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 growth of brown trout (Salmo trutta) in mild winters and summer droughts in upland Wales

SUMMARY. 1. We compared the observed annual growth of 0- and I-group trout in nine Welsh upland streams, with growth predicted from temperature assuming that this was the only limiting factor.
2. Autumn weights of second year fish were 51–67% of predicted ( G _max) values in 1988, but only 30–40% in 1989 and 1990 when drought occurred. Though initial weights of fry were unknown, simulations suggested that first year growth was also less than G _max, but with no obvious effect of drought.
3. To evaluate the possible effects of future climate change, we simulated stream temperature regimes 1.5–4.5°C above those of a recent year with temperatures similar to the long-term average. Growth was set at 60% G _max for both 0- and I-group, or at 40% for I-group to represent the effect of drought. As winter temperature increased, time to hatching and emergence decreased, for example by 56 and 49 days respectively for a rise of 3°C. 0-group growth was slightly enhanced at up to + 3°C but retarded at + 4.5°C. Simulations of I-group growth suggested that warmer winters could enhance trout growth while warmer summers would only increase growth if there were no adverse effects of drought.
4. We discuss many uncertainties in these simulations, which nevertheless suggest the magnitude of possible effects of climate change.     

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 temperature on developmental performance, survival and growth of sea lamprey embryos

This study assesses the influence of thermal regime on the development, survival rates and early growth of embryos of sea lamprey Petromyzon marinus incubated at five constant temperatures (7, 11, 15, 19 and 23° C). The time from fertilization to 50% hatching and from hatching to 50% burrowing were inversely related to incubation temperature. All the embryos incubated at 7° C died at very early stages, while those maintained at 11° C did not attain the burrowing stage. Survival from fertilization to hatching was 61, 89, 91 and 89% at 11, 15, 19 and 23° C, decreasing to 58, 70 and 70% from hatching to burrowing at 15, 19 and 23° C, respectively. Larvae reared during the first 3 months of exogenous feeding in a common environment at constant 21° C, revealed maximum survival for an incubation temperature of 15° C (43% of burrowed larvae) decreasing strongly at 19° C (16%) and 23° C (one suvivor among 240 larvae). Body length at the burrowing stage was maximum for embryos incubated at 19° C, but body mass increased in the interval 15–23° C. Mean incubation temperatures experienced by 117 broods during the embryonic development in the source river were estimated in 15·3±2·30° C and 16·7±1·76° C (mean±1 s.d .) for the periods fertilization-to-hatching and hatching-to burrowing, respectively.     

Influence of hydrologic attributes on brown trout recruitment in low-latitude range margins

Factors controlling brown trout Salmo trutta recruitment in Mediterranean areas are largely unknown, despite the relevance this may have for fisheries management. The effect of hydrological variability on survival of young brown trout was studied during seven consecutive years in five resident populations from the southern range of the species distribution. Recruit density at the end of summer varied markedly among year-classes and rivers during the study period. Previous work showed that egg density the previous fall did not account for more than 50% of the observed variation in recruitment density. Thus, we expected that climatic patterns, as determinants of discharge and water temperature, would play a role in the control of young trout abundance. We tested this by analyzing the effects of flow variation and predictability on young trout survival during the spawning to emergence and the summer drought periods. Both hatching and emergence times and length of hatching and emergence periods were similar between years within each river but varied considerably among populations, due to differences in water temperature. Interannual variation in flow attributes during spawning to emergence and summer drought affected juvenile survival in all populations, once the effect of endogenous factors was removed. Survival rate was significantly related to the timing, magnitude and duration of extreme water conditions, and to the rate of change in discharge during hatching and emergence times in most rivers. The magnitude and duration of low flows during summer drought appeared to be a critical factor for survival of young trout. Our findings suggest that density-independent factors, i.e., hydrological variability, play a central role in the population dynamics of brown trout in populations from low-latitude range margins. Reported effects of hydrologic attributes on trout survival are likely to be increasingly important if, as predicted, climate change leads to greater extremes and variability of flow regimes.     

Effects of temperature and a piscivorous fish on diel winter behaviour of juvenile brown trout (Salmo trutta)

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Timing matters: species‐specific interactions between spawning time,substrate quality,and recruitment success in three salmonid species

Substratum quality and oxygen supply to the interstitial zone are crucial for the reproductive success of salmonid fishes. At present, degradation of spawning grounds due to fine sediment deposition and colmation are recognized as main factors for reproductive failure. In addition, changes in water temperatures due to climate change, damming, and cooling water inlets are predicted to reduce hatching success. We tested the hypothesis that the biological effects of habitat degradation depend strongly on the species‐specific spawning seasons and life‐history strategies (e.g., fall‐ vs. spring‐spawners, migratory vs. resident species) and assessed temperature as an important species‐specific factor for hatching success within river substratum. We studied the species‐specific differences in their responses to such disturbances using egg‐to‐fry survival of Danube Salmon (Hucho hucho), resident brown trout (Salmo trutta fario), and migratory brown trout (Salmo trutta lacustris) as biological endpoint. The egg incubation and hatching success of the salmonids and their dependence on temperature and stream substratum quality were compared. Hatching rates of Danube salmon were lower than of brown trout, probably due to higher oxygen demands and increased interstitial respiration in spring. Increases in maximum water temperature reduced hatching rates of resident and migratory brown trout (both fall‐spawners) but were positively correlated with hatching rates of Danube salmon (a spring‐spawner). Significantly longer incubation periods of resident and migratory brown trout coincided with relatively low stream substratum quality at the end of the egg incubation. Danube salmon seem to avoid low oxygen concentrations in the hyporheic zone by faster egg development favored by higher water temperatures. Consequently, the prediction of effects of temperature changes and altered stream substratum properties on gravel‐spawning fishes and biological communities should consider the observed species‐specific variances in life‐history strategies to increase conservation success.     

Evidence for non-random spatial positioning of migrating smolts (Salmonidae) in a small lowland stream

1. The ontogenetic development of anadromous salmonids includes downstream emigration of immature individuals from freshwater towards the marine environment. Although this migration of juvenile salmonids (smolts) may be associated with severe mortalities, only limited attention has been paid to the spatial positioning of smolts in small streams. 2. Using a novel approach, this study examined the vertical and horizontal positioning of brown trout and Atlantic salmon smolts while performing downstream migration in a small lowland stream. 3. Pre‐smolts of indigenous and hatchery‐reared (F1) brown trout (Salmo trutta), and two different populations of Atlantic salmon (S. salar), were tagged with passive integrated transponder (PIT) tags and subsequently released upstream of an antenna array consisting of five circular swim‐through PIT antennas. Antennas were positioned in order to determine whether the migrating smolts were bottom or surface oriented, and if they were oriented towards the mid‐channel or the stream bank. 4. During the smolt emigration period, data describing both the detection of the migrating fish and the amount of water passing through the antennas were collected. This was accomplished in order to determine if the fish were performing active positioning behaviour independently of the vertical and horizontal discharge distributions in the stream. 5. The results showed that the smolts migrated in a non‐random spatial pattern independently of the stream discharge distributions. Vertically, the indigenous brown trout and the Atlantic salmon demonstrated a preference for the bottom orientated positions. In contrast, the distribution of the F1 brown trout was not different from the discharge distribution. The latter observation suggests random vertical positioning, which may be indicative of inferior migratory performance. Horizontally, all tested smolt populations strongly preferred the mid‐channel positions. 6. The discharge‐corrected preferences for certain spatial positions suggest that smolt emigration is not entirely a matter of passive displacement in lowland streams. 7. Anthropogenically altered channels may inhibit or delay downstream emigration of smolts resulting in increased mortalities. Given that the smolts in this study actively selected spatial positions in the mid‐channel and near the bottom, it is suggested that deep, mid‐channel furrows may be used to help guide migrating smolts past adverse habitats in lowland streams.     

Forests and the temperature of upland streams in Wales: a modelling exploration of the biological effects

SUMMARY. 1. Daily temperature data from six streams in upland Wales were used to explore the thermal effects of afforestation on stream ecology. The data were linked to published biological models to simulate fish and invertebrate development.
2. Mean daily temperatures in forest streams were lower than those of moorland streams in spring and summer, and higher in winter. These spatial comparisons were supported by the results of experimental bank-side clearance at a forest site, where there was evidence that stream temperatures fell in winter and rose in spring following treatment.
3. Simulations indicated that brown trout (Salmo trutta) could weigh over 30% more by the end of their second growing season in a moorland compared with a forest stream. Several species of insects showed slower simulated egg development at forest sites. For two ephemeropteran species simulated nymphal growth was also retarded, suggesting significant alterations to the life cycle. Two plecopteran species were affected only slightly by the different temperature regimes.
4. Overall, the simulations suggested that afforestation, by reducing summer temperatures, could lead to marked reductions in rates of development of some invertebrates and fish.     

Effect of springtime water temperature on the time of emergence and size of Pteronarcys californica in the Henry's Fork catchment, Idaho, U.S.A.

1. The emergence time of Pteronarcys californica in streams in the Henry's Fork catchment, Idaho, U.S.A. was negatively correlated with mean April water temperature. Emergence was in mid- to late May at sites influenced by groundwater, where April water temperature averaged 7.9 °C. Adults emerged in mid-June in streams receiving run-off from snowmelt (mean April water temperature 5.4 °C). Intermediate emergence times were observed in a regulated section of river where water temperature was influenced, on one bank, by dam release (mean April water temperature 4.5 °C) and, on the other, by a spring-fed tributary stream (mean April water temperature 6.3 °C).
2. During each of the three study years, emergence was earlier on the bank of the regulated section that was warmer during April and May. The mean body length of P. californica exuviae, collected from the warm side of the river, averaged 1.2 mm longer than those collected from the cold side.
3. We tested the effect on emergence of altering springtime water temperature by translocating P. californica in cages from one location to another during April. Individuals moved to sites with higher April water temperature emerged earlier than individuals that remained at the site from which they were collected.     

Estimating the success of natural spawning of salmonids in streams

A new method is described to estimate the survival of salmonids in streams from fertilization of the eggs to the emergence of the fry. The method is compared to other techniques generally used to estimate the egg-to-fry survival: fry traps, excavation of redds, construction of artificial redds and laboratory experiments. Especially designed boxes filled with substratum and freshly fertilized eggs are buried in the stream bed. Survival is determined at three different embryological development stages: eyed stage, hatching and emergence. Boxes are planted inside the stream bed using a special injector, so that substratum alterations are minimal around the boxes. This method was tested successfully in two different studies, one in Switzerland on the brown trout, the other one in Sweden on the sea trout.     

Estimates of survival of stream-dwelling brown trout using

Estimated monthly apparent survival of stream-dwelling brown trout Salmo trutta in south-east Norway was higher in winter than in summer, and lower in Alpine bullhead Cottus poecilopus sites than in allopatric sites. Apparent survival denotes true survival x local site fidelity. Emigration may also explain differences in apparent survival. All brown trout included in this Study were at least 1 year old.     

Downstream displacement of post‐emergent brown trout: effects of development stage and water velocity

Brown trout Salmo trutta were introduced at hatching into distinct sections of two parallel artificial channels, one with a constant low velocity (control) and one with velocity changes (experimental), at such times as to produce 12, 3 and 0 day old fish (age after emergences) when the velocity was changed in the experimental channel. This experimental design was repeated in 2002 and 2003 at comparable dates. Young brown trout were sensitive to an increased water velocity for 5 to 6 days after emergence. Water velocity modified the displacement patterns qualitatively but not quantitatively. Eighty per cent of fish moved downstream at all water velocities. Velocity changes, however, advanced the time by which 80% of the fish had displaced downstream.     

Temporal and spatial segregation of spawning in sympatric populations of Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L.

Based on data from Norwegian streams with sympatric populations of Atlantic salmon and brown trout, it is suggested that temporal segregation is the main mechanism segregating Atlantic salmon and brown trout during spawning. Peak spawning of trout was about 15 days earlier than that of salmon. Physical factors, such as water depth, water velocity and distance from the river banks segregate spawning sites of salmon and trout poorly. Gravel sizes of the redds of salmon and trout were significantly different, though with a considerable overlap, and mean egg depth of salmon and trout were 0.18 and 0.12 m, respectively, probably attributable to the different size of spawners of salmon and trout. None of the temporal or spatial parameters analysed segregate spawners of salmon and trout completely. Species determination of eggs and alevins from the redds showed no interspecific superimposition of redds. It is, therefore, concluded that low survival of hybrids after hatching does not explain the low frequency of hybrids observed in sympatric populations of salmon and trout.     

Interspecific competition between stream-dwelling brown trout and Alpine bullhead

Density and composition of benthic invertebrates and the diet of brown trout Salmo trutta and Alpine bullhead Cottus poecilopus were studied at two sites in one Norwegian stream. The sites were separated by an impassable waterfall, and brown trout density was five to 10 times higher at the upper, allopatric site than downstream where it lived in sympatry with the Alpine bullhead. Benthic invertebrate communities did not differ between sites; however, the size distribution of chironomids and trichopterans were skewed towards lighter individuals at the sympatric site. Diet composition suggested that sympatric brown trout foraged more on invertebrate drift and from the surface than allopatric brown trout. Alpine bullhead diet did not differ significantly from brown trout diet, except that the Alpine bullhead fed on heavier individual prey within a few taxa and did not consume chironomid pupae or surface insects. The collected data support the hypothesis that brown trout living in sympatry with Alpine bullhead feed at locations with higher predation risk, which is a probable explanation for their lower population density.     

The roles of temperature and diapause in the life history of a temperate-zone dragonfly: Argia vivida (Odonata: Coenagrionidae)

Abstract. 1. The life cycle of Argia vivida Hagen generally took longer to complete in the field than was predicted on the basis of the thermal sum accumulated in laboratory rearing.
2. The prediction of a bivoltine life-cycle from geothermal sites with either a constant annual temperature of 26°C or thermal range of 11–31°C was not borne out because the intervention of short-day induced developmental delays in later larval instars extended the life cycle to 1 year.
3. This diapause, which synchronizes adult emergence with favourable summer temperatures, was also present in larvae from sites with annual temperature ranges of 0–33°C and 5–20°C.
4. At these colder sites completion of the life cycle takes 2 and 3 years respectively and dragonflies must be in cold-resistant stages during the winter. A long-day diapause, principally affecting late-instar larvae below a certain size during the summer, achieves this.
5. Large diurnal temperature fluctuations at the 0–33°C site markedly increase the useful thermal energy available to larvae for growth over that predicted by the thermal sum equation.
6. The interaction between the effects of temperatures favourable for growth and day-length-governed diapause, synchronize the emergence of the low-temperature sensitive adult stage of this tropical dragonfly with northern-latitude summers at a variety of habitats.     

Factors Affecting Redd Site Selection, Hatching, and Emergence of Brook Charr, Salvelinus Fontinalis, in an Artificially Enhanced Site

We measured microhabitat characteristics, hatching and emergence success of brook charr, Salvelinus fontinalis, in a series of sites selected and not selected by individuals spawning in an artificially enhanced lake outlet. Differences between the physico-chemistry of surface and interstitial water were small and did not suggest the presence of groundwater seepage. The mean surface water velocity was significantly higher in selected than non-selected sites during the incubation and emergence periods. Differences in interstitial water flow were not detected. Overall, selected substrate was coarser and contained a lower proportion of fine particles than non-selected substrate, as determined by the geometric mean diameter of particles, the proportion of fine particles (<1mm), and the Fredle index. The proportion of fine particles was correlated with sediment loading in incubators. A two-way ANOVA showed no significant effect of sites (selected versus non-selected) but did show a significant effect of the incubation substrate (Astro-turf, selected substrate, non-selected substrate) on both the hatching and emergence success; the percentages of hatching and emergence were significantly higher in Astro-turf than in non-selected substrate, with selected substrate being intermediate. The results of this study suggest that redd site selection by brook charr is based on surface water velocity and substrate characteristics (granulometry and proportion of fine particles) that in turn affect egg survival. It is possible that the lower proportion of fine particles in selected sites (and incubators) is related to their higher water velocity, which could carry away fine particles that reduce the availability of oxygenated water to the embryos. In the same way, higher water velocity could act as a proximate cue in the absence of groundwater seepage or interstitial water flow for individuals to select suitable sites for spawning and egg incubation.     

Temperature-dependent growth and life cycle of Deleatidium (Ephemeroptera: Leptophlebiidae) in two high-country streams in New Zealand

1. Measurements of larval growth rates were used to produce an empirical model for examining factors influencing patterns of size structure and emergence period for populations of the leptophlebiid mayfly Deleatidium ( lillii group) in two high-country streams (South Island, New Zealand). Growth rates were measured in situ by enclosing groups of larvae and natural stream substrata in growth chambers.
2. Multiple regression analysis showed that temperature explained most variation among growth rates in both streams. Growth rates were also significantly related to larval size, although the effect was minor compared with temperature. A significant relationship between larval biomass per chamber and growth rate was shown at only one site.
3. Growth models based on multiple regression equations ( R ² = 0.70–0.84) simulated the extended emergence period and complex size structure observed for populations of Deleatidium in the field. Larvae hatching from eggs deposited before mid-February (austral summer) probably emerge as adults before May (minimum cohort duration ≈ 3 months). Larvae hatching after mid-February probably do not emerge as adults until the following summer because of low growth rates during winter (maximum cohort duration ≈ 11 months).
4. On average, there are probably two generations of Deleatidium per year (bivoltine) at the study sites. The presence of numerous overlapping cohorts throughout summer, however, results in an extended emergence period and complex size structure.     

Temperature-dependent growth and life cycle of Deleatidium (Ephemeroptera: Leptophlebiidae) in two high-country streams in New Zealand

1. Measurements of larval growth rates were used to produce an empirical model for examining factors influencing patterns of size structure and emergence period for populations of the leptophlebiid mayfly Deleatidium ( lillii group) in two high-country streams (South Island, New Zealand). Growth rates were measured in situ by enclosing groups of larvae and natural stream substrata in growth chambers.
2. Multiple regression analysis showed that temperature explained most variation among growth rates in both streams. Growth rates were also significantly related to larval size, although the effect was minor compared with temperature. A significant relationship between larval biomass per chamber and growth rate was shown at only one site.
3. Growth models based on multiple regression equations ( R ² = 0.70–0.84) simulated the extended emergence period and complex size structure observed for populations of Deleatidium in the field. Larvae hatching from eggs deposited before mid-February (austral summer) probably emerge as adults before May (minimum cohort duration ≈ 3 months). Larvae hatching after mid-February probably do not emerge as adults until the following summer because of low growth rates during winter (maximum cohort duration ≈ 11 months).
4. On average, there are probably two generations of Deleatidium per year (bivoltine) at the study sites. The presence of numerous overlapping cohorts throughout summer, however, results in an extended emergence period and complex size structure.     

Incubation temperatures, sex ratios and sex determination in a population of Nile crocodiles (Crocodylus niloticus)

A demographic study of the Nile crocodile Crocodylus niloticus at Lake Ngezi, Zimbabwe, revealed that females predominated in all size classes and among embryos. The sex of C. niloticus was shown to be determined by the temperature of egg incubation in constant temperature laboratory experiments. At 31 °C and below only females were produced. The threshold temperature for maleness was between 31 ° and 34 °C, but appeared to vary between clutches. The duration of the incubation period varied with temperature and was 110 days at 28 °C, falling to 85 days at 34 °C. Incubation temperature affected hatchling length, but not mass. Hatchlings from incubation at 34 °C were shorter on average than those from incubation at 28 °C and 31 °C, but by three months had outgrown them. There was no sex-related difference in length in a random sample of 200 two-year-old C. niloticus on a crocodile farm. Mean temperatures in wild nests were consistently lower than 31 °C and therefore the male threshold as determined in the laboratory. Embryonic development was slow and hatching success poor. The shallowest eggs in a nest had higher mean temperatures and more advanced embryos than the deepest eggs. They also experienced daily temperature fluctuations of up to 10 °C during which the maximum occasionally rose to 35 °C. Constant temperature incubation was not a good model of field conditions, but the correlation between nest temperatures and embryonic sex is consistent with temperature-dependent sex determination in the wild.