2010 NORTHEASTERN NATURALIST 17(3):357–372 Movement and Habitat Use by Transplanted Adult Brook Trout in an Appalachian Headwater Stream Kyle J. Hartman1,* and Marisa Nel Logan2 Abstract - Salvelinus fontinalis (Brook Trout) are simultaneously the subject of eradication efforts in the western US and restoration efforts in the East. Thus, knowledge of their habitat requirements are important to management as well as ecological understanding of the species. Previous studies have evaluated habitat use and movement of established, resident Brook Trout, but none had looked at how transplanted Brook Trout respond in novel environments, nor has habitat selection been evaluated under different flow regimes that may detect differential use of primary habitat. We implanted wild Brook Trout with radio tags and tracked their movement for approximately 30 days during late spring 2002 and early spring 2003 in a central Appalachian stream. The hypotheses tested were: (1) there is no difference between habitat used by novel Brook Trout and available habitat, and (2) stream discharge levels have no effect on Brook Trout habitat selection. The daily tracking of fish in this study also permitted us to quantify fish movement. Brook Trout showed a preference for pool habitats—using them in greater proportion than availability—as well as a preference for large woody debris as cover. Overall, we found stream discharge did not affect habitat use. However, under low discharge levels, a negative relationship between discharge and pool use was detected, suggesting restriction to pool habitats under low flows. Home ranges of Brook Trout derived from radio telemetry averaged 450 m—similar to values obtained in other Appalachian studies employing mark-and-recapture methods. A comparison of our results with those of other studies suggests that Brook Trout released into novel environments move and select habitat similar to fish that have local knowledge of the environment. Introduction Previous studies have documented the habitat use and movements of Salvelinus fontinalis (Mitchilli) (Brook Trout) under natural conditions (Adams et al. 2000, Flebbe and Dolloff 1995, Gowan and Fausch 1996, Hansbarger 2005, Helfrich and Kendall 1982, Riley et al. 1992). These studies observed Brook Trout within the same stream in which they were captured and did not consider how habitat use may change under different flow regimes. Under these natural conditions, the most widely used type of primary habitat for Brook Trout are pools (Flebbe and Dolloff 1995, Hansbarger 2005). Movement studies of Brook Trout have identified differing demographics within the populations: some fish appear to move very little (e.g., <250 m; Niles 2010, Sweka 2003), yet immigration rates are often very high (60–91%; Gowan and Fausch 1996, Gowan et al. 1994, Sweka 2003), suggesting some individuals roam on larger spatial scales. 1Wildlife and Fisheries Resources Program, West Virginia University, Division of Forestry and Natural Resources, 322 Percival Hall, Morgantown, WV. 2Civil and Environmental Consultants, Inc., 333 Baldwin Road, Pittsburgh, PA 15205. *Corresponding author - hartman@wvu.edu. 358 Northeastern Naturalist Vol. 17, No. 3 Although numerous studies have examined the habitat use and movement of Brook Trout, previous studies focused on the response of fishes collected within the same stream as within which the study was performed. These fish would presumably already have local knowledge of the suite of habitats and conditions and their spatial arrangement within the study stream, potentially biasing results. Although native to the eastern United States, Brook Trout are considered an invasive species in the West, where they often out-compete native trout species (Benjamin et al. 2007, DeStaso and Rahel 1994, Peterson et al. 2004). This invasive tendency in the West coupled with efforts to restore Brook Trout in its native range make knowledge of habitat use and movement of novel Brook Trout—fish transplanted into a system and without prior local knowledge—an important aspect of the species biology with key management and restoration implications. Therefore, the primary objective of this study was to evaluate the habitat use, home range, and movement patterns of transplanted Brook Trout in an Appalachian headwater stream. A secondary objective was to examine habitat use under different flow regimes to document habitat use under low and high spring flows. Methods This study was conducted on Birch Fork, a 5.6-km tributary of Rocky Run in the Middle Fork River watershed, Randolph County, WV (Fig. 1). Birch Fork has a gradient of 3% and it was chosen because it supports naturally reproducing Brook Trout and is typical of streams in the area, Figure 1. The Birch Fork study site in the Middle Fork River watershed, Randolph County, WV. 2010 K.J. Hartman and M. Nel Logan 359 particularly those within the Pottsville Group geology. The watershed encompasses 5.8 km2 at a mean elevation of 871 m. The watershed is characterized by well-drained acidic soils dominated by mid- to high-elevation deciduous forest communities. Precipitation in this area often totals more than 130 cm a year. The upper portion of the study stream is buffered with the addition of limestone sand every year by the West Virginia Division of Natural Resources. The lower 1 km is easily accessible to vehicles and anglers. Most of Birch Fork is located on the MeadWestvaco Wildlife and Ecosystem Research Forest (now owned by Penn-Virginia Corporation) and has limited public access. Movement and habitat-use studies were conducted during May–June 2002 and March–May 2003. Total pool area throughout the study stream was measured and used to systematically distribute the 20 radio-tagged fish used each year (one fish per each section of stream containing 5% of the total pool area). Fish were released within minutes of surgical procedures, necessitating the release of fish in pool areas where they could recover without displacement by stream currents. Pool areas were measured using the basinwide visual estimation technique (BVET; Dolloff et al. 1997). The BVET survey began 200 m above the confluence of Birch Fork and Rocky Run during 2002. Thus, the lower 200 m of Birch Fork (above Rocky Run) served as a buffer to minimize fish movements out of the study area. In 2003, the lower 1000 m of Birch Fork was used as a buffer against angling losses of tagged fish that occurred in 2002. Available habitat and habitat use by tagged fish was evaluated to test whether novel Brook Trout selected for any particular habitat type. Available habitat was assessed as in Petty et al. (2001), by dividing the Birch Fork site into three sections (upstream, midstream, and downstream; Table 1). Habitat measurements were taken using the representative reach extrapolation technique (Platts et al. 1983). Two reaches, each measuring 35 times the mean stream width, were randomly selected within each section. Transects within these reaches were spaced 2–3 times the mean stream width (Table 1). Microhabitat availability variables were measured at five equally spaced points across each transect. Within each reach, a total of 23–24 transects were assessed for habitat Table 1. Characteristics of stream sections used for habitat assessments in Birch Fork during 2002 and 2003. The location of stream reaches were identical in 2002 and 2003. Here, n is the number of transects measured for microhabitat per reach. Mean Mean Mean Stream Location (meter mark) stream canopy stream section Year Reach 1 Reach 2 width (m) cover (%) depth (cm) n Upstream 2002 4410–4531 5189–5310 2.78 62.3 17.5 24 2003 3.76 - 14.6 24 Midstream 2002 3045–3202 3785–3939 3.71 66.0 11.3 24 2003 4.22 - 11.8 24 Downstream 2002 735–976 1470–1712 4.86 57.3 25.1 23 2003 5.25 - 18.3 23 360 Northeastern Naturalist Vol. 17, No. 3 measures each year. A total of 1037 m of Birch Fork was sampled for habitat each year, representing 18.5% of the total stream length (5600 m). Microhabitat availability variables measured included primary habitat type (pool, riffle, run), structural association (large woody debris, boulders, root wads, etc.), distance to nearest bank, distance to and type of cover, percent canopy cover, water depth, water temperature, and current velocity. During the second season (2003), percent canopy cover was not recorded because all trees were not in leaf during habitat measurements. Percent substrate composition was visually estimated within a 400-cm2 quadrat (Petty et al. 2001) and was also measured using modified Wentworth classification for substrate particle sizes (McMahon et al. 1996). Large woody debris was defined as any piece of wood within the bank-full channel that was larger than 10 cm in diameter and 1 m in length (Overton et al. 1997). Boulders were defined as greater than 30 cm along the median axis (Young 1996). Average velocity was considered the velocity at 60% of the water depth (Petty et al. 2001). Cover type, habitat type, and structural associations were all visually classified by the same researcher to minimize variability. Discharge was measured daily at three points along Birch Fork during tracking efforts (1260 m, 2860 m, and 4375 m into the study reach). Data from the most down-stream reach was later used to classify stream flow conditions. Radio telemetry was used to monitor daily movement of 20 Brook Trout and their habitat use for 30 days each year. Brook Trout were collected by electrofishing (pulsed DC output) and weighed before being selected for the study. Each fish should weigh at least 65 g (approximately 165 mm TL) to carry the 1.3-g tag, as the tag should not exceed 2% of the fish’s total weight (Bunnel et al. 1998, Winter 1983). The radio tags (F1400 series, Advanced Telemetry Systems) were sealed in epoxy with external dimensions of 8 x 16 x 7 mm. Tags were set to pulse at 35 ppm, giving them a warranty battery life of 28 days. Each tag had a unique frequency which permitted identification of individual fish. Fish were transported to Birch Fork in 19-L buckets. Surgery and release took place within 12 h of capture. Surgery techniques were performed similar to those used by Hart and Summerfelt (1975) and Young (1995). Fish were anesthetized using a 120 mg/l clove oil solution (Anderson et al. 1997) for approximately 3–4 min or until they lost equilibrium. Fish lengths and weights were then measured, and radio tags were inserted through an 8-mm incision located immediately anterior to the pelvic girdle and about 3 mm to the right of the ventral midline (Young 1995). Three sutures, approximately 3 mm apart, were used to close the incision (Hart and Summerfelt 1975). Surgery time was kept under 3 minutes to increase survival (J. Isely, Clemson University, Clemson, SC, pers. comm.). Following surgery, fish recovered while being transported to their designated stream section for release (10–15 min). There were 20 stocking sites, which each contained 5% of the total pool area and received one randomly selected, implanted fish. Fish were placed in the nearest pool within the designated section (for recovery purposes) and observed to verify recovery from the anesthesia. Fish were given at least 1.5 d to resume normal behavior before tracking began. 2010 K.J. Hartman and M. Nel Logan 361 Tracking was conducted by walking parallel to the stream, within 50 m of the bank (Young 1995). Radio-telemetry signals for comparable tags and receivers (Advanced Telemetry Systems) were reported to be detected from approximately 100 m away for stream-dwelling trout (Matthews 1996), but we found them to be detectable at a range of 200 m. Triangulation was used to determine fish location, and visual verification was also conducted to ensure accurate locations. Each fish was located once a day. Tracking occurred between 0730 h and 1830 h, and the tracking path was randomized to avoid covering the same stream reaches at the same time each day. Upon location of a fish, microhabitat measurements (the same as those made to determine available habitat) were made to evaluate habitat use. Diel movements of fish may differ from day-to-day movements (Bunnel et al. 1998, Young et al. 1997). Therefore, to document diel movements, three fish were tracked hourly within a 24-h period in 2002. These individual fish were selected because they were located near a stream access point and were within a 300-m reach of the downstream section, affording the opportunity for hourly observations. Brook Trout movements were assessed by measuring the distance from each location point to the closest 25-m bench mark on a line parallel to the thalweg (Young 1995). Total movement by each fish was calculated in three categories: home range, net movement, and cumulative movement over time. Home range was measured as the distance between the farthest upstream and downstream locations (Young 1996). Net movement was the distance between the original release site and the final position at which an individual fish was located. Cumulative movement was a sum of all movements from day to day. Daily movement was cumulative movement divided by days at large. Comparisons of habitat use versus availability were made using all fish locations after initial analysis using only a subset of all fish location observations (because some fish had twice as many observations as others, due to losses) yielded no difference in habitat use between full and subset locations. In this preliminary analysis, the subset locations were randomly deleted until all fish had 9 locations for the first season and 8 for the second season (Rogers 1998). Seven tags were lost over the two seasons: from angling (4), predation (1), emigration (1), and tag expulsion (1). Microhabitat preferences were determined by using the log-likelihood ratio test for discrete variables (primary habitat type and structural association) and the Mann-Whitney test for continuous variables (distance to nearest bank, distance to nearest cover, water depth, and velocity). Analysis of variance was used to evaluate relationships between fish length and movement or habitat use, as well as between discharge levels and habitat use. Statistical significance was determined at ά = 0.05. Fish habitat use under different flow regimes was assessed by ranking the daily discharge levels from 2002 and 2003 and determining the relative stream flow (Fig. 2). Only dates in which 10 or more fish were detected were included in this analysis. The lower 25th percentile of observations (<0.095 m3/s, n = 10) was considered “low flow”, the upper 25th percentile of discharge observations (>0.313 m3/s, n = 10) were considered “high flow”, and the middle 50 percent of observations (0.104–0.313 m3/s, n = 20) were 362 Northeastern Naturalist Vol. 17, No. 3 considered “normal flow”. For each date and corresponding daily flow, the observed habitat use (e.g., percentage of fish in pools, riffles, or runs) was then compared within a relative stream flow category, using regression to evaluate habitat use across stream discharge levels. Results Movement Four tagged fish were lost during 2002, and the remaining 16 were located an average of 18 times (range = 10–21; SD = 3.61) and occupied an average of 15 unique locations (range = 9–20). Average home range was 637 m (range = 87–2681 m; SD = 647.7), average cumulative distance traveled by each fish was 1137 m (range = 227–4469 m; SD = 1045.5), and net movement averaged 337 m upstream (range = 250 to +1195 m; SD = 406.1). Each of the 20 fish tracked in 2003 was located an average of 19 times (range = 15–21, SD = 1.67) at an average of 14 unique locations (range = 8–18). Home range averaged 301 m (range = 4–1395 m, SD = 369.6), the average cumulative distance moved was 777 m (range = 9–2996 m, SD = 1004.3), and average net movement was 27 m downstream (range = -519 to +382 m, SD = 189.4). Only net movement was significantly different between 2002 and 2003 (t-test, 34 d.f.: t = 3.57, P = 0.0011), so we pooled 2002 and 2003 movement metrics to examine overall trends in movement by Brook Trout in novel Figure 2. Discharge measurements for the lower study reach of Birch Fork during the 2002 and 2003 tracking season. Dashed lines represent the threshold discharge levels between low flows (25th percentile, 0.095 m3/s) and high flows (75th percentile, 0.313 m3/s) used for evaluating habitat use relative to discharge. 2010 K.J. Hartman and M. Nel Logan 363 environments. Movement metrics indicated some tagged Brook Trout were relatively immobile, while others moved much greater distances (Figs. 3 and 4). Cumulative movement was <1000 m for most fish during the observational periods, but several moved over 2800 m (Fig. 3). Similarly, most fish moved less than 20 m per day, but one moved as much as 124 m per day (Fig. 3). Not surprisingly, estimates of home range were typically less than 400 m, but estimates of 800–1100 m were also common (Fig. 3). Net Figure 3. Movement characteristics for 36 radio-tagged Brook Trout in a headwater Appalachian stream, Birch Fork, WV during spring 2002 and 2003 (combined). 364 Northeastern Naturalist Vol. 17, No. 3 movement was upstream in 2002, but in 2003 when flows were higher, net movement was slightly downstream (Fig. 4). A 24-h tracking experiment was conducted on a randomly selected group of 3 fish on 18–19 June 2002. The average cumulative movement (± SD) for these three fish was 40.8 ± 27.1 m. Of their cumulative movement, 85% occurred during the hours of 0530–0930 h and 1730–2130 h. During this time, they were found in runs and riffles. Habitat use There were significant differences between habitat use and availability during 2002 (Table 2). Brook Trout did not occupy primary habitats in proportion to availability (likelihood ratio: χ2 = 72.1, 2 d.f., P < 0.0001) and tended to select for pool habitats. Within pools, trout also showed selection for LWD and to a lesser extent boulders as the pool-forming mechanism (likelihood ratio: χ2 = 8.35, 2 d.f., P = 0.015). Brook Trout also selected for cover type (likelihood ratio: χ2 = 23.6, 3 d.f., P < 0.0001), with most differences occurring in higher use of rootwads than availability. No selection of habitat on the basis of substrate composition was detected (likelihood ratio: χ2 = 7.31, 6 d.f., P < 0.29). All continuous variables (distance to bank, distance to cover, velocity, and depth,) showed significant selection (P < 0.001) as fish used faster, deeper water in areas farther from the bank and closer to cover than what was proportionally available (Table 2). Figure 4. Net movement of Brook Trout tracked during 2002 and 2003. Net movement was considered the distance from original placement to final tracking location. Negative values represent a net downstream movement, and positive values represent a net upstream movement. 2010 K.J. Hartman and M. Nel Logan 365 In 2003, Brook Trout again showed selection for primary habitats (likelihood ratio: χ2 = 150.6, 2 d.f., P < 0.0001), tending to prefer pools and avoid riffles (Table 2). Selection for pool-formation mechanism was also evident (likelihood ratio: χ2 = 20.5, 2 d.f., P < 0.0001), with an even greater proportional use of pools formed by LWD than in 2002. Selection for rootwads and LWD as cover appeared to be responsible for rejecting the null hypothesis that Brook Trout did not select for cover types (likelihood ratio: χ2 = 38.4, 3 d.f., P < 0.0001). Most results for the continuous variables were comparable to 2002, with fish preferring deeper water, farther from the bank, and closer to cover (P < 0.001). However, in 2003, trout selected slower water velocities (P < 0.001) than was proportionally available (Table 2). Fish size effects Lengths of Brook Trout used for the telemetry were not significantly different between years (t = 1.72, 37 d.f., P = 0.093), but length ranges (168–230 in 2002 [14.8 SD], 159–204 in 2003 [11.1 SD]) warranted analysis of the influence of fish size on movement and habitat selection. Analysis to see if fish length influenced cumulative movement, net movement, home Table 2. Comparison of availability of various habitat components and their use by Brook Trout as assessed by radio telemetry during 2002 and 2003. Asterisks indicate statistical significance. 2002 2003 Available Use Available Use Primary habitat type* Pool (%) 11 39 10 48 Riffle (%) 71 43 60 17 Run (%) 18 18 30 34 Pool formation* Boulder (%) 13 17 41 26 Free form (%) 84 65 54 33 LWD (%) 3 18 5 43 Cover type* Boulder (%) 72 75 75 55 LWD (%) 16 11 17 29 Rootwad (%) 2 9 2 12 Under cut bank (%) 10 5 6 4 Substrate composition Boulder (%) 19 21 16 14 Cobble (%) 30 28 27 29 Pebble (%) 28 19 28 25 Gravel (%) 10 11 15 16 Sand (%) 9 11 7 9 Silt (%) 2 1 2 3 Bedrock (%) 2 9 4 4 Organic (%) 1 0 Distance to bank (m)* 0.95 1.19* 1.16 1.36* Distance to cover (m)* 1.50 0.71* 1.32 0.71* Velocity (m/s)* 0.135 0.155* 0.255 0.164* Depth (cm)* 13.3 30.5* 14.9 38.3* 366 Northeastern Naturalist Vol. 17, No. 3 range, daily movement, and proportional use of pools and run habitats by year yielded three significant results—each related to movement metrics in 2003. Cumulative movement (F = 7.29; 1, 18 d.f.; P = 0.015), home range (F = 4.81, 1, 18 d.f.; P = 0.042), and daily movement (F = 7.28, 1, 18 d.f.; P = 0.015) were all negatively related to fish size. Use of pools and runs by individuals was not related to fish size in either year (P > 0.213). Habitat use under different stream discharge levels Patterns in habitat use by Brook Trout as a function of stream discharge showed trout typically were observed in pool habitats regardless of stream discharge levels (Fig. 5). We could not reject the null hypothesis stating that habitat use did not change with stream discharge. Overall, there was no relationship between stream discharge and percentage of fish found using pools each day (R2 = 0.01, F = 0.57, P = 0.46). However, within the low-flow observations, pool habitat use by trout declined as stream discharge increased (R2 = 0.53; F = 9.06, P = 0.017). Discussion Brook Trout habitat use Our study clearly showed that Brook Trout in novel environments preferred pool habitats over riffles and runs. Thus, we rejected the null hypothesis that Brook Trout did not prefer pool habitat types. Previous studies with resident fish have also found pools to be important habitat for trout (Berg et al. 1998, Bunnell et al. 1998, Flebbe and Dolloff 1995, Gowan and Figure 5. Daily percentage of radio-tagged Brook Trout using pool habitats relative to daily stream discharge measurements in Birch Fork during spring 2002 and 2003. 2010 K.J. Hartman and M. Nel Logan 367 Fausch 1996, Hansbarger 2005, Young 1995). In our study stream, pools represented 10–11% of available habitat, but 39–48% of observations found Brook Trout in pools. Thus, Brook Trout in novel habitat appear to select similar habitats as fish with knowledge of their environment. Other aspects of Brook Trout microhabitat preferences are likely skewed by the heavy use of pool habitats. Fish were found at greater distances to the bank, shorter distances to cover, and deeper water than was available overall in the stream. Most of these characteristics are reflective of pool habitats. Distances to banks are often greater in pools due to the typically wider stream widths in pools compared to riffles and runs. Cover is often nearby in pools as it is often associated with the pool-forming mechanism. Similarly, pools have greater depths than other habitats. Although significant statistically, some of these microhabitat preferences (e.g., distance to bank) are small and probably not biologically significant. Mean water velocities occupied were similar between years (0.155–0.164 m/s), probably representing some physiological preference or limitations (Videler and Wardle 1991, Wilson and Egginton 1994). The selection for higher than average available velocity in 2002 when stream discharge was lower, may reflect positioning of fish near higher velocity areas to increase encounters with drifting prey (Tsao et al. 1998). Our study found Brook Trout selected LWD and rootwads, and used pools formed by LWD more than would be expected based upon availability. Both findings agree with observations of Salmo trutta (L.) (Brown Trout) in Wyoming streams (Young 1995). Our findings also support the idea of LWD being important to Brook Trout habitat. LWD has been shown to increase the number of pools (Hilderbrand et al. 1997), the depth of pools (Rosenfeld et al. 2000), and the complexity of pools (Harvey 1998). LWD also increases invertebrate production by providing habitat and accumulating organic storage (Angermeier and Karr 1984, Bisson et al. 1987, Cederholm et al. 1997). Given the results of these studies and the fact that novel fish appeared to behave as resident fishes, it was not surprising to find Brook Trout associated with LWD within the stream. We were unable to fully confirm our second hypothesis that Brook Trout habitat use would differ at different stream discharge levels. Although Oncorhynchus mykiss (Walbaum) (Rainbow Trout) selected different microhabitats during low and high flows (Vondracek and Longanecker 1993), we found no overall relationship between discharge level and pool use in Brook Trout. At extremes of stream discharge (very low or very high), we expect that Brook Trout would seek out pools as they provide refuge from low water levels and possible stranding during drought, as well as refuge from high water velocities during floods. Increases in discharge and velocity often cause fish to seek refuge within pools (Bisson et al. 1987, Bozek and Rahel 1991, Cederholm et al. 1997, Rosenfeld and Boss 2001). Over the range of discharge levels we observed in the study stream, there was no overall effect of discharge level upon percent pool use. Similar findings were reported by 368 Northeastern Naturalist Vol. 17, No. 3 Heggenes et al. (1991), who found no difference in habitat use by Brown Trout during increased flows. Only when looking at the lower 25th percentile of daily discharge measures did we detect a relationship between pool use and discharge. Interestingly, pool use by Brook Trout declined sharply with increasing discharge, but only when discharge was less than 0.095 m3/s. This relationship suggests that Brook Trout either sought out pools as refuge during low flows, perhaps to avoid stranding (Bradford et al. 1995), or rapidly moved to other habitats as flows increased from low levels. Such movements out of pools and into riffles and runs as flows increased slightly could be responses to redistribute and reduce competitive interactions with conspecifics (Young 2004) following confinement to pools during periods of low flow. Movement and home range Our observations on home range and movement suggest that Brook Trout in novel environments within headwater streams are comprised primarily of relatively sedentary individuals with some mobile fish. In our study, home range averaged 301 m (2003) to 637 m (2002), and cumulative movements averaged 937 m. Similar patterns and magnitudes of movement have been reported for Brook Trout in the southern Appalachians using tagging- recapture studies with resident fish. In the Great Smoky Mountains, most fish were recaptured within a 300-m reach after one year, although about 10% of all recaptures were 900 m or more from their original capture site (Moore et al. 1985). Whitworth and Strange (1983) reported that Brook Trout generally moved less than 300 m over bi-monthly periods in a Tennessee stream. However, each of these movement studies of Brook Trout may have underestimated the magnitude of movements. Gowan et al. (1994) criticized the use of tag-recapture data to evaluate movement of stream-dwelling trout, because reliance on such data ignores those individuals that may move larger distances outside the study area and are never recaptured. However, our study found comparable movement rates to the Moore et al. (1985) and Whitworth and Strange (1983) studies while using radio telemetry to establish movement rates, thereby generally confirming their findings with tag-recapture methods. Our movement measures of transplanted Brook Trout could under represent mobile individual life histories. The telemetry studies were conducted during spring; in these Appalachian streams, most energy acquisition and growth occurs during this time (Sweka and Hartman 2008, Utz and Hartman 2006). During other seasons, most fish in headwater sites are feeding below maintenance ration on a given day (Utz and Hartman 2006). Thus, there may be little motivation for individual Brook Trout to move to locate better resources during the time we monitored fish movements. At other times of the year, such as summer when lower trout densities but higher rations are available downstream in larger stream segments (Utz and Hartman 2006), we would expect fish movements to potentially be higher as some fish, particularly those in poorer condition (Gowan and Fausch 1996, Hilderbrand and Kershener 2004, Mogen and Kaeding 2005), may increase movement in 2010 K.J. Hartman and M. Nel Logan 369 an attempt to assess better feeding conditions. Despite this possible seasonal limitation, the telemetry results represent typical spring Brook Trout habitat use and movements in these headwater streams. One surprising aspect of this study was the large differences in primary habitat type measures between 2002 and 2003 despite our sampling the same reaches each year. These differences could have effects on assessment of habitat selection by trout. Pool areas available were relatively constant each year, but riffle availability declined and run availability increased from 2002 to 2003. We believe this switch in non-pool habitat reflects higher stream discharge levels in 2003, which elevated water levels and changed primary habitat classifications from riffles to runs in 2003. Regardless, novel Brook Trout continued to select for pools in higher proportion than available in each year, so differences in habitat classification on the basis of flow regimes would not affect the conclusions of this study. Fish size Fish size has been shown to play an important role in habitat selection, feeding, and reproduction in stream-dwelling salmonids (Everest 1972, Hutchings 1994, Luecke 1986). Interestingly, in 2003 when stream discharge was higher, we found fish length was negatively related to home range, cumulative movement, and daily movement. Most of this difference in movement between years was due to higher movement rates among smaller trout rather than larger movements among larger individuals. It is unclear what the mechanism might be behind this trend, but perhaps smaller adults are either more easily displaced during high discharge, or are better able to move across stream reaches under high flows. Acknowledgments The authors thank S.A. Welsh and J.T. Petty for technical advice throughout the study, J.A. Sweka and G. Seidell for statistical support, and M.K. Cox, J. Freund, M. Gamber, D. Hartman, J. Howell, T. Jenkins, and B. Lenz for help in field surveys. 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