Fish Fauna of the Sikao Creek Mangrove Estuary, Trang, Thailand

FISHERIES SCIENCE 2002; 68: 10–17

Original Article

Fish fauna of the Sikao Creek mangrove estuary, Trang, Thailand

Prasert TONGNUNUI,1 Kou IKEJIMA,2–4* Takeshi YAMANE,4 Masahiro HORINOUCHI,4 Tomon MEDEJ,1 Mitsuhiko SANO,4 Hisashi KUROKURA4 AND Toru TANIUCHI2a

1Faculty of Science and Fisheries Technology, Rajamangala Institute of Technology, Sikao, Trang 92150, Thailand, 2Department of Aquatic Bioscience, Graduate School of Agricultural and Life Sciences, 3Asian Natural Environmental Science Center and 4Department of Global Agricultural Sciences, Graduate School of Agricultural and Life Sciences, The University of Tokyo, Bunkyo, Tokyo 113-8657, Japan

ABSTRACT: Between September 1996 and March 1999, a total of 135 fish species in 43 families were recorded from the mangrove estuary of Sikao Creek, Trang Province, Thailand, using two sizes of beach seine and a bag net. A checklist of the species is given, with preliminary descriptions of their assemblage structure. In terms of the number of species per family, Gobiidae was the most diverse (28 species), followed by Leiognathidae (11 species) and Engraulidae (10 species). In terms of individual numbers, Engraulidae, Leiognathidae and Ambassidae were the most dominant, whereby the 20 most abundant species comprised 88.5% of the total number of individuals collected. The fish assemblage structure was compared with published accounts of other tropical Indo-West Pacific mangrove estuaries, and found to be similar to those of tropical Australia. Although a grater number of species were recorded from Sikao Creek than in comparable studies in other geographic regions, all of the studies were similar in that they have relatively few species that are clearly dominant in abundance.

KEY WORDS: assemblage structure, ﬁsh, mangrove estuary, species list, Thailand.

INTRODUCTION presently subject to pressures resulting from industrial and aquacultural development, and Mangrove estuaries are considered to be important wood-chip production.4,5 Detailed knowledge of as nursery or feeding sites for many fish and prawn the role of mangrove areas for fishes is required for species, including those species of commercial the management of both the former and coastal importance. Previous studies in Australia have fisheries. However, studies on fish assemblages in shown that fish assemblages of mangrove areas mangrove areas in South-East Asia have been very differ from those in adjacent habitats, such as sea- limited to date.6–9 For example, Satapoomin and grass and sandflat, and that species’ richness and Poovachiranon have provided a check list of fishes fish abundance and biomass in mangrove areas are (including 232 species) collected from mangrove greater than those in adjacent habitats.1,2 Although areas along the west coast of Thailand, but with Robertson and Duke have reported a higher total only a brief description of their assemblage struc- species’ number in seagrass/sand habitats than in ture.9 Chua has listed 81 common fishes caught mangrove habitats, fish abundance was still much in the Ponggol estuary in Singapore, with a brief greater in the latter.3 Mangrove forests are a domi- account of those that are most predominant,6 and nant feature of South-East Asian coasts, but are Chong et al.7 and Pinto8 have reported relatively high species’ numbers from Malaysia (119) and the Philippines (128), respectively; however, only a list *Corresponding author: Tel: 81-3-5841-8115. Fax: 81-3-5841- of the 10 most abundant species was included in 5189. Email: [email protected] aPresent address: College of Bioresource Sciences, Nihon the former and an overall species’ list was omitted University, Kameino, Fujisawa, Kanagawa 252-8510, Japan. from the latter. Received 9 February 2001. Accepted 13 June 2001. The present paper provides a list of fishes col- Fish fauna of the mangrove estuary FISHERIES SCIENCE 11

lected from a mangrove estuary in Trang Province, Thailand, as well as preliminary descriptions of their assemblage structure. Similarities in assemblage structure with other Indo-West Paciﬁc mangrove estuaries are also discussed.

MATERIALS AND METHODS

The study site, Sikao Creek estuary (7∞32¢N, 99∞20¢E), which is a medium-sized (main channel length approximately 5 km) mangrove estuary, is located in Trang Province on the west coast of Thai- land (Fig. 1). The forest, which is dominated by Rhizophora apiculata, has been designated as a protected area and is relatively undisturbed. Inter- tidal mudflats with mangrove fringes and sandy- mud beaches lacking mangroves, but neither seagrass beds nor coral reefs, occur in the immedi- ate vicinity of the creek mouth. The estuary is sub- jected to semidiurnal tides with a tidal range of 1.0–2.5 m. Sampling was conducted approximately three times a year: March (dry season), August/ September (middle of wet season) and Novem- ber/December (end of wet season) at six fixed stations (stations 1–6) located from the mouth to the upper reaches of the creek (Fig. 1). Salinities at sampling varied from 10 (wet season) to 30 (dry season) in the upper reaches of the creek, but remained relatively stable at the creek mouth (28–32). Water temperatures varied from 26∞C to 34∞C, often being higher at the creek mouth than in the creek reaches, but no seasonal trends were apparent. Fishes were collected using a large and small seine, and a bag net. Size, sampling session and total number of operations for each net are shown in Table 1. The small seine net collected mainly Fig. 1 Map of Sikao Creek showing sampling sites. small-sized fishes, and the large seine net collected larger, more mobile fishes. Many small creek branches of less than a few meters width, from which the water drained completely at spring low juveniles of species otherwise identified on the tides, occurred throughout Sikao Creek. The bag basis of adult specimens. The taxonomic order net was used to catch fishes which entered such a follows Nelson.15 The number of individuals of branch at station 4, the net being set at the mouth each species was counted. As the main aim of the of the small branch at slack high tide, thereby present study was to describe the overall assem- catching the fishes which sought to leave the blage structure of the Sikao Creek estuary, varia- branch on the ebb tide. Captured fishes were pre- tions in the assemblage structure among seasons served in 10% formalin and identified to species’ or stations were not analyzed. level whenever possible, following mainly Kottelat et al.,10 Mohsin and Ambak,11 Munro,12 Nakabo,13 and Smith and Heemstra.14 A few species remained RESULTS unidentified and several juveniles could be identified only to family or genus level. Among the The list of fish species caught in the Sikao Creek unidentified juveniles, Scomberoides sp. and estuary, together with the number of individuals of Secutor sp. were not considered to be separate each species, is given in Table 2. A total of 135 species because they appeared most likely to be species in 43 families were recorded, of which 94 12 FISHERIES SCIENCE P Tongnunui et al.

Table 1 Sampling scheme and net dimensions Net type Small seine Large seine Bag net Wing length (m) 8 26 5 Mesh size of wing (mm) 3 20 5 Mesh size of cod end (mm) 3 13 3 Sampling periods Aug 1996–Mar 1999 Dec 1998–Aug 1999 Nov 1997–Mar 1999 Total no. sampling sessions 8 3 5 Sampling stations* 1,2,4 3–6 4† Total no. operations 96 64 5

*See Fig. 1. †Small branch (ª 3m wide) from the main creek at station 4.

Table 2 Number of individuals (ranks) of ﬁsh species collected from Sikao Creek estuary, Trang, Thailand Family Species SS LS BN Total (Rank) Dasyatidae Himantura imbricata 33 Elopidae Elopidae sp. 1 1 Engraulidae Encrasicholina devisi 393 1 394 (15) Stolephorus andhraensis 213 Stolephorus apiensis 314 Stolephorus chinensis 39 70 590 699 (10) Stolephorus commersonii 729 Stolephorus dubiosus 112 Stolephorus indicus 1 857 17 18 1 892 (5) Stolephorus insularis 3441158 Thryssa hamiltonii 40 878 420 1 338 (7) Thryssa setirostris 525 30 Pristigasteridae Ilisha melastoma 45 45 Clupeidae Anodontostoma chacunda 1321 34 Escualosa thoracata 17 485 502 (12) Sardinella sp. 40 40 Bagridae Mystus gulio 120 21 141 Synodontidae Trachinocephalus myops 33 Batrachoididae Allenbatrachus grunniens 10 10 Mugilidae Chelon subviridis 59 59 Ellochelon vaigiensis 68 68 Moolgarda engeli 18 16 1 35 Moolgarda seheli 67 67 Moolgarda parmatus 58 58 Atherinidae Atherinomorus duodecimalis 192 2 194 Hypoatherina valenciennei 1 584 760 208 2 552 (3) Phallostethidae Neostethus lankesteri 3 947 950 (8) Adrianichthyidae Oryzias javanicus 35 1 905 1 940 (4) Belonidae Strongylura strongylura 266 14 Tylosurus crocodilus crocodilus 73 10 Hemiramphidae Dermogenys sumatrana 22 22 Hyporhamphus afﬁnis 11 Hyporhamphus limbatus 436 34 2 472 (14) Zenarchopterus buffonis 86 133 63 282 (19) Zenarchopterus dunckeri 39 1 39 79 Scorpaenidae Vespicula trachinoides 16 21 37 Platycephalidae Inegocia japonica 11 Platycephalus indicus 22 Centropomidae Lates calcarifer 11 Ambassidae Ambassis interrupta 11 11 Ambassis macracanthus 14 14 Ambassis nalua 47 2 2 51 Ambassis urotaenia 116 17 Ambassis vachellii 1 290 1 881 649 3 820 (1) Fish fauna of the mangrove estuary FISHERIES SCIENCE 13

Table 2 Continued Family Species SS LS BN Total (Rank) Apogonidae Apogon hyalosoma 810 18 Apogon albomarginatus 88 Serranidae Epinephelus coioides 11 Sillaginidae Sillago aeolus 33 Sillago intermedius 12 12 Sillago sihama 451 44 1 496 (13) Carangidae Carangoides praeustus 24 6 Caranx ignobilis 11 Caranx sexfasciatus 66 Scomberoides commersonnianus 77 Scomberoides lysan 11 2 Scomberoides sp. 3 3 Carangidae sp. 3 6 9 Leiognathidae Gazza dentex 20 17 1 38 Leiognathus decorus 360 926 231 1 517 (6) Leiognathus equulus 86 631 7 724 (9) Leiognathus leuciscus 11 Leiognathus pan 14 14 Leiognathus rapsoni 22 Leiognathus splendens 33 272 305 (17) Leiognathus stercorarius 138 39 Secutor hanedai 34 34 Secutor insidiator 35 35 Secutor megalolepis 944 1 711 50 2 705 (2) Secutor sp. 2 2 Lutjanidae Lutjanus johnii 22 Lutjanus russellii 14 12 4 30 Gerreidae Gerres erythrourus 689 5 3 697 (11) Gerres ﬁlamentosus 20 80 4 104 Gerres limbatus 31 4 Gerres macracanthus 195 69 264 (20) Haemulidae Plectorhinchus gibbosus 11 Pomadasys kaakan 7182 27 Sciaenidae Dendrophysa russelli 11 11 Pennahia anea 226 226 Mullidae Upeneus sulphureus 29 29 Upeneus sundaicus 12 12 Upeneus tragula 48 12 Toxotidae Toxotes chatareus 22 Toxotes jaculatrix 11 Teraponidae Terapon theraps 77 Blenniidae Omobranchinii sp. 2 1 3 Callionymidae Repomucenus hindsii 55 Repomucenus schaapi 33 Eleotoridae Butis butis 15 6 65 86 Ophiocara porocephala 11 Gobiidae Acentrogobius caninus 10 6 16 Acentrogobius kranjiensis 342 1 343 (16) Acentrogobius malayanus 118 3 121 Acentrogobius viridipunctatus 52 44 3 99 Amoya moloanus 50 1 1 52 Boleophthalmus boddarti 11 2 Cristatogobius sp. 2 2 Glossogobius bichirrhosus 77 Glossogobius biocellatus 20 14 34 Glossogobius circumspectus 33 Glossogobius giuris 66 Glossogobius sparsipapillus 66 Glossogobius sp. 22 22 14 FISHERIES SCIENCE P Tongnunui et al.

Table 2 Continued Family Species SS LS BN Total (Rank) Gnatholepis baliurus 539 44 Gobiopterus brachypterus 13 13 Oxyurichthys microlepis 29 49 78 Pandaka lidwilli 235 Papillogobius melanobranchus 16 16 Papillogobius reichei 76 3 79 Pseudogobius javanicus 52 52 Pseudogobius sp. 9 9 Sillhouettea evanida 18 18 Stigmatogobius sadanundio 15657 Gobiidae sp. 1 1 1 Gobiidae sp. 2 1 1 Gobiidae sp. 3 1 1 Gobiidae sp. 4 1 1 Gobiidae sp. 5 8 1 9 Scatophagidae Scatophagus argus 3381 42 Siganidae Siganus canaliculatus 14 14 Siganus guttatus 11 Siganus javus 59 5 3 67 Siganus sp. 1 2 2 Siganus sp. 2 1 1 Sphyraenidae Sphyraena barracuda 22 Sphyraena jello 11 Sphyraena putnamae 453 12 Scombridae Rastrelliger sp. 2 287 3 292 (18) Paralichthyidae Pseudorhombus arsius 11 2 Cynoglossidae Cynoglossus puncticeps 33 Triacanthidae Triacanthus biaculeatus 13 4 Monacanthidae Monacanthus chinensis 11 Tetraodontidae Arothron reticularis 11 Chelonodon patoca 33 Lagocephalus lunaris 11 27 38 Tetraodon leiurus 161 8 Tetraodontidae sp. 1 1 Total no. families 32 35 25 43 Total no. species 94 90 50 135 Total no. specimens 10 121 9 571 5 380 25 072

SS, small seine; LS, large seine; BN, bag net. species in 32 families were caught by the small lolepis (Leiognathidae) and Gerres erythrourus seine, 90 species in 35 families by the large seine (Gerreidae); those by the large seine being A. and 50 species in 25 families by the bag net. In vachellii, S. megalolepis, Leiognathus decorus terms of the number of species per family, (Leiognathidae), Thryssa hamiltonii (Engraulidae), Gobiidae was by far the most diverse (28 species), H. valenciennei, L. equulus (Leiognathidae) the next speciose families being Leiognathidae and Escualosa thoracata (Clupeidae). Fresh and (11 species), Engraulidae (10 species), Carangidae brackish water species Oryzias javanicus (six species), Mugilidae, Hemiramphidae, (Adrianichthyidae) and Neostethus lankesteri Ambassidae, Siganidae and Tetraodontidae (ﬁve (Phallostethidae) dominated the bag net catch, fol- species each). lowed by A. vachellii, Stolephorus chinensis In terms of individual numbers, Leiognathidae, (Engraulidae) and T. hamiltonii. Of the 20 most Engraulidae, Ambassidae, Atherinidae and abundant species overall, four each belonged to Adrianichthyidae were dominant (Table 2). The Engraulidae and Leiognathidae (Table 2). These most abundant species collected using the small same 20 species also comprised 88.5% of the total seine were Stolephorus indicus (Engraulidae), number of individuals, the least abundant 100 Hypoatherina valenciennei (Atherinidae), species together making up less than 6% of the Ambassis vachellii (Ambassidae), Secutor mega- total number. Fish fauna of the mangrove estuary FISHERIES SCIENCE 15

DISCUSSION Relative abundance of each family was compared for the catch by seine nets. For the comparison The species’ richness of a mangrove system family names were revised with those presented in depends upon the size of the system, in addition to Nelson.15 The species’ richness for the most spe- the diversity of habitats, proximity to other systems ciose families (more than four species) in this and nature of the offshore environment.16 The and comparable studies is listed in Table 4. The total number of species (135) recorded during numbers of species in the families Leiognathidae, the present study was greater than the highest Ambassidae and Tetraodontidae were highest in numbers recorded previously in other mangrove the Sikao Creek estuary, followed by Australian systems of similar size (e.g. 128 species in both Alli- mangrove estuaries, and were lowest in African gator Creek, Australia17 and Pagbilao, Philippines.8 mangrove estuaries. Conversely, Synodontidae, See also Robertson and Blaber16 for review). Of the Lutjanidae, Lethrinidae and Mullidae were more species collected in the present study, at least 43 diverse in African (Kenyan) mangrove estuaries, were not listed by Satapoomin and Povachiranon,9 most of the component species being reef associ- who reported 232 species from mangrove areas ates; hence, their high species’ numbers may along the west coast of Thailand. In all, a total of reflect the proximity of coral reefs.19,20 The study 275 species are now recognized from mangrove with the greatest number of species in common areas in that region. This high species number with the present study was at Embley, Australia (35) when compared to other mangrove systems may, followed by Trinity, Australia and Mombasa, Kenya at least in part, reflect the rich shore fish diversity (16). of the East Indies, being the richest in the world.18 In terms of abundance of the fish collected using In the present study, although a high number of seine nets, Leiognathidae was the dominant family species were found, the assemblage was domi- in both the present study and that at Trinity, Aus- nated by relatively few species. This is consistent tralia (Fig. 2). A predominance of leiognathids has with the findings of similar studies in other geo- also been found in trawl samples taken in man- graphic regions.3,19,20 grove estuaries in the Philippines8 and Papua New To compare the assemblage structure of fishes Guinea.23 In contrast, gerreids and plotosids in the Sikao Creek estuary with that in other tropi- dominated the African mangrove estuaries (Fig. 2). cal Indo-Pacific mangrove estuaries, recourse was Although Robertson and Duke collected fishes in made to publications that included a full species’ four tropical Australian mangrove creeks using list and abundance of each species (number of sampling gears comparable with those used in the individuals, % composition of catch, or catch per present study (a small seine with 3 mm mesh and unit effort based on number of individuals). To a larger seine with 18 mm mesh), they listed only minimize bias arising from differences in sampling the 30 most abundant species collected from man- gear and habitat, catches from sites similar to the grove, seagrass and sand habitats.3 Nevertheless, Sikao Creek estuary, using comparable sampling the fish assemblage structure of the mangrove gear, were selected and species’ number and rela- habitat in their study appears to be similar to that tive abundance of each family noted. These in the present study. About half (13) of the species selected habitats and gear are listed in Table 3. listed by Robertson and Duke3 are also recorded in

Table 3 Location, sites and gear for the comparison of the fish assemblage structure in Sikao Creek estuary with that in other tropical Indo-Pacific mangrove estuaries Net* Location Site* Type Size (m) Mesh (mm) Reference Embley, Australia† Sandy mud beaches Seine 60 ¥ 2 25 Blaber et al.21 Mud flats Stake 240 ¥ 250 Creeks and inlets Block NS 2 Trinity, Australia Estuary Seine 75 ¥ 1.5 25 Blaber22 Estuary Stake† 300 ¥ 2.5 25 Mombasa, Kenya Creeks Seine 15 ¥ 2 6 Little et al.19 Gazi, Kenya Mangrove fringed sites Seine 45 ¥ 2 20 Kimani et al.20

*Comparable sites and gear with the present study were selected from the literature. †Only species composition was compared. NS, not shown. 16 FISHERIES SCIENCE P Tongnunui et al.

Table 4 Species’ richness of speciose families in Trang, the present study, numerically dominant families Thailand and other tropical estuaries of the Indo-West in the former study being Leiognathidae, Engraul- Pacific idae, Atherinidae, Gobiidae, Clupeidae, Chandidae Locality and Pseudomugilidae. Most of the 20 dominant Families TT EA TA MK GK species in the present study are distributed widely, at least from the eastern Indian Ocean to the West Dasyatidae 1 6 Pacific, with approximately half being abundant Engraulidae 10 4 3 5 2 3,16 Clupeidae 3 6 4 2 3 also in Australian mangrove estuaries. Thus, the Synodontidae 1 4 5 assemblage structure and dominant species in the Mugilidae 5 5 2 1 3 Sikao Creek estuary are comparable with those of Hemiramphidae 5 4 2 3 2 tropical Australian mangrove estuaries. Ambassidae 5 3 1 1 Sillaginidae 3 5 2 1 1 Carangidae 6 3 3 5 3 ACKNOWLEDGMENTS Leiognathidae 11 9 7 3 1 Lutjanidae 2 2 2 1 6 We are grateful to Y Fukuyo, K Kojima, N Gerreidae 4 5 3 3 3 Paphavasit, A Piumsomboon and P Wongrat for Lethrinidae 1 2 5 Mullidae 3 1 1 2 6 their encouragement during this study. We thank Y Scaridae 5 Iwatsuki, S Kimura, K Matsuura, T Nakabo, H Gobiidae 28 12 4 11 2 Senou, K Shibukawa and T Wongratana for their Siganidae 5 1 3 help in identifying some fishes. Thanks are also due Tetraodontidae 5 4 3 2 2 to O Baba, S Janekitkarn, K Kanou, H Kohno, K Others 38 49 10 38 42 Okamoto, and staff and students at the Faculty of Total no. species 135 118 49 84 94 Science and Fisheries Technology, Rajamangala Total no. families 43 48 26 37 44 Institute of Technology, especially T Nitiratsuwan, No. species in – 35 16 16 11 C Sudtongkong, S Tanyaros and P Wirachwong for common with TT their help in field sampling. Comments on the No. families in – 32 22 26 25 common with TT manuscript from G Hardy were very helpful. We are grateful to the National Research Council of TT, Trang, Thailand; EA, Embley, Australia (Blaber et al.21); TA, Thailand for the granting of permits to conduct Trinity Australia (Blaber22); MK, Mombasa, Kenya (Little et al.19); this research in Thailand. This study was supported GK, Gazi, Kenya (Kimani et al.20). in part by Grants-in-Aid for Scientific Research, Creative Basic Research and International Scien- tific Research, from the Ministry of Education, Science, Sports and Culture of Japan (Nos. 07- 08301, 09-NP0901 and 10041160).

REFERENCES

1. Morton RM. Community structure, density and standing crop of fishes in a subtropical Australian mangrove area. Mar. Biol. 1990; 105: 385–394. 2. Laegdsgaard P, Johnson CR. Mangrove habitats as nurseries: Unique assemblages of juvenile fish in subtropical mangroves in eastern Australia. Mar. Ecol. Prog. Ser. 1995; 126: 67–81. 3. Robertson AI, Duke NC. Mangroves as nursery sites: Com- parisons of the abundance and species composition of fish and crustaceans in mangroves and other nearshore habitats in tropical Australia. Mar. Biol. 1987; 96: 193–205. 4. Fortes MD. Mangrove and seagrass beds of East Asia: Habitats under stress. Ambio 1988; 17: 207–213. 5. Ong JE. The ecology of mangrove conservation and man- Fig. 2 Relative abundance of the dominant fish families agement. Hydrobiologia 1995; 295: 343–351. collected using seine nets in Trang, Thailand (TT) and 6. Chua T-E. An ecological study of the Ponggol estuary in other tropical estuaries in the Indo-West Pacific. TA, Singapore. Hydrobiologia 1973; 43: 505–533. Trinity, Australia; MK, Mombasa, Kenya; GK, Gazi, 7. Chong VC, Sasekumar A, Leh MUC, D’Cruz R. The fish and Kenya. prawn communities of a Malaysian coastal mangrove Fish fauna of the mangrove estuary FISHERIES SCIENCE 17

system, with comparisons to adjacent mud flats and Mangrove Ecosystems. American Geophysical Union, Wash- inshore waters. Estuar. Coast. Shelf Sci. 1990; 31: 703–722. ington, D.C. 1992; 173–224. 8. Pinto L. Population dynamics and community structure of 17. Robertson AI, Duke NC. Mangrove fish-communities in fish in the mangroves of Pagbilao, Philippines. J. Fish Biol. tropical Queensland, Australia: Spatial and temporal pat- 1988; 33 (Suppl. A): 35–43. terns in densities, biomass and community structure. Mar. 9. Satapoomin U, Poovachiranon S. Fish fauna of mangroves Biol. 1990; 104: 369–379. and seagrass beds in the west coast of Thailand, the 18. Randall JE. Zoogeography of shore fishes of the Indo-Pacific Andaman Sea. Phuket Mar. Biol. Center Tech. Paper 1997 region. Zool. Stud. 1998; 37: 227–268. 1997; No. 2: 1–63. 19. Little MC, Reay PJ, Grove SJ. The fish community of an East 10. Kottelat M, Whitten AJ, Kartikasari SN, Wirjoatmodjo S. African mangrove creek. J. Fish Biol. 1988; 32: 729–747. Freshwater Fishes of Western Indonesia and Sulawesi. 20. Kimani EN, Mwatha GK, Wakwabi EO, Ntiba JM, Okoth BK. Periplus Editions (HK), Jakarta. 1993. Fishes of a shallow tropical mangrove estuary, Gazi, Kenya. 11. Mohsin AKM, Ambak MA. Marine Fishes and Fisheries of Mar. Freshwater Res. 1996; 47: 857–868. Malaysia and Neighboring Countries. Universiti Pertanian 21. Blaber SJM, Brewer DT, Salini JP. Species composition and Malaysia Press, Serdang. 1996. biomasses of fishes in different habitats of a tropical north- 12. Munro ISR. The Fishes of New Guinea. Department of ern Australian estuary: Their occurrence in the adjoining Agriculture, Stock and Fisheries, Port Moresby. 1967. seas and estuarine dependence. Estuar. Coast. Shelf Sci. 13. Nakabo T (ed). Fishes of Japan with Pictorial Keys to the 1989; 29: 509–531. Species, 2nd edn. Tokai University Press, Tokyo. 2000. 22. Blaber SJM. Fish of the Trinity Inlet system of North 14. Smith MM, Heemstra PC (eds). Smiths’ Sea Fishes. Springer- Queensland with notes on the ecology of fish faunas of trop- Verlag, Berlin. 1986. ical Indo-Pacific estuaries. Aust. J. Mar.Freshwater Res. 1980; 15. Nelson JS. Fishes of the World, 3rd edn. John Wiley & Sons, 31: 137–146. New York. 1994. 23. Quinn NJ, Kojis BL. Annual variation in the nocturnal 16. Robertson AI, Blaber SJM. Plankton, epibenthos and fish nekton assemblage of a tropical estuary. Estuar. Coast. Shelf communities. In: Robertson, AI, Alongi, DM (eds). Tropical Sci. 1986; 22: 63–90.