Transactions of the American Fisheries Society 125:78-86, 1996 American Fisheries Society 1996

Diel Movement and Habitat Use of Golden in the , California KATHLEEN R. MATTHEWS U.S. Forest Service, Pacific Southwest Research Station Box 245. Berkeley, California 94701, USA

Abstract.—I used radio transmitter determino st diee eth l movemend habitaan e us t t patternf so California golden trout mykiss aquabonita inside and outside cattle exclosures on the South Fork , , California. Twenty-three golden trout were monitored from Septembe , 199319 o t , durin0 1 r diel-trackin6 g21 g hour t fousa r study siten si upper and lower Ramshaw Meadow. No differences in distances moved or home ranges were detected between golden trout insid outsidd ean e exclosures. Mean home ranges were 18.5-54.0m inside the two exclosure sites and 36.0-68.7 m outside the two exclosures. Most golden trout were found within 5 m of their previously recorded location at both upper (93.5% of 521 observations) and lower observations4 (9239 %f o ) Ramshaw Meadow sites. Movemen individuaf to l fish varied: five fish moved less than 5 m throughout the 10-d study period, whereas 12 fish moved 100-608 m. Golden trout were active both day and night, and I observed them feeding throughout the night, even when water temperature studl 2°Cs al a t sy w .A wer sitelo s ea s insid outsidd ean e exclosures, golden trout used three macrohabitats (pools, runs, and riffles) but selected pools in higher pro- portions tha more nth e available runs. Throughou studye th t , golden trout were associated with nine habitat features (undercut banks, willows, bare banks, collapsed banks, open channel, aquatic vegetation, sedge, boulders rootwadsr ,o wert )bu e most commonly found near sedg undercud ean t banks.

Despite the status of the golden trout Onco- cattle) has occurred for more than 100 years rhynchus mykiss s California'a s state fish, littls ei throughout most of the golden trout's range. In this known abou behavios it t habitar ro t requirements. study, I evaluated golden trout habitat associations 1960e I nth 1970sd san , golden trout wer jeopn ei - in one of the golden trout's native streams within ardy becaus f competitioo e predatiod nan n from Goldee th n Trout Wilderness, Inyo National For- nonnative fishes and degradation of their habitat est, where habitat restoration efforts (cattle exclo- (Pister 1991) thein I . r native habitat within high- sures underwaye )ar . This informatio necessars ni y elevation stream e southerth n i s n Sierra for determining the fish's habitat requirements and (Everman 1906; Fisk 1983), chemical treatments assessing whether cattle exclosure effective ar s e eradicated nonnative trout rocd an ,k barriers were in restoring required habitat features. Currently, constructe o prevent d t reinvasion from down- the only opportunity for comparative behavioral stream (Pister 1991). The systematics of golden and ecological studies of golden trout is between trout are complex, although two major subspecies stream segments within cattle exclosures (used as thought to be most closely related to the rainbow surrogates for restored or less degraded habitat) trout-redbanded trout group are generally recog- and adjacent grazed areas. nized: golden trout O. m. gilberti Using radio transmitters I determine, diee dth l and the or California gold- habitat association f goldeo s n trou foun i t r sites insid outsidd ean cattlf eo e exclosure Ramshan si w en trout O. m. aquabonita (Behnke 1992). This Meadow, South Fork Kern River. The objectives paper describes lattee worth rn k o subspecies e Th . of this study wer colleco et t informatioe th n no California Department of Fish and Game reesta- diel movement and home ranges of golden trout, blished pure California golden trout in their native describe the most common habitats used by golden streams golded an , n trou currentle ar t y self-sus- trout, and determine whether there were differ- taining; their habitat, however, is still in poor con- ence dien si l movemen habitad tan inside tus d ean dition (Albert 1982; R.A. Knap K.Rd pan . Mat- outside exclosures. thews, U.S. Forest Service, unpublished). No ecological studies were conducted when Study Site golden trout habita pristines twa virtuallo ,s y noth- I conducte e studth d t Ramshaa y w Meadow ing is known about the fish's ecology under those (36°22'N, 118°15'W; elevation 2,667 m) on the conditions. Grazing (first sheep and more recently unglaciated Kern Plateau in the Golden Trout Wil-

78 GOLDEN TROUT MOVEMEN HABITAD TAN E TUS 79

derness (GTW), (Figure 1). mately 2.4 km downstream and approximately 3.2 The South Fork Kern River flows through Ram- km long. It is a "drift exclosure"; instead of ex- shaw Meadow and is 27 km south of Mt. Whitney. closin grectangulaa r portio streame - th f de n o t i , Golden trou native ar t thin ei s upper portioe th f no fines upstream and downstream boundaries and is South Fork Kern River, where they were reesta- sidese opeth t na . Because this exclosur newers ei , blished in 1983 after nonnative Salmo less recover occurres yha d (Knap Matthewsd pan , trutta and were removed. unpublished) r exampleFo . , mean stream widts hi Ramshaw Meadow underlais i broady nb , slop- similar insid outsidd e an (31 ) e6 cm (34 2 cm)d an , ing granitic alluvium. Adjacent to the meadows the mean depth inside and outside is 22 cm. Stream are lodgepole pine Pinus murrayana and foxtail velocity averaged 30 cm/s during August-Septem- pine P. balfouriana, and streamside vegetation is ber of 1993 and did not differ among study sites. primarily willows (Salix spp.), sedge (Carex spp.), Streamside vegetation also differs insidd an e and sagebrush (Artemisia spp.). Presumably, the outside the exclosures, with more willows inside Kern Plateau (eastern Golden Trout Wilderness) both exclosures, presumably from replantind gan was once a diverse high-elevation meadow eco- lack of grazing (Knapp and Matthews, unpubli- system but is now degraded from past and current shed). Also, unvegetated sandbars were more overgrazin dominated gan sagebrusy db h (Odion abundant outside the exclosures during the sum- et al. 1988). me f 199 o r. Dennis 3(A , U.S. Forest Service- un , Becaus f longeo , harsh winter t higa s h eleva- published data). tions, meadows in the GTW are characterized by shora t growing season from throug y lateMa h Sep- Methods tember. Night air temperatures during this period Fish Tagging typically fall below freezing. Late-summer snow- I used radio trackin determino gt e each trout's storms are not uncommon but did not occur during positio streae th n nmi d relativan y timo da et f eo the radio-tracking study. habitat characteristics. Adult golden trout (>14 The South Fork Kern River is a relatively shal- cm; 3-9 years old: Knapp and Dudley 1990) were low, low-gradient stream s bottoit ; m consistf o s captured by electrofishing and immediately anes- unstable sand and occasional gravel and cobble. thetized in tricaine methanesulfonate (26 mg/L). e streaTh s mcharacterizei y areab d f inciseo s d Once a fish was anesthetized, a transmitter was channels, sparsely vegetated stream banks, active surgically implanted throug- incision ab ha e th n ni erosio absencne th sites d f instreaan e,o m cover. dominal cavity just anterior to the pelvic girdle, In nearby Mulkey Creek, which also has sparse and the incision was sutured (Supramid1 HS 23). vegetation and a widened stream channel, summer During surgery, fish were kept immobil pada n e-o water temperature fluctuaty sma e 20°C daily (Stef- d V-shapede gille dth traysd weran , e constantly ferud 1993). bathed with anesthetic (6.5 mg/L) flowing froma Cattle exclosures were established in the upper battery-operated water pump. After surgery and and lower sections of Ramshaw Meadow, each en- before release, each trout was measured (total compassin gportioa Soute th f hno Fork Kern Riv- length) and allowed to recover for approximately er. The upper exclosure was established in 1983 a water-fille n i n mi 5 d cooler containin- an o gn approximatels i d an y lon1.m 1k g (Figure 1). With- in this exclosure, some of the stream is recovering; esthetic. Each transmitter (Holohil, Canada) was 16X8 stream variables measured in 1984, 1993, and 1994 X 5 mm, weighed 1.0 g in air, and had an external, chronicl a esignifican t narrowin e streath f mo g nylon-coated, stainless steel antenna. The 20-cm- (mean widths: 345 cm in 1984, 271 cm in 1993, long external antenna trailed fro closee mth - din 1994n i m ;c analysi0 23 f variancso f log-transeo - cision. Golden trout are small fish and the smallest formed data, P < 0.0001; Knapp and Matthews, tags available were used. Because of the small tag, unpublished). In 1993, the stream was narrower (means, 271 versus 402 cm) and the water was the life expectanc transmittee th f yo s 14-2wa r 1 deeper (means, 26 versus 22 cm) inside than outstude s restricteth - ywa d dan leso dt s. d tha 0 1 n side the exclosure (Knapp and Matthews, unpub- Golden trout were captured, tagged thed - an ,nre lished). This rectangular structure enclosee th s stream and the adjacent 100 m of riparian zone on 1 Trade names and commercial enterprises are men- either side. tioned solel r informationyfo endorsemeno N . e th y b t e loweTh r exclosure, buil n 1991i t s approxii , - U.S. Forest Servic implieds ei . 80 MATTHEWS

1 kilometer

Tracking location Q i i Upper exclosure fence

Lower exclosure drift fence

Golden Trout Wilderness Inyo National Forest

FIGURE 1.—Radio-tracking study sites insid outsidand e uppeeof loweand r r cattle exclosure Ramshaat s w Meadow, South Fork Kern River, Golden Trout Wilderness, California. GOLDEN TROUT MOVEMENT AND HABITAT USE 81

TABLE 1.—Summary of golden trout tagged during Sep- primarily becaus lace f th streamsidk o f eo e com- tember 1993, including fish number, length, total distance plexity (e.g., the absence of vegetation and boul- moved estimated an , d home range size. Fish were tagged ders) coulI . d sometime taggee e sse th dn i fist hou 1d 0 an oSeptembe n9 r 199 uppet 3a lowed ran r Ramshaw Meadow, respectively. ope agaid nan n verified positio f transmitterno o st withi 2-mna 2 area. Total distance Afte transmittere th r s were deployed e folth ,- Fish number Length (cm) moved (m) Home range (m) lowing data were recorded (approximately every Upper Ramshaw, inside exclosure hour) by teams assigned to upper Ramshaw and 1 16.0 101 41.9 lower Ramshaw: (1) water temperature near each 0 6. 2 5 < 15.0 3 1 8 18.68.0 0 fish's location (wit hhandhela d thermometer)) (2 ; 4 15.0 58 16.4 longitudinal fish locatio nearese th o nt t meter (es- 5 14.0 369 137.8 timated from flagging placed at 25-m intervals on Mean 122 54.0 the bank); (3) time of observation; (4) macrohab- Upper Ramshaw, outside exdosure itat type (pool, riffle, run habita) glide)r o ,(5 d t;an 6 21.0 456 87.1 features (all habitat features where golden trout 7 16.0 301 53.2 were found were recorded; they included undercut 8 17.0 187 20.0 9 16.0 270 231.6 banks, willows, bare banks, collapsed banks, open 10 15.0 <5 1.1 channel, aquatic vegetation, sedges, boulders, and 11 15.0 67 19.0 rootwads) adjacent to fish location (within the 2-

Mean 256 68.7 m area). Trackers were also able to record precise Lower Ramshaw, inside exclosure location2 habitasand t association taggesof d fishat 12 17.0 124 6.2 night because fish were often out in the open and 13 16.5 <5 11.0 away fro streae mth m banks darks whewa t .ni 14 16.0 56 48.4 15 15.5 122 18.2 On Septembe , 199310 r , macrohabitat avail- 16 16.0 31 11.6 ability was measured in 500-m sections in the four 17 15.5 71 15.6 study sites (inside and outside exclosures in upper Mean 56 18.5 and lower Ramshaw Meadow). Within each 500- Lower Ramshaw, outside exdosure m section e lengtth , f streaho m containing each 18 18.0 299 42.6 macrohabitat was measured, and the proportions 19 17.0 608 106.2 20 16.0 394 22.4 (number of meters per 500 m) of pools, runs, rif- 21 15.0 <5 3.0 flesglided an , s were determined. 22 15.0 <5 2.0 Because of the 14-21-d life expectancy of the 23 15.0 167 44.0 transmitters unabls wa I allo,o et w more thad n2 Mean 244 36.7 of recovery before collecting data. However, fish apparently recovered quickly from surgerd an y reasoo n d taggingbelievo nt ha I d e an ,tha t tagged leased at the capture site. Equal numbers of fish fish behaved differently than untagged fish. I snor- were tagged and released into each study site (six kele streae dth m dail observo yt behavioe eth f o r inside and six outside the lower exclosure, and six tagged fish (identifie theiy b d r external antenna) outsidx insidsi d uppee eth an r exclosure) (Table and untagged fish and to determine the well-being 1). However, after release fise h,on insid uppee eth r of tagged fist movedhno thavisuad y M ha .- t ob l exclosure died. servation f taggeo s d trout indicate appareno n d t On 9 and 10 September 1993, 24 golden trout adverse effects I observefro tage d mth an , d fish were implanted with transmitters, each wit- in h swimming normally and feeding the same as un- dividually recognizable frequencies locato T . e eth tagged fish. tagsChallengeS I AT use, n da r M R400AV r 0o Each team was assigned a 12-h "day" (0800- LA12-DSE receiver and a hand-held, three-ele- 2000 hours) or "night" (2000-0800 hours) shift. ment, collapsible Yagi antenne stream'th t a a s Upper Ramshaw fish were tagge September9 n do , edge. Tagged fish m coul 0 detectee 10 db o t p du and data were recorded during 0900-1900 hours away. Locations were pinpointed through trian- on 11 September, 0800-2000 hours on 12-13 Sep- gulation beford an , e data collection, fish trackers tember, 2000-0800 hour n 14-1o s 5 September, practiced locating transmitters. Transmitters were 2000-0800 hours on 15-16 September, 0800-2000 placed in known locations in the stream, and track- hours on 16 September, 2000-0800 hours on 16- s verifieer d their location withio st 2-mna 2 area, 17, 17-18 18-1d an , 9 September 1200-160d an , 0 82 MATTHEWS

hour d n 18so an Septembery da 0 totaa 5 r f o lfo , habitae oth f t features (sedges, boulders, etc.)I , nigh0 6 t tracking hours. Lower Ramshaw fish were coul t tesdno t whether they were use signifin di - taggeSeptember0 1 n do datd an ,a were recorded cantly different proportions. Instead, I report the during 1300-1900 hour 1n 1so September, 0800- proportion of observations during which golden 2000 hours on 12 September, 0800-2000 hours on trout were associated with each habitat feature. 13 September, 2000-0800 hour n 14-1o s 5 Sep- Fish movement.—Summaries and analyses of tember, 0800-2000 hour 1n 5so September, 2000- fish movement determined whether there were dif- 0800 hours on 15-16, 16-17, 17-18, 18-19 Sep- ference homn si e range movemenr so differenr tfo t tember d 1200-160an , 0 hour n 1o 8s September, time locationsr so . Nigh defines perioe wa tth s da for a total of 46 day and 60 night tracking hours. with no solar radiation (1800-0800 hours, which lower uppeFo d ran r Ramshaw Meadow, 216 track- included dawn [0600-0800 hours d dusan ]k ing hours were completed. [1800-2000 hours]), and day as the period in be- tween (0800-1800 hours). Water and Air Temperature Distances moved between observations were In addition to the water temperatures measured classified as 0-5 m, 6-25 m, 26-100 m, and greater when fish were located, water temperatures were than 100 m for night and day periods. For the also measured at upper Ramshaw Meadow at the different diel periods, I tested the hypothesis that top, middle, bottom, and just downstream of the there was no difference in movement greater than upper exclosure. In addition, air temperatures wer e(log-likelihoom 5 d rati r contingencofo y tables, bottotakee Ramshae th th t na f m o w exclosuret A . G-test 0.05= r 1974)a , ;Za treateI . d only move- each locatio watef no r temperature collectiono tw , ment greater than 5 m as real because errors in Omnidata model ES-60 single temperature probes distance estimation were common. (accurate within ±0.25°C) were placed with the I also totale movemente dth r individuasfo l fish thermistor bottoe streame th th n f mso o . Temper- ove stude th r y period r thiFo .s summary- cu e th , ature readings were taken every half hour during mulative distances moved per individual fish were our study period, September 10-19, 1993. HereI , summed and averaged for inside and outside of report the maximum and minimum stream tem- the upper and lower exclosures. The means of the peratures durin studye gth . total distance moved over the study period were then compared (Mest, a = 0.05) to determine if Data Analysis the mean distance moved differed inside and out- Analyse summaried san s were divided into three side the two exclosures. categories: habitat use, movement d homan , e To delineate home ranges useI , "adaptive dth e range. Data for upper and lower Ramshaw were kernel" method (Silverman 1986; Worton 1989), compared separately, except where noted. To test depicting 90% of the location points recorded for whether golden trout use e macrohabitatth d n i s individual fish. First, from the 24-h fish location different proportions than what was available, I data probabilistia , c density estimate (PDEs wa ) used Mests I summarize. proportioe dth - ob f no made along the linear distance of the stream (Sil- servations over the study period for individual fish verman 1986) such that each home range estimate represente e lineath d r distance coveree th y b d within each macrohabitat type. The nI compute d PDE's central 90%. Usin centrae gth mini % 90 l - the mean proportion and standard error using the mizes possible outliers resulting froe relamth - individual fish as the unit of replication. The means tively short sampling . perioThusd) 8 ( d, each of macrohabitat observations were then compared home range represents the linear distance (m) to the known proportion of the measured macro- alon streae gth 24-e th m hf o wherlocatio % e90 n habitat within the four study sites. The null hy- data points were establishe r eacfo d h e fishTh . pothesis teste HQ:s dwa mean proportion goldef o n home range estimates (m) were compared between trout observation in habitat type = measured pro- fish inside and outside of the exclosures, testing portio f habitano t type withi m0 n50 (Za r 1974). nule th l hypothesis tha differenco t n ther s ewa n ei Individual macrohabitat comparisons were per- home range distance insid outsidd ean excloe eth - formed onl pooln yo rund san s w becauslo e th f eo sures. number of observations in riffles and glides. Mean number observationf so s averaged from individual Results golden trout in pools or runs were compared inside MacrohabitatUse and outsid exclosure eth nightd an r boty e.fo hda Runs constitute largesdthe t proportioma- nof Because I did not measure relative availability crohabitats inside and outside the exclosure in up- GOLDEN TROUT MOVEMENT AND HABITAT USE 83

TABLE 2.—Proportions of measured availability of gold f poolso - t there us bu , e wer significano en t differ- en trout macrohabitats from 500-m sections insid outd ean - ences in use and availability. side exclosures at upper and lower Ramshaw Meadow and the mean proportion of day and night observations during Use of Habitat Features which individual golden trout were foun thosn i d e macro- habitats. An asterisk indicates a significant difference (/- Throughou studye th t , golden trout were asso- test, P < 0.05) between the available proportion of mac- ciated with nine habitat features (noted in Meth- rohabitat and the mean proportion averaged over ods) which were use n differeni d t proportions individual fish. throughou trackine th t g study moste Th . common- Habitat ly used were sedge (38.5 %2,85f o 7 observations) measure Pools Runs Riffles Glides and undercut banks (30.3 f %observations)o . Upper Ramshaw, inside exclosure Upper Ramshaw pools.- up n —Duringi y da e th Available 0.31 0.65 0.05 0 Ramshaper w pools (619 observations), sedges Day use 0.88* 0.11* 0.01 0 wer predominane eth t habitat feature use goldy db - Nighe us t 0.89* 0.11* 0 0 en trout both inside and outside the exclosure (Ta- Upper Ramshaw, outside exclosure ble 3). Inside the exclosure, collapsed banks, Available 033 0.55 0.11 0 aquatic vegetation, willows, and boulders (2% of Day use 0.89* 0.11* 0* 0 observations) were the other habitats used. Outside Night use 0.76* 0.24* 0* 0 the exclosure, aquatic vegetation, willows, and Lower Ramshaw, inside exclosure rootwads wer othee eth r habitat features usey db Available 0.38 0.44 0.03 0.15 e us y Da 0.43 0.56 0 0 golden trout (Table3). Night use 0.37 0.63 0 0 At nigh pooln i t s (266 observations), sedge eus Lower Ramshaw, outside exclosure inside the exclosure increased, and aquatic vege- Available 0.33 0.57 0.1 0 tation and collapsed bank use dropped (Table 3). Day use 0.82* 0.18* 0* 0 During the night outside the exclosure, golden Night use 0.63* 0.38 0* 0 trout use of willows increased, and sedge use de- creased. Upper Ramshaw runs.—There wer 7 obser9 e - per Ramshaw, but golden trout disproportionately vation f goldeso n trou runn i t t uppesa r Ramshaw. used pools durin nighd gan botty (Mesthda < P , Insid exclosure eth observa9 (2 e y durinda -e gth 0.05; Table 2). Similarly, runs formed most of the tions), golden trout were most commonly found habitat in lower Ramshaw, but use of pools sig- near undercut banks (Table 3). Sedges, open chan- nificantly exceeded proportional availability only nel, and collapsed banks were the other commonly outside the exclosure (Mest, P < 0.05; Table 2). used habitat features. Outsid exclosure th e 5 (2 e Inside the exclosure, use of runs was greater than observations), golden trout were most commonly

TABLE 3.—Summar proportioe th f yobservationl o al f no pooln s i run d san s during which golden trout were associated with the listed habitat feature inside and outside exclosures in upper and lower Ramshaw Meadow. Bare banks and boulders, although monitored, were seldom used by the fish during observation periods.

Day Night Pools Runs Pools Runs Habitat feature Inside Outside Inside Outside Inside Outside Inside Outside Upper Ramshaw Sedges 0.58 0.39 0.19 0.0 0.80 0.29 0.46 0.21 Collapsed banks 0.15 0.02 0.07 0.19 0.10 0.02 0.0 0.12 Aquatic vegetation 0.15 0.26 0.0 0.81 0.06 0.14 0.0 0.15 Willows 0.07 0.17 0.0 0.0 0.02 0.35 0.0 0.06 Rootwads 0.0 0.10 0.0 0.0 0,0 0.04 0.0 0.0 Undercut banks 0.01 0.03 0.67 0.0 0.01 0.01 0.08 0.14 Open channel 0.0 0.0 0.07 0.0 0.0 0.0 0.46 0.39 Lower Ramshaw Undercut banks 0.35 0.53 0.58 0.51 0.39 0.42 0.48 0.46 Sedges 0.31 0.25 0.41 0.49 0.28 0.26 0.52 0.53 Aquatic vegetation 0.16 0.06 0.0 0.0 0.28 0.15 0.0 0.0 Collapsed banks 0.16 0.16 0.0 0.0 0.14 0.16 0.0 0.0 84 MATTHEWS

TABLE 4.—Movement f goldeo s n trou n uppei t d an r exclosure s comparewa s thao dt f fis o t h outside lower Ramshaw Meadow. Data are separated into distance the exclosures (r-test, P > 0.05; Table 1). categories of 0-5, 6-25, 26-100, and >100 m, and into Some one-time, unidirectional distances (6-226 day (0800-1800 hours) nighd an , t (1800-0800 hours- )ob servations. m) were detected (Table 4). At both upper and lower Ramshaw Meadow dieo , n ther ls difewa - Number of observations for ference in the occurrence of these longer-distance distance category (m): Diel . movements (G-test 0.05> P , ; Tabl. e4) period 0-5 6-25 26-100 >100 Total The single longest distance movement was 226 Upper Ramshaw y fis mb 9 froh m outsid e uppeth e r Ramshaw Day 355 11 2 0 368 Meadow exclosure. It was tagged and released at Night 414 29 12 0 455 46 m above the exclosure on 11 September and Lower Ramshaw s stationarwa y unti September3 1 l ,s whewa t i n Day 249 15 4 0 268 farthem foun 6 22 dr upstream e fishTh . stayen di Night 455 31 7 2 495 this vicinity until tracking was terminated on 19 September. In contras mosto t t fish that remained withina associated with aquatic vegetation and collapsed narrowly defined location, two golden trout (18 banks. and 19) in lower Ramshaw Meadow moved re- At night in runs (43 observations), golden trout peatedly and recurrently used two primary loca- were commonly e opefounth nn i dchanne l both tions. Fish 18 move bac fortm d 0 kan d4 h between insid outsidd ean exclosure eth e (Tabl . Inside3) e two locations; it was typically found at 92 m above the exclosure, sedges and undercut banks were also the upper drift fence during the day and 132 m used. Outsid exclosuree eth , golden trout also used above it at night. At night, this fish was observed sedges, aquatic vegetation, collapsed banks, wil- with aggregations of 20-30 other golden trout in lows, and bare banks (6%). opee th n channel. Lower Ramshaw pools.—Inside e loweth - ex r Fish 19 was caught, tagged, and released above closur pooln ei s (246 observations), golden trout uppee th r drift fenc n 1e0o Septembers wa t i d an , primarily used four habitat features during the day: relocate 1n d1o Septembe insidm exclo2 e r3 eth - undercut banks, sedges, aquatic vegetationd an , sure t useI . d primarthia s sa y area y durinda e gth collapsed banks (Tabl . Outside3) exclosure eth e and, lik t movei e , fis18 hd upstrea t mnight—ia n in pools (237 observations) same th , e habitat fea- this case 101m upstrea abov m aren a 9 a6 emo t tures wer edifferenn i uset bu d t proportionst A . fencee th . nigh pooln i t s (251 observations), undercut banks, Mean home range f goldeso n trout (linear dis- sedges, aquatic vegetation collapsed an , d banks tance embracing 90% of locations) ranged from wer e predominanth e t habitat features usey b d 18.5 to 68.7 m among the four study areas (Table golden trout inside and outside the exclosure (Ta- 1). There were no differences (/-test, P > 0.05) ble 3). between home ranges inside the exclosures (range, Lower Ramshaw runs.—In both inside (30- 4ob 6.0-137.8 m) and outside (range, 1.1-231.6 m). servations) and outside (52 observations) the low- Water and Air Temperatures er exclosure during the day, golden trout were pre- dominately found near undercut banks and sedges Durin e studth g y period, water temperatures (Table 3). Similarly, golden trout were found near ranged from 2 to 16.5°C in Ramshaw Meadow and sedge undercud san t banks both insid outsidd ean e typically underwen dieta l chang f 10°Ceo . Golden the exclosure at night (486 total observations). trout remained active durin temperaturesw glo o N . differences in water temperature were detected be- tween reaches insid outsidd ean uppee eth r exclo- Movement Patterns sure (paired r-test, P > 0.05). Air temperature dur- In both upper and lower Ramshaw Meadow, ing the study ranged from -11.8 to 27.2°C. most tagged golden trout exhibited little move- ment froobservatioe mon nexe th t o nt (Tabl . e4) Discussion Overall, most fish wer previoue eth f withio m s n5 This is the first study of movements and habitat location during the day (95%) and night (91.4%). use by California golden trout in their native hab- Cumulative distance t diffemoveno r fisd r dpe hdi itat. Daily movements and habitat associations of whe e meath n n distanc f fiso e h from inside th e golden trou e criticar understandinar tou o t l f go GOLDEN TROUT MOVEMEN HABITAD TAN E TUS 85

their habitat requirements and the ongoing resto- ripariae th par f o t n Ker e areath nn so Platea - ube ration efforts. Golden trout were most commonly fore degradation because there are no comparable foun pooln i d s near sedge d undercuan s t banks, ungrazed areas. However, sedges wer commoea n and these habitat features are typically reduced or habitat feature used by adult golden trout, and cur- lacking in grazed areas (Stuber 1985; Marcus et ren tKere worth nn ko Platea u suggests that sedges al. 1990; Plans 1991). Adult trou othef o t r species may also serve to stabilize the bank (R. Knapp, prefer pool d undercuan s t banks (Campbeld an l Aquatic Research Laboratory, per- Neuner 1985; Bjornn and Reiser 1991), consistent sonal communication). Future research plans in- with wha observetI adulr dfo t golden trout. During clude measurin availabilite gth f habitayo t features e studyth , golden trout maintained small home to determine whether golden trout are seeking and ranges and usually moved little from one obser- selecting sedge r undercuso t bank preferencn i s e vation to the next. No differences in movement to other habitat features. In recovered areas inside distances or home ranges were detected between the exclosure, dense sedge banks narroe th w golden trout insid outsidd an e e livestock exclo- stream widt increasd han e water depth (Knapd pan sures. Matthews, unpublished). Thus, for restoration, it Durin e nightgth , golden trout were activd ean is crucial to determine which native plant assem- sometimes moved long distances I als. o observed blage is needed to meet the critical habitat requi- aggregations of 20-30 golden trout in the middle rements of golden trout as well as to stabilize of the stream, where they seemed undisturbed by banks. observer presence. Underwater camera and video equipment were placed within centimeters of trout, Acknowledgments which did not swim for cover as they often did I thank Roland Knap r introducinpfo e th o t e gm durin e dayth g . Golden trouy activelma t y feed incredible golden trout. Phil Pister deserves ac- ove entire th r e diel period durin late gth e summer colade extensivs hi r sfo e wor golden ko n troud an t to optimize growth during the short high-elevation for alertin worle gth theio dt r plight. Sara Chubb, growing period. The diel movement behavior of former Inyo National Forest Fish Biologistd an , rainbow trou s variablei t , some population- ex s Del Hubbs, Range Conservationist, assisted in the hibiting nocturnal activit otherd yan s being rela- logistic f researco s h implementatio e Inyth on o n tively sedentary (Jenkins 1969; Jenkin . 1970al t se ; National Forest. Ed Ballard, Dave Kammerer, Bisson 1978; Campbel Neuned an l r 1985; Angradi Dave Azuma d Thadan , Edens performee th d and Griffith 1990; Matthew- . 1994)un al s t i se t I . grueling fieldwor assisted kan d with data analysis. cleatrouy nocturnalle rwh ar t y activet ; theno o yd Jim Baldwin, Pacific Southwest Station statisti- always exploit nocturnal increases in prey (Bisson cian, reviewed the manuscript and assisted in the 1978; Angrad Griffitd an i h 1990). Nocturna- ac l statistical analysis. Michae . YoungK l , Peter Moy- tivity of salmonids may be related to water tem anonymoun a - d lean , s reviewer provided helpful perature wated an , r temperatures a sw werlo s ea and constructive reviews projece Th .fundes wa t d 2°C during this study. Fraser et al. (1993) reported Fise byth h Habitat Relationship Prograe th f mo that salmonids switch from diurnal foraging at U.S. Forest Service's Region 5 Fish Program. Den- warmer temperature nocturnao st l foragin tems ga - nis and Jody Winchester of Cottonwood Packers peratures drop below 10°C. Littl knows ewa - nre and their mules endure d movean d d moundf o s gardin nocturnae gth l activitie f goldeso n trou- be t equipment, making wilderness fieldwork possible. fore this study, and it is still unknown whether the nocturnal activity I observed is typical or occurs References ove entire rth e year seasonaA . l analysi f goldeso n Albert, C. P. 1982. A survey of factors influencing the trout movemen neededs i t . conditio streae th f mno zon Golden ei n Trout Wil- Golden trout were found more often near sedges derness. Master's thesis. Sonoma State University, than near willows, eve arean i uppef so r Ramshaw Sonoma, California. where willows wer estabw e replanteno -e ar d dan Angradi, T R., and J. S. Griffith. 1990. Diel feeding lished insid exclosuree eth mann i s yA . other res- chronology and diet selection of rainbow trout On- toration plans (Osborne and Kovacic 1993), wil- corhynchus mykiss in the Henry's Fork of the Snake lows were replanted in the Golden Trout Wilder- River, . Canadian Journal of Fisheries and Aquatic Sciences 47:199-209. ness wit constructioe hth exclosuree th f no stao st - Behnke . J 1992 . R , . Native trou f westero t n North bilize stream banks (USFS 1988). It is unknown America. American Fisheries Society Monograph 6. whether willows were a natural or an important Bisson . 1978A R , . Diel food selectio sizeo tw f so y nb 86 MATTHEWS

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