Free Access
Issue
Aquat. Living Resour.
Volume 33, 2020
Article Number 12
Number of page(s) 10
DOI https://doi.org/10.1051/alr/2020013
Published online 25 September 2020

© EDP Sciences 2020

1 Introduction

Trophic ecology is fundamental for understanding the functional role of fish species in aquatic ecosystems (e.g. Wootton, 1990; Blaber, 1997; Hajisamae, 2009; Boaden and Kingsford, 2012; Baeta and Ramon, 2013; Kwak et al., 2015; Ende et al., 2018). Therefore, studies on fish feeding ecology have been carried out worldwide (e.g. Blaber, 1997; Lucena et al., 2000; Garrison and Linke, 2000). Food partitioning among fish community members is frequently attributed to competitive or co-operative interactions (Root, 1967; Ross, 1986; Contente et al., 2011). Diets of most fishes change as they grow, but the timing of change varies by species and is often associated with changes in habitat, season (Blaber, 2000) and morphological characteristics (Labropoulou and Eleftheriou, 1997). To maximize energy intake, enhance growth rate and minimize risk of predation are the ultimate goals of ontogenetic changes in the diets of fishes (Brown, 1985). To understand this natural phenomenon, it is important to determine not only food consumption, but also an inter-specific and intra-specific trophic relationships of target species (Elliott et al., 2002). Although, dietary ontogenetic changes have been studied for a wide range of fishes in recent years, unfortunately this includes few tropical species. Spatial and temporal conditions are essential factors structuring trophic ecology of fish in natural ecosystems (Elliott et al., 2002; Hajisamae et al., 2004). Some fishes have to choose between a habitat that provides more abundant and diverse prey, but in which prey is harder to capture, and a habitat which has less prey abundance, but better capture opportunity (Crowder and Cooper, 1982). Coastal habitats of southeast Asia are recognized as an area with a high diversity of resources and provide nursery habitats for many fish species (e.g. Blaber, 2000; Hajisamae and Chou, 2003, Hajisamae and Yeesin, 2010; Hajisamae and Yeesin, 2014; Fazrul et al., 2020), but only some studies emphasized on fish diet and trophic relationship (Bachok et al., 2004; Hajisamae and Ibrahim, 2008; Hajisamae, 2009; Hajisamae et al., 2015; Paul et al., 2018; Islam et al., 2018).

Orange-spotted grouper, Epinephelus coioides (Epinephelidae), is a commercially important species from the Indo-West Pacific and Australia (Russell and Houston, 1989; Randall and Heemstra, 1991; Lieske and Myers, 1994). Juveniles are commonly found in shallow estuarine waters over sand, mud and gravel substrates (Kailola et al., 1993; Sheaves, 1995; Grandcourt et al., 2005). Coastal areas, especially estuarine habitats, along the northern coast of Malaysia and southernmost part of Thailand are known to be important nursery grounds, specifically from December to March. Collecting live juvenile groupers from nature and selling to farmers is a common practice of local fisherman as the main source of income in these areas during the recruitment season. The recruitment of E. coioides is always associated with other fish species such as Lutjanus russellii, Epinephelus areolatus and Plectropomus leopardus. There are few studies of the feeding habits of adult E. coioides in their natural habitat yet none of their juvenile stages. However, one study of juvenile stages is concentrated on aquaculture aspects (Doi et al., 1997). Adults feed on fish, shrimp, crabs and other crustaceans (Hseu et al., 2003); however, an understanding of diet, trophic relationship and ontogenetic changes during early life history stages is needed for sound fisheries management of E. coioides in the region. This study aimed to examine diet and trophic relationship of juvenile E. coioides and its association with other juvenile fish species during recruitment in estuarine habitats in Thailand and Malaysia.

2 Materials and methods

2.1 Study sites

The work was conducted in the southern Gulf of Thailand including four locations in Thailand (Songkhla province, SK; Panare coastal area, PN; Saiburi river mouth, SR; and Bangnara river mouth, NR) and one in Malaysia at Kuala Besut (MS), which were selected based on geographic and availability to sample fish juveniles (Fig. 1). SK is a man-made wave breaker (or jetty) angled 45 degrees to the shore at the river mouth. The structure creates a weak current affected by monsoonal wind and wave, making the area suitable for the collection of juvenile fishes and is suitable nursery ground for E. coioides. PN is located along the coast near a river mouth. This area faces open waters and receives wind and waves directly. SR is at the Saiburi river mouth with a similar geographic formation and wave breaker as at SK. NR faces open waters exposed directly to wind and wave action, similar to PN. MS has similar geographic formation and wave breaker construction as SK and SR. The depths of fish collection at all sites are 2–5 meters in sandy substratum. Basic physicochemical characteristics of the water at all sites were measured in situ with a YSI multi-probe meter, recording three replicates monthly, during the duration of this study (Tab. 1).

Table 1

Physicochemical of water at all sites based on monthly measurement in the lower part of the Gulf of Thailand during December 2015-March 2016. Note: Songkhla (SK), Panarik (PN), Saiburi (SR), Narathiwas (NR), Thailand and Kuala Besut (MS), Terengganu, Malaysia.

2.2 Fish sample

Fish were collected during dark moon periods (13rd day of the waning moon to the 5th day of the waxing moon) from December 2015 to March 2016 at five study sites. A traditional live fish catching technique based on fish aggregating devices (FADs) was used. This involved a “temerang” using clusters of climbing fern (Lygodium spp.) covered with a polyethylene line net (2–3 inches mesh size), or by setting up a cluster of local pine leaves (Fig. 2). Ten to twenty FADs were tied together on the main rope to make a suspended line anchored with sandbags at a depth between 2 and 5 m. These FAD lines were left overnight, and juveniles were retrieved the next morning using a scoop net (approximately 100 × 100 cm length and width by 50 cm depth with 0.2 cm mesh), slowly placed under the FADs, shaking them and lifting the net above the water surface. Fish were immediately killed with iced water, sorted, identified and preserved in 10% formalin and transported to the laboratory for further analysis. In order to evaluate interspecific food competition, the entire catches of four other juvenile fish species which were collected in the FADs: E. areolatus (areolate grouper), P. leopardus (leopard grouper), Palutrus scapulopunctatus (scapular goby) and L. russellii (Russell's snapper).

thumbnail Fig. 1

Map of study area showing five different study sites including Songkhla (SK), Panarik (PN), Saiburi (SR), Narathiwas (NR), Thailand and Kuala Besut (MS), Terengganu, Malaysia along the Gulf of Thailand.

thumbnail Fig. 2

(a) Traditional fish aggregating devices (FADS), (b) method of arrangement a cluster of FADs for capturing juvenile fishes in water.

2.3 Fish biometry

Collected fishes were kept at least five days in formalin, soaked overnight in tap water and stored (70% ethanol). Fish were measured in standard length (SL) to the nearest millimeter with a digital caliper. Fish were cut open, stomach extracted with surgical ocular scissors, and placed them in a petri dish. E. coioides juveniles were grouped into four length classes in SL (cm): <2.00, 2.01–2.50, 2.51–3.00, 3.01–6.80. Part of the foregut of stomachs (Sutela and Huusko, 2000) were dissected and an index of fullness (FL) was scored based on amount of food in the stomach based on the following categories: 0 (empty), 1 (20%), 2 (40%), 3 (60%), 4 (80%), 5 (100%), 6 (>100%, fully distended) (Hajisamae et al., 2003; Hajisamae, 2009). Stomach content analysis for each fish was conducted under a stereo microscope and each dietary items, except amphipods and isopod, was identified to the broad category including; grouper larvae, other fish larvae, small shrimp (shrimp post larvae and acetes), crab megalopa, crablet and copepod. Amphipods and isopod were identified to species when possible using the taxonomic keys of Barnard and Karaman (1991a) and Barnard and Karaman (1991b).

For analyzing the relation between mouth opening and total length of complete prey items, juvenile samples of E. coioides were randomly selected from all locations and months. Measurements were conducted using an ocular micrometer while mouth size was calculated based on Shirota (1970).

2.4 Dietary indices

The diet was quantified according to three metrics; the numeric percentage (%N), gravimetric percentage (%V) and frequency of occurrence (%FO) of each prey item. The index of relative importance (IRI) was calculated using the following formula (Pinkas et al., 1971). IRI =(%N+%V)×%FO .

The percentage of index of relative importance (%IRI) was calculated with the following formula (Cortés, 1997). % IRI = IRI IRI × 100 .

Numerous indices of trophic attributes were used to describe the status and rank the importance of each prey item for each species. These were; (1) vacuity index (VI), which was the percentage of empty stomachs based on total number of stomachs examined, (2) average number of food items (AF), which was an average number of food item observed in each stomach, and (3) diet breadth (Bi), which was calculated using Levin's standardized index based on %IRI data (Krebs, 1989; Labropoulou and Papadopoulou-Smith, 1999). The formula for Levin's standardized index is: B i = ( 1 n 1 ) ( ( 1 i ,j=1 n P i j 2 ) 1 ) , where Bi = diet breadth;Pij’ = proportion of diet of predator ‘i’ that is made up of prey ‘j’ and ‘n’ = number of prey categories. Values of Bi values vary from 0 to 1, where 0 indicates that organism consumes a single food item or 1 when it consumes many food items in equal proportion. The Bi values can be classified as high (>0.6), moderate (0.4–0.6) and low (<0.4).

Diet overlap was calculated by Simplified Morisita index based on %IRI data (Horn, 1966). The formula for Morisita-Horn index is: C H = 2 ( p i j p i k ) p i j 2 + p i k 2 ,

where CH= Morisita-Horn is the index of overlap between species ‘i’ and ‘k’; pij= proportion of food ‘i’ of the total food quantity by species ‘j’; pik= proportion of food ‘i’ of the total food used by species ‘k’ and n = total number of food item. The degree of overlap was classified as low if the index was between 0.0 and 0.29, moderate if between 0.30 and 0.59 and high or considered significant if between 0.60 and 1.00 (Langton, 1982).

2.5 Statistical analysis

One-way ANOVA was used to compare fullness index (FL) and average food items per fish (AF) of all fish species. The ANOVA was also used to test the impacts of length class (4 classes) and study site (5 sites) on FL and AF of E. coioides. Data transformation based on log (X + 1) was applied previously to analysis to reduce non-normality.

Cluster analyses were used to show differences in the dietary composition between (1) E. coioides and other co-existing fish species, (2) E. coioides collected from different study sites, (3) E. coioides of different length classes and (4) E. coioides collected during different months. To do this, mean values of diet, randomly selected from volumetric raw data of each stomach, were calculated to form the group of 3–6 dietary samples. PRIMER statistical package version 5.0 (Clarke and Gorley, 2001) was used to perform a cluster dendrogram based on square-rooted transformed data of dietary samples and Bray-Curtis similarity matrix. A complete linkage of cluster mode was applied to create the dendrogram. A one-way analysis of similarities (ANOSIM) was used to test the differences between the sub-clusters. Similarity percentage (SIMPER) was used to identify food items contributing highly to the trophic differences between groups. In addition, regression analysis was performed on log (x + 1) transformed data to detect relationship between standard length and mouth opening of E. coioides with total length of prey items (mm).

3 Results

3.1 Food composition and trophic relationship of all species

A total of 8037 fish juveniles taken from the FADs including the species of E. coioides, E. areolatus, P. leopardus, P. scapulopunctatus and L. russellii, were examined. Of these, juveniles E. coioides showed a substantially high proportion with 7837 fishes. Trophic attributes of five juvenile fish species are provided in Table 2. The highest FL, VI and Bi were in E. areolatus, E. areolatus and P. Leopardus, respectively. Small shrimp and Grandidierella sp. were dominant food items for E. Coioides and P. scapulopunctatus, while Elasmopus sp. and fish larvae were relevant for E. areolatus, and fish larvae and shrimp for P. leopardus, and shrimp for L. russellii (Tab. 3).

Five out of ten pairs of overlap in prey items examined were significant (CH > 0.60) (Tab. 4). The overlap between E. coioides and P. scapulopunctatus was very high (CH = 0.97) indicating that juveniles of these two species fed on similar foods. This was further supported by the results of cluster analysis (Fig. 3) as they were grouped mainly in the same cluster (Group 2). The overlap of prey items between E. coioides with P. leopardus and L. russellii, though significant, is weaker than that with P. scapulopunctatus. The diet of E. areolatus showed strong overlap with P. leopardus and comparatively weaker overlap with P. scapulopunctatus.

Three dietary groups were revealed by the cluster dendrogram (Fig. 3). The first group consisted of all the dietary samples of L. russelii. The dietary samples of all E. coioides and two samples of P. scapulopunctatus were clustered in the second group. The third group was comprised of the dietary samples of E. areolatus, P. leopardus and P. scapulopunctatus. Differences in the dietary composition of these clusters were significant (P < 0.01, Global R = 0.83). Similarity percentage (SIMPER) indicated the difference was caused mainly by L. russelii which fed mainly on small shrimp (70.3% contribution) with a combination of Elasmopus sp. and fish larvae. Fishes of the second cluster, including E. coioides, fed mainly on a combination of small shrimp (27.4%) and Grandidierella sp. (25.5%) together with some contribution of Elasmopus sp. and fish larvae. The third group fed on an equal combination of fish larvae, small shrimp and Elamopus sp.

Table 2

Trophic attributes of juvenile species in the lower part of the Gulf of Thailand during December 2015–March 2016.

Table 3

Index of relative importance (%IRI) of food for juveniles species collected in the lower part of the Gulf of Thailand during December 2015–March 2016 (Note: Food item: fish larvae = fl, grouper larvae = gl, Grandidierella sp. = Gr, Cheiriphotis megacheles = Cm, Elasmopus sp.= El, Amphilochus spencebatei = As, Paradexamine reva = Pr, Goratelson sp. = Go, Ceradocus sp. = Ce, small shrimp = sh, megalop = mg, young crab = cr, copepod = cop, Sphearomatidea sp. (isopod) = Sp).

Table 4

Morisita-Horn indices (CH) for the diets of juveniles species collected in the lower part of the Gulf of Thailand during December 2015–March 2016. (Note: bold = biological overlap when the value >0.60).

thumbnail Fig. 3

Dendrogram of cluster analysis of dietary samples of juveniles Epinephelus coioides and four co-existing species collected in the lower part of the Gulf of Thailand during December 2015–March 2016. (Note: ec = E. coioides, ea = E. areolatus, pl = P. leopardus, ps = P. scapulopunctatus, Lr = L. Russellii, the numbers 1–6 followed the codes of species = each dietary samples of the species).

3.2 Food composition and trophic relationship of E. coioides

Juvenile E. coioides fed mainly on four food items including small shrimp, the amphipods Grandidierella sp.and Elasmopus sp. as well as fish larvae. Differences in food composition between length classes of this species from the different study sites were observed. Details regarding the prey composition are in Table 5.

Results of ANOVA indicated that length class of E. coioides, study site and month of collection, significantly affected both FL (P < 0.0344, P < 0.0001, P < 0.0001, respectively) and AF (P < 0.0001) (Tab. 6). Additionally, results from regression analysis indicated significantly positive correlations between length of E. coioides and prey size (r 2 = 0.022, F = 6.90, P = 0.0091) and mouth opening and prey size (r 2 = 0.03, F = 10.18, P < 0.0016).

Cluster analysis separated dietary samples of E. coioides to three clusters based on study sites (Fig. 4a). The first and second clusters (Groups 1 and 2) consisted of dietary sample from Panarik (PN) and Narathiwas (NR), respectively. Dietary samples from Songkhla (SK), Saiburi (SR) and Malaysia (MS) together formed the third cluster (Group 3). This difference between groups was significant (P < 0.01, Global R = 0.82). SIMPER indicated that Grandidierella sp. (34.5% contribution), C. megacheles (26.5%) and small shrimp (18.6%) were the main contributors to the separation of the first cluster. The second cluster had Elasmopus sp. (35.8% contribution), small shrimp (21.6%) and Grandidierella sp. (19.3%) as the main contributors. Small shrimp (29.5%) and Grandidierella sp. (25.9%) were the main contributors responsible for the third cluster.

For length classes, dietary samples of E. coioides separated into two main groups (Fig. 4b). Fish length classes of <2.0 cm, 2.01–2.5 cm and 2.51–3.0 cm together formed the first cluster (Group 1a) and separated into two different sub-groups (Group1a and 1b) with samples of fish <2.0 cm indicating a different suite of food items. The second cluster (Group 2) consisted of samples of 3.01–6.8 cm and this separation was also significant (P < 0.01, Global R = 0.97). Grandidierella sp. was the main contributor (41.9%) to the formation of the first cluster, <2.0 cm length class, together with Elasmopus sp. (26.2%) and small shrimp 23.1%. Small shrimp was the main contributor for the second cluster (33.3%), followed by Grandidierella sp., Elasmopus sp. and fish larvae with relative contributions of 20.8%, 17.7% and 12.6%, respectively. The cluster of the largest size fish had small shrimp (47.4%) and fish larvae (36.3%) as the main contributors with 7.6% of grouper larvae contribution.

Temporal factors revealed two different clusters (Fig. 4c). Dietary samples from December 2015 were clustered together as Group 1, while samples from other three months formed the second group with two different sub-groups (2a and 2b). There was a significant difference between these two clusters (P < 0.01, Global R = 0.80). Results from SIMPER indicated that small shrimp (31.5%), fish larvae (27.1%) and Grandidierella sp. (22.8%) were the main contributors to the formation of this group. The second group consisted of shrimp (27.0%), Grandidierella sp. (21.5%) and Elasmopus sp. (19.2%) as the main contributors to the group.

Table 5

Index of relative importance (%IRI) of food for juveniles Epinephelus coioides of different length classes (SL) and collected from study sites along the lower part of the Gulf of Thailand during December 2015–March 2016. (Note: Food item: fish larvae = fl, grouper larvae = gl, Grandidierella sp. = Gr, Cheiriphotis megacheles = Cm, Elasmopus sp.= El, Amphilochus spencebatei = As, Paradexamine reva = Pr, Goratelson sp. = Go, Ceradocus sp. = Ce, small shrimp = sh, megalop = mg, young crab = cr, copepod = cop, Sphearomatidea sp. (isopod) = Sp; Study sites: Songkhla (SK), Panarik (PN), Saiburi (SR), Narathiwas (NR), Thailand and Kuala Besut (MS), Terengganu, Malaysia).

Table 6

Results of analysis of variance for fullness index and total number of food items of juveniles E. coioides of different length classes and collected from different sites in different months along the lower part of the Gulf of Thailand during December 2015-March 2016.

thumbnail Fig. 4

Dendrograms of cluster analysis of dietary samples of juveniles E. coioides of (a) different sites (b) sizes and (c) months along the lower part of the Gulf of Thailand. (Note: sk = Songkhla, pn = Panarik, sr = Saiburi, nr = Narathiwat, ml = Malaysia); dec = December 2015, jan = January 2016, feb = February 2016 and mar = March 2016).

4 Discussion

Juveniles of all five fish species residing along the coastal waters and estuarine habitats in the southern Gulf of Thailand during the recruitment season have a similar suite of food items. All fish species examined in this study had a low fullness index and a high vacuity index indicating that stomachs contained low food volumes, and many had no food. However, four species including E. areolatus, P. leopardus, P. scapulopunctatus and L. russellii indicate moderate or high diet breadth values, except for E. coioides, since they fed on a diverse group of food items. The low value of diet breadth for E. coioides indicates that this species has a more specific food preference compared to the other species. Moreover, this study confirms the importance of zooplankton; small shrimp and fish larvae, and micro-zoobenthos as the main food sources for juvenile stages of marine and brackish water groupers living in shallow coastal habitats (Baldo and Drake, 2002; Elliott et al., 2002; Gning et al., 2008) as they fed mainly on small shrimp, amphipods and fish larvae. Small shrimps and amphipods especially Grandidierella sp. and Elasmopus sp. were the most common food for all species except for L. russellii. Fish larvae was the dominant item of E. areolatus and P. leopardus. Generally, estuarine fishes are characterized as omnivorous, sharing common resources, and with the flexibility to exploit temporary peaks in abundance of prey populations (Ley et al., 1994). Plant or plant materials were not found in this study which is typically reported elsewhere (Laegdsgaard and Johnson, 2001). Additionally, the importance of amphipods as a main food for groupers is highlighted. A significant amount of Grandidierella sp. and Elasmopus sp. in the diets of these fishes had not been recorded previously. Compared to other studies in tropical habitats, Hajisamae et al. (2004) reported that juveniles of Ambassis interrupta, Ellochelon vaigiensis, Escualosa thoracata, Nuchequula gerroides, Stolephorus indicus, Gerres oyena, Sillago sihama and Ambassis kopsii fed mainly on calanoid copepods with lower proportions of gammarid amphipods. Juveniles of S. sihama also fed mainly on calanoid copepod and gammarid amphipods in Thai waters (Hajisamae et al., 2006). Compared to other juvenile groupers, leopard grouper, P. leopardus consumed a high proportion of benthic dwelling crustaceans, mostly penaeid shrimps before shifting to almost entirely to piscivorous (St. John, 1999), black grouper, Mycteroperca bonaci, fed mainly on fishes and crustaceans (Brulé et al., 2005), and red grouper, Epinephelus morio, had crustaceans as the dominant prey items followed by fishes (Brulé and Rodriguez-Canché, 1993). Unfortunately, there is no other description of the diet of E. coioides to compare with.

Food partitioning can be defined as any use of food resources by the species that coexist in the same habitat (Ross, 1986). Food overlap between different species or different sizes of the same species is important to understand fish community organization (Krebs, 1989). As competition for food can lead to a dietary switch, food partitioning and diet switches contribute to the assumption that fishes are extremely adaptable to their trophic surroundings (Gerking, 1994). The diet of E. coioides is highly associated with that of P. scapulopunctatus and overlapped with L. russellii and P. Leopardus (Tab. 3). It can be postulated that taxonomic resolution of prey may influence this result, however it is still a reliable indicator of how fish share food resources. Diets of most fishes change with size as they grow (Wooton, 1990; Blaber, 2000) and this mechanism is important for the survival of a species (Baldo and Drake, 2002; Elliott et al., 2002; Braga et al., 2012; Monteiro et al., 2018). Theoretically, diet changes are due to physicochemical variations in the habitat, or biotic interaction, such as competition or predation risk (Gerking, 1994). The timing of the switch in diet usually relates to juveniles becoming sub-adults or adults prior to leaving nursery habitats.

Body form and mouth opening size determine the success of foraging in fish (Blaber, 1997; Magnhagen and Heibo, 2001; Mithun et al., 2017). It was confirmed in this study that the size of the fish and the mouth opening were positively related to the size of prey. It is clearly demonstrated that the larger fishes and mouth openings, the larger the size of the food items ingested. Although the main diet for each length class remained relatively similar, the relative composition of each food item was different. For example, the smallest size fish fed more on Grandidierella sp. and some combination of shrimp, but as they grew larger the importance of Grandidierella sp. decreased significantly with greater contribution of shrimp and fish larvae. This suggests an ontogenetic dietary shift during early stage of development in natural habitats during the recruitment season by initially feeding on amphipods and small shrimps while shifting to shrimp and fish larvae as they grow. This ontogenetic shift in the diet reflects the ‘optimum foraging theory’ in which the cost/benefit ratio of catching prey is considered (Gerking, 1994).

This study also found that juvenile E. coioides fed slightly differently in different study sites and months. Although the main food items remained unchanged in all sites, their proportions were different. It was clear that fishes from Songkhla (SK), Malaysia (MS) and Saiburi (SR) fed on a similar food composition and fishes from Panarik (PN) and Narthiwas (NR) had a different combination of items. Geographically, it is possible the habitat or ecosystem characteristics were different. The sites at Songkhla (SK), Malaysia (MS) and Saiburi (SR) are at a river mouth with a man-made wave breaker, thus creating weak currents in the vicinity of the nursery grounds. For the Panarik (PN) and Narathiwas (NR) sites, the conditions are of open waters exposed to wave action and strong currents and wind. These differences in physical or chemical oceanography or other environmental dynamics may create the preferred conditions for different types of prey that led to differences in food items ingested by juvenile E. coioides.

5 Conclusion

Juvenile fishes associated with E. coioides recruiting in the southern Gulf of Thailand ingested similar food items which were mainly small shrimp, amphipods and fish larvae. Small shrimp and amphipods especially Grandidierella sp. and Elasmopus sp. were the most common prey for all species except L. russellii. Juvenile E. coioides fed mainly on shrimp, Grandidierella sp., Elasmopus sp. and fish larvae and had the greatest trophic overlap with Palatrus scapulopunctatus. Factors including length class, collection site and months of collection, significantly affected both fullness index and number of food items of juvenile E. coioides. Fishes of larger sizes and larger mouth openings ingested larger prey items. This study confirms the ontogenetic dietary shift of juvenile E. coioides during recruitment season by starting as amphipod and small shrimp feeder and shifting to shrimp and fish larvae predator as they grow. Some differences in diet composition were observed at different sites even though the main food items remained unchanged. This work clearly identified the diet of E. coioides and its trophic association with other juvenile fish species during recruitment in a tropical coastal habitat.

Acknowledgements

We would like to express our thanks to Faculty and Science Technology, Prince of Songkla University under SAT-ASEAN research scholarship scheme for funding of this research. I also would like to express my deep gratitude to Can Tho University for allowing their staff to jointly undertaken this research. Thanks to Arun Loh-hem, Abdulroseh Cheloh, Teuku Haris Iqbal and Sakariya Samae for the field and laboratory works. A special acknowledgement to Dr. Zeehan Jaafar for her valuable comment and Dr. Koraon Wongkhamhaeng for identification of amphipods.

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Cite this article as: Chuaykaur K, Hajisamae S, Ruangchuay R, Dinh TD, Fazrul H. 2020. Diet and food partitioning between juvenile of Epinephelus coioides (Perciformes: Epinephelidae) and other co-existing juvenile fishes in shallow waters of Thailand and Malaysia. Aquat. Living Resour. 33: 12

All Tables

Table 1

Physicochemical of water at all sites based on monthly measurement in the lower part of the Gulf of Thailand during December 2015-March 2016. Note: Songkhla (SK), Panarik (PN), Saiburi (SR), Narathiwas (NR), Thailand and Kuala Besut (MS), Terengganu, Malaysia.

Table 2

Trophic attributes of juvenile species in the lower part of the Gulf of Thailand during December 2015–March 2016.

Table 3

Index of relative importance (%IRI) of food for juveniles species collected in the lower part of the Gulf of Thailand during December 2015–March 2016 (Note: Food item: fish larvae = fl, grouper larvae = gl, Grandidierella sp. = Gr, Cheiriphotis megacheles = Cm, Elasmopus sp.= El, Amphilochus spencebatei = As, Paradexamine reva = Pr, Goratelson sp. = Go, Ceradocus sp. = Ce, small shrimp = sh, megalop = mg, young crab = cr, copepod = cop, Sphearomatidea sp. (isopod) = Sp).

Table 4

Morisita-Horn indices (CH) for the diets of juveniles species collected in the lower part of the Gulf of Thailand during December 2015–March 2016. (Note: bold = biological overlap when the value >0.60).

Table 5

Index of relative importance (%IRI) of food for juveniles Epinephelus coioides of different length classes (SL) and collected from study sites along the lower part of the Gulf of Thailand during December 2015–March 2016. (Note: Food item: fish larvae = fl, grouper larvae = gl, Grandidierella sp. = Gr, Cheiriphotis megacheles = Cm, Elasmopus sp.= El, Amphilochus spencebatei = As, Paradexamine reva = Pr, Goratelson sp. = Go, Ceradocus sp. = Ce, small shrimp = sh, megalop = mg, young crab = cr, copepod = cop, Sphearomatidea sp. (isopod) = Sp; Study sites: Songkhla (SK), Panarik (PN), Saiburi (SR), Narathiwas (NR), Thailand and Kuala Besut (MS), Terengganu, Malaysia).

Table 6

Results of analysis of variance for fullness index and total number of food items of juveniles E. coioides of different length classes and collected from different sites in different months along the lower part of the Gulf of Thailand during December 2015-March 2016.

All Figures

thumbnail Fig. 1

Map of study area showing five different study sites including Songkhla (SK), Panarik (PN), Saiburi (SR), Narathiwas (NR), Thailand and Kuala Besut (MS), Terengganu, Malaysia along the Gulf of Thailand.

In the text
thumbnail Fig. 2

(a) Traditional fish aggregating devices (FADS), (b) method of arrangement a cluster of FADs for capturing juvenile fishes in water.

In the text
thumbnail Fig. 3

Dendrogram of cluster analysis of dietary samples of juveniles Epinephelus coioides and four co-existing species collected in the lower part of the Gulf of Thailand during December 2015–March 2016. (Note: ec = E. coioides, ea = E. areolatus, pl = P. leopardus, ps = P. scapulopunctatus, Lr = L. Russellii, the numbers 1–6 followed the codes of species = each dietary samples of the species).

In the text
thumbnail Fig. 4

Dendrograms of cluster analysis of dietary samples of juveniles E. coioides of (a) different sites (b) sizes and (c) months along the lower part of the Gulf of Thailand. (Note: sk = Songkhla, pn = Panarik, sr = Saiburi, nr = Narathiwat, ml = Malaysia); dec = December 2015, jan = January 2016, feb = February 2016 and mar = March 2016).

In the text

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