Kamaruddin, I.S. (2011). Diversity and feeding guilds of fish populations in Pengkalan Gawi - Pulau Dula section of Tasik Kenyir Terengganu, Malaysia. Master of science thesis. Universiti Putra Malaysia. 94p.
Cyprinids was the most abundant family sampled in Pengkalan Gawi- Pulau Dula section of Tasik Kenyir from February 2008 to January 2009. The cyprinidae is the dominant fish family in the world freshwater ecosystems (Berra, 2007), and the dominance of cyprinids in tropical reservoirs has also been observed in Sri Lankan reservoirs (Amarasinghe, 1992). This group is currently the largest family of freshwater fish in terms of its abundant, genera and species throughout Malaysia (Kamarudin and Esa, 2009). The dominance of cyprinids in Malaysian lentic water bodies were also observed by Yusoff et al. (1995), Ambak and Jalal (1998), Kah-Wai and Ali (2001), Ambak and Jalal (2006) and Kutty et al. (2009). Cyprinids are the riverine species which now can adapt quite well in the lacustrine environment of reservoirs (Yap, 1992; Zakariah and Ali, 1996; and Ambak and Jalal, 1998).
In addition, Notopterus sp. and H. macrolepidota were also present in the monthly samples throughout the study period. This clearly shows that this section was also dominated by these two species. Yusoff et al. (1995) reported that H. macrolepidota was the most abundant species in their study in Tasik Kenyir. Other species showed variation in their composition. One individual fish present such as C. anomalura, L. lineatus, P. jullieni and C. striatus were considered migratory species and non resident to the study area. However, all of the samples caught are native to Malaysia and none of them is introduced into the lake section except for the P. jullieni.
The data collected such as number of individual fish (n) and number of fish species caught (S) contributed to the measurement of species diversity index. Overall, the fish species diversity using the Shannon-Weaver index showed a mean value of 1.41±0.23. According to Kutty et al. (2009) the mean Shannon Index value in Tasik Chini was 1.59 and value which more than 2.0 indicate a stable ecosystem. The Shannon-Weaver index at stations and months in present study ranged from 0.97 to 1.67. This clearly showed that it was rather low when compared to the values obtained from Kah-Wai and Ali (2001) in Chenderoh Reservoir (value of 3.21 and 3.32) and Yap (1982) in Bukit Merah Reservoir (value of 3.12). The size of the reservoir is important as the smaller reservoir holds diverse fish than the bigger one.
The low values for the Shannon-Weaver index in this study could be due to effects from several factors such as overfishing, habitat alteration, eutrophication from cage culture and nutrients from the sewage water. Zakaria et al. (1999) found that species richness, species diversity and species survival in aquatic habitats were affected by other environmental factors such as habitat destruction, the physicochemistry of the water quality, hydrological and topographical characteristics. This study was also reduced the values of the index because only three stations were set up for fish sampling purposes. More sampling stations could describe more accurate information on the fish population in the section ecosystem. Overall, the difference values of Pielou Evenness index and Margalef Richness index in this study were low between sampling stations and months and this indicated that the lake section provide a uniform and suitable habitat for fish species to live in it.
Spatially, Sungai Ikan showed the highest value in Species Richness (value of 4.75), the highest number of species landed (11 species) and the highest in terms of number of individual fish (128 fishes). This might be due to the location of this station, as it is located at the river mouth of Ikan River. The majority of the fish species still dominate the rivers and the areas near the river mouth where nutrients are found in abundance (DOF, 1994). This condition is almost similar to the conditions in the Chenderoh Reservoir where Ali and Lee (1995) found that the most productive area in terms of fish catch was at the mainstream section of the reservoir. While, in Tasik Chini, Kutty et al. (2009) reported that the most diverse and abundance fish species was near the river mouth of Sungai Chini.
Sungai Ikan station is shallower than the other stations and the riparian zone was characterized by abundant of vegetations and surrounded with tall grasses. Aquatic plants are known as nursery grounds for small fishes and they use the plants as refuge from predatory fishes (Putz, 1997). The availability of macrophytes provides more living spaces which support a large number of fish species throughout the habitat (Lemly and Dimmick, 1982). According to Khairul-Adha et al. (2009), this characteristic could create suitable conditions for a variety of fish species, and resulted in high species richness in the area. Other than that, some of the riverine species landed in this station maybe originated from the Ikan River. O. hasselti for example was found exclusively at this station and not in other stations. This species was reported by Ambak and Jalal (1998) present only in the riverine systems of Tasik Kenyir and it was associated near the rapid and pool zones of the lake.
Species like B. schwanenfeldii, Notopterus sp. and H. macrolepidota were found to be omnivores in Sungai Ikan. However, the highest proportion of animal origin of food items ingested by these species was Aquatic insects. According to Melo et al. (2004), greater deposition of organic matter occurs in the riverine area and resulted in the establishment of aquatic insect in the sediment, thus it increase the fish proportion in that area. This could be another reason in high number of species, richness and fish individual in that station.
Although Pulau Dula was the furthest station form Pengkalan Gawi jetty, it was actually received more disturbances from the human activities. This is because Pulau Dula station was located near to the route of fishing boat trips to Taman Negara. It is believed that the boat trips activities have influenced the fish community in that area. The Shannon-Weaver index showed that the value in Pulau Dula was lower when compared to the other stations. Magurran (1988) claimed that disturbance towards aquatic ecosystem will reduce species diversity of fish in one area. The successful of one species to breed and increase their community size is influences by the interaction of that species with other organisms without disturbances (Kutty et al., 2009).
Seasonal fluctuations of water level in the lake have also influenced the fish composition and its diversity. Marco and William (1990) pointed out that variability in water levels, patterns of current flow and timing of rains can strongly influence the distribution of fish species. The lowest number of individual fish and species caught were observed during the low water level period. The effect was clearly showed in the results of this study whereby the lowest Shannon-Weaver index and Margalef index values were recorded in October. From observations, water level in Tasik Kenyir was the lowest during this month. Only three species of fish were captured and it is believed that the other species are forced to leave their habitat in this section due to the low water level condition. According to Fischer and Ohl (2005), fish species may have different strategies in responding to limited habitat availability in the low-water-level period. Different fish species found during low and high water level may be associated with the migratory movements of the fish species to another area (Renato et al., 2000).
Low availability of foods is likely to be a factor resulted in migration of fish species of this section. B. schwanenfeldii, Notopterus sp. and H. macrolepidota which captured during this month were classified as omnivore species, sharing common resources of food and flexible in exploiting the prey items. According to Crowder and Copper (1982), when food items in particular environment is abundant the food intake in fish will become narrower and more specific, thus fish avoids omnivory habits. They are several other factors that influence the structure of fish community in aquatic ecosystem. According to Ernesto et al. (2008), water temperature is the abiotic factor driving the structure of fish assemblages in one study area. This was shown on temporal basis of present study where the in situ data of the water temperature reduced beginning from November, resulted in high fish abundance, diversity and species richness after the month of October.
The variations in Trophic Level values between the six species of fish examined in this study suggested that these fishes have different feeding requirements and strategies. In general, two of them are specialist and four of them are generalist. The four species are the primer consumers which classified as omnivore species, while the other two species are secondary consumers which categorized as predators. Although the lake was considered oligotrophic with low productivity (Yusoff et al., 1995), most fishes in the lake section have a high food intake.
H. nemurus showed trend of dominant by diet item of Aquatic insect. The Aquatic insect food item was also important in the diets of three other species although they were categorized as omnivore species. Aquatic insects found in fish stomachs may be due to the location of the lake which is located in the primary rain forest on the hilltops of Terengganu state. Insects are very common in Tasik Kenyir (Hasan and Ambak, 2005). According to Rouf et al. (2008), the Tasik Kenyir catchment area was originally part of the dense of hilly forest thus resulted in abundance of insect. The numbers of insect species tend to be highest in littoral areas of a lake (David and Ananthakrishnan, 2004). The abundance of insects in fish stomachs was also reported by Sakri et al. (2010) in their study in Tasik Kenyir and listed the groups included mayflies, fishflies, water beetle, water striders and dragonfly.
According to Mason and MacDonald (1982), the abundance of this fish group is related to the easy capture of insect in aquatic ecosystem which is defenseless. However, this predator need special adaptations in order to consumed Aquatic insect as their food items. According to Tejerina-Garro et al. (1998), besides visual view, some fish species have other kind of senses that allow them to locate prey. The prey is selected due to the cost-benefit ration between ingestion and energy to capture the prey is minimized (Wootton, 1998) furthermore insects are rich in proteins and nutrients (Nico and Morales, 1994).
The use of ‘animal origin of food items’ such as Aquatic insect and Fish food items in the diet of omnivore species was almost consistent with the intake of ‘plant origin of food items’ such as Plant materials and phytoplankton. It is possible that this kind of group is more generalist, using other types of food when they are available in larger amounts (Melo et al., 2004). According to Lowe-McConnell (1987) generalist species have better chance to become widely distributed in aquatic ecosystem. This could be the reason on the dominant of omnivore species such as B. schwanenfeldii, Notopterus sp. and H. macrolepidota in this lake section.
On the contrary, only one piscivore guild was observed during this study that was C. micropeltes. Stomach contents of C. micropeltes were found with the highest percentage of Fish food item. This was almost similar to the results of the study by Sakri et al. (2010) who studied the stomach contents of fish species in Tasik Kenyir and classified this species as piscivore species. The morpohometric characteristics of C. micropeltes clearly showed that this species was a real predator. C. micropeltes was piscivorous with a strong jaw and wide mouth aperture (Wootton, 1998). Although the intake of Fish food item provides high quality of nutrients, the feeding strategy of predator required complex adaptations to overcome the defence strategy of the prey (Keenleyside, 1979).
Stomach contents of H. macrolepidota and C. micropeltes in this study were examined with fish food items (prey) that were still in good shape. This may be due to the habits of these species which swallow their prey in pieces or as a whole. The stomach plays an important role as a storage organ, particularly for carnivorous species which swallow large food items (Kapoor and Khanna, 2004). Although C. micropeltes is highly predatory species, the stomach of this species also contained Plant materials. These plant materials are believed to be incidentally ingested by the fish during the time of catching prey. According to Blaber (1997) plant materials in fish stomach sometimes is not digested and its intake could be accidental.
Different trends of feeding habits were observed seasonally and spatially. Sungai Ikan was actually demonstrated the simplest feeding habit because all of the three species (B. schwanenfeldii, Notopterus sp. and H. macrolepidota) were categorized as omnivore species. None of them was classified as Aquatic insect feeder although B. schwanenfeldii and Notopterus sp. were fed on this prey. The most complex feeding habits was observed at Pulau Pupi. Some fish species earlier known as omnivore species changed their diet to include either Plant materials and become herbivore, or include Fish food items and became carnivore species. This could be an evidence of wider variety of food items available in this habitat for fish species to select (Hajisamae et al, 2003).
Temporally, some fish species underwent changes in their feeding habits during different seasons. These were B. schwanenfeldii and H. macrolepidota. The dietary shifted from Aquatic insect feeder to herbivore species for B. schwanenfeldii, and from Piscivore species to Detritivore species for H. macrolepidota. On the other hand, carnivore species presents during February/April and May/July, however, this guild was absent during August/October and November/January. According to Hajisamae et al. (2003), the dietary shifts are probably a major feature in structuring the trophic guild in the habitat.
Although H. macrolepidota had Fish food items in the diets during November/January, they were accompanied by other food item such as Detritus, thus were not considered as piscivore species. The types of Detritus found were scale, rock, sand and mud. From this finding, it is proven that the detritivory may be one of the important feeding strategies available during this season. Caragitsou and Papaconstantinou (1994) described that the organic content of mud may play an important role in the nutrition of fishes. This is because Detritus presence large amounts of microorganisms (Keenleyside, 1979).
The feeding types during February/April and May/July showed that there were three different feeding guilds observed in each season. This indicated that the three species fed on specific type of food items due to each food items were plentiful each season. However, the specific type of feeding guild does not mean that the species feeds completely on different suite of foods. There was some overlap in food choices where Aquatic insect, Plant materials and zooplankton were found in all stomach of these fishes. The fish species in particular habitat, however, has many strategies for avoiding trophic competition and in optimizing available resources (Hajisamae et al, 2003).
Results for the Length-Weight Relationship (LWR) of three most dominant species were computed using regression analysis. The species were measured their LWR data because the sample size for each species was the highest. With this LWR data, we confirmed that the growth of individual fishes of these three species were linear between their length and weight. The value of r lies within the ranged of -1 and +1, with the extreme positive values indicating perfect linear association (Gomez and Gomez, 1984). Ayoade and Ikulala (2007) found that when the r value is high this indicates that fish increased in length when it increased in weight.
Meanwhile, the slope (b) (growth exponent) for B. schwanenfeldii was recorded within the limits (2 to 4). Earlier studied done by Tesch (1971) revealed that most fishes in aquatic ecosystems have b values ranged between 2 to 4. Additional to that, when b is equal to 3, this indicates that the fish grows isometrically and values other than 3 indicate allometric growth (Rahim et al., 2009). Arslan et al. (2004) mentioned that the positive or negative allometric growth occurred when the b value was different from 3. In this study, none of the fish species were recorded to grow isometrically because the b values were not equal to 3. The calculated b value was higher than 3.0 showed positive allometric growths for H. macrolepidota, while the value lower than 3.0 shows a signs of negative allometric growths for Notopterus sp. If the value of b is significantly larger than 3.0, this means that the larger fish are heavier for their length than the shorter fish (Carlander, 1997).
The change of b values depends primarily on the shape and fatness of the species (Cherif, et al., 2008). According to Sparre (1992) other factors may affected the differences in LWR of fish such as the quantity and quality of food in one area. In this study, food preferences of the fish species obviously contributed to the difference values of b in the LWR. The species showed negative allometric growth was the omnivore fish species which consumed mainly on Aquatic insects and Plant Materials meanwhile the species with positive allometric growth was the piscivore fish species. Schneider et al. (2000) described that the exact relationship between length and weight differs among species depending to their body shape and condition (robustness) of individual fish.
The results for the condition factor (K) of fish species in this study indicated that the weight of B. schwanenfeldii was heavy for its length, while for Notopterus sp. the weight is light for its length. According to Wootton (1998), fish species with high values of K are heavy for their length while low values of K are light for their length. The general cause of the condition factor trend is likely related to food availability (Whicker et al., 1997). However, Notopterus sp. was recorded with high value of Stomach Fullness (SF) indicated that the food availability of this species was highly available. The low K value of this species could be due to physical characteristics of the species where Mohsin and Ambak (1983) described that Notopterus sp. has a laterally compressed body and sickle shaped with an upwardly curved tail. Additional to that, Schneider et al. (2000) stated that the condition of the fish sometimes reflects also on the sex and gonad development of the fish.
Overall, the results showed that only 13 species of fishes were found in the Pengkalan Gawi- Pulau Dula section of Tasik Kenyir. This result is differs when compared to previous studies, where Yap (1992) and, Ambak and Jalal (1998) reported 36 species from 13 families caught in the Kenyir Reservoir. Meanwhile, Furtado et al. (1977) and Yusoff et al. (1995) reported that there were 35 species of fish from 13 families sampled that mainly consisted of riverine species. On the other hand, Thalathiah et al. (1993) found that there were 30 species of fish lived in the Tasik Kenyir.
Although there were differences in the number of species captured when compared to past studies, almost all of the species landed in this study were recorded in previous studies. The difference in terms of number of species caught from this study when compared to the previous studies could be due to the fact that the present study was conducted only in one section of the lake and not throughout the whole lake. Different location of sampling stations could also affect the results. The present study was conducted mostly in the littoral area of the lake, while study done by Ambak and Jalal (1998) for instance was conducted in the riverine habitats such as riffles and pools.
The different result in this study when compared to the previous studies may be also due to the method of sampling using gill nets which does not captured fish that live at the bottom of the lake. If bottom dwelling species are abundant in that area, then the gill nets would underestimate these demersal species. It is clear that each study used different fishing gears and resulted in the differences in species diversity observed (Jimmy, 2007). Similarities to the stretch mesh size of the net used which was 2.5 inches, fish bigger or smaller than this size would not be caught. It is acceptable that these traditional approaches can be influenced by sampling error that could affect the accuracy of the species available (Jimmy, 2007). Furthermore, the catch of fish was only included for the night catch where the time was between evening to the next morning and this does not include the day catch. This may influence the total number of individual fish and species captured. To reduce these problems, it is suggested that to used also other fishing gear that suitable to catch demersal fishes such as fish trap (bubu) or long line with multiple hooks (rawai). It is also recommended to use various stretch-mesh sizes of the gill nets to catch different size of fishes.
Species diversity and fish composition in Pengkalan Gawi – Pulau Dula section of Tasik Kenyir were relatively low when compared to previous studies done in the whole lake of Tasik Kenyir and in other existing reservoirs in Malaysia. The main family recorded in this study was Cyprinidae and it was dominated by B. schwanenfeldii. The other species found in abundant during the study period were Notopterus sp. and H. macrolepidota.
The Shannon-Weaver index values showed that the Sungai Ikan and Pulau Pupi habitats support diverse fish populations compared to the Pulau Dula habitat. Meanwhile, the values of Pielou evenness index and Margalef richness index indicated that the fish population was evenly distributed between stations and months and in general the lake section provide uniform environment for fish species to live in it.
On the other hand, the Length-Weight Relationship and the Condition Factor for the three dominant species obtained in this study indicated that the food sources available in the lake section contributed to a good quality of physical development of the species.
The findings of this study are expected to benefit the planning and management towards sustainable and conservation of the fisheries resources so that future studies could be conduct in understanding the fish population in Tasik Kenyir. Pengkalan Gawi– Pulau Dula section of Tasik Kenyir can be properly managed and developed for eco-tourism and fish production as a sustainable basis.
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BIODATA OF STUDENT
The author, Izharuddin Shah Bin Kamaruddin was born on 18th July 1981 in Klang, Selangor. He did his Diploma of Fishery in Universiti Putra Malaysia Terengganu in 1998 for 1 year, before he got promoted to a bachelor degree level. He received his Bachelor of Science (Human Resource Development) in Universiti Putra Malaysia in year 2003. After pursuing his degree, he worked as a customer service officer in a satellite TV company (Astro) for 4 years. In January 2008, the Universiti Putra Malaysia through the Ministry of Science and Technology Malaysia (MOSTI) granted him a scholarship to pursue his study in Master of Science.
LIST OF PUBLICATIONS
Kamaruddin, I.S., A.S. Mustafa Kamal, A. Christianus, and S.K. Daud, (2009). Species Composition and Feeding Habits of Fish Population in Pengkalan Gawi - Pulau Dula Section of Tasik Kenyir, Terengganu. In: Omar, D., L.T. Chwen, S.K. Balasundram, H.Z. Jaafar and J. Talib (eds.). Tropical Agriculture in a Changing Climate and Energy Scenario. Agriculture Congress 2009. Faculty of Agriculture. Universiti Putra Malaysia. –Proceeding.
Kamaruddin, I.S., A.S. Mustafa Kamal, A. Christianus, and S.K. Daud, (2009). Species Composition of Fish Population in Pengkalan Gawi - Pulau Dula Section of Tasik Kenyir, Terangganu. In: Palmer, R. (ed.). Sustainable Aquaculture and Quality Seafood for All. Asian Pacific Aquaculture Congress 2009. World Aquaculture Society. –Proceeding.
Kamaruddin, I.S., A.S. Mustafa Kamal, A. Christianus, and S.K. Daud, (2010). A Study on dissolved oxygen, pH and temperature of the lake water in Pengkalan Gawi – Pulau Dula Section of Tasik Kenyir, Terengganu. Towards a Healthier and Sustainable Future: International Annual Symposium on Sustainability Science and Management (UMTAS 2010). Universiti Malaysia Terengganu. –Proceeding.
Kamaruddin, I.S., A.S. Mustafa Kamal, A. Christianus, and S.K. Daud, (2010). Stomach Fullness of Fish Population in Pengkalan Gawi – Pulau Dula Section of Tasik Kenyir, Terengganu. Conservation and Sustainable Development. National Conference on Natural Resources (NCNR 2010). Universiti Malaysia Kelantan. –Proceeding.
Kamaruddin, I.S., A.S. Mustafa Kamal, A. Christianus, S.K. Daud, and L. Yu-Abit, (2011). Fish community in Pengkalan Gawi – Pulau Dula Section of Kenyir Lake, Terengganu, Malaysia. Journal of Sustainability Science and Management. Universiti Malaysia Terengganu Publisher. 6: 89-97pp – Journal.