JamesH. Johnsonr
Nez PerceTribal FisheriesDepartment
P.O. Box 365
Lapwai, Idaho 83540
Comparative Diets of Paiute Sculpin, Speckled Dace, and
Subyearling Steelhead Trout in Tributaries of the
Clearwater River, ldaho
Abstract
beld;ns;) , spe&led dace (Rbinichthls osculu\ , and,
The food habits of Peiure scL pin \ 6ttui
subyeariing steclhead t.our lSalnn gandneti) were examined during summer and carly fall when
r h e v c o e * i a i i n r i f f l e s . A r o t a l o f 1 q 6 P a i u t es c u l p i n . 1 8 6 s p e c l l e d d a . c . d n d l t 4 s u b ; e a r i n g s r c r l h t a d
trout from six tributnrier of the Cl€.arwar.c Rjver, jn North Centrat Idaho, wc.c used for analysis.
Paiute sculoin fed exclusrvely on autochthonous material whereas steelhead utilized both autochthonpercent) foods Detripefcenr) end allochthonous (10-58 percent, -:14
ou! (+2-8i percenr, i:65
ru< 14t'--- peftenr, x-oo perccnr/ was rhe major i"od ir rhc dier of the.pe(L cd dat. Oterlap io
and \ubvc.rling steelhead
diet of specllcd dace with both Paiute sculpin (0.r02-o.il6, i:0.I6rt
(0.062-0.250, i-0.161)
was very low. Diet overlap betweeo Paiute sculpin and subyearling steelhcad was moderate (0.489 0.5t1) in five oI the streams and hish (0.869) in one. The data suggest
tbat during diurnal periods there is definite dclineation io the foods caten by these three species
when they coxist in riffle habitats durirg the summer and fall.
Introduction
Competition for food resourcesmay be facilitated when differeot fish speciesoccupy
rhe same microhabitat. In rhese instances,competitive inreractionsmay be reduced if
sympatric{ishes1) feed on <lifferenttypes or sizesof Prey, 2 ) feed at differeot locations
in the water column (i.e., bottom, mid-water, surface), or 3) feed at different times
of rhe day. Of these three categories,feeding at different times of the day may reduce
interspecificinteractioosbetweenspeciesbut may not substantiallydiminish competition
sincea similar diet may st-illdeplete a sharedfood resoutce.
In tributaries of the ClearwaterRiver, Idaho, the domiflant fish assemblagein the
streamsthrough the sllmmer and early fall consistso{ Paiute sculpin (Cottas beld,ingi),
speckleddace (Rhinichthy: otculus), and,subyeadingsteelheadtrout (Salmo gaidner'i)
(Kucem e, al. \983). During this peiiod these three sPeciespredomioantly occupy
riffle areas.The utilization of tiffie habitat by Paiute sculpin, speckleddace, and subyeading sreelheadtrour has led to a number of investigationsexamining interactions
betweeo these specieswith othet coexisting tiffle inhabitaots (Dietsch 1959, Hannuksela 1973, Li aod Moyle 1976, B^ltz et ttL. L982, Finger 1982). The focr:s of this
study was ro urilize both the information gained in evaluatingsimilarity in diets between
thesespeciesin ClearwarerRivet tributaries plLrsthe availabledata io the literatute to
assessthe potential for competition *'hen these speciescoexist in riffles.
Methodsand Mateiials
Fisheswere collectedfrom I Ausust to 6 Octobet 1982 in six tributaries of the ClearlPresentaddress:Departmentof Fish and \Y/ildlife, i06 S. \{/. Mill Street,P. O. Box 1501, Porc
land, Oregon 97208.
Northwest Science,Vol. t9, No. 1, 1985
water River in North Ceotral Idaho. The six streams,Bedrock Creek,Big Canyon Creek,
Cottonwood Creek, Jacks Creek, Laprvai Creek, and Mission Creek, dischargeinto the
ClearwaterRiver within rhe Nez PerceIndian Reservation( Fig. 1). A generaldescriptioo of rhe collecrion sire of each srteam is presentedin Table 1.
Fisheswere presetvedin 10 percent formalin after being stunned with an electroshocker.Diet composition was determined on a dry weighi basis from estimatesdeDWORSHAK
EDROcx CR.
RESERV
NEZ PERCE
IND,AN
RE5ERVAIION
Figure 1. location of the study streams within the Nez Perce Indian Reservation. Idaho.
rived for individual prey taxa. \Thenever possibie,aquatic invertebrateswere identified
to family, terrestrial invertebratesto order, and fish remains to species.
Diet overlap berween Paiute sculpin, speckleddace,and subyearlingsteelheadtrout
in eachstreamwas calculatedusing the equation of Horn ( 1966) :
s
a r - )
s
)X"
i-1
S
. ! . . v
s
l-:
i:l
Yr"
vrhere C,f,: overlap val re
s: food categories
X; - proportion of the total diec o{ fish species
X conoibuted by food caregoryi
Yi.- ptoportion of the total diet of fish species
Y conrribureJby i.,o,lcaregoryi
IThen employing this equation diet ovedap vahies (C),) can range from 0 (no
overlap) to 1 (complete ovedap).
Johoson
TABLE
mouih), average stream width
1. Siream lengih (km), collection location (km from slrem
Ior siuilv sbearns antl sla(Co), a r waier velocitv (cm/s)
(m), maximum temDeraiure
at each collectioD locnlion for stream wiilih, ma'ximum t€mp€rlrure an'l
tions. Estimries
condilions
velociy represent late sumer
15
Bis Canyon Creek
Cotionwood Creek
TABLE
26
13
45
34
2
14
7
E
25
22
23"
23.
2'.1o
26"
?. Number oi tish examineal, mean leneith a-nd size ra-nae (TL,
River, Idaho.
r.om six tributaries ol the CleaNatcr
Beahock Creek
08/03/82
Steelhead 0+
Lapwai
)20 '
3.ri
9.1
3.6
2,3
5.5
3.5
c-reek
08/0:t/lJ2
Sieelheaal0+
09/23/42
Sieelheaal0+
0r/28/32
Steelhearl 0+
Bis Canyon Creek
70/01/82
CononwoodCreek
10/06/82
speckled Dace
12
40
20
55
60
83
27
10
20
43
16
2l
30
40
24
29
20
26
mm),
25.9
34.6
17.8
74.7
33.9
19_5
and date oi collection,
50-?
6E.3
?0.6
65.9
68.0
63.0
64.0
72.0
71.8
66 . 1
77.7
70.3
'1J.L
73.4
84 . 2
65.0
57.4
90.5
44-63
58,?8
59-83
53-97
54-87
53-E5
46-98
63-?9
60-92
54-89
45-94
71 92
59-70
62-83
66-100
4$ 102
4 0- E i i
?6-102
Resulls
The diets of 196 Paiute sculpio, 186 speckled dace, and 194 subyearling steelhead trout
from six ClearwatetRiver tributarieswete used for aflalysis(Table 2) The three species
were collectedin sufficient numbersto quantify thafu diets in each of the six sffeams'
Sample sizes from individual streamsconsistedof a low of 57 fish from Jacks Creek
to a high of 198 fish from I-apwaiCreek ( Table 2).
Paiute scr:lpin fed exclusively oo autochthonousmaterial. HydloPsychids (150(17.1'28-5 percent, --237
Percent)'
40.9 percent,V-27.2
Percent), chirooomids
percent) were the ma,or prey consumedaod
aod baetids (4.4-25.1 percent,x:14'5
were the only taxa that were rePresentedin the diet of Paiute scllpin in each of the
six streams ( Table 3). Additional benthic taxa which were imPoftant in the diet of
Paiute sculpin were perlodids and elmids ( four streams), tipulids (three streams),
pedids (two streams),and sialids (one stream). Fisheswere only found in the diet of
Paiute sculpin in one srearn. In Big Canyoo Creek fishes comPosed 41.4 percent of
the diet of Paiute sculpin. The speciesis apparently caooabilistic since approximately
80 perceot of the fish remains in the diet (one-third of the total diet of Paiute scu\>in
in Big CaoyonCreek) was identified as Paiute sculpin.
The majot component in the dia of speckled dace was detritus (46.1-77.0 percettt'
Percent)
x-65.5 percent) ( Table 4). Chironomidlawae (.3.1-17.5
Perceot,i:117
Comparative Diets of Paiute Sculpin
TABLE
3 r'erceni at.v weisht dietarv composiuon of paiute sculoin in rributaries oi the crerrwater
River, Tda.ho.Only those Drey taxon nhich composed at teast Z percent of the aliei of
Paiute sculpin in one or more of ihe st eams :!re inclurleal.
Bic
Carwon
Coilonwootl
Creek
8.4
Baetialae
Hepl:rseniidae
UnidebiiJied
25.0
E.1
9.1
7_4
19.4
3.3
9.2
3.4
10.4
12.7
15.9
25.1
2.9
2.0
Brachycentridir"e
clossosomatidae
Helicopsychidae
HydroDsychiilae
Limnephilidae
2.5
4.1
0.3
40.9
326
2.8
1\7
*
;
;
23-0
15.0
;
Sialidae
9.0
Elmiilae
Diplera
Chironomida€
Simuliidae
Tipulidae
3.6
(L)
21.4
17.7
12.6
28.3
0.8
1.2
13_3
0.6
7.2
19.0
26.9
2Ii.5
1.1
C. beldingi
Unirlentiiied
;
TABLE
4 Pe.ceni drv wekht ari€rarv composirion or speckred arace in tributades oI ihe clearvaier
F,iver, Idaho. Onty those Drey tdon which comDoseit ai teast 2 percen! ot lhe cliet oi
speckted itace in one or more of the streMs are incluileat.
Beilrock
Perlodidae
Baeiidae
Heplaseniidae
Unidenlilied
;
kLp'wai
6.9
10.9
1.5
24.0
1.4
4.3
14.2
9.4
Tipulialae
Detriius
77.0
7.1
57.5
1.9
rl'lxon
2.2
Johnson
Bic
Canyon
Creek
5.4
Creek
4.9
5.5
5.2
4
Mission
Creek
5.2
Hydropsychidae
Dipiera
(L)
Chironomidae
'I€rrestrirl
8.3
2.2
;
5.6
17.5
\0.7
11.3
'1.1
65_5
12.\
3.1
;
72.7
0.5
were the principal benthic prey consumed and were present in the diet of fish from
all six sueams.Baetid nymphs were the secondraokiog prey (0-24.0 perceot,x-9.9
p€rce[t), occurliog in the diet of dace in five of the sffeams. Hydropsychids ( four
sueams), perlodids and tipulids (two streams) and tabanids (one stream) also contributed substantially to the diet of speckled dace. Allochthonous foods were alrnost
completelyabsent from the diet.
Subyeading steelhead trout fed heavily on allochthonous material. Terlestrial inpercent) of the diet of subverrebrat€scomprised from l0.l-57.7 petcent (x:342
yearling steelheadin rhe six streams(Tabte 5). The main terrestrial iovertebratesconpercent), coleopterans(i:6.5
persumed by steelheadwere homopertans (x:8.7
(x-4.6
percent), and hymenocenr), arachnids
oercent), hemiprerans (x:4.0
pterans (- : 3.7 percetrt). No single order of terresffial invertebrates were represented
in the diet of steelheadin each of rhe six streams.Ctrironomids (14.1-19.7 percent,
TABLE
5- Perceni dry weisht dietaxy coq$silion
ol subyearlins steelheaal troul in iribuiaries
Liver, Idaho, Only those prey iaxon which composed at leasi 2 perceni
Clearwater
iliei ol subyearling sieelheaal trout in
ot the sirea,ms axe includeal.
Bedrock
Creek
B.],eiid,r.e
HeplaseDiialae
Unideniiiied
Hydropsychidae
Unideniili€il
Di era
(L)
Chironomidae
(P)
Chironomida€
(A)
Chironomidae
LaDwai
Creek
Mission
Creek
0-5
1.3
Bic
Canyon
Creek
12.3
30.7
1.1
5.7
1 4
2.6
13.2
8.6
2.$
9.3
2.\
9.2
16.4
0.5
11.7
12.5
L.2
16.3
3.7
0.4
19.5
7.t
2.]
7.2
$6.1
11.4
2.2
t't.7
5.5
7.3
0.9
6.9
6.5
::
;
*
42.3
56.2
12.6
10.8
32.5
1.5
3.2
3.3
0.6
2.0
15.u
44.7
2.7
79.2
1.3
;
3.6
Detrilus
55.8
6.5
69.3
1_3
E1.3
4.2
7.5
3.1
89.9
Taxon
Diplera
;
Total Terrestrial
I-LaIvae;
Coltonwooal
Creek
13.6
TiDulidae
Scaiophasialae
Terrestrtd
oi the
oI the
P-Pupa
57.7
; -A.
9.9
2.7
1.4
9.2
3.5
-
1 40
'1.6
t*
0.9
15.4
8.7
;
;
;
0.4
5.6
7.9
3.3
18.?
;
3,3
10.1
AdulL
C-omoarativeDiets of Paiute Sculoin
x:15.8
percent), tnetl.ds (5.7-19.2 percenr, x:13.8
percent), and hydropsychids
().2-16.4 petcerc, x:11.4
percent) were the primary aquatic invert-ebratesutilized
by subyeadingsteelhead,
Diet qvedap was estimated berween the three species for each of the six srrearns.
Becauseof the large contribution of derirus in the diet of speckleddace, diet overlap
values between dace and Paiute sculpin (0.102-0.336, x- 0.167) and subyeading
steelhead (0.062-0.25O,x-0.161)
were very low (Table 6). Diet ovedap between
TABLE
Paiute
6. Dietary overlap values
head lroul in lfbuLaries
Scuipin-SDecklial
(Cl.) amons Paiute sculpjn, sleckled
of the Clearwaier
River. Idrho.
Dace
Bis
Canyon
Cotlonwood
Faiute
SculDin -SLeelhead 0 +
Steelhead 0+ -SDeckleil
Creek
Creek
Cotton$-ood Creek
Big Canyon Creek
Dace
Coilonwood Creek
Big Canyon Creek
dace, and subyearlins
sreol-
0.3?6
0.207
0.12u
0.120
0.111
0.t02
_
X = 0.16?
0.869
0_553
0.552
0.5t2
0_51?
0_489
_
0.250
0.247
0.1?E
0.145
0.085
0 . 06 2
_
=
X
0-161
Paiute sculpin and subyearling sreelheadwas moderate (0.489-0.553) in five of the
streams.Mutual predation on pedodids, baetids, hydropsychids,and chironomids by
Paiute sculpin and steelheadin Cottonwood Creek resulted in the only high diet overlap value obsewed,0.869.
Discussion
Collections of Paiute sculpin, speckled dace, and subyearling steelhead in the six Cleaiwarer River tributaries generally reflected a similar pattern of distribution of these
species io the basin as had been previously reported ( Maughan 1976, Maug$at and,
Saul 1979). Observationsof microhabitat urilizarion indicated that during the period
when collectionswere made all three speciespredominated in riffle areas,The utilization of riffle areasby Paiute sc'ulpin (Jones 7972, Moyle 1976, Finger 1982), speckied
dace ( Moyle 1976, Schreiberand Miockley 1981, Baltz et al. 1982), aod subyeading
steelheadtrout (Hartman 1965, Johnsonand Ringler 1980) is well documented.
The most thorough examinationof the diet of Paiute sculpin in a lotic environmenr
was colducted in SagehenCreek,C-alifornia (Dietsch 1959). In SagehenCreek benthic
insectscomposedapproximately 99 percent of the diet of Paiute sculpin with various
prey taxa being seasonallydominant in the dier. Diersch (1959) afso found evidence
that differencesoccurred berweenthe diets of small ( ()!mm)
and large ( )80mm)
sculpins.
Johffon
Paiute sculpins in the Clearwater River tributaries, like those io SagehenCreek,
fed most heavily on aquatic insects,did not utilize terrestrial invertebrates,and except
i n B i g C r n y o nC r e e k s. e r e n o t p i s c i r o r o u . .
Becauseof the difficulty in separatingalgal {rotr detritai material from the stomachs
of speckleddace for dry weight analysis,both of these items rvete grouped as detritus.
Jhingran (1948) found detritus to be a majot comPonentin the diet of speckleddace
during summer. Io the fall, dace have been shown to feecl on unidentifiable fibrous
material (Jhingran 1948) and algae (Li and Moyle i976, Pedenand Hughes 1981).
However, nlost investigatorsexaminiog the food habits of speckleddace describethe
minnoq. as a predator on benthic insects (Jhiogran 1948, Peden and Hughes 1981,
Schrieberand Miockley 1981,llakz et al. 1982)- The principalbenthictaxa that have
been observedin the diet of dace are baetids,hydropsychids,and simuliids.
Subyearlingsteelhead(rainbow) trout feed nost heavily on benthosduring summer
and early fall (HaonLrksela1973, Tippets and Moyle 1978, Johnsonand Ringler 1980,
Johnson 1981, Johnson and Johnson 1981). Horvever,at least duting diuroal periods,
terrestrial invertebrateshave been repolted to comprise a substantial Portion of the
diet of subyeatlingsteelhead(Johnsoo and Rinlller 1980, Johnson 1981, Johnson and
Johnson 1981). Although most of the diet of subyearlingsteelheadtrout in Clearrvater
River tributaties was composedof autochthonousmaterial, allochthonousprey irems
(*- 34 percent) conrtibuted a considetableamouot to the diet.
Only one of eighteenobservationsof diet ovedaP belweeo Paiute sculpin, speckled
dace, and subyeatling steelheaduout io Clearwater Rivet tributaries exceeded0.56.
Diet overlap valuesof 0.60 or more ar geenerallythought to indicate similar cliets (Zaret
and Rand 1971. Fuller aod Stewart 1977). The data show that there is defioite delineadon io the foods eatenby these three speciesduriog diurnal petiods when they occupy
rhe same habitat in Ciearwatet River tributaries. This situation cor-rldptevail from
shortly {ollowing steeiheademergencein lune until terrestrial invertebraresbecomeless
availabie,possibly during mid to late autumfl. Becauseof the scarcity of detrital and
allochthonousmaterial, diet overlap between Paiute sculpin, speckled dace, and subyearling steelheadoverrvinteting in tiffles would be expectedto be higher than observed
in this investigation. However, since food consumption rates are diminishecl at this
time and standing crops of benthic insectsare generailyat rheir Peak (Hynes 1970),
competirion for food betrveen these benthic fishes might not be severe. By spring
juvenile steelheadrvould no longer be ptesent io high densitiesin riffles; they would
be occupyiog areashaving more covei.
Diet overlap between speckleddace and Paiute sculpin in Cleatwater River tributaries was substantiallylower than irad been previously reported betrveen dace and
two other speciesof sculpins.Li aod Moyle (i976) and Bahz et al'. (1982) for.rnda
moderate overlap in diet between speckleddace with Pit sculpins (C. P;te iist rnd
rif{le sculpins (C. gxlons). These studies reported that bodr dace and sculPins fecl
similarly on benthic insects.Basedupoo the benthic comltsjtion of the diet of speckled
dace, Jhingran (1948) judged that dace were competitors with subyearling chinook
salmot (Oncorbynchtr tibduytJcba) and steelheaduout. However, the larger contriburion of detrital material in the diet of speckleddacein this study led to low overlapsrvith
both sculpin and steelhead.Hannuksela (1973) obserwedthat the summer diets of
moftled sculpins (C, baitcti) and jrlenile tainbow trout s/ere simiiat in a tributary ot
Lake Sr:perior.Except for the larger contribution of allochthonousmatetial in thc cliet of
C,omparativeDiers of PaiuteSculpin
steelheadin Clearwater River ffibutaries, results of the two studies are comparable.
Besides seasonalvariation in diec composition, anothet factof ro conside{ when
evaluating either diet ovedap or competition between fishes is diel feeding periodicity.
Paiute scuipio are mosr acive at night (Fioger 1!g2) and ur" r"port".lly no.rornul
feeders (Moyle 1976). Li and Moyle (1976) showed speckleddace to be primarily
a_diurnal feeder with peak feeding occurring at 2000 h.
lohnson (19g2) reported an
identical peak (i.e., 1600-2000h) for blacknosedace (R-. atrarltJ/tr in a tributary
of
)
Lake Ontario. The food consLrmptionof subyearlingsreelheadhas been shown
to increasesubstantiallyfrom 1200-2000h wirh peak feeding occurring from 1600-2000
h
in a tributary of the SalmonRiver, New york (Johnsonand
Johnson 19g1). Thesedata
indicate that Paiute sculpin are nocturnal feederswhereasspeckleddace
ancl subyea ing
steelheadtrout are principally diurnal feeders.Since the peak food coosumition
oi
Paiute sculpin occursar a different time than either specklJ dace or subyeading
steelhead trout, competitive interactions for food may be reduced between these
species
and sculpins.The large contribution of terrestial invertebratesin the diet of
subyelrfing
steelheadsuggeststhat rhis speciesis feeding heavily on drift. Conversely,the
detrial
and benthic componenrsin the diet o{ speckleddaceindicate that this speciesis
primarily
a benthic forager. Differeoces in foragiog areasof dace and steelheadtherefore
could
offset any competitive interactionscrearedby diurnal feeding peaksfor
both species_
Acknowledgmenls
I wish ro thank P. A. Kucera, M A. Bear, K. Carter, D. R. Cross,and C. L.
W.ebbfor
their assistarice
in the fieid, J. A. Gould fot typing the manuscript, and E. Z.
Johnson
for reviewing the manuscript.
LileratureCited
Balrz, D. M, P. B Moyle, and N. J. Knight. 1982. co_mpedtive interactions
between benthic stueam
-e;; "i;;f:
rai g
i
llli,':;tlf l'r;:Yla::€ot ot*s'""a'piit"i-J"e;ii-;;;i;;;'";;;;;
,.*.
g;,;:%:ti;,fil";"frlf#,',oi,itfn:il
ll-'"*1*f,li';tr""::,)\il,,i,i:?iiii}:;"?l
.,"*"r,
sesrcgarion
arnongrhreespecierof iculpin r'c.,rrlr). Cop€ia1o82
,,j,i;Ud,o6rr.r.''reracrive
l"i:*:t*yi:5i#,:g#'i:illi)"n?$.if
rht#:T':il::'::::,^,.:;,,'X
; ,'ib;;;;y
;if;k" iupeior
Michi
bef,.
l$:a*::.x1.:,:R?l
B?l:,J.rjr,:
f6:;!,",.i"
Harrman, c. F. 196s. The role of behavior in the ecology aod, interaction
of urderyearting coho
t::::r:?,i#:!,lt{:"r
h;sutcb)^ndstcerhead
troii a"t*iiit,iii,i.
Hotn, H
r. risi. Res.-B;a;;
,.ovedap,
S
Measuremenr of
in comparative ecological studies. Am. Nat. 100:
17966
H . , ! . - N . . l o ^ - u . .T h e f c o ' o g y o , R u n n i n a W a r c r s . U n i \ . T o r o n r o p r e s s ,
Toronro.
lllnes.
J n r n g r a n . - v . - ( r . ^ r , ) 4 U , n c o n r f l b u - r j o nr o r h e b i " l o g l o f r h e K l a m a r h 6 l a c l d a c e ,R h ; a i , h t h ) , n , , , t l x \
rtanathen t {Lvermann & MFFkt. StdnfordUniv.. palo AIro,
Crlir.. Doctoral Diqierrarion.
"n"'?":'*#
#o'1;:B*":iiif
,f:d#iffir .lr;r:*;:*iiix+:+iiil","t"*$:il
150-161.
-;"f'"1;t?B:1#::'€:":'fi1fl^""f
-'
iz !'6tyf;:
(Rhinichthr
ctace
d'ratlhu)in a 'irib'|r'
^nd E. z. Iohnson. 1981. Feedingperrodrcitv.and,drel variation in dicr composition
of sub-
'i*u*J ii"Li;')a:'zraner'l)
ir a small
-;?lj1".1'"'Hff:'1""";',.]fin'\ixr.'Tff.:'i.:"i:;,rio*a n
T;f,lJi'j""'.j;ig;,1'?,:,11:",:l'i:#:?jlffl,l
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Comparative Diets of Paiute Sculpin