Cottus Gobio Research Articles

A comparison of swimming capacity and energy use in seven European freshwater fish species

Abstract – Migrating fish species with different swimming capacities and energy use show different capacities for passing obstacles between habitats, such as culverts and fish ladders. Here, we present an integrated study on swimming capacity and energetic use in seven European freshwater fish species with different ranges of migration (brown troutSalmo truttaL., European perchPerca fluviatilisL., roachRutilus rutilusL., common carpCyprinus carpioL., gudgeonGobio gobioL., bullheadCottus gobioL. and stone loachBarbatula barbatulaL.). Critical (Ucrit), optimal (Uopt) and maximum (Umax) swimming speed and oxygen consumption (MO2) were analysed and showed values correlated to migration capacity with highest swimming capacities in trout and roach and lowest in stone loach and bullhead. The resulting data can be used to make estimates of maximum passable water speeds in culverts. In conclusion, long‐distance migrators show higher swimming capacities and can potentially clear obstacles easier than short distance migrators with lower swimming capacities. Even small obstacles (<25 cm) could be a barrier for genetic exchange between populations in short‐distance migrators.

MODIFICATION OF HOSTS' BEHAVIOR BY A PARASITE: FIELD EVIDENCE FOR ADAPTIVE MANIPULATION

Parasites relying on trophic transmission to complete their life cycles often induce modifications of their host's behavior in ways that may increase their susceptibility to predation by final hosts. These modifications have often been interpreted as parasite adaptations, but very few studies have demonstrated that host manipulation has fitness benefits for the parasite. The aim of the present study was to address the adaptive significance of parasite manipulation by coupling observations of behavioral manipulation to estimates of trophic transmission to the definitive host in the natural environment. We show that the acanthocephalan parasite Pomphorhynchus laevis manipulates the drifting behavior of one of its intermediate hosts, the amphipod Gammarus pulex, but not of a sympatric host, the introduced amphipod Gammarus roeseli. We found a 26.3-28.3 times higher proportion of infected G. pulex in the stomach content of one of the definitive hosts of P. laevis, the bullhead Cottus gobio, than in the benthos. No such trend was observed for G. roeseli. The bell-shaped curve of mean parasite abundance (MPA) relative to host size observed in G. pulex also supported an increased predation mortality of P. laevis-infected individuals compared to uninfected amphipods. Again, no such pattern was observed in G. roeseli. Furthermore, our results indicate that the modifications induced by P. laevis are specific to the definitive host and do not increase the risk of predation by inappropriate hosts, here the adult edible frog Rana esculenta. Overall, our study is original in that it establishes, under field conditions, a direct link between parasitic manipulation and increased transmission to the definitive host, and more importantly, identifies the specificity of the manipulation both in the intermediate host species and toward the definitive host.

Differential influence of two acanthocephalan parasites on the antipredator behaviour of their common intermediate host

Fish acanthocephalans can modify the antipredator behaviour of their intermediate hosts in response to cues from fish predators. However, it is still unclear whether such behavioural changes are adaptive, or are just the consequence of infection. We addressed this question through studying two acanthocephalans, Pomphorhynchus laevis and Polymorphus minutus, and their intermediate host, the amphipod Gammarus pulex. Pomphorhynchus laevis completes its cycle in a freshwater fish, whereas P. minutus exploits waterbirds as final hosts. We first assessed vulnerability of infected and uninfected gammarids to predation by bullheads, Cottus gobio. Pomphorhynchus laevis-infected gammarids were more susceptible to predation than uninfected ones when a refuge was available, whereas no selective predation on P. minutus-infected individuals was recorded, independently of refuge availability. We then quantified refuge use when a bullhead was present in an enclosure or when the enclosure was empty. Individuals of each group significantly increased refuge use in the presence of a bullhead. However, a larger proportion of P. laevis-infected gammarids remained out of the refuge in the presence of a predator, compared with uninfected controls, whereas no such effect was observed in P. minutus-infected ones. Finally, we assessed reaction to bullhead olfactory cues, using a Y-maze apparatus. Pomphorhynchus laevis-infected gammarids spent significantly more time in the predator-scented arm, whereas the reverse was observed in uninfected ones. Polymorphus minutus-infected individuals, however, did not differ from uninfected controls. We discuss our results in relation with the adaptiveness of host manipulation by parasites.

Genetic analysis of potential postglacial watershed crossings in Central Europe by the bullhead (Cottus gobioL.)

Natural colonizations across watersheds have been frequently proposed to explain the present distributions of many freshwater fish species. However, detailed studies of such potential watershed crossings are still missing. Here, we investigated potential postglacial watershed crossings of the widely distributed European bullhead (Cottus gobio L.) in two different areas along the Rhine-Rhône watershed using detailed genetic analysis. The main advantage of studying bullheads vs. other freshwater fish species is that their distribution has been lightly influenced by human activities and as such, interpretations of colonization history are not confounded by artificial transplantations. The genetic analyses of eight microsatellite loci revealed strong genetic similarities between populations of both sides of the Rhine-Rhône watershed in the Lake Geneva area, giving strong evidence for a natural watershed crossing of bullheads from the upper Rhine drainage into the Rhône drainage in the Lake Geneva area likely facilitated by the retreat of the glaciers after the last glacial maximum some 20,000 years ago. Populations from the Lake Geneva basin were genetically more similar to populations from across the watershed in the upper Rhine drainage than to populations further downstream in the lower Rhône. In contrast, populations from Belfort, an area, which was not covered by ice during the last glacial maximum, showed strong genetic differentiation between populations of the upper Rhine and Rhône drainages. Based on our results on the bullhead, we propose that glacial retreat may have eased the dispersal of numerous European freshwater fish species across several geological boundaries.

MICROSATELLITE POLYMORPHISM WITHIN TWO LAKE BAIKAL OILFISH SPECIES (COMEPHORUS LACEPEDE, 1801)

Intraspecific genetic polymorphism of a Baikal Lake endemic, big Baikal oilfish (Comephorus baicalensis Pallas, 1776), was evaluated based on microsatellite analysis. The obtained results have compared to the results received earlier for a little Baikal oilfish (C. dybowski Korotneff, 1905). Six microsatellite loci designed for the European sculpin, Cottus gobio, were used. Big Baikal oilfish samples were tested from the three Baikal trenches (southern, middle, northern). Average values of FsubST/sub and RsubST/subindexes between the samples did not exceed 0,02 that point to weak intraspecific genetic differentiation. Big Baikal oilfish had the smaller allele variety, the greater deficit of heterozygotes and presence of low-molecular alleles in comparison with little Baikal oilfish. The values of genetic differentiation were equaled FsubST/sub = 0,138 and RsubST/sub = 0,244 for all loci between cumulative samples of little and big Baikal oilfish. Analysis of microsatellite polymorphism has showed that both a big and little Baikal oilfishes are represented by single populations. The factor promoting panmixia within the Baikal oilfish species is apparently a passive transfer of juvenile and adult fishes by water currents.

Introduced bullheads Cottus gobio and infection with plerocercoids of Schistocephalus cotti in the Utsjoki, an Arctic river in Finland

Bullheads Cottus gobio, first found in the River Utsjoki, an Arctic river in Finland, in 1979, were sampled at six sites in August 1991 (two of these sites were also sampled previously in June and July 1991). Mean total length (LT) of C. gobio was 45 mm (n= 1080 fish). Overall, there were slightly more females than males, with males larger than females (mean LT males 52·1 and females 47·6 mm). Prevalence of infection with plerocercoids of Schistocephalus cotti varied from 20·0 to 43·0% over the six sites in August. Juvenile C. gobio had a prevalence of infection of 3·5% whereas in mature fish prevalence was consistently higher in females than in males (38·8 v. 20·1%). There was one plerocercoid per infected juvenile C. gobio. In mature fish the mean number varied between 1·3 and 1·8: 49% had one worm per fish, 38% two or three worms and 13% had four to eight plerocercoids. In juvenile C. gobio, S. cotti were randomly distributed (overdispersion index 1·10), whereas occurrence was slightly aggregated in the LT groups 40–59 and ≥60 mm (overdispersion indices both 1·90). Mean mass of infected fish was higher in all LT groups compared to uninfected C. gobio. After removal of plerocercoids, mean mass of fish tissue remaining was lower compared with uninfected fish for the two LT groups 40–59 and ≥ 60 mm, but higher in C. gobio <40 mm LT. Prevalence and mean intensity of infection over the 3 months did not differ either for the two sites individually or the pooled data. Mean mass of both uninfected and infected fish increased over this period, and were both highest in August as was the mean worm burden. There was, however, no difference in Fulton’s condition factor at any time. Mean mass of plerocercoids in C. gobio of ≥40 mm LT increased significantly comparing June and July with August. Comparison of backcalculated LT of infected and non‐infected bullheads of different age‐groups did not show any significant differences in growth at any age. In each parasite infrapopulation as the number of worms increased so their mean mass decreased significantly but at the same time the total mass of worms in the fish host remained more or less constant in relation to length. Aggregation of parasites in the fish population was low, as the majority were infected by one S. cotti plerocercoid. Gonad development was suppressed in infected C. gobio.

Carry-over effects in brown trout (Salmo trutta): hypoxia on embryos impairs predator avoidance by alevins in experimental channels

Laboratory experiments on brown trout (Salmo trutta) embryos suggest that sublethal stress in the gravel nest such as hypoxia may alter the behaviour and survival of fish during the early juvenile period. Eggs and embryos were incubated at constant temperature (8.2 °C ± 0.6 standard deviation) under nonlethal dissolved oxygen (DO) concentrations (3.0 mg·L–1; 26% air saturation level) and normoxia (10.3 mg·L–1DO; 90% air saturation level). The average survival from fertilization to end of embryonic development was 70% and 85% for hypoxic and normoxic groups, respectively. Hypoxic embryos grew slowly compared with their normoxic counterparts, but similar body sizes were observed when yolk-sac absorption was completed. Fish incubated as hypoxic embryos delayed their emergence from the gravel in experimental channels. In presence of freshwater sculpin (Cottus gobio), their swimming activity was reduced by 20%, on average, and predation was enhanced by 14% compared with normoxic groups. Results support the view that subtle events early in a fish's ontogeny can have carry-over effects on later periods of its life cycle, and this phenomenon may be a significant source of variation in salmonid fitness.

Intrabasin variations in age and growth of bullhead: the effects of temperature

The variability of growth variables and age structure in bullhead Cottus gobio was studied in a single river catchment by comparing populations inhabiting river reaches with different thermal regimes. Age and growth variables were determined by examining 540 sagittal otoliths from specimens ranging between 19 and 146 mm total length. Growth of young bullhead (0+ and 1+ years) was higher in the warmest sites, especially in relation to water temperature during the growing season (spring and summer). Growth of older bullhead (2+ to 5+ years) was negatively related to mean water temperature of the site. The growth performance of bullhead over its whole life cycle was negatively correlated with water temperature, which could be partly explained by a higher energy allocation to reproduction in the warmest sites. Life span was longer in the colder sites (7 years) than in the warmer ones (4 or 5 years), and bullhead matured earlier in the warmer sites (age 1+ or 2+ years) compared to the colder ones (age 2+ or 3+ years). These results, combined with previous data on reproductive traits, suggest the possible effects that global warming could have on bullhead population dynamics.

Evaluation of passive integrated transponder tags for marking the bullhead (Cottus gobio), a small benthic freshwater fish: effects on survival, growth and swimming capacity

Abstract – This study examined potential adverse effects of surgically implanted passive integrated transponder (PIT) tags (12 × 2.1 mm) on bullhead (Cottus gobio L.) of three different length‐classes (I: 50–64 mm, II: 65–79 mm, III: 80–94 mm). During a 7‐week laboratory experiment, the rate of PIT tag loss, incision closure time, survival, growth and swimming capacity were tested. The PIT tag weight to fish body weight ratio varied between 1.04% and 4.85%. The mean incision closure time differed significantly among length‐classes and varied between 2.8 (I) and 4.3 (III) weeks. Nevertheless, PIT tag retention did not differ among length‐classes and was ≥90%. The survival of untagged, sham‐tagged and PIT‐tagged bullheads was ≥90% and did not differ within or among length‐classes. Finally, within each length‐class, there was no difference in growth and swimming capacity among treatments. Hence, these results suggest the applicability of PIT tags for individually tagging bullheads ≥50 mm.

Performance of a pool-and-weir fish pass for small bottom-dwelling freshwater fish species in a regulated lowland river

Abstract Although small benthic freshwater fish species are an important biological component of fish assemblages and free instream movement is indispensable for their survival, they are often neglected in fish pass performance studies. In this study, a capture-mark-recapture approach was used to assess whether small bottom-dwelling species, including gudgeon (Gobio gobio), stone loach (Barbatula barbatula), spined loach (Cobitis taenia) and bullhead (Cottus gobio), were able to cross a pool-and-weir fish pass in a regulated lowland river. Some tagged individuals of stone loach (18%), gudgeon (7%) and spined loach (2%) managed to successfully ascend the fish pass under study, despite the fact that water velocity levels in the different overflows of the facility (between 0.55-1.22 m/s) exceeded the critical swimming speed of all three species. Although this suggests that a pool-and-weir fish pass is a able to facilitate upstream movement of some small benthic species in a regulated river, more detailed research incorporating advanced tagging and retrieving techniques is necessary.

Fish habitat modelling as a tool for river management

This paper presents an integrated modelling approach to simulate and assess ecological effects of physical habitat changes in rivers. An ecohydraulic simulation tool was created by combining a 1D hydraulic model based on HEC-RAS software and the fish habitat module of CASiMiR, a fuzzy logic-based ecohydraulic modelling system. This tool was applied on a river stretch commonly occurring in Belgium and elsewhere in Europe. In particular the effect of weir removal on habitat suitability for bullhead ( Cottus gobio L.) was simulated. Physical conditions of the studied stretch after weir removal were simulated with a hydraulic model. CASiMiR linked these conditions to ecological expert knowledge to calculate habitat suitability for three life stages of bullhead at four different flow rates based on fuzzy logic. Results indicated that after weir removal, habitat suitability increased significantly for all life stages and all flow rates. The presented approach is promising regarding fish community assessment and ecological river engineering.

Increased susceptibility to predation and altered anti-predator behaviour in an acanthocephalan-infected amphipod

According to the ‘parasitic manipulation hypothesis’, phenotypic changes induced by parasites in their intermediate hosts are effective means of increasing trophic transmission to final hosts. One obvious prediction, although seldom tested, is that increased vulnerability of infected prey to an appropriate predator should be achieved by the parasite altering the anti-predator behaviour of its intermediate host. In this study, we tested this prediction using the fish acanthocephalan Pomphorhynchus tereticollis and the freshwater amphipod Gammarus pulex. Firstly, we estimated the relative vulnerability of infected and uninfected gammarids to predation by the bullhead Cottus gobio in the field. Second, we investigated under experimental conditions how two common anti-predator behaviours of aquatic invertebrates, refuge use and short-distance reaction to predator chemical cues, were affected by infection status. We found that the prevalence of infection in the field was 10 times higher among gammarids collected from the stomach contents of bullheads compared with free-ranging individuals collected in the same river. In a microcosm uninfected gammarids, but not infected ones, increased the use of refuge in the presence of a bullhead. Finally, a behavioural experiment using an Y-maze olfactometer showed opposite reactions to predator odour. Whereas uninfected gammarids were significantly repulsed by the chemical cues originating from bullheads, infected ones were significantly attracted to the odour of the predator. Taken together, our results suggest that the alteration of anti-predator behaviour in infected G. pulex might enhance predation by bullheads in the field. Reversing anti-predator behaviour might thus be an efficient device by which parasites with complex life-cycles increase their trophic transmission to final hosts. Further studies should pay more attention to both the increased vulnerability of infected prey to an appropriate predator in the field and the influence of parasitic infection on the anti-predator behaviour of intermediate hosts.

Food-web responses to restoration of channel heterogeneity in boreal streams

We assessed the biomass and stable-isotope composition (δ13C and δ15N) of consumers (aquatic insects and fish (Cottus gobio)) and potential food sources (detritus, biofilm, seston, algae, and mosses) in channelized and restored streams in Sweden, assessing the hypotheses that restoration enhances detritus storage and detritus-based secondary production. Restored sites stored more detritus than channelized sites, with differences (+5.4% on average) prominent in margins, i.e., the channel area within 1.4 m from the banks. The biomass of other resources was similar between restored and channelized sites. Most common aquatic insects, including several putative detritivores, showed δ13C values indicating reliance on aquatic (probably algal) carbon sources. The insectivorous fish Cottus gobio, on the contrary, appeared to be more dependent on terrestrial (detrital) carbon sources. The biomass and mean δ13C values of the consumers were similar between restored and channelized sites, suggesting that restoration did not increase net secondary production or the fraction of secondary production based on detritus. We concluded that the increase in detritus storage at restored sites was either insufficient to enhance detritus-based production or the consumers were not limited by the availability of detritus.

Schistocephalus cotti n. sp. (Cestoda: Pseudophyllidea) plerocercoids from bullheads Cottus gobio L. in an Arctic river in Finland, with a key to the plerocercoids of the Palaearctic species of the genus

We compared plerocercoids of Schistocephalus Creplin, 1829 from Cottus gobio (n = 57) and Gasterosteus aculeatus f. semiarmatus (n = 45) from the River Utsjoki, Finland, taken only from single worm infections. Segment numbers in the two populations were distinct (G. aculeatus range 55-107, average 74 (SE 1.66), median 73; C. gobio range 122-189, average 146 (SE 1.78); median 144). The mean difference between populations, 71.47, t = 28.76 with 100 degrees of freedom, two-tailed p value <0.001, was considered extremely significant. Amplification of microsatellite loci that were originally designed for Schistocephalus from G. aculeatus was positive for all larvae from G. aculeatus (n = 20), whereas in no plerocercoids from C. gobio (n = 20) were any of the six microsatellites amplified, indicating that plerocercoids from G. aculeatus and C. gobio were two distinct genetic populations of Schistocephalus. The material from C. gobio is described as S. cotti n. sp. Plerocercoids of the Palaearctic species of Schistocephalus are identified as follows: S. nemachili Dubinina, 1959 with 228-235 or more segments, specific to Barbatula spp. (Balitoridae); S. pungitii Dubinina, 1959 with 62-92 (usually 70-80) segments, specific to Pungitius pungitius; S. solidus (Müller, 1776) in two forms, one in G. aculeatus f. leiurus and f. semiarmatus, with 48-100 (usually 65-75) segments, and the other in G. aculeatus f. trachurus, with 99-138 (usually 112-122) segments; and S. cotti n. sp. with 103-189 (usually 130-159) segments, probably specific to cottids. Nearctic Schistocephalus were not considered owing to the uncertain status of some North American records. Some other species of Schistocephalus of highly doubtful status were briefly noted. Cross-infection experiments and molecular studies are recommended to further elucidate the interrelationships between the various species of Schistocephalus.

Mechanisms of density dependence in stream fish: exploitation competition for food reduces growth of adult European bullheads (Cottus gobio)

In field experiments using cage enclosures, exploitation competition for invertebrate prey reduced individual growth of adult European bullheads (Cottus gobio) despite a dietary shift that maintained total prey biomass consumption. Growth of bullheads was negatively density dependent over a range of densities found in the field and total biomass production per enclosure was unrelated to stocking density. Individuals grew faster when invertebrate densities were experimentally elevated above ambient levels, indicating that bullheads were food limited. Parallel dietary shifts in response to manipulation of both bullhead density and resource availability suggested that exploitation competition for preferred Gammarus pulex prey was the primary cause of density-dependent growth. Switching to smaller prey species did not prevent growth of bullheads declining with increasing per capita food competition, indicating that the alternative prey were less profitable. Evidence for interference competition was limited. These results caution against the assumption that demographic rates of stream fish are density independent after the juvenile phase and demonstrate instead that density-dependent growth may potentially regulate populations at the adult stage via effects on fecundity and survival.

Effects of temperature on persistence times of native and invasive gammarid species in the stomachs of Cottus gobio

Gastric lavage was used to investigate the effects of temperature on persistence time of two amphipod species, one native Gammarus pulex and one invasive Gammarus roeseli, in the stomachs of bullhead Cottus gobio. Persistence time was strongly influenced by temperature and prey type, such that G. pulex species degraded faster than G. roeseli.

Effects of temperature on persistence times of native and invasive gammarid species in the stomachs of Cottus gobio.

Effects of temperature on persistence times of native and invasive gammarid species in the stomachs of Cottus gobio.

Fish pass effectiveness for bullhead (Cottus gobio), perch (Perca fluviatilis) and roach (Rutilus rutilus) in a regulated lowland river

Abstract – The effectiveness of a pool‐and‐weir fish pass in the regulated river Laarse Beek (Belgium) was assessed for bullhead (Cottus gobio), perch (Perca fluviatilis) and roach (Rutilus rutilus) from August 2002 to April 2003. Fish were caught by electrofishing downstream, within and upstream of the fish pass. They were marked with visible implant elastomer tags and released downstream of the fish pass. Our results suggest that the fish pass is ineffective for the bullhead because none of the 1270 tagged bullheads were recaptured in the upstream study area. Passage failure most likely resulted from excessive water velocities in the fish pass. On the contrary, 8% of the tagged perch and 29% of the tagged roach were recaptured in the upstream study area. Several management strategies are proposed to allow free upstream passage of the endangered bullhead at the fish pass under study.

Influence of female age, body size and environmental conditions on annual egg production of the bullhead

A population of bullhead Cottus gobio inhabiting a mountainous river network, was examined for the influence of maternal condition (total length, LT, and age class in years) and environmental factors (water temperature and fish density) on fecundity, egg size and net annual reproductive effort (ER, defined as the ratio of total egg production mass to body mass) within several stocks. Regardless of location within the drainage network, bullhead reproduction was synchronous, with only a single batch and fecundity ranging from 69 to 896 eggs per female. As with LT fecundity increased with age, although the difference between age classes 5 and 6 years was smaller than between other age classes. Egg diameter also increased with age from age classes 2 to 4 years, but then tended to decrease for age classes 5–6 years. Multiple stepwise regression was used to examine the effect of female LT, water temperature and fish densities on bullhead egg production for each of the four age classes (2, 3, 4 and 5–6 years) separately. When statistically significant, water temperature always had a positive effect and fish densities a negative effect, on fecundity and egg size. The LT always had a positive effect on fecundity for all the age classes. In contrast, LT had a positive effect on egg size and ER only for female reaching maturity (2 years), whereas intermediate aged females (3–4 years) were insensitive and the oldest fish showed a negative response.

A genetic map ofCottus gobio(Pisces, Teleostei) based on microsatellites can be linked to the physical map ofTetraodon nigroviridis

To initiate QTL studies in the nonmodel fish Cottus gobio we constructed a genetic map based on 171 microsatellite markers. The mapping panel consisted of F1 intercrosses between two divergent Cottus lineages from the River Rhine System. Basic local alignment search tool (BLAST) searches with the flanking sequences of the microsatellite markers yielded a significant (e < 10(-5)) hit with the Tetraodon nigroviridis genomic sequence for 45% of the Cottus loci. Remarkably, most of these hits were due to short highly conserved noncoding stretches. These have an average length of 40 bp and are on average 92% conserved. Comparison of the map locations between the two genomes revealed extensive conserved synteny, suggesting that the Tetraodon genomic sequence will serve as an excellent genomic reference for at least the Acanthopterygii, which include evolutionarily interesting fish groups such as guppies (Poecilia), cichlids (Tilapia) or Xiphophorus (Platy). The apparent high density of short conserved noncoding stretches in these fish genomes will highly facilitate the identification of genes that have been identified in QTL mapping strategies of evolutionary relevant traits.