Variation in Diet Composition of the Mudskipper, Periophthalmodon Septemradiatus, from Hau River, Vietnam
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Bull Mar Sci. 96(3):487–500. 2020 research paper https://doi.org/10.5343/bms.2018.0067 Variation in diet composition of the mudskipper, Periophthalmodon septemradiatus, from Hau River, Vietnam 1 Department of Biology, Quang Minh Dinh 1 * School of Education, Can Tho Lam Thanh Tran 2 University, 3/2 Street, Xuan 3 Khanh Ward, Ninh Kieu District, Tuyet Thi Minh Tran Can Tho 900000, Vietnam Diem Kieu To 1 2 Bac Lieu University, Bac Lieu Tien Thi Kieu Nguyen 4 City, Bac Lieu 960000, Vietnam Dinh Dac Tran 5 3 Centre for Continuing Education of Soc Trang Province, Soc Trang 950000, Vietnam ABSTRACT.—Periophthalmodon septemradiatus (Hamilton, 4 An Khanh High School, An 1822) is a mudskipper of the Mekong Delta that can be found Khanh Ward, Ninh Kieu District, Can Tho 900000, Vietnam along estuaries and lower reaches of rivers. In the present study, we determined diet and feeding ecology of this species 5 College of Aquaculture and by analyzing the contents within the stomachs of 1360 Fisheries, Can Tho University, 3/2 fish samples collected from August 2017 to July 2018. Data Street, Xuan Khanh Ward, Ninh Kieu District, Can Tho 900000, analysis suggested that P. septemradiatus is a carnivorous fish. Vietnam We found six main food item categories: small fishes, prawns (Acetes spp.), crabs (Uca spp.), molluscs, ants (Dolichoderus * Corresponding author email: sp.), and detritus. Both males and females at different sizes, <[email protected]> seasons, and habitats ingest primarily Dolichoderus sp., secondarily detritus, and rarely other prey. Diet composition was similar between sexes but varied according to fish size, season, and habitat. Dolichoderus sp. and detritus regulate the spatial variation of food composition. Our findings contribute to future artificial cultivation for conservation. Guest Editor: Amy Y Then Section Editor: Rafael J Araújo Date Submitted: 2 September, 2018. Date Accepted: 15 March, 2019. Available Online: 28 March, 2019. Food and feeding ecology are necessary to understand fish biology and trophic interactions among species in a fish community (Brodeur and Pearcy 1992, Wootton 1996, Blaber 2000). Diet composition can vary according to fish size, season, and habitat (Aarnio and Bonsdor 1993, Carman et al. 2006, Brush et al. 2012). This vari- ation is driven by fish foraging behavior and food availability (Dinh et al. 2017b). Knowledge on diet and feeding ecology is limited for some gobiid species in the Mekong Delta, where they have been overfished (Trinh and Tran 2012). Therefore, it is crucial to study the feeding ecology of gobiid species in the Mekong Delta, which can be used for fishery assessment and fish population conservation. Bulletin of Marine Science 487 © 2020 Rosenstiel School of Marine & Atmospheric Science of the University of Miami 488 Bulletin of Marine Science. Vol 96, No 3. 2020 Figure 1. The sampling map in the Mekong Delta. Arrowhead: Sampling area: (1) Long Duc–Soc Trang, (2) An Lac Tay–Soc Trang, (3) Phu Thu–Can Tho, (4) Tan Hung–Can Tho, and (5) Binh Duc–An Giang. Periophthalmodon is a genus of the subfamily Oxudercinae (Gobiidae) that contains three species including Periophthalmodon freycineti (Quoy and Gaimard, 1824), Periophthalmodon schlosseri (Pallas, 1770), and Periophthalmodon septemradiatus (Hamilton, 1822) (Murdy 1989, 2011, Murdy and Jaafar 2017). However, only P. schlosseri and P. septemradiatus are recorded in Vietnam (Tran et al. 2013). Periophthalmodon septemratidatus is a burrowing and amphibious fish (Martin and Bridges 1999), and is widely distributed in the mangrove swamps and mudflat areas of eastern India, Bangladesh, Myanmar, Thailand, Malaysia, Singapore, Indonesia, and Vietnam (Murdy 1989, 2011, Murdy and Jaafar 2017). The mudskipper P. septemradiatus population has recently declined due to various factors including urbanization, environmental pollution, and climate change (Dinh et al. 2018). However, knowledge of biological characteristics and trophic interactions of this species is not sufficient to build efficient conservation strategies. This study provides knowledge of diet and feeding ecology of male and female P. septemradiatus of different sizes, and in different seasons and habitats. Dinh et al.: Diet of Periophthalmodon septemradiatus 489 Materials and Methods Study Sites.—We conducted the present study from August 2017 to July 2018 in muddy regions of tributaries of Hau River including Long Duc–Soc Trang (LD; 9°42´55.4˝N, 106°04´28.4˝E), An Lac Tay–Soc Trang (ALT; 9°49´52.4˝N, 105°59´44.5˝E), Phu Thu–Can Tho (PT; 9°59´45.06˝N, 105°48´22.73˝E), Tan Hung– Can Tho (TH; 10°12´07.17˝N, 105°34´43.89˝E), and Binh Duc–An Giang (BD; 10°24´03.54˝N, 105°25´10.82˝E; Fig. 1). Dinh et al. (2018) reported that silt, clay, sand, and organic matter characterizes the mudflat of these five regions. The distance from the river bank to the riverbed of the mudflat was about 2.5 m at the lowest tide for all sites; however, the vegetation in each site differed as follows: the vegetation in LD consisted mainly of the mangrove apple, Sonneratia caseolaris (L.) Engl., and the mangrove palm, Nypa fruticans Wurmb. The slopes of the river bank at this site were approximately 25°. ALT’s vegetation was predominantly S. caseolaris, N. fruticans, and the water trumpet, Cryptocoryne ciliata (Roxb.) Fischer ex Wydler. The slope of the river bank in ALT was also approximately 25°. Because PT was near industrial zones, none of the predominant plants were found in PT’s vegetation. Much like LD and ALT, the slope of the river bank at PT was approximately 25°. The vegetation in TH consisted mostly of the canary wood, Nauclea orientalis (L.) L., and S. caseolaris. The slope of the river banks in TH was approximately 45°. The vegetation in BD con- sisted mostly of N. orientalis, but we did not find S. caseolaris at this location. The slopes of the river bank in BD were also approximately 45°. At these five study sites, there are typically two seasons, including a dry season with little precipitation (January–May) and a wet season with roughly 400 mm pre- cipitation per month (June–December). The mean annual temperature in this region is approximately 27 °C. The tidal amplitude in our study locations is semidiurnal (Le et al. 2006). Fish Collection.—As fish usually appeared on the mudflat surface at low tide, we used fishing rods to obtain fish specimens of different sizes during low tides. We collected fish for roughly 3 hrs, which coincided with the ebb tide in all five sites. At each site, we chose an area of 30 m2 (15 m along the river bank and 2 m from the river bank to the riverbed) to collect fish monthly. Every field campaign lasted 5 d (1 d per sampling site). After fish collection, we used the external description of Khaironizam and Norma-Rashid (2003) to identify the fish. We used external morphology and genital papilla to differentiate sexes; e.g., the dorsal fin of males was longer, larger, and more colorful than that of females, but the genital papilla of males was smaller and whiter than that of females (Dinh et al. 2018). We fixed specimens in 5% formalin before transport to the laboratory. For fish conservation purposes, we released fish at juvenile stages (e.g., those with total length was <5.0 cm) and at sexual maturation stage (e.g., those with a big abdomen and red genital area, which were about to release gametes) back to their habitat after measuring them. Diet Composition Analyses.—In the laboratory, we determined fish total length (TL, 0.1 cm) and weight (W, 0.01 g), then we dissected the specimens to re- move the gastrointestinal tract. Next, we examined stomach content to determine diet composition. We used a stereomicroscope and Nguyen et al. (2013) to identify to the lowest possible taxonomic level. Finally, stomach contents were quantified by 490 Bulletin of Marine Science. Vol 96, No 3. 2020 prey occurrence in fish stomach (%Oi = 100 × Oi/N, where Oi is the number of fish consuming prey i and N is the total number of fish examined; Hynes 1950) and the gravimetric method (%Wi = 100 × Wi/Wtotal, where, Wi is weight of prey i, Wtotal is to- tal weight of all prey individuals; Hyslop 1980). Dietary composition was quantified using a combined analysis of prey occurrence and weight, also known as biovolume, preponderance index, or point of prey, and was calculated as %Vi = (100 × Oi × Wi)/ Σ(Oi × Wi), where Vi, Oi, and Wi are the percentage of biovolume, occurrence, and weight of prey i respectively. We used the biovolume value to test if diet composition varied by sex, fish size, season, and site (Natarajan and Jhingran 1961, Hyslop 1980). Feeding Strategy Analysis.—Amundsen et al. (1996) modified the Costello (1990) graphical method. We used this method to plot the percentage of biovolume vs frequency occurrence of food items to determine diet specialization (e.g., feeding strategy) and prey importance. With this graph, the most important prey items are closer to the top right corner and the prey items, which occur at low level but have important roles, are closer to the top left corner. The least important prey items are closer to the low left corner, and the prey items with high level but least important are closer to the low right corner (Adámek et al. 2007). Data Analysis.—We used a permutational multivariate analysis of vari- ance (PERMANOVA) using PRIMER v6.1.11 (Clarke and Gorley 2006) with the PERMANOVA+ v1.0.1 add-on package (Anderson et al. 2008) to analyze the bio- volume of all food items to compare diet composition according to sex, size, season, and site (Baeck et al.