Total population density during the first year of life as a major determinant of lifetime body‐length trajectory in marble trout

Abstract – The conditions experienced early in life can strongly influence life‐history trajectories in a variety of animal species. Here, we use data from four isolated populations of the endangered stream‐dwelling salmonid marble trout (Salmo marmoratus Cuvier 1817) living in the Soca and Idrijca river basins (Slovenia) to explore the influence of the total density experienced during and after the first year of life by marble trout year‐classes on body length of marble trout through the lifetime. Analyses were performed by pooling together the stream‐specific datasets to cover a wider range of densities. Mean body length of marble trout year‐classes through the lifetime (from age 1+ to 5+) was negatively related to total density of marble trout during the first year of life. The relationship between density during the first growth period and body length through the lifetime was well described by negative power curves. Total population density after the first year of life was not correlated with body length, thus suggesting that body growth trajectories are heavily determined early in life. Given size‐dependent sexual maturity and egg production in marble trout, the relationship between density early in life and lifetime individual growth may have strong implications in terms of population dynamics and regulation of population size.     

Dietary niche expansion and niche shift in native marble trout (Salmo marmoratus) living in sympatry with introduced rainbow trout (Oncorhynchus mykiss)

The non‐native rainbow trout (Oncorhynchus mykiss) has been introduced worldwide for angling purposes and has established self‐reproducing populations in many parts of the world. Introduced rainbow trout often have negative effects on the native salmonid species, ranging from decrease abundance, growth and survival, to their local extinction. Assessing the effects of introduced rainbow trout on the native species is thus crucial to better set up conservation programmes. In this study, we investigated the effects of non‐native rainbow trout on the diet of native marble trout (Salmo marmoratus) living in the Idrijca River (Slovenia). An impassable waterfall separates the stream in two sectors only a few hundred metres apart: a downstream sector (treatment) in which marble trout live in sympatry (MTs) with rainbow trout (RTs) and an upstream sector (control) in which marble trout live in allopatry (MTa). Specifically, we investigated using stable isotopes the effects of rainbow trout on dietary niche, diet composition, body condition, and lipid content of marble trout. We found dietary niche expansion and niche shift in marble trout living in sympatry with rainbow trout. Compared to MTa, MTs had higher piscivory rate and showed higher body condition and prereproduction lipid content. Our results indicate that the presence of rainbow trout did not have negative effects on marble trout diet and condition and that changes in dietary niche of marble trout are likely to be an adaptive response to the presence of rainbow trout, and further research is needed to better understand.     

Early survival of marble trout Salmo marmoratus: evidence for density dependence?

Abstract –  The role of endogenous and exogenous factors in regulating population dynamics of freshwater salmonids is still a matter of debate. The aim of the present work was to assess the relative importance of density-dependent and -independent factors in determining the survival of marble trout ( Salmo marmoratus ) yearlings in two populations living in Slovenian streams (Zakojska and Gorska). The investigation was performed by combining a classical life table analysis with Monte Carlo simulation. Size-dependent fecundity was estimated by stripping wild adults in the fish farm. A significant positive relationship was found between length of marble trout females and the number of eggs produced. Egg density was the major determinant of survival from eggs to age 1+ ( σ ₀) in both streams. Residuals of the relationship between σ ₀ and egg density were positively correlated with rainfall only in Zakojska, probably because, within a certain range, more intense rainfalls increases stream flow and, consequently, suitable habitat for trout. Our study shows how density-dependent and environmental factors can interact to determine the survival of marble trout during the first year of life.     

The effect of competition among three salmonids on dominance and growth during the juvenile life stage

Although non‐native species can sometimes threaten the value of ecosystem services, their presence can contribute to the benefits derived from the environment. In the Great Lakes, non‐native brown trout (Salmo trutta) and rainbow trout (Oncorhynchus mykiss) support substantial recreational fisheries. With current efforts underway to restore once‐native Atlantic salmon (Salmo salar) to Lake Ontario, there is some concern that Atlantic salmon will impede non‐native contributions to the recreational fishery because Atlantic salmon exhibit niche overlap with brown trout and rainbow trout, particularly during the juvenile life stage. We therefore examined competition and growth of juvenile Atlantic salmon, brown trout and rainbow trout in semi‐natural streams. We found that brown trout were the most dominant and had the greatest growth rate regardless of what other species were present. Rainbow trout were more dominant than Atlantic salmon and consumed the most food of the three species. However, in the presence of brown trout, rainbow trout fed less frequently and exhibited negative growth as compared to when the rainbow trout were present with only Atlantic salmon. These data suggest that, outside of density‐dependent effects, Atlantic salmon will not impact stream production of brown trout and rainbow trout.     

Changes in the density and growth of brown trout (Salmo trutta) after intensive removal of sympatric Arctic charr (Salvelinus alpinus) in the sub‐Arctic lake Møkkelandsvatn,Norway

This study tests the basic hypothesis that the removal of charr, Salvelinus alpinus (L.), would cause an increase in both the growth and density of a sympatric trout population, Salmo trutta L. The charr population was characterised by slow‐growing individuals, with a high proportion of mature fish, that is typical for so‐called overpopulated populations. A total of 31,000 charr was removed from the lake in the period 1990–1992, and the density of younger trout (1+, 2+), but not older trout (3+, 4+), increased. The growth of older trout (3+, 4+) increased, but the evidence for similar growth increases of younger trout (1+, 2+) was limited. From 1989 to 1990, the proportion of trout increased from 30 to only 40% of the total catch, but from 1991 to 1994, it was significantly higher (60–80%) than that of charr. Total trout biomass increased to a maximum in 1992 and then decreased so that the biomass of 1994 was nearly similar to that of 1989, that is before the start of the charr removal. Back‐calculated lengths of trout from otoliths showed that 2+ and 3+ trout caught in the pelagic were growing consistently faster over previous years than those caught in the littoral, while this was not the case for the 4+ fish. Therefore, the hypothesis was partially supported; the growth rate of trout increased (age groups 1+ to 4+), while the density of juvenile trout (1+, 2+), but not the older trout (3+, 4+), increased after the removal of charr.     

Demographic changes following mechanical removal of exotic brown trout in an Intermountain West (USA), high‐elevation stream

Exotic species present a great threat to native fish conservation; however, eradicating exotics is expensive and often impractical. Mechanical removal can be ineffective for eradication, but nonetheless may increase management effectiveness by identifying portions of a watershed that are strong sources of exotics. We used mechanical removal to understand processes driving exotic brown trout (Salmo trutta) populations in the Logan River, Utah. Our goals were to: (i) evaluate the demographic response of brown trout to mechanical removal, (ii) identify sources of brown trout recruitment at a watershed scale and (iii) evaluate whether mechanical removal can reduce brown trout densities. We removed brown trout from 2 km of the Logan River (4174 fish), and 5.6 km of Right Hand Fork (RHF, 15,245 fish), a low‐elevation tributary, using single‐pass electrofishing. We compared fish abundance and size distributions prior to, and after 2 years of mechanical removal. In the Logan River, immigration to the removal reach and high natural variability in fish abundances limited the response to mechanical removal. In contrast, mechanical removal in RHF resulted in a strong recruitment pulse, shifting the size distribution towards smaller fish. These results suggest that, before removal, density‐dependent mortality or emigration of juvenile fish stabilised adult populations and may have provided a source of juveniles to the main stem. Overall, in sites demonstrating strong density‐dependent population regulation, or near sources of exotics, short‐term mechanical removal has limited effects on brown trout populations but may help identify factors governing populations and inform large‐scale management of exotic species.     

Recruitment of the threatened mussel Margaritifera margaritifera in relation to mussel population size,mussel density and host density

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Addition of structural complexity – contrasting effect on juvenile brown trout in a natural stream

In a field experiment, we examined the effects of structural complexity in the form of added artificial plastic plants and shredded plastic bags on growth and abundance of juvenile brown trout (Salmo trutta). Just after emergence, the added complexity had a positive effect on the density, biomass and condition factor of young‐of‐the‐year (0+) brown trout. This difference in density was not present six weeks later. In contrast, both young‐of‐the‐year and older brown trout generally tended to be larger in the simple habitat. Hence, our data suggest that increased complexity initially is beneficial for young‐of‐the‐year individuals probably due to lower risk of predation and increased densities of prey. However, as density increases in the complex environment, it may induce negative density‐dependent effects, here reflected in smaller sized fish in the complex environment. This might force fish to redistribute to habitats with lower densities of conspecifics as they grow larger. We propose that habitat complexity can increase survival of yearlings in early phases and thereby also affect the overall population structure of brown trout in natural streams.     

Metabolic characteristics of early life history stages of native marble trout (Salmo marmoratus) and introduced brown trout (Salmo trutta) and their hybrids in the Soča River

Metabolic potential and respiration rate have been measured in the early life stages of native marble trout Salmo marmoratus (MT), introduced brown trout Salmo trutta (BT) and their hybrids (MH = female of MT × male of BT and BH = female of BT × male of MT) to examine the differences in their metabolic characteristics. Metabolic potential and respiration rate were lowest in each developmental stage in MT, followed by MH, BT and BH. The ratio between metabolic potential and respiration rate (ETS/R) in larvae was also lowest for MT (1.0) and MH (1.1), followed by BT (1.4) and BH (1.6), indicating more intensive exploitation of metabolic potential for actual respiration in MT (100%) and MH (~90%) than in BT (~70%) and BH (~60%). Lower ETS/R means a disadvantage in the case of enhanced metabolic activity due to insufficient enzyme machinery needed for energy production. Thus, potential and actual metabolic activity and ETS/R ratio could account for the different competitive abilities among pure trout species and their hybrids in a particular environment. This is the first report on biochemical and physiological differences between pure MT and their hybrids. The combination of metabolic and respiration rate measurements appears to be a useful tool for estimating the metabolic properties in the early developmental stages of fish.     

Effects of stocking on the genetic structure of brown trout,Salmo trutta,in Central Europe inferred from mitochondrial and nuclear DNA markers

Abstract Stocking has had a considerable effect on wild brown trout, Salmo trutta L., populations throughout Europe. To elucidate this impact and to outline further management strategies, the genetic structure of 25 wild populations and five hatchery stocks from Czech Republic and Slovakia were analysed using mitochondrial (control region) and nuclear DNA (microsatellites, LDH‐C1*) markers. Stocking practices have caused massive hybridisation between the Atlantic and Danube brown trout strains in the central Danube basin and have lead to a loss of among‐population divergence in Slovakia and the eastern part of Czech Republic. Comparison with studies from neighbouring countries revealed substantial differences in haplotype, allele frequencies and genetic diversity across Central Europe. Differences in stocking management and origin of breeding stocks appear to be crucial factors for the spatial variability of the genetic structure of brown trout.     

Effects of intra‐ and interspecific competition and hydropeaking on growth of juvenile Atlantic salmon (Salmo salar)

The effects of hydropeaking and intra‐ and interspecific competition on the growth performance (growth in length, mass and lipid content) of juvenile Atlantic salmon Salmo salar and brown trout Salmo trutta were studied in six experimental channels (three experiencing hydropeaking and three controls with a stable discharge of water). Changes in the water‐covered area in the hydropeaking channels were small to avoid fish stranding. Each channel was divided into three similar‐sized sections and stocked with either low or high density of Atlantic salmon, or a mix of Atlantic salmon and brown trout, with the density of the latter equalling the high‐density treatment of Atlantic salmon. A marked effect of competition was visible as salmon in the low‐density treatment were significantly larger (27–33%) and had a higher mass (30–38%) than salmon in both the high‐density salmon treatment and the high‐density salmon and trout treatment. Hydropeaking had only minor and insignificant effects on the growth performance: overall final length, mass and body lipid content in the salmon experiencing hydropeaking differed by ?9%, ?7% and +2% compared with controls. Furthermore, there was no indication that the competitive regime influenced hydropeaking effects. The increase in both intra‐ and interspecific competition among the juvenile salmon had a pronounced and significant effect on growth. Our study adds to the growing evidence that energetic consequences of hydropeaking are likely to be small for Atlantic salmon and that stress and mortality associated with stranding represent the main source of population impact.     

Parallelism in thermal growth response in otoliths and scales of brown trout (Salmo trutta L.) from alpine lakes independent of genetic background

Low density in natural populations of salmonids has predominantly been managed by stocking of non‐native conspecifics. Due partly to domestication, introduced non‐native fish may be maladapted under natural conditions. Interbreeding between introduced and wild individuals may therefore impair local adaptation and potentially population viability. Brown trout (Salmo trutta L.) from three headwaters (with stocked fish) and three interconnected lakes (with native fish) on the Hardangervidda mountain plateau, southern Norway, were tested for differences in thermal effects on scale and otolith growth. Otolith and scale annuli widths from immature brown trout showed positive correlation with mean annual summer temperature for all six sampled populations. In mature individuals, a similar positive thermal correlation was evident for the otoliths only. Interannuli width measurements from scales indicate a halt in somatic growth for brown trout in this alpine environment when reaching ages between 7 and 9 winters, coinciding with age at maturity. Our study indicates that otolith growth follows summer temperature even when individuals do not respond with somatic growth in these populations and that introduced brown trout and introgressed populations have similar thermal growth responses. Due to the continued otolith growth after stagnation in somatic growth and the impact of fluctuations in summer temperature, the utilisation of otolith annuli widths for back calculation of length at age should be treated with caution.     

Relationship between sea lice levels on sea trout and fish farm activity in western Scotland

The relationship between aquaculture and infestations of sea lice on sea trout, Salmo trutta L., is controversial. Here, the association between sea lice infestations on wild sea trout and characteristics of local Atlantic salmon, Salmo salar L., farms were investigated using data collected on the Scottish west coast. The proportion of sea trout with louse burdens above a critical level was positively related to the fork length of the sea trout and the mean weight of salmon on the nearest fish farm, and negatively related to the distance to that farm. The distance to the nearest fish farm did not influence the probability of infestations above the critical level beyond 31 km although there was considerable uncertainty around this cut‐off distance (95% limits: 13–149 km). The results support a link between Atlantic salmon farms and sea lice burdens on sea trout in the west of Scotland and provide the type of information required for marine spatial planning.     

Functional response and size‐dependent foraging on aquatic and terrestrial prey by brown trout (Salmo trutta L.)

Gustafsson P, Bergman E, Greenberg LA. Functional response and size‐dependent foraging on aquatic and terrestrial prey by brown trout (Salmo trutta L.).Ecology of Freshwater Fish 2010: 19: 170–177. © 2010 John Wiley & Sons A/S Abstract – Terrestrial invertebrate subsidies are believed to be important energy sources for drift‐feeding salmonids. Despite this, size‐specific use of and efficiency in procuring this resource have not been studied to any great extent. Therefore, we measured the functional responses of three size classes of wild brown trout Salmo trutta (0+, 1+ and ≥2+) when fed either benthic‐ (Gammarus sp.) or surface‐drifting prey (Musca domestica) in laboratory experiments. To test for size‐specific prey preferences, both benthic and surface prey were presented simultaneously by presenting the fish with a constant density of benthic prey and a variable density of surface prey. The results showed that the functional response of 0+ trout differed significantly from the larger size classes, with 0+ fish having the lowest capture rates. Capture rates did not differ significantly between prey types. In experiments when both prey items were presented simultaneously, capture rate differed significantly between size classes, with larger trout having higher capture rates than smaller trout. However, capture rates within each size class did not change with prey density or prey composition. The two‐prey experiments also showed that 1+ trout ate significantly more surface‐drifting prey than 0+ trout. In contrast, there was no difference between 0+ and ≥2+ trout. Analyses of the vertical position of the fish in the water column corroborated size‐specific foraging results: larger trout remained in the upper part of the water column between attacks on surface prey more often than smaller trout, which tended to seek refuge at the bottom between attacks. These size‐specific differences in foraging and vertical position suggest that larger trout may be able to use surface‐drifting prey to a greater extent than smaller conspecifics.     

Direct and indirect effects of riparian canopy on juvenile Atlantic salmon,Salmo salar,and brown trout,Salmo trutta,in south‐west Ireland

Abstract Dense riparian tree canopy is generally found to have a negative effect on salmonid populations. Canopy can affect fish both directly via trophic impacts and its effects as cover, and indirectly via its effects on the distribution of instream vegetation. This study examined the impact of riparian canopy on the ecology of juvenile Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L., in south‐west Ireland. Riparian canopy reduced the density, length and gut contents of juvenile salmon, but not brown trout. The negative relationship between canopy cover and fish size and feeding was strongly influenced by the abundance of instream macrophytes, which in turn varied as a function of catchment water chemistry. As a management strategy, the reduction of dense riparian canopy is unlikely to have the same effect on juvenile salmonids on all streams within an ecoregion because of differences in catchment‐wide factors.     

Genetic diversity within sea trout population from an intensively stocked southern Baltic river,based on microsatellite DNA analysis

Sea trout, Salmo trutta L., populations in the Slupia River basin have been affected by mass stocking with smolts and fry. This work is focused on a small‐scale differentiation in sea trout populations from one basin with a strong emphasis on the relationship between wild and hatchery origin fish. A total of 216 sea trout were genotyped at 10 microsatellite loci. Global F_ST obtained by amova was low at 0.0165. Pairwise F_ST were significant for all tests except wild and stocked adults. The highest pairwise difference was found between the hatchery sample and Kwacza (F_ST = 0.038). Analysis of the genetic structure revealed micro‐geographical differentiation with four subpopulations. The quality of the artificial spawning was found not to be adequate with a high risk of adverse effects to the whole population. All future stocking actions in the basin should consider the existing population structures.     

Seasonal movement,growth and survival of brook trout in sympatry with brown trout in Midwestern US streams

Seasonal patterns in growth, survival and movement of brook trout Salvelinus fontinalis were monitored in two southeastern Minnesota streams divided into study reaches based on brown trout Salmo trutta abundance. We estimated survival and movement while testing for effects of stream reach and time using a multistrata Cormack–Jolly–Seber model in Program MARK. Multistrata models were analysed for three age groups (age‐0, age‐1 and age‐2+) to estimate apparent survival, capture probability and movement. Survival varied by time period, but not brown trout abundance and was lower during flood events. Age‐0 brook trout emigrated from reaches with low brown trout abundance, whereas adult brook trout emigrated from downstream brown trout‐dominated reaches. Growth was highest in spring and summer and did not differ across streams or reaches for the youngest age classes. For age‐2+ brook trout, however, growth was lower in reaches where brown trout were abundant. Interspecific interactions can be age or size dependent; our results show evidence for adult interactions, but not for age‐0. Our results suggest that brook trout can be limited by both environmental and brown trout interactions that can vary by season and life stage.     

Long-term variation in the population structure of Arctic charr, Salvelinus alpinus, and brown trout, Salmo trutta

The effects of induced water level fluctuations and introduction of the mysid Mysis relicta Lovén on population structure of brown trout, Salmo trutta L., and Arctic charr, Salvelinus alpinus (L.), were studied during 1953–1995 in Limingen hydroelectric reservoir, Norway. The main response was a marked reduction in catch‐per‐unit‐effort (CPUE) for trout and charr, probably caused by reduced recruitment following increased variation in water level. For both species, mean length decreased until 1967 and increased thereafter, whereas mean mass‐at‐length increased for the whole period. Both length and mass‐at‐length were negatively correlated with CPUE. The increases in mean length and mass‐at‐length were probably because of reduced competition following the reduced recruitment. Mysis relicta has become an important food item for charr but not for brown trout, but the increases in mean length and mass‐at‐length of charr started prior to the appearance of M. relicta in the charr diet.     

The density dilemma: limitations on juvenile production in threatened salmon populations

Density‐dependent processes have repeatedly been shown to have a central role in salmonid population dynamics, but are often assumed to be negligible for populations at low abundances relative to historical records. Density dependence has been observed in overall spring/summer Snake River Chinook salmon Oncorhynchus tshawytscha production, but it is not clear how patterns observed at the aggregate level relate to individual populations within the basin. We used a Bayesian hierarchical modelling approach to explore the degree of density dependence in juvenile production for nine Idaho populations. Our results indicate that density dependence is ubiquitous, although its strength varies between populations. We also investigated the processes driving the population‐level pattern and found density‐dependent growth and mortality present for both common life‐history strategies, but no evidence of density‐dependent movement. Overwinter mortality, spatial clustering of redds and limited resource availability were identified as potentially important limiting factors contributing to density dependence. The ubiquity of density dependence for these threatened populations is alarming as stability at present low abundance levels suggests recovery may be difficult without major changes. We conclude that density dependence at the population level is common and must be considered in demographic analysis and management.     

Potential population and assemblage influences of non‐native trout on native nongame fish in Nebraska headwater streams

Non‐native trout are currently stocked to support recreational fisheries in headwater streams throughout Nebraska. The influence of non‐native trout introductions on native fish populations and their role in structuring fish assemblages in these systems is unknown. The objectives of this study were to determine (i) if the size structure or relative abundance of native fish differs in the presence and absence of non‐native trout, (ii) if native fish‐assemblage structure differs in the presence and absence of non‐native trout and (iii) if native fish‐assemblage structure differs across a gradient in abundances of non‐native trout. Longnose dace Rhinichthys cataractae were larger in the presence of brown trout Salmo trutta and smaller in the presence of rainbow trout Oncorhynchus mykiss compared to sites without trout. There was also a greater proportion of larger white suckers Catostomus commersonii in the presence of brown trout. Creek chub Semotilus atromaculatus and fathead minnow Pimephales promelas size structures were similar in the presence and absence of trout. Relative abundances of longnose dace, white sucker, creek chub and fathead minnow were similar in the presence and absence of trout, but there was greater distinction in native fish‐assemblage structure between sites with trout compared to sites without trout as trout abundances increased. These results suggest increased risk to native fish assemblages in sites with high abundances of trout. However, more research is needed to determine the role of non‐native trout in structuring native fish assemblages in streams, and the mechanisms through which introduced trout may influence native fish populations.