Diverse environmental cues drive the size of reproductive aggregation in a rheophilic fish.

BACKGROUND: Animal migrations are periodic and relatively predictable events, and their precise timing is essential to the reproductive success. Despite large scientific effort in monitoring animal reproductive phenology, identification of complex environmental cues that determine the timing of reproductive migrations and temporal changes in the size of reproductive aggregations in relation to environmental variables is relatively rare in the current scientific literature. METHODS: We tagged and tracked 1702 individuals of asp (Leuciscus aspius), a large minnow species, and monitored with a resolution of one hour the size of their reproductive aggregations (counts of sexes present at the breeding grounds standardized by the sum of individuals in the season) over seven breeding seasons using passive integrated transponder tag systems. We examined the size of reproductive aggregations in relation to environmental cues of day number within a reproductive season (intra-year seasonality), water temperature, discharge, hour in a day (intra-day pattern), temperature difference between water and air, precipitation, atmospheric pressure, wind speed and lunar phase. A generalized additive model integrating evidence from seven breeding seasons and providing typical dynamics of reproductive aggregations was constructed. RESULTS: We demonstrated that all environmental cues considered contributed to the changes in the size of reproductive aggregations during breeding season, and that some effects varied during breeding season. Our model explained approximately 50% of the variability in the data and the effects were sex-dependent (models of the same structure were fitted to each sex separately, so that we effectively stratified on sex). The size of reproductive aggregations increased unimodally in response to day in season, correlated positively with water temperature and wind speed, was highest before and after the full moon, and highest at night (interacting with day in a season). Males responded negatively and females positively to increase in atmospheric pressure. CONCLUSION: The data demonstrate complex utilization of available environmental cues to time reproductive aggregations in freshwater fish and their interactions during the reproductive season. The study highlights the need to acquire diverse data sets consisting of many environmental cues to achieve high accuracy of interpretation of reproductive timing.

Fish scale stable isotopes as potential indicators of nutrient pollution: Exploring the response of roach (Rutilus rutilus) scale δ15N and δ13C to a gradient of land use disturbance.

To examine the suitability of fish scales as potential tracers of nutrient pollution, we analysed the nitrogen and carbon stable isotope values (δ15N and δ13C) in scales of a generalist fish species, roach Rutilus rutilus, collected from 22 Czech reservoirs covering wide gradients of catchment land use and nutrient enrichment. Using generalised additive mixed models in the first step and generalised linear mixed models in the second step, we evaluated the response of roach scale stable isotope values to catchment land use variables (percentage of agricultural land and human population density) and in-reservoir water quality variables. Roach scale δ15N values varied by 15 ‰ among the reservoirs and were strongly, linearly, and positively associated with the percentage of agricultural land in the reservoir catchments, pointing to agriculture as the dominant source of nitrogen pollution in the investigated systems. Roach scale δ13C values differed by 8 ‰ among the studied reservoirs and were not related to catchment land use variables or in-reservoir primary production (chlorophyll-a levels). Possible variation in roach foraging strategies (littoral versus pelagic) between reservoirs or the contrasting effects of eutrophication-related autotrophic and heterotrophic processes on baseline δ13C values may explain the lack of relationships between roach scale δ13C values and the explanatory variables. In summary, our findings show that fish scale δ15N values are sensitive bioindicators of catchment-derived anthropogenic nitrogen inputs to freshwater ecosystems. Because scales can be sampled in a nonlethal way and δ15N analysis is relatively inexpensive, we suggest that measuring the δ15N values of fish scales could be an effective method for monitoring nitrogen pollution in aquatic environments.

Different hosts in different lakes: prevalence and population genetic structure of plerocercoids of Ligula intestinalis (Cestoda) in Czech water bodies.

Ligula intestinalis (Linnaeus, 1758) is a tapeworm parasite with a worldwide distribution that uses a wide variety of fish species as its second intermediate host. In the present study, we investigated the prevalence and population genetic structure of plerocercoids of L. intestinalis in five common cyprinoid species, roach Rutilus rutilus (Linnaeus), freshwater bream Abramis brama (Linnaeus), white bream Blicca bjoerkna (Linnaeus), bleak Alburnus alburnus (Linnaeus), and rudd Scardinius erythrophthalmus (Linnaeus), collected in six water bodies of the Czech Republic (Milada, Most, Medard, Jordán, Rímov and Lipno). Of the six study sites, the highest frequency of parasitism was recorded in Lake Medard (15%). The overall prevalence rate among the species was as follows: roach > rudd ≥ freshwater bream > bleak > white bream. Two mitochondrial genes (cytb and COI) were used to compare the population genetic structure of parasite populations using selected samples from the five fish species. The results of the phylogenetic analysis indicated that all populations of L. intestinalis were placed in Clade A, previously identified as the most common in Europe. At a finer scale, haplotype network and PCoA analyses indicated the possible emergence of host specificity of several mtDNA haplotypes to the freshwater bream. Moreover, pairwise Fixation indices (FST) revealed a significant genetic structure between the parasite population in freshwater bream and other host species. Parasite populations in roach not only showed the highest rate of prevalence but also depicted a maximum number of shared haplotypes with populations from bleak and rudd. Our results suggest that recent ecological differentiation might have influenced tapeworm populations at a fine evolutionary scale. Thus, the differences in prevalence between fish host species in different lakes might be influenced not only by the parasite's ecology, but also by its genetic diversity.

Hydropeaking causes spatial shifts in a reproducing rheophilic fish.

The hydropeaking regime below hydropower facilities represents a serious threat to riverine fauna and may cause declines in populations living under its influence. However, the knowledge on direct fish responses to the threat of hydropeaking is limited. Here, we aimed to test whether the hydropeaking generated 12 km upstream may have a negative effect on the position of actively spawning rheophilic fish, asp, Leuciscus aspius. Two passive telemetry antenna arrays were used to record fish position on the spawning ground. We monitored the position of spawning fish (545, 764 and 852 individuals) in three one-month long spawning seasons in 2017-2019 and related the changes in detection probability on the two antenna arrays to flow conditions, temperature, time of a day and individual fish ID. The fish detection on the spawning ground was negatively affected by the flow change (both increase and decrease) in time. Moreover, the probability of fish detection was also influenced by water temperature, the time of the day and, as seen from the magnitude of individual random effect variability, the detection probability was rather individual-specific. Hydropeaking resulted in the change of spawning behaviour and likely caused interruption of spawning or shifting spawning outside the optimal area for egg development. We therefore advise to reduce the hydropeaking regime during the rheophilic fish spawning season under fisheries or conservation interests.

Stable isotope evidence from archived fish scales indicates carbon cycle changes over the four-decade history of the Rímov Reservoir (Czechia).

Using archived fish scale samples together with long-term monitoring data, this study investigates the potential of fish scales to record historical changes in the aquatic environment. We analysed stable carbon (δ13C) and nitrogen (δ15N) isotopes in the scales of two planktivorous cyprinid species collected from the meso-eutrophic Rímov Reservoir, Czechia, over its entire four-decade history (1979-2016). The δ13C of the fish scales varied greatly throughout the reservoir history. The lowest δ13C values were observed immediately after the reservoir was filled in 1979, indicating that fish production at that time was likely partially supported by 13C-depleted CO2 released from the inundated soil. During the 1980s, due to the high levels of phytoplankton production stimulated by high phosphorus inputs from the catchment, the δ13C values substantially increased. However, since 1990, the δ13C values have generally decreased, reflecting a gradual reduction in reservoir primary production caused by the decreasing input of phosphorus and increasing input of dissolved organic carbon from the catchment. The δ13C of fish scales was also used to reconstruct the CO2 concentration of the surface water. The reconstructed CO2 varied significantly during the four-decade history, but it was always below the air-equilibrium concentration, suggesting that the surface water of the reservoir has consistently absorbed atmospheric carbon. The fish-scale δ15N values remained relatively stable, while slightly increasing within three years after impoundment, likely because the nitrogen supply was high throughout the studied period. Our study contributes to the growing body of literature demonstrating that stable isotope analysis of archived biological samples is a promising approach for understanding historical trends in the biogeochemistry of aquatic environments. In particular, our results highlight the potential of δ13C in archived fish scales in reconstructing carbon cycle changes and evaluating human impacts on aquatic ecosystems.

Negative feedback concept in tagging: Ghost tags imperil the long-term monitoring of fishes.

Wildlife monitoring using passive telemetry has become a robust method for investigating animal migration. With increased use, this method progressively pollutes the environment with technological waste represented by so called ghost tags (PIT tags ending in the environment due to reproductive expulsions, shedding or animal mortality). However, their presence in the environment may lead to failed detections of living individuals. We used tagging data from studies of the asp Leuciscus aspius and the bleak Alburnus alburnus collected from 2014 to 2018 and located ghost tag positions on the monitored spawning site using portable backpack reader for their detection. We modelled virtual river-wide flat-bed antennas (widths 0.2, 0.4, 0.6 and 0.8 m) representing monitoring effort and estimated the probability of the presence of ghost tags within the antenna field. Of 3724 PIT tags used in the study, we detected on the spawning ground 173 ghost tags originating from long-term monitoring. The ghost tags accumulated in the environment in time, suggesting insufficient degradation rate or shift downstream from the research site. Number of ghost tags present on the spawning ground led to high probability of disabled readings of tagged fish passing through the antenna electro-magnetic field. We demonstrate how accumulated ghost tags may cause detection failures for focal species and incomplete data acquisition. We infer that intensive long-term monitoring using PIT tag technology may encumber future data acquisition or entail additional costs for clean-up.

The pros and cons of the invasive freshwater apex predator, European catfish Silurus glanis, and powerful angling technique for its population control.

Catfish have spread across Europe and several countries out of this region within the last decades. Basic knowledge of this apex predator has revealed concerns of invasive behaviour and questions regarding its utilization as a biomanipulation species. However, a method enabling its regulation to a required level has not yet been developed. We simulated the impact of angling on the catfish population by method of hook-lines in two post-mining lakes with a monitored population consisting of tagged individuals and in two reservoirs as reference sites. Further, the efficiency of hook-lines as a reducing device was examined and the economic aspects were determined. Catfish population in localities where the species is unwanted or invasive may be efficiently reduced to a harmless level by hook-lines and angling (depending on the approach of anglers). The most efficient time of the year seems to be spring to early summer with catch efficiency of 5.4 individuals per 10 baits in one day. The catch efficiency markedly decreased during the second part of the year and did not exceed 2.8 individuals per 10 baits in one day. Mean size of catfish had negative impact whereas catfish biomass had positive impact on the catch efficiency. Trophic status and number of catfish in the locality had no impact on the catch efficiency. According to model, 11-18 bait-days per 1 ha per season is efficient to decrease catfish population to 10% of the original size. Both angling and hook-lines are very simple, they are financially and time bearable mechanisms of catfish regulation in any condition. However, catfish play an important role as a biomanipulative species in many localities. In this case where catfish is beneficial, angling presents a real threat of population collapse and loss of the biomanipulative effect.

Nocturnal spawning as a way to avoid egg exposure to diurnal predators.

Animals that do not provide parental care have to secure the survival of their offspring by ensuring a safe reproductive environment or smart timing tactics. Nocturnal spawning behaviour of many fish species is an example of the latter behaviour in the animal kingdom and is hypothesized to provide a survival advantage to the eggs spawned during the night. In order to test the efficiency of the smart timing tactics in a freshwater fish, a study was carried out of the interaction of the rheophilic spawner (asp Leuciscus aspius) and the predator of its drifting eggs (bleak Alburnus alburnus) using passive telemetry. According to a model based on acquired data, asp laid 63% of its eggs at night, while vision-oriented bleak was present in 92% of the time during the day. This study gives support to the predator avoidance hypothesis, which expects animals to reproduce in a period when the probability of offspring predation is at its lowest.

Invasive round goby Neogobius melanostomus has sex-dependent locomotor activity and is under-represented in catches from passive fishing gear compared with seine catches.

The higher proportion of males of the invasive round goby Neogobius melanostomus in samples from two activity selective passive fishing gears compared with one activity non-selective fishing gear in three Dutch lakes is related to higher male locomotory activity and is a sex-dependent trait. This difference in activity reflects the different ecology of male and female N. melanostomus.

Real-time distribution of pelagic fish: combining hydroacoustics, GIS and spatial modelling at a fine spatial scale.

Understanding spatial distribution of organisms in heterogeneous environment remains one of the chief issues in ecology. Spatial organization of freshwater fish was investigated predominantly on large-scale, neglecting important local conditions and ecological processes. However, small-scale processes are of an essential importance for individual habitat preferences and hence structuring trophic cascades and species coexistence. In this work, we analysed the real-time spatial distribution of pelagic freshwater fish in the Rímov Reservoir (Czechia) observed by hydroacoustics in relation to important environmental predictors during 48 hours at 3-h interval. Effect of diurnal cycle was revealed of highest significance in all spatial models with inverse trends between fish distribution and predictors in day and night in general. Our findings highlighted daytime pelagic fish distribution as highly aggregated, with general fish preferences for central, deep and highly illuminated areas, whereas nighttime distribution was more disperse and fish preferred nearshore steep sloped areas with higher depth. This turnover suggests prominent movements of significant part of fish assemblage between pelagic and nearshore areas on a diel basis. In conclusion, hydroacoustics, GIS and spatial modelling proved as valuable tool for predicting local fish distribution and elucidate its drivers, which has far reaching implications for understanding freshwater ecosystem functioning.

Early life-history predator-prey reversal in two cyprinid fishes.

Predator-prey relationships are often perceived simply as a situation in which a predator enhances its own fitness while reducing the fitness of its prey. However, this relationship may become reversed when the prey feeds on the juvenile predator stages. We investigated this phenomenon in a model asp (Leuciscus aspius; predator)-bleak (Alburnus alburnus; prey) relationship. The adhesive asp eggs are available for bleak predation after a spawning event for only tens of seconds before they adhere to the stones, where bleak do not forage. Gut content analysis demonstrated that eggs were utilized in high quantities, especially in the spawning peak of the asp reproductive season. Furthermore, using underwater video, we recorded the bleak feeding efficiency on naturally drifting asp eggs as the percentage of eggs eaten. Within the 40 cm egg trajectory captured by our cameras, total egg mortality was 21.2 ± 2.2% on average. The highest survival chances occurred among eggs drifting in aggregations, since the short drifting distance together with their aggregated distribution satiated bleak and part of the egg aggregation could attach to the spawning ground. This study emphasizes the potential efficiency of predator egg utilization by prey, which may have further consequences in predator-prey dynamics.

Seasonal and daily protandry in a cyprinid fish.

In polygynandrous mating systems, in which females limit reproductive success, males can increase their success by investing in courtship. Earlier arrival at the spawning ground compared to when females arrive may increase their opportunities in competitive mating systems. In this study, we used passive telemetry to test whether a male minnow known as the asp, Leuciscus aspius, times its arrival at spawning grounds relative to the arrival of females. Males arrived in a model stream approximately five days earlier than females on average and left four to five days later than females over two years. Both sexes performed a daily migration between a staging ground (standing water, low energy costs) and the fluvial spawning ground (high energy costs). Fish abundance peaked twice a day, with a major peak at sunset and a minor peak at sunrise and with the evening peak abundance for males occurring 1 hour 40 minutes earlier than that of females. The number of females on the spawning ground never exceeded the number of males. While the degree of protandry is hypothesized to be influenced by the operational sex ratio (ranging from 0.5 to 1 in our study), our data did not support this theory.

A novel upward-looking hydroacoustic method for improving pelagic fish surveys.

For ethical reasons and animal welfare, it is becoming increasingly more important to carry out ecological surveys with a non-invasive approach. Information about fish distribution and abundance in the upper water column is often fundamental. However, this information is extremely hard to obtain using classical hydroacoustic methods. We developed a rigid frame system for pushing upward looking transducers of the scientific echo sounder (38 and 120 kHz) in front of the research vessel. The efficiency of the new approach for monitoring juvenile fish at night was investigated by comparing the results with a quantitative fry trawl in the Rímov Reservoir in the Czech Republic. The experimental setup enabled comparisons for the 0-3 m and 3-6 m depth layers, which are utilized by almost all juvenile fish in summer. No statistically significant differences in the estimated abundance of juveniles were found between the two sampling methods. The comparison of abundance estimates gathered by the two frequencies were also not significantly different. The predicted mean lengths from acoustic sampling and the trawl catches differed by less than 10 mm in all comparisons. Results suggest that mobile hydroacoustic upward-looking systems can fill the methodological gap in non-invasive surveying of surface fishes.

Response of fish communities to multiple pressures: Development of a total anthropogenic pressure intensity index.

Lakes in Europe are subject to multiple anthropogenic pressures, such as eutrophication, habitat degradation and introduction of alien species, which are frequently inter-related. Therefore, effective assessment methods addressing multiple pressures are needed. In addition, these systems have to be harmonised (i.e. intercalibrated) to achieve common management objectives across Europe. Assessments of fish communities inform environmental policies on ecological conditions integrating the impacts of multiple pressures. However, the challenge is to ensure consistency in ecological assessments through time, across ecosystem types and across jurisdictional boundaries. To overcome the serious comparability issues between national assessment systems in Europe, a total anthropogenic pressure intensity (TAPI) index was developed as a weighted combination of the most common pressures in European lakes that is validated against 10 national fish-based water quality assessment systems using data from 556 lakes. Multi-pressure indices showed significantly higher correlations with fish indices than single-pressure indices. The best-performing index combines eutrophication, hydromorphological alterations and human use intensity of lakes. For specific lake types also biological pressures may constitute an important additional pressure. The best-performing index showed a strong correlation with eight national fish-based assessment systems. This index can be used in lake management for assessing total anthropogenic pressure on lake ecosystems and creates a benchmark for comparison of fish assessments independent of fish community composition, size structure and fishing-gear. We argue that fish-based multiple-pressure assessment tools should be seen as complementary to single-pressure tools offering the major advantage of integrating direct and indirect effects of multiple pressures over large scales of space and time.

Latitudinal variation in sexual dimorphism in life-history traits of a freshwater fish.

Sexual dimorphism is common across the animal kingdom, but the contribution of environmental factors shaping differences between the sexes remains controversial. In ectotherms, life-history traits are known to correlate with latitude, but sex-specific responses are not well understood. We analyzed life-history trait variation between the sexes of European perch (Perca fluviatilis L.), a common freshwater fish displaying larger female size, by employing a wide latitudinal gradient. We expected to find sex-dependent latitudinal variation in life-history variables: length at age, length increment, and size at maturity, with females showing consistently higher values than males at all latitudes. We further anticipated that this gender difference would progressively decrease with the increasingly harsh environmental conditions toward higher latitude. We hypothesized that growth and length increment would decrease and size/age at maturity would increase at higher latitudes. Our results confirmed female-biased sexual size dimorphism at all latitudes and the magnitude of sexual dimorphism diminished with increase in latitude. Growth of both sexes decreased with increase in latitude, and the female latitudinal clines were steeper than those of males. Hence, we challenge two predominant ecological rules (Rensch's and Bergmann's rules) that describe common large-scale patterns of body size variation. Our data demonstrate that these two rules are not universally applicable in ectotherms or female-biased species. Our study highlights the importance of sex-specific differences in life-history traits along a latitudinal gradient, with evident implications for a wide range of studies from individual to ecosystems level.

Who Is Who: An Anomalous Predator-Prey Role Exchange between Cyprinids and Perch.

Piscivory in cyprinids (Cyprinidae) is extremely rare. Specifically, common bream (Abramis brama) and common carp (Cyprinus carpio) are zooplanktivorous fish in deep lentic waters. Nevertheless, we observed predation by these two cyprinids under natural conditions in the Vír Reservoir, Czech Republic. We conducted diet analysis for cyprinids caught by trawling and gillnets and the large amount of young-of-the-year (YOY) perch (Perca fluviatilis), with sizes of 37-52 mm standard length, were found in their digestive tracts. In 2010, a large amount of YOY perch caused a significant decrease in Daphnia spp. size and abundance in the reservoir. Hence, a food deficit was induced for the cyprinids, apparent also from the poor nutritional condition of common bream which was much worse than the condition of those in similar reservoirs. Common carp and common bream shifted to forced piscivory, and they utilized the YOY perch as an alternative food source. In contrast, smaller species, such as roach (Rutilus rutilus) and bleak (Alburnus alburnus), widely utilized planktonic cyanobacteria. In the following year, YOY perch occurred in significantly lower numbers and conversely, Daphnia spp. size and abundance were significantly higher. The forced piscivory was not observed. Our results indicate a switch to forced piscivory by cyprinids, which was caused by a shortage of their natural food source. Moreover, this phenomenon presents an effective mechanism for reduction in the numbers of YOY perch, ensuring the stability of the ecosystem.

Biomass and abundance biases in European standard gillnet sampling.

The European Standard EN 14757 recommends gillnet mesh sizes that range from 5 to 55mm (knot-to-knot) for the standard monitoring of fish assemblages and suggests adding gillnets with larger mesh sizes if necessary. Our research showed that the recommended range of mesh sizes did not provide a representative picture of fish sizes for larger species that commonly occur in continental Europe. We developed a novel, large mesh gillnet which consists of mesh sizes 70, 90, 110 and 135mm (knot to knot, 10m panels) and assessed its added value for monitoring purposes. From selectivity curves obtained by sampling with single mesh size gillnets (11 mesh sizes 6 - 55mm) and large mesh gillnets, we identified the threshold length of bream (Abramis brama) above which this widespread large species was underestimated by European standard gillnet catches. We tested the European Standard gillnet by comparing its size composition with that obtained during concurrent pelagic trawling and purse seining in a cyprinid-dominated reservoir and found that the European Standard underestimated fish larger than 292mm by 26 times. The inclusion of large mesh gillnets in the sampling design removed this underestimation. We analysed the length-age relationship of bream in the Rímov Reservoir, and concluded that catches of bream larger than 292mm and older than five years were seriously underrepresented in European Standard gillnet catches. The Rímov Reservoir is a typical cyprinid-dominated water body where the biomass of bream > 292mm formed 70% of the pelagic trawl and purse seine catch. The species-specific relationships between the large mesh gillnet catch and European Standard catch suggested that the presence of carp (Cyprinus carpio), European catfish (Silurus glanis), tench (Tinca tinca) or bream warrants the use of both gillnet types. We suggest extending the gillnet series in the European Standard to avoid misinterpretation of fish community biomass estimates.