Salvelinus namaycush spawning substratum attracts egg predators and opportunists through chemosensory cues

Two separate field experiments were conducted in a series of small boreal lakes to test for the attraction of egg predators to lake trout Salvelinus namaycush spawning shoals and subsequently to determine whether chemosensory cues attract egg predators to these sites. In the first experiment, minnow traps set on spawning sites captured significantly more egg predators than those set on structurally similar non‐spawning sites. Captures of slimy sculpin Cottus cognatus, common shiner Luxilus cornutus, blacknose shiner Notropis heterolepis and virile crayfish Orconectes virilis were more than double on spawning sites relative to non‐spawning sites for the two study lakes. To test whether chemosensory cues could attract egg predators to S. namaycush spawning sites, paired minnow traps were placed on eight to 10 sites in each of the three study lakes; one trap contained visually concealed S. namaycush spawning substratum and the other with visually concealed non‐spawning substratum. Traps containing spawning substratum consistently captured more fish and had higher mean daily catches than those that contained non‐spawning substratum. The combined results demonstrate a greater prevalence of egg predators on S. namaycush spawning shoals that appears to be the result of chemosensory attraction to spawning substratum.     

Lake trout (<Emphasis Type="Italic">Salvelinus namaycush</Emphasis>) spawning behaviour: the evolution of a new female strategy

A video camera inside an underwater housing was used to record the spawning activities of lake trout, Salvelinus namaycush, in Kushog Lake, Ontario (Canada). Contrary to the commonly accepted belief describing lake trout as the only salmonine to spawn exclusively at night, the recordings were taken during the day. Lake trout spawning behaviour is described in detail and compared with other salmonine species. The loss of female “nest-building” and assumption of a novel itinerant strategy during which females travel across the breeding grounds accompanied by a group of males, is considered to be an important change in the evolution of the lake trout’s unique mode of spawning.     

Midwater spawning inHaplochromis chrysonotus (Boulenger) (Teleostei: Cichlidae) in Lake Malawi

Synopsis Haplochromis chrysonotus, a semipelagic shoaling cichlid from Lake Malawi, has been observed spawning in open water up to ten metres above the substratum. It is suggested that open water spawning without the establishment of a substratum-based territory eliminates competition with other species for breeding space and that it may be a stage in the evolution of a totally pelagic mode of life.     

Lake water level increase during spring affects the breeding success of bream Abramis brama (L.)

In Lake Constance, Eurasian bream Abramis brama (L.) spawn in very shallow littoral areas by the beginning of May. They attach their adhesive eggs to pebble and cobble substratum at <40 cm depth. Increasing water levels before spawning inundate bare substratum to which bream eggs may attach better than to deeper substratum covered by epilithon. Consequently, the water level increase prior to spawning should determine the amount of pristine spawning substratum available to bream and thus influence their breeding success. In order to test this hypothesis, the influence of hydrology and climate on the abundance of age-0 bream was combined with the results from field investigations on the egg survival and abundance of age-0 bream. A strong positive correlation between the mean water level increase during the spawning season of bream (April–May) and the abundance of juvenile bream was found. In contrast, the absolute water level during spawning and during the nursery stage in summer, the cumulative temperature during the egg, larval and juvenile stages and two North Atlantic Oscillation (NAO) indices did not affect the abundance of juvenile bream. The field investigations confirmed that bream eggs attach better to and have higher survival rates on bare substratum than on substratum with epilithon cover. Accordingly, eggs within a spawning habitat of bream were most abundant between 10 and 20 cm depth, where the epilithon cover was lower than at depths exceeding 30 cm. The results of this study confirm an adverse influence of epilithon cover on the attachment and subsequent survival of bream eggs and emphasize the importance of spring inundations for the successful breeding of the bream. Handling editor: J. A. Cambray     

Anadromy and marine habitat use of Lake trout (Salvelinus namaycush) from the central Canadian Arctic

Anadromy was documented in 16 lake trout, Salvelinus namaycush, from Canada's central Arctic using capture data and otolith microchemistry. For the first time, estuarine/marine habitat use was described for five individuals using acoustic telemetry. Age-at-first-migration to sea was variable (10–39 years) among individuals and most S. namaycush undertook multiple anadromous migrations within their lifetime. Telemetry data suggested that S. namaycush do not travel far into marine habitats and prefer surface waters (<2 m). These results further our collective understanding of the marine ecology of Arctic S. namaycush.     

Evaluating the effects of controlled flows on historical spawning site access,reproduction and recruitment of lake sturgeon Acipenser fulvescens

Lake sturgeon Acipenser fulvescens spawn at the base of Kakabeka Falls, a 39 m waterfall on the Kaministiquia River, a tributary to Lake Superior. Access to this historical spawning site can be restricted or delayed due to hydroelectric flow fluctuations that coincide with the A. fulvescens spawning season. The objectives of this study were to determine (a) the necessary flow conditions that facilitate spawning site access; (b) quantity and duration of flow required for successful spawning and dispersal of larvae; and (c) evaluate recruitment of juvenile A. fulvescens in relation to flow. A. fulvescens spawning migrations were tracked using a stationary telemetry receiver that logged the movements of 166 A. fulvescens fitted with radio-transmitters. Unrestricted access to the spawning site was facilitated when spawning flow was controlled at 23 m³ s⁻¹ in 2004 and 17 m³ s⁻¹ in 2006. Fluctuating (0.5–8.5 m³ s⁻¹) and delayed spawning flows resulted in restricted and delayed access to the spawning site. Flow duration for successful egg incubation, hatch and larval dispersal was determined by sampling larvae using drift nets and quantified using cumulative temperature units (CTU). Over 10 years, 10,083 larvae were captured between 31 May and 20 July with 97% of the drift occurring prior to 30 June. From the date of first spawning to the end of larval dispersal took an average of 38.6 days, and the mean CTU value was 398.6. In general, a minimum flow of approximately 14.5 m³ s⁻¹ from the date of initial spawning to the accumulation of c. 400 CTU ensured successful hatch and larval dispersal. During the timeframe of this study, recruitment was variable. This study described the complex and variable reproductive life history of A. fulvescens and defined spawning flow requirements ecologically, which can be used to develop operational provisions at hydropower facilities to ensure successful reproduction.     

Obituary of Vadim Dimitrovitch Vladykov, 1898–1986

Synopsis The behaviour and ecology of lake charr Salvelinus namaycush, brook charr S. fontinalis, and Atlantic salmon Salmo salar, are reviewed to assess the susceptibility of these species to mortality from exposure to episodic pH depressions and accompanying changes in water quality. Critical life history periods are identified from an examination of: spawning site characteristics, timing of acidic pulses, changes in the sensitivity of developing fish to acidic water, and the ability of these species to detect and avoid deleterious chemical conditions. The three species appeared to be most vulnerable to pH depressions during Snowmelt as: alevins emerging from the redd (brook charr); free embryos in the interstitial spaces of the spawning rubble (lake charr); and embryos during hatching and juveniles during smoltification (Atlantic salmon). Factors that may modify the toxic effects of or the behavioural responses to low pH included: water temperature, acclimation and body energy reserves. The effect of site-specific or episodic mortality of early life history stages on recruitment to the population could not be evaluated due to the lack of information on compensatory mechanisms.     

Effects of intracoelomic transmitter implantation on metabolic rate,swimming performance,growth and survival in juveniles of two salmonids

In this study, we investigated the effects of acoustic tag implantation on standard and routine metabolic rate (SMR and RMR, estimated via oxygen consumption), critical swimming speed (U_crit), survival and growth in juveniles of rainbow trout Oncorhynchus mykiss and lake trout Salvelinus namaycush. Tag burdens ranged from 1.8% to 7.5% across the two species. Growth rates in acoustic-tagged fish were equal to or higher than those in other treatments. Acoustic-tagged S. namaycush had a marginally lower U_crit than controls but that effect was not replicated in the O. mykiss experiment. Tagging did not have clear effects on metabolic rate but there was an interaction whereby SMR and RMR tended to increase with time since surgery in tagged O. mykiss but not in other treatments (the same trend did not occur in S. namaycush). Survival was high across treatments (mean 98% survival among O. mykiss, 97.5% among S. namaycush). There were no statistically significant effects of tag burden (percentage of body mass) except for a weak negative relationship with growth rate (across species) and a weak positive relationship with U_crit but only in the O. mykiss. Collectively, our findings suggest there were minor, context-dependent effects of acoustic tagging in juvenile S. namaycush and O. mykiss during an eight-week laboratory experiment. Further research will be required to assess whether tagging can cause meaningful behavioural effects in these species in captivity or in the wild and whether there is a tag burden threshold above which deleterious effects consistently occur.     

Male spawning success and female choice in the mottled triplefin,Forsterygion varium (Pisces: Tripterygiidae)

A detailed analysis was made of the variation in spawning success among male Forsterygion (э Tripterygion) varium. The aim was to assess whether females were mating selectively and, if so, to examine the basis of that choice. Eggs were laid on small rocks within the males' territories. Females did not distribute eggs randomly among males. Some males guarded as many as 23000 eggs in one breeding season, while others experienced little or no breeding success. Male spawning success was positively correlated with both their size and some characteristics of their territories, namely the amount of spawning substratum and the presence of a large boulder adjacent to the territory. Experimental manipulations indicated that female choice of males was based on these features. Addition of small rocks into the territories of previously unsuccessful males resulted in a significant increase in spawning success. Conversely, spawning success of the most successful males decreased when small rocks were removed from their territories. When successful males were removed, each territory was occupied by small males; however, there was no significant change in the spawning success of these males. This experiment suggested that, although territory quality was important, female mating decisions could also be influenced by male characteristics such as body size.     

The Lek-like Mating System of Halichoeres melanochir (Pisces: Labridae) at Miyake-jima,Japan1

Reproductive behavior and social organization of the labrid fish Halichoeres melanochir were studied at Miyake-jima, Izu Islands, Japan. ♂♂ set up contiguous territories from early to mid-afternoon daily during the breeding season, with ♂♂ at the center of the spawning site experiencing significantly higher spawning success than those at the periphery. Unlike other labrids with lek-like mating systems previously reported, H. melanochir is sexually monochromatic and shows only minimal sexual size-dimorphism. The spawning behavior and mating system are described and compared with current theories on labrid reproductive strategies.     

Effects of substratum restoration on salmonid habitat quality in a subalpine stream

Stream substratum restoration is a widely applied tool to improve spawning habitat quality for salmonid fishes. However, there is a lack of studies which comprehensively assess effects of the restoration on site, as well as on downstream habitats. Our study addressed effects at both locations and compared abiotic (analyses of texture, penetration resistance, oxygen concentration, redox, nitrite, nitrate, ammonium, pH, electric conductivity, temperature) with biotic (depth-specific macroinvertrebrate abundance and diversity, brown trout hatching success) indicators before and after excavation of the substratum in a highly colmated brown trout spawning site. Strong improvements of hyporheic water conditions (increased oxygen supply and redox potential, reduced concentrations of nitrite and ammonium) as well as ~50 % reductions of substratum compaction and fine sediment content were observed 1 day after the restoration measure. Improvements of habitat quality were still detectable 3 months after treatment. Consequently, the hatching success of Salmo trutta eggs increased from 0 % to 77 % after the restoration. Short-term decrease of macroinvertebrate abundance (from 13.1 to 3.9 macroinvertebrates/kg substratum) was observed within the hyporheic zone of the restoration site, but after 3 months, the number of taxa increased from 13 to 22 taxa and abundance reached 17.9 macroinvertebrates/kg. Significantly increased fine sediment deposition was detected within 1 km downstream of the restoration site and may negatively affect these habitats. Trade-offs between positive effects at restored sites and negative effects in downstream habitats need to be considered for a comprehensive evaluation of stream substratum restoration.     

Effects of Pacific salmon spawning and carcass availability on the caddisfly Ecclisomyia conspersa (Trichoptera: Limnephilidae)

1. The effects of spawning coho (Oncorhynchus kisutch) and chum salmon (Oncorhynchus keta) on the limnephilid caddisfly Ecclisomyia conspersa were evaluated by experimentally excluding salmon from the upper 14‐m stretch of a spawning channel by a wire‐meshed fence. Density, and development and growth rates, of larvae upstream of the fence (without salmon) were compared with those downstream (with salmon). 2. Larval density in the stretch with salmon declined during spawning, but increased again after spawning subsided and the carcasses of dead fish became available. In the stretch with salmon, larval density on salmon carcasses was seven to 37 times greater than on the adjacent channel substratum. The rate of larval development in the stretch with salmon was greater than that in the stretch without salmon. Two months after carcasses became available, 98% of larvae sampled from the stretch with salmon were in the fifth instar, compared to only 23% from the stretch without salmon. Body weight of E. conspersa in the stretches with and without salmon increased by an average of 3.04 and 2.38 mg, respectively, over a 6‐month period. 3. ¹⁵N values of larvae from the stretch with salmon increased following the arrival of the fish, suggesting that the larvae were feeding on salmon‐derived material, such as eggs and carcasses, which contain a high proportion of the heavier stable isotope. In contrast, ¹⁵N values of larvae from the stretch without salmon remained relatively constant throughout the experiment. The availability of salmon carcasses as a high‐quality food source late in larval development may increase survival and fecundity of E. conspersa. 4. These substantial differences were consistent with the view that they were due to the experimental exclusion of salmon and salmon carcasses from the upstream stretch, though the study was un‐replicated and thus precludes ascribing causation more definitely.     

The pre‐spawning migratory behaviour of Atlantic salmon Salmo salar in a large lacustrine catchment

The movements of adult Atlantic salmon Salmo salar were determined as they migrated to spawning habitats in a large lacustrine catchment, Lough Neagh, in Northern Ireland. The minimum average ground speed of S. salar through the lake was 2·1 km day^?1 and the mean residence time was 11 days. Tagged S. salar tended to actively migrate through the lake which represented a transitory habitat for adult S. salar. Migration time from the release site, through the lake, to a spawning tributary decreased during the migratory period. During the 4 year study period between 20·5 and 41·6% of tagged S. salar which entered the lake each year, explored at least one other channel before ascending the final spawning tributary. Exploratory behaviour was more likely in S. salar which spawned in the tributaries furthest from the sea. Exploratory behaviour was also more likely to occur during periods of reduced discharge in the natal stream. The fishery management implications of complex pre‐spawning behaviour in a mixed stock lacustrine system, are discussed.     

Interspecific variation in spawning time and male mating tactics of the parrotfishes on a fringing coral reef at Iriomote Island,Okinawa

Spawning time and male mating tactics of parrotfishes (family Scaridae) were investigated on a fringing coral reef at Iriomote Island, Okinawa. Spawning was observed in 14 species, and more frequently in more abundant species such as Chlorurus sordidus, Scarus rivulatus and Chlorurus bowersi. At the reef-edge spawning site, C. bowersi spawned at high tide, C. sordidus spawned both at high tide and in the early morning, whereas Calotomus carolinus and most of the Scarus species such as S. rivulatus spawned only in the early morning, mostly 0630–0830 h. Spawning only in the early morning irrespective of tide phase and moon age has seldom been reported from the scarid species of other localities. It is suggested that spawning in the early morning would be adaptive in species such as S. rivulatus, which migrated considerable distances (ca. 500 m) to the inshore feeding sites, in order to minimize feeding losses due to migration. For male mating tactics, pair spawning by territorial TP (terminal phase) males occurred in all 14 species, and streaking and group spawning by nonterritorial small IP (initial phase) males were seen more frequently in more abundant species. Moreover, group spawning by nonterritorial TP males, which were larger than the IP males but smaller than the territorial TP males, frequently occurred in S. rivulatus. Such mating tactics of TP males have not been reported from Scaridae.     

Timing and site selection of spawning in a landlocked population of rainbow smelt in Wisconsin

To study the site selection and timing of spawning by rainbow smelt, Osmerus modax, in a freshwater system, smelt were sampled with fyke nets during the spawning run in spring 2002 and 2003. The sex ratio of smelt favored males in the early and late segments of the run, with over 90% of smelt caught being male. At the peak in the catch, females comprised 57% of the smelt collected. The gonadosomatic index of males and females declined significantly on the day of peak catch, signaling the peak in spawning activity. Higher catches of smelt occurred on cobble-dominated substrates during the peak of the run, but no site selection was apparent earlier or later. Groundwater outflow did not appear to affect catches. Our study suggests that smelt come inshore but do not select specific spawning substrates prior to or after the peak in spawning. Smelt do select cobble sites during the highest rates of spawning. The relation between the peak in spawning and site selection could have application in planning removal strategies for the species.     

Radiocaesium concentrations in the muscle and eggs of salmonids from Lake Chuzenji,Japan, after the Fukushima fallout

Approximately 18 months (September to December 2012) after the Fukushima Dai‐ichi Nuclear Power Plant accident, elevated radiocaesium concentrations were measured in samples of muscle and eggs from masu salmon Oncorhynchus masou, kokanee Oncorhynchus nerka, brown trout Salmo trutta and lake trout Salvelinus namaycush from the Lake Chuzenji system, central Honshu Island, Japan (160 km from the station). Mean muscle concentrations were 142·9–249·2 Bq kg^?1 wet mass and mean concentrations in eggs were 38·7–79·0 Bq kg^?1 wet mass. There was no relationship between fork length and muscle radiocaesium concentration in any of the species, but there were significant relationships between individual muscle and egg radiocaesium concentrations from O. masou, S. trutta and S. namaycush.     

Field activity of lake trout during the reproductive period monitored by electromyogram radiotelemetry

Many lake trout Salvelinus namaycush spawn in shallow areas along windswept shores. However, precise determination of time and location of spawning is limited by its nocturnal occurrence, and possibly by some postulated, but unproven, spawning in deep water by certain populations. In White Pine Lake, Ontario, a well-studied experimental lake, lake trout spawn in two shallow locations where some direct observation is possible. Radiotelemetry apparatus was used in 1991 to obtain records of the electromyograms (EMGs) produced by muscle activity in one adult male and one adult female lake trout, captured from the lake before their reproductive period and returned to the lake at the beginning of the spawning period. Both fish survived, though only the male was detected as active over the main spawning site, where it showed evening EMG activity patterns indicating considerable activity. At other times of day, when not located over the spawning site, but present elsewhere in the lake, the male's EMG (i.e. muscle) values were considerably reduced. At the cessation of the spawning period, the transmitter-equipped male's EMG record showed no further pattern indicative of high activity during what had formerly been the daily spawning period. The female was tracked as she moved around the lake, but based on both her location and level of EMG activity, is thought not to have spawned. In 1992, a second male was captured, equipped with a transmitter and released again. This fish failed to show spawning activity but, as with the 1991 female, moved fairly considerable distances in the lake. In one instance, in a swim of short duration (6 min), well away from the spawning grounds, this fish displayed EMG activity levels resembling those of the high activity of spawning. The results obtained appear to demonstrate that EMG telemetry could be used to assess the level of muscular activity occurring in lake trout, especially in relation to reproductive behaviour, and when they cannot be directly observed.     

The spawning behaviour of the endangered Eastern Cape rocky, Sandelia bainsii (Anabantidae), in South Africa

Sandelia bainsii is an endangered freshwater fish species which only occurs in short sections of several Eastern Cape river systems in South Africa. The spawning behaviour of S. bainsii is described for the first time. Female S. bainsii remained a light grey colour. Males exhibited distinctive nuptial coloration consisting of an intense black with a white bar on their caudal fin and white windows on the posterior areas of their dorsal and anal fins. There was a darkened post orbital contact organ of ctenoid scales on the reproductively active males. The spawning behaviour was recorded on a video recorder and analyzed during frame by frame replay. Sandelia bainsii are uniparental substrate breeders. The male prepared a spawning area by clearing the substrate with rapid body movements (sweeps). There were a number of preliminary courtship movements, several trial embraces by the male before the actual final embrace. The male underwent a U bend around the female, anchored himself with the contact organ, squeezed the abdominal area of the female with a quick head movement. Courtship and spawning all took place near the substratum. Twenty one separate mating bouts were filmed and analyzed. The duration varied from 1.44 to 5.11 seconds with an average embrace of 3.39 seconds. During the mating clench the female was not normally turned over, as in some anabantoids, but remained upright. The male then guarded the spawning area. Eggs were light yellow, adhesive, demersal and 1.28 mm in diameter with 0.61 mm oil globules. The breeding behaviour of S. bainsii has some similarities with both breeding behaviourial groups, the bubble nesters and the free spawners, within the other African anabantid genus, Ctenopoma. Sandelia bainsii have acquired some temperate specializations with regard to their reproductive style. Free spawning and the release of buoyant eggs is the plesiomorphic anabantid reproductive style. The demersal eggs and male substratum nest sites of S. bainsii are considered apomorphic.     

Brown trout (Salmo trutta,Linnaeus, 1758) spawning habitat improvement in the Ebro River Basin (eastern Spain)

Limestone streams in Spain are subject to calcium carbonate precipitation processes, which make the substrate very cohesive. Many populations of brown trout (Salmo trutta, Linnaeus, 1758) in those rivers show reduced production due to unsuitable spawning habitats. This study presents data on S. trutta spawning habitat improvement in the Guadalope River Basin during 2011. The main objectives of this study were to (i) modify the substrate in order to make it easier for S. trutta females to dig when spawning, (ii) test the effect of the gravel quality improvement on the habitat, (iii) analyse the effect of habitat improvement on S. trutta recruitment, and (iv) monitor habitat improvement effectiveness in the long term. Three manual measures were adopted to make the gravel suitable for S. trutta breeding: the separation of the embedded gravel using an iron bar by hitting it with a mace‐hammer (‘substrate breaking’), the spreading out of the gravel loosened by the previous action with a rake (‘raking up’), and thirdly the elimination of fine sediment depositions (‘substrate cleaning’). Results of the first year of monitoring (2012) in improved and control areas were compared. Methodology effectiveness assessment requires monitoring over a longer period, but the data from the first year after habitat improvement provide encouraging results, mainly dealing with changes in spawning habitat parameters and an increase in the S. trutta age‐0 class in the improved areas.