Fish abundance and habitat relationships in forest and grassland streams, northern Hokkaido, Japan

The relationships between habitat variables and population densities of masu salmon (Oncorhynchus masou), rosyface dace (Leuciscus ezoe), Siberian stone loach (Noemacheilus barbatulus) and wrinklehead sculpin (Cottus nozawae) were examined by data collected at 55 reaches in forest and grassland streams in northern Hokkaido, Japan. Regression analysis suggested that salmon and dace densities were affected by water temperature (negative for salmon, positive for dace) and structural habitat factors (woody debris for salmon, pools for dace). Salmon density was higher in forest reaches than in grassland reaches, whereas dace density was higher in grassland reaches, suggesting that the removal of riparian forest had raised water temperature and allowed upstream invasions by dace. In contrast to salmon and dace, neither the density of loach nor sculpin differed between the forest and grassland reaches. For their densities, a negative effect of each on the other was most important, suggesting a strong effect of interspecific competition between loach and sculpin on their distributions. However, regression models also suggested that substrate heterogeneity mediated the outcome of their interspecific competition. On the basis of the results, a scenario is predicted for a fish-assemblage change with a typical land-development process in Hokkaido, and the importance of leaving or restoring riparian buffer for conservation and restoration of stream habitat is emphasized.     

Habitat structure along channel-unit sequences for juvenile salmon: a subunit-based analysis of in-stream landscapes

1. Habitat structure and habitat use by juvenile masu salmon, Oncorhynchus masou Brevoort, in small streams in northern Hokkaido, Japan, were examined by considering 'subunits' (patches within channel units) as structural elements of stream reaches.
2. Whole wetted channel surfaces of three study reaches were divided into 0.5 × 0.5 m quadrats, which were grouped into eight subunit types according to water depth and velocity, and substratum conditions by a cluster analysis. The subunit distribution showed a regular mosaic pattern corresponding to the channel-unit sequence in each of the three reaches.
3. Juvenile masu salmon exhibited a strong preference for a subunit type characterized by greater depth and moderate current velocity (deep–moderate subunit; mean depth = 0.29 m; mean velocity = 0.19 m s⁻¹). This subunit type usually occurred downstream of stretches with fast current. The preference of masu salmon for the deep–moderate subunit could be because of its usual spatial position in relation to other subunit types as well as to the characteristics of the subunit itself.
4. The results suggest that the value of a habitat is determined not only by the characteristics of the habitat itself, but also by those of adjacent habitats. Therefore, habitat use by stream fish should be studied in the context of the whole 'in-stream landscapes'.     

Effects of woody debris on the habitat of juvenile masu salmon (Oncorhynchus masou) in northern Japanese streams

1. The effects of woody debris on stream habitat of juvenile masu salmon ( Oncorhynchus masou ) were examined at two spatial scales, stream reach and channel unit, for first to thirdorder tributaries of the Teshio River in northern Hokkaido, Japan. The fortyeight study reaches were classified into three distinct types: coarsesubstrate steppool (CSP), coarsesubstrate poolriffle (CPR) and finesubstrate poolriffle (FPR) reaches. Each reach type included reaches with different riparian settings, broadly classified as forest (relatively undisturbed forest and secondary forest after fires) or grassland (bamboo bushland and pasture).
2. The reachscale analyses showed that neither total pool volume nor pooltopool spacing was correlated with woody debris abundance in any of the three reach types. Masu salmon density was positively correlated with both woodydebris cover area and total cover area, but not with total pool volume in the reaches.
3. Channelunitscale analyses revealed that woody debris reduced nonpool velocity, increased pool depth and retained fine sediment in pools in FPR reaches, where the size of woody debris was very large relative to the substrate material size. However, woody debris did not influence any of the hydraulic variables (depth, velocity, substrate) in either nonpools or pools of CSP and CPR reaches. Habitat use by masu salmon in nonpools or pools was affected by woodydebris cover area or total cover area rather than by hydraulic variables in any of the reach types.
4. The effects of woody debris on habitat at the reach and channelunit scales in the study area were less than those indicated by previous work in the Pacific Northwest, North America, owing to the relatively small size of the riparian trees. However, the overall results suggested that woody debris in the study area contributed to masu salmon habitat by providing cover at the smaller, microhabitat scale.     

Application of a bioenergetics model to estimate the influence of habitat degradation by check dams and potential recovery of masu salmon populations

Using a bioenergetics model, we examined how check dams negatively effect masu salmon (Oncorhynchus masou) populations by causing habitat loss in upstream areas and habitat degradation in downstream areas. The potential recovery of masu salmon populations in the upstream area was estimated based on the expected biomass and potential recovery area. We also determined if and how fish carrying capacity is affected by degradation of substrate conditions (armoring and compaction) in the downstream area. Recovery of upstream areas was considered to be effective in enhancing and conserving masu salmon populations. We demonstrated that the dam-induced altered substrate conditions and habitat degradation in the downstream area resulted in a considerable reduction of drifting prey. Simulation analysis revealed that a 40 % increase in the abundance of masu salmon juveniles in the downstream area could be expected if substrate conditions were restored. We concluded that both improvement of migration barriers and restoring the sediment regime would be important in enhancing and conserving wild masu salmon populations.     

Preferable summering habitat of returning adult masu salmon in the natal stream

The preferable summering habitat of returning adult masu salmon Oncorhynchus masou Brevoort was examined in 31 study pools of a small stream in northern Hokkaido, Japan. A total of 23 salmon were captured and were present in only 10 of the study pools. Female salmon were found to be significantly larger than male salmon. Habitat characteristics were expressed by six environmental variables: mean water depth, maximal cover size, mean current velocity, pool volume, substratum coarseness and cover area. There were no significant differences between the pools with salmon and those without salmon in terms of substratum coarseness and cover area. However, significant differences were found in mean water depth, maximal cover size, mean current velocity and pool volume. The analysis found that the pools with salmon were discriminated from those without salmon at a level of 93.5% which was determined by the six environmental variables. Our results revealed that the returning adult salmon selectively used the pools with greater depth, large cover and slow current velocity as their preferable summering habitat. The results also found that the summering habitat of salmon can be estimated by measuring only six environmental variables. Therefore, these variables can be considered useful parameters in determining the summering habitat components of returning adult masu salmon and can contribute to the conservation and restoration strategy of this species by providing a better understanding of the relationship between fish and their habitat.     

Effects of habitat availability on dispersion of a stream cyprinid

Synopsis We analyzed temporal changes in the dispersion of the rosyside dace,Clinostomus funduloides, (family Cyprinidae) in a headwater stream, to assess the role of habitat availability in promoting fish aggregation. The dace foraged alone and in groups of up to about 25 individuals, and dispersion varied significantly among monthly censuses conducted from May through December. In two of three study pools, dace aggregated during July, October and/or December, but spread out during other months, especially during September when dispersion did not differ significantly from random. Dispersion was not significantly correlated with the total amount of suitable habitat available to foraging dace, but during summer, corresponded to the availability of depositional areas adjacent to rapid currents. Foragers aggregated in eddies or depositional areas during high stream discharge in July, and shifted out of depositional areas when current velocities declined from July to September. During late autumn, however, aggregations formed independently of changes in habitat conditions, and dace dispersion did not vary significantly among months in a third pool. The study suggests that dace dispersion cannot be predicted from the overall availability of suitable habitat as estimated from point measurements of depth and velocity; both the occurrence of a specific habitat feature (i.e., eddies adjacent to high velocity currents) and seasonal differences in behavior more strongly influenced the spatial distribution of foragers.     

Influence of habitat structure and fish density on Atlantic salmon Salmo salar L. territorial behaviour

A laboratory stream experiment was performed to test the influence of young‐of‐ the‐year Atlantic salmon Salmo Salar density on habitat selection ( i.e . pools and riffles) and the combined effect of habitat and density on territoriality. Pools were preferred at low and medium fish density. At higher density, the saturation of the riffles led to an equal fish repartition in both habitats. Aggressive interactions increased according to fish density, and the level of aggression was significantly higher in the pools than in the riffles.     

Multi-scale effects of resource patchiness on foraging behaviour and habitat use by longnose dace, Rhinichthys cataractae

1. We examined the response of a predatory benthic fish, the longnose dace ( Rhinichthys cataractae ), to patchiness in the distribution of benthic macroinvertebrates on cobbles at three hierarchical spatial scales during summer and autumn 1996, and spring 1997 in a southern Appalachian stream. 2. At the primary scale (four to five individual cobbles separated by <1 m), the intensity of foraging was not correlated with the biomass of benthic macroinvertebrates/cobble, regardless of season. 3. At the secondary scale (i.e. foraging patches <5 m in diameter) we found that benthic macroinvertebrates were patchily distributed in summer, but not in autumn or spring. Concomitantly, in summer, longnose dace foraged on cobbles with a significantly higher biomass of benthic macronvertebrates than nearby, randomly selected cobbles with similar physical conditions (i.e. longnose dace tended to avoid low-prey foraging patches). In contrast, when benthic macroinvertebrates were distributed homogeneously (spring and autumn), dace did not select patches with a significantly higher biomass of benthic macroinvertebrates than that available on randomly selected cobbles. 4. At the tertiary scale (i.e. stream reaches 11–19 m long), the biomass of benthic macroinvertebrates (per cobble per reach) was patchily distributed (i.e. differed significantly among reaches) in all seasons. Among reaches with physical characteristics preferred by longnose dace, (i.e. erosional reaches dominated by cobble/boulder substratum and high current velocity), we detected a significant, positive correlation between the biomass of benthic macroinvertebrates/cobble and longnose dace density in all seasons. 5. Our results demonstrated that both spatial and temporal patchiness in resource availability influenced significantly the use of both foraging patches and stream reaches by longnose dace.     

Effects of habitat loss, habitat fragmentation, and isolation on the density, species richness, and distribution of ladybeetles in manipulated alfalfa landscapes

Abstract.  1. Habitat loss and fragmentation are the main causes of changes in the distribution and abundance of organisms, and are usually considered to negatively affect the abundance and species richness of organisms in a landscape. Nevertheless, habitat loss and fragmentation have often been confused, and the reported negative effects may only be the result of habitat loss alone, with habitat fragmentation having nil or even positive effects on abundance and species richness.
2. Manipulated alfalfa micro-landscapes and coccinellids (Coleoptera: Coccinellidae) are used to test the effects habitat loss (0% or 84%), fragmentation (4 or 16 fragments), and isolation (2 or 6 m between fragments) on the density, species richness, and distribution of native and exotic species of coccinellids.
3. Generally, when considering only the individuals in the remaining fragments, habitat loss had variable effects while habitat fragmentation had a positive effect on the density of two species of coccinellids and on species richness, but did not affect two other species. Isolation usually had no effect. When individuals in the whole landscape were considered, negative effects of habitat loss became apparent for most species, but the positive effects of fragmentation remained only for one species.
4. Native and exotic species of coccinellids did not segregate in the different landscapes, and strong positive associations were found most often in landscapes with higher fragmentation and isolation.
5. The opposing effects of habitat loss and fragmentation may result in a nil global effect; therefore it is important to separate their effects when studying populations in fragmented landscapes.     

Influences of microhabitat use and foraging mode similarities on intra- and interspecific aggressive interactions in a size-structured stream fish assemblage

Aggressive interactions, foraging behaviour and microhabitat use were observed among four sympatric stream fishes inhabiting the water column: ayu (Plecoglossus altivelis), white-spotted charr (Salvelinus leucomaenis), masu salmon (Oncorhynchus masou) and Japanese dace (Tribolodon hakonensis), each species being categorised into five body-size classes (species-size groups; SSG's). Aggressive interactions were observed between most pairs of SSG's, an almost linear dominance order being apparent throughout the three-month study period. Ayu were relatively subordinate in June, but became the second most dominant in July and the most dominant in August, as a consequence of a reversal in dominance order with salmon. In contrast, smaller-sized dace, which continually suffered from intra- and interspecific aggression, occupied the most subordinate ranks throughout the study period. Intensive aggression was observed among various SSG's, exhibiting same microhabitat propensity throughout the three months. The direction and frequency of aggressive interactions varied month by month due to a reversal in dominance order between ayu and masu salmon, and/or changes in density, body size and resource use of the component members. Opponent selectivity was higher within SSG's, where resource use was assumed to be highly overlapping, rather than among SSG's throughout the study period. Correlation analysis indicated that opponent selectivity in aggressive interactions among SSG's was positively correlated with similarity in microhabitat selectivity in June, but not in other months or with that in foraging habits, suggesting that intensive aggressive behaviour reflected overlapping habitat use among assemblage members during a certain period.     

The effects of introduced salmonids on two native stream‐dwelling salmonids through interspecific competition

Using an artificial stream, habitat use by two sympatric native salmonids in the presence and absence of introduced salmonid species was investigated experimentally. When only native white‐spotted charr Salvelinus leucomaenis and masu salmon Oncorhynchus masou were sympatric, they occupied different microhabitats. In the presence of introduced brown trout Salmo trutta or rainbow trout Oncorhynchus mykiss , however, white‐spotted charr and masu salmon were observed to use a similar habitat and interspecific competition between white‐spotted charr and masu salmon was initiated. The study suggested that the coexistence of native salmonids was negatively affected through interspecific competition between native and introduced salmonids.     

Stream fish assemblage and habitat structure in a tropical African river basin (Nyagui River, Zimbabwe)

Few studies have been carried out on stream ecology in southern Africa although many species are endangered. This study investigated the stream fish assemblage and their habitat associations over a period of 3 months (October 2004 to January 2005), in view of the proposal to build a dam across the Nyagui River. Twenty-four fish species were collected and were separated into groups based on preferred microhabitats. The first group, dominated by Barbus paludinosus , comprised species collected from the upstream stations with slow flow, shallow depth (pools) and fine substrate type. Species associated with riffles, which included Chiloglanis neumanni , Labeobarbus marequensis and Opsaridium zambezense , comprised the second group on the downstream. The last group comprised species preferring pools with rock substrate and slow flow such as Pharyngochromis acuticeps and Pseudocranilabrus philander . The species were consistently associated with their habitat types throughout the sampling period. This relationship may be explained by the fish's morphological adaptations. Species richness increased from nine in the upstream section to twenty in the downstream section and this was related to increasing habitat complexity downstream. The construction of the Kunzvi Dam across the Nyagui River is likely to lead to loss of rheophilic species while cichlids and introduced species may increase.     

Fish species associations in riffle habitat of streams of varying size and acidity in New Brunswick and Nova Scotia

Factors related to stream size and alkalinity were most influential in determining fish species associations in three catchments of New Brunswick (NB) and Nova Scotia (NS) Canada, as determined by discriminant function analysis of TWINSPAN stream site classification.
In the circumneutral Saint Croix (NB) catchment, the creek chub, Semotilus atromaculatus (Mitchill), brook trout, Salvelinus fontinalis (Mitchill), blacknose dace, Rhinichthys atratulus (Hermann), Atlantic salmon, Salmo salar Linnaeus, eel, Anguilla rostrata (Lesueur) and fallfish. Semotilus corporalis (Mitchill) dominated stream sites of progressively greater discharge and higher median mid-summer temperature. In the acidic Gold (NS) system, the brook trout, Atlantic salmon, and eel exhibited a distribution pattern in relation to stream size and temperature similar to that for the Saint Croix, with the eel relatively more abundant in the Gold at large stream sites. The creek chub was excluded from the smallest tributaries by low pH. The ranges of blacknose dace and fallfish do not extend to southwestern Nova Scotia. In the Medway (NS) system (slightly more acidic than the Gold), the relative abundance of Atlantic salmon is reduced, and that of eel increased as compared with the Gold and Saint Croix systems. The lower limiting mid-summer pH levels for creek chub, salmon, brook trout, and eel are 5.2, 5.0, 4.7 and <4.5 respectively.     

Flight activity of insects along a mountain stream: is directional flight adaptive?

1. Flight activity of Trichoptera, Plecoptera and Ephemeroptera was studied by sticky trapping for 12 months at five sites along a New Zealand mountain stream. Over 19 000 insects were captured by the traps, which were located in forest and grassland reaches, including a reach with intermittent flow.
2. Most species occurred predominantly in forest or grassland, although some were trapped throughout the stream. Longitudinal distributions of adults and their larvae were strongly correlated.
3. Flight periods of 24 caddisflies, three mayflies and four stoneflies ranged from 2 to 12 months. Six species were trapped in all months and 17 (55%) in more than 5 months.
4. The most abundant forest-dwelling caddisfly species were over-represented on the downstream sides of sticky traps located in, and immediately below, forest indicating a majority was flying upstream. Upstream flight compensates for downstream drift of larvae and should maximize the likelihood that forest-dwelling species will locate preferred habitat for egg, larval and/or adult development. Unlike the caddisflies, the stonefly Spaniocerca zelandica was over-represented on the upstream sides of traps, suggesting that some adults may float or fly downstream following emergence.
5. In contrast to forest-dwelling species, only one common caddisfly ( Oxyethira albiceps ) was over-represented on the downstream sides of traps at grassland sites. Unlike the forest-dwelling species, most species taken at the downstream sites probably came from a variety of sources, including a nearby stream.     

Strategies for the conservation and management of isolated salmonid populations: lessons from Japanese streams

1. Endangered native populations of stream salmonids in Japan face three major threats: (i) negative interactions with introduced hatchery‐reared fish, (ii) fragmentation of habitat by impassable dams and (iii) recreational angling. 2. To prevent imminent extinction of many local populations, we evaluated these threats and possible conservation actions for red‐spotted masu salmon (Oncorhynchus masou ishikawae) and white‐spotted charr (Salvelinus leucomaenis japonicus) in the Fuji River system in central Japan. 3. Red‐spotted masu salmon and white‐spotted charr occupied only 0.73 and 2.4% of suitable thermal habitats, respectively, with masu salmon typically occupying habitats closer to human population centres. 4. Population viability analysis resulted in a 100‐year probability of extinction of 78.1% for masu salmon and 48.1% for charr. However, extinction risk of both species was predicted to be <5% if the carrying capacity increased from 141 to 303 for masu salmon and from 94 to 125 for charr, by allowing fish passage at the lower end of the habitat, and if annual adult survival rate increased by 0.04. Adult survival rate was the principal factor associated with population persistence. 5. To conserve isolated populations of stream‐dwelling salmonids, we recommend (i) assessing the distribution of remnant native and non‐native fish populations, (ii) that fishing regulations are modified to improve adult survival and population persistence and (iii) that fragmented reaches be reconnected to adjacent habitat, for example by removing or modifying artificial barriers to increase the carrying capacity of the isolated populations. Reconnection of fragmented reaches should, however, be avoided if it results in non‐native fish invading isolated populations.     

The food web positioning and trophic niche of the non-indigenous round goby: a comparison between two Baltic Sea populations

Native species may show invasiveness toward a recipient ecosystem through increases in abundance as a result of artificial stocking events. Salmonid species are typical examples of native invaders whose abundance is increased after stocking with hatchery fish. This study evaluated the effects of hatchery chum salmon fry on sympatric wild masu salmon fry, benthic invertebrate prey, and algae, after a single stocking event in Mamachi stream, Hokkaido, northern Japan. The results suggested that the stocked hatchery chum salmon fry decreased the foraging efficiency and growth of the wild masu salmon fry through interspecific competition, and depressed the abundance of Ephemerellidae and total grazer invertebrates (Glossosomatidae, Heptageniidae, and Baetidae) through predation. Also, the hatchery chum salmon fry may increase algal biomass through depression of grazer abundance by predation (top-down effect). These results suggested that a single release of hatchery chum salmon fry into a stream may influence the recipient stream ecosystem.     

Effects of habitat structure on habitat use by Gammarus pulex in artificial streams

1. The influence of coarse substratum and flow, coarse substratum and food, and predation risk and flow on habitat use by Gammarus pulex was studied in three experiments conducted in artificial stream channels. Each stream channel consisted of a riffle and pool habitat.
2. Location of coarse substrata and food was manipulated by placing cobbles (coarse substratum) and leaf packs (food) in different habitats. Predation risk was varied by running experiments in the presence and absence of sculpins ( Cottus gobio ), and flow was varied by pumping water with one or two pumps.
3. In all experiments Gammarus were most abundant in pools but placement of cobbles in riffles increased use of the latter. An even greater percentage of Gammarus used riffles if leaf packs were also placed there. Decreased discharge and the presence of sculpins ( Cottus gobio ) also caused Gammarus to increase use of riffles. These data indicate that Gammarus is able to evaluate differences in habitat quality and respond accordingly.     

Oviposition habitat selection by the mosquito Culiseta longiareolata in response to risk of predation and conspecific larval density

Abstract. 1. Most female Culiseta longiareolata (Diptera: Culicidae) avoid ovipositing in pools that contain the predatory backswimmer Notonecta maculata . Such oviposition habitat selection has been suggested to reflect a trade-off between the risk of predation on larvae and potential density-dependent fitness costs. This putative trade-off was examined. In particular, evidence was sought in support of direct female response to local heterogeneity in habitat quality.
2. Three habitat types were established using artificial outdoor pools: predator pools, and non-predator pools with either low or high densities of Culiseta larvae. During each experimental night, females were offered one of the three possible pair-wise treatment combinations.
3. The majority (≈88%) of females oviposited in low-density pools rather than in the predator- or high-density pools. Furthermore, a substantially higher proportion of females oviposited in predator pools when faced with the high-density alternative, however this was due largely to fewer females ovipositing in high- vs low-density pools.
4. Females of a second mosquito species ( Culex laticinctus ), the larvae of which are at a lower risk of predation, were predicted to exhibit weaker aversion to N. maculata ; this prediction was supported only weakly.
5. Oviposition habitat selection by female C. longiareolata does not appear to involve a behavioural response that is based on individual assessment of local heterogeneity in relative pool quality, at least not at the spatial scale examined here; alternative explanations are discussed.     

Effects of habitat structure on habitat use by Gammarus pulex in artificial streams

1. The influence of coarse substratum and flow, coarse substratum and food, and predation risk and flow on habitat use by Gammarus pulex was studied in three experiments conducted in artificial stream channels. Each stream channel consisted of a riffle and pool habitat.
2. Location of coarse substrata and food was manipulated by placing cobbles (coarse substratum) and leaf packs (food) in different habitats. Predation risk was varied by running experiments in the presence and absence of sculpins ( Cottus gobio ), and flow was varied by pumping water with one or two pumps.
3. In all experiments Gammarus were most abundant in pools but placement of cobbles in riffles increased use of the latter. An even greater percentage of Gammarus used riffles if leaf packs were also placed there. Decreased discharge and the presence of sculpins ( Cottus gobio ) also caused Gammarus to increase use of riffles. These data indicate that Gammarus is able to evaluate differences in habitat quality and respond accordingly.     

Habitat-specific estimates of competition in stream salmonids: A field test of the isodar model of habitat selection

Summary The population densities of sympatric Atlantic salmon,Salmo salar and brook charr,Salvelinus fontinalis, were measured in riffle and pool stream habitats to test whether non-linear isodars, a multispecific model of habitat selection based on ideal distribution assumptions, could (1) predict the distribution of densities between habitats and (2) reproduce the processes postulated to underlie spatial segregation and species interactions in previous laboratory and field studies. The model provided a good fit to observed density patterns and indicated that habitat suitability declined non-linearly with increased heterospecific competitor densities. Competitive effects in riffles appeared to be due to exploitative resource use, with salmon always emerging as the superior competitor. No evidence was found for interference competition in riffles. In contrast, interspecific competition in pools seemed to occur through exploitation and interference. The specific identity of the superior competitor in pools depended on the density of both species; pools provided the charr with refuge from competition with the salmon, presumably through the adoption by the charr of density-dependent behaviours, such as schooling and group foraging, that mitigated the negative impact of the salmon. Charr were displaced from the riffles toward the pools as the total salmon density increased. The isodar analysis, based on limited density data, successfully reproduced the processes suggested to underlie spatial segregation in previous field and laboratory studies and provided new insights into how changes in competitor densities modify habitat suitability in this system.