:

Number of species (S), Species richness index (d), Species evenness index (J’) and

Shannon’s Species diversity index (H’) in Transect 1 and Transect 2 of Buntal

Beach.

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Figure 3: A) Clustering analysis of macrobenthos community in Transect 1 based on Bray-

Curtis similarity. The cluster grouping can be identified by dark black lines after

performing SIMPROF test. B) Macrobenthos assemblages based on cluster

analysis and SIMPROF separated in the Multidimensional scaling (MDS)

ordination plot.

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Figure 4: A) Clustering analysis of macrobenthos community in Transect 2 based on Bray-

Curtis similarity. The cluster grouping can be identified by dark black lines after

performing SIMPROF test. B) Macrobenthos assemblages based on cluster

analysis and SIMPROF separated in the Multidimensional scaling (MDS)

ordination plot.

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Table 1: Summary of sediment granulometry and sediment biological parameters at Transect 1

and Transect 2. Cs: Coarse sand (%) = 1 mm; Ms: Medium sand (%) = 250 µm; Fs:

Fine sand (%) =150 µm; Vfs: Very fine sand = 63 µm; SC: Silt and clay (%) = <63

µm; TOM: Total organic matter (g/g sed); Chl a: Chlorophyll a (mg/m3).

Station Cs Ms Fs Vfs SC Mean Sorting Skewness TOM Chl a

Transect

1 ST1 28.5 43.6 21.7 5.9 0.4 1.7 0.8 0.2 0.4 3.0

ST2 23.0 44.0 30.3 1.6 0.1 1.8 0.8 -0.2 1.6 29.6

ST3 25.0 51.3 18.3 4.6 0.7 1.7 0.7 0.2 2.0 25.6

ST4 32.0 48.0 16.2 2.7 0.3 1.6 0.7 0.2 2.0 21.5

ST5 25.8 47.0 21.9 4.7 0.5 1.7 0.8 0.1 1.5 3.6

ST6 29.6 53.6 14.7 2.0 0.1 1.6 0.7 0.2 0.2 19.0

ST7 21.9 56.3 18.3 3.1 0.2 1.6 0.7 0.5 3.1 5.0

ST8 28.4 42.9 24.2 2.4 0.3 1.7 0.8 -0.1 2.8 15.0

ST9 30.4 43.4 22.8 1.9 0.3 1.6 0.8 0.0 1.6 10.2

ST10 29.3 42.8 23.7 2.3 0.1 1.7 0.8 0.0 2.7 5.1

ST11 29.6 43.4 22.0 1.8 0.1 1.6 0.8 -0.1 1.1 53.5

ST12 28.7 44.3 22.5 2.8 0.2 1.7 0.8 0.0 1.5 19.0

Transect

2 ST1 29.2 45.2 19.4 4.2 1.0 1.7 0.8 0.2 2.6 45.5

ST2 32.7 52.1 12.5 0.8 0.1 1.5 0.7 0.1 2.6 2.1

ST3 21.4 40.0 31.3 6.3 0.8 1.8 0.8 0.2 0.5 8.8

ST4 15.4 35.8 39.7 7.9 0.9 2.1 0.8 -0.4 0.8 18.1

ST5 23.1 32.2 28.1 14.3 2.2 1.9 0.9 -0.1 1.7 52.1

ST6 19.3 19.4 28.1 27.4 2.9 2.1 1.0 -0.5 0.5 83.1

ST7 32.7 36.3 20.2 7.7 1.0 1.7 0.9 0.2 2.7 10.0

ST8 27.5 40.5 23.0 6.7 2.0 1.7 0.8 0.2 1.7 1.7

ST9 23.3 31.1 30.6 13.0 1.9 1.9 0.9 -0.1 3.2 2.1

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Table 2: Species responsible for similarities of macrobenthos in each station at Transect 1. It

was indicated by SIMPER procedure which based on presence/absence data and

percent contribution (percentage contribution of total similarity).

Species Percent Contribution Percent Cumulative

Group ST1 Average similarity: 61.92 %

Nephtys sp. 1 20.67 20.67

Prionospio sp. 1 20.14 40.8

Nephtys sphaerocirrata 17.51 58.31


Barantolla sp. 34.2 34.2

Carinoma sp. 4 24.54 58.74


Cumella sp. 30.12 30.12


Nephtys sp. 1 12.21 12.21


Nephtys sp. 1 20.85 20.85



Umbonium elegans 17.71 36.78


Nephtys sphaerocirrata 30.13 30.13




Tellina sp. 2 37.92 37.92


Tellina sp. 1 41.9 41.9




Tellina sp. 1 61.42 61.42

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Table 3: Species responsible for similarities of macrobenthos at each station in Transect 2. It

was indicated by SIMPER procedure which based on presence/absence data and

percent contribution (percentage contribution of total similarity).

Species Percent Contribution Percent Cumulative


Nephtys sp.2 23.56 23.56

Glycera sp.3 17.63 41.19

Nephtys sp. 1 17.63 58.82

Glycera sp.1 9.99 68.81




Ophiura sp.1 31.06 31.06

Precephalothrix sp. 24.56 55.62




Magelona sp. 45.1 45.1




Tellina sp. 1 23.98 23.98

Spiophanes sp. 14.87 38.85






Notomastus lineatus 28 66.22

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Table 4: Summary of BIO-ENV analysis based on the Bray-Curtis similarities with fourth root

transformed data performed for macrobenthos density at Transect 1 (T1) and

Transect 2 (T2).

Station No of variables Factor/-s Correlations

T1 3 1,3,15 0.450

3 1,4,15 0.450

3 1,6,15 0.450

T2 2 2,14 0.487

2 6,14 0.487

2 14,15 0.487

4 2,3,6,14 0.457

Notes:

The environment factors associated to correlation selection: 1- Dissolved oxygen, 2- pH, 3 -

Salinity, 4-Temp, 6 - Chl-a, 14 - Fine sand, 15 - Very Fine sand.

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Table 5: Spearman’s rank correlations (r) between species density, species diversity index (H’)

and species evenness index (J’) with environment parameters at Transect 1.

Species density H’ J’

Dissolved oxygen r -0.235 0.186 0.648

p-level 0.463 0.564 0.023*

Salinity r 0.380 0.162 -0.303

p-level 0.223 0.615 0.339

Temperature r -0.160 -0.178 -0.046

p-level 0.618 0.579 0.886

Chlorophyll a r -0.420 -0.329 -0.406

p-level 0.175 0.297 0.191

Fine sand r 0.196 0.140 0.161

p-level 0.542 0.665 0.618

Note: Asterik (*) indicates a significant difference at p<0.05.

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Table 6: Spearman’s rank correlations (r) between species density, species diversity index (H’)

and species evenness index (J’) with environment parameters at Transect 2.

Species density (H’) (J’)

pH r 0.261 -0.252 0.609

p-level 0.498 0.513 0.082

Salinity r 0.571 0.804 -0.281

p-level 0.109 0.009* 0.464

Temperature r -0.527 0.083 0.274

p-level 0.145 0.832 0.475

Chlorophyll a r -0.035 -0.087 -0.794

p-level 0.929 0.825 0.011*

Fine sand r -0.381 -0.848 0.299

p-level 0.311 0.004* 0.434

Note: Asterik (*) indicates a significant difference at p<0.05.

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Species recorded on intertidal of Buntal Beach (Transect 1). Values were mean density. – absent

Station

ST

1

ST

2

ST

3

ST

4

ST

5

ST

6

ST

7

ST

8

ST

9

ST

10 ST 11

ST

12 Total

PHYLUM: MOLLUSCA

Class : Bivalvia

Tellinidae Tellina sp. 1 - - - - 1.33 5.33 6.67 8.0 - 26.67 - 78.67 126.7

Tellina sp. 2 - - - - - - - 2.67 20.0 6.67 - 13.33 42.7

Solenidae Solen regularis - - - 1.33 2.67 0.67 - 1.33 5.33 6.67 - - 18.0

Siliqua sp. - - 1.33 - - - - - - - - - 1.3

Mactridae Harvella sp. 1.33 - - 1.33 - - - 8.0 - - - - 10.7

Nuculanidae Nuculana acuta - - 1.33 - - - - - - - - - 1.3

Veneridae Meretrix meretrix 1.33 - - 4.0 2.67 - - 2.67 4.0 - 4.0 - 18.7

Class: Gastropoda

Trochidae Umbonium elegans - - 2.67 2.67 12.0 36.00 14.67 20.0 24.0 4.0 - 34.67 150.7

Nassariidae Nassarius jacksonianus - - 1.33 - - - - - - 1.33 - - 2.7

Nassarius reticulatus - - - - 2.67 4.0 2.67 1.33 2.67 2.67 - - 16.0

Nassarius sp. - - - - - - 2.67 1.33 - - - - 4.0

Naticidae Natica onca - - - - 2.67 - 1.33 - - - - - 4.0

Natica sp. - - - - - - - 20.0 - - - - 20.0

Olividae Olivella minuta 2.67 - - - - - - - 9.33 - - - 12.0

PHYLUM: ANNELIDA

Class: Polychaeta

Chrysopetalidae Chrysopetalum sp. - - - 2.67 - - - 1.33 - 1.33 - 1.33 6.7

Glyceridae Glycera sp. 1 - - - 2.67 4.0 10.67 5.33 25.33 10.67 4.0 - 2.67 65.3

Glycera sp. 2 1.33 - 2.67 - - 4.0 1.33 5.33 - - - - 14.7

Glycera sp. 3 4.0 - 1.33 1.33 1.33 2.67 4.0 13.33 - - - 10.67 38.7

Continued..

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Station

ST

1

ST

2

ST

3

ST

4

ST

5

ST

6

ST

7

ST

8

ST

9

ST

10 ST 11

ST

12 Total

Goniada sp. 1 - - 1.33 2.67 - 2.67 2.67 1.33 - - - - 10.7

Maldanidae Maldanella sp. - - 5.33 - - - - - 6.67 1.33 - - 13.3

Capitellidae Heteromastus filobranchus - - - - - - - 1.33 2.67 1.33 2.67 - 8.0

Notomastus hemipodus 4.00 - - - - 5.33 - 13.33 4.0 5.33 - 1.33 33.3

Notomastus lineatus - 8.00 10.67 - - - 2.67 10.67 8.0 - 20.0 8.0 68.0

Dasybranchus sp. - - - - 4.00 - 1.33 - 1.33 - - - 6.7

Mastobranchus sp. - - - - - - - 1.33 - - - 1.33 2.7

Barantolla sp. - 37.3 8.0 6.67 - - 4.00 14.67 4.0 6.67 14.67 - 96.0

Capitellidae sp. - - - - - - - - - - 2.67 1.33 4.0

Cirratulidae Cirrulatus sp. - - - - - - - 1.33 - - - - 1.3

Eunicidae Eunice sp. - - - - - - - 1.33 - - - - 1.3

Lumbrineridae Lumbrineris latreilli - - - - 1.33 2.67 - 5.33 - - - - 9.3

Lumbrineris heteropoda - - - - - - - 5.33 - - - 2.67 8.0

Magelonidae Magelona sp. - - 6.67 - 14.67 13.33 21.33 10.67 1.33 1.33 - - 69.3

Nephtyidae Nephtys sp. - - 4.00 12.0 34.67 37.33 - 12.0 - 5.33 - 4.00 109.3

Nephtys sphaerocirrata 17.33 - 21.33 16.0 26.67 34.67 24.00 34.67 16.0 - - 26.67 217.3

Nephtys impressa - - 1.33 2.67 - - 1.33 2.67 2.67 8.0 - - 18.7

Nephtys sp. 16.0 - - - 5.33 1.33 - - - - - - 22.7

Nereididae Ceratonereis sp. 9.33 - - 2.67 - - 6.67 2.67 - - - - 21.3

Nematonereis sp - - - 1.33 1.33 - - 1.33 - - - 2.67 6.7

Neanthes sp. - - 1.33 - - - - 5.33 - - - - 6.7

Nereis sp. - - - - - - - - - - - 2.67 2.7

Orbiniidae Phylo felix - - - 1.33 2.67 1.33 - 5.33 - - - - 10.7

Arabella sp. - - 1.33 - 9.33 - - - - - - - 10.7

Leitoscoloplos sp. - - - - - 1.33 - - - - - - 1.3

Oenonidae Drilonereis sp. - - - 1.33 - - 1.33 2.67 - - - - 5.3

Continued..

Continued.

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Station

ST

1

ST

2

ST

3

ST

4

ST

5

ST

6

ST

7

ST

8

ST

9

ST

10 ST 11

ST

12 Total

Onuphidae Diopatra sp. - - - - 2.67 - 2.67 1.33 - - - - 6.7

Phyllodocidae Phyllodoce arenae - - - - - - - 2.67 - - - - 2.7

Paranaitis speciosa 4.00 - 10.67 - 1.33 - - - - 1.33 - - 17.3

Ancistrosyllis sp. 1 - - 1.33 - - - - 1.33 - - - - 2.7

Ancistrosyllis sp. 2 - - - - - 9.33 - - - - - - 9.3

Eulalia sp. - - - - 1.33 1.33 1.33 - - - - - 4.0

Sabellidae Sabellidae sp. - - - - - - 1.33 - - - 1.33 - 2.7

Spionidae Prionospio sp. 1 8.00 - 17.33 - - 42.67 36.0 60.0 8.0 5.33 - 9.33 186.7

Prionospio sp. 2 1.33 - - - 4.67 16.0 - 9.33 5.33 - - - 36.7

Prionospio sp. 3 - - - - - - - 4.0 - - - - 4.0

Spiophanes wigleyi - - - - - - - 6.67 - - - - 6.7

Microspio sp. - - - 1.33 4.0 8.0 - 17.33 - - - - 30.7

Spiophanes sp. 1 - - 2.67 1.33 9.33 16.0 16.0 25.33 1.33 8.0 - - 80.0

Spiophanes sp. 2 - - 1.33 - - - - - - - - - 1.3

Spiophanes sp. 3 - - 1.33 - - - - - - - - - 1.3

Syllidae Polybostrichus sp. - - - - - - - - - - - 1.33 1.3

Syllidae sp. - - - 1.33 - - - - - - - - 1.3

Syllis armillaris - - - - 1.33 - - - - - - 5.33 6.7

Syllis sp. - - 1.33 - - - 1.33 - - 1.33 - - 4.0

Class: Clitellata

Subclass: Oligochaeta

Naididae Monopylephorus

rubroniveus - 1.33 - - - 1.33 - - - - - - 2.7

Smithsonidrilus sp. - - - - 1.33 - - - 2.67 1.33 - - 5.3

Continued..

Continued.

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Station

ST

1

ST

2

ST

3

ST

4

ST

5

ST

6

ST

7

ST

8

ST

9

ST

10 ST 11

ST

12 Total

PHYLUM: NEMERTINEA

Class: Anopla

Cephalothricidae Precephalothrix sp. - - - - - - - 2.67 - - - - 2.7

Carinomidae Carinoma sp. 1 2.67 - - - 1.33 1.33 1.33 4.0 5.33 6.67 2.0 - 24.7

Carinoma sp. 2 17.33 - 2.67 - 12.0 12.0 5.33 2.67 1.33 5.33 - - 58.7

Carinoma sp. 3 - - - - 1.33 - - - - 1.33 - - 2.7

Carinoma sp. 4 2.67 - - - - 2.67 1.33 - - - - - 6.7

Carinoma sp. 5 - 17.3 - - 6.67 - 1.33 - - 10.67 - 1.33 37.3

Class:Enopla

0.0

Lineidae Euborlasis sp. 1.33 - 1.33 - - 2.67 - - - - 2.67 - 8.0

Amphiporidae Amphiporus sp. 1 - 13.3 - - - 4.0 1.33 1.33 10.67 - 13.33 - 44.0

Amphiporus sp. 2 - - - - 1.33 - - 1.33 - - - - 2.7

PHYLUM: ARTHROPODA

Class: Crustacea

Order: Decapoda

Anomura Unidentified anomura crab - - - - - 1.33 - 9.33 - - - - 10.7

Macropthalmidae Macropthalmus sp. 1 - - - - - - - - - 4.0 - - 4.0

Macropthalmus sp. 2 - - 1.33 - - - - - 4.0 - - - 5.3

Macropthalmus sp. 3 - - - - - 4.0 - - 2.67 - - - 6.7

Dotillidae Dotilla sp. - - - - - - - - 1.33 - - - 1.3

Camptandriidae Xenopthalmus sp. - - 1.33 - - - - - - - - - 1.3

Unidentified crab zoea - - 4.0 - - - - - - 2.67 - - 6.7

Sergestidae Acetes sp. 1 - - - 2.67 - - - 1.33 - - - - 4.0

Order: Cumacea

0.0

Nannastacidae Cumella sp. - - 16.0 2.67 - 8.0 - - - 8.0 - - 34.7

Continued..

Continued.

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Station

ST

1

ST

2

ST

3

ST

4

ST

5

ST

6

ST

7

ST

8

ST

9

ST

10 ST 11

ST

12 Total

Order: Amphipoda

Amphilochidae Apolochus sp. - - 5.33 - - - - - - - - - 5.3

Ampeliscidae Ampelisca sp. 1 - - 1.33 - - - - - - - - - 1.3

Leucothoidae Leucothoe sp. 4.0 2.67 5.33 - - - - - - - - - 12.0

Aoridae Bermlos sp. - - - - - - - 3.0 - - - - 3.0

Phylum: ECHINODERMATA

Ophiuroidea Ophiura sp. 1 4.0 5.33 1.33 - - - - - - 12.0 - 17.33 40.0

Ophiura sp. 2 - - - - - 2.67 - - - - - - 2.7

Caudinidae Paracaudina sp. - - - - - - - - - 1.33 - - 1.3

Total density

102.7 85.3 148.0 72.0 178.0 296.7 173.3 397.7 165.3 152.0 63.3 226.7 2061.0

Note: Common : >20 ind.m-²

Less common : <20 ind.m-²

Rare : <5 ind.m-²

Continued.

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Species recorded on intertidal of Buntal Beach (Transect 2). Values were mean density. – absent

Phylum Species Station

ST1 ST2 ST3 ST4 ST5 ST6 ST7 ST8 ST9 Total

PHYLUM: MOLLUSCA

Class : Bivalvia

Tellinidae Tellina sp. 1 - 4.0 - - 2.67 32.0 66.67 32.0 - 137.33

Tellina sp. 2 - - 5.33 - 1.33 12.0 - - - 18.67

Solenidae Solen regularis - - 1.33 - - - - - - 1.33

Veneridae Meretrix meretrix - 4.0 - 2.67 1.33 - 1.33 4.0 - 13.33

Mactridae Harvella sp. - 1.33 - - - - - - - 1.33

Class: Gastropoda - Nassariidae Nassarius jacksonianus - - - - 1.33 - - - - 1.33

Nassarius reticulatus - 1.33 - - - - 4.0 - - 5.33

Naticidae Natica onca - - - - 1.33 5.33 5.33 - - 12.0

Natica gualtieriana - - - - - - 2.67 8.00 - 10.67

PHYLUM: ANNELIDA Class: Polychaeta

Chrysopetalidae Chrysopetalum sp. - - 1.33 - - - 20.0 1.33 - 22.67

Capitellidae Heteromastus filobranchus - 5.33 - - - - - - - 5.33

Notomastus hemipodus - - - - - - - - 2.67 2.67

Notomastus lineatus - - - - 2.67 16.0 - 22.67 10.67 52.0

Barantolla sp. - 10.67 5.33 2.67 - 54.67 38.67 33.33 16.0 161.33

Glyceridae Glycera sp. 1 6.67 12.0 5.33 8.0 12.0 6.67 14.67 13.33 - 78.67

Glycera sp. 2 - - - - 4.0 2.67 6.67 - - 13.33

Glycera sp. 3 5.33 2.67 8.0 4 - 2.67 1.33 - - 24.0

Goniada sp. - - 1.33 - - 2.67 - 4.0 - 8.0

Glycera dibranchiata - - 2.67 - - 2.67 - - - 5.33

Continued..

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Station


Lumbrireridae Lumbrineris heteropoda 2.67 1.33 - - 1.33 - - - - 5.33

Lumbrineris latreilli 5.33 - - - - - - - - 5.33

Magelonidae Magelona sp. - 2.67 - 2.67 22.67 5.33 61.33 52.0 - 146.67

Maldanidae Maldanella sp. - - - - - 2.67 - - - 2.67

Nephtyidae Nephtys sp. 1 10.67 - - - 4.0 8.0 18.67 28.0 - 69.33

Nephtys sp. 2 12.0 - - - - - - - - 12.0

Nephtys sphaerocirrata - 16.0 6.67 4.00 4.0 18.67 38.67 26.67 8.0 122.67

Nephtys impressa - 4.0 - - 2.67 - 2.67 14.67 - 24.0

Nereididae Ceratonereis sp. - 1.33 1.33 - 4.00 1.33 8.0 6.67 - 22.67

Nematonereis sp. - - - - - 2.67 17.33 - - 20.0

Neanthes sp. - 1.33 - - 2.67 - - - - 4.0

Phyllodocidae Phyllodoce arenae - - - - - - 4.0 2.67 - 6.67

Paranaitis speciosa - - - - - - 14.67 1.33 - 16.0

Eulalia sp. - 2.67 - - - 5.33 5.33 - - 13.33

Spionidae Prionospio sp. 1 - 10.67 - - - 6.67 18.67 4.0 - 40.0

Prionospio sp. 2 - 2.67 - - - - - 4.0 - 6.67

Prionospio sp. 3 - - - - - 1.33 4.00 - - 5.33

Spiophanes sp. 1 - - - - - 18.00 42.67 6.67 - 67.33

Spiophanes wigleyi - - - - - - 4.0 8.0 - 12.0

Microspio sp. - - - - - - 5.33 6.67 - 12.0

Sabellidae Sabellidae sp. 1 - - - - - 1.33 - - - 1.33

Sabella sp. - - - - - - 1.33 4.0 - 5.33

Syllidae Polybostrichus sp. - - - - - 1.33 4.0 - - 5.33

Syllis armillaris - - - - 1.33 1.33 - - - 2.67

Class: Clitellata

Subclass: Oligochaeta Continued..

Continued.

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Station


Naididae Monopylephorus rubroniveus - - - - - - 1.33 - - 1.33

PHYLUM: NEMERTINEA

Class: Anopla

Cephalothricidae Precephalothrix sp. - 5.33 13.33 - 2.67 - - 4.0 - 25.33

Carinomidae Carinoma sp. 1 - - - - - 1.33 4.0 - - 5.33

Carinoma sp. 2 - - 2.67 2.67 - 1.33 2.67 8.0 2.67 20.0

Carinoma sp. 3 - 1.33 - - - - - 1.33 - 2.67

Carinoma sp. 4 - - - - - - - 2.67 - 2.67

Carinoma sp. 5 1.33 2.67 1.33 - - 12.00 1.33 - - 18.67

Class:Enopla

Lineidae Euborlasis sp. - - - - - 2.67 4.0 - - 6.67

Amphiporidae Amphiporus sp. 1 2.67 1.33 2.67 - 5.33 - - - - 12.0

Amphiporus sp. 2 - - - - - - 1.33 - - 1.33

PHYLUM: ARTHROPODA

Class: Crustacea Order: Decapoda

Anomura Unidentified Anomura crab 4.00 2.67 5.33 - - - - - - 12.0

Macropthalmidae Macropthalmus sp. 1 - - - - - 4.0 - - - 4.0

Dotillidae Dotilla sp. - - - - - - 5.33 1.33 - 6.67

Leucosiidae Leucosiidae sp. - 1.33 - - - - - - - 1.33

Unidentified crab zoea - 1.33 - - - - - - - 1.33

Order: Cumacea

Nannastacidae Cumella sp. - 4.0 - - 4.0 - - - - 8.0

Bodotridae Cyclaspis sp. - - 2.67 - 2.67 - - - - 5.33

Order: Amphipoda Continued..

Continued.

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Station


Amphilochidae Apolochus sp. - - - - - - 8.00 - - 8.0

Leucothoidae Leucothoe sp. - 2.67 - - - - - - - 2.67

Leucothoe sp. 1 - - - - - 4.0 10.67 - - 14.67

Aoridae Bermlos sp. - - 2.67 - 1.33 - - - - 4.00

PHYLUM: ECHINODERMATA

Ophiuroidea Ophiura sp. 1 - 8.0 14.67 - - 2.67 - - - 25.33

Ophiura sp. 2 - - 1.33 - 1.33 4.0 - - - 6.67

Total density �� 50.67 114.67 85.33 26.67 86.67 243.33 450.67 302.67 40.0 1400.67

Note: Common : >20 ind.m-²

Less common : <20 ind.m-²

Rare : <5 ind.m-²

Continued.

Page 47 of 47

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distribution of intertidal flat macrobenthos in buntal bay ... · 50 bako-buntal bay (n...

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