Tramacrbm of ths Amcrican Fkhrrirr Snrirry 12l:M7453, 1999
Amcrlcan Plshcrkr Sockty 1
W
Movement Patterm of Stream Fishes
Ouachita Highlands Stream: An Examination of the
Restricted Movement Paradigm
Deparrmenr of Biology, Universiv of Mississippi, University. Mississippi 38677, USA
and US.Foresr Servke, Southern Research Station,
Odord, Mississippi 38655, USA
U.S. Forest Service. Southern Research Sratlon,
Oxford, Mississippi 38633, USA
A6slracr.-The rcstrictcd movement paradigm (RMP), which states adult fish do not move out
of a pool or rcstricted stream reach, does not fully define the movements of stream fishes. Although
stream fishes may spend thc majority of their time in n home pool, they also make regular exploratory trips away from the home pool. Rccapturc of Ouachita Highlands atream fish marked
with passive integrated transponder (PIT) tags showcd that the majority of thc uniqucly marked
fish in this study were in the pools of initial collection. The following percentages of marked fish
were recaptured outside the pool of initial collection: 12% of creek chub S~molilusnlromnculullu-;
33% of blackspotted topminnow Fundulus olivuceus; 12% oC green sunfish LPpomis cyanellus; and
14% of longear sunfish Lepomls muguloris. Standard lengths and weights did no1 dilrer significantly
hetween nonmobile and mobile recaptured fish, and patterns of nlultiple recaptures indicated
exploratory, round-trip movements. Such regular exploratory trips suggest that the area of use by
and critical habitat for stream fishes may be greater than conlrnonly cstimatcd.
Populations of strcam fishes havc been viewed
as completely homogcncous and mobile (Linfield
1985; Fausch and Young 1995) or au relatively
discrete units exhibiting restric~ed movement
(Gerking 1953; Hill and Grossman 1987; Heggencs ct al. 1991). These two views of fish movement
havc profoundly diffcrcnt implications for the
management and conservation of stream fishes.
Therefore, a better understnnding of the patterns
and proccsscs of fish movement is necessary to
dclincatc fish populations.
Many studies of fish movement have indicated
that strcam fishes arc scdcntary, never leaving a
particular puol or stream rcach (c.g., Bcrra and
Gunning 1972). Gowan el al, (1 994) termed this
concept the "restricted movement paradigm"
(RMP). Restricted movement is defined as recapture from thc arca whcrc the fishes were initially
marked and rclcascd (Gcrking 1959). Examination
of the data from such studies shows that although
* Corresponding author: IeesmittyQaol.com
Prcscnt address: 1702 Devine Sweet, Jackuon, Mississippi 39202, USA.
Present address: Department of Fisheries and Allied
Aquaculrures. Apburn University. Auburn, Alabama
36849, IJSA.
Recejvcd January 8, 1998; acccptcd Novcmbcr 11, 1998
a majority of fishes studied are sedentary, leading
authors to conclude that movement is restricted, a
proportion of fishes in every study do move, sometimes for long distanccs (Gerking 1953; Hill and
Grossman 1987; Heggcncs ct al. 1991). Funk
(1957) proposed that populations of fishcs arc
composed of both mobile and sedentary fractions.
Few studies have addressed whether individuals
exhibit onc stratcgy or the other exclusively or
switch from mobile to sedentary behaviors. However, a study of individually marked brown trout
Sulmo trutta showed frequent-switching behavior
(Harcup et al. 1984).
Populations of fishes are not likely to be complctcly scdcntary. Studies of recolonization suggest that fishes arc rnobilc and rcspond rapidly to
habitat in which no fish arc found (Larimorc ct al.
1959; Peterson and Bayley 1993). Recolonization
of defaunated stretches of stream can begin immcdiatcly aftcr a disturbance, and sedentary species such as longcar sunfish kpornis megalotis can
recolonize within 2 wccks (Pctcrson and Bayley
1993). The identity of such colonists has been
poorly studied, but rapid colonizcrs may represent
thc mobilc cornponcnt of a nearby population.
We examined the RMP by investigating patterns
of movement of individually marked fishes in a
Ouachita Highlands stream. By using individually
SMrI'HSON A N D JOHNS'I'ON
-mphny
Boundary
Meters
marked fishes, wc wcrc alsn able L O dctcrnline if
mobile fishes wcl-c a discrete coniponcnt of the
population or i f individual tishes swi~chcdbctwcen
mobile nnd sedenlury behaviors.
Study Site
Littlc Glazypeau Creek (Ouuchitn River drninage) is in the Ouachita Highlnnds of Arkansas
(Garland Chunty; Figure I). 11 is n low-order
strcarn wilh rifflc--pool sequences. Thc substrate
rangcsfrnm small cohhle to bouldcrs and the walcr
is clear. Thc riparian zone consists of deciduous
hardwood uud pine foresl. During the dry season
of late summer to winter, pools in Littlc Glazypeau
Creek are connected by shallow riCRcs; however,
TAHL~,
I.-Wfi~er VOIUIIICS
oC Little Glazylwau Creek.
Watcr volurnc (m')
Fool Menn
SU
Kungc
and VCIOCIL~CS in sludy pools
Watcr vclocily (mly)
Mcnn
S1)
Kungc
PWI volumes (cu m
1-237
11-541
2 - 8.9
3 - 85.9
4 - 72.8
5 - 136.0
6 - 32.1
7 - 6.8
ew
the study site was isolatcd upstream o f pool 1 and
downstrcarri of pool 10 during the Cull of 1995.
Water temperature ranged from 20°C to 25'C during the study.
The study ~ i t was
c 500 ni long and hnd I 1 sludy
pools ranging in length from 10 in to 59 m. Bccause the I I th pool was added during the second
sarnpling trip, it is numerically out of sequence
with thc othcr pools. Additional 300-tn slrclchcs
were sampled upstream and downstream of ~ h c
study area 10 dctcct long-range movernenls of
marked sludy specimens. Thc lotnl sampling urcu
was 1. I k ~ nlong. During cnch sampling cpisodc.
pool Icngth, width, and depth and water velocity
wcrc rccorded for each siudy pool. ('hhlc I).
Methods
Crcck chub Scmotillrs ntrornur~ulatus, blackspottcd topminnow Fundul~rs olivurcus, longear
sunfish, and green sunfish Lrporni.~ryant~lluswere
markcd in May 1995. Nine colleclions (one per
month) were conducted during Muy, July, August,
and Octohcr of 1995 and 1996 and May 1997 LO
monitor mtwcrncnts and mal-k additional individuals. F i ~ h c swere cnllectcd by electrofishing with
n backpuck shocker. Initinl sampling indicalcd that
two passes within u pool resulted in depletion o f
thc focal specics. Wc continued to usc two passcs
in ench study pool on subsequent trips lo cnsure
-
' I ' I I S ' I01;
' KI;.STKITTEDM ~ V E M E N TP A R A D I C ~ M
sampling efl'orl was consisletit. We conductcd single sampling passes in all areas bctwccn the sludy
pools and the 300-111 strclchcs upstl-cam and downstrculn o f the study sile on each trip to satnplc for
marked lish.
Date arid locatioti of collcctIon (pool number or
intervening area) wcrc I-ccordcd for all of the study
spccics captul-cd. Fish were ariestheti~ed,weighed
(g), and measured (standard Icngtli, Sl.,, I .O mm).
Each fish collcctcd from thc sludy pools was
markcd wilh a subcutaneous injection of artist's
lalex paint (Freeman 1995). The color and posilion
of the mark rcprcscntcd a spccilic pool. l'assive
integraled transponder (PIT) tags wcrc injcclcd
into crcck chuh, green su~ifishand lonpciir sunfish
large enough to acco~riruodntcthcm (>70 mrn for
creek chub and >65 Inm for (.he sunlishes). Each
tag had a uniquc numeric code (hat iderltificd individuals. Fish were released to thc location of
colleclion i~potirecovery fmm uncsthcsia. A prior
c~peritrietitshowed the tagging PI-occdul-ccaused
no mortality (Johnslon and Smithson 1999). Study
spccics collccled on subsey~letittrips wcrc checked
for a paint mnrk nnd scanned IOI- presence of a 1'1'1'
tag. All fish that had n u paint mark were rnarked,
and untnggcd fish that lil the lagging criteria wcrc
tagged by the same proccdurcs used on the initial
collcction hip.
Fish recaptured in the pool of initial collection,
or home pool, wcrc clussilicd as nonmobile. They
were assigned a value o f zcro mclcrs moved (Freeman 1995). Fish recaplured outside of the initial
pool c>fcaplurcwere classified as mobile. Thc dislance traveled by a fish rccupturcd outside the i n tin1 collcction pool was measured as the midpoint
of the initial pool to the midpoint ol- the recaplure
pool (Slott 1967; Rruylants ct HI. 19Xh).
Dutu for all recaptured fish wcrc cxamincd LO
dctcrminc ~ c m p u r dand species-spccific dil'l'cl-enccs in lnovcincnl hchavior. Fish recaptured more
than oncc wcrc counted each time they wcrc captured. To test for differcnccs in movcmcnl brhavior betweell spccics, the numhers ofnonrnobilc and
mobilc lish recaptures were comparcd by chisquare analysis of mutual independence. Cornparisons wcrc made herween slandard Icngtlis and distnnccs moved of all recnpturcd fish to dctcrminc
i f thcrc was a correlation betwccn thcsc L W O variables. Correlation analysis was also used lo determine if thcrc was a signilicaril relationship bctwccn the number of days bctwccn tagging and
recapture of fish and the distance ~ n o v c dupon recapture. A G-test fol- mulual indeperidencc was
used to test for signilicanl differences among col-
849
lecting hips in fi-equency of recnpturcd tish showing movernetit (Sokal and Rohlf 198 1 ).
To determine if ~ h cnonmobile and rriobile
groups wcrc discrete, we cornpared chaructcristics
01- ~ h clagged fish that did not lnovc to thosc (ha1
did. We used I-tests with assumed unecl~lalvuriances to cvuluutc the cffccl of fish size on movcmcnt hchaviol: Slandard lengths of thc rccaptul-cd
fish that moved and thosc that did no1 move were
compared, as wcrc thcil- weights.
The movement patterns of fish rccuptul-cd at
least twice were cxamincd to dclcrmine individual
plasticity o f rnovcmenl. Movement behavior wus
divided into three catcgorics: n o movement, oneway movement, and complcx muvcmmt. Recaplures found only in thc homc pool were assigned
to the no-muvcmcnl ualegory. One-way movement
indicnlcd rccaplurcs were made in any pool olhcr
lhan he pool of marking. Fish ~rccapluredin one
or more nonmarking pools and s u b s e q ~ ~ e n t lin
y
their homc pools wcrc assigned to the complcxmovcmcnl calcgory.
To deterrnine if emigration firom ~ h study
c
reach
confoundcd thc ~rccap~ure
results, we c o ~ n p a r c drccapture I-atcs ut limes the study sitc wus hydrologically isolaled up- and downstrcum wilh recapture 1x11~s
a1 limes of through flow by rncuns of (;lcsls for itidependencc. Recapture rates compared
across collections cstimalcd sampling efficiency
and ovcrull aclivily of fishes.
Results
We found no evidcncc thal movcmcnt of fishes
into or out of lhc study site compromised our rcsults. Rec;~plurerates (all study spccics combined)
on sampling dutcs when the study sitc was hydrolopically isolated did no1 differ significantly h-om
I-cuplure ratcs on all the olhcr sampling dates ( G
= 0.0368; df - 5: P > O.9Y9). Overall recapture
ratcs generally ir~creascdover time: July 19958.5%; A u g ~ ~ s 1995-24.5%;
t
OcLober 199575.5%); May 199b- -40%; July-26.5%); Augusl
1996- 3 7 % ) .
The rnajority of d l four monilored species wcrc
rccapturcd in Lhc poul of initial collcction. Smaller
PI-oporlions of the rccuptu~-cdlish were collected
outside of their homc pools (T:ible 2).
There was a sjgnilicanl difference in the ratio
of nonmohilc LO mobile recaptures hctwccn the
four sludy species ( X ' = 9.458; dl'= 3; P = 0.03 I ) .
N o differcnccs wcrc found in movement behaviolbetween crcck chuh, gl-ccn sunfish, and longcalsunfish when the contingency tablc was subdivided
( x 2- 0.002; dl' = 1; P = 0.966), but 1hc proportion
850
S M I T H S O N A N D JOTINSTON
TAIH.I!
2.-Nurribcr~
of tish ninrkcd nnd rccnpnlrcd (including mulliplc recaptures) in G l w y p e a u Creek
Slxcics
Numlrcr
marked
C'rcck chulr
Blacksptted topminnow
Circcn ~ l l n i i ~ h
I ,ongcur uunliuh
112
163
679
.364
Numhcr ( 4 j ) n
I-ecoptul-ed
25 (22)
30 ( 18)
201 (30)
231 (65)
Nurnlw ( ' f i ~ ) ~ Numlrc~,(%jb
rccapturrd
recaptlired 011~
US homr pod
in homc pool
22 (XX)
2 0 (67)
176 (88)
109 (80)
3 (12)
10 (3.3)
25 (12)
32 (14)
P c r ~ c n ~ a gorc r~uml,cr ~ ~ ~ a r l c d .
"crccntapc nf total number rrcnpturcd.
of mobilc hlackspottcd topminnow was significantly greater than that of the three other study
spccics combined (x' = 7 . 6 9 ; df = I ; P = 0.006).
Nevertheless, standard lengths and weights did not
diffcr significantly different between nonmobile
and mobile fractions for any study spccics (Table
3). Moreover, correlation analysis showcd no rclationship bctwccn standard lcngth of rccupturcd
fish and distanced movcd from the pool of initial
collcclion to the pool of recapture (creek chub: N
= 25, r = 0.205. P = 0.414; blackspotted topminnow: N = 30, r = 0.041, P = 0.836; grccn
sunfish: N = 201. r = 0.117, P = 0.284; longcar
sunfish: N = 231, r = -0.073. P = 0.493). Thcrc
also was no significant corrclalion bctwccn the
numbcr of days from tagging LO rccnpturc and distance moved (creek chub: N = 21, r = -0.252, P
= 0.314; green sunfish: N = 89, r = -0.0564, P
= 0.640; longear sunfish: N = 91, r = -0.1 16, P
= 0.213).
Subsequent arialyses were restricted to sunfishes
because recaptures of other study spccics wcrc too
few nnd only su~ifisheswere recaptured more than
once. The ratio of non~nobileto rnobilc rccapturcs
did riot differ betweeu green and longear sunfishes
( G = 13.88; df = 7; 0.05 < P < 0.10), so rccnpture
data for thc two spccics wcrc comhincd.
The majority of green and longear sunfishes recaptured outside their home pools were collected
in the pools adjacent to the pools of initial collectioo (Figures 2. 3. Of thc mobilc suntishcs. 70%
moved lcss than 100 m (Figurc 4); tbc most distant
rccapturcs wcrc 453 In for a grccn sunfish and 506
m for a longcar sunfish.
Of the 19 grccn sunfish recaptured more than
oncc, 18 showed no movement from their initial
pool and only 1 showed complex movement (Table
4). In contrast, nearly one-third of longear sunfish
rccapturcd at lcast twicc showcd onc-wny or complcx movcrncnts.
TARI.E
?.-Mean
slantiurti Irnglhs ( m m ) und w c i g h h ( g ) -C hulr-95% confidence ir~tervuls(CT)Tur r~cinmohilr and
rnohilc rccnpturcd Little Cilnzylwnu Creek tish.
S p x i c s a i d variahlc
Nunrnohilc:
lncan I CI
(~'an~c)
Muhile:
mean I CI
(~'angc)
3.23
57.35
(4.3-72)
3 57 L 0.63
l1,(+7,1)
56.10
2,.24
(48-60)
3.05 L 0.47
(2.3-3.81
0.538
68.38 I 2.08
(40-1 znj
12.19 3.60
(2.1-60. I )
hH.36 1 10.50
(33-10s)
13.07
4.12
( 1 .3-39.1 )
0.W7
t'
(1-tcsr)
Creek chub
hnglh
Wr~ghl
Hluckhpollcd l~ipn~innow
Length
Wcighr
+
+
0.2 10
Green suntish
Lcnglh
Wcight
Lungeiu sunlish
I .cr~~lh
Weighl
+
+
0.694
85 1
TEST OF RESTHI(:'I'EI) M O V E M E N T F A R A D I ( . ~ M
0
1
2
3
4
5
8
7
0
8 1 0 1 1
Capture Pool
I:I(?LIRI: 3.-('cinipn1-isor of recilplurc pool with initial
I ' I ~ L I I E2.-Corriparison of rccapturt: pool wilh initinl
caplure pool lor grwn s~~ntisli
in I,ilLle Gln/.ypcnu C'reck. ciiplurc pool t o r longcar sunfish in Littlc Glnnypeau
Pool r~unihcrsnrc dung {he axes. Numhcrs nexr lo h l i ~ (:reek. Convcnlions arc lhosc (7f 1iipu1-e2.
squnrcs RI-e~iuinhcrso r Iish ~ C C R P ~ I I I -(includinl:
C~
111111tiple reci~plurrs)i n pnrticulnr pools; squaws with no
in discrele groups, but that individual fish switchcd
numbers represent siriglc I-ccoptures.
movernent be11avio1-(c.g., Harcup ct al. 1984).
The ~nultiplcrccaplure of individunls in this
Discussion
study shows that fishcs displayed complex, or exrrioverncnt hchuvior. Coupled with thc
The rcsults of this study do no1 cntirely support plo~-ulo~-y,
the rcstrictcd movernent paradigm us defitled by tinding [hat there arc no distinguishing characterGcrking (1959). Although the ~niijorilyof recap- istics of lishes that wcrc mohile, this behavior sugtured tish wcre in their homc puuls, some mobile gests that rnoverncnl plusticily was nt the individfish also wcre recaptured. Thc mobile fish wcrc ual level. At any onc timc, a proportion of the
collcctcd predominately iu pools adjacent to homc population travclcd outside of thcir homc pool arid
pools, hul some individuals traveled as many as that proportion was constantly turning over.
'I'his type of' hchavior rnay not hc cxccplional.
I0 pools away. Thcsc rcsul~sagree with othcr
rnoverncnt studics in showing that strcam tish pop- McVcy (1955) reported complex movernent bculations co~uprisca large static group and a srnall havio~-for lhree species o f ccntrarchids that wcrc
mobile group (Funk 1957; SLott 1967; Bruylants ~rccapturedscvcral limes in a lakc. Fifty percent of
ct al. 1986; Heggcncs c~ al. 199 1 ; Ereerilnn 1995). hluegills Lrpotnis rrrrrrmr~hit-usrccupturcd twice or
There was no tcmporal inlluence on the vagilily more displayed complex triovcmcnt. Fwly and 25
of fish populations in this sludy. Ratios of non- perccnt 01' multiple longear sunfish and greet1 sunrnohile to mohilc ~rccap~urecl
fish did n o t change fish, rcspcclively, exhibited complex movement.
significantly hclwccn the collecting trips of late Largc brown 1ro~ltwere dctcclcd shifting back and
May to mid-October, arid movcmcnt did not in- forth f1-om daytime covcr s t r u c l ~ ~ r etos nighttime
crease as time hrtween tagging and I-ccap~urein- covcr slruclures in u strcum (Clapp et al. 1990).
Powcr ( 1984) supgcslcd that "temporary migrant"
creased.
We found no diffcrcnccs in length or wcight loricariids left home pools to nsscss alternative
between thc nonrnohilc and rnobilc recaptures for habitats and [hen returned; shc also provided cvthe study species. I'revio~is invcstigalors desip- idcncc that fishes can rcmcmber habitats and that
natcd stalic und rnobile groups within a population, this capacity allows lishcs LO evaluate thcir home
suggcsling [hat these groups wcrc discrek (Gownn pool huhitnt in rclutjon to other pools. Asscssmenl
ct al. 1994). Prior reports indicutcd that the rnobilc of filternative homc sites during rcgulal- cxploraconiponenl consisted o f displaced intermcdiutc- lory trips provides an opportunity for recolonisized or less-fit individuals (Funk 1957; Bruylants zalion (Frccmun 1995). Ootnplcx movement e m ct al. 1986; Hcggcncs cl HI. 1991; Gowun und hles species thut are corisidercd scdcnlary, such as
Fausch 1996), that lai-gr fish were nori~nobilc(La].- longenr sunfish, lo recolonize dcfaunakd areas
Although [he number of our recaptured fishcs
ilnorc 1952). thal large fish wcrc mobile (Cierking
1953; Tyus and McAda 1984), 01- that the non- was small, we assume, using [he "slicc in lime"
mnbilc and mobile groups did not differ (Stott approach, that we obtained a represcntutivc sample
1967; Rerra arid Gunning 1972). Still olher studics of what was occurring. Because wc uscd only one
implied thal nonmobile and mobile fishes wcrc not sumpling rnethod, however, larger individuals may
SMITIISON A N D J O I I N S T O N
..............
...
........
..
......
375
....
..........................
.......
....
. .
.
.
.
....
.
..
. . .
I
.................
. . . . . .
-
>-3133
-250
-200
-150
-100
-50
0
50
100
.
.
150
.
.
200
250
>300
Distance Moved
I'loul<r.4. -1;rcqucncy dis~rihulior~
o I tlislmcr moved ( m ) by rccapturcd grccn nnd Inngcnr hnnfishrs (combined)
have bccn overrepresented in our samples. Low
rccupturc riitcs, a common characteristic of markrecapture st~ldiesof Gshcs, suggcst mnrkcd fishes
move out of the stlidy area (Gowun et al. 1994).
In our study, this hypothesis was not supported.
Rccupturc ratcs did not differ between times the
sludy site was hydrologicully isolated and times it
was 1101. Kecaplure ratcs incrcuscd us the study
progressed, supporting the theory of rcstl-iclcd
movcmcnt. A less-than-perfect recapture I-ate is
due to sumpling incfficicncy, fish mortality, arid
thc ahscncc o f thosc fish making cxploratory or
permanent trips oil1 of (he collecting Lirca. Annual
natural ~nortalityof bluegills can be 56% (Ricker
1945). The pcrccnl of sunfish making cxploratory
trips away from a home pool in our study was 1214%,so the percentage of lagged fish available for
recapture may have been as low as 30% if mortulity ratcs upproachcd that for bluegill. The final
ovcl-all ~-ccaptul.crutc obtaincd in August 1996 was
37%. This suggesls we ohtaincd good rcsults from
our sumpling efforts. We conclude lhal the samplcs
collected provided reprcscntutivc data on the
lnovcrncnt behavior of the study species.
The reslrictcd movcrncnt paradigm does not fully define movements of stream lishcs. In many
home range studies, home range is dclincd as thc
nrca truvclcd routinely (c.g., Berra and Gunning
1972; Hill and Grossman 1987) and hcncc as the
TAIII.I:
4.-Movcmcnl behaviors of Littlc (ilar.ypcau
Crcck suntishcs rccup~uretlIwo or more rimcs.
L~I-ca
whcrc the majority of recaptures occur. Somc
studies suggcst pools can be considered discrete
units (Gerking 1959; Malthcws 1986). Regular explorutory ~novcments,however, exlend lhc scale
of imporlancc of hubitut bcyond that of a home
pool. Homc rangc studies should encompass Lhc
range of movcmcnt hy the nlobilc component,
which may also rcprcscnt common movement behavior. The purpose of exploratory movcmcnts
may hc to carry out some rouline process that rcquircs a dit'fcrcut location. For example, Clapp c i
al. ( 1990) found that brown trout required a variety
of habitats for a variely o f purpcwcs. In addition
Lo dcfining the scale of habitat importance, routine
exploratory movclncnt of strcarn fishes is a mechanism for recnlonixation hccuusc it provides fishes
an opportunity to assess altcmalivc bubitats. Sheldo11 and Meffe (1994) concluded that rccolonization i s a rcsult of high mobility and "density-proportional cquilibriurn." Recolonization occurs by
irnrnigralion balancing cmigratinn with a continuing turnover.
This sludy demonstratcd rclativcly restricted
rnovctucnt of stream fishes, suggesting [hat the sclected species spcnd thc majority of their lime in
u honlc pool hut make visits inlo udjuccnt pools.
This behavior irnplies that lishcrics managers
should consider larger areas of habilal as critical
to stream lish populations. Additional information
about the frequency and cxtcnl of cxploratory
mclvcn~cntis needed in order to predict thc rcsponsc of fish populations to disturbance, thc impact of stocking fish, and the spatial requirements
nccdcd to manage a species.
Acknowledgments
We thank R. Adams, J. Barlletl, D. Curlson, J.
Chambers, K. Damon, C. Figiel, W. Haag, K. Klcincr, E. Ixighton, B. Libman, G. McWhirtc~; K.
T E S T O F KLSTRICTET) M O V E M E N T P A R A D I G M
r,l,rrki) in a srn;lll. coaslal stream. Canadian Journal
o r Fisheries and Aqualic Sciences 48:757-762.
Hill, J., and G. D. Grossman. 1987. Home range llir
thrcc Norlh A~ilcricnnstrcaln tishcs. C'opcin 1987:
376-378.
References
Johnslon, C. E . , and F.. H . Smi~hson.1999. Kelenlitin
of pnssivc intcgrntcd trnnspondcr (PIT) tags for inBerm, T. M., nnd C;. E. G u ~ ~ n i n g1972.
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tin1 slnhility o r cullhroa~ lroul ( O n c ~ o r h p r ~ r ~ h ~ ~Nalurillist
~
Overstreet, L. Smithson, and M. Villcgas for field
assistnnce. This project was fi~ndcdby thc U.S.
Forest Service.