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Marine

Freshwater

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Volume 53, 2002

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A journal for the publication of original contributions

in physical oceanography, marine chemistry,

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© CSIRO 2002 10.1071/MF01136 1323-1650/02/020419

Spatial and temporal variations of the estuarine larval fish community

on the west coast of Taiwan

Wann-Nian Tzeng, Yu-Tzu Wang, Chih-Wei Chang

Department of Zoology, College of Science, National Taiwan University, Taipei, Taiwan 106, ROC

email: wnt@ccms.ntu.edu.tw

MF01136 Variations in an estuarine larval fish community Wann-Nian Tzeng

etal.W.-N.TzengY.-T.Wang, C.-W.Chang

Abstract.

Spatio–temporal variations in the distribution and community structure of the estuarine larval fishes on

the west coast of Taiwan was examined in the estuaries of Shuangchi Creek (SC), Gongshytyan Creek (GST), Tatu

Creek (TT) and Tongkang River (TK). Fish were collected by a net (mesh 0.8–1.8 mm) set against the flood tide

at night during the new- and full-moon periods from September 1997 through December 1998; 28–49 families (56–94

species) were collected from the four estuaries. Fish larvae were abundant from spring to autumn. Fish communities

differed among estuaries: Mugilidae were the most abundant in SC, Terapon jarbua in GST, Stolephorus insularis

in TT and Ambassis urotaenia in TK. The 15 most dominant species constituted 88–94% of the total catch. The

relationship of fish abundance and species diversity to water temperature and salinity differed among estuaries.

Species composition could be classified into northern (SC, GST winter and TT winter) and southern (TK, GST

summer and TT summer) groups. The species composition of the larval fish communities was more diverse in

spring–autumn than in winter, and in southern than in northern estuaries. Monsoon-driven coastal currents may

influence seasonal dispersal and community structure of the estuarine larval fishes on the western coast of Taiwan.

Extra keywords: larval dispersal, community structure, seasonal variation, monsoon, coastal currents, estuary

Introduction

Estuaries are among the most productive ecosystems on the

earth, comparable to areas of coastal upwelling (Haedrich

and Hall 1976). They function as nursery grounds for

estuarine-dependent marine fish populations during their

early life stages because it provides food resources, shelter,

absence of turbulence, and a reduced risk of predation

(Wallace and van der Elst 1975; Blaber and Blaber 1980;

Day et al. 1981; Lenanton 1982; Robertson and Duke 1987;

Blaber and Milton 1990). The species composition and

abundance of fish larvae and juveniles in the estuary are

influenced by physico–chemical and biological factors and

are important indicators for predicting forthcoming fishing

stocks (Haines 1979; Blaber et al. 1985; Stephens et al.

1986, 1988; Tzeng and Wang 1993; Wang and Tzeng

1997).

Studies of the larval and juvenile fish communities in the

estuaries of Taiwan have been limited to specific areas where

environmental protection has been a concern, e.g. the

sewage-polluted Tanshui River estuary (Tzeng and Wang

1992, 1993, 1997) and Yenliao Bay near a nuclear power

plant under construction (Tzeng et al. 1985, 1997; Tzeng and

Wang 1986). Thus, large-scale spatio–temporal variations of

larval fish communities among estuaries in Taiwan are not

clear. This study investigated the larval fish community

structure in estuaries on the west coast of Taiwan with a view

to understanding the effect of coastal currents on the

dispersal of fish larvae and juveniles.

Materials and methods

Study area and sampling design

The coastal waters of western Taiwan comprise the Taiwan Strait, which has a wide and shallow continental shelf, suitable as a spawning and nursery area for coastal fishes. In contrast, the continental shelf on the eastern coast is narrow and strongly influenced by the oceanic Kuroshio Current (Fig. 1). The coastal currents on the western coast that transport larvae to estuaries change direction with the seasonal monsoon (Chu 1963). In winter, the China Coastal Current, induced by the eastern monsoon, flows from the coast of mainland China to the north-east and north-west coasts of Taiwan. In summer, this current is replaced by the south-western monsoon-driven coastal current, which flows from the South China Sea along the western coast of Taiwan.

A net was set against the tidal current in four estuaries, Shuangchi Creek (SC), Gongshytyan Creek (GST), Tatu Creek (TT) and Tongkang River (TK) (Fig. 1), to collect fish larvae and juveniles during the nighttime flood tide around the new- and full-moon periods from September 1997 to December 1998. The mesh size of the net ranged from 0.8 to 1.8 mm, similar to the commercial fishing gear used for collecting fish fry for restocking in Taiwan. The fish were preserved in

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95% alcohol after collection and identified to species level when possible (Leis and Rennis 1983; Ozawa 1986; Wang 1987; Okiyama 1988; Leis and Trnski 1989). Surface water temperature and salinity were measured to 0.1°C and 0.1 psu by use of a microprocessor conductivity meter during sampling.

Data analysis

The abundance of fish larvae and juveniles was estimated roughly by the number of fish per hour (CPUE). The cumulative abundance of dominant species among the four estuaries was compared by K-dominance curves. When two different K-dominance curves do not overlap, the upper curve represents the community most dominated by a few species and thus that which is less diverse (Lambshead et al. 1983; Clarke and Warwick 1994). The species diversity of each larval fish community was calculated by use of the Shannon–Wiener index of species diversity (H′) (Pielou 1966) and Simpson’s index of species concentration (Σpi2) (Peet 1974). The

Shannon–Wiener index emphasizes the contribution of rare species, whereas Simpson’s index gives greater weight to dominant species. The similarity of species compositions of fish larvae and juveniles in the four estuaries among sampling dates was analysed by clustering (Bray–Curtis similarity index; unweighted pair-group using arithmetic average) and ordination (multiple dimensional scaling, MDS). Indicator species for each of the seasonal and geographic groups and the species that discriminated groups were examined by the similarity percentage routine (SIMPER; Clarke and Warwick 1994). The data used for clustering and ordination were log (n+1) transformed and selected to span one year from December 1997 through November 1998 because the period investigated differed among estuaries. All computation of diversity indices and cluster and ordination analyses used the software ‘PRIMER 5’ (Clarke and Warwick 1994). Relationships between abiotic and biotic factors of the four estuaries were analysed by Kendall rank correlations (Siegel 1956).

Results

Water temperature and salinity

Trends in water temperature were similar among the GST, SC

and TT estuaries, but there was much less seasonal variation

in the TK estuary where winter water temperatures did not

decline to the same degree as those in the other systems. The

water temperatures in the four estuaries ranged from 14.0 to

33.0°C. Annual mean (± s.d.) water temperatures increased

from 23.2±3.9°C in SC in the north to 29.1±2.5°C in TK in

the south (Fig. 2a). Salinities in the four estuaries varied from

0.3 to 36.5 psu; the annual mean (± s.d.) salinity ranged from

17.5±10.3 in TT to 29.6±7.2 in TK (Fig. 2b). A geographic

cline in mean salinity was not obvious among estuaries;

however, the maximum value of the range was higher in both

SC and TK than in either GST or TT, because SC and TK were

influenced to a greater degree by the highly saline Kuroshio

Current than were GST and TT (Fig. 1).

Seasonal changes in number of species, abundance and

indices of community structure

In total, 30, 33, 26 and 26 samples were collected from the

SC, GST, TT and TK estuaries, respectively: 28 families and

56 species from SC, 49 families and 94 species from GST, 44

families and 73 species from TT and 46 families and 77

species from TK. The dominant species differed among

estuaries. Mugilidae were the most abundant in SC, Terapon

jarbua in GST, Stolephorus insularis in TT and Ambassis

urotaenia in TK (Appendix 1).

The number of species increased from spring to autumn

(April to September) in GST estuary, but no seasonal trend

was apparent in the other three (Fig. 2c). Marked seasonal

variation in abundance (CPUE) was noted in all four estuaries.

Two periods of peak abundance occurred in the northern

estuaries (SC and GST), one in the spring (March–June) and

the other in the autumn (September), but the autumn peak did

not occur in the southern estuaries (TT and TK). The peak

abundance in spring occurred earlier in the south than in the

north (Fig. 2d). Species diversity (H′, 0.2–2.7) and

concentration indices (Σp

i

2

, 0.1–1.0) of the larval fish

community greatly fluctuated with season and were inversely

correlated (Figs 2e, 2f). This indicates that the larval fish

community became uneven when dominant species occurred.

Fish abundance was significantly positively correlated to the

number of species except in SC, and was positively correlated

with Σp

i

2

only in TK (Table 1).

The effect of temperature and salinity on the number of

species, abundance and community indices differed among

estuaries (Table 1). In SC, there was no significant correlation

of the four biotic factors with temperature or salinity. However

in GST, number of species, abundance and species diversity

(H′) were positively correlated with temperature (P <0.05),

but not with salinity. In TT, the species diversity (H′) and

concentration indices (Σp

i

2

) were correlated with salinity

Fig. 1. Sampling sites of fish larvae and juveniles in the estuaries of Shuangchi Creek (SC), Gongshytyan Creek (GST), Tatu Creek (TT) and Tongkang River (TK). Dashed line: 200 m depth contour.

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(positively and negatively, respectively; P <0.05). In TK, only

abundance was positively correlated with temperature

(P <0.05). This may be due to different geomorphology and

hydrodynamics among estuaries.

Spatio–temporal similarity of species composition

Cluster and ordination analyses indicated that the species

composition of all estuaries except SC comprised two

seasonal groups at a similarity level of ~20% (Figs 3a–d).

These two seasonal groups were a spring–autumn group

(April–November) and a winter group (December–March).

When the species data from the four estuaries were pooled

together and analysed, the winter groups of both GST and

TT clustered with SC into a northern group, while the

summer groups of both GST and TT were more similar to

TK samples and formed a southern group at a similarity

level of ~10–15% (Fig. 3e). This indicates that the

summer larval fish communities of GST and TT, as well as

that of TK throughout the year, may originate from

southern Taiwan, whereas the winter fish communities of

GST and TT, together with that of SC, derive from

northern Taiwan.

K-dominance curves showed higher cumulative

percentage abundance in winter than in spring–autumn for

both GST and TT estuaries, indicating that the communities

of winter groups were less diverse than spring–autumn

groups and were characterized by few dominant species

(Fig. 4a). Similarly, the northern group was characterized by

fewer dominant species (Fig. 4b).

Similarity percentage of species contribution to clustering

Species contributing to the similarities within seasonal

groups shown in Figs 3a-d and to the southern and northern

groups in Fig. 3e are listed in Tables 2 and 3. The larval

Fig. 2. Seasonal changes in (a) temperature, (b) salinity, (c) number of species, (d) abundance, (e) index of species diversity and (f) index of species concentration in the four estuaries (SC, GST, TT and TK refer to Fig. 1).

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fish community of the SC estuary and the winter

assemblages of the GST and TT estuaries were dominated

by few species. The average percentage contribution of

only five species in these groups ranged from 81.3% to

94.0%. Mugilidae spp. and Mugil cephalus were the most

important, contributing 33.4–36.8% of total average

similarity (Table 2). In contrast, the communities of the TK

estuary together with the spring–autumn groups of GST and

TT were more diverse and the average similarity

contributed by the five most important species ranged from

38.3% to 70.1% (Table 2).

Similarly, the larval fish community was more diverse in

the southern than in the northern groups (Table 3). The

average similarity of the top 10 species in the northern group

was 89.7%. Mugilidae spp., Mugil cephalus and Sicyopterus

japonicus were the most important taxa contributing to this

group similarity (67.2%). In the southern group ten species

made relatively even contributions to account for only 63.1%

of the group similarity. Species accounting for the

dissimilarity between these two groups are also listed in

Table 3 and comprised many minor species because the sum

of the average dissimilarity for the top 10 species was only

37.8% (Table 3).

Discussion

An estuary is a semi-enclosed system influenced by both

fresh water and seawater, and freshwater, estuarine and

marine fishes might be expected in such a system (Blaber

and Blaber 1980; Day et al. 1981). However, most larval

fishes collected in this study were estuarine-dependent

marine fishes (e.g. Mugilidae spp., Mugil cephalus and

Terapon jarbua) and estuarine fishes (e.g. Gobiidae spp. and

Ambassis spp.). Few freshwater fishes (e.g. Cyprinidae)

were caught (Tables 2 and 3; Appendix 1). This is because

we collected fishes on flood tides that transported the marine

fishes to the estuaries. The dispersal of larval fishes from

marine spawning grounds to the estuaries is influenced by

coastal currents (Boehlert and Mundy 1988). The coastal

currents on the west coast of Taiwan change direction with

season (Chu 1963). Thus the recruitment of larval fishes to

the estuary will be influenced in spatial and temporal terms

by these coastal currents.

Larval fish communities in three of the four estuaries

were classified into winter and spring–autumn seasonal

groups, which reflected the change of seasonally

monsoon-driven coastal currents on the western coast of Taiwan (Chu

1963). In winter, the north-eastern monsoon-driven,

low-salinity, cold China Coastal Current flows southward and is

impeded by the warm Kuroshio Current in the middle of the

Taiwan Strait. In summer, the south-western

monsoon-driven, high salinity, warm South China Sea surface current

flows through the Taiwan Strait (Jan 1995; Shao et al. 1997).

Most of the larvae in the four estuaries were

estuarine-dependent marine fishes (Wallace and van der Elst 1975;

Blaber and Blaber 1980). In general, larvae of these fishes

passively migrate with the coastal currents from spawning

grounds to estuaries (Boehlert and Mundy 1988). Thus, the

monsoon-driven coastal current on the west coast of Taiwan

may play an important role in the seasonal onshore

movement of fish larvae and juveniles.

The fish communities of the SC estuary and the winter

assemblages of both the GST and TT estuaries clustered

together into a similar group, which was distinct from that

of the TK estuary and the summer assemblages of the GST

and TT estuaries. This suggests that the winter larval fish

community transported from the north by the NE

monsoon-driven China Coastal Current is distributed only as far as

the middle part of the Taiwan Strait. In contrast, the

summer larval fish community transported by the SW

monsoon current from the south reaches the northern part

of the Taiwan Strait. Shao et al. (1997) noted that the fish

fauna differed with latitude on the west coast of Taiwan

under the influence of the seasonal monsoons, and that the

boundary separating northern and southern fish faunas may

be located approximately at Penghu Island in the middle of

Taiwan Strait. Kuo et al. (1999) found that mangrove fish

assemblages on the west coast could be classified into

Table 1. Relationship between abiotic and biotic factors of four estuaries

SC, GST, TT and TK refer to Fig. 1. Temp, temperature; Sal, salinity; Sp, number of species; Ab, abundance; H′, index of species diversity;

Σpi

2, index of species concentration

Temp Sal Sp Ab HSC Sal 0.42* Sp –0.03 –0.26 Ab –0.21 0.12 0.18 H′ 0.26 –0.12 0.66*** –0.14 Σpi 2 –0.32 0.08 –0.53*** 0.25 –0.86*** GST Sal 0.16 Sp 0.77*** 0.08 Ab 0.62*** 0.22 0.59*** H′ 0.27* –0.05 0.31* 0.07 Σ pi2 –0.17 0.09 –0.22 –0.04 –0.84*** TT Sal –0.26 Sp 0.12 0.08 Ab 0.25 –0.12 0.47*** H′ 0.03 0.35* 0.39** –0.05 Σ pi2 –0.07 –0.34* –0.27 0.11 –0.85*** TK Sal –0.03 Sp 0.26 0.02 Ab 0.36* 0.04 0.49*** H′ –0.17 0.17 –0.03 –0.41** Σ pi2 0.17 –0.10 0.07 0.40** –0.83***

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Fig . 3. C lu st e r de nd ro g ra m s (upp er ) an d MD S or di na ti o n s (l o w er ), ba se d o n Br ay – Cu rtis simi la rit ie s o f la rv al a n d ju v en ile f ish c o mmu n it ie s sa mp le d in (a ) SC , (b ) GS T , (c ) TT , (d ) TK a nd (e ) o v er al l d a ta of a-d. N u m e ra ls 1 – 12 i n d ic at e t h e o rd er o f t h e m ont hs fr om J an u ar y t o D e ce m b e r. w : w int e r, s-a: sp ri ng -a ut um n .

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northern and southern groups, which probably arise as a

result of the oceanic currents around Taiwan. This study

indicated that the spatial distribution of the larval fish

community on the west coast of Taiwan changed seasonally.

In the summer, the SW monsoon current transported fish

larvae through the Taiwan Strait. In winter, the dispersal of

larvae transported by the NE monsoon current was impeded

in the middle of the Taiwan Strait by the Kuroshio Current.

The communities of the four estuaries exhibited high

spatial and temporal variations in species diversity. Fish

communities of spring–autumn groups originating in the

south were more diverse than were the winter groups from the

north. Pianka (1966, 1967) proposed that species diversity

was influenced by eight principal factors, i.e. evolutionary

time, ecological time, climatic stability, spatial heterogeneity,

productivity, stability of primary production, competition

and predation. Estuaries are highly variable and unstable

environments (Whitfield 1990), and fish communities living

in such habitats theoretically should reach an equilibrium

state and contain fewer species when interspecific

competition is prevalent (Sanders 1968; de Morais and de

Morais 1994; Harris and Cyrus 2000). The salinities of the

four estuaries fluctuated greatly, with a range of 0.3–36.5 psu.

Highly variable salinities may influence species diversity and

led to dominance of estuarine larval fish communities by a

few species. In addition, the water mass of the coastal current

from the north was more productive than that from the south.

This may lead to the fish communities originating from the

northern group being less diverse and dominated by fewer

species than fish communities originating from the south.

Furthermore, the differences in species composition among

Fig. 4. K-dominance curves for the numbers of species of fish larvae and juveniles of (a) seasonal groups [except SC, which represents all data] and (b) geographic groups (SC, GST, TT and TK refer to Fig. 1).

Table 2. Average similarity percentage contributed by the major species in the seasonal groups of four estuaries

SC, GST, TT and TK refer to Fig. 1

Rank SC % GST % TT % TK %

Whole year Winter group Winter group Winter group

1 Mugilidae spp.A 33.4 Mugil cephalus 34.3 Mugilidae spp.A 36.8 Acanthopagrus schlegeli 16.5

2 Sicyopterus japonicus 18.5 Anguilla japonica 21.3 Sardinella spp. 19.7 Ambassis urotaenia 16.3

3 Ambassis gymnocephalus 14.0 Acanthopagrus latus 17.2 Acanthopagrus spp. 18.0 Mugilidae spp.A 11.1

4 Mugil cephalus 10.6 Mugilidae spp.A 10.2 Acanthopagrus latus 17.0 Gobiidae spp. 10.6

5 Leiognathus nuchalis 4.8 Anguilla marmorata 2.9 Stolephorus insularis 2.5 Omobranshus spp. 8.6

6 Other (51 species) 18.7 Other (27 species) 14.1 Other (24 species) 6.0 Other (39 species) 37.0

Spring–autumn group

Spring–autumn group

Spring–autumn group

1 Terapon jarbua 11.0 Stolephorus insularis 38.8 Gobiidae sp.7 15.7

2 Gobiidae sp.9 7.9 Sillago sihama 9.2 Taenioides cirratus 12.8

3 Gobiidae sp.7 7.5 Gobiidae spp. 7.6 Megalops cyprinoides 8.6

4 Megalops cyprinoides 6.5 Ambassis

gymnocephalus

7.2 Gobiidae spp. 7.7

5 Rhinogobius giurinus 5.4 Mugilidae spp. A 7.2 Rhinogobius giurinus 6.3

6 Other (91 species) 61.7 Other (61 species) 29.9 Other (67 species) 48.8

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the four estuaries were not only due to the influence of these

two current systems on the recruitment of larval marine

fishes. Spatial heterogeneity within the estuaries that

provided a diversity of habitat for the estuarine fish species

probably also played a role.

Acknowledgments

The study was financially supported by the Council of

Agriculture (87-AST–1.4-FID–05(09), 88-AST–1.4-FID–

05(18) and 89-AST–1.2-FID–02(06)) and National Science

Council (M–002–006 and

NSC89–2611-M–002–039) of the Republic of China. The authors are

grateful to Messrs C. S. Jean, C. S. Lu, C. S. Chen and

C. L. Chen for collection of specimens, Mrs H. Y. Teng for

specimen sorting, and Mr Brian Jessop for helpful

comments.

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Table 3. Average similarity and dissimilarity percentages contributed by the major species within and between groups

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2 Mugil cephalus 16.9 Megalops cyprinoides 8.2 Stolephorus insularis 4.2

3 Sicyopterus japonicus 11.9 Gobiidae sp. 7. 7.5 Gobiidae spp. 4.0

4 Ambassis gymnocephalus 6.1 Rhinogobius giurinus 6.7 Gobiidae sp. 7. 3.9

5 Acanthopagrus latus 4.8 Terapon jarbua 6.3 Rhinogobius giurinus 3.9

6 Anguilla marmorata 3.1 Taenioides cirratus 5.4 Taenioides cirratus 3.7

7 Sillago sihama 2.4 Mugilidae spp.A 5.2 Terapon jarbua 3.7

8 Anguilla japonica 2.2 Stolephorus insularis 5.1 Mugilidae spp.A 3.6

9 Leiognathus nuchalis 1.9 Ambassis gymnocephalus 4.5 Ambassis gymnocephalus 3.2

10 Terapon jarbua 1.9 Sillago sihama 4.5 Mugil cephalus 2.9

11 Other (74 species) 10.3 Other (107 species) 36.9 Other (121 species) 62.2

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seasonal dynamics of the larval and juvenile fish community in the mangrove estuary of Tanshui River, Taiwan. Marine Biology 113, 481–90.

Tzeng, W. N., and Wang, Y. T. (1993). Hydrography and distribution dynamics of larval and juvenile fishes in the coastal waters of the Tanshui River estuary, Taiwan, with reference to estuarine larval transport. Marine Biology 116, 205–17.

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estuaries. Environmental Biology of Fishes 28, 295–308.

Manuscript received 21 May 2001; revised 10 December 2001; accepted 9 February 2002

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Appendix 1. Species composition, developmental stage and geographic distribution of four estuaries from December 1997 through November 1998

SC, GST, TT and TK refer to Fig. 1. Le, Leptocephalus larva; Pr, preflexion larva; Fl, flexion larva; Ge, glass eel; Po, postflexion larva; Ju, juvenile; Yo, young; Te, temperate; Tr, tropic; W, world wide. *dominant stages; –, not caught

Family and species Stage No. of fish Geog.

SC GST TT TK DASYATIDAE Dasyatis akajei Yo – 1 – – Te ELOPIDAE Elops hawaiensis Le – 217 63 337 Tr MEGALOPIDAE Megalops cyprinoides Le – 6526 1107 5988 Tr ALBULIDAE Albula vulpes Le – – – 1 Tr ANGUILLIDAE

Anguilla bicolor pacifica Ge 7 36 – 32 Tr

Anguilla celebesensis Ge – 4 – – Tr

Anguilla japonica Ge 2 204 – 5 Te

Anguilla marmorata Ge*, Yo 87 240 – 76 Tr

MURAENIDAE spp. Le, Ge* – 2 – 2

OPHICHTHYIDAE spp. Le, Ge*, Yo – 7 1 80

NETTASTOMATIDAE sp. Le – – 1 –

CONGRIDAE

Conger myriaster Yo 1 – – – Te

Congridae spp. Le – 1 1 –

CLUPEIDAE

Sardinella spp. Pr, Fl, Po*, Ju, Yo 12 33 814 4 Tr

Ilisha melastoma Pr, Fl, Po*, Ju, Yo – – 211 – Tr

ENGRAULIDAE

Encrasicholina heteroloba Ju – 1 – 2 Tr

Encrasicholina punctifer Ju*, Yo – 19 14 – Tr

Engraulis japonicus Fl, Po, Ju*, Yo,

Ad

8 2 81 1 Te

Stolephorus insularis Fl, Po, Ju*, Yo 10 57 1858 13 Tr

Thryssa chefuensis Fl, Po, Ju*, Yo – 57 643 70 Tr

CHANIDAE

Chanos chanos Po*, Ju, Yo 13 168 9 158 Tr

CYPRINIDAE Carassius auratus Yo – 3 – – Te Paracheilognathus himantegus Yo 1 – – – Te Zacco platypus Yo 6 – – – Te ARIIDAE Arius sp. Yo – – 2 – Tr PLOTOSIDAE Plotosus lineatus Yo – 1 – – Tr SYNODONTIDAE Saurida gracilis Ju 1 – – – Tr Saurida wanieso Ju – – 1 – Tr

Trachinocephalus myops Fl, Po, Ju* – 1 6 – Tr

Synodontidae spp. Po, Ju* – 1 – 2

PARALEPIDIDAE spp. Po, Ju* – 3 1 –

MYCTOPHIDAE

Benthosema pterotum Fl, Po*, Ju*, Yo – – 13 2 W

Myctophidae sp. Fl – – 1 –

BREGMACEROTIDAE

Bregmaceros neonectabanus Ju*, Yo – 1 1 1 Tr

EXOCOETIDAE sp. Yo – – – 1 Tr

BELONIDAE

Strongylura anastomella Yo – 1 – – Te

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Appendix 1. (continued)

Family and species Stage No. of fish Geog.

SC GST TT TK

HEMIRAMPHIDAE sp. Ju – – 1 – Tr

POECILIIDAE

Gambusia affinis Ju, Yo* – 49 2 1 Tr

ATHERINIDAE

Atherinomorus lacunosus Yo 1 – – – Tr

Atherion elymus Yo 2 – – – Tr

Hypotherina valenciennei Yo 5 – – – Tr

Hypotherina woodwardi Yo 4 – – – Tr

Atherinidae spp. Po, Ju, Yo* 3 – 1 1

PEGASIDAE sp. Ju – – – 1 Tr SYNGNATHIDAE Hippichthys spicifer Yo 1 9 – – Syngnathidae spp. Yo – – 3 13 SCORPAENIDAE Sebastiscus marmoratus Pr*, Yo – – 1 – Te Scorpaenidae spp. Pr*, Po, Ju – 2 1 31 PLATYCEPHALIDAE spp. Ju*, Yo – 104 8 44 Tr CENTROPOMIDAE

Ambassis gymnocephalus Po*, Ju, Yo 186 2482 530 24 Tr

Ambassis urotaenia Po, Ju*, Yo – – 46 11130 Tr

PERICHTHYIDAE

Lateolabrax japonicus Fl, Ju*, Yo – – 6 – Te

SERRANIDAE spp. Ju – 15 – 30

TERAPONIDAE

Pelates quadrilineatus Yo – 1 – – Tr

Terapon jarbua Po, Ju*, Yo 102 35770 19 723 Tr

KUHLIIDAE

Kuhlia marginata Pr, Fl, Po, Ju*, Yo 12 83 334 5086 Tr

PRIACANTHIDAE

Priacanthus sp. Fl 1 – – – Tr

APOGONIDAE

Apogon spp. Po, Ju* – 25 – 11 Tr

SILLAGINIDAE

Sillago japonica Fl, Po, Ju*, Yo 1 216 60 99 Te

Sillago maculata Po, Ju*, Yo 4 655 31 – Tr

Sillago sihama Po, Ju*, Yo 64 6738 74 67 Tr

Sillaginidae spp. Ju, Yo* – 15 – 31

CARANGIDAE

Caranx lugubris Yo 41 21 – – Tr

Caranx sexfasciatus Yo 12 7 – – Tr

Scomberoides tol Ju*, Yo 7 55 – 12 Tr

Selar crumenophthalmus Po – – 1 – Tr

Trachinotus baillonii Yo – – – 1 Tr

Carangidae spp. Po, Ju*, Yo 5 3 2 13

LEIOGNATHIDAE

Gazza minuta Yo 12 5 – 1 Tr

Leiognathus elongatus Yo – – – – Te

Leiognathus nuchalis Po, Ju*, Yo 30 353 24 12 Tr

Secutor ruconius Yo 1 7 21 13 Tr

Leiognathidae spp. Pr, Fl, Po, Ju, Yo* 1 52 20 229

LUTJANIDAE

Lutjanus argentimaculata Po, Ju* 1 143 – 1 Tr

Lutjanus fulviflamma Ju – 4 – – Tr

Lutjanus russellii Ju*, Yo – 62 – – Tr

Lutjanus vita Ju – 49 – 26

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Appendix 1. (continued)

Family and species Stage No. of fish Geog.

SC GST TT TK

GERREIDAE

Gerreomorpha japonica Po, Ju*, Yo 5 1508 – 8 Te

Gerres abbreviatus Po, Ju*, Yo 241 2040 1 8 Tr

Gerres filamentosus Ju*, Yo 89 1624 1 13 Tr

Gerres macrosoma Ju*, Yo 37 9284 173 108 Tr

HAEMULIDAE

Hapalogeny nitens Yo 2 – – – Te

Plectorhynchus sp. Ju – – – 1

Pomadasys spp. Pr, Fl, Po, Ju*, Yo 6 228 9 31

SPARIDAE

Acanthopagrus latus Po, Ju*, Yo 6 811 38 – Tr

Acanthopagrus schlegeli Fl, Po, Ju*, Yo – 281 125 240 Te

Acanthopagrus australis Pr, Po, Ju*, Yo – 4 95 1

SCIAENIDAE spp. Pr, Po*, Ju – 8 1 2

MULLIDAE sp. Ju – – – 1

SCATOPHAGIDAE

Scatophagus argus Ju*, Yo – 158 9 37 Tr

CICHLIDAE spp. Ju*, Yo 1 22 29 6 MUGILIDAE Mugil cephalus Yo 408 904 1 57 Te Mugilidae spp.A Yo 937 10006 405 493 Tr SPHYRAENIDAE Sphyraena barracuda Yo – – 1 – Sphyraenidae spp. Ju*, Yo 1 4 4 3 Tr POLYNEMIDAE Polydactylus plebeius Yo 2 – – – Tr LABRIDAE spp. Po*, Ju 5 72 – 10 Tr SCARIDAE spp. Po*, Ju – 47 1 5 Tr PERCOPHIDAE sp. Po 1 – – – BLENNIIDAE

Omobranchus spp. Po, Ju* 1 6 14 294

CALLIONYMIDAE spp. Pr, Po, Ju*, Yo – 1 13 3

GOBIIDAE

Apocryptodon madurensis Ju – – 40 –

Bathygobius fuscus Po*, Ju, Yo 15 2723 – 4 Tr

Eleotris acanthopoma Po*, Ju, Yo 3 1334 – 477 Tr

Favonigobius reichei Ju, Yo* 4 16 – –

Glossogobius biocellatus Po*, Ju, Yo – 140 – –

Mugilogobius tagala Yo – 3 – –

Periophthalmus cantonensis Yo – 1 – –

Rhinogobius brunneus Po*, Ju, Yo 6 595 55 38 Te

Rhinogobius giurinus Fl, Po*, Ju, Yo 9 5704 1644 3226 Te

Scartelaos viridis Ju – 56 10 –

Sicyopterus japonicus Po, Ju* 367 1157 – 89 Te

Taenioides cirratus Po*, Ju 1 4205 3 2755 Tr

Gobiidae sp. 1 Po*, Ju, Yo – 3119 12 122

Gobiidae sp. 2 Po, Ju*, Yo – 312 – –

Gobiidae sp. 3 Po*, Ju, Yo – – – 230

Gobiidae sp. 4 Po, Ju* – 5773 3 892

Gobiidae sp. 5 Po*, Ju, Yo – 3376 4 367

Gobiidae sp. 6 Po, Ju* – 6 – –

Gobiidae sp. 7 Po, Ju*, Yo – 804 – –

Gobiidae spp. Pr, Fl, Po, Ju*, Yo 37 1956 113 527

GOBIESOCIDAE sp. Po – – 1 –

SIGANIDAE

Siganus fuscescens Ju – 28 1 43 Te

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http://www.publish.csiro.au/journals/mfr

Appendix 1. (continued)

Family and species Stage No. of fish Geog.

SC GST TT TK

TRICHIURIDAE

Trichiurus lepturus Po, Yo* – 3 3 – W

BOTHIDAE spp. Po, Ju*, Yo – 350 1 213

PLEURONECTIDAE spp. Ju – 123 1 –

CYNOGLOSSIDAE spp. Pr, Ju*, Yo – 1 2 52

MONACANTHIDAE spp. Po, Ju*, Yo – 4 1 –

TETRAODONTIDAE spp. Ju*, Yo – 26 11 27

DIODONTIDAE spp. Yo – 6 – 14

Total (fish) 2828 113340 8838 34772

Total (family) 28 49 44 46

Total (species) 56 94 73 77

數據

Fig. 3.Cluster dendrograms  (upper) and MDS ordinations (lower), based on Bray–Curtis similarities of larval and juvenile fish communities sampled in (a) SC, (b) GST, (c) TT, (d)TK and (e) overall data of a-d
Table 2. Average similarity percentage contributed by the major species in the seasonal groups of four estuaries SC, GST, TT and TK refer to Fig
Table 3. Average similarity and dissimilarity percentages contributed by the major species within and between groups

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