Low-energy nitrogen removal from ammonium-rich wastewater is crucial in preserving the water environment. A one-stage nitritation/anammox process with two inflows treating ammonium-containing wastewater, supplied from inside and outside the wound filter, is expected to stably remove nitrogen. Laboratory-scale reactors were operated using different start-up strategies; the first involved adding nitritation inoculum after anammox biomass formation in the filter, which presented a relatively low nitrogen removal rate (0.171 kg N/m3 · d), at a nitrogen loading rate of 1.0 kg N/m3 · d. Conversely, the second involved the gradual cultivation of anammox and nitritation microorganisms, which increased the nitrogen removal rate (0.276 kg N/m3 · d). Furthermore, anammox (Candidatus Brocadia) and nitritation bacteria (Nitrosomonadaceae) coexisted in the biofilm formed on the filter surface. The abundance of nitritation bacteria (10.5%) in the reactor biofilm using the second start-up strategy was higher than that using the first (3.7%). Thus, the two-inflow nitritation/anammox process effectively induced habitat segregation using a suitable start-up strategy.

Different crustacean species can differ in their response to light. In Tanaidacea, a small group of aquatic, benthic crustaceans, previous studies suggested that several species may be positively phototactic based on their attraction to nocturnal light traps, but no experimental investigations of phototaxis had been conducted on this group. Here we show experimentally that two species in the genus Zeuxo are phototactic but exhibit opposite reactions to light; Zeuxo ezoensis, which inhabits the blades and stipes of seaweeds, was positively phototactic, whereas Zeuxo molybi, which inhabits muddy sediments overlying bedrock, was negatively phototactic. This differential response may reflect differences in photoenvironment between these species\' microhabitats.

Electrofishing and visual observations revealed contrasting riverine distribution and habitat use of the Japanese eel, Anguilla japonica, and the giant mottled eel, Anguilla marmorata, in a relatively gentle-slope river of Southern Kyushu, Japan. A. japonica was distributed in the lower and upper reaches. A. marmorata inhabited the middle reaches with relatively fast water velocity, heterogeneous water depth, coarse substrata and no adjacent paddy fields. This may have important conservation implications because a decrease in environmental diversity and/or river connectivity could result in the disappearance of A. marmorata habitats.

AbstractSpatial segregation of closely related species is usually attributed to differences in stress tolerance and competitive ability. For both animals and plants, reproductive interactions between close relatives can impose a fitness cost that is more detrimental to the rarer species. Frequency-dependent mating interactions may thus prevent the establishment of immigrants within heterospecific populations, maintaining spatial segregation of species. Despite strong spatial segregation in natural populations, two sympatric California monkeyflowers (Mimulus nudatus and M. guttatus) survive and reproduce in the other\'s habitat when transplanted reciprocally. We hypothesized that a frequency-dependent mating disadvantage maintains spatial segregation of these monkeyflowers during natural immigration. To evaluate this hypothesis, we performed two field experiments. First, we experimentally added immigrants in varying numbers to sites dominated by heterospecifics. Second, we reciprocally transplanted arrays of varying resident and immigrant frequencies. Immigrant seed viability decreased with conspecific rarity for M. guttatus but not for M. nudatus. We observed immigrant minority disadvantage for both species, but it was driven by different factors-frequency-dependent hybridization for M. guttatus and competition for resources and/or pollinators for M. nudatus. Overall, our results suggest a major role for reproductive interference in spatial segregation that should be evaluated along with stress tolerance and competitive ability.

Niche differentiation between closely related species leads to differentiation of their habitats. Segregation based on slight differences in environmental factors, that is niche differentiation on the microhabitat scale, allows more species to inhabit a certain geographic space. Therefore, such fine scale niche differentiation is an important factor in the support of species diversity. In addition, niche differentiation on the microhabitat scale and/or the differentiation of breeding seasons can be considered typical mechanisms that facilitate multispecies\' co-existence. In this study, sister species (Commonly, Ephemera japonica inhabit at upstream region and Ephemera strigata inhabit at middle stream region), which often coexist in the upper to middle reaches of river systems of the Japanese Islands, were targeted and the following aspects were investigated. First, differences in habitat preference and interspecific differences in flow distribution patterns on a geographically fine scale were tracked in detail. Subsequently, the temporal transitions of their distribution patterns were investigated in detail and seasonal changes were investigated. Finally, we thoroughly investigated the disappearance of nymphs of each species from the river due to emergence affected the distribution of each species (by conducting daily emergence surveys). Combining results of these multiple studies also suggested that there may be spatiotemporal interspecial interaction between these two species within/around their overlapping regions. Traditionally, the longitudinal distribution pattern of these two Ephemera mayflies has been thought to be established based on a difference in habitat preferences, but this study revealed that the interspecific interaction between the two species also plays an important role. This study provides new insights into species diversity and distribution pattern formation in river-dwelling species.

Incompletely reproductively isolated species often segregate into different microhabitats, even when they are able to survive and reproduce in both habitats. Longer term evolutionary factors may contribute to this lack of cross-habitat persistence. When reproductive interference reduces immigrant fitness, assortative mating, including self-fertilization, increases immigrants\' fitness in a single generation, but longer term, inbreeding depression may reduce the chance of population persistence. Two California monkeyflower species repeatedly segregate into drier and wetter areas in their zone of sympatry. To test whether inbreeding depression may contribute to the maintenance of this segregation pattern, we transplanted outbred and successively inbred Mimulus guttatus and Mimulus nudatus into their native habitats and heterospecific habitats. We measured germination, survival, and seed set and found that recurrent selfing reduced all aspects of fitness in both species, most strongly in foreign habitats. A simulation model, parameterized from the transplant experiment, found that inbreeding reduced fitness to such an extent that sequentially inbred populations of either species would be unable to persist in heterospecific-occupied habitats in the absence of continued gene flow. These results demonstrate that individual immigrants are unlikely to form persistent populations and thus, inbreeding depression contributes to the absence of fine-scale coexistence in this species pair.

Golden and Blueline Tilefish (Lopholatilus chamaeleonticeps and Caulolatilus microps) are keystone taxa in northwest (NW) Atlantic continental shelf-edge environments due to their biotic (trophic-mediated) and abiotic (ecosystem engineering) functional roles combined with high-value fisheries. Despite this importance, the ecological niche dynamics (i.e., those relating to trophic behavior and food-web interactions) of these sympatric species are poorly understood, knowledge of which may be consequential for maintaining both ecosystem function and fishery sustainability. We used stable isotope ratios of carbon (δ13C) and nitrogen (δ15N) to build realized ecological niche hypervolumes to serve as proxies for diet and production use patterns of L. chamaeleonticeps and C. microps. We hypothesized that: (a) species exhibit ontogenetic shifts in diet and use of production sources; (b) species acquire energy from spatially distinct resource pools that reflect a sedentary life-history and differential use of the continental shelf-edge; and (c) species exhibit differentiation in one or more measured niche axes. We found evidence for ontogenetic shifts in diet (δ15N) but not production source (δ13C) in both species, suggesting a subtle expansion of measured ecological niche axes. Spatial interpolation of stable isotope ratios showed distinct latitudinal gradients; for example, individuals were 13C enriched in northern and 15N enriched in southern regions, supporting the assertion that tilefish species acquire energy from regional resource pools. High isotopic overlap was observed among species (≥82%); however, when hypervolumes included depth and region of capture, overlap among species substantially decreased to overlap estimates of 15%-77%. This suggests that spatial segregation could alleviate potential competition for resources among tilefish species inhabiting continental shelf-edge environments. Importantly, our results question the consensus interpretation of isotopic overlap estimates as representative of direct competition among species for shared resources or habitats, instead identifying habitat segregation as a possible mechanism for coexistence of tilefish species in the NW Atlantic.

Maladaptive hybridization selects for prezygotic isolation, a process known as reinforcement. Reinforcement reduces gene flow and contributes to the final stage of speciation. Ecologically, however, coexistence of the incipient species is difficult if they initially use identical resources. Habitat segregation offers an alternative to species discrimination as a way to reduce gene flow: production of unfit hybrids is reduced if mate encounters become rare due to differing habitat choice. Using a modelling approach, we show that hybridization avoidance alone can select for habitat specialization, even if neither of the species is intrinsically better at using a specific niche. While habitat segregation and species discrimination both reduce the risk of producing unfit hybrids, these two isolation mechanisms differ from each other with respect to their effects on resource competition. Our model shows that, as a consequence of such differences, reinforcement evolves much more easily if hybridization is avoided based on habitat segregation than if the mechanism involves species recognition (mate choice traits). We also examine the outcomes when both isolation mechanisms evolve jointly. The establishment of one isolation mechanism a priori weakens selection for the other. However, an asymmetry persists here too. The net effect of habitat segregation on species discrimination was typically facilitative, but not vice versa. This asymmetry arises because habitat segregation, by enhancing coexistence, secures time for the subsequent evolution of species discrimination in a mate choice context (still relevant if habitat use is not perfectly segregated). Species discrimination does not have such a stabilizing effect on coexistence. Our results emphasize the importance of habitat segregation in reinforcement and offer a way to interpret findings where closely related taxa show similar performance on different resources or in different habitats. Studies of ecological generalization and specialization should therefore take into account that niche differences can be initiated and/or maintained by hybridization avoidance.

Knowledge of niche partitioning with respect to habitat is indispensable to understand the mechanism of coexistence of multiple species. Among primates, however, data are still deficient because repeated survey for a sufficiently long time, covering seasonal changes over a large area, is the only way to clarify habitat segregation within a seasonally fluctuating environment. Southeast Asia is particularly interesting because of the supra-annual, highly unpredictable seasonality in fruiting known as mast fruiting. We conducted repeated route census, habitat monitoring, and group tracking for 25 months in two study sites (ca. 10 km apart) in the largely primary lowland dipterocarp forest of the Danum Valley Conservation Area, eastern Sabah, northern Borneo, Malaysia. The five species of diurnal primates (Bornean orangutan Pongo pygmaeus, Müeller\'s gibbon Hylobates muelleri, red leaf monkey Presbytis rubicunda, long-tailed macaque Macaca fascicularis, and southern pig-tailed macaque M. nemestrina) did not show horizontal spatial segregation. Red leaf monkeys showed preferences for places with short tree height, but their distribution was not confined to such places. In response to the fruiting peak observed once during the study period, orangutans increased their numbers simultaneously in the two study sites. The average tree height used by the five species was different, but their range overlapped substantially. Compared with other primate communities, the lack of horizontal spatial segregation and the suggested long-distance movement of orangutans seem to be unique characteristics in Borneo, although the use of different forest strata is a widespread phenomenon among primate communities throughout the world.

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