Role of Stream Ice on Fall and Winter Movements and Habitat Use by Bull Trout and Cutthroat Trout in Montana Headwater Streams
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Transactions of the American Fisheries Society 127:223±235, 1998 q Copyright by the American Fisheries Society 1998 Role of Stream Ice on Fall and Winter Movements and Habitat Use by Bull Trout and Cutthroat Trout in Montana Headwater Streams MICHAEL J. JAKOBER1 AND THOMAS E. MCMAHON* Biology Department, Fish and Wildlife Program Montana State University, Bozeman, Montana 59717, USA RUSSELL F. T HUROW U.S. Forest Service, Rocky Mountain Research Station 316 East Myrtle Street, Boise, Idaho 83702, USA CHRISTOPHER G. CLANCY Montana Fish, Wildlife, and Parks 1801 North 1st Avenue, Hamilton, Montana 59840, USA Abstract.ÐWe used radiotelemetry and underwater observation to assess fall and winter move- ments and habitat use by bull trout Salvelinus con¯uentus and westslope cutthroat trout Onco- rhynchus clarki lewisi in two headwater streams in the Bitterroot River drainage, Montana, that varied markedly in habitat availability and stream ice conditions. Bull trout and cutthroat trout made extensive (.1 km) downstream overwintering movements with declining temperature in the fall. Most ®sh remained stationary for the remainder of the study (until late February), but some ®sh made additional downstream movements (1.1±1.7 km) in winter during a low-temperature (#18C) period marked by anchor ice formation. Winter movement was more extensive in the mid- elevation stream where frequent freezing and thawing led to variable surface ice cover and frequent supercooling (,08C). Habitat use of both species varied with availability; beaver ponds and pools with large woody debris were preferred in one stream, and pools with boulders were preferred in the other. Trout overwintered in beaver ponds in large (N 5 80±120), mixed aggregations. In both streams, both species decreased use of submerged cover following the formation of surface ice. Our results indicate that (1) continued activity by trout during winter is common in streams with dynamic ice conditions and (2) complex mixes of habitat are needed to provide suitable fall and winter habitat for these species. Populations of bull trout Salvelinus con¯uentus period is considered limiting to salmonids (Cunjak and westslope cutthroat trout Oncorhynchus clarki 1996). Movement of stream-resident salmonids is lewisi have declined substantially throughout their especially common in the fall as ®sh seek suitable native ranges (Liknes and Graham 1988; Rieman overwintering sites in response to declining water and McIntyre 1993). Though migratory (¯uvial temperatures (Bjornn 1971; Cunjak 1996) and suit- and ad¯uvial) life history forms were formerly able spawning sites in the case of fall-spawning common, as a result of fragmentation and loss of salmonids like bull trout (Elle 1995). Extent of habitat and of competition and hybridization with movements may vary widely, from 0 to more than nonnative salmonids, many remaining populations 100 km, depending on life history form and habitat persist as nonmigratory ``resident'' forms con®ned availability (Bjornn and Mallet 1964; West et al. to headwaters (Ziller 1992; Rieman and McIntyre 1992; Brown and Mackay 1995). 1993). Little is known about movement patterns Several investigators have hypothesized that and habitat preferences of these headwater popu- subsurface ice formation and groundwater in¯ux lations. are major factors determining habitat suitability We focused on de®ning the fall and winter and carrying capacity for salmonids in many ice- movement and habitat use in headwater popula- covered streams (Maciolek and Needham 1952; tions of bull trout and cutthroat trout because this West et al. 1992; Brown and Mackay 1995; Cunjak 1996). However, this hypothesis remains largely * Corresponding author: [email protected] untested since most winter studies have focused 1 Present address: Bitterroot National Forest, 88 Main on streams that remain relatively ice-free. In the Street, Stevensville, Montana 59870, USA. ice-covered Ram River, Alberta, Brown and Mack- 223 224 JAKOBER ET AL. ay (1995) reported that some cutthroat trout un- Creek, a major tributary to the Bitterroot River derwent complex, two-stage movements during near Hamilton, Montana. The average gradient in fall and winter. During ``®rst-stage'' movements the study area was 4.1% and mean elevation was in mid-September, ®sh moved in response to de- 1,424 m. Rif¯es and glides were the predominant clining temperature up to 3 km to overwinter hab- habitat type (77% of surface area), and LWD was itats of deep pools with cover of woody debris or about 50% less abundant than in Meadow Creek undercut banks. Later in the fall, subsurface (frazil (67 pieces/100 m). No beaver ponds were present, and anchor) ice formed in these sites, adhering to and substrate was primarily boulders (250/100 m) cover and ®lling pools, which triggered ``second- and cobble. Surface ice ¯uctuated markedly during stage'' movement by trout for several kilometers the winter in response to frequent freezing and to groundwater-fed sites and other areas lacking thawing events, and anchor and frazil ice were subsurface ice. Winter severity for salmonids may common. be determined more by degree of subsurface ice In each stream we established a 2-km study sec- formation rather than by temperature (Chisholm et tion located near the lower limit of bull trout dis- al. 1987; Berg 1994; Brown et al. 1994). Thus, the tribution. Cutthroat trout and brown trout Salmo harshest winter conditions may occur where in- trutta were the predominant salmonids down- complete surface ice cover results in extensive an- stream. There were no barriers to movement for chor and frazil ice formation (Maciolek and Need- at least 6 km upstream from study reaches. In ham 1952; Chisholm et al. 1987). Meadow Creek, ®sh had unimpeded access to the In the Bitterroot River drainage, Montana, bull mainstem Bitterroot River 16 km downstream; in trout and cutthroat trout persist in two types of Daly Creek, a low-head irrigation dam and de- low-order streams that differ markedly in habitat watering 15 km downstream created a seasonal availability and winter ice conditions. Small, high- barrier to migration. elevation streams are characterized by abundant large woody debris (LWD) and extensive surface Methods ice in winter; moderate-sized, mid-elevation Movements and habitat use were assessed by a streams are characterized by ¯uctuating surface combination of radiotelemetry and underwater ob- and subsurface ice and boulder±cobble substrate. servation to balance the advantages and limitations To better de®ne critical habitat needs to aid pro- of each sampling approach (Alldredge and Ratti tection and recovery of these species, we examined 1986). seasonal habitat use and movement, particularly Radiotelemetry.ÐWe used radio transmitters to the timing and magnitude of movements in re- monitor movements of 18 bull trout and 6 cutthroat sponse to differing stream ice conditions. trout from late August 1992 through February 1993. Because ®sh from these headwater popula- Study Area tions typically were less than 300 mm in total We examined two streams that typify sites where length, we were limited to small transmitters with both species persist in headwater populations. 90-d battery lives. We therefore monitored move- Meadow Creek was a high elevation, third-order ments of two sets of radio-tagged ®sh during two tributary to the East Fork of the Bitterroot River different time periods. To assess spawning move- near Sula, Montana. Wetted width in the study area ment of bull trout, only this species was radio- averaged 2±3 m at base ¯ow, mean elevation was tracked during the fall phase (28 August±13 No- 1,818 m, and average gradient was 2.1%. Large vember), whereas both species were tracked during woody debris (.10 cm in diameter and .2min the winter phase (31 October±24 February). length) was abundant (116 pieces/100 m) and sub- We captured ®sh by electro®shing within the 2- strate was primarily gravel and cobble with few km study reaches. Four bull trout (mean total boulders (23/100 m; Jakober 1995). Pools and length, 282 mm; range, 242±434 mm) were radio- beaver ponds were the most common habitat types tagged in each stream in late August 1992 to assess (36% and 40%, respectively, of the total surface fall movements. In late October 1992, ten more area). The stream was almost entirely covered with bull trout (four in Meadow Creek and six in Daly surface ice from early November through late Creek; mean length, 281 mm; range, 231±317 mm) March. and six cutthroat trout (four in Meadow Creek and Daly Creek, located about 33 km north of Mead- two in Daly Creek; mean length, 273; range, 234± ow Creek, was a moderate-sized (5±7 m wide), 330 mm) were radio-tagged to assess winter move- mid-elevation, fourth-order tributary to Skalkaho ments. ROLE OF STREAM ICE ON TROUT MOVEMENT 225 Fish were anesthetized with tricaine methane- release and ®nal location (net distance). Movement sulfonate (MS-222), weighed, and measured, and data were grouped as the mean total movement of radio transmitters were surgically implanted via a all relocations per week. We compared total and 1±2-cm incision in the peritoneal cavity immedi- net distances moved among seasons by using a t- ately anterior to the pelvic girdle. Fish were also test and determined the proportion of ®sh that tagged with colored Floy tags. Surgery lasted moved more than 100 m with a x2 test (Zar 1984). about 10 min, and no mortalities occurred during Bull trout were recorded as having spawned if they surgery or the 1-h recovery period prior to release were observed spawning or near a known redd site. near the capture site. Habitat was classi®ed according to type, as mod- Transmitters (dimensions, 40 3 13 mm; Custom i®ed from Bisson et al. (1988): rif¯e, glide, pool Telemetry, Athens, Georgia) had an internal loop with LWD, pool with boulders, pool lacking LWD antenna and emitted a unique frequency from 40.4 or boulders, pocket water (small pools formed by to 40.5 mHz.