Is water temperature an adequate predictor of recruitment success in cyprinid fish populations in lowland rivers?

SUMMARY 1. Higher than average ambient water temperature in the first year of life may be responsible for strong cohorts of adult cyprinid fish. Whilst temperature explains much of the variation in year‐class strength (YCS), however, it is not the only influential factor as high temperature does not inevitably yield strong year‐classes. Furthermore, years in which a strong year‐class is prevalent in one species do not necessarily result in strong year‐classes in other coexisting species, suggesting other biotic and abiotic factors are important in regulating recruitment success. 2. The relationships between water temperature, river discharge, the position of the Gulf Stream, 0‐group fish growth and recruitment success (YCS) were examined in three cyprinid fish species in an English lowland river, using a 15‐year data set. 3. Mean length of 0‐group fish at the end of the summer was positively correlated with water temperature (cumulative degree‐days >12 °C) and negatively correlated with river discharge (cumulative discharge‐days above basal discharge rate). Water temperature was negatively correlated with river discharge. 4. YCS was positively correlated with mean 0‐group fish length at the end of the summer and with the position of the North Wall of the Gulf Stream. 5. ’Critical periods’ (i.e. periods in the first summer of life when fish may be more susceptible to increases in river discharge) were difficult to discern because of interannual variations in river discharge relative to the timing of fish hatching. YCS of roach and chub was most strongly correlated with discharge in the period from June to September inclusive, while YCS of dace was most significantly correlated with discharge in August. 6. River discharge (rather than water temperature) may be the key factor in determining YCS, either directly (through discharge‐induced mortality) or indirectly (via reduced growth at lower water temperatures, discharge‐associated increases in energy expenditure or reduced food availability). It could be that, in effect, water temperature determines potential YCS while discharge determines realised YCS.     

Fish recruitment in a large,temperate floodplain: the importance of annual flooding,temperature and habitat complexity

1. Large river floodplains are considered key nursery habitats for many species of riverine fish. The lower Volga River floodplains (Russian Federation) are still relatively undisturbed, serving as a suitable model for studying the influence of flooding and temperature on fish recruitment in floodplain rivers. 2. We examined the interannual variability in recruitment success of young‐of‐the‐year (YOY) fish in the lower Volga floodplain in relation to flood pulse characteristics and rising water temperatures in the spring. We sampled four areas with different flooding regimes, in three consecutive years (2006–2008). 3. Extensive areas with a long duration of flooding accommodated high densities of young fish. This suggests that extended inundation improves the recruitment success of river fish. In areas with extensive flooding, the biomass of YOY of most fish species was about three times higher in 2006 and 2007 than in 2008. We hypothesise that low spring temperatures in 2008 may have caused this reduced recruitment and that a flood synchronised with rising temperature enhances recruitment success. 4. Extensive flooding was particularly favourable for species characterised by large body size, delayed maturation, high fecundity and low parental investment, such as pike Esox lucius, roach Rutilus rutilus and ide Leuciscus idus. Gibel carp Carassius gibelio, a species tolerant of high temperature and hypoxia, did particularly well in small waterbodies in the driest parts of the floodplain. 5. Structural characteristics of floodplain waterbodies explained much of YOY fish density. These species–environment associations varied from year to year, but some species such as common bream Abramis brama, roach and gibel carp showed consistent relationships with structural habitat characteristics in all years, despite large interannual fluctuations in flood pulse and spring temperature.     

Characterizing the trophic niches of stocked and resident cyprinid fishes: consistency in partitioning over time,space and body sizes

Hatchery‐reared fish are commonly stocked into freshwaters to enhance recreational angling. As these fishes are often of high trophic position and attain relatively large sizes, they potentially interact with functionally similar resident fishes and modify food‐web structure. Hatchery‐reared barbel Barbus barbus are frequently stocked to enhance riverine cyprinid fish communities in Europe; these fish can survive for over 20 years and exceed 8 kg. Here, their trophic consequences for resident fish communities were tested using cohabitation studies, mainly involving chub Squalius cephalus, a similarly large‐bodied, omnivorous and long‐lived species. These studies were completed over three spatial scales: pond mesocosms, two streams and three lowland rivers, and used stable isotope analysis. Experiments in mesocosms over 100 days revealed rapid formation of dietary specializations and discrete trophic niches in juvenile B. barbus and S. cephalus. This niche partitioning between the species was also apparent in the streams over 2 years. In the lowland rivers, where fish were mature individuals within established populations, this pattern was also generally apparent in fishes of much larger body sizes. Thus, the stocking of these hatchery‐reared fish only incurred minor consequences for the trophic ecology of resident fish, with strong patterns of trophic niche partitioning and diet specialization. Application of these results to decision‐making frameworks should enable managers to make objective decisions on whether cyprinid fish should be stocked into lowland rivers according to ecological risk.     

Fish recruitment in rivers with modified discharge depends on the interacting effects of flow and thermal regimes

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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.     

Spatially variable response of native fish assemblages to discharge,predators and habitat characteristics in an arid‐land river

1. Fish assemblages and habitats were sampled annually at fixed sites in three tributaries of the Gila River catchment over a 21‐year span that included prolonged low‐ and high‐flow periods. Model selection was used to evaluate responses of seven native fishes with variable ecological traits (four small‐bodied cyprinids, one large‐bodied cyprinid, and two large‐bodied catostomids) to mean annual discharge and predacious non‐native fishes across the three sites. We also compared habitat use and overlap of native and non‐native fishes to identify potential for negative interactions among species. 2. Assemblage structure (species abundance and richness) and recruitment of native species was strongly and primarily affected by mean annual discharge and secondarily by location and densities of non‐native predators (mainly the centrarchid Micropterus dolomieui). 3. Densities of age‐0 catostomids and small‐bodied cyprinids were positively associated with discharge, and this pattern was strongest in the tributary with the lowest densities of non‐native predators. Absence or extreme low abundance of natives during low‐flow years was most pronounced at the sites where non‐native predators were comparatively common. Densities of adults of large‐bodied native species also varied by site, but often were positively associated with densities of non‐native predators. 4. Spatially variable responses of native fish assemblages indicated that the persistence of native fishes could be jeopardized if key habitats were lost or flow regimes unnaturally altered, particularly during low‐flow conditions when recruitment of native fishes is low and predation by non‐natives is high. Large‐bodied species may be less vulnerable to multiple years of poor conditions because adults are able to avoid predation by non‐natives and thus can rely on occasional high discharge years for successful recruitment. 5. As in other arid‐land streams, native fish assemblages of the Gila River Basin continue to decline. Our results indicate that conservation requires specific knowledge and consideration of physical influences as well as life‐history attributes of native and non‐native fishes.     

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.     

Molecular phylogeny of two lineages of Leuciscinae cyprinids (Telestes and Scardinius) from the peri-Mediterranean area based on cytochrome b data

We examined phylogenetic relationships in two lineages of Leuciscinae cyprinid fishes based on the sequence data of the complete mitochondrial DNA region coding for the cytochrome b gene (1140 bp). Telestes includes obligate riverine, moderately cold water-adapted species whereas Scardinius comprises warm-adapted species living in lowland lakes and still waters of rivers and streams. We also analysed selected representatives of Leuciscus and Phoxinellus because the taxonomic status of some species belonging to these genera is dubious and they could be placed in the genus Telestes. The study includes 18 species, 43 populations, and 111 individuals from 9 of the 14 peri-Mediterranean ichthyogeographic districts. Clades recovered from the phylogenetic analyses do not support previous taxonomic assumptions based on morphology. Telestes, Leuciscus, and Phoxinellus do not form monophyletic assemblages; phylogenetic analyses suggest that L. polylepis, L. turskyi, P. croaticus, and P. metohiensis should be included in Telestes. Similarly, populations of Scardinius erythrophthalmus do not cluster together and the endangered S. scardafa, endemic to central Italy and surviving in a single locality, is nested within them. The radiations of Telestes and Scardinius occurred in different time periods. A major diversification of Telestes is consistent with a sea dispersal during the freshwater Messinian "Lago Mare" phase of the Mediterranean Sea. Cladogenetic events within Scardinius are likely related to the extension and confluence of river drainages in lowlands following multiple lowering of the sea level during the Quaternary glaciations.     

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.     

Environmental change drives long‐term recruitment and growth variation in an estuarine fish

How individuals respond to environmental change determines the strength and direction of biological processes like recruitment and growth that underpin population productivity. Ascertaining the relative importance of environmental factors can, however, be difficult given the numerous mechanisms through which they affect individuals. This is especially true in dynamic and complex estuarine environments. Here, we develop long‐term otolith‐based indices of recruitment and growth for estuary perch Percalates colonorum (Bemm River, Australia), to explore the importance of intrinsic (individual, demographic) and extrinsic (hydrologic, climatic, density‐dependent) factors in driving estuarine fish productivity. Analyses involved a novel zero‐inflated specification of catch curve regression and mixed effects modelling. The 39 years of recruitment and 46 years of growth data, spanning a period of environmental change including severe drought, displayed considerable inter‐annual variation. Recruitment success was strongly related to high freshwater inflows during the spawning season, suggesting that these conditions act as spawning cues for adults and potentially provide favourable conditions for larvae. Individuals displayed age‐dependent growth, with highest rates observed at younger ages in years characterized by warm temperatures, and to a lesser degree, greater magnitude base inflow conditions. We detected systematic among‐year‐class growth differences, but these were not attributable to year class strength, suggesting that environmental conditions experienced by individuals as juveniles can have long‐lasting effects of greater importance to population productivity than density‐dependent growth responses. The primacy of temperature in driving growth variation highlights that under‐appreciated climatic variation can affect estuarine fish productivity through direct physiological and indirect food web mechanisms. We predict that climatic warming will promote individual growth in southerly populations of P. colonorum but concurrently limit recruitment due to forecast reductions in spawning season river discharge. Disparate trait responses are likely in other fishes as they respond to multiple and changing environmental drivers, making predictions of future population productivity challenging.     

Comparative use of side and main channels by small‐bodied fish in a large,unimpounded river

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

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Distributional patterns and community structure of Caribbean coral reef fishes within a river-impacted bay

This study examined how riverine inputs, in particular sediment, influenced the community structure and trophic composition of reef fishes within Rio Bueno, north Jamaica. Due to river discharge a distinct gradient of riverine inputs existed across the study sites. Results suggested that riverine inputs (or a factor associated with them) had a structuring effect on fish community structure. Whilst fish communities at all sites were dominated by small individuals (<20 cm), diversity and total biomass were reduced with increased proximity to the river mouth. The abundance of all fishes, but particularly small-bodied, juvenile and herbivorous fishes was reduced in turbid water when compared to clear-water sites. Results strongly suggest that fluvial sediment inputs may play an important role in structuring fish assemblages even under intense fishing pressure.     

Hierarchical analysis of genetic structure in the habitat‐specialist Eastern Sand Darter (Ammocrypta pellucida)

Quantifying spatial genetic structure can reveal the relative influences of contemporary and historic factors underlying localized and regional patterns of genetic diversity and gene flow – important considerations for the development of effective conservation efforts. Using 10 polymorphic microsatellite loci, we characterize genetic variation among populations across the range of the Eastern Sand Darter (Ammocrypta pellucida), a small riverine percid that is highly dependent on sandy substrate microhabitats. We tested for fine scale, regional, and historic patterns of genetic structure. As expected, significant differentiation was detected among rivers within drainages and among drainages. At finer scales, an unexpected lack of within‐river genetic structure among fragmented sandy microhabitats suggests that stratified dispersal resulting from unstable sand bar habitat degradation (natural and anthropogenic) may preclude substantial genetic differentiation within rivers. Among‐drainage genetic structure indicates that postglacial (14 kya) drainage connectivity continues to influence contemporary genetic structure among Eastern Sand Darter populations in southern Ontario. These results provide an unexpected contrast to other benthic riverine fish in the Great Lakes drainage and suggest that habitat‐specific fishes, such as the Eastern Sand Darter, can evolve dispersal strategies that overcome fragmented and temporally unstable habitats.     

The reproductive biology of an endemic cyprinid,Zacco pachycephalus,in Taiwan

Synopsis Reproductive parameters including body weight-length relationship, condition factor, gonadosomatic index (GSI) and fecundity of an endemic cyprinid,Zacco pachycephalus, were studied from October 1991 to January 1993. Although the condition factor was similar between both sexes, GSI values of female (3.72 to 14.24%) were significantly larger than that of males (0.21 to 2.23%) during the major breeding season. The number of yolky eggs per fish ranged from 459 to 2059. Ovum size distribution, annual GSI value, and recruitment of juvenile fish indicated thatZ. pachycephalus conducted two major spawning activities in spring (February to April) and summer (June to August). The larger GSI value and number of yolky eggs in spring compared to those in summer implies thatZ. pachycephalus has evolved a reproductive style by investing more energy in spring to avoid the damage of summer freshet on its recruitment success. The mating behavior ofZ. pachycephalus is similar to that of otherZacco fishes.     

On the Size of African Riverine Fish Faunas

SYNOPSIS. An apparent anomaly exists in the size of riverinefish faunas of the Nile and Zaire; the Nile while considerablylonger, appears depauperate when compared with the Zaire. Thisparadox is explained when discharge, not length, is used asa measure of river size. Indeed, the number of freshwater fishspecies in African rivers is more closely related to dischargethan to length or catchment area. Discharge is directly proportionalto terrestrial productivity of a river basin, which in turnaffects total biomass of fish and number of species. Changesin the size of rivers during the geologic past affected theircapacity to sustain diverse fish faunas. Rivers flowing throughespecially arid lands during the late- Pleistocene were reducedin discharge, and concomitantly, fish faunas. Immigration offish from refuge rivers during the Holocene partially restoredthese diminished faunas. We propose that fish are more mobilethan they seem, and that the distinctiveness of riverine fishfaunas may be maintained by competitive pressure from establishedresidents, rather than by limited dispersal abilities of fish.Theories of the distribution of fish in Africa are considered,and we suggest that discharge as affected by climatic stabilityis largely responsible for the size of African riverine fishfaunas.     

Hydrological connectivity for riverine fish: measurement challenges and research opportunities

1. In this review, we first summarize how hydrologic connectivity has been studied for riverine fish capable of moving long distances, and then identify research opportunities that have clear conservation significance. Migratory species, such as anadromous salmonids, are good model organisms for understanding ecological connectivity in rivers because the spatial scale over which movements occur among freshwater habitats is large enough to be easily observed with available techniques; they are often economically or culturally valuable with habitats that can be easily fragmented by human activities; and they integrate landscape conditions from multiple surrounding catchment(s) with in‐river conditions. Studies have focussed on three themes: (i) relatively stable connections (connections controlled by processes that act over broad spatio‐temporal scales >1000 km² and >100 years); (ii) dynamic connections (connections controlled by processes acting over fine to moderate spatio‐temporal scales ～1–1000 km² and <1–100 years); and (iii) anthropogenic influences on hydrologic connectivity, including actions that disrupt or enhance natural connections experienced by fish. 2. We outline eight challenges to understanding the role of connectivity in riverine fish ecology, organized under three foci: (i) addressing the constraints of river structure; (ii) embracing temporal complexity in hydrologic connectivity; and (iii) managing connectivity for riverine fishes. Challenges include the spatial structure of stream networks, the force and direction of flow, scale‐dependence of connectivity, shifting boundaries, complexity of behaviour and life histories and quantifying anthropogenic influence on connectivity and aligning management goals. As we discuss each challenge, we summarize relevant approaches in the literature and provide additional suggestions for improving research and management of connectivity for riverine fishes. 3. Specifically, we suggest that rapid advances are possible in the following arenas: (i) incorporating network structure and river discharge into analyses; (ii) increasing explicit consideration of temporal complexity and fish behaviour in the scope of analyses; and (iii) parsing degrees of human and natural influences on connectivity and defining acceptable alterations. Multiscale analyses are most likely to identify dominant patterns of connections and disconnections, and the appropriate scale at which to focus conservation activities.     

Variations in the spawning periodicity of eight fish species in three English lowland rivers over a 6 year period, inferred from 0+ year fish length distributions

The spawning periodicity of eight fish species was investigated in three English lowland rivers over a 6 year period from patterns in 0+ year fish standard length ( L _S) distributions. A single cohort of 0+ year dace Leuciscus leuciscus , roach Rutilus rutilus and perch Perca fluviatilis was observed each year, suggesting that these species spawned only once annually. By contrast, populations of chub Leuciscus cephalus , bleak Alburnus alburnus , bream Abramis brama , gudgeon Gobio gobio and minnow Phoxinus phoxinus were inferred to spawn on more than one occasion each year. Annual and intercatchment variations occurred in the L _S distribution patterns of some of the fish species. In chub, for example, although a minimum of two 0+ year cohorts occurred in all years in the River Trent, 'multiple' spawning (either at the individual or population level) was most apparent in 1999, 2003 and 2004. By contrast, 'multiple' spawning events were not evident in all years in the Warwickshire Avon and Yorkshire Ouse, with recruitment presumably based upon a single spawning event in some years. There is effectively a trade-off between early spawning (extended growing season), and the possibility that environmental conditions will impact upon recruitment success, and the potential for reduced overwinter survival of smaller individuals with lower lipid resources from later spawning events. Notwithstanding, fishes as small as 15 mm L _S survived the winter in some years, suggesting that progeny from later spawning events may make important contributions to fish recruitment success.     

Isolation and characterization of microsatellites in Leuciscus cephalus (Cypriniformes,Cyprinidae) and cross‐species amplification within the family Cyprinidae

Thirteen polymorphic microsatellites were isolated from Leuciscus cephalus, a widespread cyprinid species with great ecological tolerance. Together with the cross‐species amplification of six additional loci originally published for three cyprinid fish species, we optimized a multiplex panel for L. cephalus allowing the genotyping of 19 polymorphic loci. Number of alleles and heterozygosity per locus in a sample of 20 fish individuals ranged from two to 16 and from 0.05 to 0.90, respectively. These primers will be useful in determining the population structure of L. cephalus. In addition, successful cross‐amplification was obtained for several species of Cyprinidae.