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Juvenile Salmon Winter Habitat Characteristics in Large Glacial Rivers: 2014‐2015 Final Report for: Prepared by: Jeffrey C Davis and Gay A Davis P.O Box 923, Talkeetna AK 99676 September 2015

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Page 1: Juvenile Salmon Winter Habitat Characteristics in Large Glacial Rivers…arrialaska.org/reports/2014-2015 Winter Fish in Glacial... · 2017-09-07 · 2015 Winter Fish Habitat September

Juvenile Salmon Winter Habitat Characteristics in Large Glacial Rivers: 2014‐2015  

 

Final Report for: 

Prepared by:

 

Jeffrey C Davis and Gay A Davis 

P.O Box 923, Talkeetna AK 99676 

September 2015 

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Acknowledgements: Funding for this project was provided by National Marine Fisheries Service. We appreciate the field support and comments on the report provided by the NMFS Project Manager Susan Walker and field support provided by Rachel Burns.

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Contents 1.0 Summary ................................................................................................................................................. 1 

2.0 Introduction ............................................................................................................................................ 2 

2.1 Chinook Salmon .................................................................................................................................. 2 

2.2 Coho Salmon ....................................................................................................................................... 3 

2.3 Project Objectives ............................................................................................................................... 5 

3.0 Methods .................................................................................................................................................. 6 

3.1 Sampling Locations and Dates ............................................................................................................ 6 

3.2 Water Quality and Physical Habitat .................................................................................................... 6 

3.3 Fish Collection Methods ...................................................................................................................... 7 

3.4 Data Analyses ...................................................................................................................................... 7 

4.0 Results ..................................................................................................................................................... 8 

4.1 Fish Distribution and Abundance ........................................................................................................ 8 

4.2 Habitat Associations .......................................................................................................................... 14 

5.0 Discussion .............................................................................................................................................. 18 

6.0 References ............................................................................................................................................ 20 

Appendix A. Site Photographs .................................................................................................................... 26 

 

Figures Figure 1. Size frequency distribution of juvenile coho salmon from the Susitna and Talkeetna Rivers in 

February of 2015. .......................................................................................................................................... 9 

Figure 2.  Size frequency distribution of juvenile Chinook salmon at 3 mm intervals for February Susitna 

and Talkeetna River samples. ....................................................................................................................... 9 

Figure 3. Size frequency distribution of juvenile sockeye salmon for February Susitna and Chulitna River 

samples. ...................................................................................................................................................... 10 

Figure 4.  Juvenile coho CPUT for the 17 sampling locations. Sites within brackets are not significantly 

different. ..................................................................................................................................................... 11 

Figure 5.  Average juvenile coho salmon CPUT by macrohabitat type from February 2015 samples. Error 

bars are on standard deviation. .................................................................................................................. 11 

Figure 6. Average coho CPUT in paired beaver pond and downstream habitats. ...................................... 12 

Figure 7. Coho salmon CPUT in beaver ponds as a function of dissolved oxygen showing the low relative 

abundance at low oxygen concentrations. ................................................................................................. 12 

Figure 8. Average juvenile Chinook CPUT for the 17 sampling sites. Sites where CPUT was not 

significantly different are bracketed. .......................................................................................................... 13 

Figure 9.  Average juvenile Chinook CPUT by classification type. Error bars are one standard deviation. 14 

Figure 10. Coho CPUT at 5 cm/s water velocity ranges standardized by the number of samples collected 

within range. ............................................................................................................................................... 16 

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Figure 11. Coho CPUT at 1°C water velocity ranges standardized by the number of samples collected 

within each range. ...................................................................................................................................... 16 

 

Tables Table 1. Location of 2015 sampling sites and macrohabitat‐mesohabitat classification. ............................ 6 

Table 2.  Average 2015 site juvenile coho and Chinook (Chin) and average site physical and water quality 

characteristics. Cond in conductivity, Port is portion, LWD is large woody debris, Turb is turbidity, Ice D is 

ice depth, S is shrub, M is macrophyte, and W is wood. ............................................................................ 17 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

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1.0 Summary Juvenile Chinook and coho salmon migrate from spawning areas in tributaries of the Susitna River to rearing and overwintering habitats in off-channel sloughs, side channels, and tributary mouths. However, little is known about the distribution of juvenile coho and Chinook salmon during the winter months or the characteristics of those habitats. Understanding the distribution and habitat relationships of overwintering juvenile salmon is necessary to evaluate and mitigate for any proposed changes in habitat conditions due to development or a changing climate. This is a report of the third year of studies investigating juvenile salmon overwintering habitat in the glacial Susitna, Talkeetna and Chulitna Rivers.

During February 2015 sampling was conducted at 17 sites. Sampling sites were selected in upland sloughs with and without beaver ponds, side channels and side sloughs. Juvenile salmon were sampled using baited minnow traps, and physical and water quality habitat characteristics measured at 10 sampling locations within each sampling site. Results were analyzed to test for relationships between habitat characteristics and catch per unit trap. These relationships were used to determine if habitat preferences could be used to predict the differences in overwintering juvenile salmon among sampling sites. We tested for differences in overwintering juvenile salmon among macrohabitat classification type and in juvenile salmon catch between beaver ponds and sites located immediately downstream or comparable locations.

A total of 965 coho, 62 Chinook, and 18 sockeye salmon were captured during winter sampling. Juvenile coho salmon were present at 14 of the 17 sampling locations and catch per trap ranged from 0 to 26. Coho salmon catch was negatively related to water velocity at the trap location with >70% captured in velocities <5 cm/s. Coho salmon were more likely to be present in warmer water temperatures (1.5 compared to 1.0°C); however, water temperatures were 0.1°C at the site with the highest coho CPUT. The highest juvenile coho catch were in an upland slough beaver pond and a side slough. There was no significant difference in juvenile coho catch when beaver ponds were compared to sampling locations immediately downstream, which was due to the high variability among ponds and extremely low concentrations of dissolved oxygen in some beaver ponds.

Juvenile Chinook salmon were present at 8 of the 17 sampling sites and catch per unit trap ranged from 0 to 2.5. The highest juvenile Chinook salmon catch was in an upland slough and a side channel. Juvenile Chinook salmon were absent from upland slough beaver ponds. Overwintering juvenile Chinook salmon catch was related to water velocity at the trap location, cover provided by woody debris, macrophytes, or submerged shrubs and gravel substrate.

Habitat preferences identified for overwintering juvenile coho and Chinook salmon were unable to explain the variability in catch among sampling sites. Juvenile salmon were absent from some sampling sites even though they contained characteristics indicative of quality overwintering habitat. We hypothesize that conditions during fall low flows and ice development may exclude juvenile salmon from overwintering locations even if site conditions improve as winter progresses. This has implications for assessing overwintering habitat using the instream flow approach.

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2.0 Introduction Juvenile Chinook and coho salmon have been shown to migrate from natal tributaries to rearing and overwintering habitats within glacial rivers or their tributaries. For example, Murphy et al. documented Chinook and coho salmon rearing and overwintering in off-channel habitats of the glacial Taku River (Murphy et al. 1984). Juvenile Chinook salmon have been documented rearing in non-natal tributaries to the Yukon River (Bradford et al. 2001), and juvenile Chinook and coho migrate from spawning tributaries to rearing habitats in the Susitna River and its tributaries (ADFG 1981, 1983, 1986). Migration is associated with declining water temperatures but may be linked to changes in flow, or light levels (Bjornn 1971, McMahon and Hartman 1989). Migration from summer rearing habitat may be initiated by low fall flows, winter freshets, and the loss of open water as small tributaries freeze to the bottom (Prowse 1994). Juvenile salmon generally select overwintering habitats with low water velocity, cover, and relatively warmer water from springs or upwelling groundwater (Giannico and Hinch 2003, Hillman et al. 1987, Cunjak 1996). Winter habitat selection is based on the need to minimize energy expenditure and to avoid adverse physical or chemical conditions (anchor ice, floods, low oxygen) (Cunjak 1996).

2.1 Chinook Salmon Chinook and coho salmon have different winter habitat preferences. Substrate with interstitial spaces that provide cover and lower water velocities may be important for overwintering Chinook salmon (Hillman et al. 1987, Bjornn 1971). Bjornn (1971) found fewer juvenile Chinook salmon migrated out of streams with large cobble substrate than those with gravel or finer substrate. Juvenile Chinook salmon were found in association with macrophytes and undercut banks during winter and the addition of cobble substrate increased overwinter abundance (Hillman et al. 1987) in the Lemhi River (Idaho). Juvenile steelhead and Chinook were found overwintering in deep pools and the interstitial spaces of riprap cover in a large river in British Columbia (Swales et al. 1986). Bustard and Narver (1975) found juvenile coho salmon and steelhead trout in waters with velocity < 0.15 cm/s when water temperatures were below 8°C, while Hillman et al. (1987) found Chinook salmon in water velocities < 20 cm/s during winter with larger fish using higher water velocities. Winter sampling was conducted at multiple sampling locations in sloughs, side channels, and tributaries of the Susitna River during the winters of 1981, 1982, and 1983 (ADFG 1981, 1983, 1986). Juvenile Chinook were the most abundant salmon species captured during these sampling events. Juvenile Chinook salmon were present within multiple different habitat types including tributaries and tributary mouths, side channels, and sloughs; however habitat preferences were not determined due to unequal effort among sampling locations. In 1984/1985, sampling was conducted at four locations representing three habitat types: a tributary, side slough, and two upland sloughs. Juvenile Chinook salmon catch was highest in the side slough and lowest in the tributary. Marked fish were recaptured at the tagging location; however, catches in the tributary site declined during January and increased in downstream sloughs, suggesting that juvenile Chinook salmon were migrating from tributary to slough habitat. Chinook salmon were more often found in faster water velocity sites with cobble substrate and woody debris. Sampling conducted within the Susitna River mainstem and off-channel habitats during February/March 2013 found that the relative abundance of overwintering Chinook salmon was

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greater in main channel backwaters and side slough habitats (Davis et al. 2013). However, these results are based upon samples collected from two mainstem and one side slough habitat. Only five Chinook salmon were captured during sampling conducted in January and February 2014 at nine sampling locations representing tributary mouths, side sloughs and upland sloughs. Four of these Chinook salmon were captured in tributary mouth habitats and one at the confluence of an upland slough and mainstem habitats (Davis et al. 2015a). Chinook salmon relative abundance in 2013 was positively correlated with water depth and cobble substrate. Juvenile Chinook salmon may move downstream under the ice as they grow during the winter season (Levings and Lauzier 1991; Litchfield and Flagg 1988). The precise timing of these movements and the length requirements to initialize this behavior has not been identified in southcentral Alaska. However, there is some indication that downstream redistribution of Chinook salmon may begin as early as January in the Susitna River drainage (ADFG 1986). The relative abundance decreased significantly between January and March in tributary mouth habitats in 2013 and Chinook salmon were captured in side slough habitat in March 2014 where they were previously absent (Davis et al. 2013, 2015a). These differences in relative abundance over time also support the hypothesis that juvenile Chinook salmon are emigrating during winter under the ice. Additional effort needs to be conducted to determine the macrohabitat locations and microhabitat characteristics important for Chinook salmon overwintering in glacial rivers. Initial results suggest that main channel, side slough, and tributary mouths may provide overwintering habitat for Chinook salmon. However, the number of macrohabitats sampled is limited and the preference for side slough habitat is based on the results from one location (Whiskers Creek side slough), which may not be representative of this habitat type. Additional sampling needs to be conducted to test for relationships between small scale habitat characteristics and Chinook salmon relative abundance. Studies also should be developed to determine migration timing of Chinook salmon smolt.

2.2 Coho Salmon Overwintering juvenile coho salmon are found consistently in slow-water off channel habitats with abundant cover. Overwintering coho salmon habitat in British Columbia is in side channels fed by ground water (Giannico and Hinch 2003). Ground water fed side channels and ponds provide stable water flows, higher temperatures and invertebrate production allowing fish to forage and continue to grow during winter (Peterson 1982; Brown 1985 in Giannico and Hinch 2003). In coastal streams of Vancouver, juvenile coho salmon moved into side channels and beaver ponds and were associated with cover provided by woody debris or overhanging vegetation (Bustard and Narver 1975). Juvenile coho salmon emigrated from streams as light levels decreased in the fall unless they were in locations with low water velocity and abundant cover from woody debris and overhanging vegetation. Use of cover decreased with continuous ice cover (McMahon and Hartman 1989). On the Olympic Peninsula coho salmon have been found to move large distances to off channel ponds with a large amount of cover for overwintering (Petersen 1982). Similarly in British Columbia, side channel and off channel habitats with abundant cover provide overwintering habitat for juvenile coho salmon (Swales et al. 1986). In colder side channels with a surface water source, woody debris increased coho salmon carrying capacity and smolt output, but in the relatively warmer side channel with a

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groundwater source, woody debris had no effect or a negative effect (Giannico and Hinch 2003). The authors believed that cover was more important in the cold water side channel as fish swimming ability decreased with lower temperatures. Warmer groundwater was determined to be a key characteristic of productive overwintering habitat for coho salmon (Giannico and Hinch 2003). There is a paucity of information regarding the value or preference of beaver-created habitat as an overwintering location for juvenile salmonids. Pools and ponds created by beaver dams provide slow-water habitat, as well as warmer temperatures and a productive environment for vegetation and aquatic invertebrates that may not exist in an unimpounded stream (Pollock et al 2003). Bustard and Narver (1975) found that beaver ponds were an important overwintering area for coho salmon, with a survival rate almost two times that of the stream system. Although beaver activity can reduce habitat connectivity and change hydrological dynamics, dams can increase storage capacity for water, which may be critical at times of low flows (Malison et al 2014; Collen and Gibson 2011). In a southeastern Alaska stream, juvenile coho were able to use additional habitat created by beaver ponds, suggesting that the importance of the ponds was their role in increasing stream habitat heterogeneity (Bryant 1984). Swales and Levings (1989) found that coho upstream of a beaver dam were larger, more abundant, and exhibited faster growth than those downstream of the beaver dam. Unfavorable habitat physicochemical characteristics (low dissolved oxygen, high temperatures) may exclude fish or some fish species from areas created by beaver dams. The dominance of coho salmon in beaver ponds is likely due to habitat preference, although other factors have not been eliminated and Chinook juveniles are not always absent (Swales and Levings 1989). In a winter study in Oregon coast streams, alcoves and beaver ponds made up only 31% of the area sampled, but accounted for 66% of coho salmon observed, leading researchers to conclude that coho smolt production is limited by the availability of adequate winter habitat (Nickelson et al. 1992). Tributaries, tributary mouths, side channels and sloughs provide overwintering habitat for juvenile salmon in large glacial rivers. Side channels and sloughs provide slow-water habitats and are often fed by groundwater or tributary sources (Curran et al. 2011). Whiskers Creek (tributary), Whiskers Slough (side slough) and Slough 6A (upland slough) within the middle Susitna River were used for overwintering by age 1+ and 2+ coho salmon (ADFG 1984). Significant overwintering of juvenile coho salmon in the Talkeetna-to-Devils Canyon reach of the Susitna River occurs in side sloughs and upland sloughs (ADFG 1984) and some coho may also use the mainstem and side channels for overwintering (ADFG 1981). Deep water habitats and beaver ponds were found to have the highest abundance of overwintering coho salmon in the Middle Susitna River. Initial results from studies conducted on the glacial Susitna and Talkeetna Rivers during the winter of 2012/2013 documented juvenile coho salmon present within all identified macrohabitats: mainstem, tributary mouths, upland sloughs, and side sloughs. The relative abundance of juvenile salmon in October and February/March differed among sampling locations but not among classified macrohabitats (Davis et al. 2014). Coho salmon tended to be more abundant in side slough and upland slough habitats. In October with initial ice formation, correlations were strongest between water depth and the relative abundance of juvenile coho salmon. In February/March, Chinook salmon abundance and indicators of growth (average fork

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length) were correlated with water depth, but coho salmon abundance and growth indicators were correlated with water temperatures. More frequent (~monthly) winter sampling however showed a large difference in the relative abundance of juvenile salmon between December and March suggesting possible emigration or mortality between sampling events. Evaluation of the importance of winter habitat characteristics, based on relationships with fish abundance may be influenced by sample timing. Sampling conducted during the winter of 2013/2014 confirmed the coho salmon preference for upland slough and tributary mouth habitats (Davis et al. 2015). Within these habitats coho salmon were more abundant at sites with woody debris and slow water velocities (< 10 cm/s). Water temperature and water depth were not significant predictors of juvenile coho salmon abundance among these sites with water temperatures up to 2°C. Coho salmon were absent or in low abundance in side slough habitats even though these locations contained abundant woody debris and low water velocities. We hypothesize that conditions prior to sampling (low fall flows or high water velocities during freeze up) resulted in the emigration of juvenile salmon from these locations. In 2012 Chinook and coho salmon were abundant in the one side slough (Whiskers Creek) sampled; however, this side slough habitat is unique in that, due to its short length, backwater during rising stage heights as the river freezes may prevent high water velocities during breaching flows, and the outlet is controlled by Whiskers Creek not the Susitna River, so the site would not dewater during declining main channel flows. Coho salmon are most abundant in off-channel habitats with little or no water velocity and are not subject to high water velocities during river freeze up or breakup. Beaver ponds are abundant in off channel habitats of glacial rivers and appear to provide conditions of low velocity and consistent water depths. Previous studies have documented the importance of beaver dams for overwintering; however, these studies have been conducted in locations with limited ice cover. Ice cover from October through March could reduce light and oxygen diffusion resulting in low winter concentrations. Low concentrations of dissolved oxygen during winter may preclude their use by juvenile coho salmon.

2.3 Project Objectives The main objective of this project is to identify and measure the variability in habitat characteristics important for juvenile salmon overwintering in the flood-prone area of the Susitna and other similar large glacial rivers and test for relationships between these characteristics and fish population metrics. The objectives of winter 2014/2015 studies are to:

1. Improve our understanding of habitats important for coho salmon overwintering by measuring the relative abundance of juvenile salmon in beaver ponds and test for relationships between coho salmon abundance and site physical and water quality characteristics, and

2. Further investigate the use of mainstem and mainstem influenced side channels and side sloughs for Chinook salmon overwintering and test for relationships between Chinook salmon relative abundance and site physical and water quality characteristics.

 

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3.0 Methods 3.1 Sampling Locations and Dates The study was conducted at sampling sites located on the Susitna and Talkeetna Rivers, near the village of Talkeetna, approximately 100 miles north of Anchorage. The Susitna and Talkeetna Rivers are large glacial rivers located in Southcentral Alaska, with drainage areas of 6,288 mi2 and 2,021 mi2, respectively. Sites were located near the confluence of the Susitna and Talkeetna Rivers (Table 1).

A total of 17 sites were sampled in 2015 representing 4 macrohabitat types: main channel, side channel, side slough, and upland slough. Sampling units were selected in four upland and one side slough beaver pond, upland and side slough habitat downstream from ponds, and in some cases slough habitat upstream of beaver dams and ponds. Sampling units were selected to provide 5 paired beaver pond and slough habitat below beaver ponds. Sampling sites also were selected in one main channel, four side channels, and the upper end of one upland slough (Table 1). Each sampling site was sampled on one sampling date between February 7 and February 27, 2015

Table 1. Location of 2015 sampling sites and macrohabitat‐mesohabitat classification.  

Site  Habitat Classification  Latitude  Longitude 

Susitna near Chase  Main Channel  62.43554  ‐150.13142 

Billion Slough  Side Channel  62.33612  ‐150.12752 

Talkeetna (Billion)  Side Channel  62.32607  ‐150.12073 

Talkeetna (Mahays)  Side Channel  62.32647  ‐150.11615 

Talkeetna (near Town)  Side Channel  62.31970  ‐150.11745 

Whiskers Slough  Side Slough  62.37838  ‐150.16824 

Susitna near Whiskers‐BP  Side Slough‐Beaver Pond  62.38137  ‐150.16382 

Susitna near Whiskers  Side Slough‐Below Pond  62.37966  ‐150.16309 

Chulitna Slough‐UP  Upland Slough‐Upstream from Pond  62.36455  ‐150.16383 

Susitna near Whiskers‐UP  Upland Slough‐Upstream from  Pond  62.38424  ‐150.15715 

Chulitna Slough‐BP  Upland Slough‐Beaver Pond  62.35786  ‐150.17506 

Slough 5‐BP  Upland Slough‐Beaver Pond  62.45731  ‐150.12831 

Slough 6A‐BP  Upland Slough‐Beaver Pond  62.51907  ‐150.12311 

Wiggle Slough‐BP  Upland Slough‐Beaver Pond  62.34953  ‐150.10081 

Chulitna Slough  Upland Slough‐Downstream from  Pond 62.35776  ‐150.17523 

Slough 6A  Upland Slough‐Downstream from Pond  62.51606  ‐150.12434 

Wiggle Slough  Upland Slough‐Downstream from Pond  62.34953  ‐150.10081 

 

3.2 Water Quality and Physical Habitat Stream water physical and hydraulic characteristics were measured at each trap location. We measured stream water depth and ice thickness. Total depth (stream bottom to top of water or ice surface when ice present) and ice thickness was measured at each sampling location and water depth calculated by subtracting ice thickness from total depth. Water velocity was measured at

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0.6 x depth using a Flow Tracker (YSI Inc.). Stream water pH and temperature were measured at mid-water depth with a YSI 63 meter and probe. This same meter was used to measure conductivity, which could not correct for water temperatures that were generally between 0° and 2°C. Dissolved oxygen concentration and percent saturation was measured with a YSI ODO meter and probe. Water samples were collected at each site and returned to the laboratory where they were tested for turbidity and apparent color. We documented the presence of large woody debris (> 10 cm in diameter and 1 m length) if adjacent to trap and could provide cover or influence water velocities at the trap location. We recorded the presence of other cover types including shrub debris, and macrophytes. Substrate size distribution was qualified as silt/sand, gravel, or cobble based on visual observations. We used an underwater camera (SeaViewer) to look for woody debris and substrate when visibility was limited by ice or water depth.

3.3 Fish Collection Methods Fish were sampled using baited minnow traps at each sampling location and on each sampling date. Ten traps were distributed at approximately 10 m intervals along the lateral channel margins. When ice was present, an ice auger was used to cut a 10 inch diameter hole through the ice. The traps (6.4 mm (1/4 inch mesh) Gee minnow traps), baited with salmon roe inside of perforated whirl-pak bags were secured in the flowing water under the ice. A wetted depth of approximately 23 cm (9 inch) between the bottom of the ice and stream bed was necessary for trap placement. The ice opening was covered with Styrofoam insulation to aid in the recovery of the traps. Where ice was absent, traps were placed in adequate water depths to ensure the trap openings were below the water surface.

The following day, 20 to 24 hours following traps placement, traps were removed from the ice and all fish from each trap were released into separate buckets of water. Fish from each trap were identified to species and the fork lengths (FL) of all salmonids were measured, to provide catch per species per trap (catch per unit trap (CPUT)) with 10 traps per site.

3.4 Data Analyses 3.4.1 Small scale habitat relationships 

Correlation and linear regressions were used to test for relationships between habitat characteristics that were continuous variables at each trap location and coho and Chinook CPUT. T-tests and ANOVA were used to test for significant relationships between CPUT and discontinuous variables (cover and substrate size). Data were only used from sites where at least one juvenile salmon was captured to avoid data from sites where fish were absent due to potential migration barriers or other unmeasured variables. An alpha of 0.05 was used for all tests.

3.4.2 Sampling Sites 

ANOVA was used to test for significant differences in juvenile coho and Chinook salmon among sampling sites. Tukey’s test was used for post-hoc comparisons. Correlation and linear regression was used to test for significant relationships between average site CPUT for juvenile coho and Chinook salmon and for average site physical habitat and water quality characteristics and portion of sampling locations within a site that contained LWD, shrub or macrophyte cover, or fine substrates.

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3.4.3 Macrohabitats 

ANOVA was used to test for significant differences in juvenile coho and Chinook salmon among upland slough beaver pond, upland slough, side slough, and side channel macrohabitat types.

3.4.4 Beaver Ponds 

Paired t-tests were used to test for significant differences in juvenile coho salmon CPUT between beaver ponds and sites below beaver ponds. Regression analyses were used to test for significant relationships between average juvenile coho beaver pond CPUT and average site physical and water quality characteristics.

4.0 Results 4.1 Fish Distribution and Abundance Juvenile coho salmon, Chinook salmon, sockeye salmon, three spine stickleback, and sculpin were captured during winter sampling. Coho salmon were most abundant, with 965 coho captured at the 17 sampling locations. A total of 62 juvenile Chinook salmon and 18 juvenile sockeye salmon were captured. Three-spine stickleback were abundant with a total of 578 captured.

Juvenile coho salmon fork lengths in February samples ranged from 39 to 148 mm. The size distribution suggest at least two size classes with fish less than ~70 mm age 1 (Figure 1). There were 65 coho salmon < 50 mm in February, 54 of these fish were captured in Whiskers Creek side slough. These small juvenile coho suggest early emergence or slow growth during the first summer.

Juvenile Chinook salmon fork lengths in February samples ranged from 46 to 93 mm and likely represented a single age class (Figure 2). Only 1 juvenile Chinook was less than 50 mm fork length which suggests either slow summer growth or early emergence from 2014 spawning.

Juvenile sockeye salmon fork lengths ranged from 45 to 76 mm (Figure 3). The size distribution suggests a single size class. The majority of the juvenile sockeye salmon (14 of 17) were captured at the upper portion of the Chulitna River upland slough (Chulitna Slough-UP).

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Figure 1. Size frequency distribution of juvenile coho salmon from the Susitna and Talkeetna Rivers in February of 2015.  

 

Figure 2.  Size frequency distribution of juvenile Chinook salmon at 3 mm intervals for February Susitna and Talkeetna River samples.  

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Figure 3. Size frequency distribution of juvenile sockeye salmon for February Susitna and Chulitna River samples.  

4.1.1 Coho Salmon 

Juvenile coho salmon were captured at 14 of the 17 sampling locations (Figure 4). There were significant differences in juvenile coho salmon among sampling sites. Coho salmon were most abundant with >26 CPUT with Whiskers Creek side slough and Wiggle Slough upland slough beaver pond. Juvenile coho CPUT was significantly less than these two sites within Billion Slough side channel and the three sampling sites in the Chulitna upland slough. CPUT within these four sampling sites ranged from 7.3 to 13.7. Juvenile coho CPUT was significantly less at the remainder of the sites and ranged from 0 to 2.4.

There were no significant differences in coho salmon CPUT by macrohabitat type. Juvenile coho salmon relative abundance tended to be higher in upland and side sloughs; however, CPUT was highly variable among sites representing the different macrohabitat types. Coho CPUT ranged from 0 to 13.7 in upland sloughs without beaver ponds, 1 to 27 in side sloughs, and 0 to 9.9 in side channels.

There were no significant differences in paired CPUT in beaver ponds and similar habitat without ponds (Figure 6). Coho salmon CPUT in beaver ponds was highly variable ranging from near 0 to >26. Physical and water quality characteristics also were variable among beaver ponds. Average water temperature ranged from 0.1 to 2.3°C among beaver pond sites and ice thickness from 3.7 to 77 cm. Water velocity was 0 within all ponds and water depth ranged from 44 to 85 cm. The concentration of dissolved oxygen and pH were correlated and ranged from a low of 7% saturation and a pH of 6.4, to 71% saturation and a pH of 7.25. The variability in dissolved oxygen and pH in beaver ponds explained most the variability in juvenile coho CPUT (Figure 7), with low CPUT at sites with low dissolved oxygen concentrations. However, juvenile coho salmon were captured in a trap where dissolved oxygen was measured at 0.78 mg/L or 5.6% saturation.

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Figure 4.  Juvenile coho salmon CPUT for the 17 sampling locations. Sites within brackets are not significantly different.  

 

 

Figure 5.  Average juvenile coho salmon CPUT by macrohabitat type from February 2015 samples. Error bars are on standard deviation. 

 

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Figure 6. Average coho salmon CPUT in paired beaver pond and downstream habitats.  

 

 

Figure 7. Coho salmon CPUT in beaver ponds as a function of dissolved oxygen showing the low relative abundance at low oxygen concentrations.  

4.1.2 Chinook salmon 

Juvenile Chinook salmon were present in samples from 8 of the 17 sites. These sites represented all of the macrohabitats sampled; however, juvenile Chinook salmon were not captured in upland slough beaver ponds. There were significant differences in juvenile Chinook CPUT among sampling sites. The highest mean CPUT of 2.5 was at the Billion Slough side channel sampling site. CPUT at this side channel site was did not differ significantly from the next three highest

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sites with where CPUT was greater than 0.6. These included sites located in an upland slough, side channel, and side slough beaver pond (Figure 8). In addition to upland slough beaver pond habitat, juvenile Chinook salmon were absent from the two side channel sites (Talkeetna (Billion) and Talkeetna (Mahays)), and the upland slough site (Slough 6A).

There was no significant difference in juvenile Chinook CPUT among macrohabitats (Figure 9), due to large variability among macrohabitats sampled. Juvenile Chinook CPUT ranged from 0 to 2 in the upland sloughs, 0.1 to 0.6 in side sloughs, and 0 to 2.5 in side channels.

 

 

 

 

Figure 8. Average juvenile Chinook salmon CPUT for the 17 sampling sites. Sites where CPUT was not significantly different are bracketed.  

 

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Figure 9.  Average juvenile Chinook salmon CPUT by classification type. Error bars are one standard deviation. 

 

4.2 Habitat Associations 4.2.1 Small Scale: Coho salmon 

Coho salmon CPUT in traps at sites where at least one juvenile coho was present was related to water velocity and water temperature. None of the other habitat variables measured were related to coho salmon presence or relative abundance. There was no significant difference in coho CPUT among substrate types or between sites with and without cover provided by woody debris, shrubs, or aquatic macrophytes. Juvenile salmon relative abundance was higher at lower water velocities. Juvenile coho CPUT was negatively related to water velocity (cm/s) (R = 0.28; p = 0.001), with a slope of -0.66. Over 70% of the juvenile salmon captured, standardized by the number of samples at each velocity, were captured at water velocities <5 cm/s (Figure 10).

Juvenile coho salmon showed a slight preference for sites with higher water temperatures (Figure 11). The regression relationship between juvenile coho salmon relative abundance at trap locations and water temperature was not significant (p = 0.08). However, there was a significant difference in water temperatures between traps with coho salmon and traps without coho salmon (t-test, p = 0.001). Average water was 1.5 C at trap locations where coho salmon were present and 1.0 C at sites where they were absent. However, average water temperature was 0.1°C in Wiggle Slough beaver pond, one of the sites with the highest coho CPUT (>26).

4.2.2 Small Scale: Chinook salmon 

There were no significant linear relationship between juvenile Chinook salmon CPUT and any of the continuous habitat variables. There were 61 juvenile Chinook salmon captured at velocities < 5 cm/s and 1 captured at velocities greater than this value; however, differences in juvenile

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Chinook salmon CPUT between these two groups was not significant (p = 0.10). Similarly, 58 of the 62 juvenile Chinook salmon were captured where ice thickness was < 5 cm; however, there was not a statistically significant difference in juvenile Chinook salmon CPUT when comparing traps set where ice was < 5cm and >5 cm (p = 0.07). These data suggest that these variables may be important for predicting juvenile Chinook salmon CPUT, and could be significant with a larger sample size.

There was a significantly greater juvenile Chinook salmon CPUT at sites with cover provided by woody debris, shrubs, or macrophtyes (average CPUT 1.5) compared to sites without cover (average CPUT 0.5)(p=0.05). Chinook salmon were more abundant in traps placed on gravel substrate (average CPUJT = 2.1) compared to cobble or sand (average CPUT of 0.3, and 0.9, respectively).

4.2.3 Sampling Sites: Coho 

Juvenile coho CPUT was significantly related to water velocity and water temperature measured at the trap location; however, coho salmon presence or relative abundance at a sampling site could not be predicted by these variables. That is, coho salmon were absent from a number of sites with average velocities preferred by overwintering coho salmon. These included sampling sites in beaver ponds with low concentrations of dissolved oxygen and side channel and side slough sites where oxygen concentration similar to other sampling sites that supported overwintering coho salmon.

Average juvenile coho CPUT at each site was not related to average site water velocities when all sites were included. The relationship improved, but still was not significant when only sites with at least 1 coho salmon were included in the analyses. However, there was a significant relationship between coho CPUT, at sites where coho were present, if both water velocity and dissolved oxygen were used in the analyses (p = 0.04), but not if all sites were included. Including dissolved oxygen explained some of the differences in coho salmon CPUT observed in beaver ponds; however there were still sampling sites with low velocity and adequate dissolved oxygen where coho CPUT was zero. For example, juvenile coho were not captured at three sampling sites: slough 6A, Talkeetna (Billion), and Talkeetna (Mahays) (Table 2). However, all of the water quality or physical measures (depth, water velocity, dissolved oxygen) are within the range of conditions found at other locations where juvenile coho were present.

4.2.4 Sampling Sites: Chinook 

Similar to coho salmon, the habitat variables that were significantly associated with juvenile Chinook CPUT could not be used to predict juvenile Chinook CPUT at the site level. That is, juvenile Chinook salmon were more abundant in traps placed next to woody debris or macrophyte cover and gravel substrates and tended to be more abundant at trap locations where water velocity was < 5 cm/s and ice thickness < 5 cm. However, sites where these conditions were prevalent did not always have a greater average CPUT.

Average site juvenile Chinook CPUT was not significantly related to the average of any of the habitat variables measured within that site when all sites were included in the analyses. When we removed the upland slough beaver dam sites, where Chinook salmon were absent, there was a

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significant positive relationship between average site juvenile Chinook salmon CPUT and the portion of woody debris and macrophyte cover within the site. Average site juvenile Chinook CPUT was not related to average site water velocity, substrate type, or ice thickness. Similar to juvenile coho salmon, there were sites where habitat conditions appeared favorable, with woody debris, low water velocities, and open water, but Chinook salmon were absent (e.g. Susitna River main channel near Chase) (Table 2).

 

Figure 10. Coho CPUT at 5 cm/s water velocity ranges standardized by the number of samples collected within range. 

 

 

Figure 11. Coho CPUT at 1°C water velocity ranges standardized by the number of samples collected within each range. 

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Table 2.  Average 2015 site juvenile coho and Chinook (Chin) and average site physical and water quality characteristics. Cond is conductivity, Port is portion, LWD is large woody debris, Turb is turbidity, Ice is ice thickness, S is shrub, M is macrophyte, and W is wood. 

Site  Class  coho CPUT 

Chin CPUT 

Depth (cm) 

Velocity (cm/s) 

DO % Temp C 

pH Cond  Port Cob 

Port LWD 

Turb(NTU) 

Color Ice(cm) 

Port(S,M,W) 

Wiggle Slough beaver pond 

US BP  26.33  0.0  85.1 0.00 71 0.10 7.25 47.6  0.40 0.1 3.3 13.6 46 0.0

Whiskers slough 

SS  26.90  0.1  24.9 0.00 37 2.30 6.68 71.8  0.44 0.0 13.0 53.4 8.5 0.3

Chulitna Slough‐UP 

US  13.70  0.2  55.9 0.00 61 2.13 6.83 83.2  0.00 0.2 2.2 13.2 8 0.3

Billion Slough  SC  9.90  2.5  27.5 0.10 74 1.59 7.40 43.3  0.00 0.0 3.4 15.5 0.3 0.4

Chulitna Slough 

US  7.40  2.0  54.1 0.01 37 1.74 6.58 44.0  1.00 0.0 1.5 10.3 2.1 1.0

Chulitna slough‐BP 

US BP  7.30  0.0  74.4 0.00 27 1.93 7.13 60.9  0.30 0.0 0.0 2.5 3.7 0.9

Slough 5‐BP  US BP  2.67  0.0  44.0 0.02 8 0.69 6.41 37.0  0.00 0.0 3.8 23.7 30.2 0.7

Susitna near Whiskers‐BP 

SS BP  1.70  0.6  59.1 0.00 53 2.34 7.00 89.5  1.00 0.2 1.2 7.0 19.2 0.0

Wiggle Slough  US  1.10  0.1  17.7 0.43 81 0.20 7.02 60.7  0.40 0.0 0.0 8.9 7.7 0.3

Susitna near Whiskers 

SS  1.00  0.3  36.1 0.15 62 2.27 6.67 83.1  0.00 0.3 3.5 20.9 0 0.0

Slough 6A‐BP  US BP  0.60  0.0  59.8 0.00 7 0.44 6.93 86.9  1.00 0.0 0.9 2.6 77.3 0.0

Susitna near Chase 

MS/ SC 

0.43  0.0  24.3 0.20 72 1.09 7.07 59.2  0.30 0.6 1.3 7.1 0 0.0

Whiskers Slough‐UP 

US  0.30  0.0  35.8 0.19 48 1.31 7.35 121.0  0.40 0.4 0.0 0.0 1.5 0.6

Talkeetna (near town) 

SC  0.20  0.8  31.6 0.27 86 0.56 7.25 138.5  1.00 0.0 0.0 0.0 0.0 0.6

Slough 6A  US  0.00  0.0  68.5 0.00 81 0.33 6.87 29.7  0.00 0.0 3.6 15.3 65.6 0.0

Talkeetna (Billion) 

SC  0.00  0.0  43.2 0.00 71 0.88 7.23 123.5  0.90 0.0 0.0 0.0 10.3 0.0

Talkeetna (Mahays) 

SC  0.00  0.0  35.4 0.18 82 0.58 7.17 103.0  0.50 0.2 0.0 0.0 4.8 0.0

 

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5.0 Discussion Juvenile coho and Chinook salmon were found overwintering at multiple different sampling sites representing all of the macrohabitat classification types found in glacial rivers. Physical and water quality habitat variables were related to their relative abundance at sites where present, consistent with other studies. However, their distribution among sampling sites could not be explained by the predominance of these same variables within a site. This suggests that other variables influence the distribution of juvenile salmon regardless of site physical and water quality habitat conditions measured during mid-winter. This could reduce the effectiveness of analyses that model winter distribution and abundance over time (e.g. IFIM Bovee et al. 1988), unless these other variables are identified and accounted for. We hypothesize that hydraulic conditions during early winter, prior to and following initial ice development, could result in the displacement of juvenile salmon from sites that would otherwise provide favorable overwintering habitat conditions.

The habitat variables that were associated with juvenile coho and Chinook salmon CPUT during winter are consistent with previous studies. Juvenile coho salmon preference for low water velocities was the variable that explained most of the difference in juvenile coho salmon CPUT among trap locations, with most juvenile Chinook and coho salmon captured in velocities <5cm/s. Results from 2014 sampling also documented average velocities at trap locations where coho were captured at 3.6 cm/s (Davis et al 2015). Overwintering juvenile salmon preference for low water velocities has been documented previously (see review by Morgan and Hinojosa 1996).

Cover provided by woody debris was not found to be a variable that affected juvenile coho salmon habitat selection based on results from 2015 sampling, but was determined to be an important variable in 2014 (Davis et al. 2015). This is likely due to the function of large wood and the differences in sites sampled between these two years. Providing areas of low water velocity is one of the primary functions of large wood. For example, McMahon and Hartman (1989) found that juvenile coho salmon stayed within a stream channel under increasing water velocity and decreasing water temperatures when cover was provided by a large root wad. Giannico and Hinch (2003) also showed that as water temperatures decreased in a surface fed channel, the presence of wood increased the carrying capacity for overwintering coho salmon. Taylor et al. (1998) also noted that overwintering juvenile coho salmon sought cover of logs during conditions of rising water velocities and decreasing temperatures. Results from sampling in 2014 in side sloughs and tributary mouths of the Susitna River found that juvenile coho salmon abundance was only related to water velocity when wood was absent. Large wood therefore mitigated the influence of water velocity on coho distribution. During 2014 a large number of the sampling locations where coho salmon were captured were in sites with low water velocities (beaver ponds and upland sloughs), since water velocities were already low, the presence of large wood would be less likely to affect juvenile coho distribution.

Overwintering juvenile salmon use of beaver ponds was highly variable ranging from the highest coho salmon CPUT among all sites to beaver ponds with CPUT <1. This likely reflects the large variability in habitat conditions within beaver ponds during the onset of winter and during winter

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sampling. The older beaver ponds at slough 6A and Slough 5, located in upland sloughs without an upstream connection to the mainstem, had a thick surface layer of ice and extremely low dissolved oxygen concentrations. A few juvenile salmon were present at these low oxygen concentrations in beaver pond 6A and at the inlet to the pond (Slough 5) where oxygen concentrations were slightly higher (10% saturation and 1.4 mg/L). However, the Wiggle Slough beaver pond with the highest coho CPUT was also old and in an upland slough with a complete and thick ice cover but with dissolved oxygen concentrations >70% saturation. The Chulitna upland slough beaver pond is located in a channel that often receives mainstem flow during spring break-up and ice dams downstream. The upstream end of the slough is within ~200 m of the Susitna River and the upstream Chulitna Slough-UP site, ground water discharge results in clean cobble/gravel bed and dissolved oxygen saturation is near mainstem values at 61%. Complete ice cover occurs within 30 m and continues to the Chulitna Slough beaver pond site, ~500 m downstream, where dissolved oxygen had dropped to 27% saturation. However, it may be that upwelling from the mainstem provided oxygenated water to this beaver pond site sufficient to support the relatively high juvenile coho salmon average CPUT of 7.0. The beaver pond in the Susitna near Whiskers side slough site was well oxygenated at 53% and supported overwintering coho salmon, but was a new pond constructed some time during the summer of 2014. We believe that the relatively low coho CPUT in this beaver pond is likely because this pond did not provide preferred habitat conditions, and had low juvenile coho salmon abundance through the summer and early winter.

Juvenile Chinook salmon were associated with large wood, shrubs, and macrophytes. Juvenile Chinook salmon were most abundant in the Chulitna upland slough, downstream from the beaver pond. All of the sampling locations within this site were associated with large wood, submerged tree branches (shrubs), or macrophytes. Wood was common in the Billion Slough side channel, another site where juvenile Chinook were abundant. In 2013, juvenile Chinook salmon were most abundant at two main channel sites and the mouth of Wiggle Creek which also contained a large amount of woody debris with 26, 41, and 24 pieces/100 m respectively (Davis et al. 2013). Hilman et al. (1987) found that overwintering juvenile Chinook salmon preferred habitats with undercut banks and woody debris. Swales et al. (1996) also demonstrated that overwintering juvenile Chinook preferred deep pools with log debris.

Overwintering juvenile Chinook salmon appear to prefer areas without ice cover. In both 2013 and 2015 juvenile Chinook salmon were more abundant in sites without ice cover, usually associated with moderate velocities and groundwater discharge. Brandford et al. (2001) demonstrated a negative association between juvenile Chinook overwintering survival and the development of aufeis in an upper Yukon River tributary. However, the development of a 1 m thick layer of ice in that study may have eliminated or severely reduced the amount of flowing water habitat during winter, which is different from a behavioral avoidance of sites with ice cover. In 2013 juvenile Chinook salmon were captured under the ice in side slough habitat near Whiskers Creek, and in 2015 juvenile Chinook salmon were captured in Susitna River side slough habitat upstream from Whiskers Creek. However, at both of these sites emigration was restricted due to either a dewatered channel or a beaver dam.

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In all three years of sampling, juvenile salmon have been absent or in low abundance at sampling sites that contain the preferred habitat conditions for overwintering juvenile coho and Chinook salmon measured during the middle of winter. We hypothesize that habitat conditions during early winter prior to and during ice development are not those preferred by overwintering juvenile salmon and that these sites are not recolonized as habitat conditions improve.

Susitna River flows and water surface elevations generally decline during autumn and early winter. Average monthly flow at the USGS Gold Creek gauging station declines from 14,000 cfs in September to 6,400 cfs in October prior to ice development. Flow in sloughs and side channels with outlets that are controlled by the mainstem stage height will decrease to residual groundwater discharge. Without an outlet control provided by bed elevations or beaver dams, total rearing area will be reduced considerably or eliminated. Once ice develops in the main channel and main channel water surface elevations increase, these sites can receive mainstem backwater flow, and habitat area and conditions (velocity) can improve. Rising water surface elevations during ice development also could overtop the upstream ends of side cannels and side sloughs (breaching flows). This could increase water velocity in these sites causing juvenile salmon emigration. Only very slight increases in water velocity have resulted in juvenile coho salmon emigration (Giannico and Healy 1998, McMahon and Hartman 1989, Tschaplinshki and Hartman 1983). Rearing juvenile salmon will need to migrate out of the site to the main channel or migrate upstream in the slough, side channel, or tributary to search for overwintering habitat. Since winter site fidelity is high, and main channel conditions during winter are not conducive to juvenile salmon migration, it is unlikely that these sites will be recolonized during mid-winter. This hypothesis, if supported, will affect estimates of habitat availability using instream flow methods.

Early winter conditions and conditions during ice development may present a bottle neck in estimating overwintering habitat availability in glacial rivers. Methods that assess overwintering habitat area by modeling or measuring hydraulic and habitat conditions may overestimate overwintering habitat area, if short-term changes exclude juvenile salmon use of sites for the remainder of the winter. In this case minimum estimates of habitat area and habitat quality from October through March may provide a more accurate assessment of juvenile coho and Chinook salmon overwintering habitat.

6.0 References ADFG (Alaska Department of Fish and Game). 1981. Juvenile anadromous fish study on the lower Susitna River. Phase I Final Draft Report. ADFG/Su Hydro Aquatic Studies Program, Anchorage.

ADFG (Alaska Department of Fish and Game). 1983. Resident and juvenile anadromous fish studies on the Susitna River below Devil Canyon, 1983. Phase II Basic Data Report, Vol. 3. ADFG/Su Hydro Aquatic Studies Program, Anchorage.

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ADFG (Alaska Department of Fish and Game). 1981b. Susitna Hydro Aquatic Studies - Phase I Final Draft Report: Juvenile anadromous fish study on the lower Susitna River. Prepared for Acres American, Inc, Buffalo, NY. 121 pp.

ADFG (Alaska Department of Fish and Game). 1984. Susitna Hydro Aquatic Studies, Report No. 2: Resident and juvenile anadromous fish investigations. May - October 1983. Dana C Schmidt, Stephen S. Hale, Drew L. Crawford, Paul M. Suchanek (eds.), Prepared for: Alaska Power Authority, Anchorage, AK. 396 pp.

ADFG (Alaska Department of Fish and Game). 1984b. Susitna Hydro aquatic studies report no. 1. ADF&G, Susitna Hydro Aquatic Studies Report Series, Susitna Hydro Document No. 1450, Anchorage, Alaska.

Alaska Department of Fish and Game. 1986. Report No. 11, Winter studies of resident and juvenile anadromous fish (October 1984-May 1985) Part 2. ADFG/Susitna River Aquatic Studies Program. Anchorage, Alaska.

Alfredsen, K. and E. Tesaker. Winter habitat assessment strategies and incorporation of winter habitat in the Norwegian habitat assessment tools. Hydrological Processes 16: 927-936.

Barrett, B.M., F.M. Thompson, and S.N. Wicks. 1985. Adult salmon investigations: May-October 1984. Susitna Aquatic Studies Program. Report No. 6. Alaska Department of Fish and Game, Anchorage, Alaska. APA Document #2748.

Bell, E. 2001. Survival, growth and movement of juvenile coho salmon over-wintering in alcoves, backwaters, and main channel pools in Prairie Creek, California.

Bjornn, T.C. 1971. Trout and salmon movements in two Idaho streams as related to temperature, food, stream flow, cover, and population density. Transactions of the American Fisheries Society 100(3):423-438.

Bovee, K. D., B. L. Lamb, J. M. Bartholow, C. B. Stalnaker, J. Taylor and J. Henriksen. 1998. Stream habitat analysis using the instream flow incremental methodology. U.S. Geologicial Survey, Biological Resources Division Information and Technology ReportUSGS/BRD-1998-0004. viii +131 pp.

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Appendix A. Site Photographs  

Photograph A1. Wiggle slough sampling site facing the Talkeetna River. 

 

 

Photographs A2. Wiggle slough beaver dam from Wiggle Slough. 

   

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Photograph A3. Wiggle slough beaver pond.  

 

 

Photograph A3. Susitna River near Whiskers upstream from pond upland slough.  

   

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Photograph A4. Slough 6A beaver pond. 

 

 

Photograph A5. Slough 6A downstream from beaver pond.  

   

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Photograph A6. Slough 6A.  

 

 

Photograph A7. Slough 5 beaver pond.  

   

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Photograph A8. Slough 5A beaver pond inlet.  

 

 

Photograph A9. Susitna River main channel near Chase. 

   

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Photograph A10. Susitna River near Whiskers side slough.  

 

 

Photograph A11. Susitna River near Whiskers side slough beaver pond. 

   

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Photograph A12. Billion slough Talkeetna River side channel. 

 

 

Photograph A13. Whiskers side slough. 

   

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Photograph A14. Hole in ice from groundwater upwelling in Whiskers Creek side slough.  

 

 

Photograph A15. Talkeetna (Billion) side channel. 

   

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Photograph A16. Talkeetna (near town) side channel. 

 

 

Photograph A17. Chulitna upland slough beaver pond. 

   

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Photograph A18. Chulitna upland slough upstream from beaver pond. 

 

 

Photograph A19. Chulitna upland slough upstream from beaver pond. 

   

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Photograph A20. Chulitna upland slough downstream from beaver pond.  

 

 

Photograph A21. Aquatic macrophytes and iron flocculants in Chulitna slough downstream from beaver pond. 

   

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Photograph A22. Talkeetna River (Mahays) side channel.