Marie Selby Botanical Gardens, Inc

CANOPY STRUCTURE AND INSECT COMMUNITY DISTRIBUTION IN A TROPICAL RAIN FOREST

OF WEST KALIMANTAN
Author(s): Fumito Koike, S. Riswan, T. Partomihardjo, E. Suzuki and M. Hotta
Source: Selbyana, Vol. 19, No. 2, CANOPY PROCEEDINGS (1998), pp. 147-154
Published by: Marie Selby Botanical Gardens, Inc
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 19(2):147-154
Selbyana

CANOPY STRUCTURE AND INSECT COMMUNITY

DISTRIBUTION IN A TROPICAL RAIN FOREST OF
WEST KALIMANTAN

Fumito Koike1
of BiologicalScience,Facultyof Life and Environmental
Department Science,Shimane
Matsue690, Japan
University,
S. Riswan, T. Partomihardjo
CenterforBiology,LIPI, Bogor,Indonesia
Researchand Development

E. Suzuki and M. Hotta

Science,Facultyof Science,KagoshimaUniversity,
of EarthEnvironmental
Department
Kagoshima890, Japan

Abstract. Thecanopy structure

andspatialdistribution
offlying insects
ina riverside
forest
canopy
werestudied.Basedoninsect thecanopy
composition, wasdividedintofourareas:upperopenspaceofa
canopygap,insidethecanopy, forest andriversurface.
floor, Thysanoptera wereabundant intheupper
openspace,whileColeoptera wereabundant neartheforest
floor.
Insectcompositionwasintermediate
insidethecanopy.Thehorizontal
canopystructure
affected
insectdistribution
as didthevertical
structure.
Thecommunity attheriver wasrichinaquatic
surface insects.
However, thedistribution
ofaquaticinsects
waslimitedonlytotheriversurface.

Introduction ture(Koike & Syahbuddin1993,Koike 1994),

andthehorizontal distribution oftheinsectcom-
Many ecologicalactivitiesincludingflower- munityin thefoliage-canopy has notyetbeen
ing,pollination and herbivory
occurin thecan- studied.A betweenthespa-
quantitative analysis
opy of a forest.These activitiesare affected by tialdistribution offoliagedensity andinsectdis-
environmental factorsdeterminedbythecanopy tribution has not yetbeen made. This maybe
structure (Koike et al. 1990). For some insect caused by the difficulties in measuring foliage
taxa, the foliageof the canopysuppliesfood, densitydistribution in a tallcanopy.
andthoughdensefoliagemaybe an obstaclefor
Manytrapmethodshave been used to study
it also may be a shelterfrompredators insect
flight, distribution in a forestcanopy.Although
and strongwinds.Thus thefoliagedistributionultra-violet are oftenused to assess
shouldbe consideredas a kindof map of the insect fauna in light traps
canopies (Sutton& Hudson
canopy,whendescribing thespatialdistribution
1980,Suttonet al. 1983,Casson & Hodkinson
of insectactivities. 1991, Kato et al. 1995), lightspreadsover a
Manystudieshave been conductedon insect wideareaifthereis openspacearoundthetrap,
distribution in a forest
canopy(Sutton& Hudson and a wide rangeof insectswill be trapped.
1980,Suttonetal. 1983,Rubik1993,Fukuyama Since lightmay disturbthe spatialdistribution
etal. 1994,Katoetal 1995).Toda (1977, 1987) of
studiedthe Drosophilidaecommunity insects,it is difficultto obtaina highspatial
in forest resolutionof insectdistribution by thismethod.
canopiesincool temperate forests
withreference Insecticide
fogging is also used to assess insect
to thecanopystructure. He foundtwocommu- fauna
1973, Casson & Hodkinson
nitytypes:an uppercanopycommunity of sap 1991, (Roberts et al. 1993),but spatialresolu-
feedersand a forestfloorcommunity Kitching
of fungus tionis also low. We therefore used a two-di-
feeders.Most earlierstudies,however,used mensional of stickytrapsto mea-
arrangement
heightfromthegroundto indicatethetrappo- surethe spatialdistribution of flyinginsectsin
sitionin thecanopy.Tropicalrainforests usually a tropicalrainforestcanopy.
havea horizontally heterogeneouscanopystruc-
1Currrent
address:
DepartmentofVegetation
Sci- Methods
ence,Institute
of Environmental
ScienceandTech-
nology,Yokohama NationalUniversity,
79-7Toki- A naturaltropicalrainforestalongtheDaid
Yokohama
wadai,Hodogaya-ku, 240-8501,Japan. River in West Kalimantan(0°43'N, 110°12'E,
(koike@kan.ynu.ac.jp). alt. 60 m) was studiedin August1991 (Suzuki
147

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148 SELBYANA [Volume19(2)

Figure2. Canopy diagramofthestudiedsection.

Smalltreesandshrubswereneglected.
Trapswerear-
rangedatthepositions
ofopencircles.
Hemispherical
photographsweretaken
attheplusmarks todetermine
canopystructure
andlightconditions.

Figure1. Topography ofthestudysite(Suzukiet

areat 1 mintervals.
al. 1997).Contours DaidRiveris
60 m abovesea level.P1-P7arehorizontal
positions
oftraps.

et al. 1997). The meanannualprecipitation at

Sangau,nearthe studysite,is 3,111 mm,and
themeantemperature is 26.4°C.The studywas
conductedat the end of the dry season. No
prominent flowering was foundduringthispe-
riod.The studysitewas a fragmented primary
forestsurrounded by a partiallylogged area.
Shorea stenoptera(tengkawangtungkul)and
Tristaniaobovatawereabundant alongtheriver.
Slopes behindthisforestwerecoveredby low-
landdipterocarp forest.
A 40 m linewas set acrosstheriver(Figure
1), and thecanopyabove thisline was studied
(Figure2). A ropesystem toholdtrapsandtake
photographs to determinefoliage-canopy struc- Figure3. Ropesystem toadjusttrapandcamera
turewas established (Figure 3). At first
a ring Horizontal
positions. positionis adjusted bythering
rope A, vertical
positionby B-E. F is a camera with
rope,A, was set.Thisropecouldbe maneuvered gyro H a tapemeasure
horizon, andG anelectric cable
to determine thehorizontalposition ofotherver- toreleasecamera shutter.
Sevenvertical ropes(i.e.,fi-
ticalropes,B-E. Verticallines wereplaced at fi)at 5 m intervals wereusedinthestudy. Alltraps
approximately 5 m intervalsin a horizontaldi- weresetsimultaneouslywiththesesevenropes. A sim-
rection. ilarropesystem foruseinSarawak,
is available Lam-
The foliagewas measuredby canopytomog- birHillsNational Park(Koike1994).

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1998] KOIKE ET AL.: CANOPY STRUCTURE AND INSECT DISTRIBUTION 149

raphy(Koike 1985). A camerawitha hemi- The firsttwo 'pseudo-species'are commonbe-

sphericallens on thegyrohorizonwas maneu- tweenbothtraps,butthe'pseudo-species' of4-
veredwiththeropesystem,whichwas used to 6 individuals levelis different.
Thusit is possi-
collect insects.Upward hemispherical photo- ble to considerequalityof taxonomiecomposi-
graphswere takenat 2.5 m verticalintervals. tionand differences in quantity.
Gap fractions on thephotographs weremeasured
and totalfoliageamountsin variousdirections Results and Discussion
werecalculatedfromthedata.The two-dimen-
sionaldistribution of foliagedensitywas deter- A canopygap occurredat the centerof the
minedusingtheleast squaresmethod.Relative sectionabove theriver(Figure4a). Dense fo-
lightintensity was also determined fromthe liage of thelowerlayerbelow 15 m was found
samephotographs (Anderson1964). in thestudiedsection.Lowerlayerfoliagewas
Stickytraps(tradename HaitoriRibbon)of absentjust above theriver.Lightintensity gen-
size 3.8 cm by 70.0 cm weremade of double- erallydecreasedtowardstheforestfloor(Figure
sidedadhesivepaper.Theywereseton thever- 4b), butlightstreams downintothecanopygap.
tical rope at 5 m intervals(Figure 1) for24 Such a canopystructure withdensefoliagebe-
hours.No rainfellduringtrapping. The sticky low 15 m and sparsedistribution of theupper
trapswerethenwashedwithkerosene,and the canopycrownis commonin tropicalrainforests
trapped insectswerekeptin ethanol.Sincecom- in SoutheastAsia (Koike & Syahbuddin1993,
pleteidentification of insectspecies was diffi- Koike 1994).
cult,identification was made at theorderlevel. Spatialdistribution
of thenumberof trapped
Body size was also measuredforclassification.individualsin the studiedsectionis shownin
Bodysize class was classifiedin logarithmic in- Figure5. ManyColeopteraandDipteraindivid-
tervals,0-1 mm,1-2 mm,2-4 mm,4-8 mm, uals occupiedthelowerlayer.Hymenoptera and
and over 8 mm.Insectsof different body size Thysanoptera wereabundantin theupperlayer
classes were treatedas different taxa in com- andin thegap. Trapsampleswereusuallycom-
munity classificationanalysis. posed of diversespecies.The exceptionwas a
The two-way indicator species analysis species of Chrysomelidae(Coleoptera), of
(TWINSPAN,Hill 1979) was used to classify which 84 individualswere collectedsimulta-
insectcommunities. This procedure aimsto au- neouslyat theriversurfacetrap,probablydue
tomatetheplantsociologicalmethodthatclas- to migration flight.
sifiesplantcommunities (Mueller-Dombois & Correlation
coefficients were calculatedbe-
Ellenberg1974).In thiscomputer program, traps tweenenvironmental factorsand theindividual
are ordinated by reciprocalaveraging.Distinc- numbersof majortaxa (Table 1). Since thein-
tivetaxa,whichappearon onlyone side of the dividualnumbers usedin theanalysiswereclas-
firstordination axis, are chosen.The taxa dis- sifiedbyorderlevelidentification, theindividual
tributing throughout theordination axis are ne- numberof thedominantspeciesin each order
glected,andonlythedistinctive taxaareusedin shouldcontribute mostto theanalysis.Thysan-
the subsequent ordination.This ordination opteraandHymenoptera wereabundant in areas
makesit easyto separatetrapsintotwogroups. withhighlightintensity and in open space in
TWINSPAN can essentiallyuse only 'exis- theuppercanopy.Dipteraand Coleopterawere
tenceand absence'data of taxonomiecomposi- abundantin the lower layer.Foliage showed
tion. However,to considerthe differences in negativecorrelation
in Thysanoptera, Hymenop-
quantity andto avoidtheeffect ofaccidentalap- tera and Diptera.Open space withoutfoliage
pearancesof taxon,the analysis oftenused maybe a corridor fortheseflying insects;how-
'pseudo-species.' The sametaxonwithdifferentever,thenegativecorrelation was notsignificant
individualsnumbersare countedas differentforHemipteraand Coleoptera.These taxa usu-
'pseudo-species'.We setthefivelevelsof 'pseu- allyhavehardforewings and maynotflyas of-
do-species'in thisanalysis:1 individuallevel, tenas previously mentioned taxa.The masscol-
2-3 individuals level,4-6 individualslevel,7- lectionof a Chrysomelidae speciesat theriver
15 individualslevel,andover16 individuals lev- surfacesuggeststhatopenspace abovetheriver
el. Forexample,in a giventrap,if6 individuals mayfacilitate migrationflightevenforterrestrial
of 2-4 mm body size class Coleopterawere insects.
caught,this trap had four 'pseudo-species'. The insectcommunities in thestudiedsection
Theseare 1 individual level,2-3 individualslev- were dividedinto fourtypesby TWINSPAN.
el, and4-6 individuals level.In anothertrap,if The TypeA community was in theupperopen
2 individuals of thesame size class Coleoptera area, especiallyin the canopygap (Figure 6)
werecaught,thistraphad two 'pseudo-species' where Thysanopteraand Hymenoptera were
of 1 individuallevel and 2-3 individualslevel. abundant(Table 2). Type C was near forest

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150 SELBYANA [Volume19(2)

Figure4. Foliagedensity
(FD,a) andrelative (RLI,b) ofthestudied
lightintensity canopysection.
Back-
groundcontour
linesinb indicate
foliage distribution.
density is theexpected
Foliagedensity numberofleaves
bya 1 m line.Thespatial
penetrated resolution
offoliage was5 m by5 m inthehorizontal
distribution and
TheRLI obtained
vertical. byhemispherical
photographsrepresents ofthecanopy
openness abovetheposition.

floorwhereColeopterawereabundant. TypeB choptera,weredistinctive.Aquaticinsectswere

was in themidcanopybetweenforestfloorand
uppercanopysurface.Insectcomposition was
betweentheopen space and forest
intermediate
floor.Type D was only at the riversurface,
whereaquaticinsects,Ephemeroptera and Tri-
not distributedfarfromtherivereitherin the
horizontalor verticaldirections.
Althoughopen space above 40 m was not
studiedin thisresearch,the open space in the
uppercanopygap, wheresampleswere taken,

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1998] KOIKE ET AL.: CANOPY STRUCTURE AND INSECT DISTRIBUTION 151

of individual
Figure5. Spatialdistribution numbers contour
pertrap.Background linesindicate
foliage
distribution
density (Figure4a).

Table 1. Correlation
coefficients
betweenenvironmental
factors
andpertrapindividual
number ofmajor
taxa.
Thetotalindividual
number ineachtaxonwasusedinthisanalysis. variesinsomeorders
Foliagedensity
ofmagnitude,
butlogarithmscouldnotbe usedbecausefoliage
densityhaszerovalueatsomesites.The
transformation
xmwasusedtoconsider theeffect
oflowdensity foliage.
Relative
light
Taxon Height intensity Foliage
density
Thysanoptera 0.4706* 0.5881* -0.4011*
Hymenoptera 0.4535* 0.5183* -0.2864**
Hemiptera 0.1687 0.1441 -0.0421
Diptera -0.3238** -0.0948 -0.2387f
Coleoptera -0.4126* -0.2463t -0.1573
* P < 0.01,**P < 0.05,fP< 0.1.

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152 SELBYANA [Volume19(2)

Figure6. Two-dimensional ofinsect

distribution community A toD, inthestudied
types, canopysection.
Community weredetermined
types byTWINSPAN levelidentification
usingorder ofinsect
taxaandbodysize
contour
class.Background linesindicate
foliage (Figure4a).
density

extendedto the space above the canopy.The taxon had a maximumabundancewithinthe

representative taxaof TypeA community, Thy-
sanopteraand Hymenoptera, showed positive
correlation to heightand lightcondition.
These
resultssuggestthattheremaybe a TypeA com-
munity in thespace over40 m.
Fromstudiesby Suttonand Hudson(1980),
Suttonetal. (1983) andKato etal. (1995), gen-
eral insectdistributionpatterns determinedby
lighttrapsin tropicalrainforestsmayshowthat
individual numbers arelargeattheuppercanopy
surface, andthata smallpeakoccursjustaround
theforestfloorin sometaxa(Kato et al. 1995).
The resultsof thisstudywere similar,and no
canopyat intermediate height(Týpe C, Table
2). Some taxa showeddifferences amongsites
of individualnum-
in the verticaldistributions
bers.Coleoptera, forexample,wereabundant in
theuppercanopyin manysites(Sutton& Hud-
son 1980,Suttonet al. 1983,Kato et al. 1995);
buttheywereabundantalso at theforestfloor
at thissite and in two forestsin Panamaand
Brunei(Suttonet al. 1983). As shownin this
study,small scale canopystructure greatlyaf-
fectsinsectdistribution.Since quantitativedata
ofthefoliagewas notavailableinpreviousstud-
ies, comparison of theresultsis difficult.

individual
Table2. Average weredivided
pertrap.Insectcommunities
number A toD, with
intofourtypes,
TWINSPAN (Hill1979).
TypeA TypeB Type C Type D
Upper
openspace Insidecanopy Forestfloor Riversurface
Taxon (11sites) (32sites) (5 sites) (1 site)
Thysanoptera 5.5 1.0 1.2 0.0
Hymenoptera 17.6 8.8 5.8 0.0
Hemiptera 6.2 4.4 6.0 2.0
Lepidoptera 0.9 0.3 0.8 0.0
Diptera 29.7 20.0 37.2 154.0
Collembola 0.0 0.2 0.4 0.0
Coleoptera 8.7 8.5 22.4 98.0
Ephemeroptera 0.0 0.1 0.0 30.0
Trichoptera 0.0 0.0 0.0 5.0

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1998] KOIKE ET AL.: CANOPY STRUCTURE AND INSECT DISTRIBUTION 153

Mass collectionof migrating insectssome- terainsectcommunities oftropical rainforest in

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