Streamflow Regulation and Fish Community Structure

Author(s): Mark B. Bain, John T. Finn and Henry E. Booke

Reviewed work(s):
Source: Ecology, Vol. 69, No. 2 (Apr., 1988), pp. 382-392
Published by: Ecological Society of America
Stable URL: http://www.jstor.org/stable/1940436 .
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Ecology, 69(2), 1988, pp. 382-392
? 1988 by the Ecological Society of America

STREAMFLOW REGULATION AND FISH

COMMUNITY STRUCTURE'

MARK B. BAIN2 AND JOHN T. FINN

Department
ofForestry
and Wildlife
Management,University
ofMassachusetts,
Amherst,
Massachusetts
01003 USA

AND

HENRY E. BooKE
Massachusetts
Cooperative
Fishery
ResearchUnit,
3 University
ofMassachusetts,
Amherst, Massachusetts
01003 USA

Abstract. Many regulatedstreamsare characterizedby highlyvariable and unpredict-

able flowregimes.Since changes in streamflowdirectlymodifyphysicalhabitat,streams
withsuch highlyvariable flowsprovide highlyunstableaquatic habitats.We evaluated the
effectof artificialstreamflowfluctuationson streamfishcommunitiesby comparingfish
densities,in species and habitatgroups,betweentwo riversdiffering in daily flowregime:
one witha naturalflow,and one withhighlyregulatedflows.
We developed a simple model describingthe relationshipbetween available stream
habitatand its use by 15 species or size classes of fishin the naturalriver.Species and size
classes that used a specificset of microhabitatconditions were identifiedby comparing
habitatcharacteristicsforsamples withand withouteach typeof fish;forfishthat used a
particulartypeof microhabitat,we groupedspecies and size classes accordingto similarity
in microhabitatuse. Next, we categorizedstreamhabitatsamples in both the naturaland
regulatedriversinto groupson the basis of fishhabitatuse criteria.Fish densitiesforeach
fishand habitatgroupwere thenindividuallycompared betweenthe two rivers.
An abundant (>90% of all fish)and diverse (nine species) group of small-fishspecies
and size classes wererestrictedto microhabitatthatwas characterizedas shallow in depth,
slow in currentvelocity,and concentratedalong streammargins.This group of fishwas
reduced in abundance in the regulatedriverand absent at the studysite withthe greatest
flow fluctuation.Anotherfishgroup included species and size classes that used eithera
broad rangeof habitator a microhabitatthatwas deep, fast,or both,and was concentrated
in midstreamareas. The densityof fishin thisgroupwas higherin the regulatedriverand
peaked at the siteswiththegreatestfluctuationsin flow.Highlyvariable and unpredictable
flowregimesappear to be a high-frequency disturbancethateffects fishdifferentlydepending
on the way theyuse streamhabitatand acts to reduce communitycomplexity.
Keywords: community ecology;disturbance;environmental multi-
microhabitat;
variabilityfishes;
variateanalysis;streamflow
regulation;streamhabitat.

INTRODUCTION a changein streamflow(total streamdischarge)trans-

Flow modificationis one of the most widespread
human disturbancesof stream environments(Fraser
1972, Ward and Stanford 1983), and its effectsare
poorlyunderstood(Holden 1979, Sale 1985). The flow
regimesin manyregulatedstreamsand riversare high-
ly variable and unpredictable;typical seasonal flow
variations may occur weekly or even daily, with no
For anyspecificlocationin a stream,
regularperiodicity.
' Manuscript received 20 October 1986; revised 15 June
1987; accepted 17 July1987.
2 Presentaddress: Alabama Cooperative Fish and Wildlife
Research Unit, 331 Funchess Hall, Auburn University,Au-
burn,Alabama 36849-4201 USA.
3Cooperators: United States Fish and Wildlife Service,
MassachusettsDivision of Fisheriesand Wildlife,Massachu-
settsDivision ofMarine Fisheries,and theUniversityofMas-
sachusetts.
lates into a change in the water depth and velocity.
Also, as dischargechanges,streamsubstrateand cover
objectsbecome associated withdifferentcombinations
of waterdepthand velocity.Consequently,changesin
streamflowcan be regarded as modificationsin the
physicalcomposition of the aquatic habitat. In addi-
tion, changes in the patternof flow variability(flow
regime)can be viewed as an artificialdisturbanceof
streamhabitatstability.
Zonation, or downstreamchangein fishcommunity
compositionwithinriversystems,has been associated
withgradientsof habitatchange (Kuehne 1962, Shel-
don 1968, Hynes 1970,Hawks 1975, Barilaet al. 198 1).
In streamsof similar size, physicalhabitat character-
istics(waterdepth,currentvelocity,substrate)are im-
portantfactorsinfluencingfishcommunitycomposi-
tion (Gorman and Karr 1978, Moreau and Legendre

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April 1988 FISH COMMUNITIES
1979, Savard and Moreau 1982, Schlosser 1982a, b,
Moyle and Vondracek 1985). Studies of the role of
habitatstabilityin determiningfishcommunitystruc-
ture have affirmedthe significanceof both seasonal
(Kushlan 1976, Schlosser 1985) and long-term(Ho-
rowitz 1978) steamflowvariability.In general,stream
fishcommunitystructure is stronglyinfluencedby both
habitat composition and stability,where habitat sta-
bilityis a functionof flowregime.
We compared the fishcommunityand habitat re-
lationshipsin two rivers:one withartificialshort-term
flowfluctuationsand one with a natural flowregime.
The comparisonwas made on thebasis offishdensities
by species and habitatgroups.This approach enabled
theidentification ofdifferencesamong streamfishden-
sitiesthataccounted forboth the directeffectsof flow
variability(i.e., habitat variability)and the way dif-
ferentfishused streamhabitat. Our work contributes
to researchon the effectsof artificialdisturbanceson
structureand functionof natural biological systems
(the "disturbancephenomena" of Pickettand White
1985). In one of our rivers,where disturbancesof a
natural magnitudeoccurredon an artificialtemporal
scale, we used community-levelstructuralcharacter-
isticsto detectfishresponses.
STUDY AREAS
This studywas conductedon the lower segmentsof
two fifth-order tributariesof the ConnecticutRiver.
The West River, in southernVermont,is an unregu-
lated streamwitha mean annual dischargeof 29 m3/s
and a basin area of 1092 km2at the studyreach (2.8-
10.5 km upstreamfromthe mouth [42?52' N, 72033'
W]). The WestRiverwas selectedas a "natural" stream
typical of Connecticut River tributariesin physical
structureand fishspecies composition.The Deerfield
River, in westernMassachusetts,has an artificialflow
regimewith a mean annual dischargeof 41 m3/sand
a basin area of 1407 km2at the studyreach (13.2-20.2
km fromthe mouth [42034' N, 72?35' W]); short-term
flowregulationhas been extremeforat least 50 yr.
The West and Deerfieldrivers drain adjacent wa-
tershedsof similar climate and geology (Brooks and
Deevey 1963). Althoughthe two studystreamsdiffer
in mean annual dischargeand drainagearea, theyare
similarin generalsize and appearance and fall in the
medium-size river categoryof the river continuum
framework ofVannote et al. (1980). Both studyreaches
have a highgradient(3 m/km),primarilycoarse sub-
strate(bedrock, boulder, cobble), and littleinstream
cover such as logs and debris. Water in both streams
is highlyoxygenated,soft(10-27 mg/Ltotal hardness,
30-9 5,gSconductivity),and warm(maximumsummer
temperature27QC). Both studyreaches were immedi-
ately upstream of a transitionfrom a high-gradient
streamto a low-gradientfloodplainriver(Connecticut
River) characterizedby low-velocitychannelswithfine
substrate,backwaters,and abundant instreamcover.
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IN REGULATED STREAMS 383
The same fishspecies occurredin bothofthetwo study
streams or in the nearby section of the Connecticut
River,withthreeexceptions:browntrout(Salmo trut-
ta) and rainbow trout(S. gairdneri)were stocked an-
nuallyonlyin the DeerfieldRiver,and parrofAtlantic
salmon (S. salar) were stocked only in the Deerfield
River in 1983. A detailed comparison of the physical
conditions,chemistry,and fishfauna of the two rivers
and theirdrainagebasins was given by Bain (1985).
Althoughboth streamshad essentiallythe same an-
nual flowregime(bimodal; dischargepeaks in spring
and fall)and responseto basin runoffevents,the study
reachescontrastedsharplyin dailyflowvariability.Flow
fluctuations on theDeerfieldRiver occurredfrequently
(several times daily) and varied greatlyin amplitude
(Fig. 1). They were caused by unpredictableand vari-
able-volume water releases (a response to short-term
electricalsupplyand demand in New England) froma
series of fourhydroelectricdams (?21 m high) with
shallow,short(?2 km) impoundments.Flows fluctu-
ate throughoutthe year except in springand briefly
afterheavy rainfallor snowmelt,when riverdischarge
exceeds hydroelectricplant capacities. The flowfluc-
tuationsare typicallymost pronouncedduringthe dry
period of summerand early fall when no wateris re-
leased duringthe frequent3-h shutdownswhen the
impoundmentsare being refilled.During these shut-
downs, dam leakage and tributariesmaintaina small
flowin the studyreach. The severityof flowfluctua-
tions becomes attenuateddownstreamfrom the hy-
droelectricfacilitiesbecause an abruptrise and fallat
the downstreamdam (km 21.2) becomes a relatively
gradual change at the end of the studyreach (Fig. 1).
Consequently,there is a gradientof decreasing flow
variabilityin the study reach. Physicochemicalcon-
ditions do not vary substantiallyin response to flow
changes (Bain 1985) since impoundmentshold little
waterand are regularlydrained and refilled.
The West River has a naturaldaily flowregimethat
appears essentiallyconstantwhen compared withthat
in theDeerfieldRiver. In theabsence ofa runoff event,
thereis almost no detectablechangein riverdischarge
over a 24-h period (Fig. 1). When runoffevents do
affectriverdischarge,the rate of streamflowchange is
much more gradual than the artificialflowchanges in
the DeerfieldRiver.
METHODS
Field sampling
Four studysites(81-231 m long)wereselectedalong
each studyreachin each streamto representriverhab-
itatin thevicinity.At each site,six permanenttransects
were selected to proportionatelysample the general
within-sitehabitat(e.g.,riffles,pool). On each transect,
fourevenlyspaced sample locations(23 M2) wereiden-
tifiedbeforesamplingbegan:one adjacentto each bank,
one-thirdacross thetransect,and two-thirdsacross the
384 MARK B. BAIN ET AL. Ecology,Vol. 69, No. 2

8
WEST RIVER (km 20.2)

4_

01 I , l - I I

32 DEERFIELD RIVER (km 21.2)

E 12

2
32 _a
DEERFIELD RIVER(km14.8)

22 -

12-

21 A
2400 1200 2400 1200 2400 1200 2400

6 AUG 7 AUG 8 AUG

AUGUST 1983
FIG. 1. A representative3-d recordof streamdischargeforthetwostudyrivers.The WestRivercurve(toppanel)is
basedon data froma UnitedStatesGeologicalSurveystreamgagelocatedupstream fromthestudyreach.The WestRiver
was smallerat thislocationthanat thestudyreach,buttheflowpattern was thesame.The DeerfieldRivercurveswere
constructedfromhydroelectric generationrecordsofthedam upstreamfromthestudyreach(middlepanel)and data from
a streamgagein thelowerportion(bottompanel)ofthestudyreach.The largerwaterreleasein theDeerfield Riveron 8
August1983corresponded to a heavygenerationperiodthatoccurred
at irregular
intervals
duringmostweeks.WestRiver
dischargeremainednearlyconstantduringtheperiod,sincerainrunoff had no significant
effects
on eitherriver.River
kilometresin parenthesesindicatethedistancefromtherivermouthto thedischarge measurement point.

transect.Althoughthe sampling design was not ran-
domized, potential investigatorbias was reduced by
this presitingof samples.
The 24 sampling locations at each study site were
sampled between midmorningand late afternoonon
days of typical seasonal flow conditions. Periods of
very large water releases in the DeerfieldRiver were
avoided to maintain samplingconsistency.The eight
studysiteswere sampled once duringeach of fourpe-
riods: 6-22 October 1982 and 1983, 30 June-30 July
1983, and 14-23 August1983, and thefourWestRiver
sites were also sampled once between21 Julyand 15
August 1982.
At each samplinglocation, we collected fishin the
23-iM2 area, eitherby using a propositionedarea elec-
trofishing device (described and evaluated by Bain et
al. 1985a) or, in deep-waterareas, by snorkeling(11%
of total samples). The electrofishingdevice consisted
of an electrodeframe(7.6 x 3.0 m) poweredby a 460-
volt alternating-currentgenerator.The electrodeframe
was positionedby manipulatingropes attachedto the
cornersand then leftundisturbedforat least 15 min.
In collectingfish,two personswearingwaders and car-
ryingvarious dip nets approached the frame from
downstreamto a pointat whichthe frameinteriorwas
visible.A thirdpersonthenactivatedthepowersupply
froma remotelocation.All fishoriginating fromwithin
the framewere collectedand the interiorof the frame
was thoroughlyinspectedat least twice. All fish >20
mm total lengthwere identifiedand measured.
When water exceeded wader depth (0.9 m) we vi-
sually counted fishin a 23-iM2 area. Wearing a face
mask and lyingon a surfmattress,one of us (M. B.
Bain) positionedhimselfovera samplinglocationwhile
attachedto a steelcable suspendedacross theriver.He
countedthefishbyspeciesand estimatedlengths,which
werelateradjustedby usinga regressionequation (r2=
0.98, 28 pairsofknownand estimatedlengths).During
all snorkeling,visibilitywas more thanadequate to see
thestreambottomclearly(maximumwaterdepth,230
cm). Althoughwe did not compare electrofishing and
snorkeling,otherinvestigatorshave shown that,when

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April 1988 FISH COMMUNITIES IN REGULATED STREAMS 385

visibilityis good, visual countsyieldresultsconsistent 1982, Finger 1982, Orth and Maughan 1982, Moyle
with those obtained by other sampling methods and Vondracek 1985), we divided some species into
(Northcoteand Wilkie 1963, Goldstein 1978, Sale and size classes by inspectingall possible bivariateplots of
Douglas 1981), especially when discrete areas are habitatvariables vs. fishlength.
searched (Keast and Harker 1977). First,we determinedwhichfishspeciesor size classes
Four habitat variables were measured at the center used a specificset of microhabitatconditions,usinga
ofeach 23-iM2samplinglocationafterthefishhad been multivariateanalysis of variance (MANOVA) test to
sampled. Waterdepthwas measuredto thecentimetre detectdifferences in habitatcompositionbetweentwo
with a weightedfiberglasstape measure. Surfacecur- groupsofsamples:thosewitha particularspecies(pres-
rent velocity was measured by timingthe float of a ent) and those withoutthatspecies (absent). However,
wood chip (38 x 19 x 19 mm) attached to a 1-m such a testposes an importantproblem; the presence
lengthof nylon twine. Surface currentvelocitymea- of a species in a sample indicatessuitable habitat,but
surementswereconverted(log/logregressionequation its absence does not necessarilyimplyunsuitablehab-
[r2 = 0.81], 119 pairs of measurementswitha surface itat. It seems reasonable to expect that not every 23-
wood chip and a Price AA currentmeter set at 60% m2 sample of suitable habitat will be occupied at the
depth) to mean velocity(velocityat 60% of depth),a time of sampling.Consequently,a species that uses a
formthathas oftenbeen reportedin otherstreamhab- specificmicrohabitatwill be found in a less variable
itat studies (Probst et al. 1984, Glova and Duncan set of conditions than that of the available habitat.
1985). Two streamsubstratevariables,coarsenessand While MANOVA is conservativewhen one group(ab-
heterogeneity, were quantifiedwith a 2-m lead-core sentsamples) has both a largesample size and disper-
rope divided into 20 10-cm sections(method of Bain sion (Green 1979), we checked all test resultsby dis-
et al. 1985b) by the same personthroughoutthe study. criminantanalysis group classification(as in Rice et
The 20-segmentrope was stretchedacross a sampling al. 1983) and logisticregression(Fienberg1980). Only
location (perpendicularto the current),and the dom- the MANOVA resultsare reportedbecause theywere
inant substratein each 10-cm section was recorded supportedby the alternativestatisticalproceduresin
visually as one of the followingtypes(modifiedfrom all tests.
Cummins 1962): 1 = smooth horizontalsurfacessuch Next we formedgroups of species and size classes
as bedrock,sand, or silt; 2 = gravel (2-16 mm diam- that used similar microhabitatsbased on mean posi-
eter);3 = pebble (17-64 mm); 4 = cobble (6 5-256 mm); tions (habitatspace centroids)in the available habitat
5 = boulder (>256 mm); 6 = irregular bedrock. The space. Correlationsamong the fourhabitat variables
mean of the 20 dominantsubstratevalues was used as were removed by principal components transforma-
an index of substratecoarseness. The standarddevia- tionofthehabitatdata because thistechniqueprovides
tion was used as an index of substrateheterogeneity. an objectiveformatforinterpreting species-habitatuse
patterns(Rotenberry and Wiens 1980,Carnesand Slade
Identificationoffishcommunityand 1982). Groups were identifiedby usinggroupaverage
habitatrelationships clusteranalysis of species centroidsin all fourtrans-
We grouped fishand habitat to produce a simple formeddimensions.
model describingthe relationshipbetween the fish Our finalstepin describinga fishcommunity-habitat
communityand available streamhabitat.The purpose model was to categorizehabitat on the basis of fish
of this dual categorizationapproach was to enable us grouppresenceor absence by directdiscriminantanal-
to make comparisons between the two riverson the ysis (Nie et al. 1975). Using West River data, we de-
basis of fishdensitieswithinhabitatcategories.To de- veloped a discriminantfunctionand then classified
velop the fish community-habitatmodel, we con- samples fromboth rivers.Althoughthis approach ap-
ducted three separate sequential analyses to answer pears circular,we wanted to compare quantitatively
threequestions: Which fishused a specificmicrohab- fishdensitiesbetweenriversand among sitesin a man-
itat? Can the various typesof fishbe grouped on the nerthataccountsfordifferences in habitatavailability
basis of similarityin microhabitatuse? Can the sam- and use ratherthan analyzing differencesby species
ples of streamhabitatbe groupedto reflectthe typeof along.
fishthat use the habitat?This approach is analogous
to categorizingstreamhabitatintopool and riffletypes Differencesbetweenriversand among sites
to analyze separatelythe abundance of pool and riffle Fish densities (number captured per 23 M2) were
fishes,except that we relied on statisticalresults to compared quantitativelybetween rivers and among
identifyfishand habitatcategories. sites, independentlyfor each fish-habitatcategory.
We used fishsampling and habitat data fromthe Densities were computed by studysite and sampling
West River to identifythe fishcommunity-habitat re- period foreach group of fishin each habitatcategory.
lationships,since it had a natural flow regime.Inas- Tests for differences between riversand among sites
much as microhabitatuse patternsrelatedto body size withinriverswere made by nonparametrictwo-way
have been reportedformany streamfishes(Bain et al. (riversand sites) layouttests(Kruskal-Wallis)because

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386 MARK B. BAIN ET AL.
therewere largedifferencesin groupvariances.A pos-
teriorimultiple-rangetests,based on Kruskal-Wallis
mean ranks,were used to identifydifferences among
sites withineach river.
RESULTS
During this study,we captureda total of 3039 fish
and coded names of Z
of 23 species (common, scientific,
fishesare givenin theAppendix).Three species (small- 0
mouthbass, rockbass, whitesucker)weredivided into
two lengthgroups(largeand small; size rangesin Ap- C.)~~~~F
pendix)because theiruse ofhabitatwas relatedto body
size. Of the resulting26 species and size classes, 15
SLOW
were abundant enough to allow statisticalanalysis of Z
habitatuse characteristics. P-.41
Fish communityand habitatrelationships
A significantdifference(MANOVA, P < .05) was
foundin habitatcharacteristics(some combinationof
depth,velocity,and substrate)betweenfishpresentand
fishabsent samples for 13 of the 15 fishspecies or size
classes fromthe West River. These 13 species or size
classes werethereforeidentifiedas usingsome specific
set of the microhabitatconditionswithinthe available
rangeof habitat conditions.We consideredthese fish
to be microhabitatspecialists.Because no differences
(P > .10) in habitatcompositionwere foundbetween
presentand absent samples forlargesmallmouthbass
or American eel, we identifiedthese fishas habitat
generalists.The analyses for bluegills and pumpkin-
seeds werelimitedto thelate summerand fallsampling
periodsand to the two sitesfarthestdownstreamsince
these fishwere presentin the lower West River only
duringthesetimes.All MANOVA testsincludedsam-
pling period as a factor(two-wayMANOVA) to test
for seasonal differences.Because all seasonal and in-
teractiontermswerenonsignificant (P > .29), we pooled
the data forall samplingperiods.
Clusteranalysisoffishcentroidsforthe 13 specialist
species and size classes indicatedthatnine used a sim-
ilartypeofmicrohabitat:small smallmouthbass, small
rock bass, bluegill,pumpkinseed,small whitesucker,
blacknose dace, fallfish,mimic shiner,and tessellated
darter.To illustratetheclusteranalysisresults,we offer
a two-dimensionalprincipalcomponentsplot (Fig. 2)
in which the available habitat is bounded, primary
originalaxes are superimposed,and fishcentroidsare
plotted.The nine groupedfishused a restricted habitat
characterizedby very shallow and slow water, and
coarse substrate(i.e., the boulder- and cobble-strewn
edge of the river).In addition, this group constituted
an importantportion of the fishcommunitysince it
was veryabundant(>90% ofall fishcaptured),diverse
(9 of the 15 common species), and composed exclu-
sivelyof small fish(see Appendix). The analysisto this
pointidentifiedan abundantand diversegroupofsmall
fishthat used shallow, slow microhabitat(hereinafter
called the SS GUILD).
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Ecology,Vol. 69, No. 2
WEST RIVER
+4
+4~~~~~~~ DEEP
FINE I
FAST
LRB
TD *+
SWS LMB
~ ~ B *LND
S
COARSE
C 4 PRINCIPA
SHALLOW
CMMSH N I+
-4
-4 +
~PRINCIPAL COMPONENT I
FIG. 2. Distributionof 13 microhabitatspecialists(habitat
centroids)withinthe total available West River habitat (en-
closed by heavy solid line). The extremesof the originalhab-
itataxes primarilycomposingprincipalcomponentsI and II
are superimposedto assist in interpretation.The substrate
heterogeneity axis is not shown because it is the least impor-
tantofthefouroriginalaxes and orientednearlyperpendicular
to the two-dimensionalplane illustratedhere.The nine types
offishidentifiedbyclusteranalysisas theshallow-slowhabitat
guild (SS GUILD) are enclosed by a lightsolid line. Because
in the clusteranalysis of centroidswe used substratehetero-
geneityas well as the three habitat variables shown, the
relativeproximityofcentroidsis notfullyshownin thisfigure.
The fourfishspecies and size classes that were not
partof the SS GUILD (largerockbass, longnosedace,
largemouthbass, largewhite sucker)used microhabi-
tats otherthan the shallow, slow-watertype (Fig. 2).
For example, largewhite suckersused a microhabitat
characterizedby verydeep and fairlyswiftwaterwith
relativelyfine substrate(i.e., at the heads of pools).
American eel and smallmouthbass, the habitat gen-
eralists,werenotincludedin theclusteranalysisor Fig.
2 because theyused habitatbroadlyand therefore over-
lapped all otherspecies and size classes.
To group habitat,we classifiedthe 480 West River
samples by direct discriminantanalysis into habitat
used by SS GUILD fish(shallow and slow, SS HAB-
ITAT) and habitatnot used by SS GUILD fish(deep
or fast,or both,DF HABITAT). We repeatedthisanal-
ysis usinga stepwiseprocedureto identifythe habitat
variablesmostimportantin distinguishing betweenthe
two groups. The stepwise entryorder,group means,
and discriminantfunctioncoefficientsindicated that
waterdepthand currentvelocityweretheprimaryvari-
ables distinguishingthe two typesof habitat(Table 1).
In summary,we simplifiedthefishcommunity-habitat
relationshipsby placingboth fish(the SS GUILD and
April 1988 FISH COMMUNITIES IN REGULATED STREAMS 387

TABLE 1. Means, discriminantfunctioncoefficients, and relative importanceof habitat variables for distinguishingWest
River habitatused (fishpresent)and not used (fishabsent) by SS GUILD fishes.

Group means Discriminant

function
Habitat variables* Guild present Guild absent coefficient Relative importance
Water depth (cm) 30.17 58.61 0.754 veryimportant
Currentvelocity(cm/s) 8.14 21.45 0.756 veryimportant
Substratecoarsenessindext 3.74 3.42 -0.371 less important
Substrateheterogeneityindex? 1.18 1.07 0.018 not important
* Habitat variables are listed in the orderselectedby stepwisediscriminantanalysis.
t Mean of 20 coded dominant substrateobservations.
? Standarddeviation of 20 coded dominant substrateobservations.

fishof otherspecies and size classes [non-SS GUILD])
and habitat (DF HABITAT and SS HABITAT) into
two categories.
The discriminantfunctiondeveloped withthe West
River data forclassifyinghabitat as SS HABITAT or
DF HABITAT was used to classifyDeerfield River
samples. Classificationresultswereinspectedby study
site and samplingperiod. In both rivers,slightlymore
samples were classifiedas SS HABITAT than as DF
HABITAT, althoughboth categorieswerealways well
representedat each site.
Comparisonsbetweenriversand among sites
For each studysite on each river,the densityof fish
was computed by type of fish (SS GUILD, non-SS
GUILD) and habitat(SS HABITAT, DF HABITAT).
Comparisons betweenriversand among sites on each
riverwere made independentlyforeach fishand hab-
itat combination.Samples (23 m2each) were counted
once foreach type of habitat,and each fishtype was
counted (numbersper 23-iM2 sample) separately.
For SS GUILD fishin SS HABITAT (Fig. 3A), fish
densitywas greaterin theWest River thanin theDeer-
field River (P ' .001). There were differences(P =
.012) among West River sites; at site 4 (the farthest
upstream site) the densityof SS GUILD fishin SS
HABITAT was lowerthanat theotherthreesites.This
resultwas largelyaccounted forby the movement of
largenumbersof bluegillsand pumpkinseedsinto the
lower West River duringlate summerand fall.A sim-
ilar migrationdid not occur in the Deerfield River,
even thoughthese fishwere also presentin the lower
sectionof thatriver.However, SS GUILD densityre-
mained higher(P = .001) in the West River than in
the DeerfieldRiver even withthese species excluded.
Thereweredifferences (P = .008) amongDeerfieldRiv-
er sites; the upstreamtwo sites with the greatestfluc-
tuationsin flow(sites 3 and 4; Fig. 3A) had fewerSS
GUILD fishin SS HABITAT than the downstream
two sites. The SS GUILD fishwere rare at Deerfield
River site 3 and absent at site 4. In contrastto these
sites,the densityof SS GUILD fish(primarilyfallfish)
at site 2 in the DeerfieldRiver was similarto that in
the West River (Fig. 3A). A large tributarywith high
densitiesof SS GUILD fish(M. B. Bain et al., personal
observation)emptied into the downstreamend of a
pool partlyincludedin site 2. The highdensitiesof SS
GUILD fishat DeerfieldRiver site 2 may have been
largelydue to thisimmediatesourceofSS GUILD fish.
For SS GUILD fishin DF HABITAT, there were
greaterdensities(P = .039) in the West River than in
the DeerfieldRiver, althoughthe densityof these fish
in thishabitatwas low in both rivers(Fig. 3B). There
wereno differences (P ? .142) in densityof SS GUILD
fishin DF HABITAT among siteson eitherriver.Since
the SS GUILD and DF HABITAT combinationis a
mismatch of fishand habitat type,we expected low
densitiesat least in the West River. However, the lack
of significantlygreaterdensityof SS GUILD fishin the
DeerfieldRiver clearlyindicated that these fishwere
not shiftingto DF HABITAT. Such a shiftin habitat
use, had it been found,could have explained the re-
duced densityofSS GUILD fishin SS HABITAT iden-
tifiedearlier(Fig. 3A).
The densityof otherspecies and size classes (non-
SS GUILD fish)in SS HABITAT (Fig. 3C) was greater
(P = .003) in the Deerfield River than in the West
River. There were no differences(P = .597) among
West River sites,althoughthereappeared to be mar-
ginaldifferences (P = .091) amongDeerfieldRiversites.
In the DeerfieldRiver, the trendtoward highernon-
SS GUILD densityin SS HABITAT at the sites with
the greatestflow fluctuations(3 and 4; Fig. 3C) was
primarilyattributableto American eels.
The densityofnon-SS GUILD fishin DF HABITAT
(Fig. 3D) was greater(P < .001) in the DeerfieldRiver
thanin theWest River. There wereno differences (P =
.105) among West River sites,althoughthereweredif-
ferences(P = .010) in theDeerfieldRiver. In theDeer-
fieldRiver, the densitiesof non-SS GUILD fishwere
high at the most severelyflow-fluctuating upstream
sites (3 and 4; Fig. 3D). The high densitiesat sites 3
and 4 resultedprimarilyfroma combinationof rela-
tivelylargenumbersof Americaneels, and largewhite
suckers.
In summary,the most notable resultswere compar-
ativelyhighdensitiesof SS GUILD fishin SS HABI-
TAT in the West River and the eliminationof these
fishin the DeerfieldRiver at the site withthe greatest
fluctuationsin flow.The reduced densityand absence

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All use subject to JSTOR Terms and Conditions
388 MARK B. BAIN ET AL.
of these fishin SS HABITAT in the DeerfieldRiver
could notbe accountedforbya shiftto DF HABITAT.
For fishotherthan those of the SS GUILD, densities
were greaterin the DeerfieldRiver than in the West
River regardlessof habitat type, and densities were
highestat the upstream sites with the greatestflow
fluctuations.
DISCUSSION
and streamhabitat
Fishcommunity
Althoughstreamfishcommunitystructurehas been
summarizedon the basis of trophicgroups(Grossman
et al. 1982, Angermeier and Karr 1983), foraging modes
(Horowitz 1978), life historycharacteristics(Mahon
1984), and ecomorphology(Gatz 1979, 1981, Mahon
1984), our habitat-basedapproach provides a frame-
work for assessing the effectof artificialhabitat dis-
turbances.Habitat has been identifiedas the primary
basis on which many biological communitiesare or-
ganized (Schoener 1974), and numerous studies have
supportedthisgeneralizationforfishcommunities(e.g.,
Gibbons and Gee 1972, Wernerand Hall 1976, Werner
et al. 1977, Tallman and Gee 1982, Schlosserand Toth
1984). Our statisticallyderived fishcommunity-hab-
itat model reflectsa simple relationshipbetween the
West River fishcommunityand the available stream
habitat.An abundant and diverse group of small-fish
species and size classes (SS GUILD) were restrictedto
microhabitatcharacterizedprimarilyas relativelyshal-
low in depth,slow in currentvelocity,and concentrated
along stream marginsin rifflesand pools. Fish that
were not part of the SS GUILD were relativelyscarce
and used eithera broad rangeof habitator some com-
binationof deep or fastwatersthatoccurredprimarily
in midstreamareas of riffles(shallow but fast),pools
(slow but deep), or riffle-pooltransitions(deep and
fast).
The fishcommunity-habitatmodel developed here
conflictswiththetraditionalview ofstreamsas a linear
sequence of riffleand pool habitats. It emphasizes a
bank-midstreamorientation,since shallow and slow
habitatis concentratedalong the streamedge in both
riffles and pools. In many studies(e.g., Sheldon 1968,
Gibbons and Gee 1972, Finger1982, Schlosser 1982a,
b, Tallman and Gee 1982, Angermeierand Karr 1983,
Herbold 1984), fishuse of streamhabitatwas consid-
ered withrespectto riffles (generallyshallow and fast),
pools (deep and slow), and transitionzones (interme-
diate). Althoughthe importanceof a bank-midstream
habitat orientationhas been noted for invertebrates
(Cummins 1964, Peckarsky1983) it has not been di-
rectlyidentifiedforfishesin high-gradient streams.The
difference in fishand habitat orientationbetweenthe
presentstudy and those cited above may be due to
differences in streamsize or the relative size of sam-
plingunits. In headwaterstreams(orders 1-3 of Van-
note et al. [1980]), small fishuse most or all oftheriffle
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Ecology,Vol. 69, No. 2
SS HABITAT DF HABITAT
(shallowandslow) (deepandfast)
12 5
A B
56
/ '
o I A. .~ 0
1 2 3 4 1 2 3 4
5 5
C D
0
1 j
1 2 3 4 1 2 3 4
SITE SITE
FIG. 3. Medianfishdensities rangesfor
and interquartile
fishandhabitattypebysiteand river.@-@ datafromthe
WestRiver,and0- - -O datafromtheDeerfield River.The
Y axis is on a largerscale fortheupperleftfigure (A), and
theseverity fluctuationsin flowon theDeerfieldRiverin-
creasewithsitenumber, i.e.,in themoreupstream sites(X
axis).
habitat, since midstreamareas generallyhave mod-
erate velocities and depths; as stream size increases
(medium-sized streamsand larger,orders >4), large
concentrationsof small fishmay become increasingly
restrictedto streammarginsbecause midstreamareas
are eithertoo deep or too fast,or both. Glova and
Duncan (1985) describedthispatternforvarious-sized
channelsin a largebraided river:many small fishwere
evenlydistributedacross small channelriffles but were
restrictedto stream margins in large channels. Al-
thoughthe pool vs. riffleapproach has been usefulin
many studies, it would have resulted in an entirely
differentand less distinctpatternin our study.
Althoughour generalapproach differed fromthatin
most otherstudies,many details of the fishcommu-
nity-habitatmodel that we developed here are sup-
ported by the work of others. Like Sheldon (1968),
Gorman and Karr (1978), and Moyle and Vondracek
(1985), we found water depth and current velocity to
be the most importanthabitat variables affecting fish
distribution.Studies in Illinois (Schlosser 1982a, b),
New York (Finger 1982), and California(Moyle and
Baltz 1985) identifieda patternsimilar to that in the
WestRiver:shallowand slowhabitatwas used bysmall,
youngfishof several species, and deep areas were pri-
April 1988 FISH COMMUNITIES IN REGULATED STREAMS 389

marilyinhabitedby larger,older fish.Also, our data that had the greatestfluctuationsin flow.This differ-
and analyses were sensitive enough to place fish in ence representsa major change in species, size class,
distinctcategories(SS GUILD and others) that were and numericalcompositionof a riverfishcommunity,
previouslyidentifiedas distinctonlyin termsofhabitat especiallysincethisgroupconstituted> 90% ofthefish
use: small and large white sucker (Finger 1982) and in the unregulatedriver.Some evidence suggestshow
small and large smallmouthbass (Probst et al. 1984). the fluctuating habitataffectsthis guild. Flow changes
The separategroupingof blacknose dace and longnose displace the shallow shorelinezones, forcingfishre-
dace, whichwereidentifiedby Gibbons and Gee (1972) strictedto these areas (the SS GUILD) to relocate to
as distinctlydifferent in microhabitatuse, was partic- maintaintheirspecificset of habitatconditions.Rapid
ularly notable because the longnose dace is a small decreases in flowpose a threatforany fishnot able to
species that uses fasterwater than that used by the move withtherecedingshoreline.On severaloccasions
abundantsmall SS GUILD fish.Consequently,thefish duringthe fieldworkin the DeerfieldRiver,we saw SS
community-habitatmodel developed here incorpo- GUILD fishstrandedand trappedin isolated puddles
rates many recognized stream fishcommunitychar- afterrapid decreasesin riverdischarge.Kroger(1973)
acteristics,even thoughwe did not relyon a rifflevs. reportedsimilarobservationsforsmall fishin a highly
pool habitat classificationbut instead used statistical fluctuatingsection of the Snake River, Idaho. Also,
methodsto identifygroupings. rapidflowincreasesmightexpose fishoftheSS GUILD
Althoughmost fishtend to stronglyprefera specific to increased predation,since shallow shorelineareas
typeof habitat(Hynes 1970, Gorman and Karr 1978, become accessible to larger piscivores as depth in-
Moreau and Legendre 1979, Moyle and Li 1979, Sa- creases, much as fluctuatingreservoirlevels increase
vard and Moreau 1982, Moyle and Vondracek 1985), thesusceptibility of shallow-waterfishto consumption
our resultssuggestedthattwo (large smallmouthbass, by piscivores (e.g., Heisey et al. 1980). Power et al.
American eel) were habitat generalists.In the West (1985) and Schlosser(1987) have clearlydocumented
River, the other 13 of 15 species or size classes used thatlargepiscivores(Micropterusspp.) forcesmall fish-
specifictypesof microhabitatthat could be quantita- es into shallow-waterrefugiain streams.If unstable
tivelyand effectively (a < .05) described.Consequent- shallow-waterhabitatsprovide less effective refugefor
ly, it cannot be assumed thatphysicalhabitat charac- small fish,then the functionalvalue of the habitat is
teristicsare importantin determiningthe distribution lostand a reductionin SS GUILD densitywould occur.
of all fishspecies in streams. In contrastto SS GUILD fish,the densityof other
species and size classes responded positively to in-
Effectsofflowfluctuations creased habitatvariability.As a group,these fishwere
Hydroelectricpowergenerationhas imposed a high- comparativelydense in theregulatedriverand densest
ly artificialflow regime on the Deerfield River. The at the siteswhereflowfluctuationswere greatest.This
frequency(up to severaltimesa day) and rate(minutes positiveresponsewas similarin SS HABITAT and DF
to hours,dependingon location of changein the Deer- HABITAT, and was a reflectionof both habitat spe-
fieldRiver) of fluctuationswere most uncharacteristic cialists, such as the large white suckers,and habitat
of a natural stream.Also, flowfluctuationswere un- generalists,such as the American eel and large small-
predictabledue to the complex mix of factors(such as mouth bass. Increased abundances of relativelyold,
electric demand, regional power supply, short-term large fishalso are associated with periods of frequent
nonhydroelectric capacity,upstreamreleases) thatde- naturalflowchanges(Schlosser 1982a). Increasedden-
terminepower generationschedules in the Deerfield sities of generalistfisheshave been associated with
River basin. Consequently,the aquatic habitat in the large flow changes (Harrell 1978) or other riverine
DeerfieldRiver was highlyunstable. Because natural modifications(Goldstein 1981). In theDeerfieldRiver,
environmentalinstabilityplays a criticalrole in shap- t~hehabitatof non-SS GUILD fishis changedless than
ing communitystructure(Wiens 1977, Connell 1978), thatofSS GUILD fish,fora givenflowchange,because
increased environmentalinstabilityshould non-SS GUILD fishuse eithermidstreamhabitats(deep
artificially
also affectcommunitystructure.Our analyses dem- or fastareas in riffles, pools, and transitionareas) or
onstratedclear differences in the fishcommunitiesbe- all types of habitat. However, we developed no evi-
tweenthe unregulatedWest River and the highlyfluc- dence to indicate which factorsaccount forincreased
tuating Deerfield River, and these differences densityof this group: food availability mightbe in-
correspondedqualitativelyto differences among sites creased as a resultof reduced consumptionof inver-
along the gradientof flowvariabilityin the Deerfield tebratesby the SS GUILD, increased availability of
River. invertebratesdue to flow fluctuations,or increased
The shallow-and slow-waterfishes(SS GUILD), an availability of the small longnose dace (a non-SS
abundant and diverse group of exclusivelysmall fish, GUILD fish)as the preyof piscivores.
were adversely affectedby the artificiallyhigh vari- The rivercontinuumconceptofVannoteet al. (1980)
abilityin flow.This group was reduced in abundance characterizes streamsof the size in the presentstudy
in the DeerfieldRiver and was absent at the studysite (fifthorder)as providingmoderatelyvariable environ-

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All use subject to JSTOR Terms and Conditions
390 MARK B. BAIN ET AL. Ecology,Vol. 69, No. 2

ments.Connell's (1978) intermediatedisturbancehy- Massachusetts

Divisionof MarineFisheries,and theUni-
pothesis associated high community diversitywith versity
ofMassachusetts.
Thispaperis contribution
number
105oftheMassachusetts
CooperativeFisheryResearchUnit.
moderatelevels of environmentalvariability.Connell
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APPENDIX
TABLE Al. Common name, abbreviationcode, scientificname, and rangeoftotallengthforspecies capturedin the Deerfield
River and West River, 1982-1983.

Code Total-lengthrange
Common name (used in Fig. 2) Scientificname (mm)
Common species
Americaneel Anquillarostrata 130-670
Blacknose dace BLKND Rhinichthysatratulus 21-54
Bluegill BG Lempomismacrochris 30-109
Fallfish FF Semotiluscorporalis 20-148
Largemouthbass LMB Micropterussalmoides 40-110
Longnose dace LND Rhinichthyscataractae 20-144
Mimic shiner MMSH Notropisvolucellus 43-61
Pumpkinseed PS Lepomisgibbosus 42-161
Rock bass Ambloplitesrupestris 30-202
Small, <100 mm SRB
Large, > 100 mm LRB
Smallmouthbass dolomieui
Micropterus 20-411
Small, <?100 mm SSMB
Large, >100 mm
Tessellated darter TD Etheostoma
olmstedi 26-98
White sucker Catostomus
commersoni 22-444
Small, < 150 mm SWS
Large, > 150 mm LWS
Uncommon species
Atlanticsalmon Salmo salar 124-192
Brown bullhead Ictalurusnebulosus 100-126
Brown trout Salmo trutta 189-356
Common shiner Notropiscornutus 73-78
Longnose sucker Catostomus catostomus 32-98
Rainbow trout Salmogairdneri 124-192
Sea lamprey Petromyzon marinus 62-111
Spottail shiner Notropishudsonius 43-63
Yellow bullhead Ictalurusnatalis 34
Yellow perch Percaflavescens 101-188
White perch Moroneamericana 49

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