Shigeru Nakano’s research while affiliated with Kyoto University and other places

Coexistence of species requires equalizing mechanisms that minimize fitness differences, which are balanced by stabilizing mechanisms that enhance negative intraspecific interactions versus interspecific ones. Here, we develop a simple theoretical framework that allows measuring the relative strength of intraspecific versus interspecific competition in dominance hierarchies. We use it to evaluate mechanisms promoting coexistence between two congeneric charr that compete for foraging positions, which strongly influence density‐dependent growth and survival. Agonistic interactions (n = 761) among 71 Dolly Varden Salvelinus malma and whitespotted charr Salvelinus leucomaenis were measured by snorkelling in two pools in the sympatric zone of a Hokkaido stream during two summers. Interspecific dominance hierarchies, analysed using three methods, were closely correlated with fish length but the species treated each other equally. Ranks for the most dominant fish in each pool, determined directly by knockout experiments, were also virtually identical to ranks by length. Similarly, exponential random graph modelling of the social networks provided no evidence that either species was dominant over the other. Instead, larger fish were more likely to win contests, especially over fish of the next lower ranks. These results demonstrated that the two species were nearly ecological equivalents in accessing key resources in this sympatric zone. Nearly identical growth and stable densities over 4 years further supported this inference, although Dolly Varden were a minority (29% of the assemblage), a sign of some fitness difference. Detailed foraging observations coupled with two concurrent studies revealed an effective stabilizing mechanism. Dolly Varden shifted to feeding directly from the benthos when drifting invertebrates declined, a behaviour enhanced by morphological character displacement, thereby partitioning food resources and enhancing intraspecific competition while avoiding agonistic encounters with whitespotted charr. The plurality of evidence indicates that fitness differences between these ecologically equivalent species are small in this local assemblage, and balanced by resource partitioning, a modest stabilizing mechanism that promotes coexistence. The theoretical framework presented here is a useful tool to evaluate the strength of interspecific versus intraspecific competition, which combined with information on trade‐offs in ecological performance can contribute to a mechanistic understanding of species coexistence.

Networks of direct and indirect biotic interactions underpin the complex dynamics and stability of ecological systems, yet experimental and theoretical studies often yield conflicting evidence regarding the direction (positive or negative) or magnitude of these interactions. We revisited pioneering data sets collected at the deciduous forested Horonai Stream and conducted ecosystem‐level syntheses to demonstrate that the direction of direct and indirect interactions can change depending on the timescale of observation. Prior experimental studies showed that terrestrial prey that enter the stream from the adjacent forest caused positive indirect effects on aquatic invertebrates during summer by diverting fish consumption. Seasonal and annual estimates of secondary production and organic matter flows along food web pathways demonstrate that this seasonal input of terrestrial invertebrate prey increases production of certain fish species, reversing the indirect effect on aquatic invertebrates from positive at the seasonal timescale to negative at the annual timescale. Even though terrestrial invertebrate prey contributed 54% of the annual organic matter flux to fishes, primarily during summer, fish still consumed 98% of the aquatic invertebrate annual production, leading to top‐down control that is not revealed in short‐term experiments and demonstrating that aquatic prey may be a limiting resource for fishes. Changes in the direction or magnitude of interactions may be a key factor creating nonlinear or stabilizing feedbacks in complex systems, and these dynamics can be revealed by merging experimental and comparative approaches at different scales.

Similar species that overlap in sympatry may diverge in characters related to resource use as a result of evolution or phenotypic plasticity. Dolly Varden charr (Salvelinus malma) and whitespotted charr (S. leucomaenis) overlap along streams in Hokkaido, Japan, and compete by interference for invertebrate drift-foraging positions. Previous research has shown that as drift declines during summer, Dolly Varden shift foraging modes to capture benthic prey, a behaviour facilitated by their subterminal jaw morphology. We compare body and jaw morphology of Dolly Varden in sympatry vs. allopatry in two locations to test for character displacement. Statistical analysis showed significant divergence in characters related to foraging, which was correlated with variation in individual charr diets. Dolly Varden in sympatry had shorter heads and lower jaws than in allopatry, and even within sites charr with these characteristics fed less on drifting terrestrial invertebrates but more on benthic aquatic invertebrates. Those in allopatry had longer heads and lower jaws, and fed more on terrestrial invertebrates. The close proximity of sites in one stream suggests that Dolly Varden may display phenotypic plasticity similar to other charr, allowing rapid responses in morphology to the presence of competitors. These morphological shifts probably help them maintain positive fitness when competing with whitespotted charr in Hokkaido streams.

The importance of leaf litter quality for colonization and consumption by two caddisflies (Goerodes satoi and Hydatophylax festivus) and an amphipod (Sternomoera yezoensis) was examined by laboratory and field experiments in a forested headwater stream in northern Japan. Initial values of leaf litter toughness and the carbon to nitrogen (C : N) ratio, both of which varied among the three most common tree species, oak (Quercus crispula), maple (Acer mono) and alder (Alnus hirsuta), decreased dramatically in the field experiment. The density of Hydatophylax was greatest in alder leaf litter characterized by the lowest C : N ratio among the three, with C : N ratio being the only predictor of their density. In contrast, lesser toughness was the only factor facilitating colonization by Sternomoera, the density of which was greatest in the softest maple litter. Neither toughness nor C : N ratio could be used to predict colonization by Goerodes. In the laboratory experiment, in which the shredders were reared on the aforementioned types of leaf litter, following conditioning periods of 1 week and 1 month (six separate trials), the litter consumption rate decreased with toughness in both Sternomoera and Hydatophylax, and with C : N ratio in the latter. However, the consumption rate of Goerodes was influenced by neither of the above, indicating that Sternomoera and Hydatophylax, but not Goerodes, selectively colonized the leaf litter available as a food resource in the stream. Both chemical and physical qualities of leaf litter play an important role in determining species-specific colonization by invertebrate shredders.

Riparian zones serve several ecological functions for bats. They provide a source of prey and likely provide favourable structural habitats and shelter from predators. Many studies have shown that bats use the space above streams, ponds or riparian vegetation as feeding habitat. These studies, however, have never distinguished between the effects of habitat structure and prey availability on the foraging activities of bats. Such effects can only be distinguished by an experimental approach. We predicted that bat activity along a stream is influenced by the number of emerged aquatic insects. We evaluated the response of terrestrial consumers, insectivorous bats, to changes in the abundance of emergent aquatic insects by conducting a manipulative field experiment. In a deciduous riparian forest in Japan, aquatic insect flux from the stream to the riparian zone was controlled with an insect‐proof cover over a 1·2 km stream reach. We estimated the abundance of emergent aquatic and flying terrestrial arthropods near the treatment and control reaches using Malaise traps. The foraging activity of bats was evaluated in both treatment and control reaches using ultrasonic detectors. The insect‐proof cover effectively reduced the flux of emergent aquatic insects to the riparian zone adjacent to the treatment reach. Adjacent to the control reach, adult aquatic insect biomass was highest in spring, and then decreased gradually. Terrestrial insect biomass increased gradually during the summer at both treatment and control reaches. Foraging activity of bats was correlated with insect abundance. In spring, foraging activity of bats at the control reach was significantly greater than at the treatment reach, and increased at both sites with increasing terrestrial insect abundance. Our result suggests that the flux of aquatic insects emerging from streams is one of the most important factors affecting the distribution of riparian‐foraging bats. As is the case with other riparian consumers, resource subsidies from streams can directly enhance the performance or population density of riparian‐dependent bats. To conserve and manage bat populations, it is important to protect not only forest ecosystems, but also adjacent aquatic systems such as streams.

In northern Japan, an alien crayfish species (signal crayfish; Pacifastacus leniusculus) is replacing Japan's only native crayfish species (Cambaroides japonicus) that is endemic to the country. We tested experimentally if these two crayfish species had similar roles in terms of ecosystem functioning in stream habitats. Experiments were performed to examine the impacts of alien and endemic crayfish on detritus-based food webs. During the experiment, neither crayfish species affected rates of breakdown of microbially conditioned oak leaves. In contrast, the amphipod Jessogammarus jesoensis, which dominated the non-crayfish invertebrate biomass, was significantly reduced in the presence of the two crayfish species. Thus, effects of crayfish on leaf breakdown were probably masked by the leaf processing activities of these large amphipods in the absence of crayfish. Overall, per-unit biomass impacts of Pacifastacus on leaf processing and invertebrate colonisation of leaves were comparable to those by Cambaroides.Laboratory experiments were performed to test whether rates of leaf processing of 10 riparian plant species, particulate organic matter (POM) production and nutrient excretion differed between crayfish species in the absence of other invertebrates. Although POM production rates did not differ between crayfish species, crayfish impacts on leaf processing differed depending on plant species reflecting differential leaf preference. Comparisons of individual nutrient excretion rates revealed that ammonia and phosphate excretion rates were higher in Cambaroides compared with Pacifastacus. Further, ammonia excretion rates were proportional to crayfish biomass in Cambaroides but not in Pacifastacus.Taken together, the two crayfish species had similar roles in stream food webs when biomass was similar. However, their roles as leaf processors and excretion rates of nutrients differed when there was a choice among different riparian species as sources of leaf litter. Species replacement between alien and endemic crayfish may therefore result in differential consequences for ecosystem processes.

We experimentally measured thermal changes in the gastric evacuation rate (GER) of the freshwater sculpin Cottus nozawae Snyder under three water temperature regimes (2°, 7°, and 12°C). Laboratory experiments showed that the GER was accelerated with increasing water temperature. This result suggests that the daily food rations of fish are more likely to be underestimated at higher water temperatures if estimation is simply based on the stomach content weight alone. By comparing the GER for various fish species from subfrigid to temperate streams, we found a general pattern that the GER increases with water temperature, regardless of taxonomic group or foraging mode. However, the reaction norms of the GER against water temperature showed considerable interspecific variation. This means that stomach content weight is not comparable as a simple measure for determining the daily rations among fish species when water temperature regimes are different. To consider the temperature-dependent pattern of such a physiological phenomenon is important in understanding the feeding ecology of fishes and their roles in material cycles through food webs in aquatic ecosystems.

1. We examined effects of water temperature on the community structure of a three trophic level food chain (predatory fish, herbivorous caddisfly larvae and periphyton) in boreal streams. We used laboratory experiments to examine (i) the effects of water temperature on feeding activities of fish and caddisfly larvae and on periphyton productivity, to evaluate the thermal effects on each trophic level (species-level experiment), and (ii) the effects of water temperature on predation pressure of fish on abundance of the lower trophic levels, to evaluate how temperature affects top-down control by fish (community-level experiment). 2. In the species-level experiment, feeding activity of fish was high at 12 °C, which coincides with the mean summer temperature in forested streams of Hokkaido, Japan, but was depressed at 3 °C, which coincides with the mean winter temperature, and also above 18 °C, which coincides with the near maximum summer temperatures. Periphyton productivity increased over the range of water temperatures. 3. In the community-level experiments, a top-down effect of fish on the abundance of caddisfly larvae and periphyton was clear at 12 °C. This effect was not observed at 3 and 21 °C because of low predation pressure of fish at these temperatures. 4. These experiments revealed that trophic cascading effects may vary with temperature even in the presence of abundant predators. Physiological depression of predators because of thermal stress can alter top-down control and lead to changes in community structure. 5. We suggest that thermal habitat alteration can change food web structure via combinations of direct and indirect trophic interactions.