Yoichiro Sakai

Phytoplankton community reorganization driven by eutrophication and warming in Lake Biwa

We compiled and analyzed long-term data, including chemical, physical and phytoplankton community data, for the Lake Biwa ecosystem from 1962 to 2003. Analyses on environmental data indicate that Lake Biwa had experienced intensified eutrophication (according to total phosphorus concentration) in the late 1960s and returned to a less eutrophic status around 1985, and then exhibited rapid warming and thus increased water column stability since 1990. Total phytoplankton cell volume largely followed the trend of total phosphorus concentration, albeit short-term fluctuations existed. However, phytoplankton community shifted dramatically in response to those changes of environmental states. These shifts were cause by changes in trophic status driven by phosphorus loadings and physical properties in the water column driven by warming. Moreover, most phytoplankton species did not show a strong linear correlation with environmental variables, suggesting nonlinear transitions among different states.

Is the relationship between body size and trophic niche position time-invariant in a predatory fish? First stable isotope evidence.

Characterizing relationships between individual body size and trophic niche position is essential for understanding how population and food-web dynamics are mediated by size-dependent trophic interactions. However, whether (and how) intraspecific size-trophic relationships (i.e., trophic ontogeny pattern at the population level) vary with time remains poorly understood. Using archival specimens of a freshwater predatory fish Gymnogobius isaza (Tanaka 1916) from Lake Biwa, Japan, we assembled a long-term (>40 years) time-series of the size-dependence of trophic niche position by examining nitrogen stable isotope ratios (δ 15N) of the fish specimens. The size-dependence of trophic niche position was defined as the slope of the relationship between δ 15N and log body size. Our analyses showed that the slope was significantly positive in about 60% of years and null in other years, changing through time. This is the first quantitative (i.e., stable isotope) evidence of long-term variability in the size-trophic relationship in a predatory fish. This finding had implications for the fish trophic dynamics, despite that about 60% of the yearly values were not statistically different from the long-term average. We proposed hypotheses for the underlying mechanism of the time-varying size-trophic relationship.

Carbon and nitrogen stable isotope ratios of macroinvertebrates in the littoral zone of Lake Biwa as indicators of anthropogenic activities in the watershed

Carbon and nitrogen stable isotope ratios (δ13C and δ15N) of macroinvertebrates inhabiting littoral zones of lakes can serve as useful indicators of material loading from the watershed. We collected snails (Semisulcospira spp.) and bivalves (Unio douglasiae biwae Kobelt) from 29 littoral sites in Lake Biwa near the mouths of river tributaries with various human population density (HPD) and land-use patterns. The δ13C and δ15N signatures were determined for three potential food sources: particulate organic matter in the pelagic zone (PPOM), riverine particulate organic matter from tributaries (RPOM) and epilithic organic matter in the littoral zone (EOM). The stable isotope mixing model revealed that snails relied mainly on EOM, and bivalves on PPOM and RPOM. Multiple regression analysis showed that intersite variation in δ15N for snails was best explained by HPD, while variation in δ15N of EOM and nitrate was explained to a lesser extent by HPD. Comparison with isotope signatures of their food sources and riverine nutrients revealed that snails assimilated anthropogenic nitrogen from wastewater in the watershed. Our results also showed that the δ13C value of bivalves was marginally related to the fraction of paddy fields in the watersheds. In conclusion, the isotope signatures of macroinvertebrates inhabiting the littoral zone can be useful indicators of anthropogenic impacts from the watershed.

Long-Term Changes in the Diet of Gymnogobius isaza from Lake Biwa, Japan: Effects of Body Size and Environmental Prey Availability

Body size and environmental prey availability are both key factors determining feeding habits of gape-limited fish predators. However, our understanding of their interactive or relative effects is still limited. In this study, we performed quantitative dietary analysis of different body sizes of goby (Gymnogobius isaza) specimens collected from Lake Biwa between 1962 and 2004. First, we report that the diet was composed mainly of zooplankton (cladocerans and copepods) before the 1980s, and thereafter, shifted to zoobenthos (gammarids). This foraging shift coincided with, and thus can be linked to, known historical events in the lake at that time: decrease in zooplankton abundance with the alleviation of eutrophication, increase in fish body size resulting from fish population collapse, and increase in gammarid abundance due to reduced fish predation pressure. Supporting this view, our data analyses revealed how the long-term changes in the diet composition would be co-mediated by changes in fish body size and environmental prey availability. Specifically, while zoobenthos abundance strongly affected the fish diet composition, larger (smaller) fish preferred zoobenthos (zooplankton). Furthermore, the body size effects were stronger than those of prey availability. These results provide the best long-term evidence that fish feeding habits vary over decades with its body size and prey community due to anthropogenic disturbances.

Mountain-climbing bears protect cherry species from global warming through vertical seed dispersal

In a warming climate, temperature-sensitive plants must move toward colder areas, that is, higher latitude or altitude, by seed dispersal [1]. Considering that the temperature drop with increasing altitude (-0.65°C per 100 m altitude) is one hundred to a thousand times larger than that of the equivalent latitudinal distance [2], vertical seed dispersal is probably a key process for plant escape from warming temperatures. In fact, plant geographical distributions are tracking global warming altitudinally rather than latitudinally, and the extent of tracking is considered to be large in plants with better-dispersed traits (e.g., lighter seeds in wind-dispersed plants) [1]. However, no study has evaluated vertical seed dispersal itself due to technical difficulty or high cost. Here, we show using a stable oxygen isotope that black bears disperse seeds of wild cherry over several hundred meters vertically, and that the dispersal direction is heavily biased towards the mountain tops. Mountain climbing by bears following spring-to-summer plant phenology is likely the cause of this biased seed dispersal. These results suggest that spring- and summer-fruiting plants dispersed by animals may have high potential to escape global warming. Our results also indicate that the direction of vertical seed dispersal can be unexpectedly biased, and highlight the importance of considering seed dispersal direction to understand plant responses to past and future climate change.

Seasonal and site-specific variability in terrigenous particulate organic carbon concentration in near-shore waters of Lake Biwa, Japan

Identifying sources of particulate organic matter (POM) is important for clarifying fundamental mechanisms by which lake food webs are sustained. We determined carbon and nitrogen stable isotope ratios of POM in near-shore waters of Lake Biwa, a large, meso-eutrophic lake in Japan, to estimate relative contributions of terrigenous particulate organic carbon (T-POC), plankton-derived POC (P-POC) and epilithon-derived POC (E-POC) to POC in near-shore waters. Samples were collected during different months (November, February, May and July) at 29 sites located near the mouth of tributary rivers with different discharge and catchment land use. The data revealed that POC mainly consisted of P-POC and T-POC, with relative contributions varying widely over season and among locations. E-POC generally contributed little to the near-shore POC. Path analyses revealed that the concentration of riverine POC whose isotopic signatures were similar to those of rice straws increased with a larger %paddy field area in the catchment of tributary rivers, which subsequently enhanced T-POC inputs to near-shore waters through riverine transportation. Furthermore, our results suggested that T-POC contribution was influenced, with a time lag, by wave-driven turbulence and shore topography, which appear to affect sedimentation and resuspension of T-POC.

Food web properties of the recently constructed, deep subtropical Fei-Tsui Reservoir in comparison with the ancient Lake Biwa

Using carbon and nitrogen stable isotope analysis, we characterised food web properties of the deep subtropical Fei-Tsui Reservoir (FTR), which was recently altered from a lotic to a lentic system after dam construction. In the littoral zone, zoobenthos showed strong reliance (83.9%) on benthic algal production. Zoobenthos were never found in the profundal zone because of anoxia. Zooplankton depleted 13C more than that of particulate organic matter as their putative food source, suggesting a contribution of methane-derived carbon to pelagic food webs. Excluding juveniles, non-native and fluvial species, adult fish showed strong reliance (on average 80.9%) on benthic production, weakly coupled with pelagic food webs. These results contrast low benthic production reliance (on average 27.4%) for a fish community in Lake Biwa, which is also classified as a subtropical lake. Both lakes are characterised by deep pelagic waters but quite different in their geological ages, suggesting that the aquatic communities in the FTR have fluvial origins, and their lacustrine history was too short for them to adapt to newly emerged deep pelagic habitat. Our isotope data are useful as a reference of newly established lentic food webs to monitor ongoing ecological and evolutionary dynamics as a result of anthropogenic disturbances.

Spatial genetic structure and body size divergence in endangered Gymnogobius isaza in ancient Lake Biwa

Abstract Gymnogobius isaza is a freshwater goby endemic to ancient Lake Biwa, the largest lake in Japan. The species is now listed as ‘Critically Endangered’ in the Red Data Book of Japan. Nevertheless, it remains subject to fishing without any specific management strategies. Previous studies using mitochondrial DNA markers showed that this fish species has two cryptic lineages. However, little is known about spatial genetic structure and ecological differences across the broad lakescape. In this study, we collected fish samples at nine locations along the lakeshore during the breeding season and tested for the presence of spatial heterogeneity in the lineage’s composition while measuring body size as the most fundamental biological trait. The results showed that the major lineage dominated all the sampling locations whereas the minor lineage consisted of only 11% (16/143) of samples. Furthermore, although their spatial distributions overlapped (i.e. the two lineages may be well mixed), we found it possible that the minor lineage may have a potentially narrower distribution than the major lineage. In addition, we found that the two lineages differ in body size; specifically, the minor lineage is smaller in size. From the viewpoint of genetic diversity conservation and sustainable resource use, this fish should be managed as two genetic stocks and spatial and/or body size-based fishery management is desirable, with particular attention to the minor (smaller sized) lineage.

Integrated trophic position decreases in more diverse communities of stream food webs

The relationship between biodiversity and ecosystem functioning is an important theme in environmental sciences. We propose a new index for configuration of the biomass pyramid in an ecosystem, named integrated trophic position (iTP). The iTP is defined as a sum of trophic positions (i.e. the average number of steps involved in biomass transfer) of all the animals in a food web integrated by their individual biomass. The observed iTP for stream macroinvertebrates ranged from 2.39 to 2.79 and was negatively correlated with the species density and the Shannon–Wiener diversity index of the local community. The results indicate a lower efficiency of biomass transfer in more diverse communities, which may be explained by the variance in edibility hypothesis and/or the trophic omnivory hypothesis. We found a negative effect of biodiversity on ecosystem functioning.

The effect of human activities on benthic macroinvertebrate diversity in tributary lagoons surrounding Lake Biwa

In aquatic ecosystems, tributaries play an important role in maintaining the populations and diversity of aquatic organisms throughout the drainage basin, but because of their small catchments, these ecosystems are often more susceptible to the effect of land-use changes and flow-regime alterations. Here, we examined anthropogenic effects on benthic macroinvertebrate diversity in the tributary lagoons, collectively called “Naiko,” surrounding Lake Biwa. We sampled macroinvertebrates and determined the environmental characteristics of 20 tributary lagoons. We identified the environmental factors determining the diversity of macroinvertebrates and found that turbidity significantly diminished species diversity. We assessed the anthropogenic stressors that contributed to the increase in turbidity and found that human population density and the proportion of paddy fields in the watershed area were positively correlated with turbidity, most likely caused by erosion of terrigenous organic matter from the paddy fields and urban areas. In addition, the presence of sluice gates and the lengths of channels connecting to the main lake were positively correlated with turbidity, suggesting that these factors lowered hydrologic connectivity and retained organic matter. We concluded that changes to the landscape and decreased hydrologic connectivity caused by human activity increased turbidity, which in turn decreased macroinvertebrate diversity. The identification of these factors in tributaries is vital for developing a strategy for habitat restoration to conserve the entire ecosystem of the Lake Biwa basin.