Behavioural ecology of Erythemis plebeja (Burmeister) at a small pond in southeastern Brazil (Anisoptera: Libellulidae)

8 Pages·2002·1.7 MB·English

by  De Marco, P

Odonatologica31(3):305-312 September1, 2002
BehaviouralecologyofErythemis plebeja
(Burmeister) at asmallpondinsoutheastern
Brazil (Anisoptera: Libellulidae)
P. DeMarco+Jr.¹,A.O.Latin²iandP.H.E.Ribeiro³
1Laboratorio deEcologiaQuantitativa,DepartamentodeBiologiaGeral,Universidade Federal de
Vigosa,BR-36571-000Vigosa,MG,Brazil;e-mail: [email protected]
2Graduate programinEcology,ConservationandWildlifeManagement,FederalUniversityofMinas
Gerais,BeloHorizonte,MG,Brazil;e-mail: [email protected]
3GraduatePrograminEntomology,FederalUniversityofVigosa,Vigosa,MG,Brazil
ReceivedMay15,2001/Revised andAcceptedOctober20,2001
Ananalysis ofthetime-budgetandadescriptionofreproductivebehaviouratasmall
pondin Vigosa, Minas Gerais,Brazil arepresented. The observations supportthe
classification ofthis sp.asapercher.Italso conformstothepredictionsofotherstudies
thatlargeperchersareusuallymoreaggressive towardconspeciftcs.E.plebejaSSwere
usuallyobserved simulatingoviposition in the areasthey previously defended,and
evolutionaryaspectsofthisbehaviour arediscussed.
INTRODUCTION
Theevolutionary ecologyofaspecies isoftenrelatedtotheadaptations thatallowits
persistence andsuccess inits habitat. Twomaincomponentsoftheadaptation traits
thatecologists frequently investigate are the morphology andbehaviour(WOLF &
WALTZ, 1993;WAINWRIGHT,1994;McPEEK,1997). Morphological traitshave
dominatedthegeneral adaptationist program(GOULD&LEWONTIN, 1979)since
the sixties, because thesetraitsare more reliably measured, and many theoreticians
have produced testable models that instigate research (HUTCHINSON &
MacARTHUR, 1959; MacARTHUR & LEVINS, 1964). Now is the time for
behaviouralecology, mainlybecausemanyresearchers haveclaimedthatbehavioural
adaptations are more flexibleand couldbeanimportant wayto adapt toachanging
environment(WAINWRIGHT, 1994;McPEEK, 1995).
Odonatesare oneofthebestanimalmodelsforbehaviouralstudiesduetotheirsize
andtothecomplexreproductive behavioursthathavebeing studiedinthepastyears
306 P. deMarco,A.O.Latini&P.H.E.Ribeiro
(JACOBS, 1955;CAMPANELLA&WOLF,1974;ALCOCK, 1987;MILLER,1991;
CONRAD&PRITCHARD, 1992).Ingeneral,odonatespecies areclassifiedintofliers
andperchers basedontheirthermoregulatory adaptations (CORBET, 1962; MAY,
1976).Theflierspecies arealsoendothermicandcancontroltheirthoracictemperature
byhovering andwing whirling,whilepercherspecies are exclusively exothermicbut
canminimiseproblemsofhighthoraxictemperaturebyposture adjustmentandactivity
timeshifts(CORBET, 1999).
Ageneralunderstanding ofodonatebehaviourandtheways thatevolutionhasshaped
theactualbehaviouralcharacteristicsofthis groupcouldonlybeachievedby acareful
comparative method(HARVEY&PAGEL, 1991).Thisapproach needsthedescription
ofthemostrelevantaspects ofthebehaviourandgeneralbiology ofthespecies inthe
groupandasupported phylogenetic study. In spite oftheintensestudy ofodonates,
thereare onlyfewphylogenetic treesatthespecies level(CHIPPINDALE etal., 1999;
WESTMANetal., 2000)and fewgenuine comparative studies(McPEEK, 1995).
Herewepresentthedescriptivedataonbehaviourofaneotropicallibelluliddragonfly,
aspartofa general programdesignedtoallowforfuturecomparative studiesintothe
genusErythemis. WedescribethebehaviourofmaleandfemaleE. plebeja (Burnt.)
neara small pond in Visosa, MG, southeastern Brazil, paying special attentionto
reproductivebehaviourandtryingtoexplainsomeuncommonbehavioursweobserved
duringthisstudy.
METHODOLOGY
WestudiedapopulationofErythemisplebejaatasmallpondwitha70mperimeter,approximate20m2,
attheFederal UniversityofVigosacampus(20°45’S,42°51 W),Vigosa,Brazil. TheareaisnearanAtlantic
Forestresidual fragment,withasub-tropicalclimate(KbppenCWAclassification).Thepondwascovered
bySalvinia molestaandNymphaeaspp.overapproximatedlyonethirdofitstotalarea, andborderedby a
strip ofEleocharis,sp. It also had adominant presence ofthe submersed macrophyteMiriophyllum
brasiliense. Thepondwasinhabitedbysomepossibledragonfliespredators,inparticularadensepopulation
ofRanacatesbaeana Shawandblackbass (Micropterussalmoides).
Weobservedthebehaviour ofthisspeciesinthedryseasonfrom4to 12August 1994,withthe aidof
binoculars (7x35)orbyeye,from7:00a.m.to6:00 p.m.Thetotalobservationeffortwas77 hours. Dueto
thesmalldimensionsofthepondwewereabletocounttheabundance ofeachspeciesinanareaupto2m
aroundthepondevery 15 min.We also recorded theair temperature.Toimprovethemeasurements we
usedindividual marksonthe wingsusinganon-toxicblackpaint.
Fortheanalysisofthetemporalbudgetofthisspeciesweusedthefocalobservationmethod(ALTMANN,
1974).The samplingunitoftheseobservations isthe sequenceofbehavioursreportedduring 1min of
focalobservation ofanindividual. Themomentwhen eachbehaviour startswasnoted,andthe timespent
ineachactivitycalculated.Itis similartothemethodusedbyMAY(1977)butwereducedthetimespent
followingasingleindividual,duetothe frequentloss ofthe individualsduringobservation. Indeed,a
shorterinterval also hadstatistical advantagesbecause it allowedalargernumber ofdifferentindividuals
tobesampledineach occasion.
Thebehaviouralactivitieswere classifiedfollowingDEMARCO(1998),viz.:
(1) normalperching(NP);
(2) droop-wingedperching:the wingtipsareheld below thebody and/orthe wings shade the thorax
(DWP);
Behavioural ecologyofErythemisplebeja 307
(3) obeliskperching: thewingtipsareheldbelow thebody andtheabdomenwithananglecloseto90°
tothesubstrate (OB).
(4) transitionflight:passingthroughanareawithoutpatrolling(TRAN);
(5) foragingflight:fast short-distanceflightsreturningtotheperches andinterpretedasforaging(FO);
(6) territorialdefence: chasinganotherindividual (TD);
(7) copula(CP);
(8) oviposition(DS).
RESULTS
TheactivityofmaleErythemisplebejawasrestrictedtoasmallperiodoftimebetween
10:00and 16:00h, with peak abundance at 14:00 at the timeofthehighest air
temperatures(Fig. 1).However,the presenceofterritorialmales is notexplained by
temperaturealonebecause temperaturescompatible withtheiractivity occurearlierin
the morning (09:00-10:00 h),buttherewerefewor noindividualsatthepond. After
theperiodofintenseterritorialandreproductive activities(14:00-15:00) theabundance
atthepond decreasedsharply.
This species was always observedperched on bare soil near thepond bank. The
mostcommon postureobservedwas the droop-wing perch (Fig.2).Thisspecies flies
nearly11%ofthetotaltime,usuallyintransitionflightsbetweenperch sites.Weobserved
almost 15%ofthetotal timespentinreproductive activities, with a concentrationof
activitiesbetween 12:00and 14:30h.
In63%oftheterritorialdisputesobserved(n=27) maleE.plebejaengagedinintra-
specific disputes, with90%ofwinsby theresidentmale.Intheinter-specific disputes,
14,8%wereagainstTrameabinotataand22,2%againstErythrodiplax fusca. T.binotata
always wonagainstE.plebeja,butthelatteralways wonthedisputes againstE.fusca.
Fig. 1.AbundanceofmaleErithemisplebejaand airtemperatureat differenttimesofthe dayatasmall
pondin Viçosa,MG,southeasternBrazil. — [Barsrepresent standarderrors]
308 P. deMarco,A.O. Latini&P.H.E.Ribeiro
Fig.2.Time-budgetanalysisforErithemisplebejaatasmallpondinVi$osa,MG,southeasternBrazil. —
[Bars represent standarderrors; — NP=normalperching; —DWP=droop-wingedperching; — FO=
foraging; — TRAN = transitionflight; — TD = territorialdefence;— OV=oviposition; — COP =
copula]
Thisdominanceorderreflectsthedifferencesinbodysizeofthesespecies.Theterritorial
defencesoccurredmainlybetween 12:00hand 14:00h(Fig. 3),butweremoredispersed
during the day.
We followedlong sequences ofreproductive behaviours, including copula and
oviposition six timesduringthatseason. TableIshowsacomplete enumerationofthe
largest sequence observed. Thegeneral sequenceincluded: (a) the malefoundthe
femalewhileshe flew nearby; (b) they initiatedthecopula while inflight; (c) they
perched incopula; (d) themalewenttoaregionofthepond whilethefemalestaysat
thepondbankor patrolsnear themale;(e) themaleinitiatedasequencethatresembled
the oviposition behaviourofthefemale(‘simulatedoviposition movement’); (f) the
malewent to the femaleand initiatedoviposition in theregion were the malewas
earlier.
In50% (n = 18)of thecases, the malefollowedafemalethatentereditsterritory,
accompanying ittothemargin. Theycoupled, usuallyperched onthemarginalvegetation
oronthesoilfrom48to99seconds.Whendisturbed,they sometimesflewaway fora
fewsecondsandperched again. Afterspermtransfer, themalefliestotheoviposition
site while the femaleremains perched. The male then “simulated” oviposition
movements inthearea,sometimestouching thewater. Afterabout40secondsthemale
returned tothefemale,touched it,andthey flew together totheoviposition sitewere
the femalebegan oviposition. The maleexecutednon-contact guarding movements
aroundthe femalewithrapid linearmoves.
Inthe otheroviposition events observed, the femalewas aloneand they occurred
when there are few individualsinthe pond, mainly inthe evening. The ovipositing
Behavioural ecology ofErythemisplebeja 309
behaviourwas very distinctfromwhen the malewas together with thefemale.She
deposited eggs withoutadefinedarea,exploring withshortpauses alarge areaofthe
pond.
We observed on one occasion that, while themale was doing its simulationof
oviposition movements,aRanacatesbeianaindividualattackedhimbutfailed.
DISCUSSION
Erythemisplebejapresentsaclearbehaviouralpatternwhichagreestoitsclassification
as apercherspecies. P.DeMarcoJretal. (unpublished) discussedbody-size variation
and behaviouralvariations in neotropical percher species. Thegeneral pattern that
emerged fromthisanalysis was thatlargerperchers areusuallymoreaggressive, with
alarger proportion oftheirtimebudgetspentinterritorialdefenceactivities.Thedata
on E.plebeja supportthis claim. Itisoneof thelarger perches inthis pond andwas
generally observedinvolvedinterritorialcontests.
Oneproblem ofsuch generalisation is thatthebehaviouralenvironmentcouldbea
counfounding factorforthe determinationof the degreeofaggressiveness ofa given
species. Indifferentcommunities,wheretheysuffercompetitionfromdifferentspecies,
theirbehaviourwouldbeexpected tochange ifthe species chooseoptimal territorial
defence strategies. There are only a few studies that describe the behavioural
characteristics,especially theirtimebudgets (DEMARCO,1998).Ofcourse,replicated
studiesofthesamespecies indifferentbehaviouralenvironmentsaremuchrarer.These
studiesweregenerally neglected duetoarelatively typological viewofbehaviour(e.g.
percher vs flier)thathidtherealevolutionary nature ofthis discipline.
Fig. 3.Proportionoftime spentin territorial defence by male Erithemisplebejaat different times ofthe
dayatasmallpondinViçosa,MG,southeasternBrazil.— [Bars represent standarderrors]
310 P,deMarco,A.O.Latini &P.H.E.Ribeiro
TableI
Behavioural sequenceofamale ErithemisplebejaatsmallpondinVi?osa,MG,southeasternBrazil. The
observationstarted ata 14:00 hwithanairtemperatureof26,0°C
PPoossttuurreeoobbsseerrvveedd oonnssppeecciimmeennss SSeeccoonnddss CCuummuullaattiivvee
eellaappsseedd ttiimmee
SSttaarrttiinnggoobbsseerrvvaattiioonn:: <3Jiinnoovviippoossiittiioonnpprroocceedduurree 1100 1100
25 aanndd 33iinnccooppuullaattiioonnfflliigghhtt 1144 2244
29 aanndd<3Jiinn ccooppuullaattiioonnppeerrcchheedd 5544 7788
63 ssttaarrttsstthhee ““oovviippoossiittiioonnssiimmuullaattiioonn ppoossttuurree””aanndd tthhee 29iissppeerrcchheedd 4466 112244
AAfftteerrtthhee““oovviippoossiittiioonnssiimmuullaattiioonn”” ooff63iissffiinniisshheedd,, rreessttaarrttss tthheeoovviippoossiittiioonn
pprroocceedduurreewwiitthhtthhee â6ppaattrroolllliinnggaattttuurrnn 1100 113344
33 cchhaasseessaaEE..ffuussccaaaannddwwiinnss,, 29 oovviippoossiittiioonniinngg 22 113366
29 oovviippoossiittiioonniinnggaanndd 33ppaattrroolllliinnggaattttuurrnn 5599 119955
29 aanndd63iinn ccooppuullaattiioonnfflliigghhtt 44 119999
33 ppeerrcchheeddaanndd 29oovviippoossiittiioonniinngg 1111 221100
33cchhaasseess aannEE..ffuiissccaaaaggaaiinnaannddwwiinnss,, 29 oovviippoossiittiioonniinngg 99 221199
33 ppeerrcchheeddaanndd 29oovviippoossiittiioonniinngg 7744 229933
29 aanndd 33 iinn ccooppuullaattiioonnfflliigghhtt 44 229977
29 aanndd 33iinnccooppuullaattiioonnppeerrcchheedd 6677 336644
33iissppaattrroolllliinnggaannddtthhee 29iissppeerrcchheedd 1144 337788
33 ssttaarrttsstthhee““oovviippoossiittiioonnssiimmuullaattiioonn”” aanndd tthhee 29 iissppeerrcchheedd 4433 442211
AAfftteerrtthhee““oovviippoossiittiioonnssiimmuullaattiioonn”” iissffiinniisshheedd,,tthhee 36 iissppeerrcchheeddnneeaarraanndd
lliikkeetthhee 29 2211 444422
63cchhaasseess aannootthheerrEE..ffuussccaa aannddwwiinnss,,tthhee 29rreessttaarrttsstthheeoovviippoossiittiioonnpprroocceedduurree 22 444444
29iinn oovviippoossiittiioonnpprroocceedduurree,,wwiitthh tthhee 33 ppaattrroolllliinnggaattttuurrnn 3399 448833
AAssppeecciimmeennooffRRaannaaccaatteessbbeeiiaannaa iinnvveessttss oovveerrtthhee 33;;tthhee 33fflliigghhttiinnttoo tthhee
eeddggee ooffttrreeeess aannddtthhee 29iissnnoottoobbsseerrvveeddnneeaarrtthheeppoonndd 11 448844
Actually, wehaveonly general behaviouralinformationfromE. peruviana (P. De
Marcoetal., unpublished), E. vesiculosa (DEMARCO, 1998)andE. simplicicollis
(McVEY, 1988). The first two ofthese studies produced a time-budget thatcould
allowustotentativelycomparethesespecies. E.plebejaismoresimilartoE.peruviana,
thatalsospentnearly 10%oftheirtimeinterritorialdisputes.Bothspeciesappearvery
differentfromE. vesiculosa, thatspends alarge proportionoftimeinpatrolactivities.
Thelattercomparison is subjectto criticismbecausethedataforE. vesiculosacame
fromaforaging areawhiletheotherspecies wereobservednearreproductive sites.
As alargelibellulid, E.plebeja isalsomoreefficientatthermoregulating insunny
sites atlow air temperature,due to itscapacity toheat by irradiation(MAY, 1979,
1991). Compared to otherdragonfly species, weobservedamuch largerpartof the
daywithterritorialdefenceofsites, probablyas aconsequenceofits thermoregulatory
capabilities.
JACOBS(1955)reported maleexaminationofoviposition sitesbeforethefemale’s
proper oviposition in Perithemistenera. He noted that predators, including frogs,
sometimeskill malesexamining sites.As thepond duringthisstudy wasinhabitedby
BehaviouralecologyofErythemisplebeja 311
adensepopulationofRanacatesbeiana,thatwereoftenobservedattackingthisspecies,
thisbehaviourclearly increasetherisk of malesbeing predated.Itis very difficultto
propose anexplanation forthis apparentaltruistic behaviour.We consider thatthis
behaviourcouldonly beevolvedifthedecreaseinmalefitnesswas accompanied by an
increasedsurvivaloffemalesand,consequently, theiroffspring. Inallcases,themale
firstcopulatesandtheninitiatestheexplorationoftheoviposition site. Thisbehaviour
probably secures itsfitness, evenifheiskilledduringtheexaminationofthesite.We
donotobserve thisbehaviourintheotherspecies ofthegenera,anditclearlydeserves
furtherstudiesto understandthepossible impact onmalefitnessand alsoitsspread
amongodonates.
ACKNOWLEDGEMENT
This workispartlysupportedbyFAPEM1G,CNPq andIPEMA.
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