Effects of fish species richness and assemblage composition on stream ecosystem function

Abstract –  Biodiversity is declining across aquatic ecosystems because of biological invasions and species extinctions. Because fishes have pervasive effects on ecosystems through species-specific food web interactions, alterations to species and functional richness, and composition of natural assemblages could have negative effects on aquatic ecosystem function. In this study, I tested the effects of fish species and functional richness, and assemblage composition on primary production (PPR), benthic invertebrate density, and benthic particulate organic matter (BPOM) in a 42-day experiment in artificial stream mesocosms. I found that fish species richness and assemblage composition were important predictors of PPR in stream mesocosms. However, the effect of species richness on PPR increased with time, suggesting that richness-related effects might strengthen as the magnitude of community-level interactions increases in ecosystems. There was no effect of fish species or functional richness or assemblage composition on benthic invertebrate densities or BPOM. These data provide additional support that fishes can be important regulators of ecosystem function in aquatic systems, and suggest that positive effects of fishes on ecosystems can be strengthened by increased species richness and composition of the assemblage. This study broadens the applicability of the biodiversity ecosystem-function literature to a new suite of taxa, supporting the overall hypothesis that ecosystem functions and services are likely to decline in response to species extinctions.     

Feeding territory and variations in behavioural modes of algae‐grazing fish Plecoglossus altivelis ryukyuensis (Ryukyu‐ayu) in subtropical island streams

Abstract – Social and foraging modes in fish often vary between individuals with different body sizes and between populations under different ecological conditions. We studied social and foraging behaviour of algae‐grazing fish Plecoglossus altivelis ryukyuensis (Ryukyu‐ayu) inhabiting Japanese subtropical island streams. Ryukyu‐ayu exhibited four behavioural modes: territorial, schooling, solitary and floating. Their body sizes differed among these modes. Territorial fish predominantly foraged on benthic algae, whereas schooling and solitary individuals frequently consumed drifting materials as well. Schooling fish were smaller than territorial fish, but, unlike some other algae‐grazing fish species, did not use schooling to gain access to food within territories. Territorial fish attacked smaller conspecifics but exhibited lateral display towards larger fish and schooling fish while occasionally attacking grazing gobies. Larger fish maintained larger feeding territories and occupied territories for longer periods than smaller fish did. This suggests that, in Ryukyu‐ayu, intra‐ and interspecific food competitions and relative body size can influence diverse behavioural modes and duration of territory occupation. We also found that Ryukyu‐ayu foraged more often and had larger feeding territories than ayu P. a. altivelis inhabiting temperate streams. We conclude that foraging strategies of Ryukyu‐ayu may have adapted to subtropical island streams, where algal productivity is much lower than that in temperate regions.     

Density dependence of herbivorous central stoneroller Campostoma anomalum in stream mesocosms

Herbivorous fish can have strong effects on stream ecosystem function by consuming primary producers and excreting limiting nutrients, but it is unclear whether they are resource limited. Thus, understanding factors regulating abundance of these fish might help predict ecosystem function. We used stream mesocosms to test whether populations of central stoneroller Campostoma anomalum exhibit density dependence across a range of typical densities and resource abundance found in Great Plains streams. We predicted that incrementally increasing stocking biomass from 3·7 to 24·9 g·m^?2 would reduce standing stocks of resources resulting in lower growth of stocked fish. Fish growth (over 41 days) was compared to initial stocking biomass and primary production as well as standing stocks of algae and invertebrates using regression analysis. Mean growth of individuals was negatively associated with stocking biomass ( = 0·55; P = 0·02), as predicted. Contrary to our prediction, increases in fish biomass led to increased primary productivity ( = 0·31, P = 0·07), but resulted in no relationship among algal filament lengths ( = 0·00; P = 0·34), algal biomass ( = 0·12; P = 0·19) or invertebrate biomass ( = 0·03; P = 0·30). Thus, density dependence occurred without an apparent reduction in food resources. We hypothesised that stoneroller growth was possibly limited by competition for high‐quality algae or invertebrates, or behavioural interactions causing interference competition.     

Are invasive and native trout functionally equivalent predators? Results and lessons from a field experiment

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Use of first‐order tributaries by brown trout (Salmo trutta) as nursery habitat in a cold water stream network

Many investigators have examined the importance of suitable in‐stream habitat and flow regime to salmonid fishes. However, there is much less known about the use of small (<5 l·s^?1 discharge) first‐order streams within a larger stream network by salmonids. The purpose of this study was to evaluate the use of small headwater streams by juvenile brown trout Salmo trutta in the Emmons Creek stream network in Wisconsin, USA, and to determine whether abundance was related to habitat variables in these streams. Fishes in eight spring‐fed first‐order streams were sampled during a 7‐month period using a backpack electroshocker and measured for total length. Habitat variables assessed included stream discharge, water velocity, sediment composition and the abundance of cover items (woody debris and macrophytes). Densities of YOY trout ranged from 0 to 1 per m² over the course of the study and differed among first‐order streams. Stepwise multiple regression revealed discharge to be negatively associated with trout density in spring but not in summer. All other habitat variables were not significantly related to trout density. Our results demonstrate the viability of small first‐order streams as nursery habitat for brown trout and support the inclusion of headwater streams in conservation and stream restoration efforts.     

Electrofishing efficiency in low conductivity neotropical streams: towards a non‐destructive fish sampling method

Rotenone sampling is the most efficient method for assessing the fish assemblage structure and species abundance of low conductivity Amazonian streams. It does, however, cause fish mortality and disturb aquatic ecosystem. The aim of this study was to search for a non‐destructive alternative. The efficiency of electrofishing was compared against complete removal using rotenone. This procedure was repeated in 12 streams dispersed throughout French Guiana to test for environmental and biological effects such as water conductivity, stream depth, fish family membership and body size. This study revealed that the efficiency of electrofishing was influenced by stream conductivity and stream depth, but not by fish family or body size. The electrofishing method might constitute an efficient alternative to using rotenone in smaller streams (below 25‐cm depth and above 43 μS cm^?1), whereas in deeper and/or slightly conductive streams, rotenone still remains the only method able to provide a quick and comprehensive picture of the fish assemblage.     

Conserving fluxes of terrestrial invertebrates to trout in streams: A first field experiment on the effects of cattle grazing

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Trait–environment relationships in Amazon stream fish assemblages

Studies of trait–environment relationships provide important tools for the prediction of the response of biological communities to environmental alterations. The Amazon basin presents enormous potential for the development of research on this type of relationship, given the diversity of both its fish fauna and the aquatic ecosystems this fauna inhabits. The present study investigated the association between local environmental variables and the functional traits of fish in 54 streams of six major Amazonian basins. We identified the relationship between the characteristics of the streams (channel morphology, channel habitat units, riparian vegetation cover, large woody fragments and instream cover for aquatic organisms) and fish traits related to locomotion, habitat use and feeding behaviour. The fish fauna of the broader, deeper and more slowly flowing streams was dominated by nektobenthic species that exploit autochthonous resources such as fish and invertebrates. In narrow, fast‐flowing streams, by contrast, there was a predominance of benthic fishes with varying feeding habits, including periphytivorous and invertivorous species. Narrow, shady streams were inhabited by nektonic species adapted for the exploitation of resources from the marginal vegetation. The results of this study contribute to the understanding of the association between the functional structure of fish assemblages and local environmental variables in Amazonian streams. We hope that these findings will stimulate further research into the natural variation in stream fish assemblages that will ensure the development of more effective management strategies that better protect these important aquatic ecosystems.     

Fish use of ecohydraulic‐based mesohabitat units in a low‐gradient Illinois stream: implications for stream restoration

• 1. A classification scheme for ecohydraulic‐based mesohabitat units was developed for a summer low‐flow period. Mesohabitat unit designations were based on the integration of three‐dimensional channel hydraulics, geomorphic maintenance processes of bed morphology, and biological resource needs of fish. Ecological relevance of the units was evaluated by a study of fish mesohabitat use patterns, and species relationships to feeding guild. By portraying the stream as a mosaic of hydraulic habitat patches that provide specific biotic resource needs, this study's aim was to advance how ecological information may be incorporated into the stream restoration design process.
• 2. Nine mesohabitat units were designated, including pool‐front, ‐mid, and ‐rear units, scour pool, simple and complex riffles, glide, submerged point bar, and channel expansion marginal deadwater. Physical habitat structure differed among the nine mesohabitat units by length, water depth, and bed slope and complexity. Fish were collected in specific unit volumes by use of prepositioned areal electrofishing devices, in which distinct patterns of fish mesohabitat use were observed.
• 3. A key finding was the differences in fish assemblages among the pool units, in which fish densities were greatest in the pool‐front and scour pool units. Also, fish density in the pool‐front unit was positively correlated with pool entrance slope. Biomass was greatest in the pool‐front and ‐mid units, and it was correlated with maximum mid‐pool depth. Density and biomass were generally lowest in the pool‐rear unit. Other unique relationships were also observed among the mesohabitat units.
• 4. Based on feeding guild, patterns of fish mesohabitat use were observed for this summer low‐flow period; insectivores dominantly used pool‐front and scour pool units, herbivores dominantly used complex riffle units, and piscivores used pool‐front and ‐mid units.
• 5. Useful ecological information was derived from fish species‐habitat relationships observed in this study, linking mesohabitat units with species requirements for food resources. Such findings support advancements to ecological design strategies for stream restoration that promote hydraulic habitat diversity.

Copyright © 2008 John Wiley & Sons, Ltd.     

Biological effectiveness of an inexpensive nature‐like fishway for passage of warmwater fish in a small Ontario stream

Few studies have been conducted to evaluate the effectiveness of nature‐like fishways, particularly in low gradient warmwater streams with diverse fish communities. We evaluated a nature‐like fishway that was installed to facilitate upstream passage at a low head dam on Indian Creek near Spencerville, Ontario, Canada. A passive integrated transponder (PIT) array was used to quantify attraction and passage efficiency for 391 PIT tagged warmwater fish, represented by seven species. Attraction efficiency for the three most common species, common shiner (Luxilus cornutus), creek chub (Semotilus atromaculatus) and white sucker (Catostomus commersonii), was 63.3%, 83.7% and 65.6%, respectively, and passage efficiencies were 5.1%, 38.4% and 25%, respectively. Creek chub were able to locate the fishway in less time than white sucker and common shiner; however, took longer to successfully pass. Manipulation of creek chub release locations was used to separate issues of attraction and passage and revealed that passage efficiency was highest (76.2%) for those released within the fishway and intermediate for those released at the entrance (42.1%). This multispecies fishway improved stream connectivity, but additional work is needed to fine tune its configuration. Similar projects that engage stakeholders in nature‐like fishway construction are a promising approach for the thousands of small dams that occur on low gradient streams around the globe, but those studies should incorporate a biological evaluation to ensure that attraction and passage efficiency are optimised.     

Associations between stream hydrogeomorphology and co‐dependent mussel–fish assemblages: evidence from an Ohio,USA river system

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Patterns in young‐of‐year smallmouth bass microhabitat use in multiple stream segments with contrasting land uses

Abstract Young‐of‐the‐year (YOY) smallmouth bass, Micropterus dolomieu Lacepéde, were evaluated in streams from eight catchments with two contrasting land uses to determine their use of microhabitats under a variety of stream conditions. Step‐wise discriminant function analyses revealed patterns of habitat use by discriminating used from available microhabitat conditions. Velocity was significant in 88% of streams sampled, whereas depth was significant in only the smallest stream in the forest‐dominated catchments and 75% of stream segments located in pasture‐dominated catchments. Mean velocities used by YOY bass were lower than available velocities, and mean depths used were greater than mean availability in all cases. Substrata varied significantly with availability in different stream segments. Error rates associated with classification ranged from 5 to 39%. Results indicate that YOY smallmouth bass are somewhat opportunistic, but use low‐velocity habitats in most cases and deeper water when streams are impacted by pasture land use and associated physical changes.     

Effects of food supply and stream physical characteristics on habitat use of a stream‐dwelling fish

Identifying the environmental factors that affect freshwater fish can be crucial for their conservation and management. Despite the widespread investigation of relationships between fish habitat use and environmental variables, there is a paucity of knowledge on how abiotic and biotic factors jointly influence stream fish habitat use. Three New Zealand South Island streams were selected to investigate the habitat preference of a stream‐dwelling galaxiid, banded kokopu (Galaxias fasciatus). Fish abundance in several permanent pools was determined by spotlighting at night once a month from June 2008 to May 2009. Drifting invertebrates and key physical features of each pool were measured at the time of fish sampling. An information‐theoretic approach (AIC) indicated that the most parsimonious candidate model to predict banded kokopu biomass was the one that included pool area, undercut banks, water velocity, overhanging vegetation, invertebrate drift density and an interaction term between invertebrate drift density and water velocity. Banded kokopu biomass was positively related to pool area, undercut banks, overhanging vegetation and invertebrate drift density. Our study suggests that fish resource use patterns need to be understood in the context of multiple interacting ecological factors, including prey abundance.     

Relationships between juvenile salmon, Salmo salar L., and invertebrate densities in the River Tana, Norway

Abstract –  Juvenile salmon density was related to invertebrate density in 13 streams within the River Tana, northern Norway. There were only small, nonsignificant, differences in benthic density between streams with and without juvenile salmon. All streams with a high density of juvenile salmon had low benthic densities at the stream mouth. Juvenile salmon were not found, or were in very low densities, in streams where the benthic density at the stream mouth was as high or higher than that in the stream. A multiple regression model showed that parr density was related negatively to benthic density at the stream mouth, water velocity and pH, and positively to benthic density within the stream and the proportion of the substratum covered by moss. The amount of overhanging cover in the different streams explained 93% of the variation in the drift density of terrestrial invertebrates in August. The highest densities of juvenile salmon were found in streams with riparian vegetation, and were thus associated with an abundant supply of drift food, especially terrestrial invertebrates.     

Influences of local habitat and stream spatial position on fish assemblages in a dammed watershed,the Qingyi Stream,China

Abstract – Identifying the underlying mechanisms that explain the spatial variation in stream fish assemblages is crucial for the protection of species diversity. The influences of local habitat and stream spatial position on fish assemblages were examined from first‐order through third‐order streams within a dammed watershed, the Qingyi Stream, China. Based on linear regression models, the most important environmental variables influencing fish species richness were water temperature and wetted width, but stream spatial position variables were less important. Using canonical correspondence analysis, five environmental variables were identified to significantly influence fish assemblages, including three habitats (elevation, substrate and water depth) and two spatial variables (C‐link and Link). Our results suggest that, in a heavily dammed watershed, by blocking the migration routes of fishes, dams weaken the influence of stream spatial position on fish species richness. However, fish species compositions are significantly influenced by both local habitat environment and stream spatial position, which is perhaps owing to the distribution of fish species according to ecological requirements not related to spatial processes.     

Effect of fine wood on juvenile brown trout behaviour in experimental stream channels

In‐stream wood can increase shelter availability and prey abundance for stream‐living fish such as brown trout, Salmo trutta, but the input of wood to streams has decreased in recent years due to harvesting of riparian vegetation. During the last decades, fine wood (FW) has been increasingly used for biofuel, and the input of FW to streams may therefore decrease. Although effects of in‐stream FW have not been studied as extensively as those of large wood (LW), it is probably important as shelter for small‐sized trout. In a laboratory stream experiment, we tested the behavioural response of young‐of‐the‐year wild brown trout to three densities of FW, with trout tested alone and in groups of four. Video recordings were used to measure the proportion of time allocated to sheltering, cruising and foraging, as well as the number of aggressive interactions and prey attacks. Cruising activity increased with decreasing FW density and was higher in the four‐fish groups than when fish were alone. Foraging decreased and time spent sheltering in FW increased with increasing FW density. Our study shows that juvenile trout activity is higher in higher fish densities and that trout response to FW is related to FW density and differs from the response to LW as reported by others.     

Density‐dependent territory size and individual growth rate in juvenile Atlantic salmon (Salmo salar)

Whether territoriality regulates population size depends on the flexibility of territory size, but few studies have quantified territory size over a broad range of densities. While juvenile salmonids in streams exhibit density‐dependent mortality and emigration, consistent with space limitation, there has been relatively little study of how territory size and individual growth rate change over a broad range of densities, particularly in field experiments. Consequently, we manipulated the density (range = 0.25–8 m^?2) of young‐of‐the‐year (YOY) Atlantic salmon (Salmo salar) in mesh enclosures erected in a natural stream to test whether (i) territory size is fixed, decreases continuously or decreases towards an asymptotic minimum size as density increases; and (ii) individual growth rate decreases as a negative power curve with density as in observational field studies. Territory size decreased with increasing density, consistent with an asymptotic minimum size of about 0.13 m² for a 5‐cm fish. Individual growth rate also decreased with density, although the magnitude of decrease was steeper than in observational studies. Our results suggest a limit to how small territories can be compressed, which will set the upper limit to the local density in a habitat. The density‐dependent changes in territory size and individual growth rate will both play a role in the regulation of stream salmonid populations.     

The complementary role of lentic and lotic habitats for Arctic grayling in a complex stream‐lake network in Arctic Alaska

Lakes can be important to stream dwelling fishes, yet how individuals exploit habitat heterogeneity across complex stream‐lake networks is poorly understood. Furthermore, despite growing awareness that intermittent streams are widely used by fish, studies documenting the use of seasonally accessible lakes remain scarce. We studied Arctic grayling (Thymallus arcticus) in a small seasonally flowing (June–October) stream‐lake network in Alaska using PIT telemetry. Overall, 70% of fish visited two lakes, 8% used a single lake, and 22% used only stream reaches. We identified five distinct behavioural patterns that differed in dominant macrohabitat used (deep lake, shallow lake or stream reaches), entry day into the network and mobility. Some juvenile fish spent the entire summer in a shallow seasonally frozen lake (average 71 days), whereas others demonstrated prospecting behaviour and only entered the stream channel briefly in September. Another group included adult and juvenile fish that were highly mobile, moving up to 27 km while in the 3‐km stream‐lake network, and used stream reaches extensively (average 59 days). Lentic and lotic habitats served differing roles for individuals, some fish occupied stream reaches as summer foraging habitat, and other individuals used them as migration corridors to access lakes. Our study emphasises the importance of considering stream‐lake connectivity in stream fish assessments, even to shallow seasonally frozen habitats not widely recognised as important. Furthermore, we demonstrate that individuals may use temporary aquatic habitats in complex and changing ways across ontogeny that are not captured by typical classifications of fish movement behaviour.     

Fragmentation and drying ratchet down Great Plains stream fish diversity

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