Diverse migratory histories of Japanese Trachidermus and Cottus species (Cottidae) as inferred from otolith microchemistry

The migratory histories of Japanese freshwater sculpins, one Trachidermus and four Cottus species, were studied by examining strontium (Sr) and calcium (Ca) concentrations in their otoliths using wavelength dispersive X‐ray spectrometry on an electron microprobe. The Sr : Ca ratios in the otoliths changed with salinity of the habitat. The otoliths of Cottus nozawae showed consistently low Sr : Ca ratios, with an average of 3·37 × 10^?3 from the core to the edge, suggesting a freshwater resident life cycle. In contrast, the otolith Sr : Ca ratios for Trachidermus fasciatus and Cottus kazika changed along the life history transects possibly in accordance with their migration patterns from sea to fresh water. The ratios of T. fasciatus and C. kazika averaged 5·4 × 10^?3 and 5·3 × 10^?3 respectively, in the otolith region from the core to the points 450–890 μm, and changed to the lower levels, averaging 2·0 × 10^?3 and 2·7 × 10^?3, in the outer otolith region. These data suggest that both the species have a catadromous life cycle. The otoliths of Cottus hangiongensis had low Sr : Ca ratios in the two regions from the core to the points 15–30 μm and the points 415–582 μm to the edge, averaging 2·0 × 10^?3 and 1·9 × 10^?3, with significantly higher ratios in the narrow area between these regions, averaging 4·6 × 10^?3. Similar ontogenetic changes in otolith Sr : Ca ratios were found in the otoliths of Cottus amblystomopsis, suggesting their amphidromous life cycle. These findings suggest that otolith Sr : Ca ratios reflect individual life histories and that Japanese Trachidermus and Cottus species have diverse migratory histories.     

The trophic niche of sculpins Cottus spp. in forage fish assemblages of boreal lakes

We compared the trophic niches of freshwater sculpins Cottus spp. with those of other co-habiting forage fishes in two groups of boreal lakes with distinct habitats and fish communities. Near North Lakes (45° 00′ to 47° 30′ N) were deeper, stratified and contained lake trout Salvelinus namaycush as the apex piscivore, whereas Far North Lakes (51° 10′ to 52° 20′ N) were shallower, did not stratify and contained pike Esox lucius and walleye Sander vitreus as the apex piscivores. Trophic niches of sculpins and other forage fishes were compared based on niche metrics calculated from muscle stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope ratios. In Near North Lakes, sculpins were found almost exclusively in deep, offshore waters and their niche positions reflected a greater reliance on pelagic production (lower δ¹³C) and a higher trophic elevation (higher δ¹⁵N) compared with most other forage fishes. Furthermore, sculpins in Near North Lakes tended to have larger trophic niches (occupied greater area in δ¹³C– δ¹⁵N space), particularly in the food chain (δ¹⁵N) dimension, than other cohabiting forage fishes. In contrast, sculpins in Far North Lakes were commonly found in the nearshore and had trophic niche positions and sizes that were similar to those of the other cohabiting forage fishes. This study illustrates the flexibility in the realised trophic niches of sculpins in relation to habitat availability and fish community composition in boreal lakes.     

Comparative ecology of prickly sculpin,Cottus asper,and coastrange sculpin,C. aleuticus,in the Eel River,California

Synopsis We documented species' distributions, size structure of populations, abundance in mainstem and tributary streams, habitat use, and diets of prickly sculpin, Cottus asper, and coastrange sculpin, C. aleuticus, in the Eel River drainage of California, to determine the processes allowing coexistence of these very similar fishes. We observed prickly sculpins at 43 sites and coastrange sculpins at 34. The species co-occurred at 26 sites. Young-of-year coastrange sculpins were only observed within 42 km of the ocean, but young-of-year prickly sculpins were present throughout the species range. Mean, maximum, and minimum lengths of coastrange sculpins were positively correlated with distance from the ocean but no significant relationships were found for prickly sculpins. Absolute abundance of both species was highest in mainstem habitat (prickly sculpins = 0.6 sculpins m^–2 and coastrange sculpins = 0.4 sculpins m^–2) . Tributary densities of both species tended to be less than 0.1 sculpins m^–2. The species inhabited very similar habitats and had very similar diets. Coastrange sculpin populations in upstream areas were maintained by immigration from downstream areas in contrast with prickly sculpin populations that produced young-of-year fish throughout their range. Densities were probably not high enough for interspecific interactions to be important. The factors limiting the upstream distribution of the species may include high water temperatures, stability of the stream bed, and behavior of the fish. In the past, the range of sculpins within the Eel River drainage probably fluctuated with changing physical conditions. Recent introductions of exotic species that compete with and prey upon sculpins, and ongoing human activities in the drainage could result in major reductions in the distribution and abundance of one or both species.     

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 (L_T) of C. gobio was 45 mm (n= 1080 fish). Overall, there were slightly more females than males, with males larger than females (mean L_T 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 L_T groups 40–59 and ≥60 mm (overdispersion indices both 1·90). Mean mass of infected fish was higher in all L_T 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 L_T groups 40–59 and ≥ 60 mm, but higher in C. gobio <40 mm L_T. 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 L_T increased significantly comparing June and July with August. Comparison of backcalculated L_T 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.     

Common sculpin Cottus gobio as a natural paratenic host of Proteocephalus longicollis (Cestoda: Proteocephalidae), a parasite of salmonids, in Europe.

Common sculpins Cottus gobio L. (Pisces: Cottidae), from the Mlynsky Brook near Ceské Zleby in the Sumava National Park, southwestern Bohemia, Czech Republic, were found to harbour in their intestines juvenile cestodes Proteocephalus longicollis (Zeder, 1800), a common parasite of holarctic salmonids, with a prevalence of 60% and intensity of 1 to 11 (mean 5) parasites per fish; undoubtedly, these prey fish serve as paratenic hosts. In this locality, the definitive host of P. longicollis is the brown trout Salmo trutta m. fario L., large specimens of which apparently acquire infection of this parasite by feeding on infected sculpins. C. gobio is the first known natural paratenic host of P. longicollis in Europe.     

Shifts in abundance and growth of slimy sculpin in response to changes in the predator population in an arctic Alaskan lake

Lake trout (Salvelinus namaycush) in Toolik Lake are tightly coupled to the benthos, since they have no pelagic forage fishes. Slimy sculpins (Cottus cognatus) are a prey of lake trout and the soft sediment chironomids are an important prey for the sculpin. Our previous work showed that the median size of lake trout in Toolik Lake had decreased significantly between 1977 and 1986, and smaller lake trout are likely to be less effective as sculpin predators. Using our historic data on the slimy sculpin population from 1978, we took advantage of the recent change in the predator community to examine for subsequent changes in the sculpin community. Between 1978 and 1987, the percentage of slimy sculpin caught in the soft sediments has increased (25% to 39.5%). In 1987 there was a significant difference in the mean weight of sculpin caught on different substrates. The mean individual weight of sculpins increased from the nearshore rock area to the rock/soft-sediment interface to the soft sediments. There was no difference in mean individual weight with habitat in 1978. The mean total length at age for slimy sculpins during this time has also increased significantly. We suggest that the risk of predation while foraging in the soft sediments has declined. The increased use of the soft sediment area appears to have contributed to their increased growth, likely due to greater food abundance.     

Presence of sculpins (Cottus gobio) reduces drift and activity of Gammarus pulex (Amphipoda)

Introduction of sculpins into a stream previously devoid of these predators significantly reduced drift rate of Gammarus pulex. The drift of insect larvae was not affected. High amounts of exudates after implantation of sculpins were probably responsible for the low number of drifting G. pulex specimens. Laboratory experiments confirmed reduced locomotory activity of G. pulex when exposed to caged sculpins, an observation that excludes reduced drift activity as a result only of predation.The average size of drifting G. pulex specimens was larger during the night than during the day. This result is in accordance with the hypothesis that large individuals should, in relation to small ones, turn nocturnal because of greater predation risk during daytime. Presence of sculpins did not alter the size composition of drifting G. pulex.     

Intraspecific interactions influence size specific depth distribution in Cottus bairdi

Synopsis We tested depth selection by different sizes of mottled sculpin, Cottus bairdi, in a southern Appalachian stream. Field observations indicated that, during one hour periods, both small (<50 mm SL) and large (≥55 mm SL) individuals move within an area less than 0.50 m². Individuals of both sizes, placed in field enclosures, preferred deep microhabitat. When large fish were placed in cages with small fish, small fish initially spent more time in slope and shallow microhabitats. Average interfish distances were not correlated with their absolute size differences, suggesting C. bairdi interactions may involve both predation and competition. In streams, size-related differences in microhabitat depth may result more from intraspecific interactions than from size-specific depth preferences.     

Distribution of Cottus poecilopus Heckel and C. gobio L (Pisces) in Scandinavia

Andreasson. S. (Department of Animal Ecology, Lund University, Lund. Sweden.) Distribution of Cottus poecilopus Heckel and C. gobio L. (Pisces) in Scandinavia. Zool. Scripta 1(2): 69–78, 1972. –The distribution of the freshwater sculpins Cottus poecilopus Heckel and C. gobio L. in Scandinavia is presented. In southern Scandinavia there are rather defined limits between the species with C. poecilopus in the upper parts of the water courses and C. gobio in the lower reaches. There are few cases of sympatry. In northern Scandinavia there is a broad overlap and coexistence is common in the large rivers. The distribution pattern seems to be partly a result of an interaction between the species. In the South, the sharp limits are set by falls blocking the upstream dispersal of C. gobio thus indicating an exclusion of C. poecilopus in the lower reaches below the falls by the presence of C. gobio there. Where the species meet or overlap there is a habitat segregation.     

Distribution,production, and age structure of slimy sculpin in an Arctic lake

Synopsis Slimy sculpins (Cottus cognatus) were caught in the littoral region of Toolik Lake, an arctic lake. These sculpins grew slower and lived longer (71 mm at 8⁺ years) than more southerly populations. Sculpin distribution along the slope of the rocky littoral zone was greatest at the 3.5 m depth and coincided with the rock-mud interface. The hypothesis that this preferred area provided both increased prey and reduced predation was advanced. Yearly production estimate of littoral sculpins was 0.40 g · m^–2 · yr^–1 with a P/B ratio of 0.36. This is considerably less than estimates for more southerly populations of the same species.     

Partitioning of food resources by sculpins in Lake Michigan

Synopsis Resource partitioning was studied in two benthic Lake Michigan fishes, the deepwater sculpin,Myoxocephalus thompsoni, and the slimy sculpin,Cottus cognatus, that exhibit nearly disjunct distributions along a hypolimnetic depth gradient. Fish were collected in an area of sympatry over two 24 h periods. These sculpins exhibited food segregation—slimy sculpins ate primarilyPontoporeia affinis, deepwater sculpins ate bothP. affinis andMysis relicta — and their depth segregation was associated with the reported abundance of these prey. Different feeding behaviors may be responsible for this association, since slimy sculpins consumed intermediate size ranges ofPontoporeia and deepwater sculpins consumedPontoporeia in a pattern similar to the ambient size distribution. Neither fish showed a daily activity cycle, and both appear food-limited.     

A benthic predatory fish does not cause selection on armour traits in three‐spined stickleback Gasterosteus aculeatus (Gasterosteiformes: Gasterosteidae)

Predation can promote divergence between prey populations and contribute to ecological speciation. In theory, predators can also constrain prey population divergence. In coastal British Columbia, Canada, Gasterosteus aculeatus (three‐spined stickleback) species pairs only occur in lakes with a single species of predatory fish: Oncorhynchus clarkii (the cutthroat trout). Similar lakes containing additional predatory fish species (Cottus asper, prickly sculpins; Oncorhynchus mykiss, rainbow trout) contain only single species of morphologically intermediate stickleback, suggesting that these predators prevent the coexistence of stickleback species pairs. We conducted a mesocosm experiment to investigate how prickly sculpins might constrain divergence, by quantifying their impact on survival and natural selection on antipredator (armour) traits in F₂ stickleback from a cross between ecologically divergent populations. We tested three hypotheses: (1) sculpin predation on sticklebacks reduces survival in a way that could result in their exclusion from certain niches; (2) sculpins compete with stickleback; (3) sculpins respond to prey vulnerabilities in similar ways to cutthroat trout, tending to constrain rather than to enhance divergence. We found that sculpins significantly reduce stickleback survival, that their presence per se does not reduce growth in stickleback, and that predation did not result in selection on any of the armour traits measured, or on gill raker length, which is an important trophic trait. These results tend to refute hypotheses (2) and (3), while supporting hypothesis (1). © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 877–885.     

Influences of slimy sculpin (Cottus cognatus) predation on the rocky littoral invertebrate community in an arctic lake

We tested the role of the slimy sculpin (Cottus cognatus), a benthic fish, in structuring the rocky littoral invertebrate community in Toolik Lake, Alaska. Comparisons of sculpin gut contents and prey community structure indicated that these fish forage selectively, eating proportionally more large and motile prey, and proportionally fewer small and sessile forms. Field experiments compared the effects of natural, reduced and elevated sculpin densities on benthic community structure. At natural levels of sculpin density, biomass of trichopteran larvae were reduced by more than 50%, and predatory chironomid larvae by 27%, in comparison to areas where sculpin were excluded. Tube-dwelling and small free living chironomid larvae were unaffected at normal sculpin densities. Under artificially high sculpin densities, there was some reduction of tube-dwelling chironomids, but the small free living ones remained unaffected. There appears to be a threshold length of about 3.5 mm, below which chironomid larvae are free form sculpin predation. Tube-dwelling chironomids may be longer than this threshold, but still avoid predation by having most of their body hidden in their tubes.     

Effects of fish predation on larval chironomid (Diptera: Chironomidae) communities in an arctic ecosystem

Chironomid communities were analyzed in systems with three types of predator regimes to determine hierarchical effects of predation; ponds without fish present, lakes with slimy sculpin (Cottus cognatus) present, and lakes with slimy sculpin present along with burbot (Lota Iota) and lake trout (Salvelinus namaycush). Samples were collected by coring bare sediment habitats in 4 systems of each type near the Toolik Lake field station in northern Alaska. Lakes with burbot and lake trout present in addition to slimy sculpin displayed significantly higher (P<0.05) biomass, density, richness, and diversity. This is likely due to the increased complexity of the predator regime. Ponds without fish had a significantly greater percentage of predacious chironomids present.     

Non-visual feeding behavior of the mottled sculpin,Cottus bairdi,in Lake Michigan

Synopsis Field and laboratory experiments indicate that the mottled sculpin, Cottus bairdi, feed in the dark. Blinded sculpins feed on a variety of motile prey in the laboratory and show stereotyped responses to prey stimuli. The sculpins bite at moving inert objects, even if buried in substratum, indicating that they use their lateral line system to detect prey. Covering portions of the lateral line with an inert paste eliminates response to objects near the covered region of the lateral line. The sculpins can also detect prey (including inert objects) in a stream if the prey is upstream. Collection from two series of presunset, postsunset, presunrise, postsunrise, dives in Lake Michigan indicate nocturnal feeding by the mottled sculpin.     

Ectodysplasin signalling genes and phenotypic evolution in sculpins (Cottus)

Despite their deeply conserved function among vertebrates, ectodysplasin (Eda) signalling genes are involved in microevolutionary change in humans and sticklebacks. If such a dual role is common, Eda signalling genes constitute hotspots for morphological evolution. Variation in sculpin (Cottus) skin prickling and body shape resembles patterns caused by variation in Eda signalling in sticklebacks. We mapped Eda signalling genes and performed quantitative trait locus mapping in crosses between Cottus rhenanus and Cottus perifretum. A genomic region containing the Eda receptor (Edar) was strongly associated with prickling and contributed to shape. The expression of Edar in developing prickles and skeletal elements in Cottus was confirmed by in situ hybridization. Coding sequence changes between Edar alleles in C. rhenanus and C. perifretum exceeded sequence differentiation in other vertebrates. However, it is likely that additional genetic elements besides coding changes affect the phenotypic variation. Although the phenotype in a natural hybrid lineage between C. rhenanus and C. perifretum resembles C. perifretum, the respective coding Edar alleles are not fully fixed (88.6%). Hence, our results support an involvement of Eda signalling in microevolutionary changes, but imply that the Edar gene is affected by multiple evolutionary processes that vary among freshwater sculpins.     

Feeding rate of slimy sculpin and burbot on young lake charr in laboratory reefs

Synopsis Predation and contaminants are two possible factors in the poor recruitment of young lake charr Salvelinus namaycush in the Great Lakes. We measured the feeding rate of slimy sculpins Cottus cognatus and burbot Lota lota on young lake charr (uncontaminated young from eggs of a hatchery brood stock and contaminated young from eggs of Lake Michigan lake charr) in laboratory test chambers with a cobble substrate. The median daily consumption rate of sculpins for all tests was 2 lake charr eggs (N = 22 tests; 95% confidence interval, 0–13) and 2 lake charr free embryos (N = 31 tests; 95% confidence interval, 0–10). Feeding rate did not differ between hatchery and contaminated prey. Slimy sculpins continued to feed on lake charr when another prey organism, the deepwater amphipod Pontoporeia hoyi, was present. Feeding by burbot on free embryos (4–36 d^–1) increased as the mobility of young increased, but burbot consumed about 10% of their body weight weekly in free-swimming young (140–380 d^–1). Predation on lake charr eggs by sculpins could be considerable over the 100 to 140 d incubation period, and burbot could eat large numbers of free-swimming lake charr as the young fish left the reef. Predation pressure on young lake charr may inhibit rehabilitation of self-sustaining populations of lake charr on some reefs unless a critical egg density has been reached.     

Contemporary hybrid speciation in sculpins (Cottus spp.)

Natural hybridization between closely related taxa is frequent in many organismal groups, yet it has long been perceived as a force preventing diversification and speciation, especially so in animals. In recent years, growing evidence in favour of hybridization facilitating adaptive divergence has accumulated ( Mallet 2007 ; Mavárez & Linares 2008 ; Nolte & Tautz 2010 ). Homoploid hybrid speciation (the formation of hybrid lineages without changes in chromosome number) occurs when distinct species come into contact, hybridize, and at least in part of their range, produce hybrid swarms. If the hybrid genotypes can then colonize areas of the adaptive landscape inaccessible to ancestral species, they may eventually form new distinct lineages, reproductively isolated from their ancestors. Invasive sculpins (Cottus sp.) are one of a few good examples of homoploid hybrid speciation in animals. In this issue, Stemshorn et al. (2011) identified three distinct hybrid lineages, which have emerged out of a secondary contact situation of Cottus rhenanus and Cottus perifretum. Hybrids have recently invaded large river habitats unsuitable to ancestral species. Through the use of genetic mapping, the authors established that contrary to expectations, chromosomal rearrangements were not apparent in the hybrid lineages. In addition, different population genetic models were tested and the results suggest that contemporary gene flow from ancestral species represents an important component of the system. As such, recent and ongoing hybridization appears to be promoting the appearance of phenotypes adapted to novel environments. The examination of partially isolated lineages such as invasive hybrid sculpins should permit to identify early adaptive genetic changes before they become confounded by differences arising once speciation is complete.     

Comparative life histories and microhabitat use in three sympatric sculpins (Cottidae:Cottus) in northeastern California

Synopsis Life histories of three sculpins endemic to the Pit River system, northeastern California, are detailed.Cottus pitensis is widely distributed throughout the drainage. This species is typically found in rapidly flowing, shallow water on coarse substrate. It grows to a large size, >100 mm, and lives to 5 years. Spawning occurs annually, in early spring. Fecundity is relatively low, <320 oocytes.Cottus asperrimus is restricted to a large population in Fall River and smaller concentrations in Hat Creek and Pit River. It is a relatively small fish, <80 mm, and lives to 5 years. It occurs in deep, slow-moving, often spring-fed streams, typically on fine substrates. Spawning lasts from autumn to early spring. Males establish nests on hard substrates and guard several clutches of eggs. Fecundity ranges from 140–580 oocytes.Cottus klamathensis macrops is relatively rare in the drainage. The largest concentration is in the Hat Creek system, smaller populations exist in Fall River and Pit River. This sculpin is found on coarse substrate, often where aquatic vegetation is abundant. It is typically found in slow, relatively shallow water. It grows to >100mm and lives to 5 years. Males guard eggs during the short spawning season beginning in late winter. Fecundity in this species reaches a maximum of 650 oocytes. Life history strategies vary within the genus, yet the life histories of these sculpins are more similar to each other than they are to many of their congeners. This within-group similarity may have resulted from exposure of all three species to similar environmental pressures. This is predicted by life-history theory.     

Life-history variations in the fluvial sculpin, Cottus nozawae (Cottidae), along the course of a small mountain stream

Life-history variations in male and female fluvial sculpins, Cottus nozawae, were studied in a small mountain stream in Hokkaido, Japan, primarily by using capture-mark-recapture methods. At three study areas established along the stream course, the majority of marked sculpins were recaptured in their original location over one or more years, indicating their long-term occupation of each restricted habitat area. Sculpin densities increased toward the upstream habitats, whereas individual growth rates were more rapid downstream. In both sexes, sculpins distributed downstream matured at a larger body size and later in life than upstream sculpins, clearly demonstrating a clinal variation in these respects. A comparison of life-history variations in C. nozawae with those in amphidromous C. hangiongensis suggests that intrapopulational life-history variations in the former might be environmentally induced, and that one of the most important determinants for the variations in Cottus species might be population density.