1 Original Article

2 Study of Diversity and Morphometry in Edible Bivalves and Gastropods from a Coastal

3 Wetland in Sarawak

1* 3 1 1 4 Mohd Hanafi Idris , Hadi Hamli , Abu Hena Mustafa Kamal , Roslizawati Ab Lah , Nik

5 Mohd Shibli Nik Jaafar2

6

7 1Faculty of Fisheries and Food Science, Universiti Malaysia Terengganu, 21030 Kuala Nerus,

8 Terengganu, Malaysia

9 2Faculty of Science and Marine Environment, Universiti Malaysia Terengganu, 21030 Kuala

10 Nerus, Terengganu, Malaysia

11 3Department of Science and Fishery, Faculty of Agriculture and Food Sciences

12 Universiti Putra Malaysia Bintulu Sarawak Campus

13 Nyabau Road, P.O. Box 396, 97008, Bintulu, Sarawak, Malaysia

14 *Corresponding author, Email address: [email protected]

15

16 Abstract

17 Diversity and morphometry of edible, bivalve and gastropod classes of mollusc were 18 investigated from July e82010 to January 2012, for a particular set of bivalve , at 19 selected coastal divisions of Sarawak including Kuching, Sibu, Mukah, Bintulu, Miri, 20 Limbang, and Lawas. A total of 41 edible species were recorded and identified; comprising 21 11 families of gastropods represented by 21 species, and 12 family of bivalves represented by 22 20 species. For bivalves, species of the family Cyrenidae (Geloina erosa, G. expansa and G. 23 bengalensis) were more widely distributed than bivalve species. The gastropod, family of 24 Potamididae (Cerithidea obtusa, C. quadrata, C. rizophorarum and Telescopium 25 telescopium) were also commonly observed along the entire seven divisions. Species 26 diversity was found to be highest in Bintulu division for both classes, and consisted of 13 27 species of gastropod and 11 species of bivalve. Jaccard’s index showed high similarity 28 amongst edible bivalves for Limbang division versus Miri division (0.75), and for Limbang 29 versus Lawas (0.75). For gastropods, Sibu versus Mukah showed higher similarity (0.50). 30 Morphometric study of bivalves indicated that VPM/SL and LCT/SL features were 31 applicable as an identification key to differentiate between Geloina spp. This morphometric 32 identification key could be developed further for identification of other species in the mollusc 33 family.

34

35 Keywords: Edible gastropod, bivalve, diversity, distribution, Sarawak

36

37 1. Introduction

38 The State of Sarawak, Malaysia contains vast areas of wetlands, including brackish

39 water, coastal marine, and mangrove areas. The state wetlands comprise contains 1.24 million

40 hectares which covers 13% of the state’s total land area (Page, 2011). This wetland provides

41 support for extensive floral and faunal biodiversity. Molluscs are part of the faunal

42 composition of the wetlands of Sarawak, and one of the most important protein resources to

43 surrounding human the communities. Apart from providing good nutrition, molluscs play an

44 important economic role in Sarawak.

45 The contribution of edible mollusc to local economies within Sarawak is high,

46 notwithstanding, this resource has yet to be well documented. To the best of our knowledge,

47 only a few studies have documented species of bivalves and gastropods occurring in

48 Malaysia; (Hamli et al., 2015a; Hamli et al., 2015b; Idris et al., 2011; Abdullah, Sidi, & Aris,

49 2007; Sallih, 2005; Abu Hena et al., 2004; Nakao, Nomura & Satar, 1989). Previous study

50 has recorded edible mollusc distribution in the tropical region of Thailand (Nateewathana,

51 1995), but it is difficult to develop an accurate figure for local consumption of edible bivalve

52 and gastropod species in Sarawak, Malaysia. This paper aims to investigate the diversity, 53 morphological characteristics and habitat of edible mollusc to improve information for the

54 selected division in Sarawak. The majority of the bivalve and gastropod species found in

55 wetland coastal waters of Sarawak, in the coastal divisions, hence this regional study provides

56 valuable information on bivalves and gastropods, which could be used to further develop bivalve and

57 gastropod fisheries in future.

58

59 2. Materials and Methods

60 2.1 Study Area

61 Samples were collected from native markets, and fish markets from the selected seven

62 divisions of Kuching, Mukah, Sibu, Bintulu, Sibu, Bintulu, Miri, and Lawas, Sarawak.

63 (Figure 1). Samples were preserved in ice-filled boxes and transferred to the laboratory for

64 further analysis and study.

65 2.2 Species Identification and Morphometric Study as a Demarcation Tool

66 The identification of bivalves was conducted following procedures common to a

67 number of researchers, namely, Morton (1984), Poutiers (1998), Mass, O’ Mullan, Lutz, and

68 Vrijenhoek (1999), Yuh-Wen, Hon-Cheng, Sin-Che and Chaolun (2001), and Okutani

69 (2017). Gastropod samples were identified following the studies by Nateewathana (1995),

70 Kohler and Glaubrecht (2001), Poutiers (1998), Perez, Clark and Lydeard (2004), Tan and

71 Clements (2008), and Mujiano (2009). Specimens of bivalve and gastropod were measured

72 according to morphometric characteristics using digital Mitutyo vernier calliper (0.01±mm)

73 (Hamli et al., 2015a and Idris et al., 2008). A total of 20 characteristics (15 for bivalve and 5

74 for gastropods) were used to differentiate the species (Table 1; Figure 2).

75 The current morphometric focused only on a single genus of bivalve from Geloina

76 genus (G erosa, G. expansa, G. bengalensis.) with a total of 436 matured individuals. The 77 sum of 15 shell morphometrics, numbered one through fifteen (Table 1) had been measured,

78 which were then divided by shell length to convert the measurements to proportionate ratio

79 by size, for each Geloina species.

80 2.3. Statistical Analysis

81 Measurements of collected species from the selected Sarawak divisions were

82 correlated with geographical source to establish a distribution pattern. Similarities in species

83 numbers within the respective divisions were evaluated as either present (1), or absent (0)

84 based on Jaccard’s Index (Ludwig & Reynolds, 1988).

85 Jaccard’s Index = A/(A+B+C)

86 Where

87 A = total number of species present in both communities

88 B = the number of species present in community 1 but not at community 2

89 C = the number of species present in community 2 but not at community 1

90 The samples from the Geloina species, from the study areas in the present study were

91 chosen for morphometric differentiation within species by scrutinising shell proportion ratio

92 using analysis of variance (ANOVA) with general linear model (GLM), and using statistical

93 analysis computer software (SAS) version 9.1 (SAS Institute, 2004).

94

95 3. Results and Discussion

96 3.1 Distribution

97 A total of 41 edible species comprised 12 families of bivalves representing 20 species

98 (Table 2), and 11 families of gastropods representing 21 species (Table 3) were recorded, and 99 identified from seven divisions of Sarawak (Figures 3 and 4). Cyrenidae (Bivalve) and

100 Potamididae (Gastropod) families were widely distributed in seven and six divisions,

101 respectively. Bintulu division presented the greatest variety of bivalves and gastropods with a

102 total of 24 species (13 gastropods, and 11 bivalves) recorded, while Sibu division presented

103 only minor variation.

104 Mollusc species presented in this study were low when compared to Printrakoon,

105 Wells and Chitramvong (2008), and other studies, due to the current study focus on edible

106 species alone. Printrakoon et al. (2008), Frith, Tantanasiriwong and Bathia et al. (1976) and

107 Brandt (1974) recorded 47, 43 and 56 species of mangrove, coastal, and estuarine mollusc,

108 respectively in Thailand. Macintosh, Ashton and Havanon (2002) found 34 species of

109 mollusc in the Ranong biosphere reserve in Thailand. Jiang and Li (1995) recorded 52

110 species of in the Jiulong river estuary, China. However, the above studies were taxonomic,

111 diversity and distribution studies, rather than studies specific to edible molluscs in specified

112 coastal, tropical wetlands. The current study indicates that the diversity of edible mollusc in

113 this region is less, although Somchai (1995) reported that gastropods were a major class of

114 mollusc with high market value on Phuket Island, Thailand.

115 The marsh clams, Geloina erosa and G. bengalensis were both within the sampling

116 area. Within the Indo-Pacific area, they can be described under the subgenus Geloina

117 (Morton, 1984), which subgenus is also distributed in the mangrove area of China (Han et al.,

118 2003), Australia (Wells, 1986) and Thailand (Printrakoon et al., 2008). Species Geloina

119 expansa (), Cerithidea rizophorarum, and C. obtusa () were found

120 distributed within four divisions in Sarawak. Other gastropod and bivalve species were found

121 within one to three divisions only. Presence of G. erosa and G. bengalensis at almost study

122 area was equivalent with to presence of mangrove area at Sarawak. 123

124 3.2 Similarity Indices

125 Analysis of the similarity index, described above, showed high similarities for

126 bivalves in Limbang versus Miri (0.75), and in Limbang versus Lawas (0.75), and high

127 lower similarity for edible gastropod in Sibu versus Mukah (0.50). Similarities as low as 0.00

128 was observed for bivalves in Kuching versus Mukah divisions and in Lawas versus Kuching.

129 Similarities were also as low as 0.00 for edible gastropod in Lawas versus Sibu, Lawas versus

130 Mukah, Lawas versus Bintulu, Lawas versus Miri, Lawas versus Limbang, Bintulu versus

131 Sibu and Bintulu versus Mukah divisions. The similarity index value for edible bivalve and

132 gastropod between each division could probably account for or be related to the psence of

133 wide mangroves and muddy area supporting suitable habitats these molluscs. Furthermore,

134 Puri, Namboothri and Shanker (2014) described that increasing similarity of gastropod

135 communities showed a relationship with decreasing geographical distance between study

136 locations.

137 3.3 Morphometrics

138 Study on 14 calculated ratios of morphometric characteristics on Geloina erosa, G.

139 expansa and G. bengalensis indicated significant difference (p<0.05) for VPM/SL and

140 LCT/SL (Table 4). Length of cardinal tooth proportion with SL was recorded as significantly

141 higher (p<0.05) for G. bengalensis when compared with other species. For G. erosa,

142 characteristics recorded no significant difference (p>0.05) when compared with G. expansa.

143 Geloina bengalensis recorded significant difference (p<0.05) for VPM/SL since this

144 characteristic is absent in G. erosa and G. expansa.

145 Previous study of different populations of Corbiculidae (Corbicula fluminea) showed

146 shell height to be significantly correlated with shell length (Araujo, Moreno, & Ramo, 1993). 147 Morton (1984) reported that width and height of G. erosa and G. expansa were extremely

148 variable which caused these characteristics to be unreliable for analysis of dimensional

149 relationships. There are many reasons that can cause such outcome. Some bivalves grow to

150 become higher and wider to overcome the turbulence and currents in the surrounding area

151 (Hinch & Bailey, 1988). With respect to the mangrove habitat of this species, they are always

152 affected by tide. Bivalve shell, especially from a mangrove area was vulnerable to the erosive

153 effects of the acid mangal soil (Morton, 1984). Moreover, the presence of predators can also

154 affect shell characteristics, as reported by Preston and Roberts (2006) on Calliostoma

155 zizyphinum. Therefore, the current study did not depend on major dimensions (length, height

156 and width) alone, but an additional characteristic was used to support differentiation between

157 species. The supplementary features chosen were significant in morphometric assessment for

158 gastropod and bivalve to reduce or remove attribution to impacts of the environment, or and

159 predators.

160 4. Conclusion

161 The higher number of edible bivalves and gastropods in Sarawak could be due either

162 to habitat preferences of molluscs, or to over exploitation of the natural resource by local

163 fishermen, e.g., due to rapid urbanization and, customer demand. This may affect the total

164 edible mollusc diversity within Sarawak. Diversity, abundance, and shell features of edible

165 molluscs are significantly affected by changes in habitat. Therefore, fluctuation in

166 environmental conditions, generates changes shell formation, which in turn influence species

167 identification, while the current study was able to identify new shell characteristics to support

168 discrimination between species exposed to abiotic variations. Further study is required to

169 identify, monitor and classify the diversity and population status of edible mollusc in this

170 region in support of conservation and management. 171

172 Acknowledgments

173 The research was supported by funding of project No. 5523703 FRGS, from the

174 Ministry of Education Malaysia. Technical and logistic support was provided by Universiti

175 Putra Malaysia Bintulu Sarawak Campus. The authors would like to thank Mr. Azwandi and

176 Awangku Nizam for their help during data collection in Sarawak, which made this study

177 possible.

178 References

179 Abdullah, M. H., Sidi, J., & Aris, A. Z. (2007). Heavy Metal (Cd, Cu, Pb, Zn) in Meretrix

180 meretrix Roding, Water and Sediments from Estuaries in Sabah, North Borneo.

181 International Journal of Environmental and Science Education, 2 (3), 69–74.

182 Abu Hena, M. K., Hishamuddin, O., Misri, K., Abdullah, F., & Loo, K. K. (2004). Benthic

183 Faunal Composition of Penaeus monodon Fabricius Culture Pond West Coast of

184 Peninsular Malaysia. Journal of Biological Science, 4 (5), 631–636.

185 Araujo, R., Moreno, D., & Ramos, M. A. (1993). The Asiatic Clam Corbicula fluminea

186 (Muller 1774) (Bivalvia: Corbiculidae) in Europe. American Malacological Bulletin, 10

187 (1), 39–49.

188 Brandt, R. A. M. (1974). The Non Marine Aquatic Mollusc of Thailand. Archiv Fur

189 Molluskenkunde, vol. 105 (I-iv), 1–423.

190 Frith, D. W., Tantanasiriwong, R., & Bhatia, O. (1976). Zonation of Macrofauna on a

191 Mangrove Shore, Phuket Island. Phuket Marine Biological Center Research Bulletin,

192 10, 1–37. 193 Hamli, H., Abd. Rahim, A., Idris, M.H., Abu Hena, M.K. & Wong, S.K. (2015a).

194 Morphometric Variation Among Three Local Mangrove Clam Species of Corbiculidae.

195 Songklankaria Journal of Science Technology. 37(1), 15-20.

196 Hamli, H., Idris, M.H., Rajaee, A.H. & Abu Hena, M.K. (2015b). Reproductive Cycle of

197 Hard Clam, Meretrix lyrata Sowerby, 1851 (Bivalvia: ) from Sarawak,

198 Malaysia. Tropical Life Science Research, 26(2), 59-72.

199 Han, W., Lui, J., He, X., Cai, Y., Ye, F., Xuan, L., & Ye, N. (2003). Shellfish and Fish

200 Biodiversity of Mangrove Ecosystems in Leizhou Peninsula, China. Journal of Coastal

201 Development, 7(1), 21–29.

202 Hinch, S. G., & Bailey, R. C. (1988). Within- and Among-Lake Variation in Shell

203 Morphology of the Freshwater Clam Elliptio complanata (Bivalvia: Unionidae) from

204 South-Central Ontario Lakes. Hydrobiologia 157, 27–32.

205 Idris, M. H., Arshad, A., Amin S. M. N., Japar Sidik, B., Daud, S. K., Mazlan, A. G., Zakaria,

206 M. S., & Yusoff, F. M. (2011). Age, Growth and Length-Weight Relationships of

207 Pinna bicolor Gmelin (Bivalvia: Pinnidae) in the Seagrass Beds of Sungai Pulai

208 Estuary, Johor, Peninsular Malaysia. Journal Applied Ichthyology, 28, 597-600.

209 Idris, M. H., Arshad, A., Japar Sidik, B., Mazlan, A.G. and Daud, S.K. (2008). Morphometric

210 Analysis as Application Tool to Differentiate Three Local Pen Shells Species.

211 Pertanika Journal of Tropical Agriculture Science, 31(2), 287-298.

212 Jiang, J. X., & Li, R. G. (1995). An Ecological Study on the Mollusc in Mangrove Areas in

213 the Estuary of Jiulong River. Hydrobiologia, 295, 213–220.

214 Kohler, F., & Glaubrecht, M. (2001). Toward a Systematic Revision of the Southeast Asian

215 Freshwater Gastropod Brotia H. Adams, 1866 (Cerithiodea: Pachychilidae): An

216 Account of Species from Around the South East China Sea. Journal of Molluscan

217 studies, 67, 281–318. 218 Ludwig, J. A., & Reynolds, J. F. (1988). Statistical Ecology: A Primer on Methods and

219 Computing. A Wiley-Interscience Publication. 338 p.

220 Macintosh, D. J., Ashton, E. C., & Havanon, S. (2002). Mangrove Rehabilitation and

221 Intertidal Biodiversity: A Study in the Ranong Mangrove Ecosystem Thailand.

222 Estuarine, Coastal and Shelf Science, 55, 331–345.

223 Mass, P. A. Y., O’ Mullan, G. D., Lutz, R. A., & Vrijenhoek, R. C. (1999). Genetic and

224 Morphometric Characterization of Mussels (Bivalvia: Mytilidae) from Mid-Atlantic

225 Hydrothermal Vents. Biological Bulletin, 196, 265–272.

226 Morton, B. (1984). Review of Polymesoda (Geloina) Gray 1842 (Bivalvia: Corbiculacea)

227 from Indo-Pacific Mangroves. Asian Marine Biology, 1, 77–86.

228 Mujiano, N. (2009). Mudwhelks (Gastropoda: Potamididae) from Mangroves of Ujung Kulon

229 National Park, Banten. Jurnal Biologi, 13(2), 51–56.

230 Nakao, S., Nomura, H., &. Satar. M. K. B. A. (1989). Macrobenthos and Sedimentary

231 Environments in a Malaysian Intertidal Mudflat of the Cockle Bed. Bulletin of the

232 Faculty of Fisheries Hokkaido University, 40(4), 203s–213.

233 Nateewathana, A. (1995).Taxanomic Accaount of Commercial Edible Molluscs, Excluding

234 Cephalopodas, of Thailand. Phuket Marine Biological Center, 20, 93–116.

235 Okutani, T. (2017). Marine Mollusks in Japan. 2nd edition. Tokai University Press. 1375p.

236 Page, S., 2011. Biodiversity Information on Peat swamp Forest in S.E. Asia.

237 Available:http://www.strapeat.alterra.nl/download/6%20biodiversity%20information.p

238 df.

239 Perez, K. E., Clark, S. A., & Lydeard, C. (2004). A Primer to Freshwater Gastropod

240 Identification. In Freshwater Gastropod Identification Workshop. pp 1–61. 241 Poutiers, M. (1998). The Living Marine Resources of the Western Central Pacific: FAO

242 Species Identification Guide for Fishery Purpose, K.E. Carpenter and V.H. Niem, Ed.

243 Rome, Italy: Food and Agriculture Organization of the United Nations, 1, 123–362.

244 Preston, S. J., & Roberts, D. (2007). Variation in Shell Morphology of Calliostoma

245 zizyphinum (Gastropoda: ). Journal of Molluscan Studies, 73(1), 101-104.

246

247 Puri, M., Namboothri, N., & Shanker, K. (2014). Multi‐Scale Patterns in Co‐occurrence of

248 Rocky Inter‐tidal Gastropods along the West Coast of India. Oikos, 123(3), 345-355.

249 Printrakoon, C., Wells, F. E., & Chitramvong, Y. (2008). Distribution of Molluscs in

250 Mangrove at Six Sites in the Upper Gulf of Thailand. The Raffles Bulletin of Zoology,

251 18, 247–257.

252 Sallih, K. (2005). Mussel Farming in the State of Sarawak, Malaysia: A Feasibility Study.

253 Fisheries Training Programme. pp. 44.

254 Somchai, B. (1995). The Market Value of Rare and Common Molluscs, Phuket Island,

255 Thailand. Phuket Marine Biological Center, 20, 35–38.

256 Tan, S. K., & Clements, R., (2008). and Distribution of Neritidae (:

257 Gastropoda) in Singapore. Zoological Studies 47(4), 481–494.

258 Wells, F. E. (1986). Distribution of Molluscs Across a Pneumatophore Boundary in a Small

259 Bay in North-Western Australia. Journal of Molluscan Studies, 52, 83–90.

260 Yuh-Wen, C., Hon-Cheng, C., Sin-Che, L., & Chaolun, A. C. (2001). Morphometric Analysis

261 of Shell and Operculum Variation in Viviparid Snail, Cipangopaludina chinensis

262 (Mollusca: Gastropoda), in Taiwan. Zoological Studies, 41(3), 321–331.

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264

265 266

267

268

269 270 Figure 1 Sampling location from seven divisions of Sarawak.

271

272

273

274

275

276

277 278

279 Figure 2 Measurements of shell characters for bivalvia and gastropoda (adopted from 280 Mass et al., 1999; Ramesh and Ravichandaran, 2008). Note: 1(SL), 2(SH), 3(SW), 4(LL), 281 5(PL), 6(AL), 7(UL), 8(LCT), 9(LPAS), 10(PW), 11(PW), 12(PVM), 13(PAPM), 282 14(AAAM), 15(VPM) 16(sl), 17(sw), 18(al), 19(aw), 20(op).

283 284 285 Figure 3 Edible bivalve species recorded from Sarawak: (A) Geloina expansa, (B) 286 Geloina erosa, (C) Geloina bengalensis (D) Meretrix meretrix (E) Meretrix lyrata (F) 287 undulatus (G) Circe scripta (H) Solen regularies (I) Solen lamacrkii (J) 288 Pharella acutidens.

289

290

291

292 293

294

295 Figure 3 (Cont’d) (K) Amusium pleuronectes, (L) Tegillarca granosa, (M) Pholas 296 orientalis, (N) Gluconome virens, (O) Placuna placenta, (P) Crassostrea lugubris, 297 (Q) ephippium, (R) Sinanodonta woodiana, (S) Pilsbryoconcha exilis 298 (T) Arcuatula arcuatula.

299 300 301

302 Figure 4 Edible gastropods from Sarawak; (A) Cerithidea obtusa, (B) Cerithidea 303 rizophorarum, (C) radiatus, (D) Cerithidea quoyii, (E) Telescopium 304 telescopium, (F) Monodonta labio, (G) Nerita articulata, (H) Nerita chamaeleon, 305 (I) Melo melo, (J) Nerita albicilla, (K) Clithon ritropictus, (L) Planaxis sulcatus.

306

307 308 309

310 Figure 4 (continued) (M) Brotia costula, (N) Melanoides costellaris, (O) Tylothais virgata 311 (P) Blanocochlis glandiformis, (Q), Tylomelania helmuti (R),Turbo crasus, (S) Pilla 312 ampullacea, (T), Ellobium aurisjudae and (U) Pomacea bridgesii.

313

314

315

316

317

318 319 Tables

320 321 Table 1 Morphometric characteristic used for bivalve identification with the abbreviation 322 (*=Additional characteristic used during study period) 323 Measurement Abbreviation Details Number

1 SL Distance from the anterior margin to posterior margin

2 SH Distance between ventral margin and umbo

3 SW Distance from left shell to the right shell

4 LL Ligament length

5 UL Distance from umbo to posterior margin

6 AL Distance from the posterior part of umbo to the anterior margin of the shell

7 PL Distance from the posterior part of umbo to the posterior margin of the shell

8 LCT Length from anterior cardinal tooth to posterior cardinal tooth

9 LPAS Distance from anterior adductor muscle scar to posterior adductor muscle scar

10 PW Wide of posterior adductor muscle scar

11 AW Wide of anterior adductor muscle scar

12 PVM Distance from ventral margin to pallial line

13 PAPM Distance from posterior adductor muscle scar to posterior margin of the shell

14 AAAM Distance from anterior adductor muscle scar to anterior margin of the shell

15 VPM Ventral posterior margin length

16 sl Shell length

17 sw Shell width

18 al length

19 aw Aperture width

20 op Outer thickness

324 325 Table 2 Checklist and distribution of edible bivalve species from eight different divisions in Sarawak, Malaysia.

Family Species Habitat Kuching Sibu Mukah Bintulu Miri Limbang Lawas

Veneridae Circe scripta Salt water - - - + - - -

Paratapes undulatus Salt water - - + - - - -

Meretrix meretrix Salt water - - - + - - -

Meretrix lyrata Salt water + ------

Solenidae Solen regularies Salt water + ------

Solen lamarckii Salt water + ------

Brackish Pharella acutidens + ------water

Brackish Cyrenidae Geloina bengalensis + + - + + + - water

Brackish Geloina erosa - + + + + + + water

Brackish Geloina expansa - - - + + + + water

Unionidae Sinanodonta woodiana Fresh water - - - + + - -

Pilsbryoconcha exilis Fresh water - - - - + - -

Arcidae Tegillarca granosa Salt water + - - + - - -

Myoida Pholas orientalis Salt water + ------

Gluconomidae Gluconome virens Salt water + ------

Pectinidae Amusium pleuronectes Salt water + - - + - - -

Placunidae Placuna plcenta Salt water - - - + - - -

Ostreidae Crassostrea lugubris Salt water - - - + - - -

Mytilidae Arcuatula arcuatula Salt water ------+

Isognomonidae Isognomon ephippium Salt water - - - + - - -

326 Note: (+) = Present, (-) = Absent

327

328

329

330

331

332

333

334

335

336 337 338 Table 3 Checklist of gastropod species existing from eight different divisions in Sarawak

Family Species Kuching Sibu Mukah Bintulu Miri Limbang Lawas

Neritidae Nerita chamaeleon Salt water - - - + - - -

Nerita articulata Brackish water + - + - - - -

Nerita albicilla Salt water - - - + - - -

Clithon retropictum Brackish water - + - - - - -

Cerithidea Brackish water Potamididae rizophorarum + + + - - + -

Cerithidea obtusa Brackish water + + + - + - -

Cerithidea quoyii Brackish water ------+

Telescopium Brackish water telescopium ------+

Ampullariidae Pomacea bridgesii Fresh water - - - + - - -

Pilla ampullacea Fresh water + - - + + + -

Trochidae Trochus radiatus Salt water - - - + - - -

Monodonta labio Salt water - - - + - - -

Pachychilidae Brotia costula Fresh water - - - + - - -

Tylomelania helmuti Fresh water - - - + + - -

Thiaridae Melanoides costellaris Fresh water + ------

Blanocochlis Fresh water glandiformis - - - + - + -

Planaxide Planaxis sulcatus Salt water - - - + - - -

Turbinidae Turbo crasus Salt water - - - + - - -

Muricidae Tylothais virgata Salt water - - - + - - -

Volutidae Melo melo Salt water - - - + - - -

Melampidae Ellobium aurisjudae Brackish water + ------

339 Note : (+) = present; (–) = absent

340

341

342

343

344

345

346

347 348

349

350

351 Table 4 Analysis of one way ANOVA with general linear model (GLM) for 14 352 morphometric characteristic proportion with shell length of three Geloina 353 species

Morphometric Geloina erosa Geloina expansa Geloina F value p characteristic bengalensis N=194 N=139 N=103

SH/SL 0.89±0.21a 0.89±0.01a 0.93±0.01a 0.39 0.678ns

SW/SL 0.53±0.03a 0.54±0.01a 0.55±0.01a 0.11 0.895ns

UL/SL 0.75±0.04a 0.78±0.01a 0.80±0.00a 0.58 0.564ns

LL/SL 0.32±0.02a 0.31±0.00a 0.34±0.01a 0.91 0.409ns

AL/SL 0.56±0.03a 0.58±0.01a 0.60±0.01a 0.66 0.523ns

PL/SL 0.72±0.04a 0.75±0.01a 0.60±0.01a 0.48 0.625ns

PAPM/SL 0.05±0.00a 0.05±0.00a 0.05±0.00a 1.99 0.149ns

AAAM/SL 0.05±0.00a 0.05±0.00a 0.05±0.00a 1.22 0.305ns

PVM/SL 0.10±0.01a 0.10±0.00a 0.09±0.00a 1.39 0.261ns

LPAS/SL 0.60±0.03a 0.63±0.00a 0.63±0.00a 0.68 0.535ns

AW/SL 0.12±0.01a 0.12±0.00a 0.13±0.00a 1.78 0.180ns

PW/SL 0.17±0.01a 0.15±0.00a 0.17±0.00a 1.91 0.160ns

LCT/SL 0.16±0.01ab 0.14±0.00a 0.17±0.00b 4.45 0.017

VPM/SL NA NA 0.36±0.00 NA NA

354 SL=Shell length, SH=Shell height , SW=Shell weight, UL=Umbo length, LL=Ligament 355 length, AL=Anterior length, PL=Posterior length, PAPM=Posterior adductor scar to posterior 356 margin, AAAM=Anterior adductor scar to anterior margin, PVM=Pallial line to ventral 357 margin,LPAS=Length of posterior adductor scar to anterior adductor scar, AW=Anterior 358 adductor scar width, PW=Posterior adductor scar width, LCT=Length of cardinal tooth, 359 VPM=Ventral posterior margin length, NA=Not available.

360

361 Note: difference superscripts indicate a significantly difference at level p<0.05 using Tukey 362 mean comparison, ns: no significantly difference

363

364

365 366

367

368

369