Shigeru Nakano - an uncommon Japanese fish ecologist

Kurt D. Fausch
Department of Fishery and Wildlife Biology, Colorado State University, Fort Collins, Colorado 80523, USA

Shigeru Nakano with some present and former graduate students at the Center for Ecological Research at Kyoto University in January 2000.  Left to right are: Yukihiro Kohmatsu, Yoshinori Taniguchi, Yoichi Kawaguchi, Shigeru Nakano, Motoharu Konishi, and Tomoya Iwata.
 

On 27 March 2000 Shigeru Nakano was lost in the Sea of Cortez off Bahia de Los Angeles in Baja California, when the research vessel that he and eight others were using to return from nearby islands capsized in an unexpected storm.  Nakano, Takuya Abe, and Masahiko Higashi, all faculty of the Center for Ecological Research (CER) at Kyoto University in Japan were visiting island research sites where Dr. Gary Polis of the University of California-Davis was studying food webs, and were accompanied by five other researchers and students.  Nakano and his two Japanese colleagues, Polis, and Michael Rose, a postgraduate researcher, drowned.  Survivors reported that Shigeru Nakano repeatedly pulled others back to the capsized boat when they were washed away by the raging sea, and strapped his own life jacket onto one of his colleagues who could not swim, literally giving his own life to save the lives of others.  Nakano was a superb diver and field biologist, the best I have ever known, but I know from personal experiences during grueling field work in the mountains of Japan and Montana that he would never have left his friends to swim to the nearest island more than a kilometer away and save himself.  Nakano's body was not recovered despite an extensive search effort.  He was 37 years old and is survived by his wife and three children, and his parents and brother.

Shigeru Nakano was born 25 November 1962 and raised in Kamioka, a small town in the mountains of Gifu Prefecture in central Japan.  He earned his B.S. degree in 1985 at Mie University in Tsu in south central Japan and entered graduate school there, earning his M.S. degree under the direction of Dr. Makoto Nagoshi in 1987.  His first job was as a biologist in the Hida Nature Education Center in Kamitakara Village near his home town.  In 1989 he moved to become an Assistant Professor (the rank of Joshu in Japan) at the Nakagawa Experimental Forests of Hokkaido University, located in a small town in northern Hokkaido, the northern island in the Japanese archipelago.  He earned his Ph.D. from Hokkaido University in 1991 while working there.  In 1995 he moved to the Tomakomai Experimental Forests in south central Hokkaido where he was promoted to Associate Professor (Jokyoju) in 1997, and became Director in 1998.  In 1999 he moved to the Center for Ecological Research at Kyoto University as Associate Professor to help start a new National Institute of Global Environmental Science with Masahiko Higashi and Eitaro Wada.  Shigeru Nakano's meteoric rise through the Japanese academic hierarchy from relatively modest beginnings to the top institute for ecology is legendary in Japan.

I had the great fortune to first meet Shigeru Nakano during the 1988 Symposium on Charrs and Masu Salmon (see Noakes 1989) in Sapporo, Hokkaido during the period when he was conducting his Ph.D. research.  During the last decade I collaborated closely with him and his colleagues.  Nakano began his research as an undergraduate, traveling to Hirakura Stream, a small mountain stream near Mie University, to snorkel and measure ecology and behavior of the native landlocked redspotted masu salmon (Oncorhynchus masou ishikawae) during extended periods from spring through fall.  This early work was strongly influenced by previous studies on individual behavior of stream fishes by Hiroya Kawanabe (see Yuma et al. 1998) and his student Tetsuo Furukawa-Tanaka of Kyoto University, and resulted in Nakano's first publication (Nakano & Nagoshi 1985).  In 1985 Nakano traveled to Lake Tanganyika with a group of researchers led by Kawanabe to study brood defense in a cichlid fish for his M.S. research, again using underwater observation to study detailed individual behavior and territoriality (Nakano & Nagoshi 1990).  During 1985 through 1990 he conducted the research he published to earn his Ph.D., on ecology and behavior of native whitespotted charr (Salvelinus leucomaenis), Dolly Varden (S. malma), and masu salmon (O. m. masou) in mountain streams in both central Honshu (the main island) and Hokkaido (e.g., Nakano 1994b, 1995a, 1995b, Nakano & Furukawa-Tanaka 1994).  Several of these papers are now cited in one of the leading ecology textbooks as among the best examples of intraspecific and interspecific dominance hierarchies (Ricklefs & Miller 2000).

The 1988 Charr Symposium and the 1990 International Ecological Congress (INTECOL) and satellite conferences held in Japan afforded Shigeru Nakano and many other Japanese ecologists the opportunity to introduce international scientists to their work.  Based on these important contacts Nakano began to travel to international meetings and collaborate with foreign scientists.  During the next decade he conducted research in Alaska (Maekawa et al. 1993), Montana (Nakano et al. 1992, 1998, Kitano et al. 1994), Kamchatka (Nakano & Kaeriyama 1995), the Amur River basin (Nakano 1999), and Borneo, all but the most recent of which is published.  He also developed an extensive network of collaborators within Japan, supervising and publishing research with graduate students from other laboratories in addition to his own (e.g., Nakano et al. 1996, Yamamoto & Nakano 1996, Inoue & Nakano 1997, 1998, 1999, Nisikawa & Nakano 1998).  During 1994 and for the rest of the decade while he worked at the Tomakomai Experimental Forest he expanded his horizons from ecology and behavior of stream fishes to consider the interactions of entire food webs in streams and their riparian zones.  He and his colleagues recently published (Nakano et al. 1999) results from a large-scale manipulation of the flux of terrestrial insects falling into Horonai Stream that drains the Tomakomai Experimental Forest.  They demonstrated that eliminating this subsidy from forest to stream during summer caused the native Dolly Varden to trigger a top-down trophic cascade via intense predation on aquatic insects that graze algae.  Moreover, Nakano and his colleague Masashi Murakami have submitted a manuscript showing that this subsidy shifted from stream to forest in early spring and fall when birds and spiders were subsidized by emerging aquatic insects, to the opposite direction in summer when fish were subsidized by terrestrial insects.  This work, which Mary Power of the University of California - Berkeley described as "...the best demonstration in any system, terrestrial, freshwater, or marine, of seasonal shifts of cross-habitat resource subsidies", is what led Nakano to interest both Power and Gary Polis, among the most prominent food web ecologists, in the research he and his colleagues were conducting in Japan.

As can be judged from his scientific productivity, Shigeru Nakano was an incredibly creative and hard working ecologist.  He combined a deep intuition about streams and fish borne of years of angling and snorkeling in the Hida Mountains of his boyhood, with the Japanese penchant for detail and accuracy, to measure and publish data on charr and salmon behavior and ecology that are better than any other I know of (e.g., Nakano 1994b, 1995a, 1995b).  Field research I conducted with Nakano and his graduate student collaborators in Hokkaido and Montana in 1991 and 1992 were intense expeditions where he taught me methods of capturing fishes (e.g., by angling while snorkeling, and using cast nets in streams) and measuring their behavior and microhabitat that I had never seen.  In turn, I taught him about field experiments and some field methods not known in Japan.  Nakano was also an excellent chef, having grown up in his parent's restaurant where large banquets were served.  After long days of cold snorkeling he could quickly prepare the most delicious traditional Japanese meals with only a gas stove.  While Director at Tomakomai Experimental Forest and later at CER he drove his graduate students and postdocs to be the best they could be, often by his own example.  He was famous for his all-night sessions with them to design research or complete the writing of manuscripts in English (no small feat for Japanese scientists), and often invited former students back for focused retreats to finish papers for publication (ronbun gasshuku, in Japanese).  Despite his strong personality he was also well liked by his students and staff for his warm humanity.  Shigeru Nakano's boundless energy and enthusiasm for life and his work was infectious, and he refused to be constrained by the strict Japanese hierarchical system.

Overlayed on this great skill as a field biologist and supervisor was Nakano's unique vision of a much broader picture about how stream fishes interacted with food webs to influence, and be influenced by, ecosystem processes.  In a sense he found a way to apply the Oriental philosophy of holism to his ecological research, and developed a series of studies with his students that investigated the complex interactions among members of both the aquatic and terrestrial food webs (e.g., Kuhara et al. 1999, in press, Miyasaka & Nakano 1999, Nakano et al. 1999a, 1999b, Kawaguchi & Nakano in press, Murakami & Nakano in press, Taniguchi & Nakano in press).  As of this writing, 12 more manuscripts are in review from this and other work by Nakano and his collaborators.

As the memories of our intensive field work, writing papers, and visits to various field research sites and laboratories in Japan and the U.S. during the past 13 years flood back into my mind, some of the most vivid are from 1994 when Nakano arranged for me to travel to Japan on a Research Fellowship from the Japanese Society for the Promotion of Science.  During our visit Nakano and his family hosted my family for several days of traveling in the mountains near his home town.  After the first day he could see that our two young children could not eat much of the strange food, such as the small squid that stared back at them from the tiny bowls.  When we reached Kamioka he took us to his parents' home, and for dinner they prepared a feast in their banquet room of as many Western foods as they could find including beef steak, fried pork and fish, bread, milk, and ice cream.  During these and other experiences our families developed a bond that led to sharing more visits, and children's books and toys.  Nakano's generosity and outgoing personality were well known by the many foreign scientists he and others hosted in Japan, and by everyone I spoke to that Nakano and his colleagues encountered at Berkeley and Davis during their final visit.

Shigeru Nakano had reached a pinnacle in his career at the age of only 37, which in the age-conscious Japanese academic system is barely old enough to qualify even the brightest and most productive scientists to become an Associate Professor.  His research had grown in scope from understanding how stream salmonids and African Rift Lake cichlids use and defend habitats, to work on several continents and at the scale of whole linked terrestrial-aquatic food webs in ecosystems.  His legacy is not only a unique body of ecological research that met the highest standards for science worldwide, but also a group of graduate students and scientists, including those like myself in other countries, that learned new ways of thinking about, and working in, stream ecosystems.  Along with many other aquatic ecologists in North America and elsewhere, I hope that Nakano's work can be continued by his collaborators in Japan and other countries.  All of us were privileged to have known and worked with him, and we will miss him greatly.
 

Acknowledgements

Yoshinori Taniguchi, Hitoshi Miyasaka, and Hiroya Kawanabe were very helpful in providing information for this tribute.
 

References cited

Noakes, D. L. G.  1989.  Symposium to be remembered.  Env. Biol. Fish. 24:313-317.
Ricklefs, R. E. & G. L. Miller.  2000.  Ecology, 4th edition.  W. H. Freeman & Company, New York, New York.  822 pp.
Yuma, M., I. Nakamura, & K. D. Fausch (ed.).  1998.  Fish biology in Japan: an anthology in honour of Hiroya Kawanabe.  Kluwer Academic Publishers, Dordrecht, The Netherlands.  405 pp.
 

Publications of Shigeru Nakano

 1. Nakano, S. & M. Nagoshi.  1985.  Density regulation and growth of a redspot masu-trout, Oncorhynchus rhodurus, in a mountain stream.  Physiol. Ecol. Japan 22:1-16.
 2. Nagoshi, M., S. Nakano & Y. Tokuda.  1988.  Changes in the utilization of microhabitat and food with growth of amago Oncorhynchus rhodurus in a Japanese mountain stream.  Nipp. Suisan Gakk. (Bull. Japan Soc. Sci. Fish.) 54:33-38 (in Japanese with English summary).
 3. Nakano, S., & T. Nakashima.  1989.  Notes on the reproduction of tilapiine fish, Oreochromis niloticus, in a hot spring area in Gifu Prefecture.  Res. Bull. Gifu Pref. Museum 10:9-12 (in Japanese with English summary).
 4. Nakano, S, S. Taguchi, Y. Shibata & T. Furukawa-Tanaka. 1989.  Red-spotted masu salmon.  pp. 169-179.  In: H. Kawanabe, N. Mizuno, and A. Sakurai (ed.) Freshwater Fishes of Japan, Yamatokeikokusha, Tokyo. (in Japanese)
  5. Nakano, S., T. Kachi, & M. Nagoshi.  1990.  Restricted movement of the fluvial form of red-spotted masu salmon, Oncorhynchus masou rhodurus, in a mountain stream, central Japan.  Jap. J. Ichthyol. 37:158-163
 6. Nakano, S., K. Maekawa & S. Yamamoto.  1990.  Change of the life cycle of Japanese charr following artificial lake construction by damming.  Nipp. Suisan Gakk. (Bull. Japan Soc. Sci. Fish.) 56:1901-1905.
 7. Nakano, S. & M. Nagoshi.  1990.  Brood defence and parental roles in a biparental cichlid fish Lamprologus toae in Lake Tanganyika.  Japan. J. Ichthyol. 36:468-476.
 8. Kitano, S., & S. Nakano.  1991.  Growth, sexual maturity and food habit of the Dolly Varden charr (Salvelinus malma) in the Horobetsu Stream, Shiretoko Peninsula.  Res. Bull. Shiretoko Mus. 13:1-12 (in Japanese with English summary).
 9. Nakano, S., T. Hino, S. Natsume, M. Hayashida, Y. Inaba, & A. Okuda.  1991.  Notes on nesting trees of Japanese flying squirrel, Pteromys volans orii, in Hokkaido during winter.  Res. Bull. Coll. Exper. Forests Hokk. Univ. 48:183-190 (in Japanese with English summary).
 10. Nakano, S., T. Kachi & M. Nagoshi.  1991.  Individual growth variation of red-spotted masu salmon, Oncorhynchus masou rhodurus, in a mountain stream.  Jap. J. Ichthyol. 38:263-270.
 11. Hino, T., & S. Nakano.  1992.  Breeding bird community of a deciduous broad-leaved forest in northern Hokkaido, Japan.  Res. Bull. Coll. Exper. Forests Hokk. Univ. 49:195-200 (in Japanese with English summary).
 12. Nakano, S.  1992a.  A preliminary report on food habit of Japanese huchen Hucho perryi in northern Hokkaido.  Biol. Inland Waters 7:20-23 (in Japanese with English summary).
 13. Nakano, S.  1992b.  Competition for space and coexistence in Japanese charr and masu salmon.  Asahi Animal Encyclopedia 87:76-77 (published by Asahi Shinbunsha, Tokyo, in Japanese).
 14. Nakano, S., K. D. Fausch, T. Furukawa-Tanaka, K. Maekawa & H. Kawanabe.  1992.  Resource utilization by bull char and cutthroat trout in a mountain stream in Montana, U.S.A.  Japan. J. Ichthyol. 39:211-217.
  15. Yamamoto, S., S. Nakano & Y. Tokuda.  1992.  Variation and divergence of the life-history of Japanese charr Salvelinus leucomaenis in an artificial lake-inlet stream system.  Japan. J. Ecol. 42:149-157 (in Japanese with English summary).
  16. Goto, A. & S. Nakano.  1993.  Distribution and ecology of freshwater fishes in Hokkaido.  pp. 113-126.  In: S. Higashi, A. Osawa & K. Kanagawa (ed.)  Biodiversity and Ecology in the Northernmost Japan.  Hokkaido Univ. Press, Sapporo.
 17. Iwamoto, Y., & S. Nakano.  1993.  An example of a check dam considering the natural environment.  Shinsabo 46:32-34 (In Japanese).
 18. Kitano, S., S. Nakano, M. Inoue, K. Shimoda & S. Yamamoto.  1993.  Feeding and reproductive ecology of exotic rainbow trout Oncorhynchus mykiss in the Horonai Stream in Hokkaido, northern Japan.  Nipp. Suisan Gakk. (Bull. Japan Soc. Sci. Fish.) 59:1837-1843 (in Japanese with English summary).
 19. Maekawa, K., T. Hino, S. Nakano & W. W. Smoker.  1993.  Mate preference in anadromous and landlocked Dolly Varden (Salvelinus malma) females in two Alaskan streams.  Can. J. Fish. Aquat. Sci. 50:2375-2379.
 20. Nakano, S., K. Shimoda, S. Kitano & M. Inoue.  1993.  Records of the river sculpin, Cottus amblystomopsis, and the flathead goby, Luciogobius guttatus, from the Shiretoko Peninsula, Hokkaido.  Res. Bull. Shiretoko Mus. 14:33-36 (in Japanese with English summary).
 21. Shimoda, K., S. Nakano, S. Kitano, M. Inoue & Y. Ono.  1993.  Present condition of stream fish assemblage in the Shiretoko Peninsula with special reference to human impacts.  Bull. Env. Sci. Hokk. Univ. 6:17-27 (in Japanese with English summary).
  22. Fausch, K. D., S. Nakano & K. Ishigaki.  1994.  Distribution of two congeneric charrs in streams of Hokkaido Island, Japan:  considering multiple factors across scales.  Oecologia 100:1-12.
 23. Inoue, M. & S. Nakano.  1994.  Physical environment structure of a small stream with special reference to fish microhabitat.  Japan. J. Ecol. 44:151-160 (in Japanese with English summary).
 24. Kitano, S., K. Maekawa, S. Nakano & K. D. Fausch.  1994.  Spawning behavior of bull trout in the upper Flathead drainage, Montana, with special reference to hybridization with brook trout.  Trans. Am. Fish. Soc. 123:988-992.
 25. Maekawa, K. & S. Nakano. 1994a. From anadromous to riverine life.  pp. 206-220.  In: A. Goto, K. Tsukamoto & K. Maekawa (ed.) Diadromous Fishes: Life History and Evolution, Tokai University Press, Tokyo (in Japanese).
 26. Maekawa, K. & S. Nakano.  1994b.  Non-oviposition of mature eggs by female fluvial red-spotted masu salmon.  Fish. Sci. (Japan) 60:37-39.
 27. Maekawa, K., S. Nakano & S. Yamamoto.  1994.  Spawning behaviour and size-assortative mating of Japanese charr in an artificial lake-inlet stream system.  Env. Biol. Fish. 39:109-117.
 28. Nakano, S.  1994a.  Fish that inhabit forests.  pp. 200-209. In: Mombusho (Ministry of Education, Science, Sport, and Culture)(ed.), School Textbook for Modern Japanese Literature, Kadokawa Shoten, Tokyo (in Japanese).
 29. Nakano, S.  1994b.  Variation in agonistic encounters in a dominance hierarchy of freely interacting red-spotted masu salmon (Oncorhynchus masou ishikawae).  Ecol. Freshwater Fish.  3: 153-158.
 30. Nakano, S. & T. Furukawa-Tanaka.  1994.  Intra- and interspecific dominance hierarchies and variation in foraging tactics of two species of stream-dwelling chars.  Ecol. Res. 9:9-20.
 31. Nakano, S. & K. Maekawa.  1994.  Charrs - life history variation and environmental alterations by human activities. pp.86-100.  In: A. Goto, K. Tsukamoto & K. Maekawa (ed.) Diadromous Fishes: Life History and Evolution, Tokai University Press, Tokyo (in Japanese).
 32. Yamamoto, S., N. Iida & S. Nakano.  1994.  Freshwater fish fauna in Rebun Island, Hokkaido.  Rishiri Studies 13:13-17 (In Japanese).
 33. Kitano, F., S. Nakano, K. Maekawa & Y. Ono.  1995.  Effect of stream temperatures on longitudinal distribution of fluvial Dolly Varden and potential habitat loss due to global warming.  Wildlife Conservation (Japan) 1:1-11 (in Japanese with English summary).
 34. Nakano, S.  1995a.  Competitive interactions for foraging microhabitats in a size-structured interspecific dominance hierarchy of two sympatric stream salmonids in a natural habitat.  Can. J. Zool. 73:1845-1854.
 35. Nakano, S.  1995b.  Individual differences in resource use, growth and emigration under the influence of  a dominance hierarchy in fluvial red-spotted masu salmon in a natural habitat.  J. Anim. Ecol. 64:75-84.
 36.  Nakano, S., M. Inoue, T. Kuwahara, T. Toyoshima, H. Hojyo, E. Fujito, H. Sugiyama, S. Okuyama, & K. Sasa.  1995.  Freshwater fish fauna in the Teshio and Nakagawa Experimental Forests and adjacent areas with reference to damming effects on their distribution.  Res. Bull. Hokk. Univ. Forests 52:95-109 (In Japanese with English summary).
 37. Nakano, S. & M. Kaeriyama.  1995.  Summer microhabitat use and diet of four sympatric stream-dwelling salmonids in a Kamchatkan stream.  Fish. Sci. (Japan) 61:926-930.
 38. Kurashige, Y., T. Toyoshima, & S. Nakano.  1996.  Topographical change in a channel bed after habitat improvement.  Jap. J. Limnol. 57:193-197.
 39. Nakano, S., F. Kitano & K. Maekawa.  1996.  Potential fragmentation and loss of thermal habitats for charrs in the Japanese Archipelago due to climatic warming.  Freshwater Biol. 36:711-722.
 40. Nakano, S., & Y. Taniguchi.  1996.  Interspecific competition and coexistence in freshwater-dwelling salmonids: a review.  Jap. J. Ichthyol. 43:59-78 (In Japanese).
 41. Takami, T., F. Kitano & S. Nakano.  1996.  High water temperature influences on foraging responses and thermal deaths of Dolly Varden Salvelinus malma and white-spotted charr S. leucomaenis in a laboratory.  Fish. Sci. (Japan) 63:6-8.
 42. Taniguchi, Y., H. Urabe, & S. Nakano.  1996.  Natural reproduction of coho salmon Oncorhynchus kisutch population introduced in a pond-associated stream in Hokkaido, Japan.  Fish. Sci. (Japan) 62:992-993.
 43. Toyoshima, T., S. Nakano, M. Inoue, Y. Ono & Y. Kurashige.  1996.  Fish population responses to stream habitat improvement in a concrete-lined channel.  Jap. J. Ecology 46:9-20 (in Japanese with English summary).
 44. Yamamoto, S., & S. Nakano. 1996.  Growth and development of a bimodal length-frequency distribution during smolting in a wild population of white-spotted charr in northern Japan.  J. Fish Biol. 48:68-79.
 45. Fausch, K. D., S. Nakano, & S. Kitano. 1997.  Experimentally induced foraging mode shift by sympatric charrs in a Japanese mountain stream.  Beh. Ecol. 8:414-420.
 46. Inoue, M., S. Nakano, & F. Nakamura.  1997.  Juvenile masu salmon (Oncorhynchus masou) abundance and stream habitat relationships in northern Japan.  Can. J. Fish. Aquat. Sci. 54:1331-1341.
   47. Fausch, K. D. & S. Nakano. 1998.  Research on fish ecology in Japan: a brief history and selected review.  Env. Biol. Fish. 52:75-95.
 48. Inoue, M. & S. Nakano.  1998.  Effects of woody debris on the habitat of juvenile masu salmon (Oncorhynchus masou) in northern Japanese streams.  Freshwater Biol. 40:1-16.
 49. Nakano, S., S. Kitano, K. Nakai, & K. D. Fausch.  1998.  Competitive interactions for foraging microhabitat among introduced brook charr, Salvelinus fontinalis, and native bull charr, S. confluentus, and westslope cutthroat trout, Oncorhynchus clarki lewisi, in a Montana stream. Env. Biol. Fish. 52:345-355.
 50. Nisikawa, U. & S. Nakano.  1998.  Temporal variation in foraging group structure of a size-structured stream fish community.  Env. Biol. Fish. 52:357-370.
 51. Urabe, H. & S. Nakano.  1998.  Contribution of woody debris to trout habitat modification in small streams in secondary deciduous forest, northern Japan.  Ecol. Res. 13:335-345.
 52. Usio, N. & S. Nakano.  1998.  Influences of microhabitat use and foraging mode similarities on intra- and interspecific aggressive interactions in a size-structured stream fish assemblage.  Ichthyol. Res. 45:19-28.
 53. Inoue, M. & S. Nakano.  1999.  Habitat structure along channel-unit sequences for juvenile salmon: a subunit-based analysis of in-stream landscapes.  Freshwater Biol. 42:597-608.
 54. Kuhara, N., S. Nakano & H. Miyasaka.  1999.  Interspecific competition between two stream insect grazers mediated by non-feeding predatory fish.  Oikos 87:27-35.
 55. Maekawa, K., S. Nakano & K. Iguchi.  1999.  Changes in freshwater fish distributions due to global warming.  pp. 204-218.  In: S. Kawano & O. Imura (ed.) Populations in Changing Environments, Kaiyusha, Tokyo (in Japanese).
 56. Miyasaka, H. & S. Nakano.  1999.  Effects of drift- and benthic-foraging fish on the drift dispersal of three species of mayfly nymphs in a Japanese stream.  Oecologia 118:99-106.
 57. Nakano, S.  1999.  Diet differentiation in polymorphic Brachymystax lenok in streams of southern Primor'e, Russia.  Ichthyol. Res. 46:100-102.
 58. Nakano, S., K. D. Fausch & S. Kitano.  1999.  Flexible niche partitioning via a foraging mode shift: a proposed mechanism for coexistence in stream-dwelling charrs.  J. Anim. Ecol. 68: 1079-1092.
 59. Nakano, S., Y. Kawaguchi, Y. Taniguchi, H. Miyasaka, Y. Shibata, H. Urabe, & N. Kuhara.  1999.  Selective foraging on terrestrial invertebrates by rainbow trout in a forested headwater stream in northern Japan.  Ecol. Res. 14:351-360.
 60. Nakano, S., H. Miyasaka & N. Kuhara.  1999.  Terrestrial-aquatic linkages: riparian arthropod inputs alter trophic cascades in a stream food web.  Ecology 80:2435-2441.
 61. Urabe, H., & S. Nakano.  1999.  Linking microhabitat availability and local density of rainbow trout in low-gradient Japanese streams.  Ecol. Res. 14:341-349.
 62. Saito, T., & S. Nakano.  1999a.  Differences in the impacts of a weir on the reproductive activities in white-spotted charr and Dolly Varden in a Japanese pond-associated stream system.  Fish. Sci. (Japan) 65:898-903.
 63.  Saito, T., & S. Nakano.  1999b.  Reproductive-timing-dependent alteration of offspring life histories in female threespine sticklebacks.  Can. J. Zool. 77:1314-1321.
 64. Kuhara, N., S. Nakano, & H. Miyasaka.  2000.  Flow rate mediates the competitive influence of a grazing caddisfly on mayflies.  Ecol. Res. 15:145-152.
 65. Murakami, M. & S. Nakano.  2000.  Species-specific bird functions in a forest canopy food web.  Proc. Royal Soc. London B 267:1597-1601.
 66. Taniguchi, Y., Y. Miyake, T. Saito, H. Urabe, & S. Nakano.  2000.  Redd superimposition by introduced rainbow trout, Oncorhynchus mykiss, on native charrs in a Japanese stream.  Ichthyol. Res. 47:149-156.
 67. Taniguchi, Y. & S. Nakano.  2000a.  Complex effects of global warming and local environmental disturbance on freshwater fish communities: the mechanisms, predictions, and repercussion effects.  Jap. J. Limnol. 61:79-94 (in Japanese with English summary).
  68. Taniguchi, Y. & S. Nakano.  2000b.  Condition-specific competition: implications for the altitudinal distribution of stream fishes.  Ecology 81:2027-2039.
 69. Kawaguchi, Y., & S. Nakano.  in press.   The contribution of terrestrial invertebrates to the annual resource budget for salmonids in forest and grassland reaches of a headwater stream.  Freshwater Biol.
 70. Nakano, S., H. Miyasaka & T. Furukawa-Tanaka.  in press.  Food resource divergence between white-spotted charr and masu salmon in Japanese mountain streams: circumstantial evidence for competition.  Ecol. Res.
 

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