The influence of groundwater withdrawal and land use changes on brook charr (<Emphasis Type="Italic">Salvelinus fontinalis</Emphasis>) thermal habitat in two coldwater tributaries in Michigan,U.S.A.

Brook charr (Salvelinus fontinalis) is a sentinel fish species which requires clean, cold water habitats. As such, many jurisdictions in the United States where brook charr are present use this species as an indicator of ecosystem health. In Michigan, groundwater-dominated streams are currently being impacted by increased groundwater withdrawal and land use/land cover changes which alter stream temperatures and their flow, and thus have the potential to significantly influence brook charr production and behavior. We quantified the influence of groundwater withdrawal and land use alteration on thermal habitat availability for brook charr using a groundwater modeling tool that estimated changes in baseflow to a stream segment based on changes to the groundwater system due to groundwater withdrawal and changes in rates of recharge related to landscape changes within a watershed. Projected stream temperature changes were calculated using a stream temperature modeling tool and compared to the range of temperature preferenda for brook charr in order to evaluate the potential impact of policy decisions regarding water extraction and land use/cover changes. The models predicted relatively small changes in both stream baseflow and consequently, stream temperature, with increased groundwater withdrawal rates. Land use/land cover alterations which we analyzed were shown to either mitigate or enhance the loss of brook charr thermal habitat as a result of groundwater withdrawal, depending on its relationship to recharge dynamics. This study emphasizes the importance of collaboration between water, land, and fisheries managers to ensure brook charr population viability, productivity, and sustainability in the face of environmental change, increasing water use and development in the watershed.     

Genetic Diversity and Hybridisation between Native and Introduced Salmonidae Fishes in a Swedish Alpine Lake

Understanding the processes underlying diversification can aid in formulating appropriate conservation management plans that help maintain the evolutionary potential of taxa, particularly under human-induced activities and climate change. Here we assessed the microsatellite genetic diversity and structure of three salmonid species, two native (Arctic charr, Salvelinus alpinus and brown trout, Salmo trutta) and one introduced (brook charr, Salvelinus fontinalis), from an alpine lake in sub-arctic Sweden, Lake Ånn. The genetic diversity of the three species was similar and sufficiently high from a conservation genetics perspective: corrected total heterozygosity, H’_T = 0.54, 0.66, 0.60 and allelic richness, A_R = 4.93, 5.53 and 5.26 for Arctic charr, brown trout and brook charr, respectively. There were indications of elevated inbreeding coefficients in brown trout (G_IS = 0.144) and brook charr (G_IS = 0.129) although sibling relationships were likely a confounding factor, as a high proportion of siblings were observed in all species within and among sampling locations. Overall genetic structure differed between species, Fst = 0.01, 0.02 and 0.04 in Arctic charr, brown trout and brook charr respectively, and there was differentiation at only a few specific locations. There was clear evidence of hybridisation between the native Arctic charr and the introduced brook charr, with 6% of individuals being hybrids, all of which were sampled in tributary streams. The ecological and evolutionary consequences of the observed hybridisation are priorities for further research and the conservation of the evolutionary potential of native salmonid species.     

Effects of Large Woody Debris Addition on Stream Habitat and Brook Trout Populations in Appalachian Streams

Large woody debris (LWD) was added to eight streams in the central Appalachians of West Virginia to determine if stream habitat could be enhanced and brook trout (Salvelinus fontinalis) populations increased. Brook trout populations were assessed one year prior to habitat manipulation and 3 years post-habitat manipulation. LWD was added by felling approximately 15 trees per 300 m stream reach. Four of the streams had LWD added to one 300 m reach with 300 m unmanipulated reaches upstream and downstream of the manipulated reach to observe within-stream effects of LWD additions on brook trout density. The remaining four streams had LWD added to three 300 m reaches and these streams were compared to those with only a single 300 m manipulated reach to observe the effects of the extent of habitat manipulation on brook trout density. New pools were formed by the addition of LWD, but overall pool area did not increase significantly in reaches where LWD was added. The relatively high gradient and coarse substrate of these streams may have precluded the added LWD from having a significant influence on stream channel morphology and habitat complexity. No pools were formed in the highest gradient stream, while the stream with the most pools formed had the lowest gradient. Brook trout populations fluctuated following habitat manipulations, and there was no overall effect of the LWD additions on within-stream variability in brook trout density. When there were significant differences among-streams with different extents of LWD additions, those streams receiving LWD additions over a large extent had the greatest brook trout densities. The full potential of added LWD to change stream habitat and influence on brook trout populations may take more time to develop than the 3 years post-manipulation period of this study.     

Influence of spring and summer water temperature on brook charr, Salvelinus fontinalis, growth and age structure in the Ford River, Michigan

Synopsis The influence of late spring and summer water temperatures on brook charr, Salvelinus fontinalis, growth and age structure was evaluated from 1984 to 1991 in the Ford River, Michigan. Temperature was monitored and brook charr sampled for vital statistics from late May through September using fyke nets and weirs at four locations within a 25.8 km section of stream. Scale analysis was used to determine captured brook charr age, past length at age and relative annual growth rates. Late spring and summer water temperature patterns varied between years with the greatest variability occurring in May and June. Age and size structure also varied between years and was significantly related to temperature. Years with cooler late spring and summer temperature patterns were dominated by older (age 2 and 3), larger brook charr, while years with warmer spring and summer temperature patterns were dominated by younger (age 1), smaller brook charr. Spring and summer temperature did not appear to have a significant effect on the growth of age 0 or age 1 brook charr. However, temperature was negatively related to brook charr growth from age 2 on. As spring and summer water temperatures are critical to brook charr growth and survival, it is important that a streams thermal regime be considered when establishing management goals for this species.     

Does life-history variability in salmonids affect habitat use by juveniles? A comparison among streams open and closed to anadromy

1. Migratory and resident forms of salmonids coexist in many river systems. Although such coexistence is widespread, little is known about its ecological basis and no studies have compared the habitat use of premigratory juveniles and residents. 2. We employed a comparative approach to explore the differential habitat use of juvenile anadromous and resident brook trout. This required the investigation of habitat use in streams closed to anadromy, containing only resident brook trout Salvelinus fontinalis ('resident-only' streams) and streams open to anadromy, containing coexisting Atlantic salmon Salmo salar and anadromous and resident brook trout ('migrant-resident' streams). 3. We demonstrate that fast habitats (riffles) are occupied more frequently in streams with migratory brook trout relative to riffle habitats of streams with only resident brook trout. In contrast, occupation of slow current velocities (pools) was observed in both migrant-resident and resident-only streams as both stream types contain resident brook trout. The net effect is a wider distribution of occupied habitats (pool and riffles) in migrant-resident streams relative to resident-only streams, resulting in few, if any, unused habitats. 4. These results are consistent with previously reported bioenergetic, morphological and stable isotope differences observed between anadromous and resident brook trout. 5. Our findings suggest that a link exists between juvenile habitat use, metabolic costs and life-history strategies.     

Shifting thermal regimes influence competitive feeding and aggression dynamics of brook trout (Salvelinus fontinalis) and creek chub (Semotilus atromaculatus)

The natural distributions of freshwater fish species are limited by their thermal tolerances via physiological constraints and increased interspecific competition as temperatures shift toward the thermal optima of other syntopic species. Species may mediate stress from temperature change physiologically, behaviorally, or both; but these changes may compromise competitive advantages through effects on feeding and social behavior. In the Appalachian Mountains of North America, creek chub (Semotilus atromaculatus) are found in warm‐water and cold‐water streams and overlap in range with brook trout (Salvelinus fontinalis) across lower thermal maxima, where they compete for food and space. As stream temperatures continue to increase due to climate change, brook trout are under increasing thermal stress which may negatively affect their ability to compete with creek chub. To examine the influences of temperature on competitive interactions between these species, we observed feeding behavior, aggression, and habitat use differences at three temperatures approaching brook trout thermal maxima (18°C, 20°C, and 22°C) among dyad pairs for all combinations of species in experimental flow‐through tanks. We also examined feeding and habitat use of both species under solitary conditions. We found as temperature increased, feeding and aggression of brook trout were significantly reduced in the presence of creek chub. Creek chub pairs were more likely to occupy benthic areas and refugia while brook trout pairs used surface water more. Space use patterns significantly changed by pairing treatment. Aggression and space use shifts allowed increased exploitative and interference competition from creek chub when paired with brook trout that was not present in conspecific pairs. The decreased dominance of a top predator may lead to diverse impacts on stream community dynamics with implications for the future range restriction of brook trout and demonstrate possible mechanisms to facilitate competitive advantages of warm water generalist species under thermal stress.     

A historical perspective on drift foraging models for stream salmonids

Thirty years ago, Fausch (Can J Zool 62:441–451, 1984) proposed a simple model of optimal positions for drift-feeding salmonids in streams, whereby fish maximize their net energy intake (NEI) by selecting focal points in low water velocity near faster currents that deliver abundant drifting invertebrates. The theory was based on earlier observations in artificial and natural streams describing characteristics of salmonid positions and a conceptual model by Chapman (Am Nat 100:345–357, 1966). A test of this simple drift foraging model in a laboratory stream showed that the growth rate of juvenile trout and salmon increased with NEI, and that the rank of NEI at positions held by coho salmon (Oncorhynchus kisutch) correlated nearly perfectly with their rank in the dominance hierarchy. Fausch (1984) inferred from these findings that positions that optimize NEI, within the constraints of the dominance hierarchy, are the resource for which these stream salmonids compete. In turn, the model was used to test the effects of interspecific competition by coho salmon on the foraging positions held by brook trout (Salvelinus fontinalis) and brown trout (Salmo trutta), and these results were used to infer potential effects of the introduced salmon on resident trout in Lake Michigan tributaries. Though the goals for this model were originally modest, it was tested in the field and further refined by Hughes and Dill (Can J Fish Aquat Sci 47:2039–2048, 1990) and others. During the last 20 years, the general theory has been incorporated into other models, which have been applied widely to analyze salmonid distribution and abundance in streams and rivers and used for management and restoration of habitat and flow regimes to benefit these fishes.     

The effects of introduced salmonids on two native stream‐dwelling salmonids through interspecific competition

Using an artificial stream, habitat use by two sympatric native salmonids in the presence and absence of introduced salmonid species was investigated experimentally. When only native white‐spotted charr Salvelinus leucomaenis and masu salmon Oncorhynchus masou were sympatric, they occupied different microhabitats. In the presence of introduced brown trout Salmo trutta or rainbow trout Oncorhynchus mykiss , however, white‐spotted charr and masu salmon were observed to use a similar habitat and interspecific competition between white‐spotted charr and masu salmon was initiated. The study suggested that the coexistence of native salmonids was negatively affected through interspecific competition between native and introduced salmonids.     

Hybridization between native white-spotted charr and nonnative brook trout in the upper Sorachi River,Hokkaido, Japan

Invasion status and impacts of nonnative brook trout (Salvelinus fontinalis) in a Hokkaido stream were investigated with field surveys and genetic analyses. Nonnative brook trout was detected in nine (41 %) of the 22 sampled reaches in three tributaries of the Sorachi River, Hokkaido, Japan. Based on the external pigmentation, twelve putative hybrids between brook trout and native white-spotted charr (Salvelinus leucomaenis) were collected in two reaches. Microsatellite and mitochondrial DNA data established that 58% of these hybrids were first-generation (F₁) progenies between male brook trout and female white-spotted charr. Our results suggest potential negative impacts of nonnative brook trout on native charr populations in Hokkaido through interspecific interactions.     

Diel Habitat Partitioning by Bull Charr and Cutthroat Trout During Fall and Winter in Rocky Mountain Streams

We used underwater observation to determine diel habitat partitioning between bull charr, Salvelinus confluentus, and cutthroat trout, Oncorhynchus clarki, during fall and winter (0.1–8.3°C) in two Rocky Mountain streams that differed in habitat availability. The majority (>70%) of both species emerged from concealment cover at night, though bull charr exhibited a greater tendency for nocturnal behavior than cutthroat trout. Differences in day and night counts were most pronounced at temperatures <3°C, when very few fish of either species were observed in the water column during the day, but both species were common at night. Both species used concealment cover of large woody debris and boulder substrate crevices in deep pools during the day. At night, fish emerged from cover and habitat use shifted to shallow water with low cover. Microhabitat partitioning among species and size classes occurred at night, cutthroat trout moving into shallower, faster water that was farther from cover compared to bull charr. Smaller fish of both species occupied focal positions in slower, shallower water closer to the substrate than larger fish. Large, mixed-species aggregations also were common in beaver ponds both day and night. High variation in diel and site-specific winter habitat use suggests the need for caution in developing habitat suitability criteria for salmonids based solely on daytime observations or on observations from a few sites. Our results support the need to incorporate nocturnal habitat use and partitioning in studies of salmonid ecology.     

Global warming threatens the persistence of Mediterranean brown trout

Current climate change exacerbates the environmental restrictions on temperate species inhabiting low latitude edges of their geographical ranges. We examined how temperature variations due to current and future climate change are likely to affect populations’ persistence of stream‐dwelling brown trout Salmo trutta at the vulnerable southern periphery of its range. Analysis of 33 years of air temperature data (1975–2007) by time‐series models indicated a significant upward trend and a pronounced shift in air temperature around 1986‐1987. This warming is associated with an ongoing population decline of brown trout, most likely caused by a loss of suitable thermal habitat in lower latitudes since the 1980s. Population decrease may not be attributed to physical habitat modification or angler pressure, as carrying capacity remained stable and populations were not overexploited. We developed regional temperature models, which predicted that unsuitable thermal habitat for brown trout increased by 93% when comparing climate conditions between 1975–1986 and 1993–2004. Predictions from climate envelope models showed that current climate change may be rendering unsuitable 12% of suitable thermal habitat each decade, resulting in an overall population decrease in the lower reaches of around 6% per year. Furthermore, brown trout catches markedly decreased 20% per year. Projections of thermal habitat loss under the ecologically friendly B2 SRES scenario showed that brown trout may lose half of their current suitable habitat within the study area by 2040 and become almost extinct by 2100. In parallel to the upstream movement of brown trout thermal habitat, warm water species are increasing their relative abundance in salmonid waters. Empirical evidence was provided of how current climate change threatens some of the most healthy native brown trout populations in Southern Europe and how forthcoming climate change is expected to further decrease the conservation status of the species.     

What predicts the use by brook trout (Salvelinus fontinalis) of terrestrial invertebrate subsidies in headwater streams?

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Diffusion in the littoral zone: scoping emergence times and movement to essential habitat for young‐of‐year brook trout in lakes

Factors governing the process of dispersal of lake‐spawned brook trout ( Salvelinus fontinalis ) young‐of‐year (YOY) appear drastically different from those governing dispersal in more commonly studied stream salmonids. Rather than dispersal being highly density‐dependent and the result of territoriality and aggression, in brook trout it may be density‐independent and driven by a common need for coldwater habitat for summer survival. Emerging fish travel great distances from single spawning sites on lakes to cold groundwater habitat. Movement is a gradual, one‐dimensional diffusion around lake margins and represents a very unique and simplified natal dispersal pathway allowing for dispersal rates, distances, and factors controlling these parameters to be measured. We predicted timing of the emergence period for brook trout alevins from lake spawning sites using the known relation between water temperature and emergence times for salmonids. Emergence and dispersal was then observed in the field by visually estimating YOY density in segments of the littoral zone throughout the dispersal period. Fluxes in density in these sections were used to estimate rate of YOY spread. YOY behaviour and body condition was also assessed across the wave of spatial spread. We are working towards a spatially explicit model to identify critical groundwater rearing habitat needing protection from forestry activities.     

Salmonid biomass and habitat relationships for small lakes

Lake morphometry and water chemistry were analyzed as predictors of brook trout and total salmonid biomass (brook trout, Atlantic salmon and Arctic char) in water bodies of Newfoundland. Lake morphometric variables included surface area, depth, perimeter and catchment area while water chemistry variables included conductivity/TDS and total Phosphorus. The broadly used Morpho-Edaphic Index was also assessed. Fish biomass/ha varied by an order of magnitude in the study lakes, despite similar water chemistry. Interactive stepwise multiple regression analysis suggested that Basin Permanence Index best explains brook trout biomass while surface area best explains the variance of total salmonid biomass in the relatively small, unproductive water bodies of Insular Newfoundland. In contrast, the Morphoedaphic Index performed poorly. The results suggest that biomass relationships with easily measured habitat variables such as surface area provide useful information, especially in management of water bodies under similar climate and nutrient regimes.     

Failure of interspecies androgenesis in salmonids

Androgenetic development of salmonid embryos was induced in recipient oocytes from the same or other species (intra- or interspecies androgenesis). Parameters for induced androgenesis were investigated in brown trout Salmo trutta and brook trout Salvelinus fontinalis . Reciprocal androgenetic and control crosses were conducted among fishes from three genera: Oncorhynchus (rainbow trout, O. mykiss ), Salmo (brown trout) and Salvelinus (brook trout), and within two genera: Salmo (brown trout and Atlantic salmon, S. salar ) and Salvelinus (brook trout and Arctic charr, S. alpinus ). Live hatched androgenetic progenies were obtained in all intraspecies variants, where oocytes and sperm originated from the same species. Interspecies androgenesis resulted in no viable larvae, despite the fact that most hybrid controls and intraspecies androgenetic individuals were viable. When recipient oocytes originated from other genera (interspecific intergeneric androgenesis), embryonic development ceased in early developmental stages, except for haploid controls of brook trout produced in eggs of brown trout. Survival of embryos to the eyed-egg stage was relatively high in the intrageneric androgenesis experiment. Nevertheless, none of these embryos survived to hatching. Some of the presumed Atlantic salmon individuals developing in brown trout eggs contained maternal DNA, questioning the accuracy of enucleation using irradiation. The inability to induce interspecific androgenesis among the examined salmonid species may have been the result of substantial kariotypical and developmental differences between spermatozoal donors and oocyte recipients, causing an incompatibility between maternal cytoplasmic regulatory factors and the paternal nuclear genome.     

Evaluating a science-based decision support tool used to prioritize brook charr conservation project proposals in the eastern United States

Declines in brook charr (Salvelinus fontinalis) throughout its historic eastern United States of America (U.S.) range prompted the formation of the Eastern Brook Trout [Charr] Joint Venture (Joint Venture) whose mandate is to restore these charr by working cooperatively at a range-wide, regional-scale. Joint Venture habitat projects in the U.S. are initiated by local community-based organizations, assisted by fisheries management institutions, and funded through the U.S. Fish and Wildlife Service. In order to assist with ranking these community-based proposals for available funding, the Joint Venture designed a quantitative scoring method as a decision support tool that addressed range-wide threats that were documented to limit brook charr production. These threats include habitat modification (e.g., dams, urbanization, and agricultural landuse) and interactions with antagonistic non-native species [i.e., brown trout (Salmo trutta) and rainbow trout (Oncorhynchus mykiss)]. This article evaluated the prioritization method used by comparing the quantitative rankings of the proposals with the final reports for 12 completed projects using paired t tests for unequal sample sizes and variance. Two of the 12 comparisons revealed statistically significant differences (P < 0.05) between project proposals and final reports; namely, the scoring criteria for the proposals did not accurately reflect the outcome of these projects in relation to brook charr management. Large inter-quartile ranges between reviewer scores were common, indicating that the quantitative scoring method includes qualitative elements. In addition to the science-based criteria, subjective value judgments by individual reviewers have factored into the Joint Venture’s decision making process. By evaluating the decision support tool, this analysis aims to assist the Joint Venture in structuring its approach to conservation and rehabilitation of brook charr populations in the eastern U.S. To better improve the Joint Venture’s ability to address brook charr management on a regional level, we recommend that future proposal prioritization scoring criteria provide more explicit guidance for accurate scoring, separate scoring methods for policy and assessment work, and a weighting factor for larger than sub-watershed projects and for scoring criteria for which information is unknown.     

Seasonal changes in stream salmonid population densities in two tributaries of a boreal river in northern Japan

We examined seasonal changes in population densities of stream salmonids (masu salmon Oncorhynchus masou, white-spotted charr Salvelinus leucomaenis, and rainbow trout O. mykiss) in two tributaries of the Shoro River, eastern Hokkaido, Japan. In one small tributary, water temperature was relatively high during the winter, and populations of salmon and trout increased through immigration at this time of the year, becoming dominant components of the salmonid assemblage; the density of charr in this stream decreased during the winter, but charr was dominant during the summer. In another medium-sized tributary, the water temperature fell to close to 0°C during the winter, and densities of salmon and charr decreased in this season, through emigration; trout were very rare in this stream. Seasonal patterns of stream salmonid densities vary among species and between localities, resulting in seasonal changes in species composition. For a comprehensive understanding of population processes, a whole-river survey across seasons will be necessary.     

Adapting streams for climate change using riparian broadleaf trees and its consequences for stream salmonids

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More than a corridor: use of a main stem stream as supplemental foraging habitat by a brook trout metapopulation

Coldwater fishes in streams, such as brook trout (Salvelinus fontinalis), typically are headwater specialists that occasionally expand distributions downstream to larger water bodies. It is unclear, however, whether larger streams function simply as dispersal corridors connecting headwater subpopulations, or as critical foraging habitat needed to sustain large mobile brook trout. Stable isotopes (δ¹³C and δ¹⁵N) and a hierarchical Bayesian mixing model analysis was used to identify brook trout that foraged in main stem versus headwater streams of the Shavers Fork watershed, West Virginia. Headwater subpopulations were composed of headwater and to a lesser extent main stem foraging individuals. However, there was a strong relationship between brook trout size and main stem prey contributions. The average brook trout foraging on headwater prey were limited to 126 mm standard length. This size was identified by mixing models as a point where productivity support switched from headwater to main stem dependency. These results, similar to other studies conducted in this watershed, support the hypothesis that productive main stem habitat maintain large brook trout and potentially facilitates dispersal among headwater subpopulations. Consequently, loss of supplementary main stem foraging habitats may explain loss of large, mobile fish and subsequent isolation of headwater subpopulations in other central Appalachian watersheds.     

Assessing the suitable habitat for reintroduction of brown trout (Salmo trutta forma fario) in a lowland river: A modeling approach

Huge efforts have been made during the past decades to improve the water quality and to restore the physical habitat of rivers and streams in western Europe. This has led to an improvement in biological water quality and an increase in fish stocks in many countries. However, several rheophilic fish species such as brown trout are still categorized as vulnerable in lowland streams in Flanders (Belgium). In order to support cost‐efficient restoration programs, habitat suitability modeling can be used. In this study, we developed an ensemble of habitat suitability models using metaheuristic algorithms to explore the importance of a large number of environmental variables, including chemical, physical, and hydromorphological characteristics to determine the suitable habitat for reintroduction of brown trout in the Zwalm River basin (Flanders, Belgium), which is included in the Habitats Directive. Mean stream velocity, water temperature, hiding opportunities, and presence of pools or riffles were identified as the most important variables determining the habitat suitability. Brown trout mainly preferred streams with a relatively high mean reach stream velocity (0.2–1 m/s), a low water temperature (7–15°C), and the presence of pools. The ensemble of models indicated that most of the tributaries and headwaters were suitable for the species. Synthesis and applications. Our results indicate that this modeling approach can be used to support river management, not only for brown trout but also for other species in similar geographical regions. Specifically for the Zwalm River basin, future restoration of the physical habitat, removal of the remaining migration barriers and the development of suitable spawning grounds could promote the successful restoration of brown trout.