Microhabitat use by two small benthic stream fish in a 2nd order stream

Direct observation of two benthic fish species — the stone loach (Barbatula barbatula) and the gudgeon (Gobio gobio) — in the field revealed a spatial segregation between the species in a study area with shallow riffle and moderately deeper pool sections. Stone loach generally inhabited shallow, more current-exposed locations and gudgeon preferred deeper, mostly sandy areas. The small individuals of both species were confined to shallow muddy locations and the larger individuals were found in deeper and more current exposed areas. The main factor affecting microhabitat choice was the flow regime:

it was a limiting factor for the stone loach, where the discharge rates controlled the presence of fish in the study area.

distribution patterns of both species were influenced by discharge and by fluctuations in discharge.

The following mechanisms regulating the distribution of stone loach and gudgeon are hypothesized:

1. Gudgeon: They prefer high water depth, low current velocities and sandy substrate, which strongly limits their spatial niche. Mainly relatively small individuals (size class 2; 6—9 cm) changed microhabitat in relation to environmental parameters. Size class 3 (approx. 9—12 cm) may be interpreted as a rather unpredictable transitory period between juvenile and adult stage. Large gudgeon entered the observation area mainly when discharge rates were high and variable. Increasing discharge rates increased the spatial niche of the large gudgeon.
2. Stone loach: At low flow rates, the observation area seemed to be an optimal place for the stone loach. Changes in environmental conditions are reflected in the distribution patterns. The transition between juvenile and adult microhabitat use takes place in size class 2. Under high and/or variable flow regime the species was not found in the observation area.

     

The response of stream fish to local and reach‐scale variation in the occurrence of a benthic aquatic macrophyte

1. The aquatic macrophyte Podostemum ceratophyllum has been shown to increase stream productivity, abundance and biomass of benthic invertebrates, and local occurrences of some stream fishes. However, experimental evidence that fishes preferentially associate with Podostemum is lacking, and the value of Podostemum as a predictor of stream fish assemblage composition has not been studied. 2. We conducted two short‐term (2 week), small‐scale (36 m²) experimental manipulations of Podostemum cover in the Conasauga River (Georgia and Tennessee, U.S.), and found higher abundances of benthic insectivorous fishes in patches with augmented (>80%) compared to reduced (7%) Podostemum cover. In an observational study, we quantified associations among percent cover of Podostemum, fish species richness, land cover, shoal length and base‐flow turbidity at 20 randomly selected shoals from a 39‐km reach that spanned a gradient of decreasing forest land cover. 3. Richness of all fish species and of lotic fishes peaked in the centre of the study reach, and richness was weakly correlated with predictor variables. Occupancy models for individual species also indicated that longitudinal position was a strong covariate for 13 of 19 species examined, with little support that Podostemum cover influenced occupancy. 4. Local associations may reflect choices by benthic fishes to utilise Podostemum, whereas downstream decline in fish species richness and Podostemum cover may reflect altered capacity of the system to support native species.     

Predictability of phenotypic differentiation across flow regimes in fishes

Fish inhabit environments greatly varying in intensity of water velocity, and these flow regimes are generally believed to be of major evolutionary significance. To what extent does water flow drive repeatable and predictable phenotypic differentiation? Although many investigators have examined phenotypic variation across flow gradients in fishes, no clear consensus regarding the nature of water velocity's effects on phenotypic diversity has yet emerged. Here, I describe a generalized model that produces testable hypotheses of morphological and locomotor differentiation between flow regimes in fishes. The model combines biomechanical information (describing how fish morphology determines locomotor abilities) with ecological information (describing how locomotor performance influences fitness) to yield predictions of divergent natural selection and phenotypic differentiation between low-flow and high-flow environments. To test the model's predictions of phenotypic differentiation, I synthesized the existing literature and conducted a meta-analysis. Based on results gathered from 80 studies, providing 115 tests of predictions, the model produced some accurate results across both intraspecific and interspecific scales, as differences in body shape, caudal fin shape, and steady-swimming performance strongly matched predictions. These results suggest that water flow drives predictable phenotypic variation in disparate groups of fish based on a common, generalized model, and that microevolutionary processes might often scale up to generate broader, interspecific patterns. However, too few studies have examined differentiation in body stiffness, muscle architecture, or unsteady-swimming performance to draw clear conclusions for those traits. The analysis revealed that, at the intraspecific scale, both genetic divergence and phenotypic plasticity play important roles in phenotypic differentiation across flow regimes, but we do not yet know the relative importance of these two sources of phenotypic variation. Moreover, while major patterns within and between species were predictable, we have little direct evidence regarding the role of water flow in driving speciation or generating broad, macroevolutionary patterns, as too few studies have addressed these topics or conducted analyses within a phylogenetic framework. Thus, flow regime does indeed drive some predictable phenotypic outcomes, but many questions remain unanswered. This study establishes a general model for predicting phenotypic differentiation across flow regimes in fishes, and should help guide future studies in fruitful directions, thereby enhancing our understanding of the predictability of phenotypic variation in nature.     

Temporal variation in foraging group structure of a size-structured stream fish community

Temporal variation in foraging group structure of a fish assemblage was examined in a flood-prone stream in southern Hokkaido, Japan. Foraging behaviour was observed underwater for four species which inhabit the water column: ayu, Plecoglossus altivelis, white-spotted charr, Salvelinus leucomaenis, masu salmon, Oncorhynchus masou, and Japanese dace, Tribolodon hakonensis, with each species being categorized into five size classes (species-size group; SSG). Based on foraging behaviour, each SSG of the fish assemblage was classified into one of four foraging groups: algae grazers, drift foragers, benthos-drift foragers, and omnivores, defined as SSG exhibiting similar foraging behaviour. All size classes of ayu, and of charr and salmon were categorized as algae grazers and drift foragers, respectively, throughout the study period. In contrast, size classes of dace were categorized as drift foragers, benthos-drift foragers, or omnivores with the same size classes often assigned to different foraging groups from month to month. Digestive tract contents of the fishes in the four foraging groups reflected their observed foraging behaviour, and foraging groups were therefore regarded as representing trophic groups. Abundance and membership of each foraging group varied in accordance with changes in abundance of SSG due to their growth, immigration, emigration, and/or mortality. Moreover, due to numerical dominance within the assemblage, plasticity in foraging behaviour of small- and medium-sized dace also played a key role in determining variability in the foraging group structure. Relative frequencies of two types of foraging behaviour, algae nipping and benthos foraging, of the small-sized dace were significantly correlated with the level of each resource, whereas no significant relationship was detected between foraging frequencies of the medium-sized dace and either resource. Fluctuations in foraging group structure within this assemblage occurred through niche shifts of some component members and by changes in SSG composition.     

Local modification of benthic flow environments by suspension-feeding stream insects

Larval black flies often exhibit spatially aggregated distributions, and individuals within patches can potentially reduce the supply of suspended food particles to downstream neighbors by modifying local flow characteristics. We used hot-film anemometry to quantify the magnitude and spatial extent of flow modifications downstream from feeding Simulium vittatum larvae in a laboratory flume, and to determine whether temporal patterns of flow variation are related to movements of the larval feeding appendages. Mean velocity 1 mm downstream from feeding larvae was reduced by 75%, and the percent reduction in velocity diminished asymptotically with downstream distance. Reduced velocities were evident as much as 60 mm downstream from, and 3 mm to either side of, larvae. Turbulence intensity (i.e., the SD of the velocity time series) was generally higher in this region relative to control flow conditions. Three results demonstrate the major contribution of the larval feeding appendages (i.e., labral fans) to such flow modification. First, there was a minimal reduction in mean velocity 5 mm downstream from non-feeding larvae (i.e., with closed labral fans), whereas mean velocity at the same location was reduced markedly when larvae were feeding. Second, the power spectrum of the velocity time series exhibited greatest power at frequencies that corresponded to the frequency of labral fan motions. Third, fan flick times accounted for most of the variance in the velocity power spectrum. The large local flow modifications that we documented have potentially important consequences for the feeding performance and growth of individuals located within larval aggregations, and are likely to influence behavioral interactions and spacing patterns.     

Population genetic structure across dissolved oxygen regimes in an African cichlid fish

Ecological isolation is a process whereby gene flow between selective environments is reduced due to selection against maladapted dispersers, migrant alleles, or hybrids. Although ecological isolation has been documented in several systems, gene flow can often be high among selective regimes, and more studies are thus needed to better understand the conditions under which ecological gradients or divergent selective regimes should influence population structure. We test for ecological isolation in a system in which high plasticity occurs with respect to traits that are adaptive in alternate forms under different environmental conditions. Pseudocrenilabrus multicolor victoriae is a widespread haplochromine cichlid fish in East Africa that exploits both normoxic (normal oxygen) rivers/lakes and hypoxic (low oxygen) swamps. Here, we examine population structure, using mitochondrial DNA and microsatellites, to determine if genetic divergence is significantly increased between dissolved oxygen regimes relative to within them, while controlling for geographical structure. Our results indicate that geographical separation influences population structure, while no effects of divergent selection with respect to oxygen regimes were detected. Specifically, we document (i) genetic clustering according to geographical region, but no clustering according to oxygen regime; (ii) higher genetic variation among than within regions, but no effect of oxygen regime on genetic variation; (iii) isolation by distance within one region; and (iv) decreasing genetic variability with increasing geographical distance from Lake Victoria. We speculate that plasticity may be facilitating gene flow between oxygen regimes in this system.     

Microhabitat preferences of benthic fauna in a woodland stream

Estimates of numbers, biomass, and diversity of benthic macroinvertebrates were made quarterly over a two-year period to investigate microhabitat preferences. Although biomass of most taxa was significantly different among sampling times, physical factors also appeared to be important in determining abundance of many taxa. Optimum depth, velocity, substrate type, and turbulence were determined for major taxa. Optimum conditions for diversity appeared to be 34 cm depth, 60 cm s^?1 velocity, and rubble and boulder substrate type. Habitat preference functions were derived for several taxa based on significant polynomial regressions of biomass on depth, velocity, substrate, and Froude number (turbulence). The relationship between abundance and physical habitat conditions was tested by using the product of the preference factors (range: 0–1) for depth, velocity and substrate type as a measure of habitat suitability (joint preference factor). There were significant correlations between biomass [transformed by log_e (x + 1)] of 10 benthic species and the joint preference factor. The joint preference factors accounted for from 11 to 61% of the variation of biomass of the 10 benthic species. The intercepts of the relationships between biomass of individual species and the joint preference factor were not significantly different from zero for any species. Therefore, the joint preference factors appear to be valid indicators of biomass. The preference functions have utility in habitat assessment studies, specifically with regard to minimum instream flow determinations.     

Prey selection by two benthic fish species in a Mato Grosso stream, Rio de Janeiro, Brazil

Key to understand predator choice is the relationship between predator and prey abundance. There are few studies related to prey selection and availability. Such an approach is still current, because the ability to predict aspects of the diet in response to changes in prey availability is one of the major problems of trophic ecology. The general objective of this study was to evaluate prey selection by two species (Characidium cf. vidali and Pimelodella lateristriga) of the Mato Grosso stream, in Saquarema, Rio de Janeiro, Brazil. Benthos and fishes were collected in June, July and September of 2006 and January and February of 2007. Fish were collected with electric fishing techniques and benthos with a surber net. Densities of benthic organisms were expressed as the number of individuals per/m2. After sampling, the invertebrates were fixed in 90% ethanol, and, in the laboratory, were identified to the lowest taxonomical level. Approximately, seventy individuals from each species were selected randomly in each month. Fishes were fixed in 10% formalin in the field and transferred to 70 degrees GL ethanol in the laboratory. Fishes had their stomachs removed for subsequent analysis. Fish diet was described according to the numeric frequency method. The Manly Electivity Index was applied in order to verify prey selection. The most abundant families in both benthos and diet of both fish species were the same, indicating that these species consume mainly most abundant prey in the environment. We concluded that prey selection occurs even for preys that had small abundance in the environment. However, it is the availability of the macroinvertebrate resources that determines the major composition of items in diet of fish, demonstrating that the abundance is the factor that most influences the choice of prey.     

Recruitment variation in a stream galaxiid fish: multiple influences on fry dynamics in a heterogeneous environment

1. The relative importance of density‐dependent and density‐independent processes in explaining fluctuations in natural populations has been widely debated. In particular, the importance of larval supply and whether it may control the type of regulatory processes a population experiences has proved contentious. 2. Using surveys and field experiments conducted in streams in Canterbury, New Zealand, we investigated how variation in the survival of non‐migratory Galaxias vulgaris fry was affected by density‐dependent and density‐independent processes and how this variation influenced recruitment dynamics. 3. Fry populations with high settlement densities experienced a 70–80% reduction in population size from density‐related mortality during the first fourteen days after peak settlement but thereafter the influence of density‐dependent processes on fry was weak. The impact of environmental conditions on fry populations was dependent on fry size and the magnitude of the perturbation, such that flooding effects on fry survival were most severe when fry were small. 4. In streams not affected by flooding, the size and density of introduced trout (Salmo trutta and Oncorhynchus mykiss) were the most significant factors determining the abundance of eventual recruits. A field experiment manipulating brown trout access to fry populations revealed that trout as small as 110 mm may be capable of greatly reducing and possibly preventing galaxiid recruitment. 5. Overall, the results indicated density‐dependent population regulation was only possible at sites with high native fish densities because trout were likely to be suppressing the number of potential recruits at sites with low native fish numbers. Whilst density‐dependent processes had a strong effect on fry survival following the period of peak fry abundance, density‐independent processes associated with flow and predatory trout influences on fry survival largely determined recruitment variability among galaxiid populations. Focusing conservation efforts on improving habitat to increase fry retention and reducing the impacts of trout on galaxiids would ensure more native fish populations reached their potential abundance.     

Giving‐up densities and ideal pre‐emptive patch use in a predatory benthic stream fish

1. We used observational and experimental field studies together with an individual‐based simulation model to demonstrate that behaviours of mottled sculpin (Cottus bairdi) were broadly consistent with the expectations of Giving‐Up Density theory and an Ideal Pre‐emptive Distribution habitat selection model. 2. Specifically we found that: (i) adult mottled sculpin established territories within patches characterised by significantly higher prey densities and prey renewal rates than patches occupied by juveniles or randomly selected patches; (ii) patches abandoned by adult sculpin possessed significantly lower prey densities than newly occupied patches, although this was not true for juveniles; (iii) the observed giving‐up density (GUD) for adult sculpin (i.e. average prey density in patches recently abandoned) increased linearly with increasing fish size up to the average prey density measured in randomly selected patches (i.e. 350 prey items per 0.1 m²) and decreased with increasing sculpin density and (iv) juveniles rapidly shifted their distribution towards the highest quality patches following removal of competitively dominant adult sculpin. 3. These results provide the first evidence of the applicability of GUD theory to a stream‐dwelling organism, and they elucidate the underlying factors influencing juvenile and adult sculpin habitat selection and movement behaviours. Furthermore, optimal patch use, ideal pre‐emptive habitat selection and juvenile ‘floating’ provide behavioural mechanisms linking environmental heterogeneity in the stream benthos to density‐dependent regulation of mottled sculpin populations in this system.     

Microhabitat use in a stream fish assemblage

We examined microhabitat use among fishes in a 37-m section of Coweeta Creek, North Carolina. Numerical abundances of species changed substantially during the 17-month study period. Microhabitat availability, however, did not change markedly during our investigation. Seasonal principal component analyses of microhabitat availability and fish microhabitat use data extracted two main patterns of non-random microhabitat use. Clinostomus funduloides, Notropis eoccogenis and Semotilus atromaculatm were significantly over-represented in deep areas with low to intermediate velocities and depositional substrata. Campostoma oligolepis, Coitus bairdi, Etheostoma blennioides, Rhinichthys cataractae and Salmo gairdneri all occurred in intermediate to deep microhabitats with moderate to high velocities and erosional substrata. Five of seven species exhibited seasonal variation in microhabitat utilization, whereas six species displayed size-related variation in use. Size-related variation was probably ontogenetic. We attributed most seasonal changes in microhabitat use to variations in microhabitat availability.
We used canonical analysis of discriminants to identify factors maximizing interspecific differences in microhabitat use. This analysis indicated that species could be assigned to either a benthic or a water column guild. Species within a guild generally could not be differentiated statistically, whereas members of different guilds were readily separable. These patterns persisted throughout the study, despite changes in numerical abundances of assemblage members. There was no evidence of either exploitation or interference competition for microhabitat, consequently it is unlikely that spatial resources were limiting during our study.     

Spatial and temporal variation of fish assemblages in a subtropical small stream of the Huangshan Mountain

Spatial and temporal variation of fish assemblages were investigated seasonally from May 2007 to February 2008 across 11 study sites in a subtropical small stream, the Puxi Stream, of the Huangshan Mountain. Along the longitudinal gradient from headwater to downstream, fish species richness and abundance increased gradually, but then decreased significantly at the lower reaches. The highest species richness and abundance were observed in August and the lowest in February. Based on analysis of similarities (...     

Population size,habitat fragmentation,and the nature of adaptive variation in a stream fish

Whether and how habitat fragmentation and population size jointly affect adaptive genetic variation and adaptive population differentiation are largely unexplored. Owing to pronounced genetic drift, small, fragmented populations are thought to exhibit reduced adaptive genetic variation relative to large populations. Yet fragmentation is known to increase variability within and among habitats as population size decreases. Such variability might instead favour the maintenance of adaptive polymorphisms and/or generate more variability in adaptive differentiation at smaller population size. We investigated these alternative hypotheses by analysing coding-gene, single-nucleotide polymorphisms associated with different biological functions in fragmented brook trout populations of variable sizes. Putative adaptive differentiation was greater between small and large populations or among small populations than among large populations. These trends were stronger for genetic population size measures than demographic ones and were present despite pronounced drift in small populations. Our results suggest that fragmentation affects natural selection and that the changes elicited in the adaptive genetic composition and differentiation of fragmented populations vary with population size. By generating more variable evolutionary responses, the alteration of selective pressures during habitat fragmentation may affect future population persistence independently of, and perhaps long before, the effects of demographic and genetic stochasticity are manifest.     

Annual variation of community biomass is lower in more diverse stream fish communities

Anthropogenic influences have disproportionally affected freshwater ecosystems, and a loss of biodiversity is forecasted to greatly reduce ecosystem function and services. Loss of species may destabilize communities by limiting the stabilizing forces of compensatory dynamics and/or statistical averaging, both of which are effects that can buffer variation in aggregate community properties. Currently, support for positive diversity‐stability relationships stems from experiments with simple communities at small spatial and temporal scales, and application to natural communities is limited. Using a long‐term dataset of 35 stream fish communities matched with hydrologic data, we show that community stability (annual variation of standing biomass of fishes) was less variable in more species‐rich communities and was not associated with stream hydrology. Only the statistical averaging model of community stability was consistent with observed patterns of lower biomass variation in more species‐rich communities. Our findings suggest anthropogenically induced extirpation of vertebrate consumers may lower community biomass stability in complex ecosystems.     

Associations between stream fish and benthos across environmental gradients in southern Ontario, Canada

1. The relationship between fish and benthic invertebrate communities in wadeable streams in southern Ontario, Canada, was examined using three independent and spatially distinct data sets. 2. Associations between fish and benthos were always significant when benthos were identified to family and often at the level of phylum. Identification to genus improved the strength of the fish–benthos association in one case. In contrast, identification to species did not improve the strength of the fish–benthos association. Associations between benthos and fish were weaker for one study utilizing a ‘rapid’ bio-assessment protocol involving field sorting and identification, and a second study which utilized winter benthic collections. 3. In two surveys, stream temperatures were important to the distributions of both fish and benthos, while in a third survey, fish and benthos were primarily influenced by stream size. In all three studies, fish and benthos were associated with similar suites of environmental variables, suggesting that the fish–benthos associations in these streams was driven by corresponding environmental tolerances. 4. Although there was significant variation in the strength of the fish–benthos association which could be attributed to differences in sampling methodologies, the findings from the present study confirm that stream fish and benthos are significantly associated. Therefore, surveys of benthos can be used to make inferences on the condition of fish community composition.     

Individual and combined effects of fish predation and bed disturbance on stream benthic communities: a streamside channel experiment

1. The composition and spatiotemporal dynamics of biological communities are influenced by biotic processes, such as predation and competition, but also by physical disturbances, such as floods in running waters. However, the interplay of disturbance with predation is still poorly understood, especially in frequently disturbed streams. Further, different predator species can affect prey communities in different ways depending on their feeding mode and efficiency. 2. We investigated the individual and combined effects of flood‐induced bed disturbance and fish predation on the benthos for 4 weeks in 18 streamside channels fed by a flood‐prone New Zealand river. Bed movements caused by floods were simulated by tumbling the substratum in half the channels. Six channels each were stocked with introduced brown trout (Salmo trutta) or native upland bully (Gobiomorphus breviceps) or had fish excluded. We studied algal biomass and both invertebrate density and daytime activity on surface stones on several dates after the disturbance, invertebrate community composition in the substrata of the entire channels on day 28 and leaf decomposition rates over the 28‐day period. 3. Disturbance affected algal biomass and density, richness and activity of surface stone invertebrates, and overall density and richness of channel invertebrates. Presence or absence of fish, by contrast, did not influence overall invertebrate standing stocks when subsurface substrata were included but did affect invertebrate densities on surface stones in 45% of all analysed cases and invertebrate activity on surface stones in all cases. Leaf decomposition rates were not influenced at all by the experimental manipulations. 4. Native upland bullies featured more often than exotic brown trout in causing invertebrate density changes and equally often in causing changes to grazer behaviour. Overall, our results imply that fish predation can have strong effects on the benthic invertebrate community in frequently disturbed streams, especially via behavioural changes.     

Seasonal changes in morphological structuring in a guild of benthic stream fishes

Summary The ecomorphological model which attempts to identify important variation in the total morphology of groups of animals has been used with some success in communities of fishes. The applicability of the model to subsets of the community, such as guilds of ecologically similar species, is uncertain, and the effects of the possible changes in correlations among morphological variables caused by demographic changes across seasons is unknown. An ecomorphological model is applied to a guild of benthic stream fishes from the Tombigbee River, Mississippi. The model is found to describe structuring in the guild based on aspects of the trophic morphology and the length of the body. Morphologically distinct groups of species are described by the model. The effect of indeterminate growth seems not to be of sufficient magnitude to change groupings across seasons. However, species associations within morphological groups change seasonally. I conclude that ecomorphological models may be properly applied to community subsets, but the interpretation of the model must consider the nature of the data set with respect of seasonality.     

Habitat-Specific variation in eyespot size of a benthic cladoceran

Summary Pigmented eyespot size of the benthic cladoceran Simocephalus exspinosus was measured in individuals sampled from four freshwater ponds that differed in the extent of visually-oriented predation. In ponds with such predation (from fish, salamander larvae, and dragonfly nymphs), eyespot size was found to be significantly smaller, relative to body size, than in a pond without visually-oriented predation. Reduction in pigmented eyespot size may represent an adaptation to reduce vulnerability to predation.