A laboratory study of the effects of shelter availability and invasive crayfish on the growth of native stream fish

1. The alteration of stream habitats by urbanisation reduces the availability of shelter. Reduced shelter availability may increase both predation risks and metabolic costs, negatively affecting the growth performance of stream fish. Although urbanised streams often allow the establishment of invasive species, the additive or interactive effects of shelter availability and invasive species are rarely explored. The invasive red swamp crayfish (Procambarus clarkii) is a strong shelter competitor and predator in streams. 2. We conducted a stream‐channel experiment to investigate how shelter availability and this invasive crayfish species affect the growth performance (measured as the loss in body mass under starved conditions) of two native benthic species, Japanese crucian carp (Carassius auratus complex) and an endangered cobitid fish (Lefua echigonia). 3. We found that both the absence of shelter and the presence of crayfish increased body mass loss in the crucian carp. However, the growth performance of the crucian carp was improved by the presence of shelter except in the presence of crayfish, in which case the shelter available did not ameliorate the negative effect of the crayfish on growth performance (i.e. an interactive effect). This result suggests that crayfish may affect the growth performance of the crucian carp through chemical cues as well as through shelter occupancy. 4. In contrast, shelter availability is the primary factor affecting the body mass loss of the cobitid fish. However, the growth performance of these fish was unaffected by the presence of crayfish. The change in the growth performance of the cobitid fish in response to the absence of shelter was greater than that of the crucian carp. This finding suggests that the cobitid fish may be more vulnerable than the crucian carp to a reduction in shelter availability. 5. Our study demonstrates that reduced shelter availability and/or crayfish invasion can have significant, indirect negative effects on the growth performance of native fish, but whether those effects are interactive might vary depending on the species. Our findings have clear implications for stream restoration and habitat assessment.     

Shelters and their use by fishes on fringing coral reefs

Coral reef fish density and species richness are often higher at sites with more structural complexity. This association may be due to greater availability of shelters, but surprisingly little is known about the size and density of shelters and their use by coral reef fishes. We quantified shelter availability and use by fishes for the first time on a Caribbean coral reef by counting all holes and overhangs with a minimum entrance diameter ≥3 cm in 30 quadrats (25 m(2)) on two fringing reefs in Barbados. Shelter size was highly variable, ranging from 42 cm(3) to over 4,000,000 cm(3), with many more small than large shelters. On average, there were 3.8 shelters m(-2), with a median volume of 1,200 cm(3) and a total volume of 52,000 cm(3) m(-2). The number of fish per occupied shelter ranged from 1 to 35 individual fishes belonging to 66 species, with a median of 1. The proportion of shelters occupied and the number of occupants increased strongly with shelter size. Shelter density and total volume increased with substrate complexity, and this relationship varied among reef zones. The density of shelter-using fish was much more strongly predicted by shelter density and median size than by substrate complexity and increased linearly with shelter density, indicating that shelter availability is a limiting resource for some coral reef fishes. The results demonstrate the importance of large shelters for fish density and support the hypothesis that structural complexity is associated with fish abundance, at least in part, due to its association with shelter availability. This information can help identify critical habitat for coral reef fishes, predict the effects of reductions in structural complexity of natural reefs and improve the design of artificial reefs.     

Spatial distribution of limited resources and local density regulation in juvenile Atlantic salmon

1. Spatial heterogeneity of resources may influence competition among individuals and thus have a fundamental role in shaping population dynamics and carrying capacity. In the present study, we identify shelter opportunities as a limiting resource for juvenile Atlantic salmon (Salmo salar L.). Experimental and field studies are combined in order to demonstrate how the spatial distribution of shelters may influence population dynamics on both within and among population scales. 2. In closed experimental streams, fish performance scaled negatively with decreasing shelter availability and increasing densities. In contrast, the fish in open stream channels dispersed according to shelter availability and performance of fish remaining in the streams did not depend on initial density or shelters. 3. The field study confirmed that spatial variation in densities of 1-year-old juveniles was governed both by initial recruit density and shelter availability. Strength of density-dependent population regulation, measured as carrying capacity, increased with decreasing number of shelters. 4. Nine rivers were surveyed for spatial variation in shelter availability and increased shelter heterogeneity tended to decrease maximum observed population size (measured using catch statistics of adult salmon as a proxy). 5. Our studies highlight the importance of small-scale within-population spatial structure in population dynamics and demonstrate that not only the absolute amount of limiting resources but also their spatial arrangement can be an important factor influencing population carrying capacity.     

Density-dependent refuge use among over-wintering wild Atlantic salmon juveniles

Sheltering behaviour of wild juvenile Atlantic salmon in an indoor stream was found to be density dependent; the proportion of fish sheltering decreased significantly with increasing population density. The mean number of fish occupying refugia was ≤1.5 fish per refuge even at very high densities (potentially 5 fish per shelter). These results suggest that shelter availability has potentially important consequences for the carrying capacity for natural populations of salmon in streams.     

Effects of discharge and local density on the growth of juvenile Atlantic salmon Salmo salar

The study explored the combined effects of density, physical habitat and different discharge levels on the growth of juvenile Atlantic salmon Salmo salar in artificial streams, by manipulating flow during both summer and winter conditions. Growth was high during all four summer trials and increased linearly with discharge and mean velocity. Differences in fish densities (fish m^?3) due to differences in stream volume explained a similar proportion of the variation in mean growth among discharge treatments. Within streams, the fish aggregated in areas of larger sediment size, where shelters were probably abundant, while growth decreased with increasing densities. Fish appeared to favour the availability of shelter over maximization of growth. Mean growth was negative during all winter trials and did not vary among discharge treatments. These results suggest that increased fish densities are a major cause of reduced summer growth at low discharge, and that habitat‐mediated density differences explain the majority of the growth variation across habitat conditions both during summer and winter.     

Does Channel Incision Affect In‐stream Habitat? Examining the Effects of Multiple Geomorphic Variables on Fish Habitat

Incised river channels are dynamic components of fluvial systems, represent geomorphic degradation, and are encountered worldwide. Ecological effects of incision can be far‐reaching, affecting habitat availability and channel processes. Although incision can reflect habitat degradation, some studies suggest that important in‐stream habitats do not differ with the degree of incision. Therefore, we tested whether in‐stream habitat variables that are important to imperiled fishes differ in river reaches with varying degrees of incision. Because incision (measured using entrenchment ratio) had no discernable effect on in‐stream habitat characteristics (i.e., proportion fines, gravel, cobble, and macrophyte occurrence and length), we expanded our analysis to assess the effects of 29 additional geomorphic variables on in‐stream habitat. These analyses indicated that bank height, bed mobility, D₈₄, cross‐sectional area, bankfull width, and wetted perimeter accounted for 42% of macrophyte occurrence and 64% of macrophyte length variance. Postflood surveys indicated that macrophyte occurrence on cobble declined as bank height and bed mobility increased, and sediment size decreased, suggesting that sediment size and bed mobility have a stronger influence on in‐stream habitat than incision. Although channel incision often indicates environmental degradation, important aspects of habitat are not described by this measurement. Strategies that depend on incision to identify restoration sites may have limited habitat benefits in Southeastern Piedmont streams and rivers. Instead, landscape or shoal‐scale restoration approaches that increase coarse sediment proportions may increase macrophyte occurrence, length, and persistence. Sediment budgets that identify coarse and fine sediment sources and transport may be useful to prioritize restoration approaches.     

Short‐term Costs and Benefits of Habitat Complexity for a Territorial Fish

An increase in habitat complexity is thought to decrease visibility and the territory size of visually oriented animals. Hence, the addition of physical structure has been viewed as a useful restoration technique to increase the density of territorial species, particularly in stream fishes. However, a decrease in territory size may have a negative effect on the fitness of individual organisms. We attempted to quantify some of the positive and negative effects of increasing habitat structure on the behaviour and growth rate of wild young‐of‐the‐year (YOY) Atlantic salmon. Fish were exposed to one of two habitat treatments in mesh enclosures in Catamaran Brook, New Brunswick: a gravel substrate (low complexity) or a gravel substrate with boulders added (high complexity). Wild‐caught individuals were tagged, weighed and measured before being stocked at densities of 1·m^?2 for 7‐d trials. While fish from high‐complexity treatments benefited from lower rates of aggression, they also had lower foraging rates and smaller territories compared to those in low‐complexity treatments. Specific growth rate, however, did not differ significantly between treatments. While the addition of structure to a habitat may be beneficial at the population level in terms of an increase in population density, our results suggested that individual fish may pay some short‐term costs in these environments. Further research is needed to evaluate the longer term costs and benefits of adding structure to improve the habitat quality for stream salmonids.     

Effects of food availability on temporal activity patterns and growth of Atlantic salmon

1. Patterns of sheltering and activity are of fundamental importance in the ecology of animals and in determining interactions among predators and prey. Balancing decreased mortality risk when sheltering with increased feeding rate when exposed is believed to be a key determinant of diel patterns of sheltering in many animals. 2. Despite lower foraging efficiency at night than during the day, Atlantic salmon Salmo salar parr are nocturnal during winter and at low summer temperatures. Nocturnal activity also occurs at warm water temperatures during summer, but little is known about the functional significance of this behaviour. 3. This study aimed to determine: (1) the preferred activity and shelter pattern of Atlantic salmon parr during warm summer months, and (2) their response to variations in food availability when balancing growth rate (G) and mortality risk (M), as expressed through time out of shelter. We differentiated among four potential responses to reduced food availability: (1) no response; (2) G decreases but M remains constant; (3) G remains constant but M increases; and (4) G decreases and M increases. 4. Time exposed from shelter was inversely related to food availability. Fish subject to high food availability were significantly less active during the day than those with restricted rations. However, food availability had no significant effect on the extent to which fish were active at night. There was no evidence of variation in growth rate with food availability. 5. Salmon were predominantly nocturnal at high ration levels, consistent with their previously reported behaviour during winter. Rather than switching to diurnal behaviour at high temperatures per se, as previously was supposed, it appears that the fish are diurnal only to the extent needed to sustain a growth rate, and this extent depends on food availability. 6. Atlantic salmon parr modulate the amount of time they are active rather than growth when responding to variations in food availability over an order of magnitude.     

Linking fish trait responses to in‐stream habitat in reconstructed valley‐plugged stream reaches of the Coastal Plain,U.S.A.

Valley‐plug formation is a challenging consequence of stream channelization especially in physiographic regions with highly erodible soils. Upstream channel degradation and incision results in accelerated sediment delivery processes wherein downstream aggradation decreases stream power and creates sand‐clogged channels. Channel reconstruction is now meeting hydrogeomorphic goals related to valley‐plug remediation, yet there exists a need to understand how this practice also facilitates ecological restoration. We evaluated fish trait response to in‐stream habitat conditions in channelized, recently restored, and “least‐disturbed” reference reaches of Coastal Plain streams in West Tennessee. Restored reaches were ecologically similar to channelized reaches, having higher proportions of nest‐guarding omnivores that were correlated with higher percentages of pool habitats and lower wetted width:depth ratios compared to reference reaches. Reference reaches had higher proportions of fast‐water dwelling and specialized insectivores that were correlated with high abundance of large woody debris, high‐wetted width:depth ratios, and low percentages of pool habitats. We conclude that in‐stream habitats in reconstructed channels have yet to promote reach‐scale ecological restoration relative to fish assemblage organization because trait‐habitat associations were not similar to reference conditions. However, our results lend to the development of ecological restoration targets that can be incorporated in future channel reconstruction projects in valley‐plugged, channelized streams of the Coastal Plain.     

Shelter selection in juvenile Atlantic salmon,or why do salmon seek shelter in winter?

During winter, juvenile Atlantic salmon Salmo salar become nocturnal and seek refuge during the day in the stream bed gravel interstitial spaces. The function of this behaviour is unclear, but two major types of hypothesis have been proposed. One is that the fish are hiding from something (e.g. a predator) and the other is that the fish are seeking shelter from the water current. These hypotheses were tested by examining the selection of juvenile salmon for refuges that offered different degrees of concealment or shelter. The fish clearly preferred refuges that allowed them to hide (i.e. they were dark and opaque) but offered little shelter from the current. Therefore, it can be assumed that the primary function of this nocturnal behaviour during winter is most likely to hide from diurnal predators.