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Vertical Stratification ol Bat Activity in an Old-Growth Forest in Western

Washington

Article in Northwest Science · March 2000

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John P Hayesand JeffreyC. Gruver,Department
ol ForestSc ence 32lDRichardsonHa,OregonStateUnversty,
Corva s, Oregon9733T5752

VerticalStratificationol Bat Activity in an Old-GrowthForestin Western

Washington

Abstract
We exannnedthc amounl and lcnporal paltemsof bai acti\'ity at fouf differenr heightsin an old growth coniler lbrest at the Wind
Iliver Canopy Cra.e RcscarchFacilit) in soulh'ceDtralWashington.Anal]sis of 2,304 bat passesshosed thll| rnouni of activity
dificrcd antong !crticel slruta.For Mrorir bats,actiritl' was grearestin the lower canopy.lbllowed by the ground-lelel and uppcr
canopy.rcspeclilely. \Ve did nol detectactivity ofM_\rrlr abovethe canop!. Non rl rrir bals usedlowerand uppercanopiesmorc
licquend)- than grouDd-leveland abovethe canopy.Temporalpattcms ofacd!i1) generully e\hibited a bimodal disiriburion. bul
fie exlerl ol bimodalit) and the time and relative siTe of peakr difcrcd $jth speciesSroup and arnong heighls. Acli\'ity $as
grcatcslat Sround level earl) in ihe night and latefshifted tohigherstrala. Pallcmsoluse oi old growth fofesls bv batsmavrcilccr
Ihc corrrplc\ !crdcal slruclureoflhe vegetationin thoseforests.Exclusive rse ofground bascdequipnent can fesult in an incom
pleie picturc of lhc aclj\ iry ol bal\ in c|)lrplex tbrest stands.

Introduction within lbreststands.Becausebatsusespacethree-

The verticalandhorizont;ilspatialpatternsot trees
and other vegetationin forests (foresl structurc;
Oliver and Larson I 996) influencepatternsof bat
activity.In the Pacific Nofihwest,Thomas(1988)
foundold growthforcstssupponedhigherlevels
ofbat activitv than did young stands.and hypoth-
esizedthatdiftcrencesin levelsofactivity resulted
fron increased availabilityofroosts(largediam-
eter snags)in older forests.Humeset al. (1999)
reportedsimilarrcsults.and alsotbundthat lev-
els of activity diliered among young forests in
r e l r l i o nt o t r e eJ e n . i t Sl:r n o u n tl l ' h J t u ( t i ! i l ) i n
) i , u n ! . l . r n Ju\ i t h l o w d e n ' i t i e s o l t r e uc a s' in-
temrediatebetwccn young densestandsand old-
growth tbrests.Erikson( 1998)Iound that amount
olbat rctivity differedamongage classesofyoung
managedforcsts.HayesandAdam ( 1996)reponed
that activity of bats in riparian aleas in thc Or-
< g o nC o r . t R c n g er ru . i n l l r r rt r c e Jh 1 r e g e t a t i o n .
with clrarnatically lower ler,elsofactivity of nyotis
bats (Mt.rli.r sp.) in areasthat had been clearcut
and trrorcactivity of lalger bats(prim;rrily silver-
hairedbats,l,arlor_r'.rcrirr1.rclil,1rg.r?.r) in logged
areas.Sinilarly, in othcf regiens,activitvof bats
hasbeenshown to be influencedby forcst struc-
turc(e.g.,Fenlonct al. 1992.GrindalandBrigham
I991J,Kalcouniset al. 1999,KavanaghandBamkin
1995,Krusic et al. 1996,Zubaid 1993).
l n r d d i t r n t t h , . . i r r l l r r e r ru
r el si b r c ' t . t r u .
ture on activity of bats among forcst stands.fbr-
cst structure may also inlluence pattems of usc
dimensionally.lbrest structuremay influence
ve ical patlernsof bat activity as u'ell as hori-
zontal pattemsof use. Kalcouniset al. (1999)
c \ a m i n e d\ e r l i c : rnl r l l e r n \o l u \ eo f b l s i n r s p e n
(Poltulus tremuloi&:s). black spruce (Plcea
muriuntt).andjack-pine(PinuSbanksiuruilfor-
estsin the borealforest of centralSaskatche$,an.
ln aspcntbrests,they tbund thatM_r.rllswerc most
activewithin andabovethecanopyof aspenstands
and that there \vas more activity of hoary bats
(Lasiunrscinereus)abovethan below the canopy.
However,activitydid not significantlydi1l'erarnong
heights in spruce or jack-pine fbrests. Vertical
patternsof bat activity in coniferous tbrests of
the Pacific Nonhwest have not previously bcen
reported.
We examinedamountsand pattemsof activ-
ity of bats in difl-erentvertical strataof an old-
growth conittrous forcst in westernWashington.
BccauseoJd-growthfbrestsin theregionarestruc-
turally complex(Franklinand Spies l99l). we
hypothesizedthat useby batswould difler among
\c ical strata.In addition, we hypothcsizedthat
p a t t e mo' f r c t i ri t 1 u , ' u l JJ i l l e r a m o n g . p e c i c . .
Methods
This study was conductedat the Wind Rivcr
Canopy Crane Facility, located in the Thornton
T. Munger ResearchNatural Area and the Wind
River ExperimcntalForestin the Gifford Pinchot
National Forest.The sitc is in the southernWash-

102 Northn'estScience.Vol.7:1.No.2.2000
alnl)0h) rhc\rnh\e{S.eftili.A*o.ir!o. Alt'Llhsr...rr.d
ingtonCascadeMountains.nearCarson,Wash
ington.rt ln elevationof J55 m. Thc arcarcccircr
approximately250 cm of precipitationper year,
mostly occurdng as rain falling between Octo-
ber and May. The study site is an dd-growth fbr-
est with an ove$tory dominatedby Douglas-fir
(Pseudotsugamenaiesii) and westem hemlock
(.Tsugaheteroph La),with westemredcedar(Thl7a
pllczrta).Pacific silvcr lir (Ahiesamabilis). grand
fi (A. grandis), and westem white pine (Pinas
montit:oltt)being less abundant.Overstory trees
rangefiom 200 to ,100years old with maximum
heightsof 65 m. Understorytrcesinclude Pacific
yew (Ia,rns brevifolia) and Pacific dogwood
(Cornusnuttallii).T he most abundantunderstory
shrubs are vine maple (Acer circinatum), salal
(Gat theria sftrrllon). Oregon grape (.Berberis
nenusa).andCalifom i^h^zd'nut(.Coryluscomuta
tar. caliJornica).Thecanopy str-ucture ofthe site
has been describedas "bottom-heavy." indicat-
ing thatthemajorityofthe vegetationoccurswithin
the lower one quarter of the height of the stand
(Parker1997).The studyareais oenteredon the
Wind Rirer canop) errne.r con.lru(lion crrne
measuring74.5m in heightat thejib. The jib of
the craneextends85 m with a counterweightjib
extending35 n in the oppositedirection.Bat spe-
cies that are likely to use the site include
Townsends big-eared bat (Cory'norh!nus
townsendii). big brown bat (Eptesicus fuscus).
silver hairedbat. hoary bat. Calitbrnia myotis
(Mtotis caliJbrnicus),long-earedmyotis (M.
4,rtir),little brown myotis (M. lrcfirgr.r), fiinged
myolis (M. th)-sctnodes), longlegged myotis (M.
r,rlars), and Yuma myoLis(.M.)-umqnensis).
We usedAnabatIl bat detectors(Titley Elec-
tronics,Ballina,NSW,Australia),automatcdwith
dclay switches,to record echolocationcalls of
batsonto audiotapes(HayesandHounihan 199.1).
To account fbr potential diflerencesin sensitiv
ity of detecton (Larson and Hayes in prcss),we
calibrated detectors using a Transonix IX pest
repeller(Ryansby Mai1,LagunaHills. CA).
We establishedecholocationnonitoring sta
tionsat two pointsfor eachoffourheighh: ground-
level, thc lower canopy( l7 to 25 m abovethe
ground),the upper canopy(35 to,+0m abovethc
ground), and abovc the canopy (75 rn above the
ground). Locations of echolocationmonitoring
stationsabove the canop)' were constrainedby
lhe \tructureol thc crn('p) (ritne:\lilljons \\'ere
positionedat thc end ofthejib andcounterweight
jib (85 rn and 35 m from the crane's center.re-
spectively). Becauscthecranewasallowedto freely
"weathervane,"
spin. or during the night to pre-
vent damagewhen an operatorwas not prcsent,
locatiol] of stationsabovethe canopycould vary
both \'",ithinand among nights. We used the dis-
tancesto the ends of the jib and counterweight
iib to describetwo concentriccircles surround-
ing the baseof the crane.We located one moni-
todng station at randomly generatedcompass
bearingsalong each circle on the ground. lower
canopy,anduppercanopy.Stationson the ground
wereplacedunderplywood shelten to protectthe
equipmentfiom rain (HayesandHounihan1994).
Stationsin thelower anduppercanopywereplaced
in plywood shelterssuspendedbetweentwo trees
by rope. Stationscould not alwaysbe placedpre
cisely at the randomly generatedpoints becausc
of the positionof tree\hureach.tation\\ asposi
tioned within l0 m of the randomly generlted
point. Detectorson the ground and in the lower
anduppercanopywereorientedwith microphones
liL ing tou ard thearcr u ith thegre:rtcrropening.
Those above the canopy were orjentedwith mi-
crophonestacing perpendicularto the axis of the
craneboom.All detecbrs were tilted upward at a
30-degreeangle.
We conductedour study during nine nights
between 8 July and 1,1August 1996.Gencrally
two, but at least one station at each height was
operationaleach night. Simultaneoussampling
at eachheighl allowed us to acoountfor temporal
variabilityamongnights(Hayes1997).
We downloadedand digitized all calls using
thc ZCAIM Interlace irnd ANABAf 5 soliware
package(TitleyElcctronics. Ballina.NSW Aus-
tralia). Time and location were rccorded in llle
headerscreatedfbr eachcall. We examinedeach
passand noted the minimum and maximum fie-
quency and the duration of calls. Calls were
groupedinto three broad categoriesfor analysis:
I t c r l l s i d e n t i l ' i ead. h a \i n gc h a r a c l e r i \ r r c : : i r n i -
lar to most speciesof bats in the genusMl.rli.t
(typicallyhighfi'equency. shondurationcalls)were
placed in the Mlolls group (O'Fanell 1997), 2)
calls with characteristicssimilar to the big brown
bat.silver-hairedbat.hoarybat,andfringedmyotis
(all with typically low'frequency. long duration
calls) were placedinto the non-M).rtir group,and
3) fuagmentarycalls that did not yield sufficient
VerticalStratification
of Bat Activiry 103
infbrmationrcgardingfrequencyanddwationwerc
classifiedas undetermined. Furthertaxonomic
relinement was not possiblebecausea high per-
centageof calls were fragmentary.
We ex;imined diffelenccs in log-transformed
n u r n b eor l p a . : e sb 1 \ e f li c a l: r r a t u r n
u s i n ga t u o -
u ay ANOVAusing SAS PROC GLM. We gener
ated least squarcsestimatesof log-transfbrmed
meansfor numberofpassesin eachvertical stra-
tum andconductedmultiple comparisons fbr num-
berof passesamongstratausingFisher'sprotected
LSD. To assesstemporal pattemsof activity, we
partitioned the period betwcen sunsetand sun-
rise into 20 equallengthperiods(mean= 27.1
min.. raoge25.5to 29.6min.) to accountfor dif
ferenccsin length of nights during rlle study pe-
riod. We then dctermined the number of passes
occunin-tin each5 percentileperiod(Hayes1997).
Wc usedthe Kolmogorov-Smimov Two-Sample
Test (Sokal and Rohlf 1995) to examine differ-
ences1ntemporalpattemsof useby bats among
vertical stlata. P-r'aluesfbr the Kolmogorov
Smimov Test were takcn from tables provided
by Rohlf and Sokal( l98l).
Results
We recordcd2,30.1bat passcs.Of these,1,207
(52.,17r)wele classitiedasM_r'otls,154(.6.7a/t)
as
non-M\'.tlr,and 943 (40.97r)as undelcnnined.
Significantly diffcrent nurnbersof passesby
Mr'rtir were detectedat eachvertical stratum(Tablc
I ). The greatestnumbcr of passeswere recorded
in the lower canopy.tbllowed by the ground level
and uppercanopy,respectively.No calls recorded
above the caDopywere classitied as Mrotis. al-
though it is possiblethat somc 0f the calls rc-
corded abor,ethe canopv and classificdas unde
temined mrv havebeenproducedby Ml.rrri.rbats.
Thc patterusobservcdlbr undeterminedpasses
TABLE I NlcdiannuDrbefolpasscs rcro\s rllnighls. Confidenceintervals(in parentheses)r'erc calculatedfor log ransibmred
numbersrnd back tran\fornredti)r easeofjntcryreirtion. \,ledian !alues lbr number ofpasseshaving rhc samesupef-
scripl \Lithln l column arc nol significantl,vdifcrent (P > 0.05) from one another Total passesare the sLnnof Dasses
catcgoized as,U\..rir. non Mlo/rr. and undetemrined.
{)
L ppcr clnoP) 8 . 86 . 5 - 1 r . 8 f
2 1. l \ 1 9 . 1 - 3 7 . I l
Ground lc\el 15.0(108-10.7).
10.:1 Hayes and Gruver
andfor total passeswere similar to that for Ml'rliJ,
aswould be expectedgiven the predominanceof
use by M)?li.r bats in the study area.
The largestnumber of passesfbr rlon M);oti.l
batswererecordedin the uppercanopy,fbllowed
by the lower canopy,ground leveJ.and abovethc
canopy(Table1).Numberofpassesof non,Mt,olis
batsat the ground-level$'asnot signiticantly dif-
fercnt than numberof passesrecordedabovethe
canopy,andnumberofpasses recordedin thelower
canopy did not difter from that recordedin the
upper canopy.
Over all vertical stratacombined, activity of
My,otisexhibitedabimodaldistriburionwith peaks
duringthefifth andninetiethperccntileof thenight.
However,tcmporalpattem of activity for Mt,otl"t
at the groundlevel differcd signiflcantlyftom that
in the lower (P < 0.005)and upper(P < 0.001)
canopy.Thcre was a strong pelk in activity of
M].r/iri at the groundJevel in the first fifth per'
centile of the night with a smaller and less pro-
nouncedpeak during the last 15% of the night
(Figure 1): 49.77eof all Mlotl.r calls recordedat
the groundlcvel were recordedduring thc first
56loof the night. In contrast.the largestpcak in
activity in the lower canopyoccurreddufing the
last25% (roughly2-112hours)ofthe night:with
53.4Voof all Myotis passesrecordcdin the lower
canopyduringthis period(Figurel). Relarivcto
the pattem observed at ground-level. thc early
eveningpeak in activity in the lower canopy was
nuch smaller and was shified later by roughly
one-half hour (one 5 perccnt interval). The pat-
tem of activjty for Mtotis in the uppcr canopy
shou,eda slight bimodal tendency,with somc-
what more calls recordedduring the lafter por-
tion ofthe night thanin thc earlierpofiionofthe
night. Temporalpattemsof actiyity in the lower
and upper canopy were not significantly differ
CJIe!,,r]of tr\.c\
0 . 6( 0 . 2 - 1 . 2 1 0.9(1..1-1.6f
1 . 8( r . l - l . l r 2r.6 (16..r-t0.9)h
t . 1( 1 . 0 - 2 . 1 ) " ,r6.5(32.6-65.7).
0 . 5( 0 . 1 - 1 . 0 ) . 33.8(23.1-,18.1)h.
 Non-Myotis Myotis

Abovethe Canopy Abovethe Canopy

0.1
=
.9 0.0 0.0
=
(|, L25 2.O
.L
o 1.00
o
IA
UpperCanopy
tt
.q 0.75
4
o 0.50

o 0.25
tt
E
= 0.00
=
E 1.25
.E
o
= 1.00
LowerCanopy
0.75

0.50

0.25

0.00 0

4 m
't6
3 Ground Ground
12
2
8
1
a
0 0
100 0 1 0

Percentage
of night

Figure L Tempofal distribution of rctivity fin prs\es cl.rssified.rsMr.r/i.r of as non'rulrtii lbr each \erlic.rl slratum. Lelel ol
a c t i ! i r yi s s c r l c dd i l l c r c n l l ] l b r c a c hs E a l u ma n dt a x o n o i c g r o u p r o i l l u s t r a r e t e m p o r a l p a r t e r n s . P a s s e s a t 0 a n d 1 0 5 %
ofthe night representall tasses recordedpfior to sunsetand alief sunrise,re\pectiveiy.

VcrticalStratificalionof Bat Activitv 10-5

ent (P > 0.10). Diffefencesin pattemsof activity
amonglevelsretlecta shift in useof theareaover
the cou$e of the night. During the initial 5oloof
the night, over 90% of the Mxrtl.e passeswere
recordedat ground level (Figure2). Activity then
shifted to higher vertical stratawith most of the
activity occurringin thelowercanopyduringmost
of thc rcst of the night; the predominanceof ac-
tivity in the lo$'er canopy was most pronounced
duringthe last 2070of the night.
: {
9
Percentage of Night
Figure 2. Tenporal patrcm ofrclativc amounrof Mlolir ac-
tility recordedal ground ]e!el. &e lower canop,"".
and the upper canop). Passesat 0 and 1057. of the
l flor l^ .Jr.el
r ! h r r c p r ( ' c n rJ l l 0 d . . . r c ( ^ r J c c p
and alier sunrise,respectivel!.
Non-M}otir batsalso showeda strongpeakin
activityduringthe first 5% ofthe night.although
a secondpeak later iD the night was not evident.
The earlyeveningpeakwasmost pronounced at
thc ground-lcvcl(Figure1);only onepassofnon
Mt.rti.r bats was recorded at groundJevel alter
the first 570of the night. Less pronouncedpeaks
in non-M_rotisactivity were observedduring the
fir'st5Trof the night in the lo$er anduppercanepy.
and a low, but relatively consistent,amount of
activity was recordedthroughout the rest of the
night.Activity was consistentlylow throughout
the night above the canopy and no distinct mo-
dality was evident. although patternsmay have
beenobscuredby thc cxtrcnely small numberof
calls recordedat that level. The patternof actjv-
it) JI gr,\un(l-1.\u f ol r n o n - M r , , i i b
. ' a t su r . s i g
nificantly different than that observedat each of
the othcr thrcc lcvels (P <0.001);temporalpat
lemsof activityfor non Ml.rtiJ at thelo\\"ercanopy,
upper canopy, and above the canopy were not
signilicantlydifterent(P > 0.l0).
106 Hayesand Gruver
Discussion
Studiesthat monitorecholocation callsto assess
activity ofbats typically are conductedusing bat
detectorsplaced on or very near to the ground.
We found pattems of use differed substantially
amongvefiical strata,and that thesepattemsdif:
feredbetweenspeciesgroups.Thus.exclusiveuse
of ground-based equipmentmay rcsultin an in-
complete p i c t u r eo f t h er c t i r i t 1 o f h a t si n c o m -
plex foreststands.Resultsof echolocation-noni-
toring studiesconductedin tbrestedenvrronments
usingonly ground-based equipmentshouldthus
be interpretedwith recognitionof potentiallimi
tationsin inferencethat resultliom samplingonly
one verticitlstratum(Hayesin press).
The bimodal patternof activity obscrvcdat
ground level lor Mlotis bats is consistentwith
pattemsobservedin other studies(Erkert 1982.
Hayes1997.Kunz 1973,Maier 1992,Taylorand
O'Neill 1988,Thomas1988),althoughthe rela-
tive magnitudcof the peakobservedduring the
early pofiion of the night in this study exceeds
that typically observed.The bimodal pattern of
activity of bats in old gro$'th forests of the Pa
cific Nofthwesthasbeeninterpretedto resultliom
an initial pulseofactivity asbatsleavetheirroost
sites.tbllowed by decreased activity in the rniddle
ol the night as bats tbragc in other arcas.and a
secondpeakofactivity fesulting asbatsretun] to
their roostsitesduring early morning hours(Tho-
mas1988).Others(Hayes1997,Kunz 1974.Kunz
et al. 1975) have attributed the bimodal pattern
of activity ofbats to an initial period of foraging
and roosting alter energence from day roosts.
reduceductivityin the middleof the night when
bats are at night roosts,and a final bout of forag-
ing andcommutingbeforeretumingto day rcosts.
Kalcouniset al. (1999)suggested tbatvertical
shilis in activity may accountfor temporal pat
terns of activity observedat ground-level.We
obseNeda shift in activity from groundlevel to
higher vertical strata through the night (Figure
2), consistentwith this hypothesis.The mecha-
nismsresponsible for thcscvcfticalshiftsin ac-
tivity are not clear,but may be relatedto vertical
shiftsin abundance oravailabilityofinscctsthrough
rhenight.Shiftsin heightofacrivirydo nor,hou,-
ever accountfor all of the obscrvedbimodality
in activity, as a bimodal pattem is evident even
when all venical strataarecombined.Night-roost-
ing probablyaccountstbr someof this bimodality,
asnight-roosting comprisesa substantial
of the activity budgetof some species(Anthony
et al. 1981.Barclay 1982,Perlmeter1996)and
periodsof night-roostinggenerallyoverlap with
pcriodswhenrccordcdactivityis lowcst(Adam
andHayesin press,Kunz 1974,Perlmeterl996).
Movement from day-rooststo lbraging habitat
undoubtedly occursas well (Waldien1998).and
m l ) r c c o u n ti o r r p o r t i o no f t h ep r l t e mi n : o m e
habitats.
Mtolls and non-Mloti.r bats exhibited differ-
ent pattemsof use of vefiical strata.Differences
in useof vertical stratabetweenM}.oti.rand non-
M_}.oti.r
bats in our study may have resultedtiom
a numberoffactors. Differencesin useofhabitat
amongspeciesof batshave beenpostulatedto be
related,in part, to dil'ferencesin abilitics to cx-
ploit resourceswith more maneuverablespecies
concentratinguse in more clutteredhabitatsand
Iess maneuverablespeciespredominantlyusing
less cluttered environments (Aldridge and
Rautenbach1987,Brigham ct al. 1997,Crome
andRichards1988.KalcounisandBrigham 1995,
Kalcouniset al. 1999.McKenzieandRohe 19136,
andNorbergandRayner l987); this may account
tbr the diffbrent pattemsof use we observed1br
MloliJ andnon M).rlir specles.
The complex vertical structureof old-growth
forestsinthe PacificNofihwest(FrunldinandSpies
l 9 9 l r p r , ' r i d e .r h eo p p r , n u n i t l)u r p a n i l i , ' n i n !
and exploitation of diverse niches by aerial in-
sectivores.such as bats.Other taxa, including
epiphytes(McCune et al. 1997)and birds (Shaw
portion andFlickin press),differin presence.abundance,
oruse ofhabitat at differentheightsin old-growth
coniferous forests.Forestswith simple vertical
structure su. c ha s) o u n gc o n i l e p
r l a n t ai o
t n . .m a 1
notprovide a diversityoffomging nichcsforbats;
this may, in part. accountfor differencesrn activ-
ity of bats in older and youngerforests(Thomas
1988,Humeset al. 1999).We wercunableto test
this hypothesisbecauseour work was limited to
a single site. and we could not comparepatterns
of activity among forests diffcring in structural
complexity.Comparativcstudiesto evaluatethis
hypothesis throughexamination of differences in
activity ofbats at different heightsamongforests
in the PacificNorthwestdiffering in verticalcom-
plexity wouldbe valuable.
Acknowledgements
We thank David Shaw. Buzz Baker, and Mark
Creighton of the Wind River Canopy Cranc for
their logisticaland field suppon,enablingus to
deploy and maintain bat detectorsin and above
the canopy.Mike Adam and Dave Larson pro
vided field assistance. ManuelaHuso and Shay
Howlin provided statistical advice. This paper
benetittedfrom commentsby Tim Reynolds.David
Waldien, and an anonymousrevicwer on earlier
drafts.Fundingfor this researchwasprovided b"v
the Wind River Canopy CraneResearchFacility
and the Coastal Oregon Productivity Enhancc-
ment (COPE) Program.College of Forestry.Or-
egon StateUniversity.

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