Factors affecting the duration of nest defense and reproductive lifespan of female sockeye salmon, Oncorhynchus nerka

Reproductive success of female animals is often affected by a combination of fecundity and parental care. In female salmonid fishes, acquisition of nest (redd) sites and prevention of their use by other females are critical to reproductive success. These factors are particularly important for stocks that spawn at high densities. Body size is positively correlated with fecundity and egg size, and has been hypothesized to control the outcome of intrasexual competition and longevity. We tested this hypothesis by evaluating the influences of body size, intrasexual aggression and arrival date on duration and success of redd guarding by female sockeye salmon, Oncorhynchus nerka, in a small Alaskan creek. Contrary to the hypothesis, larger females guarded their redds no longer than smaller females, and did not live as long in the stream. Aggression was not related to body size or overall longevity but was positively correlated with residence period on the redd. Females that entered the creek earlier lived longer, spent longer on their redds, and spent more time guarding their redds after spawning than females that entered the creek later. However, despite their longevity, early-arriving fish were more likely to have their redds reused by another female because they died before all the females had selected redd sites. The small average body size in this stock is consistent with weak selection for large size, and with our evidence that size provided little if any advantage in nest guarding.     

Spawning migration and redd construction of Sakhalin taimen, Hucho perryi (Salmonidae) on northern Hokkaido Island, Japan

Spatial and temporal patterns of redd construction by Sakhalin taimen, Hucho perryi , were studied in small streams on northern Hokkaido, Japan. Spawning took place in late April with close relation to discharge patterns. Observed redd size (mean ± 95% C.I.) was 145 ± 11·8 cm in length and 84 ± 5·8 cm in width. The number of egg pockets per redd was either one or two, which is fewer than that of other salmonids reported in the literature. Spawning ground distributions in 1991 and 1992 were similar to each other in terms of the distribution mean and dispersion of the redds. Unlike other salmonids Sakhalin taimen probably constructed multiple redds during one spawning season.     

Tradeoffs between homing and habitat quality for spawning site selection by hatchery-origin Chinook salmon

Spawning site selection by female salmon is based on complex and poorly understood tradeoffs between the homing instinct and the availability of appropriate habitat for successful reproduction. Previous studies have shown that hatchery-origin Chinook salmon (Oncorhynchus tshawytscha) released from different acclimation sites return with varying degrees of fidelity to these areas. To investigate the possibility that homing fidelity is associated with aquatic habitat conditions, we quantified physical habitat throughout 165?km in the upper Yakima River basin (Washington, USA) and mapped redd and carcass locations from 2004 to 2008. Principal components analysis identified differences in substrate, cover, stream width, and gradient among reaches surrounding acclimation sites, and canonical correspondence analysis revealed that these differences in habitat characteristics were associated with spatial patterns of spawning (p?<?0.01). These analyses indicated that female salmon may forego spawning near their acclimation area if the surrounding habitat is unsuitable. Evaluating the spatial context of acclimation areas in relation to surrounding habitat may provide essential information for effectively managing supplementation programs and prioritizing restoration actions.     

Spawning tactics of summer chum salmon Oncorhynchus keta in relation to channel complexity and hyporheic exchange

Vertical hydraulic gradient, channel hydraulic, and substratum data were collected at active chum salmon redds to evaluate their influence on redd site selection in the Kwethluk River, a wandering gravel-bedded tributary of Alaska’s Lower Kuskokwim River. Two distinct summer chum salmon spawning tacitcs: primary- and off-channel groups spawning in different hydraulic and thermal environments. Channel hydraulics, ground-surface water exchanges, and bed sediment data were collected throughout the range of topographic conditions at sites occupied and unoccupied by spawners. Analyses of these data showed that redd site selection was incoherent with channel hydraulics and substratum size, when all channel types were considered. Rather, spawning was associated with the exchange of river and groundwater at four spatial scales. In primary channels, chum salmon spawned only in reaches where the regional hydraulic gradient (regional scale) promoted the penetration of river water, or downwelling, into the bed sediments. Within these reaches, primary-channel spawning was influenced by bed topography (fluvial scale) that promoted strong localized downwelling; a logit model, fitted to constituent hydraulics, discriminated between selected and unselected sites with 89 % accuracy. In contrast, spawners in off-channel habitats (flood and spring channels) selected areas of upwelling groundwater, regardless of channel hydraulics and the regional hydraulic gradient. In parafluvial flood channels, spawners selected localized upwelling associated with short groundwater flow paths (parafluvial scale). In spring channels, within the floodplain’s forested riparia (orthofluvial scale), spawners eschewed areas of localized downwelling and uniformly selected upwelling zones feeding these channels. Thus, redd selectivity differentiated two spawning tactics (groups) in this complex channel network where ground and surface water exchange is an important predictor of spawning habitat.     

Population regulation in adult, but not juvenile, resident trout (Salmo trutta) in a Lake District stream

1. Unlike a neighbouring sea-trout population that showed strong density-dependent survival, a resident trout population ( Salmo trutta L.) showed simple proportionate survival in the early life-stages. However, this persistent population fluctuated within narrow limits. Mature adults, especially during spawning, were the only possible life-stage left in which regulation might occur. 2. An October census, just prior to spawning, was made at five sites (total area 300 m ² ) from 1965 to 1983. Gravel nests (redds) associated with females of known size were excavated outside these sites to obtain a power-function relationship between egg density per redd and female length (range 181–280 mm, n = 26). This relationship and the census data for females (range 186–284 mm) were used to estimate egg densities in each year-class. 3. The census data for the early life-stages (0+, 1+, 2+ trout) confirmed proportionate survival with no evidence for density-dependent regulation. In contrast, the number of spawning females produced in each year-class was strongly density dependent on the initial number of females that laid eggs at the start of the year-class. Similarly, total egg production in each year-class was density dependent on initial egg density. 4 Both relationships were well described by the Ricker and Beverton-Holt stock-recruitment models (P < 0.001) and the goodness-of-fit was similar for both models. This study is probably the first to provide clear evidence for fish population regulation in the adult, rather than the juvenile, stage.     

Observations on interbreeding between dwarf and normal arctic charr,Salvelinus alpinus,from Stora Rösjön,central Sweden

Synopsis The spawning behaviour of dwarf and normal arctic charr, Salvelinus alpinus, was studied through trough observations in October 1988. The experimental fish consisted of reared, three-summer old progeny of artificially bred dwarf and normal charr from an isolated body of water in central Sweden. The spawning behaviour of three normal female charrs was studied and video-recorded. The study showed that the arctic charr is potentially polyandric both within and between forms. The normal female was courted and mated both by normal and dwarf males. It was also noted that antagonistic behaviour on behalf of the female was much more common towards dwarf males than normal males. Guarding tactics were only observed in normal males, whereas sneaking tactics were employed both by dwarf and normal males. The dwarf as well as normal males mated the female charr either by courting the female or by joining the spawning fish at oviposition. Only dwarf males were observed resting silently on the bottom close to the redd.     

The structure and dimensions of redds and egg pockets of the endangered salmonid, Sakhalin taimen

Each redd of the Sakhalin taimen Hucho perryi had a V–shaped pot that is specific to this species. The shape of each multiple–egg–pocket redd was a combination of one–egg–pocket redds, and the number of egg pockets in the redd could be estimated from the shape. False redds were small and did not have any V–shaped pots. The factors affecting redd size were examined numerically; consequently, numbers of egg pockets and eggs and female fork length were correlated significantly with tail length. Each egg pocket contained 11–920 eggs (mean: 546·7±95% CL 81·9) and the egg pocket depth was correlated significantly with female fork length. Mean number of redds per female was three, and the number of females could be estimated as one third of the redds.     

Spawning success in Atlantic salmon (Salmo salar L.): a long-term DNA profiling-based study conducted in a natural stream

Spawning success of Atlantic salmon (Salmo salar L.) was investigated, under near-natural conditions, in the Girnock Burn, an 8-km long tributary of the River Dee in Scotland. Employing minisatellite-based DNA profiling, mating outcomes were resolved over three spawning seasons by assigning parentage to progeny samples removed from spawning nests ('redds'). While individual spawning patterns differed markedly, consistent trends were present over the 3 years studied. Multiple spawning was found to be prevalent. More than 50% of anadromous spawners of both sexes contributed to more than one redd. Up to six redds for a single female and seven for a single male were detected. Both sexes ranged extensively. Distance between redds involving the same parent varied from a few metres to > 5 km. Distances > 1 km were common. Both males and females ranged to a similar extent. Range limit was not correlated to fish size. Pairs were not monogamous, both males and females mating with different partners at different sites. Size assortative mating was apparent among 1991 spawners but was not detected for 1992 or 1995. Redd superimposition was found to be common (17-22% of redds over the 3 years), although it was not correlated to the number of anadromous spawners present. High levels of nonanadromous mature parr mating success (40-50% of total progeny sampled) were recorded, and these likely contribute greatly to the effective population size. The relevance of these findings at the individual and population level is discussed, with particular reference to management implications.     

The effect of salmon redd excavation on stream substrate and benthic community of two salmon spawning streams in Canterbury,New Zealand

The effects of redd excavation by female quinnat salmon (Oncorhynchus tshawytscha walbaum) on the bentic invertebrate communities of two stable springfed spawning streams in the headwaters of the Rakaia River, South Island, New Zealand, were studied during January to September 1982. The gross effects of salmon spawning on the streams appear to be: the loss of instream macrophytes, algae and mosses, a loss of fines and detritus, and a modification of the pool-riffle character. Redd excavation may be associated with a reduction in benthic standing crop of up to 83.6%, with only a minor effect on the benthic community structure. Benthic recolonisation rates were similar for both spawning streams, with almost complete recovery after 95 days.     

Spawning behaviour of taimen (<Emphasis Type="Italic">Hucho taimen</Emphasis>) from the Uur River,Northern Mongolia

This study presents the first observations of Hucho taimen spawning in the wild based on underwater video recordings. One pair of taimen was monitored during a 19 h period, supplemented with visual observations from two other spawning pairs. We recorded two full spawning events performed in two different locations separated by approximately 30 m. The absence of an established male hierarchy along a nesting female was the most important difference between taimen and other salmonine breeding biology. Taimen spawning, based on our observations, is a single pair event. The male prevented the approach of other males by launching intense attacks that extended for several meters away from the redd. Our data suggests that taimen females, differently from other salmonines, do not cover their eggs immediately after having spawned but rest for a variable number of minutes before covering them.     

Alternative tactics in spawning site selection by brook trout (Salvelinus fontinalis) related to incubation microhabitats in a harsh winter environment

1. Species with distributions that span a broad range of latitudes may have populations that exhibit distinct life history traits associated with environmental gradients. The majority of previous studies have indicated a strong association between spawning site selection by brook trout (Salvelinus fontinalis) and the presence of upwelling groundwater, but does this generalisation extend to the thermal regimes experienced at northern sites? 2. We investigated the role of hyporheic flow in redd site selection by brook trout in a relatively high‐latitude boreal system. Hyporheic flows through streambed substratums can be dominated by groundwater or surface water and may be influenced by the presence of morphological features. For autumn spawners such as brook trout, embryos situated in microhabitats where hyporheic flow in the shallow substratum is groundwater dominant (i.e. warmer in winter) are likely to experience accelerated development rates relative to embryos in redds where there is downwelling surface water (i.e. colder in winter). 3. We measured vertical hydraulic gradients (VHG) at the microhabitat scale and spatial and temporal variation in upwelling/downwelling flow and thermal regimes in brook trout spawning/incubation habitats. Additionally, we noted the proximity of redd sites to stream morphological features (e.g. riffle crests). 4. Our results indicate that upwelling flow was not a decisive cue in redd site selection at the microhabitat scale (10⁰ m) as an approximately equal number of redds were situated in microhabitats with upward flow as compared to microhabitats with downward flow through the redd. Redds situated on bedforms with convex longitudinal profiles (e.g. riffle crests, log steps) were associated with downward flow, whereas redds not immediately adjacent to these bedform features were associated with upward flow. Winter streambed temperatures confirmed that both sites with steady upwelling (i.e. warm incubation regime) and downwelling (i.e. cold incubation regime) were indeed selected by spawners. 5. Our observations that spawners utilised both cold‐regime and warm‐regime sites suggests the existence of distinct reproductive tactics related to hyporheic flow patterns in this boreal system. As temperature is the dominant factor controlling rates of embryonic development, the use of spawning microhabitats with distinct thermal regimes implies substantial differences in the timing of hatching and the phenology of emergence.     

The evolution of redd site selection in brook charr in different environments: same cue,same benefit for fitness*

1. Despite the great interest in characterising fish reproductive habitat, the relationship between the selection of a given spawning site and individual fitness has not been experimentally tested. 2. In this study, we used an in situ experimental approach to determine (i) the relative contribution of substrate characteristics as well as hydrological and physicochemical variables to small‐scale redd site selection by brook charr, Salvelinus fontinalis (Mitchill) and (ii) if hatching and emergence success, used as a surrogate of fitness, are improved in selected compared to non‐selected sites in both lake and stream habitats. 3. Our results show that upwelling groundwater flow was always significantly higher in selected than in non‐selected sites in both lake and stream habitats. We found no significant difference in the mean geometric substrate diameter and no consistent differences in substrate composition between selected and non‐selected sites. Oxygen concentration was higher (significantly so in three of four comparisons) and conductivity tended to be lower in selected than in non‐selected sites, while temperature showed no significant or consistent variations. We found a significant positive relationship between the selection of a given spawning habitat and hatching and emergence success in these systems. 4. These results show that the main cue that brook charr use to select their spawning sites is upwelling groundwater in lake and stream habitats, and that active selection of these sites increases individual fitness. This suggests that natural selection acted on the same cues in lentic and lotic environments; this could have been highly adaptive in a species that used both habitats as colonisation routes after the last glaciation.     

Spawning tactics of paired males of the dark chub,Zacco temmincki,reflect potential fitness costs of satellites

Synopsis Paired males of the dark chub, Zacco temmincki, buried released eggs by vibrating their anal fin, but this behavior was prevented and eggs were cannibalized when many satellites (males and females) were present. A number of satellite males also caused a loss to paired males in sperm competition on spawning grounds far from shelters. Paired males followed repeating tactics, which were defined as successive spawning acts at the same redd, in most cases, but occasionally did shifting tactics which refer to spawning acts conducted successively at different redds. The proportion of the shifting tactic was not correlated with the dominance status of paired males. The shifting tactic was not advantageous in performing spawnings frequently. The calculation of the total advantage of both tactics indicated that the shifting tactic itself was not more beneficial than the repeating tactic at any density of satellites. Since a number of satellites stayed around the redd when no spawning pair was present, or pursued a pair or a single dominant male moving between spawning grounds, the occasional shifting tactics of paired males functioned to confuse and disperse satellites. The spawning tactics of paired males apparently reflected potential fitness costs of satellites. Paired males mainly spawned at the redds near shelters, despite the fact that more satellites were present to devour eggs, presumably because they could obtain many females and monopolize fertilizations.     

Fecundity and egg density in the redd for sea trout

The relationship between fecundity and length for sea trout Salmo trutta from three streams in NW England was curvilinear and, after logarithmic transformation of both variables, was well described by a linear regression model. Regression equations for the fish from the three streams were not significantly different ( p >0.05). The relationship between estimated eggs per redd and female size in a tributary of one of the streams was also well described by a linear regression model after logarithmic transformation. Fecundity was usually higher than estimated eggs per redd, and this difference increased with female size so that the mean number of eggs per redd expressed as a percentage of mean fecundity decreased from 100% for a female with a length of 240 mm to 79% for one of 650 mm. The power function in its logarithmic form has now been fitted to fecundity-length data for sea trout from 12 streams and rivers in England, Scotland, Ireland and Norway. Comparisons between fecundity estimates for the same sizes of females revealed a large variation between populations and possible reasons for this are discussed.     

Effects of fish farm effluents on egg-to-fry development and survival of brown trout in artificial redds

Egg-to-fry development and survival of brown trout Salmo trutta were compared in two rivers of the Pyrenean piedmont, the Nive d'Arnéguy, with few human activities, and the Nive des Aldudes, with many anthropogenic activities including 11 fish farms and two sewage treatment plants (STPs). Survival was estimated between spawning (early December) and emergence time (early March) by means of capsules, 7 cm³ in volume, filled with green eggs then inserted into the gravel of artificial redds at spawning sites. In the Nive des Aldudes, three sets of conditions were studied: a fish farm near the springs, a fish farm and an STP in a village, and a series of fish farms and an STP in a village. In each situation, two artificial redds were created upstream and two others downstream from the fish farms. In the Nive d'Arnéguy, four sites were equipped: each of the two downstream sites with two artificial redds, and the two upstream sites with one redd. Substratum characteristics (proportion of fine particles) and the quality of surface and interstitial water (oxygen content, ammonia and nitrite nitrogen) were periodically measured. There was no redd substratum difference between sites upstream and downstream of fish farms. Survival to fry emergence was higher in the Nive d'Arnéguy (63·5%) than in the Nive des Aldudes (47·7%). In this latter river, the nitrogen released from the STPs was 0·5% that from fish farms. Fish farming impaired survival close to fish farm effluents (31·6%), as compared to survival upstream (63·6%), and induced a development delay during the yolk-sac fry stage. These differences were mainly linked to a drop in the dissolved oxygen content in interstitial water induced by the nitrogen flow in surface and interstitial waters.     

Preliminary study on female choice of mates versus sites in a wrasse, Cirrhilabrus temminckii

Female choice of mates versus sites was studied in a wrasse, Cirrhilabrus temminckii. Males had territories within a restricted area on a rocky slope at which females visited and pair-spawned pelagic gametes. Females visited several males or territories before spawning, suggesting the opportunity of female choice. Of the four characteristics of territorial males examined—body size, ratio of pelvic fin length to body size, courtship, frequency and territory depth—only territory depth was significantly correlated with daily mating success of males. The former three male characters were not related to territory depth. These results suggest that female C. temminckii chooses deep sites rather than specific mates in mating.     

Observations on siting, dimensions and structure of salmonid redds

Redds of trout, Salmo trutta L., Atlantic salmon, S. salar L., and rainbow trout, S. gairdneri Richardson, were studied at sites in north-east England, south-west Wales and southern England (Dorset). Spawning females preferred gravels of 20–30 mm diameter and water velocities at 0·6 depth of 15 cm s^-1 and < c. 2·0 body lengths s^-1. After logarithmic transformation the major horizontal dimensions of redds could be related to one another by linear regressions, and redd tail length could, similarly, be related to female fish length. Egg burial depth in upland stony streams in north-east England and south-west Wales could be predicted from a linear regression of mean depth upon female length. Information is also given on gravel composition and redd structure.     

Mating success of alternative maturation phenotypes in male Atlantic salmon,Salmo salar

Summary Mature male parr successfully fertilized eggs of anadromous female Atlantic salmon, Salmo salar, in competition with anadromous males under simulated natural conditions. Mating situations were created in which mature male parr and anadromous males competing for the opportunity to spawn with an anadromous female differed in allelic forms of the same enzyme. Females deposited eggs into a sequence of 3 or 4 eggs nests. The mean proportion of eggs in a redd fertilized by parr increased with increasing numbers of parr present at a redd, reaching 23% at male parr: anadromous male ratios of 20:1. Single male parr fertilized, on average, 5% of the eggs in a redd. The proportion of eggs in an egg nest fertilized by parr also depended upon the order of egg nest construction, such that parr mating success was highest at the initial nest constructed and lowest at the final nest. Parr have relatively high fertilization success for their size when compared with the smaller maturation phenotype of other salmonids.     

Female spawning migrations of the protogynous wrasse,Halichoeres marginatus

Spawning sites and spawning migration paths of tagged females of the protogynous wrasse,Halichoeres marginatus, were studied on the shallow reefs at Kuchierabu-jima Island, Japan. Males set up mating territories above prominent rocks on the offshore reef slope in the late afternoon, and pair-spawned with females, which had migrated there from their home ranges located in inshore areas. Small females migrated to the spawning sites near their home ranges, whereas large females migrated to various spawning sites located within a wide area, including downcurrent sites. Spawning at the downcurrent sites favors transport of eggs offshore, thereby increasing the female’s fitness. The spawning sites where an individual had spawned as a female were subsequently used for mating after it had changed sex. It is suggested that the wide migration of females to various spawning sites, enables the storing up of information on those sites, which later helps in the acquisition of mating territories after changing sex.