(Adult Lateolabrax Latus) on a Temperate-Subtropical Rocky Shore
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Foraging ecology of a large opportunistic predator (adult Lateolabrax latus) on a temperate-subtropical rocky shore Seiji Arakaki, Neil Hutchinson, Mutsunori Tokeshi Coastal Ecosystems, 2014, vol 1, 14-27 © 2014 Society for Coastal Ecosystems Studies - Asia Pacific Arakaki S et al. Foraging by a large opportunistic predator Regular Paper Foraging ecology of a large opportunistic predator (adult Lateolabrax latus) on a temperate-subtropical rocky shore Seiji Arakaki1, Neil Hutchinson1,2 and Mutsunori Tokeshi1 1AMBL, Kyushu University, Amakusa 863-2507, Japan 2JCU Singapore, TropWATER - Centre for Tropical Water & Aquatic Ecosystem Research, Centre for Tropical Water & Aquatic Ecosystem Research James Cook University, Singapore 57442 Corresponding author : S. Arakaki, e-mail: [email protected] Abstract Foraging ecology of a large opportunistic predator (adult Lateolabrax latus) was investigated through intensive, targeted sampling of the species on an exposed shallow rocky shore of south-western Japan. The abundance of L. latus increased in autumn and spring, whilst no individual occurred in summer when water temperatures were >24˚C. In spring larger sized individuals (>70 cm TL) tended to occur, resulting in a wide range of body sizes. L. latus consumed a wide variety of organisms, of which fishes and benthic crustaceans were particularly important. Overall, the relative importance of invertebrates as food increased in spring, which may be related to seasonal fluctuations of macro-algae and algae-associated faunas. The fact that a large number of adult fish was collected at high tide in intertidal areas and their guts contained intertidal organisms suggests that L. latus relies on the rocky intertidal habitat as a major foraging ground. This in turn points to the potential importance of large, highly mobile predators such as adult L. latus in the food web dynamics of shal- low-water (including the intertidal) assemblages. Keywords: diet, intertidal, piscivore, sea bass, seasonal occurrence, stomach contents assemblages of organisms. Whilst migration of fishes Introduction into intertidal habitats has been reported under various environmental settings (e.g., Burrows et al. 1994; Rangeley Shallow coastal areas are generally characterised by & Kramer 1995; Sheaves 2001; Faria & Almada 2006; high productivity and provide a large amount of resources Castellanos-Galindo, Krumme & Willis 2010), most studies to diverse marine organisms. In addition, shallow water dealt with juveniles or relatively small fishes and information habitats including intertidal zones constitute important on large-sized foragers is scarce. Intermittent use of nursery grounds for larval stages of marine taxa, generally intertidal areas as foraging sites is a potentially important being considered as ‘refugia’ with lowered risks of predation ecological topic that could link subtidal and intertidal (e.g. Blaber & Blaber 1980; Kneib 1987; McIvor & Odum ecosystems. Although some previous studies suggested 1988, Ruiz, Hines & Posey 1993). Therefore, shallow a possibility that migratory foraging may have a significant coastal areas play important ecological functions for different impact on intertidal communities (e.g. Edwards, Conover & © 2014 SCESAP - 14 - Coastal Ecosystems, 2014, Vol 1, 14-27 Arakaki S et al. Foraging by a large opportunistic predator Sutter 1982; Lubchenco et al. 1984; Castellanos-Galindo, Wong 2003; Islam & Tanaka 2006; Islam, Yamashita & Krumme & Willis 2010), our understanding of such situations Tanaka 2011), studies on L. latus have largely been limited is limited. Thus, the present study focused on the temperate to investigations of their early life stages in surf zones or sea bass, Lateolabrax latus, as one of the potentially estuaries and other soft-bottom environments (Fujita et al. important large predators in shallow rocky shore habitats of 1988; Kinoshita & Fujita 1988; Inoue, Suda & Sano 2005, south-western Japan. Yube et al. 2006). The apparent paucity of ecological Sea basses of the genus Lateolabrax are common information on adult L. latus is due mainly to the difficulty coastal fishes that constitute important elements of of specimen collection and observation on exposed rocky commercial and recreational fisheries in East Asia including shores where adult individuals are known to occur. In Japan, Korea and China. The blackfin sea bass, Lateolabrax order to build a more comprehensive understanding of the latus, is considered to have a relatively limited distribution, ecological roles of a predatory species in shallow coastal mainly occurring on rocky shores of southern Japan, while waters, it is vital to elucidate their patterns of resource a congener L. japonicus occurs widely in both open and utilisation, particularly at the adult stage when the level and inner coastal areas including estuaries, and occasionally in impacts of resource consumption are higher. rivers from northern Japan to southern coasts of the Korean The objective of the present study is to quantitatively Peninsula (Hatooka 2002). analyse the foraging ecology of adult L. latus on a While a good number of studies have dealt with temperate-subtropical rocky shore. We relied on intensive different ecological aspects of L. japonicus (e.g., Mito 1957; line-fishing surveys to obtain numerically adequate data on Hatanaka & Sekino 1962; Matsumiya, Mitani & Tanaka adult individuals. 1982; Matsumiya, Masumoto & Tanaka 1985; Nip, Ho & sizes of the artificial lure baits used, 70-140 mm in length, Materials and Methods meant that fish individuals targeted were skewed towards larger sizes (i.e. adult stage), as intended in the present Study site and fish collection study. Fish collection was conducted at least once in three weeks. Collection effort was raised once a fish was captured Fish samples were collected within a radius of 3 km of the or some responses (biting/chasing) were obtained and Amakusa Marine Biological Laboratory, Kyushu University discontinued when there was no sign of fish (with a minimum (32˚32'N, 130˚02'E), located in the north-western corner of effort of once in three weeks). Time of capture, total body the Amakusa-Shimoshima Island, south-western Japan. length (TL, to the nearest 0.5 cm) and body weight (to the This area has a variety of shore habitats at the scale of nearest 0.1 kg) were recorded. several kilometers with complex and occasionally strong currents at the entrance of the Ariake Bay facing the East Gut contents China Sea (Tokeshi and Arakaki 2011). The sampling sites were moderately exposed rocky shores with a gentle A total of 112 individuals of Lateolabrax latus (i.e. 60% slope and a tidal amplitude of about 3.5 m: the shores were of those captured in April 2004 - March 2007, size range exposed for a distance of about 80-120 m seaward at the 39.0 - 73.0 cm) were subjected to gut content analysis. low tide (Fig 1). The substrates were variable and consisted Samples were preserved in 10% formalin and gut contents mainly of rocks and boulders with patchy occurrence of were sorted and identified to broad taxonomic categories. seaweeds, especially in winter to spring season. Water Samples were rinsed in 70% ethanol before being labelled, temperature was monitored with a data logger (HOBO Water dried to constant weight at 60˚C. Preliminary observation Temp Pro v2, onset) at 7 m depth. Using this record, 3-day indicated that both ripe and spent gonads were observed mean temperature trends were worked out to determine the in March while no spent gonad was found before March. annual minimum and the maximum temperature with the Fish samples were therefore divided into two groups, “pre- period from the former to the latter being defined as the ‘rising’ spawning” (October to February, n = 53) and “post-spawning” period and the opposite as the ‘declining’ period. (March to July, n = 59) (see Supplementary Information). Lateolabrax latus individuals (Fig 1) were collected Mean sizes of fishes for gut contents analyses were not during daylight hours (including twilight) between January significantly different between the pre-spawning and the 2000 and March 2007 (except January-February 2006) by post-spawning group (Mann-Whitney U test, P > 0.05). line fishing from the shore using artificial lure baits. The The percentage occurrence of each prey item (%O) was © 2014 SCESAP - 15 - Coastal Ecosystems, 2014, Vol 1, 14-27 Arakaki S et al. Foraging by a large opportunistic predator Fig. 1 (a) Study site at low tide: a moderately-exposed shallow rocky shore, (b) Lateolabrax latus in the pre-spawning season (November), an individual with ‘average’ (common) body size (50 cm TL) in this study and (c) samples of a large (upper, 71 cm TL) and a small (lower, 38.5 cm TL) individual of Lateolabrax latus in the post-spawning season (April). © 2014 SCESAP - 16 - Coastal Ecosystems, 2014, Vol 1, 14-27 Arakaki S et al. Foraging by a large opportunistic predator calculated as the proportion of the total number of guts (of L. latus individuals) containing it, while abundance and Results dry weight data of prey were used to calculate the mean percentage number (%N) and weight (%W) of each prey Fish occurrence item in the guts. One way ANOSIM and nMDS were then used to identify A total of 409 individuals of Lateolabrax latus were collected differences in gut contents between the two sampling in the course of this study. There were two clear modes of seasons based on Bray-Curtis similarities with log(x+1) fish occurrence in spring and autumn, while no individual transformed