Exploring particulate foraging as adaptive tactics amidst competitive grazing dynamics in Ryukyu-ayu Plecoglossus altivelis ryukyuensis

We examined monthly occurrence patterns and feeding habits of larvae and juveniles of the critically endangered Ryukyu-ayu Plecoglossus altivelis ryukyuensis in the lower reach of the Kawauchi River, Amami-oshima Island, southern Japan, for two successive years. The study area was classified into a brackish lake (Lake Uchiumi) and a coastal zone (Sumiyo Bay) based on monthly variations in water temperature and salinity. Larvae and juveniles occurred from December to April in the first year and from January to February in the second year, respectively. Smaller-sized and earlier developmental-staged P. altivelis ryukyuensis individuals tended to emerge in larger numbers in Lake Uchiumi than in Sumiyo Bay. The present data suggest that Lake Uchiumi, where the physical environment is characterized by lower temperature and salinity than in Sumiyo Bay, would be suitable as a nursery ground for this fish, especially for earlier developmental-staged larvae. No individuals ≥25.0 mm body length, which is the size at which they begin their upstream migration, occurred in Lake Uchiumi or Sumiyo Bay in April or May, when upstream migration is assumed to reach a peak based on previous studies, in either year. The peak of upstream migration, therefore, may have occurred several months earlier in the years covered in this study due to higher water temperature than in typical years. To assess the feeding habits of this fish, the gut contents were observed. In both areas, copepods were abundant in P. altivelis ryukyuensis diet with calanoids and Oithona spp. being the most abundant. The diet composition differed between Lake Uchiumi and Sumiyo Bay and clearer ontogenetic variation in the composition of the diet was observed in Lake Uchiumi. Small individuals fed on tintinnid ciliates, but the proportion of these organisms in the diet gradually decreased with growth. Large individuals fed exclusively on harpacticoid copepods and insects including demersal species. Overall, larval and juvenile P. altivelis ryukyuensis exhibited a generalist feeding habit that tended to increase with growth. It is indispensable to adequately manage the environmental conditions in the lower reach of the Kawauchi River, especially Lake Uchiumi, to conserve P. altivelis ryukyuensis population.

1912～2007年の96年間に出版された文献に基づき, 日本産アユPlecoglossus altivelis altivelisの寄生虫に関する情報を寄生虫-宿主リストに整理して目録を作成した。本目録には, 29種の寄生虫(微胞子虫類1種, ミクソゾア類1種, 吸虫類13種, 単生類3種, 条虫類1種, 線虫類1種, 鉤頭動物3種, カイアシ類4種, エラオ類2種)に加えて, 学名がまだ決定していない寄生虫の情報が含まれる。寄生虫-宿主リストでは, 各寄生虫は高位分類群ごとに配列され, 最新の学名, シノニム, 寄生部位, 地理的分布および報告者の情報が示されている。アユの亜種, リュウキュウアユPlecoglossus altivelis ryukyuensisからは寄生虫の記録はない。ミクソゾア類のMitraspora plecoglossiをHoferellus属に移すとともに, ハスコウトウチュウAcanthocephalus opsariichthydis Yamaguti, 1935の学名が動物命名規約に則って正しい綴りと命名年であることを確定した。また鉤頭動物のEchinorhynchus oblitusをカジカコウトウチュウEchinorhynchus cottiのシノニムとした。

Evaluation of the Genetics of Growth and Resistance to Vibriosis in Ryukyu Ayu Plecoglossus altivelis ryukyuensis by Hybridization

Cultured Ryukyu ayu Plecoglossus altivelis ryukyuensis was found to be infected with a microsporidian parasite on Amami Oshima Island in January 2009. The parasite formed numerous cysts on the visceral organs and skin of the diseased fish. High temperature seemed to suppress the development of the parasite; in 2009 and 2010, the disease occurred in the 0-year fish from April to August, and afterwards the prevalence decreased dramatically in September and October. These characteristics well resembled glugeosis in ayu P. altivelis altivelis in other areas of Japan. This parasite seems to be indigenous to Amami Oshima Island, because it was detected in the museum specimens of wild Ryukyu ayu caught in 1991. Average dimensions of spores from Ryukyu ayu were slightly shorter and wider than those of Glugea plecoglossi, but the ranges of the measurements were overlapped. From the above characteristics and molecular analysis of SSU rRNA, it was concluded that the present microsporidian was included genus Glugea, but the specific identity remained to be clarified.

Maintaining a certain number of spawning grounds for endangered fish is particularly important in rivers and streams where the environment changes drastically through floods and exploitations. Ryukyu-ayu Plecoglossus altivelis ryukyuensis is an endangered annual fish inhabiting the few streams and rivers flowing through the Amami-Oshima Island, southern Japan, and spawns in the lower reaches at sites with the specific riverbed conditions. In the island, resident volunteers along with researchers modify riverbed manually to provide conditions suitable for the natural spawning of the fish every year. This study investigated the effects of these activities on the riverbed conditions and consequently on the spawning by Ryukyu-ayu in two rivers. Our results showed that riverbed hardness and the amount of silt significantly decreased in the modified areas, but not in the adjacent unmodified areas. In one of the two rivers, the modification suppressed the settlement of net-spinning caddisfly larvae Stenopsyche schmidi that promote consolidation of the riverbed. The riverbed conditions in the modified areas were similar to those in the sites where P. a. ryukyuensis indeed spawned. These results suggest that manual modification is efficient in improving riverbed conditions for the spawning of Ryukyu-ayu. However, the eggs of the fish were not found at the modified areas in both rivers. It will be necessary to develop effective and practical ways of selecting sites for modification without obstructing the natural spawning in this fish.

The occurrence and body size of drifting land-locked Ryukyu-ayu, Plecoglossus altivelis ryukyuensis, larvae were investigated in the San-numata River, Okinawa-jima Island from November 2000 to March 2001. The water temperature in the river fluctuated from 13.8 to 17.9°C during the sampling period. The drifting larvae occurred from mid-December to mid-March with an estimated spawning peak during late November and early December. The notochord length of the drifting larvae ranged from 4.9 to 6.7 mm (5.7 ± 0.3 mm, mean ± SD), and became longer as the spawning months progressed. These results imply that the land-locked population possesses the same maturation attributes as the original amphidromous population.

ABSTRACT: The development of embryos, larvae and juveniles of the Ryukyu-ayu, Plecoglossus altivelisryukyuensis are described based on laboratory-reared specimens.The eggs were spherical, 0.98–1.18 mm (mean 1.06 mm)in diameter with an adherent membrane. The incubation period after fertilizationwas approximately 155 h at a water temperature of 19.7–22.0°C(mean 20.7°C). Newly hatched larvae were 5.0–5.9 mm(mean 5.5 mm) in body length (BL) with 59–62 myotomes.Within 5 days after hatching, the larvae had attained 6.4–8.2 mm(mean 7.8 mm) BL and had completely consumed their yolk.Notochord flexion began at 13.7 mm BL and was completedby 16.3 mm BL. The rudimental dorsal, anal, pelvic andadipose fins appeared at 14.7, 16.3, 21.8 and 21.8 mm BL, respectively.All fin rays reached the same fixed number of adult fish at about28 mm BL. The comb-like teeth began to form at approximately30 mm BL and were fully developed at about 40–50 mm BL.The proportions of P. a. ryukyuensis specimens,which were consistent with adult fish, occurred at approximately40 mm BL.

The countable characters of Qingdao, Zhejiang, Liaoning and Japanese Ayu are compared. There are no substantial different characters among the former three Ayu, so the comprehensive values of them are taken as representative of Chinese Ayu, which is in turn compared with Japanese Ayu (Plecoglossus altivelis altivelis and Plecoglossus altivelis ryukyuensis). By the coefficient of difference test, 3 and 4 characters are found to be beyond the subspecies level between Chinese Ayu and nominate subspecies (Plecoglossus altivelis altivelis) and between Chinese Ayu (Plecoglossus altivelis chinensis) and Ryukyus subspecies (Plecoglossus altivelis ryukyuensis), respectively, which shows that they belong to different subspecies. The China mainland Ayu is a new subspecies, Plecoglossus altivelis chinensis Wu & Shan, the establishment of which and its forming causes are discussed.

Survival rates of larval Ryukyu-ayu Plecoglossus altivelis ryukyuensis under differing experimental conditions of temperature and salinity

Seasonal appearance and size of newly-hatched larvae of Ryukyu-ayu Plecoglossus altivelis ryukyuensis in the Yakugachi and Kawauchi Rivers, Amami-oshima Island, southern Japan

The behavior of Ryukyu-ayu Plecoglossus altivelis ryukyuensis larvae during downstream migration from the Yakugachi River flowing into Sumiyo Bay, Amamioshima Island, southern Japan.

The diet shift of larval and juvenile Ryukyu-ayu Plecoglossus altivelis ryukyuensis was investigated for the landlocked population stocked in the Fukuji Reservoir, Okinawa Island. Gut contents of 274 specimens (5.1–31.4 mm in body length) including 31 yolk-sac larvae (5.7–8.0 mm) were examined. The diet of larval and juvenile landlocked P. a. ryukyuensis was mainly composed of zooplankton such as dinoflagellates, rotifers, copepods, and cladocerans. Feeding incidence was 48% in total and it gradually increased with growth. Diet analysis clearly demonstrated a diet shift during the larval and juvenile periods. For yolk-sac larvae and 5.0–19.9 mm fish, the diet was dominated by dinoflagellates Gymnodinium sp., and Peridinium sp., both numerically and by frequency, followed by rotifers that mainly consisted of Polyarthra spp. Diet breadth (Levin’s standardized index BA) diversified with growth, and feeding on copepods and cladocerans gradually increased in 10.0–14.9 mm fish. Diet overlap (Morisita’s index: C) showed that diet drastically changed in 20.0–24.9 mm fish with a specialization on cladocerans Bosminopsis deitersi and Diaphanosoma brachyurum. This diet shift probably corresponded with an increase of feeding ability resulting from an enlargement of the mouth, increased swimming ability, and the development of sense organs.

Variation in the life cycle of diadromous fishes can be explained by differential food availability between marine and freshwater habitats, since migration is often interpreted as a mechanism for exploiting food resources. Theoretically, a migration pattern of mainly remaining in freshwater occurs in tropical and subtropical habitats where fluvial productivity possibly exceeds marine productivity. However, in Yakugachi River, Amami-Oshima Island, southern Japan, low nutrient concentrations in the river suggest that food availability is limited for the subtropical Ryukyu-ayu Plecoglossus altivelis ryukyuensis. Since Ryukyu-ayu is an amphidromous fish that mainly grows in rivers after spending 2 months in the sea, limited food availability in rivers would force this species to migrate to other habitats with better food availability. Otolith increment and Sr:Ca analyses of 48 adult Ryukyu-ayu collected from the Yakugachi River revealed that all individuals visited estuaries more than three times after moving upstream. Although the specific growth rates of this species in the river had no correlation with the salinity profile in the fluvial period, this movement may be an adaptive choice because the salinity profile significantly affected the body size at maturity. Our results highlighted individual-based variations in amphidromous migration for utilizing estuaries, which could be explained by relatively higher productivity in estuarine than in freshwater and marine habitats.

In dispersive species with continuous distributions, genetic differentiation between local populations is often absent or subtle and thus difficult to detect. To incorporate such subtle differentiation into management plans, it may be essential to analyse many samples from many localities using adequate numbers of high-resolution genetic markers. Here, we evaluated the usefulness of dense locality sampling in resolving genetic population structure in the ayu (Plecoglossus altivelis), a dispersive fish important in Japanese inland fisheries. Genetic variability in, and differentiation between, ayu populations around the Japan-Ryukyu Archipelago were investigated in 4746 individuals collected from 120 localities by genotyping 12 microsatellite markers. These individuals represented the two subspecies of ayu, namely the Ryukyuan subspecies (Plecoglossus altivelis ryukyuensis) and both amphidromous and landlocked forms of the nominotypical subspecies (P.a.altivelis) along the archipelago. We successfully detected an absence of genetic differentiation within the landlocked form and subtle but significant differentiation and clear geographic patterns of genetic variation among populations of the amphidromous form, which had been considered genetically homogeneous. This suggests that dense locality sampling effectively resolves subtle differences in genetic population structure, reducing stochastic deviation in the detection of genetic differentiation and geographic patterns in local populations of this dispersive species. Resampling analyses based on empirical data sets clearly demonstrate the effectiveness of increasing the number of locality samples for stable and reliable estimations of genetic fixation indices. The genetic population structure observed within the amphidromous form provides useful information for identifying management or conservation units in ayu.

In lower vertebrates, the bone-mineralizing hormone "calcitonin" is secreted from ultimobranchial glands, which assume various shapes, e.g., follicles, cellular masses, or cell strands. Histological observations support the view that in some teleosts, the glands increase in size when females maturate. We determined the exact volume of the gland in juveniles and adult males and females of a teleost, Ryukyuayu (Plecoglossus altivelis ryukyuensis). Furthermore, we examined plasma Ca, Na and K levels. In this species, the gland was fundamentally composed of a single follicle. The gland volume was converted to numerical data under a certain condition. It thus became clear that the value of the follicle wall was significantly increased only in maturing and mature females with high plasma Ca levels and that the value of the lumen did not show any statistically significant changes during growth and maturation.