Western North American Naturalist Volume 60 Number 1 Article 2 1-20-2000 Home-range fidelity and use of historic habitat by adult Colorado pikeminnow (Ptychocheilus lucius) in the White River, Colorado and Utah David B. Irving U.S. Fish and Wildlife Service, Fish and Wildlife Management Assistance Office, Roosevelt, Utah Timothy Modde U.S. Fish and Wildlife Service, Colorado River Fishery Project, Vernal, Utah Follow this and additional works at: https://scholarsarchive.byu.edu/wnan Recommended Citation Irving, David B. and Modde, Timothy (2000) "Home-range fidelity and use of historic habitat by adult Colorado pikeminnow (Ptychocheilus lucius) in the White River, Colorado and Utah," Western North American Naturalist: Vol. 60 : No. 1, Article 2. Available at: https://scholarsarchive.byu.edu/wnan/vol60/iss1/2 This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Western North American Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact scholarsarchive@byu.edu, ellen_amatangelo@byu.edu.
Western North American Naturalist 60(1), 2000, pp. 16 25 HOME-RANGE FIDELITY AND USE OF HISTORIC HABITAT BY ADULT COLORADO PIKEMINNOW (PTYCHOCHEILUS LUCIUS) IN THE WHITE RIVER, COLORADO AND UTAH David B. Irving 1 and Timothy Modde 2 ABSTRACT. Twelve wild adult Colorado pikeminnow (Ptychocheilus lucius), captured in the tailwaters of Taylor Draw Dam on the White River, Colorado, were implanted with radio transmitters and their movement patterns monitored from 1992 to 1994. The spawning migration of these fish was extensive. In 1993, the only full year of the study, the fish migrated an average of 658 km from the White River to spawning sites in the Yampa or Green rivers and back to the White River. Eight of these fish were translocated in the river upstream of the dam in April 1993. These fish and the 4 others below the dam remained in the river until May 1993. All 12 had migrated down the White River to spawning sites in the Green and Yampa rivers by July 1993. The fish that were located above the dam successfully passed over the dam during their downstream migration. Seven fish migrated upstream toward the Yampa River Canyon spawning site and 5 migrated downstream toward the Green River Desolation/Gray Canyon spawning site. Five of 7 Yampa River fish were found at the spawning site. The other 2 were found 5 8 km downstream of the site. One of 5 Green River fish was found at the spawning site, the other 4 between 16 and 62 km upstream of the site. All fish migrated back to the White River by August 1993 and were found near the dam by October 1993. Two fish were recaptured and translocated above the dam in September 1993. Five fish were located below the dam and 2 above the dam in April 1994. By July 1994 seven of the same fish that had migrated toward the Yampa River in 1993 were found at the Yampa Canyon spawning site. At the same time, 3 of 5 fish that migrated toward the Green River in 1993 were found at the Desolation/Gray Canyon spawning site. This included 2 fish that had been found upstream of the site in 1993. The 12 fish traveled an average of 6 km d 1 (range: 4 10 km d 1 ) during the migration period from May through October 1993. Generally, fish moved faster to the spawning site than back from the site to the White River. These fish moved very little within their home ranges in the White River. Six fish tagged in 1992 moved only 0.1 2.3 km in the tailwater reach below Taylor Draw Dam from September 1992 through April 1993. All fish, after their spawning runs, had moved up to or near the dam by October 1993. These fish were not tracked again until April 1994. Their movement patterns in April 1994 were similar to those observed in April 1993. The greatest amount of fish movement in the White River was displayed by the 8 fish placed above Taylor Draw Dam in April 1993 and the 2 placed in Kenney Reservoir in September 1993. They moved 1.1 40.6 km in the river before and after their spawning migration in spring and autumn 1993. These spawning migrations suggest that adult Colorado pikeminnow in the White River were recruited from both Green and Yampa river spawning populations and were presumably imprinted to these respective spawning sites. Those fish placed above Taylor Draw Dam established home ranges in habitats previously occupied by Colorado pikeminnow before the dam was completed. They remained there until they migrated downstream during the spawning period. Although we did not study fish passage, our study demonstrates that adult Colorado pikeminnow will use habitat if access is provided. Translocation of wild adult fish into historic but unoccupied habitats may be a valuable recovery option. Key words: Colorado pikeminnow, Ptychocheilus lucius, migration, telemetry, home-range fidelity. Colorado pikeminnow, Ptychocheilus lucius, is a large, warmwater cyprinid endemic to the Colorado River basin of western United States and Mexico (Jordan and Everman 1896, Minckley 1973). Once widely distributed in the main channel and major tributaries, the long-lived species has been extirpated from 80% of its historic range following the construction of mainstem impoundments and establishment of nonnative predators (Tyus 1991). Colorado pikeminnow was listed as endangered by the U.S. Fish and Wildlife Service in 1967 and given protection under the Endangered Species Act in 1974 (Federal Register 39[3]:1175). Selfsustaining populations exist only in the upper Colorado River basin, with greatest numbers in the Green River subbasin (Tyus 1991). As with several other large-river fishes in the American Southwest, the Colorado pikeminnow is potamodromous, with spawning migrations 1 U.S. Fish and Wildlife Service, Fish and Wildlife Management Assistance Office, 855 East 200 North (112 13), Roosevelt, UT 84066. 2 U.S. Fish and Wildlife Service, Colorado River Fishery Project, 266 West 100 North Suite 2, Vernal, UT 84078. 16
2000] COLORADOPIKEMINNOW IN WHITE RIVER 17 initiated by changes in discharge, temperature, and photoperiod (Tyus 1986, 1990). Using an extensive telemetry and mark-recapture data set, Tyus (1990) described factors initiating migration and spawning of Colorado pikeminnow in the Green River subbasin, including the White River. Some fish migrated 1-way distances >300 km to 1 of 2 known spawning areas (Yampa Canyon in Yampa River or Gray/ Desolation Canyon in Green River), and individuals were captured on the same spawning sites in multiple years. Of 153 fish implanted with radio transmitters, 41% migrated 1 times to 1 of the 2 known spawning areas, with an additional 11% suspected of doing so (Tyus 1990). Spawning migration was not detected in approximately half the fishes implanted with radio transmitters. Lack of movement was presumably because fish were either immature or nonannual spawners (Tyus 1990). Ryden and Ahlm (1996) found similar behavior in Colorado pikeminnow radio-tracked over a 3.5-yr period on the San Juan River in New Mexico, Colorado, and Utah. Movement of 12 of 13 fish they studied averaged 17.7 km (range: 1.8 32.8 km). The other fish moved a total of 93.0 km and was the only one thought to display migratory behavior. Although factors initiating spawning migrations and spawning-site fidelity have been well described (Tyus 1986, 1990), movement patterns of individual fish in consecutive years are not well understood. Previous studies have found no early life stages and few juvenile Colorado pikeminnow in the White River; most fish in the river are adults (Tyus 1986). Taylor Draw Dam, constructed in 1985 on the White River, created a barrier to upstream movement, preventing access to about 32% (77.8 km) of the habitat in the White River historically used by adult Colorado pikeminnow (Carlson et al. 1979, Wick et al. 1985, Trammell et al. 1993). It is assumed that after closure of the dam those Colorado pikeminnow upstream of the dam migrated over the dam to downstream spawning areas. Post-spawning fish moving back upstream were blocked from returning to their previously occupied home range above Taylor Draw Dam. As a result, fish congregated below the dam in densities up to 37 adults per 0.4 km as recently as 1993 (Irving and Modde 1994). These densities may also be the result of recent high recruitment of Colorado pikeminnow throughout the Green River basin (McAda et al. 1998). This study examined migratory movements of 12 adult Colorado pikeminnow in the White River through 2 successive spawning periods and determined where these fish spawned. It also tested whether some of these fish that were translocated into formerly occupied habitats above Taylor Draw Dam would remain in that habitat, move into the reservoir, or return to home ranges below the dam. STUDY AREA The White River drainage encompasses 1.3 million ha of arid pinion-juniper and sagebrush desert in northwestern Colorado and northeastern Utah (Fig. 1). The river drains into the Green River, a major tributary to the Colorado River in southeastern Utah. The high-gradient, cool-headwater, canyon-bound reaches consist of riffles, runs, and rapids with boulder, cobble, and gravel substrates. The low-gradient warmwater reaches are characterized by deep eddies, pools, and runs that meander through slower, turbid waters, with vegetated shorelines and gravel, sand, and silt substrates. Summer water temperatures often reach 20 C. Peak spring discharge ranges between 100 and 170 m 3 s 1. Taylor Draw Dam is operated as a run-of-the river facility and provides water storage, flood control, and hydroelectric power. Kenney Reservoir, above the dam, is 275 ha and provides recreational boating and fishing. MATERIALS AND METHODS We captured 12 wild adult Colorado pikeminnow by electrofishing and trammel-netting in a 0.5-km reach (km 163 168) of the White River below Taylor Draw Dam. Six fish (#1 6) were caught in September 1992 and 6 more (#7 12) in April 1993. Each fish was measured for total length (TL), tagged with a PIT (passive integrated transponder), and surgically implanted with a 24-month (16-g) radio transmitter. Fish were anesthetized with tricaine methanesulfonate and radio transmitters (internal loop antennas) placed into the body cavity via a surgical incision as quickly as possible following capture. Fish were released within 10 min after surgery. Eight fish, 2 recaptured from the 1992 group and 6 from the 1993 group, were translocated 30 km upstream of the dam to determine if fish would remain in habitats previously occupied
18 WESTERN NORTH AMERICAN NATURALIST [Volume 60 Fig. 1. Radio telemetry study area for 12 wild adult Colorado pikeminnow (Ptychocheilus lucius) tracked on the White, Green, and Yampa rivers, upper Colorado River basin, Colorado and Utah, 1992 1994. by Colorado pikeminnow (historic habitat) before dam completion. We also translocated 2 fish recaptured in September 1993, 1 from the 1992 group and 1 from the 1993 group, into the lower reach of Kenney Reservoir (km 169) to determine if they would remain in the reservoir, pass through the reservoir, or access upstream habitats. Locations of all 12 fish were monitored on the ground using a radio search-receiver with bidirectional paddle antennas and from the air with fixed-wing aircraft with omni-directional loop antennas. Fish tracked on the ground were triangulated to the nearest 0.25 m; fish locations tracked by air were estimated to the nearest 0.4 km (based on transmitter pulse rate, signal
2000] COLORADOPIKEMINNOW IN WHITE RIVER 19 strength, and aircraft speed). From September 1992 to April 1993 (before placing any upstream of the dam), we monitored fish locations monthly, and approximately weekly between 15 April and 30 September 1993. Fish locations during 1994 were monitored in late April and on 2 dates during the migration and spawning period (1 and 13 July). Although radio-tracking was not monitored on a continuous basis, travel rates (km d 1 ) were calculated for each fish for the 1993 migration period. RESULTS Twelve Colorado pikeminnow (409 743 mm TL, 1000 3750 g) were captured in the 0.4-km tailwater reach below Taylor Draw Dam (km 168.2) in 1992 and 1993 (Table 1). The 6 fish (#1 6) captured and implanted with radio transmitters in September 1992 remained in the White River within 1 km below Taylor Draw Dam through fall 1992 and spring 1993 (Fig. 2). The 6 fish (#7 12) captured in April 1993 were located within a 0.8-km reach below the dam. It was assumed that all fish survived because each transmitter remained active throughout the 2-yr monitoring period. The spawning migration of these fish was extensive. Colorado pikeminnow tagged in the White River migrated an average of 658 km from the White River to spawning sites in the Yampa or Green rivers and back to the White River (Table 1, Fig. 2). All 12 radio-tagged fish migrated downstream in the White River in 1993 and entered the Green River. Seven of these fish moved upstream in the Green River, 5 fish (#1, 3, 8, 9, 10) into the Yampa River spawning area (Yampa Canyon km 0 32), and 2 (#2, 7) located 11 and 16 km downstream of the spawning area (Fig. 2). These 7 fish migrated between 548 and 951 km from mid- May through late October 1993. During the same year 1 fish (#12) moved downstream in the Green River and was located at the Green River spawning site (Gray/Desolation Canyon km 232 256). Four other fish (#4, 5, 6, 11) were found 27 100 km upstream of the spawning area (Fig. 2). The Green River migrants traveled between 437 and 687 km from mid- May through late September 1993 (Table 1). All fish returned to the White River between mid-august and late October 1993, thus showing home-range fidelity. The 8 fish (#1, 4, 7, 8, 9, 10, 11, 12) relocated in the White River (km 198.2) above Kenney Reservoir in the spring of 1993 remained in historic riverine habitats above the reservoir until initiation of the downstream spawning migration (Fig. 3). One fish (#1) moved downstream on May 15 (Table 1), the others between mid-june and early July (Fig. 2). On 9 and 14 September 1993, two fish (#4, 9, respectively) were relocated to the lower end of Kenney Reservoir (km 170.6). These fish moved upstream of the reservoir (Fig. 3) within 8 d and were located in historical riverine habitats the following spring (20 April 1994). Five fish (#1, 3, 6, 10, 12) were located just below Taylor Draw Dam (km 167.4 168.2) and 2 fish (#4, 9) above the dam (km 180.2 185.1) the following year on 20 April 1994 (Fig. 2). The 5 fish below the dam, located 3 15 km downstream of the dam the previous October, moved back upstream to the dam in spring 1994. The 2 fish relocated into Kenney Reservoir in September 1993 moved an additional 5 8 km further upstream. On 1 and 13 July 1994 seven fish (#1, 2, 3, 7, 8, 9, 10) located in April were found within the Yampa River spawning reach: three (#5, 6, 12) were found at the Green River spawning site, and two (#4, 11) were found between 43 and 105 km upstream of the Green River site. All fish moved to or near the same spawning area they used the previous year. The 12 fish traveled an average of 6 km d 1 (range: 4 10 km d 1 ) during the migration period from May through October 1993 (Table 1). Generally, fish moved faster to the spawning site than back to the White River. Aside from their spawning migration, these fish moved very little within their home ranges in the White River. For example, the 6 fish tagged in 1992 moved only 0.1 2.3 km in the tailwater reach below Taylor Draw Dam from September 1992 through April 1993. All 12 fish, after migrating to and from their respective spawning sites, migrated back to the White River by late August and September 1993. Five of these fish (#1, 4, 6, 9, 11) moved back upstream to Taylor Draw Dam and then redistributed themselves downstream 2.4 10.1 km by October 1993. The other 7 fish (#2, 3, 5, 7, 8, 10, 12) did not appear to move up to
20 WESTERN NORTH AMERICAN NATURALIST [Volume 60 TABLE 1. Length, weight, and telemetry data of 12 radio-tagged wild adult Colorado pikeminnow in the White, Green, and Yampa rivers, upper Colorado River basin, Colorado and Utah, 1992 1994. (Fish number refers to order in which fish was captured). 1992 1993 1994 Fish Length Weight Date monitored No. days No. km Date monitored No. days No. km Dates migrated No. days No. km Rate Date monitored No. days No. km No. a (mm) (g) Start End tracked traveled Start End tracked traveled Start End migrated traveled (km d 1 ) Start End tracked traveled 1 647 9/30 11/30 3 0.1 1/6 10/29 20 755.9 5/15 8/31 108 720.3 6.7 4/20 7/13 3 339.7 2 633 9/30 11/30 3 0.0 1/6 9/22 8 631.0 5/19 8/31 104 548.1 5.3 7/1 7/13 2 525.6 3 409 10/1 11/30 3 0.1 1/6 10/29 13 722.4 5/19 10/29 163 722.4 4.4 4/20 7/13 3 395.7 7 484 1000 4/9 10/29 12 678.4 5/15 10/29 167 646.4 3.9 7/1 7/13 2 381.1 8 743 3750 4/12 9/22 9 985.1 6/15 9/22 99 951.4 9.6 7/1 7/13 2 419.7 9 654 2750 4/14 9/22 18 787.5 6/15 9/8 85 710.7 8.4 4/20 7/13 3 348.9 10 604 4/26 9/22 11 768.2 6/15 9/22 99 710.7 7.2 4/20 7/13 3 354.2 n 7 3 3 3 3 3 7 7 7 7 7 7 7 7 7 7 7 7 7 Min 409 1000 9/30 11/30 3 0 1/6 9/22 8 631 5/15 8/31 85 548.1 6.4 4/20 7/13 2 339.7 Mean 596.3 2500 9/30 11/30 3.0 0.1 3/3 10/7 13.0 761.2 5/29 9/24 117.9 715.7 6.1 5/20 7/13 2.6 395.0 Max 743 3750 10/1 11/30 3 0.1 4/26 10/29 20 985.1 6/15 10/29 167 951.4 5.7 7/1 7/13 3 525.6 Std 112.9 1391.9 4 632 10/1 11/30 3 0.1 1/6 9/22 17 679.0 5/15 8/16 93 635.0 6.8 4/20 7/13 3 313.8 5 451 10/1 10/15 2 0.1 1/6 9/22 12 541.4 5/19 9/22 126 541.2 4.3 7/13 7/13 1 304.2 6 430 10/2 10/15 2 0.1 2/3 10/29 13 447.4 5/19 8/31 104 437.1 4.2 4/20 7/13 3 315.9 11 624 1960 4/28 10/29 15 642.5 6/15 9/8 85 589.4 6.9 7/1 7/13 2 251.9 12 652 2100 4/28 10/29 11 733.5 5/27 8/31 96 686.5 7.2 4/20 7/13 3 315.9 n 5 2 3 3 3 3 5 5 5 5 5 5 5 5 5 5 5 5 5 Min 430 1960 10/1 10/15 2 0.1 1/6 9/22 11 447.4 5/15 8/16 85 437.1 5.1 4/20 7/13 1 251.9 Mean 557.8 2030 10/1 10/30 2.3 0.1 2/25 10/14 13.6 608.8 5/25 9/3 100.8 577.8 5.7 5/21 7/13 2.4 300.3 Max 652 2100 10/2 11/30 3 0.1 4/28 10/29 17 733.5 6/15 9/22 126 686.5 5.4 7/13 7/13 3 315.9 Std 107.8 99 a Fish 1, 2, 3, 7, 8, 9, and 10 migrated to or near the Yampa Canyon Yampa River spawning site (km 0 32). Fish 4, 5, 6, 11, and 12 are those that migrated to or near the Grey/Desolation Canyon Green River spawning site (km 232 256). Fish 1, 4, 7, 8, 9, 10, 11, and 12 are the 8 fish translocated in the White River 30 km upstream of Taylor Draw Dam. Fish 4 and 9 were translocated into Kenney Reservoir 4 km upstream of Taylor Draw Dam.
2000] COLORADOPIKEMINNOW IN WHITE RIVER 21 Fig. 2. Movement patterns of 12 radio-tagged wild adult Colorado pikeminnow (Ptychocheilus lucius) that migrated from the White River to spawning sites in the Yampa River (7 fish, Yampa Canyon at km 0 32) and Green River (5 fish, Gray/Desolation Canyon at km 232 256), upper Colorado River basin, Colorado and Utah, 1992 1994. (Larval drift dates are adapted from Bestgen et al. 1998.)
22 WESTERN NORTH AMERICAN NATURALIST [Volume 60 Fig. 3. Locations of 12 radio-tagged Colorado pikeminnow (Ptychocheilus lucius) translocated above Taylor Draw Dam and subsequently contacted in Kenney Reservoir and the White River, Colorado, above Taylor Draw Dam (km 168). the dam but had located themselves 5.6 26.6 km below the dam by October 1993. Fish movement in September and October 1993 was 3- to 6-fold greater than the movement exhibited by these fish during the same period in 1992. It is unknown why these fish moved downstream of Taylor Draw Dam in 1993 when they congregated below the dam during the same period in 1992. The average discharge at the dam during the months of August, September, and October was more than 1.5 times greater in 1993 than in 1992. In addition, a new hydroelectric generator was installed and in operation by summer 1993 and caused some redirection and fluctuations of flows in the river channel directly below the dam. Although these fish were not tracked again until April 1994, they showed movement patterns similar to those in April 1993. The greatest amount of fish movement in the White River was displayed by the 8 fish placed above Taylor Draw Dam in April 1993 and the 2 fish placed in Kenney Reservoir in September 1993. The 8 fish moved 1.1 40.6 km between April and July 1993; the other 2 fish, which overwintered in the river above the dam, moved 10.5 19.3 km between September 1993 and April 1994. DISCUSSION Colorado pikeminnow in the White River moved long distances annually. All 12 Colorado pikeminnow monitored from 1992 to 1994 migrated from the White River and were located at or near the spawning sites, or apparently en route to the sites in the Green and Yampa rivers. They then returned to areas near their original capture sites below Taylor Draw Dam. All demonstrated movements to spawning areas previously described by Tyus (1990): 7 migrated toward the Yampa Canyon spawning site and 5 toward the Gray/Desolation Canyon site. They also displayed fidelity to a single site and migrated the same direction in successive years. Our observations indicate that many wild adult Colorado pikeminnow in the White River undergo annual migrations to spawning sites. It is unknown whether these fish actually spawn each year. Although we contacted all migrating fish within a few kilometers of the known spawning sites, telemetry was limited, and we were unable to locate each fish at its respective site. Either they reached the sites between our contacts or they might have migrated to other concentration points near the spawning sites.
2000] COLORADOPIKEMINNOW IN WHITE RIVER 23 This study showed 5 of 7 Yampa River fish were within the spawning area in 1993 and 1994. In 1994 one of the 2 fish found just below the site in 1993 was found on the site and the other just above the site. One of 5 Green River fish was at the site in 1993 and 3 were there in 1994. Of the 4 fish located above the site in 1993, two were at the site in 1994 and the remaining 2 fish above the site in both years. The only way to confirm that there are more spawning sites in the upper Green River drainage is to tag more fish and follow their migration patterns more closely over successive years. Presumably, 100% of the radio-tagged Colorado pikeminnow survived over the 2-yr study. This rate is higher than the 85% annual survival rate reported by Osmundson et al. (1997) for adult Colorado pikeminnow in the upper Colorado River. Our study confirms and expands the findings of other researchers. From 1980 through 1990, radio telemetry detected 37 Colorado pikeminnow moving in and out of the White River (Tyus et al. 1981, Tyus 1990, Trammell et al. 1993). Twenty-three of these pikeminnow remained in the White River, or at least were there on all monitoring dates. However, some were not contacted each time, and several months sometimes elapsed between tracking dates. It is thus possible that individuals migrated from the White River to spawning sites in the Green or Yampa rivers and returned undetected. Of the remaining 14 telemetered pikeminnow, 9 moved to the Yampa River spawning area and 5 to the Gray/Desolation Canyon spawning area in the Green River. Only 2 of the 14 are known to have returned to the White River. All 12 of the Colorado pikeminnow we studied exhibited migratory behavior in 1993. This is in contrast with previous studies in the Green River where about 50% were nonmigratory. This suggests that White River stocks are composed entirely of reproductively active individuals that utilize the White River exclusively for adult habitat. The long-distance migrations of these fish to spawning areas in the Green and Yampa rivers suggest that adult habitats may be limiting for such an energetically costly mechanism to have evolved. In addition, if natal imprinting is important in homing to spawning destinations (Tyus 1990), the White River population of Colorado pikeminnow represents a mixed stock recruited from both upstream and downstream spawning populations. Tyus (1985, 1989, 1990) noted that pikeminnow in the middle Green and Yampa rivers tended to spawn at the Yampa River spawning site, and fish from the lower Green River (i.e., Desolation and Gray canyons) tended to migrate to the Gray Canyon spawning site. Although the White River confluence is located approximately equidistant from both spawning sites (confluence Yampa River site 159 km, confluence Green River site 140 km), it is much nearer to known nursery areas upstream of the White River (Irving and Burdick 1995, McAda et al. 1998). Thus, it is surprising that about half the fish studied by us and others utilized the Gray Canyon site. This suggests that occupation by subadult/adult fish moving upstream, not the presence of nursery areas for juvenile fish, is the mechanism for colonizing the habitat in the White River. Presumably, juvenile fish may move upstream in the mainstem Green River for some time before exploring tributary streams. Spawning migration dates of this study also match well with Colorado pikeminnow larval drift dates calculated by Bestgen et al. (1998). He found that larvae drifted downstream of both spawning sites between late June and mid-august 1993 and between mid-june and late July 1994 (Fig. 2). This study found that most tagged Colorado pikeminnow in the White River began their migration to the Yampa and Green river sites by mid-may or mid- June and then migrated back to the White River by mid- to late August (Table 1). Eight fish relocated upstream of Kenney Reservoir remained there 2 3 months before migrating downstream. Two fish relocated just above the dam in Kenney Reservoir in September 1993 moved through the reservoir to historic riverine habitats above the reservoir where they overwintered before migrating downstream to spawning areas the following summer. Adult Colorado pikeminnow we studied responded differently from hatchery-reared juvenile and adult pikeminnow previously stocked into Kenney Reservoir. Trammell et al. (1993) stocked 96,597 juvenile Colorado pikeminnow into the reservoir between April 1988 and September 1990, but none remained in the impoundment following stocking. Results from 4 hatchery-reared adult fish implanted with
24 WESTERN NORTH AMERICAN NATURALIST [Volume 60 transmitters (Trammell et al. 1993) were inconclusive: 3 died within 36 d, and the survivor, found in the impoundment on 12 June, moved downstream of the reservoir and was located 12 km below the dam on 9 August 1990. To date, no hatchery-reared fish have displayed the type of spawning migration documented for the White River, nor have they utilized unoccupied habitats upstream of Kenney Reservoir. However, behavior of wild fish was similar to our results. One of 3 wild adults implanted with transmitters died within 8 d, but the remaining 2 behaved similarly to the fish we studied: 1 moved upstream of the reservoir to occupy riverine habitats and the other moved downstream over the reservoir spillway. Contact was lost with the latter fish from 12 June to 11 September; thus, it could have migrated to a spawning area during this time. Apparently, there is a net upstream movement of subadult fishes into preferred habitats, but post-spawning Colorado pikeminnow often exhibit home-range fidelity by returning to the area (i.e., home range) they occupied before migration (Tyus 1986, 1990). Because of these behaviors, efforts to restore access (i.e., fish ladders, etc.) of adult fish to historic habitats may not be productive. Instead, younger fish would slowly colonize as they mature, thus increasing the time necessary to occupy restored habitats. However, our study demonstrates that adult fish will use habitats if access is provided them. Migration and habitat use of White River fish indicate that powerful selection mechanisms have developed over perhaps thousands of years of evolution. This is evident in migration patterns and habitat use. When provided access, wild adult Colorado pikeminnow utilized historic, unoccupied habitats rather than returning to sites below the dam where they had been restricted following closure of Taylor Draw Dam. This assumes that these study fish and other wild fish present before the dam was built have been attempting to ascend the White River since dam completion in 1985. On the other hand, hatchery-reared fish exhibited a different behavior. They did not show the same tendency to occupy riverine habitats upstream of Kenney Reservoir nor undertake such migrations. If restoration efforts connect occupied habitats with historic reaches via fish passages, wild adult fishes may access historic habitats, establish new home ranges, and continue successful reproduction by using highquality habitats. However, hatchery-reared fishes may not. This study suggests that wild adult Colorado pikeminnow might use a fish passage facility if it were in place at Taylor Draw Dam. Research conducted by Burdick and Pfeifer (1999) shows that Colorado pikeminnow will use the fish passage structure at Redlands Diversion Dam on the Gunnison River near Grand Junction, Colorado, to access riverine habitats upstream of this 12-ft dam. Since 1996, when the fish passage facility was opened, 47 subadult/adult (TL 383 763 mm) Colorado pikeminnow have passed upstream through this structure. Six fish that used the structure in July and August 1997 and 1998 successfully passed downstream over the dam after that date and then used the passage structure again in either 1998 or 1999. One fish has been found upstream as far as 49 km. This information can help guide present recovery efforts in areas where historic habitats have been blocked. Further, translocation of wild fish offers another feasible alternative to stocking hatchery-reared fish whose behavior may be problematic, such as not being imprinted to a successful spawning area nor being able to congregate with juvenile fish reared in high-quality nursery habitats. Finally, recovery efforts can be more successful if lifehistory needs of Colorado pikeminnow are better understood in areas where fish are most abundant and least disturbed. ACKNOWLEDGMENTS This study was initially proposed by the Colorado Division of Wildlife. We thank T. Nesler and B. Elmblad for their assistance in developing the study. A. Brady, Rio Blanco Water Conservation District, and M. Caddy, Colorado Division of Wildlife, provided local assistance in data collection and landowner permission. Assistance in field data collection was provided by H. Husband, B. Hilbert, D. Beers, J. Baker, Q. Bradwich, B. Sheffer, R. Arment, and H. Hines. We are especially grateful for thorough and thoughtful reviews by H. Tyus, K. Bestgen, M. Trammell, and K. Irving. Finally, we wish to thank Dinosaur National Monument of the U.S. National Park Service, Ouray National Wildlife Refuge of the U.S.
2000] COLORADOPIKEMINNOW IN WHITE RIVER 25 Fish and Wildlife Service, and Utah Division of Wildlife Resources for permits and permission to collect fish in and fly over their respective areas. This study was supported by the Recovery Implementation Program for the Recovery of Endangered Fish in the Upper Colorado River Basin. LITERATURE CITED BESTGEN, K.R., R.T. MUTH, AND M.A. TRAMMELL. 1998. Downstream transport of Colorado squawfish larvae in the Green River drainage: temporal and spatial variation in abundance and relationships with juvenile recruitment. Final report to Colorado River Recovery Implementation Program, Project 32. Larval Fish Laboratory, Department of Fishery and Wildlife Biology, Colorado State University, Fort Collins. BURDICK, B.D., AND F.K. PFEIFER. 1999. Evaluation of the effectiveness of the fish passage structure at Redlands Diversion Dam on the Lower Gunnison River. Colorado River Fishery Project, Grand Junction, Colorado. U.S. Fish and Wildlife Service. Recovery Implementation Program for the Endangered Fishes of the Upper Colorado River Basin. CARLSON, C.A., C.G. PREWITT, D.E. SNYDER, E.J. WICK, E.L. AMES, AND W.D. FONK. 1979. Fishes and macroinvertebrates of the White and Yampa rivers, Colorado. U.S. Bureau of Land Management. Biological Science Series 1, Denver, CO. IRVING, D.B., AND B.D. BURDICK. 1995. Reconnaissance inventory and prioritization of existing and potential bottomlands in the upper Colorado River basin, 1993 1994. Final report submitted to the Recovery Implementation Program for the Endangered Fish Species in the Upper Colorado Basin, U.S. Fish and Wildlife Service, Denver, CO. IRVING, D.B., AND T. MODDE. 1994. Assessment of Colorado squawfish in the White River, Colorado and Utah, 1992 1994. Final report. Recovery Implementation Program, Upper Colorado River Basin. US. Fish and Wildlife Service, Denver, CO. JORDAN, D.S., AND B.W. EVERMAN. 1986. The fishes of North and Middle America. Bulletin of U.S. Natural Museum 47 (4 parts), I-IX:1 3313. MCADA, C.W., W.R. ELMBLAD, K.S. DAY, M.A. TRAMMELL, AND T.E. CHART. 1998. Interagency standardized monitoring program: summary of results, 1997. Recovery Implementation Program for the Endangered Fishes of the Upper Colorado River Basin, U.S. Fish and Wildlife Service, Denver, CO. 21 pp + appendices. MINCKLEY, W.L. 1973. Fishes of Arizona. Arizona Game and Fish Department, Phoenix. OSMUNDSON, D.B., R.J. RYEL, AND T.E. MOURNING. 1997. Growth and survival of Colorado squawfish in the upper Colorado River. Transactions of the American Fisheries Society 126:687 698. RYDEN, D.W., AND L.A. AHLM. 1996. Observations on the distribution and movements of Colorado squawfish, Ptychocheilus lucius, in the San Juan River, New Mexico, Colorado, and Utah. Southwestern Naturalist 41:161 168. TRAMMELL, M.A., E.P. BERGERSEN, AND P.J. M ARTINEZ. 1993. Evaluation of Colorado squawfish stocking in a mainstem impoundment on the White River. Southwestern Naturalist 38:362 369. TYUS, H.M. 1985. Homing behavior noted for Colorado squawfish. Copeia 1985:213 215.. 1986. Life strategies in the evolution of the Colorado squawfish (Ptychocheilus lucius). Great Basin Naturalist 46:656 661.. 1990. Potamodromy and reproduction of Colorado squawfish Ptychocheilus lucius. Transactions of the American Fisheries Society 119:1035 1047.. 1991. Ecology and management of Colorado squawfish. Pages 379 402 in W.L. Minckley and J.E. Deacon, editors, Battle against extinction: native fish management in the American West. University of Arizona Press, Tucson. TYUS, H.M., AND G.B. HAINES. 1991. Distribution, abundance, habitat use, and movements of young Colorado squawfish Ptychocheilus lucius. Transactions of the American Fisheries Society 120:79 89. TYUS, H.M., AND C.A. KARP. 1989. Habitat use and streamflow needs of rare and endangered fishes, Yampa River, Colorado. U.S. Fish and Wildlife Service, Biological Report 89(14). 27 pp. TYUS, H.M., C.W. MCADA, AND B.D. BURDICK. 1981. Radiotelemetry of Colorado squawfish and razorback suckers, Green River system of Utah. Transactions of the Bonneville Chapter, American Fisheries Society 1981:19 24. WICK, E.J., J.A. HAWKINS, AND C.A. CARLSON. 1985. Colorado squawfish and humpback chub population and habitat monitoring 1981 1982. Draft. Colorado Division of Wildlife, Endangered Wildlife Investigations Job Progress Report SE3-6. Denver, CO. Received 31 March 1998 Accepted 30 November 1998