Multiyear Use for Spawning Sites by Crucian Carp in Lake Biwa, Japan
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Journal of Advanced Marine Science and Technology Society, Vol. 24, No. 2, pp. 19–30, 2018 Original Paper Multiyear use for spawning sites by crucian carp in Lake Biwa, Japan Yoshio Kunimune*1, Yasushi Mitsunaga*2 *1 Graduate School of Agriculture, Kindai University, 3327-204 Naka-machi, Nara, Nara, 631-8505, Japan *2 Faculty of Agriculture, Kindai University, 3327-204 Naka-machi, Nara, Nara, 631-8505, Japan, [email protected] Received: July, 10. 2018, Accepted: October, 19. 2018 Abstract From April 2008 to July 2009, an acoustic telemetry study on crucian carp was conducted in Lake Biwa, Japan. Twenty Nigorobuna, ten gengoroubuna and one ginbuna captured during spawning seasons were released with ultrasonic transmitters in the South Basin. Signals from the tagged fish were detected by 28 ultrasonic-receiver stations installed around Lake Biwa. Seasonal preference of habitats was examined by distance-based analysis. Seasonal distributions of nigorobuna and gengoroubuna in Lake Biwa were observed including consecutive two spawning seasons. It was revealed that seasonal migration patterns of nigorobuna and gengoroubuna were different. Nigorobuna appeared mainly around the release site all the year round, while gengoroubuna migrated to the North Basin after their spawning season, where they stayed during the non-spawning season, and moved to the South Basin in the next midwinter, where they stayed until the end of the next spawning season. It was also revealed that gengoroubuna preferred lower water temperature than nigorobuna. Gengoroubuna moved faster than nigorobuna and ginbuna. There were high-degree correlations in distance ratios of each station during spawning seasons in the release year and the next year in both nigorobuna and gengoroubuna, suggesting a multiyear use for the same spawning site. Through these results, deeper understanding of these endangered fish species in Lake Biwa and information to help the development of effective conservation programs would be provided. Keywords: crucian carp, acoustic telemetry, seasonal migration, reproductive homing 1. Introduction or submerged-plant zone along the shoreline (Yamamoto et Lake Biwa is the largest natural lake in Japan and can be al., 2006; Suzuki et al., 2007). The spawning sites extend into divided into North Basin and South Basin by the narrowest influent streams, satellite lakes, irrigation channels or paddy region. Lake Biwa is one of a small number of ancient lakes waters connected to the lake and so on. There are many in the world and provides habitats to many endemic species spawning sites along the coast of the South Basin (Kohashi et (Rossiter, 2000). Of 71 fish species living in Lake Biwa, 13 al., 1994; Yamamoto et al., 2006). Larva and early juvenile of species are endemic including 7 cyprinid fish species (Yuma these crucian carp feed on zooplankton as its initial feed et al., 1998). Crucian carp living in Lake Biwa consist of two (Nakamura, 1969; Hirai, 1972; Nagoshi, 1977). Juveniles of endemic species, nigorobuna Carassius auratus grandoculis nigorobuna and gengoroubuna form schools when they reach (CAG) and gengoroubuna Carassius cuvieri (CCU), and one up to 1.6 cm and 1.8 cm of body length respectively, and epidemic species, ginbuna Carassius auratus langsdorfii move away from the shoreline (Hirai, 1972; Nagoshi, 1977; (CAL). Although the endemic species are endangered, they Usuki et al., 1994). Around this period, their feeding habits are also important fishery resources. These crucian carps are begin to diverge. Nigorobuna and ginbuna mainly feed on medium-sized cyprinoid fish (maximum body length: BL benthos and periphyton, while gengoroubuna mainly feed on > 40 cm) (Nakamura, 1969). Nigorobuna is a high-value fish- phytoplankton (Hirai, 1972; Nagoshi, 1977). These fish spe- ery resource as the main ingredient of Funa-sushi, a traditional cies are familiar in Japan, but there is little knowledge about fermented food. Gengoroubuna have been cultured and intro- where and how they live after this period because Lake Biwa duced into lakes and ponds all over Japan. It is now one of the is too large to get the information about their locations. Basic most familiar targets of recreational fishing. However, the knowledge in their distribution by life cycle is critically population of these endemic fish species in Lake Biwa recently important for conservation of these resources. Recently it has has decreased drastically (Yuma et al., 1998; Yamamoto et al., been possible to acquire information about the locations of 2006) and they got registered in Threatened Category IB of fish living in vast and deep water areas by using acoustic the Red List of Ministry of the Environment, Japan in 2007. telemetry (Lucas and Baras, 2000). Kunimune et al. (2011) The spawning season of crucian carp in Lake Biwa is April reported that nigorobuna which spawned in the South Basin to June (Hirai, 1972; Nagoshi, 1977). They spawn in emergent stayed near their spawning sites all the year round and that 19 Yoshio Kunimune, Yasushi Mitsunaga gengoroubuna migrated seasonally between the North and rowest region. There are as many as 120 influent rivers, while South Basins based on their telemetry experiment in 2007. Seta River is the only effluent river located at the south end of They also discussed the relation between the difference in sea- Lake Biwa. The area and the water volume of the North Basin sonal distribution patterns and high water temperature in the are 11 times and 99 times greater respectively than that of the South Basin in summer. Nigorobuna stayed in the South South Basin. Most of the rainwater in the catchment area is Basin, while gengoroubuna migrated to the North Basin to collected in the North Basin, flows down into the South Basin, avoid the high water temperature. and then outflows into Seta River. The North Basin is deep Natal homing for spawning has great adaptive significance (average depth: 43 m, maximum depth: 104 m) and has a to reproductive strategy (Leggett, 1977; Dittman and Quinn, steeply angled topography and few observed littoral zones. In 1996; Odling-Smee and Braithwaite, 2003). It has been contrast, the South Basin has a mean water depth of only 4 m reported that many fish species including cyprinoid fish use the and the entire South Basin is classified as littoral (Tsujimura et same spawning site every year (L’Abee-Lund and Vbllestad, al., 2000). The North Basin is mesotrophic and the South 1985; Lucas and Batley, 1996). Many of such fish appear to Basin is eutrophic. Emerged and submerged plants grow along have a habit of natal homing to maintain its strains and genetic the shoreline and in the littoral zones. In spring and summer, diversity (Odling-Smee and Braithwaite, 2003). these plants grow in large areas of the South Basin and pro- The objects of this study were to confirm reproducibility of vide spawning grounds to many fish species. the telemetry experiment performed by Kunimune et al. (2011) 2.2. Experimental fish to review the seasonal distributions of crucian carps, including Nigorobuna and gengoroubuna captured using eight set nets migration patterns, habitat preferences and moving speeds, belonging to Otsu Fisheries Cooperative off the west coast of furthermore, to characterize the usage of spawning sites in the South Basin from 21 April to 26 May 2008 were used for consecutive two spawning seasons. Through this work, deeper the experiment (Fig. 1, Table 1). This series of capture had understanding about these endangered fish species in Lake been granted exceptional permission by the governor of Shiga Biwa and information to help development of effective con- prefecture. Body measurements and inspection of gonads of servation programs would be provided. the fish were performed during the surgeries for acoustic telemetry. All of the experimental fish were supposed to be not 2. Methods younger than 3 years old based on the previous study of these 2.1. Study area fish species (Tomoda, 1960; Nagoshi, 1977; Nishimori, 1994). Lake Biwa having an area of 670 km2 is the largest natural Fish species was identified by body shape and density of the lake in Japan (Fig. 1). It can be divided into the northern part first gill raker before surgery (Tomoda, 1960) and a small (North Basin) and the southern part (South Basin) by the nar- specimen was obtained from anal fin of each fish for subse- quent random amplified polymorphic DNA (RAPD) assay (Suzuki et al., 2007). If the results of fish species identifica- tion by external appearance and RAPD assay were inconsis- tent, RAPD assay was repeated once more. When the assay provided the same result as the previous assay, the result was adopted. In this way, one fish identified as nigorobuna by external appearance was considered to be ginbuna according to the result of RAPD. Thus twenty nigorobuna (CAG 24–43), ten gengoroubuna (CCU 12–21) and one ginbuna were pro- vided for the acoustic telemetry. 2.3. Acoustic telemetry According to the method described by Komeyama et al. (2007), an ultrasonic transmitter (V13-1L; Vemco Ltd., Canada) was surgically implanted into the abdominal cavity of each specimen. The fish were anesthetized by immersing them in lake water including 0.05% 2-phenoxyethanol for 3 to 5 min- utes. Each surgery took approximately the same time of 5 minutes. The ultrasonic transmitter with a weight in water of Fig. 1 Study area. Each solid circle indicates a station (St.) where 6 g sent out 69-kHz coded ultrasonic signals. These signals a receiver was installed. The numerals on each map are the had individual ID information and were transmitted randomly station numbers. The receivers at St. 13 and St. 14 were installed on the boundary between the North and South at intervals of 30 to 90 sec for about 450 days until the batter- Basins.