A shady phytoplankton paradox: when phytoplankton increases under low light.

Light is a fundamental driver of ecosystem dynamics, affecting the rate of photosynthesis and primary production. In spite of its importance, less is known about its community-scale effects on aquatic ecosystems compared with those of nutrient loading. Understanding light limitation is also important for ecosystem management, as human activities have been rapidly altering light availability to aquatic ecosystems. Here we show that decreasing light can paradoxically increase phytoplankton abundance in shallow lakes. Our results, based on field manipulation experiments, field observations and models, suggest that, under competition for light and nutrients between phytoplankton and submersed macrophytes, alternative stable states are possible under high-light supply. In a macrophyte-dominated state, as light decreases phytoplankton density increases, because macrophytes (which effectively compete for nutrients released from the sediment) are more severely affected by light reduction. Our results demonstrate how species interactions with spatial heterogeneity can cause an unexpected outcome in complex ecosystems. An implication of our findings is that partial surface shading for controlling harmful algal bloom may, counterintuitively, increase phytoplankton abundance by decreasing macrophytes. Therefore, to predict how shallow lake ecosystems respond to environmental perturbations, it is essential to consider effects of light on the interactions between pelagic and benthic producers.

Effects of sediment replenishment on riverbed environments and macroinvertebrate assemblages downstream of a dam.

Riverbeds downstream of dams are starved of sediment, impacting habitat structure and ecological function. Despite the implementation of sediment management techniques, there has been no evaluation of their conservational effectiveness; the impacts on high trophic level organisms remain unknown. This study examined the effects of sediment replenishment on riverbeds and macroinvertebrates in a dammed river before and after sediment replenishment. We evaluated the particle sizes of replenished sediments and the case material of a case-bearing caddisfly. We observed significant changes in macroinvertebrate assemblages before and after sediment replenishment, and between the upstream and tributary references and downstream of the dam. The percentages of Ephemeroptera, Plecoptera, and Trichoptera, and the number of inorganic case-bearing caddisflies downstream of the dam following sediment replenishment, were significantly higher than the upstream and tributary reference sites. The particle size of case materials used by case-bearing caddisfly corresponded to the size of the replenished sediment. Dissimilarity results after replenishment showed that assemblages downstream of the dam differed from upstream sites, although they were similar to the tributary sites. The dissimilarity between the tributary and downstream of the dam was the same as that between the upstream and tributary. Sediment replenishment was observed to reduce the harmful effects of the dam, and partly restore benefits such as increasing species diversity and altering community assemblages, similar to the effects of tributary inflows.

A unified framework for herbivore-to-producer biomass ratio reveals the relative influence of four ecological factors.

The biomass ratio of herbivores to primary producers reflects the structure of a community. Four primary factors have been proposed to affect this ratio, including production rate, defense traits and nutrient contents of producers, and predation by carnivores. However, identifying the joint effects of these factors across natural communities has been elusive, in part because of the lack of a framework for examining their effects simultaneously. Here, we develop a framework based on Lotka-Volterra equations for examining the effects of these factors on the biomass ratio. We then utilize it to test if these factors simultaneously affect the biomass ratio of freshwater plankton communities. We found that all four factors contributed significantly to the biomass ratio, with carnivore abundance having the greatest effect, followed by producer stoichiometric nutrient content. Thus, the present framework should be useful for examining the multiple factors shaping various types of communities, both aquatic and terrestrial.

Effects of sampling seasons and locations on fish environmental DNA metabarcoding in dam reservoirs.

Environmental DNA (eDNA) analysis has seen rapid development in the last decade, as a novel biodiversity monitoring method. Previous studies have evaluated optimal strategies, at several experimental steps of eDNA metabarcoding, for the simultaneous detection of fish species. However, optimal sampling strategies, especially the season and the location of water sampling, have not been evaluated thoroughly. To identify optimal sampling seasons and locations, we performed sampling monthly or at two-monthly intervals throughout the year in three dam reservoirs. Water samples were collected from 15 and nine locations in the Miharu and Okawa dam reservoirs in Fukushima Prefecture, respectively, and five locations in the Sugo dam reservoir in Hyogo Prefecture, Japan. One liter of water was filtered with glass-fiber filters, and eDNA was extracted. By performing MiFish metabarcoding, we successfully detected a total of 21, 24, and 22 fish species in Miharu, Okawa, and Sugo reservoirs, respectively. From these results, the eDNA metabarcoding method had a similar level of performance compared to conventional long-term data. Furthermore, it was found to be effective in evaluating entire fish communities. The number of species detected by eDNA survey peaked in May in Miharu and Okawa reservoirs, and in March and June in Sugo reservoir, which corresponds with the breeding seasons of many of fish species inhabiting the reservoirs. In addition, the number of detected species was significantly higher in shore, compared to offshore samples in the Miharu reservoir, and a similar tendency was found in the other two reservoirs. Based on these results, we can conclude that the efficiency of species detection by eDNA metabarcoding could be maximized by collecting water from shore locations during the breeding seasons of the inhabiting fish. These results will contribute in the determination of sampling seasons and locations for fish fauna survey via eDNA metabarcoding, in the future.

Effects of stream grazers with different functional traits on the spatial heterogeneity of periphyton mats.

BACKGROUND: Grazing activity on periphytic mats determines mat structure and spatial heterogeneity. Spatial complexity in stream periphyton is highly divergent and may depend on the functional traits of the different primary consumers species (i.e., grazers) such as mouthpart morphology, feeding behavior, and feeding activity. We evaluated the effect of grazing by three species having different functional traits on periphytic mat structure with a focus on mohthpart morphology. METHODS: An enclosure experiment was conducted in a stream located in the Nara Prefecture of Japan using two caddisflies with scraping mouthparts, Micrasema quadriloba and Glossosoma, and one mayfly, Epeorus, with brushing mouthparts. The spatial heterogeneity of chlorophyll a(Chl a) was evaluated, and the periphytic mat was analyzed using scanning electron microscopy (SEM) after a 12-d feeding experiment. RESULTS: Our results showed the differences in the spatial heterogeneity of the periphytic mats, such as differences in Chl a levels, grazed by the different grazing species. The strongest effect on the spatial heterogeneity and periphytic abundance was observed for Micrasema quadriloba, a caddisfly species with scraping mouthparts. Epeorus mayfly, with brushing mouthparts and high-mobility behavior, produced the weakest effect on spatial heterogeneity. Glossosoma caddisflies had moderate effects on periphytic spatial heterogeneity and abundance. Our results suggest that differences in grazing effects are largely dependent on grazer mouthparts and behavior.

Environmental DNA detection and quantification of invasive red-eared sliders, Trachemy scripta elegans, in ponds and the influence of water quality.

Environmental DNA (eDNA) is a powerful tool for monitoring the distribution of aquatic macro-organisms. However, environmental factors, including the water temperature and water quality, can affect the inhibition and/or degradation of eDNA, which complicates accurate estimations of eDNA concentrations and the detection of the presence/absence of species in natural habitats. Further very few eDNA studies have been conducted for reptiles, especially with respect to estimating their biomass and/or abundances. Here we examined the relationship between the visually-observed number of red-eared sliders (Trachemys scripta elegans) and eDNA concentrations across 100 ponds. Additionally, we evaluated the effect of water quality on red-eared slider eDNA concentration in these ponds. We found that there was a significant positive correlation between the observed number of red-eared sliders and the eDNA concentration in the ponds. On comparing various water quality indicators, including dissolved nitrogen, dissolved phosphorous, organic matter, and chlorophyll a (Chl. a), we found that only Chl. a had a negative correlation with the red-eared slider eDNA concentration, while we did not find any inhibition in the quantitative PCR. We conclude that concentrations of eDNA can potentially be used for estimating the abundance of the red-eared slider. Additionally, Chl. a might indirectly influence the degradation of eDNA through the microorganisms bonded to the phytoplankton in the ponds, as microbial activity is thought to decrease eDNA persistence.

Distribution and drift dispersal dynamics of a caddisfly grazer in response to resource abundance and its ontogeny.

Stream grazers have a major impact on food web structure and the productivity of stream ecosystems; however, studies on the longitudinal (upstream versus downstream) and temporal changes in their drift dynamics and resulting distributions remain limited. Here, we investigated the longitudinal and temporal distributions and drift propensity of a trichopteran grazer, the caddisfly, Micrasema quadriloba, during its life cycle in a Japanese stream. The distribution of larvae significantly shifted downstream during the fifth instar larval stage during late winter; with periphyton abundance (i.e. their food source) showing similar shifts downstream. Therefore, our results show that the drift dispersal the caddisfly occurs in response to decline in available food resources (i.e. food-resource scarcity) and an increase in food requirements by growing individuals. Furthermore, our results show that this observed longitudinal shift in larval distribution varies through their life cycle, because the drift dispersal of fifth instar larvae was greater than that of immature larvae. The correlation between periphyton abundance and drift propensity of fourth instar larvae was not statistically significant, whereas that of fifth instar larvae was significantly negative. In conclusion, we detected an ontogenetic shift in drift propensity, which might explain the longitudinal and temporal distributions of this species.

Environmental DNA method for estimating salamander distribution in headwater streams, and a comparison of water sampling methods.

Environmental DNA (eDNA) has recently been used for detecting the distribution of macroorganisms in various aquatic habitats. In this study, we applied an eDNA method to estimate the distribution of the Japanese clawed salamander, Onychodactylus japonicus, in headwater streams. Additionally, we compared the detection of eDNA and hand-capturing methods used for determining the distribution of O. japonicus. For eDNA detection, we designed a qPCR primer/probe set for O. japonicus using the 12S rRNA region. We detected the eDNA of O. japonicus at all sites (with the exception of one), where we also observed them by hand-capturing. Additionally, we detected eDNA at two sites where we were unable to observe individuals using the hand-capturing method. Moreover, we found that eDNA concentrations and detection rates of the two water sampling areas (stream surface and under stones) were not significantly different, although the eDNA concentration in the water under stones was more varied than that on the surface. We, therefore, conclude that eDNA methods could be used to determine the distribution of macroorganisms inhabiting headwater systems by using samples collected from the surface of the water.

Detection of an endangered aquatic heteropteran using environmental DNA in a wetland ecosystem.

The use of environmental DNA (eDNA) has recently been employed to evaluate the distribution of various aquatic macroorganisms. Although this technique has been applied to a broad range of taxa, from vertebrates to invertebrates, its application is limited for aquatic insects such as aquatic heteropterans. Nepa hoffmanni (Heteroptera: Nepidae) is a small (approx. 23 mm) aquatic heteropteran that inhabits wetlands, can be difficult to capture and is endangered in Japan. The molecular tool eDNA was used to evaluate the species distribution of N. hoffmanni in comparison to that determined using hand-capturing methods in two regions of Japan. The eDNA of N. hoffmanni was detected at nearly all sites (10 eDNA-detected sites out of 14 sites), including sites where N. hoffmanni was not captured by hand (five eDNA-detected sites out of six captured sites). Thus, this species-specific eDNA technique can be applied to detect small, sparsely distributed heteropterans in wetland ecosystems. In conclusion, eDNA could be a valuable technique for the detection of aquatic insects inhabiting wetland habitats, and could make a significant contribution to providing distribution data necessary to species conservation.

Reach-scale distribution dynamics of a grazing stream insect, Micrasema quadriloba Martynov (Brachycentridae, Trichoptera), in relation to current velocity and periphyton abundance.

Reach-scale temporal shifts in the distribution of larvae of a grazing caddisfly, Micrasema quadriloba (Brachycentridae), were investigated in a Japanese mountain stream. The larvae showed an aggregated distribution within the reach at the beginning of the immigration, then became randomly dispersed throughout the reach as the immigration progressed. The abundance of periphyton in the reach decreased dramatically with increasing dispersal of the larvae. Simple regression analyses revealed that the stream's flow regime was the most important environmental factor that determined the reach-scale distribution of the larvae and that the relationship between the flow regime and the distribution of the larvae shifted temporally. In addition, our results suggest that only this species of grazing insect, which was dominant in the study reach, controlled the reach-scale abundance of the periphyton.

Stream grazers determine their crawling direction on the basis of chemical and particulate microalgal cues.

This study aimed to determine the association between herbivore behavior and cues from producers. We used stream grazer Glossosoma larvae and determined their crawling direction in relation to chemical and visual cues from microalgae. The experimental treatments included control (no cue), particulate (chemical and particulate cues), and dissolved (chemical cue) cues from microalgae. The experimental water samples were randomly placed into either arm of a Y-shaped channel, and the crawling direction of the grazers was determined. Although the grazers crawled toward the arm containing either particulate or dissolved cues, they preferred the arm with particulate cues. This suggested that grazers responded well to both particulate (i.e., drifting algal cells) and chemical (algal smell) cues, and that particulate cues were more important for foraging. In natural habitats, grazers detect cues from producers and change their behaviors to maintain a balance between top-down and bottom-up cues.

Drifting plankton from a reservoir subsidize downstream food webs and alter community structure.

Subsidy between ecosystems has been considered in many natural ecosystems, and should alter food webs and communities in human-impacted ones. We estimated how drifting plankton from a reservoir contribute to downstream food webs and showed that they alter community structures over a 10-km reach below the dam. To estimate the contribution of the drifting plankton to macroinvertebrates, we used C and N isotopes and an IsoSource mixing model. In spring and autumn, contributions of plankton to collector-filterer species were highest 0.2 km downstream of the dam, and clearly decreased from 0.2 to 10 km. At 0.2 km, the contribution of plankton to a predator stonefly was remarkably high. These results indicated that drifting plankton from a dam reservoir could subsidize downstream food webs and alter their energy base, but the importance of this subsidy decreased as distance from the reservoir increased. The general linear models indicated that the abundance of collector-filterers and predators was related positively to zooplankton density in stream water. Thus, food source alteration by drifting plankton also influenced the community structures downstream of the dam.