Residence and territorial characteristics of Libellulidae species in a neotropical assemblage (Anisoptera)

2008·2.2 MB·English

by  Resende, D C

Odonatologica37(3):213-220 September1,2008
Residence andterritorial characteristics of
Libellulidaespecies ina neotropicalassemblage
(Anisoptera)
D.C. Resende¹*and P.De Marco+Jr².
1 Laboratorio deEcologiaeSolos,CursodeCiencias Biologicas,
Centro Universitario do LestedeMinas Gerais,R. BarbaraHeliodora,725 Bom Retiro,
BR-35160-215,Ipatinga,MG, Brazil
[email protected]
2Laboratorio deEcologiaTeorica eSintese,DepartamentodeBiologiaGeral,
Universidade Federal deGoias,BR-74001-970,Goiania,Goias,Brazil
[email protected]
ReceivedFebruary25, 2007/Revised andAcceptedDecember30,2007
Duringterritorial behaviour,aggressive attacksamongheterospecificodon. 33
arecommonand maycauseaseparationofniches, based onthepreferredsites for
territorial defence. Here,territorial behaviourand thecharacteristics ofterritories in
<JdofErythrodiplaxmedia,Micrathyriacatenataand M.hesperisaredescribed and
theirterritorial fidelity, capturingandmarkingof<3 3arediscussed. Inallspp. stud-
ied,there wasacleardistinctionamongthe microhabitatsdefendedasterritories. In
both Micrathyriaspp., 3 3 seemtodefendterritories with defined resources.InE.
media,thedefended resourcesarelessevident.Its3 3arehighlyaggressiveandshow
highterritorial fidelitybut,apparently,theylosethe territoryiftheystay awayfrom
waterforatleast oneday.
INTRODUCTION
During territorialdefence, aggressive attacks amongheterospecific odonate
malesare common(HASSAN, 1978; MAY, 1980)andcompetition and aggres-
sive interactionscouldcausesomespecies tochange theirhabitatselection(MAY,
1980;RUSSELL etal., 1998). Theseaggressive interactions may cause asepa-
rationofniches amongspecies. Thusthe co-occurring species, Ischnuraelegans
(VanderL.),Coenagrion pulchellum (VanderL.), Enallagma cyathigerum(Charp.)
andErythromma najas(Hans.), forexample, haveenough differentiationintheir
oviposition sites(JOHANNSSON, 1978). Sympetrumflaveolum (L.)malesuse
*Correspondingauthor
214 D.C. Resende& P.DeMarcoJr
lowerperches than thoseof S.sanguineum (Mull.) butbothspecies perch lower
whentheirneighbour is amaleoftheotherspecies (REHFELDT & HADRYS,
1988).
Differences in territorial qualitybetween odonate species are littleevident.
In somespecies, malesdefendedterritories selected by the femalebasedon the
quantityofoviposition resourceavailable(TSUBAK1&ONO, 1986;ALCOCK,
1987a; ALCOCK, 1987b; KOENIG, 1991; MEEK & HERMAN, 1991;BAT-
TIN, 1993),whereasinotherspecies, territorial defenceseemstobemorerelated
to thechanceofmales detecting femalesarrivingatthepond(ALCOCK, 1987a,
1987b; VAN BUSKIRK, 1986; BATTIN, 1993; DE MARCO& RESENDE,
2004)andinthis case,thesizeoftheterritory candeterminereproductivesuccess
of somespecies, sinceit increases thechanceofmalevisualizing females(VAN
BUSKIRK, 1986).Theperches orsubstrates forthemaleperching inaterritory
canalsobe diverse(PARR, 1983;OSBORN & SAMWAYS, 1996),andchange
mainly as afunctionoftheiravailability.
The objective ofthisstudy was to determinethe criteria forselection of ter-
ritoriesby males of Erythrodiplax mediaBorror, Micrathyria catenata Calvert,
and M. hesperis Ris, by observing theirterritorialbehaviour, territorialfidelity,
andthe characteristicsoftherespective territories.
METHODS
STUDY AREA - The work wascarried out duringFebruary-April in the Centrode Estudos
deFlorestasNaturaisof Universidade Federal deViposa,MinasGerais(20°45’S,42°5TW).There-
gionalclimate isCwa (Kdppenclassification),moderatelyhumid sub-tropicalwith ahidric deficit
between May andSeptember(GOLFARI, 1975).Theaverageannualrainfall is1450mm,relative air
humidityca80%and thetemperature varyingfromamaximum of26,1°Ctoaminimum of 14,0°C
(VALVERDE, 1958).
Duetoalocal dam,Nympheacf.amplaandEleocharis areveryabundantinthecoastalregion.At
the shoreline therearesticksand grassesthat areused asperchesbysomedragonflyspecies.
BEHAVIOUR ANDTERRITORIALCHARACTERISTICS - Duringthe studyperiod,all
activities relativetoterritorial defence,matingritual and ovipositionwererecorded. The emphasis
wasondescriptionsofthebehaviour,courtshipritualbefore mating,maledefencebehaviourduring
ovipositionand ontheovipositionsites.
Tocharacterize theterritories defended by E. media, M. catenata and M. hesperis, wewalked
aroundtheshoreline ofthepondandrecorded,foreach individual,thetypeand heightofitsperch,
thedistancefromtheperchtothe shoreline,thevegetationsurroundingtheperchand thedistance
betweenperches oftheconspecificterritorialmales. Todetermineinterspecificdifferencesintheter-
ritorial characteristics,weperformedanalyses ofChi-squareforcategorical variables and analyses
ofvariance (ParametricANOVAand Kruskal Wallis)forquantitativevariables.
TERRITORIALFIDELITY— Forthesakeoftheenquireintotheuseoftheterritoriesbymales
ofthethreespecies, 12adjacentsectionsoffivemeterslengthweremarked in thelittoralzoneofthe
pond.Maleswerecapturedand markedonthewingswith non-toxicpaint.Thedateofcaptureand
thesectionin whichthemalewasencounteredwererecorded. Duringthesubsequentdays,between
10:00and 14:00.thesegmentswerechecked, allmarkedindividuals andanynewonesmarked.
Intheanalysesofmaleterritorial fidelity,maleswith atleast fourrecaptures wereused forE.media
Territorial characteristicsin neotropicalLibellulidae 215
andthosewith threerecapturesforMcatenataandM.hesperis.Thisdifferencein criteriawasdueto
thelower frequencyofrecapturedindividuals ofthe lattertwospecies.Aperiodoffiveconsecutive
dayswasused todetermine whethertheindividuals absentfromthedam foratleast onedaycould
losetheirdefendedterritories. Itwasassumedthatamale hadchangeditsterritorywhen itappeared
in adifferent section,notcontiguoustotheoriginally defended segment. Therelationshipbetween
territorial changeand theabsenceofmaleswastestedbyaChi2analysis.
RESULTS
BEHAVIOUR AND TERRITORIALCHARACTERISTICS
Erythrodiplax mediaseemstobemoreaggressive thantheMicrathyria species.
Thefemaleswere abundantatthepond andcouples incopula wereoftenseen,
butnocourtshipritualwaseverobserved.Aftercopulation, thefemalesremained
forsomesecondsperched onbranchesnearthemaleterritory,generallyguarded
bythemale.Finally, they oviposited neartheshorelineofthepond, inlocations
withsubmerged vegetation. Oviposition was notrestricted totherespective male
territories, although themalesguarded the femalesduringtheentireoviposition
period, attacking othermales thatapproached.
Micrathyria catenatais not avery aggressive species, though thelargest ofthe
threespecies studied. Maleswereobserved, butnofemalesandmatingswereever
seen.
M. hesperis is asmall and likewise notvery aggressive species. A single mat-
ing was observed andthere was neitheracourtship ritualnor apause between
mating andoviposition. Thefemaleoviposited inthemale’sterritoryanditwas
guarded by themaleflying above.However,weobservedanotherfemaleovipos-
itingunguarded, perched onaleaf of Nymphea andsubmerging herabdomen.
Thetwo Micrathyria species perched morefrequently onaquatic macrophytes
(73.6%;n= 53), whencompared withE. mediamales, whichperched onbranch-
es,grassesorevenonthesoil (78.9%; n= 19;Chi2 = 15.76;df = 1;p<0.001; Fig.
1) inadditionto using macrophytes. Comparing only Micrathyria species, M.
catenataperched mainly on Eleocharis, whereas M. hesperisperched morecom-
monlyon thebud oronthe leafof Nymphea (74% and48%,respectively; Chi2
= 15.34;df= 1; p<0,01; Figs lb, 1c).
Theperch’s height was also differentamongthe threespecies (ANOVA; F =
12,85;df=2;p<0,01). M.catenata perched higherthanM.hesperis(Fig. 2; Tukey
test; p<0,01). Theperch’s height forE.mediamaleswas intermediate, not differ-
ing significantly fromthatineitherofthetwo Micrathyriaspecies(Fig.2; Tukey
test;p= 0,13).
Furthermore,thevegetationaroundtheterritoriesofthethreespecieswasdifferent
(Chi2=50,90;df=8; p<0,01). E. mediadefendedterritorieswithEleocharisand
grasses(37%and42%,respectively; Chi2= 19,89;df=4;p<0,01),whereasM. cat-
enatadefendedmostlythosewithEleocharis(76%;Chi2=28,09;df= 1;p<0,01),
216 D.C. Resende&R De MarcoJr
and M. hesperis thosewith Nymphea (94%;Chi: = 18,1;df = 1;p<0,01).
E.mediamalesdefendedterritoriesalongtheshorelineandperched onbranch-
es, grasses, directly onthe soil or on Eleocharisat anaverage height of 14cm.
Thetwo Micrathyria species perched furtherover thepond thanE. media,par-
ticularlyinthecaseofM. hes-
peris (Chi2 = 18,83; df = 1;
p<0,01)forcomparison with
M. catenata (Kruskal-Wal-
lis ANOVA;Chi2=21,62;df
= 1; p<0,01). In M. catena-
ta, thecentre of the territo-
ries averaged onemeterfrom
the shoreline;males perched
higherthantheotherspecies,
primarily on Eleocharis.The
territoriesof M. hesperis ex-
tendedeven furtherout over
the water than those of M.
catenata (Kruskal-Wallis
ANOVA; Chi2 = 5,37; df =
1;p=0,021;Fig. 3)andthey
mainly perched ontheleaves
orflowersofNymphea.How-
ever, thedistance of M. hes-
peristerritoryfromtheshore-
line variedand there was no
homogeneity of variance
in the data(Fig. 3; Bartlett
Test; Chi2 = 27,128;df = 2;
p<0,001).
There was some differ-
encein thedistancebetween
perches usedby malesofthe
three species (F = 11,06; df
=2; p<0,01; Fig. 4). Thedis-
tance between the perches
used by E. media males(x
=229 cm) was similarto the
distancebetweenthesein M.
Fig. I.Relative frequencyofthe typeofperches inViqosa, hesperis (.v = 338cm; Tukey
MG.Brazil, usedby malesof:(a)E.media, —(b)M.catena- test; p=0,87). However, M.
ta, - and(c) M.hesperis. catenatamalesperched near-
Territorialcharacteristics inneotropicalLibellulidae 217
er theirneighbours (x = 103
cm) thanthe other two spe-
cies(Tukey test; p<0,001).
TERRITORIAL FIDELITY
Thedisplacementof males
of the three species between
sectionsofthepondwassim-
ilar (F = 2,24; df = 2; p =
0,13; Fig. 5).However, there-
capturerate wasverylow, es- Fig. 2.Mean heightofperchesused formales ofE.media,
pecially, for the Micrathyria M. catenataand M.hesperisinViijosa, MG,Brazil. Barsin-
species. Ofthe 151markedE. dicate thestandard deviation.
mediamalesonly35wererecaptured atleastfourtimes,whereasof the114marked
M. catenataand24M. hesperis males,onlysevenandfour,respectively, werere-
captured atleast threetimes. Thusonly the E. mediadatawere consideredfur-
ther.Thefrequency distributionofdisplacement inthisspecies showedthatmost
malesdefendedterritoriesinthesameplace,moving about 10to20meters,with
onlyafewindividualsmoving morethan35meters(Fig. 6).Furthermore, in77%
oftheoccasionswhenE. me-
dia males stayed away from
thepond foratleastone day
they losttheirsectionofter-
ritorialdefence, whereasthis
occurred inonly 34%ofthe
cases when males remained
atthepond(Chi2=9,19; n=
38; df = 1;p<0,05).
DISCUSSION
Fig.3.Distance ofperchestothepondmargininmales ofE. Thepartition ofmicrohab-
media.M. catenataand M. hesperis, in Vi?osa,MG,Brazil. itatamong the species of a
pPoonsidti.veBavrasluinesd,icinattehethpeonsdta;n—danrdegdaetviviaetvioanlu.es,awayfromthe communityisacommonphe-
nomenonandcan occur due
to interspecific competition. Females of many Micrathyria species oviposit on
aquaticplants,suchasNymphea (PAULSON, 1969),henceitishardtosaywhether
themicrohabitatsselected by Micrathyria speciesas describedinthisstudy are a
consequenceofcompetitionwithE.mediaora characteristicofthegenus.Libel-
lulidaemostly oviposit directlyonthe surface ofthewater oronthe surfaceof
wetfloating objects, including plant material— exophytic oviposition — (COR-
218 D.C. Resende& P. DeMarcoJr
BET, 1999). Theoviposition
on plant materialcan possi-
bly result in (1) an increase
in the parental care survival
as theeggs wouldbeless ex-
posed to predators such as
fish andotheraquaticorgan-
isms, or(2) adecreaseinhar-
assmentoftheovipositing fe-
maleby malestryingtocopu-
late.
Fig.4. Distance between perchesused bymales ofE.media, Males with a high degree
M. catenataand M.hesperisin Viposa,MG,Brazil. Bars in- of residence should be dis-
dicate thestandard deviation. placed lessoften thanmales
that continually lose territo-
rialdisputes(PARR, 1983;HARVEY&CORBET, 1985;CONVEY, 1989).The
timeofabsenceofa territorial malefromwater canberelatedto theenergyres-
torationortheperiod offoraging(FRIED& MAY,1983;ANHOLTetal., 1991;
MARDEN &ROLLINS, 1994;DE MARCO, 1998). Thiscan cause confusion
inthe residenceofa territory(GRIBBIN & THOMPSON, 1991), resulting ina
greater displacement of residentmales.DUNHAM (1994) hasalready empha-
sizedthatthe departure forforaging can makeitdifficultto maintaingood ter-
ritoriesand,consequently, malescan lose matingopportunities. Nevertheless, if
territorialdefencedemandsahighenergeticcost(FRIED & MAY, 1983),larger
males, with larger energetic reserves, are supposed to show a higherreproduc-
tivesuccess, sincethey stay fora longer timein theterritories withoutforaging
(HARVEY &CORBET, 1985).
The distance between in-
dividualsof aspecies canbe
directly relatedto the degree
of aggressiveness of territo-
rialmales(VAN BUSKIRK,
1986). Differences in the re-
source abundancecan also
affect male distribution
(MEEK & HERMAN,
1991). At our locality Nym-
phea covers a large area of
the dam, therefore M. hes-
perismalesare intheposition
Fig. 5.MeanmobilityofmalesofE.media, M. catenataand
toestablishterritoriesrather M. hesperisin Viyosa, MG, Brazil. Bars indicate the stand-
distantfromeachother,per- ard deviation.
Territorialcharacteristics inneotropicalLibellulidae 219
haps minimising therewith
the intraspecific competi-
tion. Nevertheless, nodiffer-
enceintheresourceavailabil-
ity was evident in E. media
and M.catenata atthepond,
the greaterdistancebetween
perches in E. mediaterrito-
rial males was probably due
to the fact that this species
performs moreaggressive in-
teractions thandoes M. cat-
enata. Fig. 6.Frequencydistribution ofmobilityofmarkedE.me-
Thespatialarrangementof diamales inVigosa-MG,Brazil.
species intohabitatunitsisacomplex resultofbionomiccharacteristics, includ-
ing oviposition resources and perch choice, andinteractionswith otherspecies
inthecommunity. Thebionomicfeaturesare probably themostimportant force
driving thechoiceofhabitatusedby aspecies atalocality, despite thefactthat
the co-occurring species can affect, atleast withincertain limits, the choiceof
microhabitat.
ACKNOWLEDGEMENTS
We thankANDERSON LATINIforfieldassistance andTANIASANTOS, MARILIAGAIA
andMARIANA MELOforvaluable revisions andcomments.Thisstudyreceived financial support
fromtheBrazilian Conselho Nacional dePesquisas(CNPq).
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