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Ecological Notes of the Fishes and the Interspecific Relations Among Them in Lake Biwa

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Ecological Notes of the Fishes and the Interspecific Relations Among Them in Lake Biwa Ecological Notes of the Fishes and the Interspecific Relations among Them in Lake Biwa Taizo MIURA Introduction Lake Biwa has been much interested by biologists, specially by ichthiologists because of the richness and variety of the lake fish fauna and of their diverse origin. Therefore, a good number of papers on taxonomic researches of the fishes have been published. However ecological studies of the fishes have been poorly made. Freshwater environments as well as terrestrial ones have been practically reformed, accompanied with development of human society and specially of industries and agriculture. Lake Biwa is also not an exception. Recently the ministry of construction is planning to construct a dam between the north basin and the south basin of the lake, and to pump up water from the north basin to the south one for the purpose of holding a continuous water supply to the outlet river. By such programme the two basins will nearly be disconnected and the water level of the north basin is often expected to be lowered. It can be easily expected that production of the organisms in the lake will be affected by the reformation, so that it 'is important to predict how the organic production will be affected, and to propose how the programme should be made in order to benefit the local people, especially the fishermen. Thus, about sixty scientists including biologists, physicists, chemists and agricultural economists had formed a co-operative research team and have investigated the biotic re- sources of the lake from 1962 to 1965. Through this investigation ecological studies of some of the fish species have been intensively made. This paper presents, in the first place, descriptions of ecological elements of the fishes including spawning, habitat, feeding and growth etc., and introductions of some other ecological works. Secondly, by comparison of their ecologies, Contribution from the Animal Ecological Research Group, Department of Zoology, Kyoto University. Contribution from the Otsu Hydrobiological Station, Foreign Language Series, No. 182. Contribution from the B. S. T. (Research Group for Biotic Resources in Lake Biwa), No. 64. 50 interspecific relationships are considered. The materials presented here are inadequate for discussing the fish community of the lake, however, the author believes that this article is usable as a step of further investigation of the fishes in this lake. History and environmental descriptions of the lake Lake Biwa (35•‹15'N., 136•‹05'E.), the largest lake in Japan, is situated just on the isthmus which seems to be the distored portion of arcs building the Honshu Island (HORIEE, 1961). Through analysis of Post-Pliocene sediments around Lake Biwa and of fossils from the deposits, TAKAYA (1963) has established the follow- ing three paleolimnological stages of of Paleo-Biwa Lake ; "Old Closed Lake", "Open Lake" "Young Closed Lake" . According to TAKAYA (1963) the first stage of late Pliocene age has provided the immigration of the fauna and flora of continental element to the lake under a warm climate. At the second stage of the earliest Pleistocene, the lake, "Open Lake", had a broad channel leading to the sea at the southwestern end of the lake without any distinct topographicar barriers, though it was thoroughly lacustrine. The channel is supposed to have prepared a quite different condition for marine dwellers and to have given them the opportunity of invasion into the lake. When the channel was disappeared by a very strong crustal movement at Early Pleistocene the third stage started and has continued till the present time. Perhaps, almost all the species which had invaded before and during the channel stage were compelled to become perfect land-locked species. Passing through several occasions of water levelling which might be responsible for terrace formations during the "Young Closed Lake" period, the present lake basin had been formed. The lake is about 68 km long and 22.6 km in width. The surface area is approximately 674.4 km2, the length of shore line 188 km, the maximum depth 104 m, the volume 22.6 km3. The lake is devided into two basins; the north basin and south one. The former occupies approximately 92% in area and 99% in volume. The latter is small and shallow, the maximum depth 7.5 m and mean depth 2.8 m. The lake has numerous inlet streams which is occasionally dried up in dry periods. The main tributaries are located around the north basin. There is only one outlet, the Uji River, which has a high control dam at the southwestern end of the south basin. 51 According to MORIKAWA and OKAMOTO (1963) and MORIKAWA (1964), the surface temperature of offshore water seldom rises 30•Ž in summer. The lake begins to show thermal stratification in May and thermocline gradually becomes remarkable toward August (Fig. 1). It develops between the depths of 10 m and 30 m in summer. After then it is gradually reduced by cooling of the surface water and disappears in the end of November. In winter and spring thermal stratification entirely disappears and the water temperature is nearly the same from the surface to the lake bottom (Fig.2). The lowest temperature of offshore water seldom shows less than 6•Ž in the northern basin. FOJINAOAet al. (1966) indicates that even during the summer stagnation a fair amount of dissolved oxygen can be observed in the hypolimnion, and oxygen deplession never observed in the entire area of the south basin and of the north basin. The same authors describe that the chemical nature of the north basin is intermediate between oligotrophic and mesotrophic and of the south basin is eutrophic, as far as contents of nutrients are considered. Such differences of chemical nature as well as lake topography more or less reflect upon organisms. Density of phyto-and zooplankters are observed greater in the south basin than in the north basin, with the exception of winter. Typical example can be seen in larger aquatic plants. IKUSHIMAet al. (1962) state that the small bottom area of the south basin which is only 8.5 % of the total area sustains 40% of the total yield of aquatic vegetation. Fish fauna of the lake Lake Biwa and its tributaries have been recorded to contain 62 fish species and 2 subspecies. However, during the intensive investigation of the lake by the Research Group for Biotic Resources in Lake Biwa, 49 species and 2 subspecies observed as the inhabitants of the lake today (Table 1). Among these species are several that include endemic ones, namely, nigorobuna, gengorobuna, hon- moroko, hasu, iwatokonamazu, biwako-onamazu and isaza. Hasu and isaza have been considered as a relict (UENO, 1943, HORIE 1961, TOMODA1963 a, b), while the rests may be due to autochthonous intra-lacustrine speciation (TOMODA1962,1963a, b, 1965). 52 Table 1. Species list of the fishes in Lake Biwa. 53 54 Ecological notes of the fishes in the lake Ecological notes of 9 species, which have been intensively investigated in recent years because of their ecological as well as economic importance will be described below. Besides these, brief notes of spawning period, spawning place, growth, habitat and food of fairly abundant species are listed in Table 2. Biwa-masu (Oncorhynchus rhodurus) Biwa-masu appear to . spawn in inlet streams. Spawning occurs from the middle of October until late November. The main spawning streams are the River Ado, R. Ishida, R. Chinai, R. Shiozu-okawa, R. Ane and R. Inukami. The majority of spawning migrants are caught by weirs and traps before they arrive at the spawning place. Eggs of the spawners caught are artificially fertilized before marketing and the eggs are sent to the hatchery. The youngs reared in the hatchery are stoecked into the lake in spring of the succeeding year. It is likely, therefore, that the majority of recruits are hatchery fish. The main part of the male is covered by 2+ and 3+ year fish and of the female by 3+ and 4+ ones. The 5+ spawners are extremely rare (MIURA,unpublished M.S.) In warm season, both the youngs and adults inhabit in the hypolimnion and in cold season they expand the habitat into the epilimnion. The result of growth estimation by the scale method indicates that mean body length attains 24.0 cm by the second year., 36. 5 cm by the third and 42.0 cm by the fourth (MIURA,unpublished M. S.). The types of food organisms vary with the size of the fish. The youngs eat planktonic crustaceans and gammarids, while the adults take small fishes such as ayu and isaza (Shiga Pref. Fish. Res. Lab. 1942). Ayu (Plecoglossus altivelis) AZUMA(1964) summarized many features of the ecology of ayu in Lake Biwa. He reveals that there exist two groups of ayu which differ in their mode of life as well in morphological characters. Either of them spawn in the inlet streams in Autumn (September-October) and die there. The larvae hatched out drift with the current toward the lake at night. After then they appear on the coastal bottom near the streams. In October the larvae (40 mm BL) can be collected at night in the surface water of the off-shore area. Further developed larvae (50 mm EL) seem to have a habit of aggregation around the bottom layer of shelf during the daytime and migrate toward the inshore area during the 55 nighttime. Such distributional and migratory patterns are continued untill late winter. Food organisms observed in the alimentary tract of the fry and juvenile are water fleas and copepods. The fish of these stages likely feed on these zoopla- nkters during the daytime and unlikely feed during the nighttime, because the majority of alimentary tracts of the fish caught during the daytime are filled with the zooplankters but of the fish caught at night at the inshore area are entirely empty.
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