Trophic consequences for riverine cyprinid fishes of angler subsidies based on marine‐derived nutrients

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Utility of fish scales from stock assessment surveys in stable isotope analysis for initial assessments of trophic relationships in riverine fish communities

The utility of using fish scales collected during stock assessment exercises to assess the trophic relationships of riverine fishes using their stable isotopes of d¹³C and d¹⁵N was tested using three riverine fish communities in England (Rivers Great Ouse, Ivel and Goyt). In each river, European barbel Barbus barbus was an important species, with other cyprinid species, including chub Squalius cephalus, present. Stable isotope analyses was completed using relatively small sample sizes per species (<11) from fish samples collected in 2001, 2005 and 2006 when up to 5 scales were collected from each fish. The calculation of standard ellipse areas (as a measure of trophic niche size) revealed that relative to other fishes, B. barbus occupied high trophic positions with minimal overlap in their trophic niche with other species, especially S. cephalus. As the analysed fish samples comprised species of various length ranges and as length has strong ontogenetic consequences for fish diet composition, generalized linear models were developed in which length was the covariate; model outputs included length‐adjusted mean δ¹³C and δ¹⁵N for each species. In each fish community, significant differences in δ¹³C and δ¹⁵N were apparent between B. barbus and S. cephalus, but were less apparent between B. barbus and other fishes. Thus, whilst the utility of using fish scales from stock assessments in stable isotope analyses are limited due to the differing length ranges of the sampled fishes, they can be useful in identifying trophic differences between species when methods such as stomach content analyses are unavailable.     

Using stable isotopes to test for trophic niche partitioning: a case study with stream salamanders and fish

1. Stream salamanders and fish often co‐occur even though fish prey on and outcompete salamanders. However, the mechanisms that allow palatable salamanders to coexist with fish are unknown. 2. We tested mechanisms in the field that promote coexistence between Idaho giant salamanders (Dicamptodon aterrimus) and stream salmonid fishes in headwater streams. Previous research in this system indicated that salamander dispersal did not promote coexistence with fish. We tested the hypothesis that D. aterrimus shift their diet when they occur with fish, facilitating coexistence through local niche partitioning. 3. We used nitrogen and carbon stable isotopes to describe the trophic niche of D. aterrimus and fish in three co‐occurring populations of salamanders and fish and three populations of salamanders without fish. We used two approaches to quantify trophic niche partitioning with stable isotopes: 95% kernel density estimators and isotopic mixing models. 4. We found that salamanders and fish were generalists that consumed aquatic invertebrates primarily, but both species were also cannibalistic and predatory on one another. We also found no support for trophic niche partitioning as a coexistence mechanism because there were no differences in the trophic niche metrics among salamander populations with and without fish. 5. Although we did not identify mechanisms that facilitate salamander and fish coexistence, our empirical data and use of novel approaches to describe the trophic niche did yield important insights on the role of predator–prey interactions and cannibalism as alternative coexistence mechanisms. In addition, we found that 95% kernel estimators are a simple and robust method to describe population‐level measure of trophic structure.     

Spawning strategies in cypriniform fishes in a lowland river invaded by non-indigenous European barbel Barbus barbus

Spawning strategies of lowland river fishes include single spawning, where reproduction generally occurs in early spring to provide 0+ fish with an extended growth season through the summer, but with a high risk of stochastic mortality events occurring, such as early summer floods. This risk can be reduced by multiple or protracted spawning strategies, where 0+ fish are produced over an extended period, often into mid-summer, but with the trade-off being a shorter growth season. The spawning strategies of cypriniform fish were explored in the River Teme, a spate river in Western England, which has non-indigenous European barbel Barbus barbus present. Sampling 0+ fish in spring and summer and across three spawning periods, B. barbus, chub Squalius cephalus and minnow Phoxinus phoxinus always revealed multiple spawning events, with 0+ fish of < 20 mm present in samples collected from June to August. Fish below 20 mm in August remained relatively small by the end of their growth season (October). For dace Leuciscus leuciscus, only single spawning events were evident, but with 0+ dace always being relatively large. Therefore, multiple spawning appears to be a common strategy that provides resilience in 0+ fish against stochastic mortality events in lowland rivers.

     

Trophic consequences of non‐native species for commercial fish in a backwater bay of the Three Gorges Reservoir,Southwest China

Trophic niche overlap in native and alien fish species can lead to competitive interactions whereby non‐native fishes outcompete indigenous individuals and eventually affect the viability of natural populations. The species Erythroculter mongolicus and Erythroculter ilishaeformis (belonging to the Culterinae), which are two commercially important fish species in the backwater bay of the Pengxi River in the Three Gorges Reservoir (TGR), were threatened by competition from the non‐native Coilia ectenes (lake anchovy). The latter is an alien species introduced into the lower reaches of the Yangtze River in China and now widespread in the TGR. The trophic consequences of non‐native lake anchovy invasion for E. mongolicus and E. ilishaeformis were assessed using stable isotope analysis (δ¹³C and δ¹⁵N) and associated metrics including the isotopic niche, measured as the standard ellipse area. The trophic niche of native E. mongolicus had little overlap (<15%) with the alien fish species and was significantly reduced in size after invasion by lake anchovy. This suggests that E. mongolicus shifted to a more specialized diet after invasion by lake anchovy. In contrast, the trophic niche overlap of native fish E. ilishaeformis with the alien fish species was larger (>50%) and the niche was obviously increased, implying that fish in this species exploited a wider dietary base to maintain their energetic requirements. Thus, marked changes for the native E. mongolicus and E. ilishaeformis were detected as the trophic consequences of invasion of non‐native lake anchovy.     

Comparative trophic impacts of two globally invasive cyprinid fishes reveal species‐specific invasion consequences for a threatened native fish

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Post‐stocking pallid sturgeon Scaphirhynchus albus growth,dispersal, and survival in the lower Missouri River

Hatchery augmentation has been used to mitigate declines in fish populations worldwide, especially for sturgeon species. Information regarding stocking success including survival, dispersal, and growth of sturgeon post‐stocking may refine sturgeon augmentation programs and facilitate recovery. Pallid sturgeon Scaphirhynchus albus populations have been supplemented by hatchery‐reared stocks for 25 years in the Missouri River, USA. We assessed survival, dispersal patterns, and growth characteristics of post‐stocked pallid sturgeon in the lower Missouri River. Hatchery‐reared pallid sturgeon stocked at age‐1 (4.1%) and > age‐1 (2.9%) were recaptured at a higher frequency than fish stocked at age‐0 (0.3%). Post‐stocking dispersal patterns suggested dispersal range increase as age increased, but individuals tended to remain in the same river segment as their stocking location. Growth rates varied by year class with younger year classes having truncated growth trajectories compared to older year classes. Post‐stock survival of pallid sturgeon varied by age‐at‐stocking and suggest age‐1 survival has declined through time. Augmentation of pallid sturgeon may benefit from considering dispersal from stocking location and by stocking older individuals which appear to have increased survival post‐stocking. A better understanding regarding environmental drivers of growth and specific habitat features used is needed to better predict optimal timing and location of future stockings.     

Niche partitioning and the effect of interspecific competition on microhabitat use by two sympatric galaxiid stream fishes

1. Numerous interacting abiotic and biotic factors influence niche use and assemblage structure of freshwater fishes, but the strength of each factor changes with spatial scale. Few studies have examined the role of interspecific competition in structuring stream fish assemblages across spatial scales. We used field and laboratory approaches to examine microhabitat partitioning and the effect of interspecific competition on microhabitat use in two sympatric stream fishes (Galaxias‘southern’ and Galaxias gollumoides) at large (among streams and among sites within streams) and small (within artificial stream channels) spatial scales. 2. Diurnal microhabitat partitioning and interspecific competition at large spatial scales were analysed among three sympatry streams (streams with allotopic and syntopic sites; three separate catchments) and four allopatry streams (streams with only allotopic sites; two separate catchments). Electro‐fishing was used to sample habitat use of fishes at 30 random points within each site by quantifying four variables for each individual: water velocity, depth, distance to nearest cover and substratum size. Habitat availability was then quantified for each site by measuring those variables at each of 50 random points. Diet and stable isotope partitioning was analysed from syntopic sites only. Diel cycles of microhabitat use and interspecific competition at small spatial scales were examined by monitoring water velocity use over 48 h in artificial stream channels for three treatments: (i) allopatric G. ‘southern’ (10 G. ‘southern’); (ii) allopatric G. gollumoides (10 G. gollumoides) and (iii) sympatry (five individuals of each species). 3. One hundred and ninety‐four G. ‘southern’ and 239 G. gollumoides were sampled across all seven streams, and habitat availability between the two species was similar among all sites. Galaxias‘southern’ utilised faster water velocities than G. gollumoides in both the field and in channel experiments. Both species utilised faster water velocities in channels at night than during the day. Diet differences were observed and were supported by isotopic differences (two of three sites). No interspecific differences were observed for the other three microhabitat variables in the field, and multivariate habitat selection did not differ between species. Interspecific competition had no effect on microhabitat use of either species against any variable either in the field (large scale) or in channels (small scale). 4. The results suggest that niche partitioning occurs along a subset of microhabitat variables (water velocity use and diet). Interspecific competition does not appear to be a major biotic factor controlling microhabitat use by these sympatric taxa at any spatial scale. The results further suggest that stream fish assemblages are not primarily structured by biotic factors, reinforcing other studies de‐emphasising interspecific competition.     

Fish, climate and the Gulf Stream: the influence of abiotic factors on the recruitment success of cyprinid fishes in lowland rivers

1. Climatic effects are increasingly being recognised as an important factor causing inter‐annual variability in organism abundances in aquatic and terrestrial ecosystems. This study investigated the relationships between water temperature (cumulative degree‐days >12 °C), river discharge (cumulative discharge‐days above basal discharge rate), the position of the North Wall of the Gulf Stream (NWGS), and the 0+ growth (September mean length) and recruitment success (year‐class strength, YCS) of three species of cyprinid fishes in two contrasting English lowland rivers, using a 21‐year dataset. 2. Contrary to the majority of studies on 0+ fishes, growth in the Yorkshire Ouse was most significantly correlated with river discharge, with water temperature of less importance. By contrast, temperature was more influential than discharge in the River Trent, possibly because of its regulated hydrological regime, although none of the relationships were statistically significant for this river. 3. Year‐class strength of roach (Rutilus rutilus) was positively correlated with the position of the NWGS, and there was evidence of synchrony in recruitment success between rivers, but the relationships were poorer for chub (Leuciscus cephalus) and dace (Leuciscus leuciscus). The strongest relationships between YCS and discharge during specific time periods were for when the fish were in their early (especially larval) developmental stages, although none of the relationships were statistically significant because of inter‐annual variations in river discharge relative to the timing of fish hatching. 4. Fishes are key predators in the majority of aquatic ecosystems and, as such, fluctuations in their abundances can have implications for ecosystem functioning as a whole. This study has demonstrated an underlying influence of broad‐scale climatic effects on the recruitment of riverine fishes, in spite of local variations in biotic and abiotic conditions. The relative importance of various abiotic factors on the recruitment success of riverine cyprinid populations varies spatially and temporally. For example, river discharge is likely to be of relatively greater importance in poorly‐structured rivers or those that are prone to large and rapid fluctuations in flow, while temporal variations occur because of inter‐annual differences in river discharge relative to the timing of fish hatching. Biotic factors may also be important determinants of fish recruitment success, especially in rivers with stable and predictable flow regimes.     

River habitat homogenisation enhances trophic competition and promotes individual specialisation among young of the year fish

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Predicting the contributions of novel marine prey resources from angling and anadromy to the diet of a freshwater apex predator

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Eutrophication mediates rapid clonal evolution in Daphnia pulicaria

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Aquatic food‐web dynamics following incorporation of nutrients derived from Atlantic anadromous fishes

Changes in the isotopic composition (δ¹³C and δ¹⁵N) in biofilm, macro‐invertebrates and resident salmonids were used to characterize temporal dynamics of marine derived nutrients (MDNs) incorporation between stream reaches with and without MDN inputs. Five Atlantic rivers were chosen to represent contrasting MDN subsidies: four rivers with considerable numbers of anadromous fishes; one river with little MDN input. Rainbow smelt Osmerus mordax, alewife Alosa pseudoharengus, sea lamprey Petromyzon marinus and Atlantic salmon Salmo salar, were the primary anadromous species for the sampled rivers. Regardless of the spatial resolution or the pathway of incorporation, annual nutrient pulses from spawning anadromous fishes had a positive effect on isotopic enrichment at all trophic levels (biofilm, 1·2–5·4‰; macro‐invertebrates, 0·0–6·8‰; fish, 1·2–2·6‰). Community‐wide niche space shifted toward the marine‐nutrient source, but the total ecological niche space did not always increase with MDN inputs. The time‐integrated marine‐nutrient resource contribution to the diet of S. salar parr and brook trout Salvelinus fontinalis ranged between 16·3 and 36·0% during anadromous fish‐spawning periods. The high degree of spatio‐temporal heterogeneity in marine‐nutrient subsidies from anadromous fishes lead to both direct and indirect pathways of MDN incorporation into stream food webs. This suggests that organisms at many trophic levels derive a substantial proportion of their energy from marine resources when present. The current trend of declining anadromous fish populations means fewer nutrient‐rich marine subsidies being delivered to rivers, diminishing the ability to sustain elevated riverine productivity.     

Swimming with the giant: coexistence patterns of a new redfin minnow Pseudobarbus skeltoni from a global biodiversity hot spot

Ecological niche theory predicts that coexistence is facilitated by resource partitioning mechanisms that are influenced by abiotic and biotic interactions. Alternative hypotheses suggest that under certain conditions, species may become phenotypically similar and functionally equivalent, which invokes the possibility of other mechanisms, such as habitat filtering processes. To test these hypotheses, we examined the coexistence of the giant redfin Pseudobarbus skeltoni, a newly described freshwater fish, together with its congener Pseudobabus burchelli and an anabantid Sandelia capensis by assessing their scenopoetic and bionomic patterns. We found high habitat and isotope niche overlaps between the two redfins, rendering niche partitioning a less plausible sole mechanism that drives their coexistence. By comparison, environment–trait relationships revealed differences in species–environment relationships, making habitat filtering and functional equivalence less likely alternatives. Based on P. skeltoni's high habitat niche overlap with other species, and its large isotope niche width, we inferred the likelihood of differential resource utilization at trophic level as an alternative mechanism that distinguished it from its congener. In comparison, its congener P. burchelli appeared to have a relatively small trophic niche, suggesting that its trophic niche was more conserved despite being the most abundant species. By contrast, S. capensis was distinguished by occupying a higher trophic position and by having a trophic niche that had a low probability of overlapping onto those of redfins. Therefore, trophic niche partitioning appeared to influence the coexistence between S. capensis and redfins. This study suggests that coexistence of these fishes appears to be promoted by their differences in niche adaptation mechanisms that are probably shaped by historic evolutionary and ecological processes.     

Patterns and drivers of fish extirpations in rivers of the American Southwest and Southeast

Effective conservation of freshwater biodiversity requires spatially explicit investigations of how dams and hydroclimatic alterations among climate regions may interact to drive species to extinction. We investigated how dams and hydroclimatic alterations interact with species ecological and life history traits to influence past extirpation probabilities of native freshwater fishes in the Upper and Lower Colorado River (CR), Alabama‐Coosa‐Tallapoosa (ACT), and Apalachicola‐Chattahoochee‐Flint (ACF) basins. Using long‐term discharge data for continuously gaged streams and rivers, we quantified streamflow anomalies (i.e., departure “expected” streamflow) at the sub‐basin scale over the past half‐century. Next, we related extirpation probabilities of native fishes in both regions to streamflow anomalies, river basin characteristics, species traits, and non‐native species richness using binomial logistic regression. Sub‐basin extirpations in the Southwest (n = 95 Upper CR, n = 130 Lower CR) were highest in lowland mainstem rivers impacted by large dams and in desert springs. Dampened flow seasonality, increased longevity (i.e., delayed reproduction), and decreased fish egg sizes (i.e., lower parental care) were related to elevated fish extirpation probability in the Southwest. Sub‐basin extirpations in the Southeast (ACT n = 46, ACF n = 22) were most prevalent in upland rivers, with flow dependency, greater age and length at maturity, isolation by dams, and greater distance upstream. Our results confirm that dams are an overriding driver of native fish species losses, irrespective of basin‐wide differences in native or non‐native species richness. Dams and hydrologic alterations interact with species traits to influence community disassembly, and very high extirpation risks in the Southeast are due to interactions between high dam density and species restricted ranges. Given global surges in dam building and retrofitting, increased extirpation risks should be expected unless management strategies that balance flow regulation with ecological outcomes are widely implemented.     

Trophic overlap between fish and riparian spiders: potential impacts of an invasive fish on terrestrial consumers

Studies on resource sharing and partitioning generally consider species that occur in the same habitat. However, subsidies between linked habitats, such as streams and riparian zones, create potential for competition between populations which never directly interact. Evidence suggests that the abundance of riparian consumers declines after fish invasion and a subsequent increase in resource sharing of emerging insects. However, diet overlap has not been investigated. Here, we examine the trophic niche of native fish, invasive fish, and native spiders in South Africa using stable isotope analysis. We compared spider abundance and diet at upstream fishless and downstream fish sites and quantified niche overlap with invasive and native fish. Spider abundance was consistently higher at upstream fishless sites compared with paired downstream fish sites, suggesting that the fish reduced aquatic resource availability to riparian consumers. Spiders incorporated more aquatic than terrestrial insects in their diet, with aquatic insects accounting for 45–90% of spider mass. In three of four invaded trout rivers, we found that the average proportion of aquatic resources in web‐building spider diet was higher at fishless sites compared to fish sites. The probability of web‐building and ground spiders overlapping into the trophic niche of invasive brown and rainbow trout was as high as 26 and 51%, respectively. In contrast, the probability of spiders overlapping into the trophic niche of native fish was always less than 5%. Our results suggest that spiders share resources with invasive fish. In contrast, spiders had a low probability of trophic overlap with native fish indicating that the traits of invaders may be important in determining their influence on ecosystem subsidies. We have added to the growing body of evidence that invaders can have cross‐ecosystem impacts and demonstrated that this can be due to niche overlap.     

Mortality induced by electrofishing and handling in five young‐of‐the‐year cyprinids: effect of the fish size,species and anode size

Immediate mortality induced by electrofishing and handling of young‐of the‐year (YOY) fish of five cyprinid species (bitterling Rhodeus amarus, chub Leuciscus cephalus, bleak Alburnus alburnus, barbel Barbus barbus, roach Rutilus rutilus) was quantified during routine YOY fish monitoring (point abundance sampling with backpack electrofisher, in situ identification and measurement). Sampled, identified and measured fish were held in screened containers immersed in the river for 5 h after electrofishing. Bleak exhibited significantly higher mortality (17%) than all other species (1–3%). Mortality decreased significantly with fish length; fish <35 mm exhibited the highest mortality. Anode size (10, 20, and 30 cm diameter) had no direct effect on mortality. Point abundance electrofishing with in situ identification and measurement is a safe method for sampling riverine YOY cyprinids.     

Temperature,invaders and patchy habitat interact to limit the distribution of a vulnerable freshwater fish

Interacting global‐change drivers such as invasive species and climate warming are likely to have major and potentially unexpected influences on aquatic ecosystems. In river networks, modified water temperature combined with patchy physical conditions will likely cause shifts in the amount and distribution of suitable habitat, with influential invasive species further altering habitat availability. We examined how distributions of a thermally sensitive galaxiid fish native to the alpine rivers of New Zealand, Galaxias paucispondylus, were influenced by these drivers using spatially extensive presence–absence electrofishing surveys of 46 sites spread over four subcatchments. A unimodal response to water temperature and an interaction with substratum size meant G. paucispondylus were limited to streams with average summer water temperatures between 10.6 and 13.8 °C and were absent when average substratum sizes were <36 mm, regardless of temperature. In addition, non‐native trout >150 mm long excluded G. paucispondylus, but were only found in streams with average summer water temperatures <10.6 °C. These influences of trout likely strengthened the unimodal temperature response of G. paucispondylus and led to a very small G. paucispondylus realized niche. When predicted temperature increases were applied to catchment models, G. paucispondylus distributions were patchy and variable across subcatchments. Moreover, local physical characteristics of river networks were particularly important because of the non‐linear and interactive influences of temperature and substratum size on the outcome of species interactions. Therefore, substratum sizes, water temperature and a non‐native predator combined to influence the distribution of this thermally sensitive fish, illustrating how the effects of climate warming will likely be strongly context‐dependent and interactive.     

Prey availability and selective feeding of sea trout (Salmo trutta L., 1758) fry stocked in small forest streams

The food base and stomach contents of stocked sea trout (Salmo trutta) fry were determined in the first 4 months (April–August 2010) of life in two small lowland streams after resorption of the yolk sac. In each stream, 600 individual trout were released in a 200 m stocking‐section. The macrozoobenthos given as a food base for the fry were collected once a month from the streams using a bottom scraper. Three subsamples of macrozoobentos were considered as one sample from the stocking‐section. On the same day, 50 stocked fry were captured in each stream using electric fishing gear. Preferred food components were usually the taxa represented by number in a given stream in a particular month. Width size range of the prey in fish stomachs in May was from 0.10 to 1.47 mm, and from 0.11 to 3.78 mm in August. All food items found in both streams during the study months were also represented in intensity in the fish: Cyclopoida, and larvae of Baetidae and Chironomidae. Almost all Helodidae and Simuliidae larvae as well as the majority of Limoniidae and Nemouridae larvae were available as food each month.     

Direct and indirect effects of predatory young‐of‐year fishes in a dryland river food web

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