Yukio ONODA. Abstract

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Rikunomizu(Limnology in Tokai Region of Japan)74 : 29-34(2016) Report Rediscovery of Japanese charr in the Denjogawa River and its tributary in 2016 after a disturbance from the Ontake Landslide in 1984: significance of a tributary as a refugium from disturbance 1), 2), 3) Yukio ONODA Abstract Japanese charr (Salvelinus leucomaenis japonicus) was rediscovered in Denjogawa River and its tributary in 2016 against reports of its extinction following huge sediment inflows caused by the Ontake Landslide associated with the Western Nagano Prefecture earthquake in 1984. All of the 27 individuals of the charr were observed in pools more than 30 cm deep. Most (25 of 27) of the fish were observed in the tributary, and relative density (represented as count per unit effort [CPUE]) was higher at upstream sites in the tributary. Moreover, the fish in the tributary had wider ranges of body size. In Denjogawa River, the species was observed downstream of the confluence but not upstream although both sites had pools of depths more than 30 cm. As relative density gradually decreased from upstream of the tributary to downstream of the confluence, the population in the tributary should be considered as the source population. Furthermore, the population in the Denjogawa River should have been maintained by the individuals drifting from the tributary. Additionally, the wide ranges of the body size of fish in the tributary suggest that the population in the tributary was maintained through reproduction. Because the small population in the tributary may function as a source population for that in the Denjogawa River, it will be significant to secure migration pathways between the mainstream and the tributary for continuity of the population in the Denjogawa River system. Key words: mountain stream, disturbance regime, refugia, Mt. Ontake, the Western Nagano Prefecture earthquake (Received: 20 November 2016; Accepted: 25 November 2016) Introduction As mountain rivers tend to undergo massive disturbances such as from landslides due to high gradient, habitats of fishes in the rivers tend to be unstable and fail to sustain their populations (Geertsema et al., 2009). However, mountain rivers have many tributaries (Horton, 1945; Allan, 1995), which can function as refugia from such disturbances (Sedell et al., 1990; Koizumi et al., 2013). Accordingly, monitoring of the recovery processes of the fish population after disturbances is significant for understanding the mechanisms that sustain fish populations in these rivers. Nevertheless, such monitoring is difficult because massive disturbances sometimes cause extinction of fish populations and because records of changes in fish population are scarce. Fortunately, I had the rare opportunity of finding a population of the Japanese charr (Salvelinus leucomaenis japonicus), which had been reported to be extinct in the Denjogawa River system after the Ontake Landslide caused by the Western Nagano Prefecture earthquake in 1984 (Takeda, 1985). As the population was found about the confluence of the Denjogawa River and a tributary, this provides opportunities to examine the significance of tributaries for resilience of the fish population in the mountain rivers. In this study, 1) Mie University Research Center, 1577 Kurimamachiya-cho Tsu city, Mie 514-8507, Japan 2) Mount Ontake Volcanic Eruption Research Group of the Tokai Branch of the Japanese Society of Limnology (with Drs. Kentaro NOZAKI, Takashi TASHIRO, and Tomomasa TANIGUCHI) 3) Kiso Biological Station, Kyoto University, 6388 Fukushima Kisogun Kisomachi, Nagano 397-001, Japan 29 C2016 The Tokai Branch of the Japanese Society of Limnology

Yukio ONODA the significance of the tributary for resilience of fish population was evaluated by comparing fish density and size distribution among sites around the confluence. Methods Japanese charr was searched at five sites located near the confluence of the Denjogawa River and a tributary located on the southern side of Mt. Ontake in central Japan (Figs. 1 and 2A). The watershed areas of the Denjogawa River and the tributary were estimated at 8.52 and 1.95 km 2, respectively, by a spatial analyst tool (hydrological analysis) (ArcView 10.2; ESRI Japan) using fundamental geospatial data from Geospatial Information Authority of Japan as digital elevation model (DEM) data (T. Tashiro, unpublished data). Water quality were measured in the Denjogawa River (electrical conductivity: 41.40 ms m -1, water temperature: 14.1 C, ph: 7.5, turbidity: 0.36 NTU, and chromaticity: 1.54) and the tributary (ibid. 2.30 ms m -1, 12.8 C, 6.8, 0.17 NTU, and 2.62, respectively) at 11:00 and 12:45 h on September 15, 2016, respectively (K. Nozaki, unpublished data). Each site was chosen to include one or more pools with a depth greater than 20 cm, which is preferred by the charr (Yagyu, 2009) and considered as potential habitat of the fish. All the sites were separated by placing weir(s), except between sites 2 and 5 (Fig. 1B). Fish censuses were performed at all sites on September 15 and 16, 2016, by snorkeling for 15 or 30 min according to the observed fish density. The fish species was identified under water as Salvelinus leucomaenis japonicus based on the patterns of colors and spots (Nakabo, 1993). Because adults of the subspecies S. l. pluvius were absent, young fish with white spots on the sides of body but not on their back or vertex of the head were also identified as S. l. japonicus, moreover (Naruse and Yoshiyasu, 1983). The size distribution was approximated by measuring the total length of the fish to the nearest multiple of 5 cm by visual estimation, corrected using a ruler under water. The numbers observed were divided by the searching times to convert counts into relative density as counts per unit effort (CPUE), according to the method described by Onoda and Kayaba (2016). Results A total of 27 individuals belonging to one taxon (Japanese charr, Salvelinus leucomaenis japonicus) were observed in this study (Table 1, Fig. 2B, C, D). Relative density (CPUE) ranged from 0 to 14 individuals per 30-min observation and decreased gradually along the tributary from upstream to its confluence with the Denjogawa River. In the Denjogawa River, some fish were present at the downstream site (site 2) but not at the upstream site (site 1). The body size distribution of the fish was wider at the sites in the tributary than in the Denjogawa River (Table 1). Noticeably, among all the sites, the body size distribution of the fish was widest (range: 10 30 cm) at the most upstream site of the tributary (site 3). Discussion Fig. 1. Location of the Denjogawa River and a tributary (A), and study sites around the confluence (B). Fig. 1B gives an enlarged view of the inset in Fig. 1A. Area that had received debris from the Ontake Landslide in 1984 is redrawn in gray, according to Mizuyama and Hara (1991). Each number (gray) near a transverse structure in Fig. 1B indicates its height (in meters) from the water surface of the site immediately downstream to it. Possibility of survival of the charr population after a large disturbance The fish populations observed in this study might have survived a large-scale disturbance (i.e., the Ontake Landslide in 1984). This possibility is supported by some collateral evidences. First, all the observed fish were identified as C2016 The Tokai Branch of the Japanese Society of Limnology 30

Figure 2 Significance of a tributary as a refugium for Japanese charr from disturbance A B C D E Fig. 2. Photographs of the study sites near the confluence (A), adult (B, C) and young (D) of Japanese charr, and a dead small snake (Rhabdophis tigrinus) with some bite marks (E). The Denjogawa River (left) and the tributary (right) meet near the mid-bottom of the photograph (A). The photograph (A) was taken by T. Tashiro and the other photographs (B-E) were taken in pools of the tributary by the author. 31 14 2016 The Tokai Branch of the Japanese Society of Limnology

Yukio ONODA Table 1. List of observed fish at the sites in the Denjogawa River and its tributary. Site no. *1 Date in 2016 Observation Time (min) Species Total body length (cm) Observed fish number CPUE *2 Maximum depth (cm) *3 Denjogawa River 1 Sep. 16 15 0 0 55 2 Sep. 15 15 Salvelinus leucomaenis 20 1 2 70 10 1 2 70 Tributary 3 Sep. 16 30 Salvelinus leucomaenis 10 4 4 90 15 3 3 90 20 2 2 90 25 4 4 90 30 1 1 90 4 Sep. 15 30 Salvelinus leucomaenis 10 3 3 120 25 3 3 120 30 1 1 120 5 Sep. 15 30 Salvelinus leucomaenis 10 1 1 30 15 1 1 70 25 2 2 70 *1 Detailed information of the sites is presented in the text and in Figure 1. Table 1. Fish observed at the sites in the Denjogawa River and its tributary. *2 Counts per unit effort *3 Maximum depth of the pool where fish were observed or searched. Salvelinus leucomaenis japonicus, a species that was distributed in the Denjogawa River before the landslide in 1984 (Takeda, 1985). Second, artificial release of the fish in the Denjogawa River is not known since the landslide occurred (Y. Sawada, personal communication). Moreover, cultivation of the species is known to be much less preferred than that of the subspecies S. l. pluvius (Tsuchida et al., 2008), suggesting that the observed fish could not have been released. Third, according to aerial photographs taken after the landslide, the tributary studied seems to be located on the margin of a range of the landslide (T. Tashiro, unpublished data, also see Fig. 1B). Given with the above information, the fish population found in this study is likely to have survived the disturbance caused by the landslide, although the possibility that the fish were artificially released should be considered carefully. Maintenance of the local population of the charr in the tributary through reproduction Even if the observed population of charr consisted of released individuals, it is noteworthy that the population has maintained itself through reproduction. The population in the tributary had a wider range of body sizes (10 30 cm), which are equivalent to the body size of one- (body length: 13 16 cm), two- (16 22 cm), and three-year fish (17 25 cm), respectively (Miyadi et al., 1976). Of them, larger charrs, corresponding to two and three years of age, are known to reproduce (Miyadi et al., 1976). Yamamoto et al. (1994) reported reproductive activity in male and female charr that had body lengths of more than 17 and 21 cm, respectively, based on gonado somatic index (GSI). Similarly, Kitano et al. (2005) found reproductive activity in male and female charr that had fork lengths of more than 20 and 22 cm, respectively. Additionally, the charr intrinsically prefer tributaries to mainstreams as habitats for reproduction (Nakamura, 1999). Furthermore, the entire watershed area of the tributary (1.95 km 2 ) is larger than 1.01 km 2, which is needed to maintain charr populations (Endou et al., 2006) although the effects of fragmentation on the population by weirs in the tributary should also be discussed circumstantially in future studies. Therefore, the wide ranges of body size in the tributary suggest that the population has been maintained through reproduction. C2016 The Tokai Branch of the Japanese Society of Limnology 32

Significance of a tributary as a refugium for Japanese charr from disturbance Significance of local population in the tributary as a source population This study suggests that local population in the tributary functions as a source population in the Denjogawa River because relative density decreased along the tributary and the confluence (Table 1). A higher density in the tributary might be caused by the migration for reproduction from the mainstream, because this study was performed in the early reproductive season of the charr (late September to early November, Miyadi et al., 1976). However, the fish are unlikely to have migrated upstream in the tributary because high weirs existed between the study sites (Fig. 1B) and the charr could migrate only downstream in the tributary. As the upper site in the Denjogawa River had no charr, the possibility that the charr population in downstream site of the Denjogawa River being caused by a drift from upstream is quite unlikely. Therefore, the charr population in the mainstream of the Denjogawa River is likely maintained by the charrs provided by the tributary. the resilience of the metapopulation. Changes in the charr population of the Denjogawa River system, including its tributaries, should be surveyed continuously and extensively in further studies to comprehensively understand the effects of tributaries as refugia and the recovery processes from huge disturbances such as landslides. Acknowledgments The author thanks T. TASHIRO for providing data, cooperation in the field survey, and valuable suggestions, Y. SAWADA of the Education Board of Otaki Village for valuable information, Kiso District Forest Office, Forest Agency of Japan for permission to use a forest road for the access to the Denjogawa River, M. YAMADA of Kiso Biological Station, Kyoto University for her hospitality, and Editage (www.editage.jp) for English language editing. This study was partly supported by grants from the Water Resources Environment Center to K. NOZAKI (2015-04). Implications for conservation of sustainable population of the charr It is important to secure the Denjogawa River and its tributaries to allow re-colonization and conservation of a sustainable charr population in the Denjogawa River system. Although the studied tributary is expected to have a watershed area larger than that necessary for the sustainability of the charr population, the usable area for the population may have decreased by isolation due to transverse constructions such as weirs (Endou et al., 2006). A decrease in the usable area should mean a decrease in the carrying capacity. Indeed, a dead small snake (Rhabdophis tigrinus) with bite marks, assumed to have been received from a charr (Miyadi et al., 1976), was found in a pool of the tributary (Fig. 1E), likely reflecting severe shortage of prey resources in the small tributary. As the re-colonization by the charr population increases usable area and the carrying capacity at the watershed scale, it should contribute to conservation of the sustainable population. Additionally, the re-colonization will act to disperse the risks encountered during the next disturbance because the tributary, which functioned as a refugium in this event, might actually receive greater disturbances, such as landslides, in the next event. Moreover, the re-colonization can resolve the isolation, which causes decrease in genetic diversity (Morita et al. 2009). Therefore, securement of the pathways for migration and re-colonization will be significant to improve References Allan, A. D. (1995): Stream Ecology: structure and function of running waters. Chapman & Hall, London. Endou, S., J. Tsuboi and T. Iwata (2006): Effects of damming on the persistence of white-spotted charr and redspotted masu salmon populations. Japanese Journal of Conservation Ecology, 11: 4-12 (in Japanese with English abstract). Geertsema, M., L. Highland and L. Vaugeouis (2009): Environmental impact of landslides. In: Landslides Disaster Risk Reduction, pp. 589-607, Springer Berlin Heidelberg. Horton, R. E. (1945): Erosional development of streams and their drainage basins; hydrophysical approach to quantitative morphology. Bulletin of the Geological Society of America, 56 (3): 275-370. Kitano, S., M. Yagyu, H. Akiyama, S. Kobayashi, H. Ichikawa, S. Endo and S. Yamamoto (2005): Fish fauna of Outaki stream of the Kiso River in central Nagano Prefecture. Bulletin of Nagano Environmental Conservation Research Institute, 1: 9-13 (in Japanese with English abstract). Koizumi, I., Y. Kanazawa and Y. Tanaka (2013): The fishermen were right: experimental evidence for tributary refuge hypothesis during floods. Zoological Science, 30 (5): 375-379. 33 C2016 The Tokai Branch of the Japanese Society of Limnology

Yukio ONODA Miyadi, D., H. Kawanabe and N. Mizuno (1976): Coloured illustration of the freshwater fishes of Japan. Hoiku-sha, Osaka (in Japanese). Mizuyama, T. and Y. Hara (1991): Ontake kuzure ni tomonau dosyadoutai (Sediment dynamics associated with Ontake Landslide). Research Report of Public Works Research Institute, 182: 195-205 (in Japanese). Morita, K., S. H. Morita and S. Yamamoto (2009): Effects of habitat fragmentation by damming on salmonid fishes: lessons from white-spotted charr in Japan. Ecological Research, 24 (4): 711-722. Nakabo, T. (1993): Fishes of Japan with pictorial keys to the species. Tokai University Press, Tokyo (in Japanese). Nakamura, T. (1999): Comparison of physical characteristics of the spawning redds between the fluvial Japanese charr Salvelinus leucomaenis and the masu salmon Oncorhynchus masou masou in the headwater of the Kinu River, central Japan. Nippon Suisan Gakkaishi, 65 (3): 427-433 (in Japanese with English abstract). Naruse, T. and K. Yoshiyasu (1983): Iwana, Salvelinus leucomaenis, from view point of its white spots in head region. In: The Freshwater Fishes Vol. 9. Ed. by E. Kimura, Tansuigyohogokyoukai, Osaka, pp. 109-126 (in Japanese). Onoda, Y. and Y. Kayaba (2016): Comparison of fish fauna in a river that received pyroclastic flow from the volcanic eruption of Mt. Ontake in 2014 with that in neighboring rivers. Rikunomizu (Limnology in Tokai Region of Japan), 74: 23-28. Sedell, J. R., G. H. Reeves, F. R. Hauer, J. A. Stanford and C. P. Hawkins (1990): Role of refugia in recovery from disturbances: modern fragmented and disconnected river systems. Environmental Management, 14: 711-724. Takeda, K. (1985): Ohtakigawa siryuu Denjougawa no zetsumetsusita yamatoiwana (Extinction of Salvelinus leucomaenis japonicas in the Denjougawa River, a tributary of the Ohtakigawa River). In: The Freshwater Fishes Vol. 11. Ed. by E. Kimura, Tansuigyohogokyoukai, Osaka, pp. 151-153 (in Japanese). Tsuchida N., J. Tsuboi and K. Mitsui (2008): Zairai iwana syubyou seisan shiken (A test for seedling production of native Japanese charr). Report of Yamanashi Prefectural Fisheries Technology Center, 35: 7-12 (in Japanese). Yagyu, M. (2009): Foraging microhabitat of Japanese charr, Salvelinus leucomaenis, in a mountain stream. Japanese Journal of Ichthyology, 56: 111-118 (in Japanese with English abstract). Yamamoto, S., Y. Sawamoto and M. Kohara (1994): Total length of maturation on wild charr Salvelinus leucomaenis in Nagano Prefecture. Bulletin of Nagano Prefectural Fisheries Experimental Station, 3: 5-7 (in Japanese). (Editor: Dr. Kentaro NOZAKI, School of Education, Sugiyama Jogakuen University) 摘 1984 年に発生した御嶽崩れ ( 山体崩壊 ) による撹乱後の伝上川とその支流における2016 年のヤマトイワナの再発見 : 撹乱からの避難場所としての支流の重要性 要 小野田幸生 王滝川水系の伝上川では,1984 年の長野県西部地震に伴う御嶽崩れ ( 山体崩壊 ) による土砂流入によって, ヤマトイワナの個体群が絶滅したと報告されていた しかしながら, 2016 年 9 月末に潜水目視調査によって, イワナ属魚類が27 個体発見された 確認された地点は水深が30 cm 以上の落ち込み型の淵であった 確認個体の多くは, 伝上川の左岸に流入する支流で確認され, 上流ほど単位努力量当たりの確認個体数が多かった 伝上川本川では水深が30 cm 以上の淵があったにも関わらず, 支流の合流点の下流側では2 個体が確認され, 上流側では確認されなかった 多くの調査地点の間に治山ダムがあることを合わせて考慮すると, 調査対象の支流がソース個体群であり, 流下した個体群が伝上川で確認されたと考えられる さらに, 伝上川本川と比べて, 支流の個体群の体サイズは幅広いサイズで構成されたことから, 支流の個体群は再生産を通じて維持されてきたと考えられる 支流などを単位とした局所個体群は小規模であるため, 他の個体群との交流が確保されるような移動を保障することが, 山地河川に生息するイワナの保全に重要であると考えられる キーワード : 山地河川, 撹乱体制, 避難場所, 御嶽山, 長野県西部地震 C2016 The Tokai Branch of the Japanese Society of Limnology 34