Reproductive behaviour of Leucorrhinia albifrons (Burmeister) in a non-territorial situation (Anisoptera: Libellulidae)

11 Pages·1998·1.2 MB·English

by  Reinhardt, K

Odonatologica27(2):201-211 June 1, 1998
ReproductivebehaviourofLeucorrhinia
albifrons (Burmeister) inanon-territorial
situation(Anisoptera: Libellulidae)
K.Reinhardt
Institute ofEcology,Friedrich Schiller University,DornburgerStr. 159,D-07743 Jena,Germany
Received May9, 1997/RevisedandAcceptedAugust20, 1997
The reproductivebehaviour in ahigh-densitysituation ofthe sp.wasinvestigated
innorthernPolandin 1993 and 1995. Becauseoftheir highdensity,males werenon-
-territorial.Afteraverybrieftandemflight,copulationstookplace eitherontheground
or in the surroundingpine trees and lasted,onaverage, 640 s. Afterwards the male
guardedthe femalewhile she wasonpost-copulatoryrest (PCR).DuringPCR (which
occurred in 11outof 13observed cases),the malebent hisabdomen tipuptothe basal
segments after a meantimeof 81s. Thisbehaviour wasinterpretedas intra-male
sperm translocation (ST). Ovipositiontook placeoveropenwater. During mating
time,from 10:00h to about 17:00CEST, almostnoneofthe observed ovipositions
was completedundisturbed. Outside thistime,half oftheovipositions observedwas
completedundisturbed.The latter lasted 34 s, in which the female had anaverage
frequencyof0.6 to 1.9dipsper s.Elevenhand-held ovipositionsrevealed ameanegg
number of327 eggsper female and amean egg flowrate of4,6eggs s'.Itiscon-
cludedthat the matingsystem ofL. albifronsis best described asa combination of
resourcelimitation and female control. Someknown effects ofhighmaledensity on
thereproductive behaviour oftheLibellulidae arediscussed.
INTRODUCTION
Libelluliddragonflies areamongthebest studiedanimals in regardto theirre-
productive biology, including speciesrecognition (SINGER, 1990),materecogni-
tion (PAJUNEN, 1964a),territorialbehaviour (PARR, 1983;WOLF & WALTZ,
1988),alternative mating behaviour(WALTZ & WOLF, 1984; CONVEY, 1989;
WOLFetal„ 1989;WOLF& WALTZ, 1993),sperm competition(SIVA-JOTHY,
1984;MICHIELS&DHONDT, 1988),genital morphology (MILLER, 1991)and
underlyingfactorsofoviposition siteselection(WOLF &WALTZ,1988). Manyof
theinvestigations werecarriedout inthegenusLeucorrhinia.Generaldescriptions
202 K. Reinhardt
ofthematingbehaviourhavebeenmadeforL. rubicunda(MUNCHBERG, 1931;
PAJUNEN, 1963;PAJUNEN, 1966;RUPPELL, 1990),L. dubia(MUNCHBERG,
1931; STEINER, 1948; SCHIEMENZ, 1954; PAJUNEN, 1962a, 1964a), L.
pectoralis (KIAUTA, 1964)and.L.hudsonica(HILTON, 1984).Detailedinforma-
tion on aggressive interactionsbetween males are available for L. caudalis
(PAJUNEN, 1964b)and L. dubia(PAJUNEN, 1962b). PAJUNEN (1964a) and
SINGER (1990)showedthatspeciesrecognition is notperfect withinthesympatric
Leucorrhiniaspecies L. dubiaand L. rubicunda, andwithinL. intacta,L.frigida
andL.proxima,respectively. Allspecies studiedsofarhavebeenshownto defend
territories,eitherfromperches (PAJUNEN, 1966),by patrolling,or by bothtactics
(WOLF&WALTZ,1984,1993).Perching mightbecostly.First, SMITH&COOK
(1991) showedthatterritorialmalesofL.frigida havehigherloadsoftheparasitic
water mite Limnocharesamericana.Second, in high- density populations of L.
dubia(PAJUNEN, 1962a,1962b),L.caudalis(PAJUNEN, 1964b)andLrubicunda
(PAJUNEN, 1966),theterritorialsystem istoo costly andbreaksdown.Theaimof
thepresentpaperis to describeaspects ofthereproductive behaviourofthehith-
ertounstudiedspecies L.albifrons in such ahigh density situation.
MATERIALANDMETHODS
The study wasundertaken atonepeat bogpondinachainoffourinthe KaszubLakeDistrict, 18
kmnortheast ofChojnice,northeastern Poland(53°46‘N, 17°42‘ E, approx.200 m alt.).The pond's
diameterwasabout 100m.FloatingSphagnummosses,leavesofthewaterliliesNupharandNymphaea,
and stems ofPeucedanumpalustrewerethe dominantvegetation.The moss vegetationextendedall
aroundthebanks and atawaterdepthofabout3m wasathick layerofmud.The pH ofthe waterwas
5.5.The studysite anditsclose surroundingsharbourarichdragonflyfaunathatisdetailedelsewhere
(BROCKHAUS, 1990; LABIjDZKI, 1994; REINHARDT, 1994).The study wasconducted between
18-VIand 1-VII,1993 and between 29-VII and 4-VIII,1995 when dailyobservationstookplace.The
weatherwasrather changeableinthe 1993 observationperiodandwarmand sunny in 1995.
Individualswereobserved unaidedorbybinocularsandasmany variables aspossibleweremeas-
ured tothenearest second accordingtothefollowingprotocol: the duration oftandem, copula and
intra-malespermtranslocation (ST),thetimeofST after copulaand duration ofpost-copulatory rest
(PCR-timefromcopulatotheonset ofovipositionflightby thefemale).Furthermore,thenumberof
copulationattempts by the male, location ofcopulation,the duration ofoviposition and whether or
notmate-guardingtookplacebythemale werenoted.Duringoviposition,thenumber ofdips,distur-
bancesbymales andthe distancefromthepondmarginwerealso noted.Intotal,27datasetscouldbe
usedfromthe 1993 seasonand 34fromthe 1995 season.Additional datawerederivedfrom 12incom-
pletevideosequences.Forallevents,date andtimeofdaywerenoted.Elevenfemaleswerecaughtfor
hand-heldovipositions (STEINER, 1948; McVEY, 1984)and their duration wasmeasuredineight
females. Theyweredefined asbeing finished when noeggs werelaid for60s.Each female’s eggs
werestoredseparatelyandwerecountedin thelaboratory.
Ata certain stage very shortly aftercopulationthemalebent his abdomentip upto thebasal
segments (see Results), exactly as hasbeen describedfor theintra-malespermtranslocation(ST, see
UTZERI, 1985;UTZER1&OTTOLENGHI, 1992).Todetermineifthis wasST(whichusuallyoccurs
beforecopulation,UTZERI,1985;UTZERI&OTTOLENGHI, 1992)orserved,forinstancetoclean
penis ofhisown orforeign sperm,twelve males (three in copula, two afterST, and sevenin an
Reproductivebehaviourin Leucorrhinia albifrons 203
unknown stage)werecapturedin 1995. Theyweredried and laterinvestigatedforthe presence ofa
sperm crustonthepenis.Ifpenis cleaningoccurred duringthis abdomenbending,onewould expect
tofindno spermcrustatthepenis ofthose malesthatwerecapturedinanunknown stage,oratleast
notinthose malescapturedimmediatelyaftertheyshowed this abdomen bending.
Inordertodescribethetemporalpatternofmaleandfemale arrival,onepondbankabout 50m long
wassearched formales,femalesandcopulationwheels.Countsweremadein 15minintervals andthe
respective ambient temperature wasmeasured inthe shadetothe nearest 1°C.Altogether,34 counts
werecarriedoutin 1993 and64in 1995.Datawerethenpooledforhoursandanaveragevaluesgiven.
Within-yeardataarecomparedby
non-parametricstatistics using the
statistic programmeSPSS 6.0.1.In
the following,means± 1 S.D. with
therange and samplesizesinbrack-
etsaregiven.
RESULTS
MATINGACTIVITY
Under the warm weather
conditionsof1995,nodistinct
diel mating pattern was de-
tected. In the more change-
able period of 1993, mating
seemedtopeak around 13:00
CEST (Fig.l).At the mating
site, there was usually a2.5
to 5 foldbias towards males
withatotalofapproximately
10males per50m shoreline
(Fig.l).However,thereisalso
a strong correlationofmale
and female presence at the
water. Atmaximummaleac-
tivity, nearly all females ob-
served were in copula. The
percentageofsolitary females
was higher only in early
morning andlateevening.
Fig. 1.Presence ofmalesandfemales atonebank (length:
DESCRIPTIONOFTHEREPRO- approximately50 m)oftheovipositionpond.The percent-
age offemales in copulais also given. Data are based on
DUCTIVEBEHAVIOUR single counts (atleast 15min apart)andare pooledfor4
dayseach inboth 1993and 1995.Weathersituationsin 1993
Fromtheparametersmeas- preventedfurtherobservations. See Methods forfurtherex-
ured, only wheel perching planation.
204 K. Reinhardt
height, oviposition durationandthenumberofdipspersecond differedbetween
thetwo years.Thus,allother variablesare combined.
Themalegraspedthefemaleusually duringthefirst orsecondattemptalthough
thirdattemptswerealsoobserved.Thetandempositiondidnot lastlongerthan 1 s
if mating occurred. No ST couldbe detected during this shorttandem time.If
copulation was refusedafterthe tandemattempt, tandemdurationsofup to 40s
wereobserved.In 1993,copulation tookplace atheights of0to 10cm(n=19)but
wereobserved atheights of0to 400cm(average 114± 120cm,n= 22)in 1995.
Matings, on average,lastedfor 640±426 s(73 - 1827s,n=19). Iftwo extraordi-
narily shortcopulations of73 and80 s areexcluded,themean wouldbe706.5±
400.0s.Abdomenpumping, duringwhichrivalspermisremovedfromthefemale
bursa copulatrix, was only recorded 3 times. Inthese cases itendedafter326 s
(copulation duration:511 s), after246 s(copulation duration663s), andafter117
s(copulation duration291 s). During mating, thepairchanged itssite0to 5 times
(n=25). Theseflights, whichwere usually overshortdistancesonly, occurred spo-
radically orwere inducedby approaching antsorotherinsects. Whenothermales
ofL. albifrons, Lepidoptera orasilidfliesapproached acopulating pairthe copu-
lating malebriefly opened his wings,resulting ina retreat ofthe disturbing indi-
vidualinall cases. During copulationboth sexes groomed theheadwith thefront
legs. Usually, thefemaleused allthreepairs oflegs tohold themale’sabdomen.
After theendofcopulation, whenmales andfemalesseparated, bothsexesrested
ontheground, closetoeachother(onaverage50±44cm, 15to 120cm,n= 13).In
one case theterminationofcopulation was preceded by astrong vibrationofthe
wholefemalebody including thewings.During PCR, themalebenthisabdomen
tip to thebasal segments. This occurredafter 81 ±71 s (range 18 to 279 s) and
lastedfor2to3 s(n=12),inonecase for6s.Twice,thisabdomenbending was not
observeduntilthemaleescaped after300s and376 s.
Ofthetwelvemalescaptured toinvestigate thepurposeoftheabdomenbending
behaviour, only one(in anunknown stage) didnot haveaspermcrustatthepenis
head. All others, including the two males captured immediately after abdomen
bending hadspermcrusts.
PCRoccurred in 11 of 13 cases. Inthetwoothercases, thepairflew in copula-
tionwheelto theopenwater. Bothtimesthefemales weregrasped by othermales
immediately afterbeing releasedby thefirstmale.Afteranaverage timeof 152±
98 s (27 - 376 s, n= 11)ofPCR, females flew to oviposit. Inthree out ofthese
elevencases themaletriedto initiateoviposition by flying infrontofthefemale.
Occasionally, femalesleftthesiteofPCRandflewtothesurrounding forest.There
wasnorelationbetweencopulation durationandPCRduration(rs=0.429,p=0.397,
n= 11, N.S.). In thehours from 10:00hto 17:00h CEST,only two undisturbed
ovipositions were observed (outofsomehundredobserved). Before orafterthis
time, (8:00h to 10:00h and 17:00hto20:30 h CEST),7 out of 14 (50%) were
undisturbed.They lasted, onaverage, 34.4± 16.6 s(14 to53 s,n=5) in 1995but
Reproductivebehaviourin Leucorrhiniaalbifrons 205
were significantly longerin 1993(Mann-Whitney U- test,U= 1.0,p= 0.010) when
they lastedfor91.8 ± 49.8s (range 51 - 180s, n=6). I severaltimes observed a
male remating with the samefemaleimmediately after acopulation. When fe-
males arriving at the oviposition site were disturbedthey were chasedby up to
seven males.
In 1993, during single undisturbedoviposition bouts, females dipped 82 ± 49
times (range26to 154,n=5) onthewater surfacebut42±25times (range 14-69,
n= 6) in 1995.In contrast, therewas also a differencein the dipping frequency
betweenthetwoyears. In 1993,ameanof0.82±0.23dipspers(range0.6to 1.12,
n=6) was observedbutin 1995,theobserved meanwas 1.41± 0.31 (range 0.6to
1.90, n= 13). The differencebetween the two years was statistically significant
(Mann-Whitney- U- test,U= 7.5, p= 0.003). In 1995,five completed ovipositions
had thesame dip frequency as ten disturbedones, whereonly thefirstpartofthe
dipping sequencecouldbeobserved (Mann- WhitneyU- test,U= 14.5,p= 0.196,
N.S.). Thecomplete sequences show apositive correlationofdipping frequency
with temperature(rs= 0.900,p= 0.037, n=5). Nosuchrelationship was evidentin
1995(rs= -0.041, p= 0.889, n=14). The eggswere laid in openwater,usually at
least5 to 10m fromthebank. Oviposition flights were carriedoutinlong loops.
TheotherLeucorrhiniaspeciespresent,L. dubia,,alwayslaiditseggswithintwo m
ofthe margin.
HAND-HELD OVIPOSITIONS
Egg flowduringhand-heldovipositions atair temperaturesof24 - 28°Clasted
69.2± 20 s(range 36- 100s, n=8), whereby 326± 146eggs (range 184 to723
eggs,n= 11)were laid.Fromthesedata,ameaneggflowrate of4.6±2.5eggsper
s (range 2.3to9.9 eggspers, n= 8) canbe derived. Noambienttemperature de-
pendent egg release rate occurred (rs= -0.156, p= 0.711, n= 8). However,hand-
-heldovipositions lastedlongerthanundisturbedones(Mann-Whitney U-test,U-
2.0,p=0.008),indicating somesortofunnaturalpatternduringhand-heldoviposition
inthisspecies, resulting inanunderestimationof theeggflowrate.
DISCUSSION
OVIPOSITION
Givenanabdomenlength of24- 27mminL.albifrons (SCHIEMENZ,1981)an
eggflow rate of6- 11 eggsper swouldhavebeenexpected at32 ±3°C(McVEY,
1984).Ifound no differencein the dipping frequency between undisturbedand
disturbedovipositions, indicating thatit is fairlyconstant within one oviposition
bout.During hand-heldovipositions, L.albifronsfemalescaught duringor shortly
206 K.Reinhardt
aftercopulation laid327eggs.Givenameanof82dipsperundisturbedoviposition
in 1995,approximately 4eggs were laidper dip. This is, however, a veryrough
estimateas littleisknown abouttheeggflowpatternwithincreasing dipnumber.
REHFELDT(1991)foundthatundisturbedovipositions werecarriedout withhigher
diprate thandisturbedonesin Crocothemiserythraea atoneoftwo sites investi-
gated. However, anotherimportant factor inegg releaserate,the female’spresent
eggload(WATANABE&HIGASHI,1993),hasrarely beeninvestigatedand might
interferewith suchresults.
Table I
Effectsofhighmale densityon aspectsofthe reproductivebehaviour oflibellulid dragonflies
BBeehhaavviioouurraassppeecctt EEffffeecctt SSppeecciieess RReeffeerreennccee
nnuummbbeerrooffoovviippoossiittiinnggffeemmaalleess ddeeccrreeaassiinngg LLeeuuccoorrrrhhiinniiaaiInnttaaccttaa WWoollff&&WWaallttzz,, 11998888
tteerrrriittoorriiaalliittyy iinnccrreeaassiinngg LLiibheelllluullaajJuulliiaa HHiillttoonn,, 11998844
LLiibheelllluullaalluuccttuuoossaa MMoooorree,, 11998877
NNeesscciiootthheemmiissnniiggeerriieennssiiss PPaarrrr,, 11998833
AAcciissoommaappaannoorrppooiiddeess HHaassssaann,, 11997788
ddeeccrreeaassiinngg LLiibheelllluullaassaattuurraattaa DDeeBBaannoo,, 11999933
LLeeuuccoorrrrhhiinniiaa rruubbiiccuunnddaa PPaajjuunneenn,, 11996666
LLeeuuccoorrrrhhiinniiaa dduubbiiaa PPaajjuunneenn,, 11996622aa
LLeeuuccoorrrrhhiinniiaa ccaauuddaalliiss PPaajjuunneenn,, 11996644aa
LLeeuuccoorrrrhhiinniiaaaallbbiiffrroonnss tthhiissssttuuddyy
iinnddiivviidduuaallmmaalleemmaattiinnggssuucccceessss ddeeccrreeaassiinngg LLiibheelllluullaalluuccttuuoossaa MMoooorree,, 11998899
iinnccrreeaassiinngg PPaacchhyyddiippllaaxxlloonnggiippeennnniiss MMccKKiinnnnoonn&&MMaayy,, 11999944
ffeemmaalleerreejjeeccttiioonnrraattee ddeeccrreeaassiinngg LLiihbeelllluullaalluuccttuuoossaa MMoooorree,, 11998899
ccooppuullaattiioonn dduurraattiioonn nnoocchhaannggee LLiihbeelllluullaalluuccttuuoossaa MMoooorree,, 11998899
mmaatteegguuaarrddiinngg lleessss lliikkeellyy LLiihbeelllluullaalluuccttuuoossaa MMoooorree,, 11998899
iinnccrreeaassiinngg SSyymmppeettrruummppaarrvvuulluumm UUeeddaa,, 11997799
PPaacchhyyddiippllaaxxlloonnggiippeennnniiss SShheerrmmaann,, 11998833
oovviippoossiittiioonndduurraattiioonn ddeeccrreeaassiinngg LLiihbeelllluullaalluuccttuuoossaa MMoooorree,, 11998899
oovviippoossiittiioonnffrreeqquueennccyy ddeeccrreeaassiinngg PPaacchhyyddiippllaaxxlloonnggiippeennnniiss MMccKKiinnnnoonn&&MMaayy,, 11999944
ooppeerraattiioonnaallsseexx rraattiioo mmaalleebbiiaasseedd LLiihbeelllluullaa lluuccttuuoossaa MMoooorree,, 11998899
uunnddiissttuurrbbeedd oovviippoossiittiioonnss ddeeccrreeaassiinngg LLeeuuccoorrrrhhiinniiaarruubbiiccuunnddaa RRiiiippppeellll,, 11999900
PPaajjuunneenn,, 11996666
PPaacchhyyddiippllaaxxlloonnggiippeennnniiss RRoobbeeyy,, 11997755
LLeeuuccoorrrrhhiinniiaaaallbbiiffrroonnss tthhiiss ssttuuddyy
ffeemmaalleeaarrrriivvaalliinn ccooppuullaattiioonnwwhheeeell iinnccrreeaassiinngg LLeeuuccoorrrrhhiinniiaarruubbiiccuunnddaa RRiiiippppeellll,, 11999900
ssiittttiinnggoovviippoossiittiioonn ooccccuurrrriinngg LLeeuuccoorrrrhhiinniiaarruubbiiccuunnddaa RRiiiippppeellll,, 11999900
mmaalleeppeerrcchhiinnggttiimmee ddeeccrreeaassiinngg LLeeuuccoorrrrhhiinniiaarruubbiiccuunnddaa RRiiiippppeellll,, 11999900
hheetteerroossppeecciiffiiccttaannddeemmffoorrmmaattiioonn iinnccrreeaassiinngg LLeeuuccoorrrrhhiinniiaarruubbiiccuunnddaa RRiiiippppeelll,l, 11999900
SSyymmppeettrruumm ddeepprreessssiiuussccuulluumm RReehhffeellddtt,, 11999933
mmaalleeiinntteerraaccttiioonn dduurraattiioonn iinnccrreeaassiinngg LLiibheelllululalassaattuurraattaa DDeeBBaannoo,, 11999933
ssaatteelllliitteemmaalleebbeehhaavviioouurr iinnccrreeaassiinngg LLiibheelllluullaaqquuaaddrriimmaaccuullaattaa CCoonnvveeyy,, 11998899
Reproductivebehaviour in Leucorrhinia albifrons 207
MATINGSYSTEM
Thedifferencesbetweenthetwo yearswithamore distinctmaleactivitypeak in
the cooler 1993 season support a temperature dependent mating activity rather
than a dependence on the timeof day. It is interesting thatmaximum density in
1993(12malesper50mofshore)was closetotheplateau reachedin1995(9to 11
malesper50mofshore) indicating thatthiswas themaximumdensity.
Thehigh densityofmalesresultedinahighpercentageofdisturbedovipositions.
Therefore, very few mating sequencescouldbe observed completely. Neverthe-
less, thematingpatternobserved inLeucorrhiniaalbifrons corresponds wellwith
other findings withinthegenus (PAJUNEN, 1962a, 1962b, 1963, 1964a, 1964b,
1966;KIAUTA,1964;HILTON, 1984;WALTZ&WOLF,1984;WOLF&WALTZ,
1984, 1988, 1993), including the lack ofcourtship, the duration ofcopulation,
repeated mating inboth sexes,temperaturedependence ofmating peaks, common
occurrence ofPCR, highpercentageofinterrupted ovipositions whendensitiesare
high, dippingfrequency, eggnumber,andeggflowrate. Thepresentinvestigation
showed forboth years that males donot defendterritories. This is in contrast to
casual observationsofintra- and inter-specific male aggression atneighbouring
ponds andatdifferentlocalitiesineastern Germany wheredensitieswerelow(less
thanthreemalesper50mofshore, unpubl. data).Incontrast toPAJUNEN(1962a)
Iobserved amaletopair withthesamefemaleseveraltimes.Thispossibly was a
resultofthe high population densityandthemale’s inability torecognize thepre-
viousmate. Othertypesoflibellulidreproductive behaviourassociatedwith high
density are shown inTableI, indicating thatthereis no typical libellulidresponse.
Anotherbehaviourmost likely connectedwith high densitieswasthe occurrence
ofSTjust aftercopulation. Up tonow, thishasbeen reported foronly one other
dragonfly species,i.e.Pachydiplax longipennis (ROBEY, 1975).NeitherPAJUNEN
(1963)from film sequence analyses ofL. dubiaand L. rubicunda nor KIAUTA
(1964) inL.pectoralisandHILTON(1984)inL.hudsonica didobserve STbehav-
iourand concludedthat spermtransfer in Leucorrhiniais generally not immedi-
ately followedby copulation. At least in PAJUNEN’s (1963) study, the species
showed similarhigh densitiesas doesL. albifrons inthepresentstudy. Itherefore
assume thatST immediately aftercopulation is an advantage for a malein high
densitypopulations. Itenableshimtobeimmediatelyready forfurthercopulations.
Maximizing his mating bouts wouldincrease his paternity chances. ST immedi-
ately aftercopulation wouldthen beadaptive.
PCRhasbeenobservedinmanylibellulids(e.g.PRENN, 1930;PAJUNEN, 1963;
MILLER, 1983;MILLER& MILLER, 1989;REHFELDT, 1989;LEE, 1994;
WILDERMUTH,1994).Although listingonly afewofthemandincorrectlyciting
MILLER & MILLER’S (1989) observations on the proportion of Orthetrum
coerulescenscopulations withPCR, LINDEBOOM (1996) concludedthat PCR
nearlyalwaysoccurs. REHFELDT(1989)observedthatonly86%ofthecopulations
208 K.Reinhardt
were followedby PCR, and also PRENN (1930) mentionscopulations both with
andwithoutPCR inL. dubia.MILLER& MILLER(1989)andREHFELDT(1989)
summarizedpossible functionsofPCRforthefemales:testing themale’sguarding
quality,avoiding maleinterference,assessing predationpressure byawaitingother
ovipositing females, preparing eggs for deposition, or sperm handling. While
MILLER & MILLER’S (1989) data favour the sperm handling hypothesis,
REHFELDT’s (1989)dataindicateasignificant influenceofthepresenceofother
males, bank vegetation, and disturbance (see however LEE, 1994). For the
Calopterygidae LINDEBOOM (1996) clearly confirmeda sperm handling func-
tionandassumed that the spermexpelled is that fromthe previous male. Ifthis
holdstruealso for theLibellulidae, onlycopulations with virgin femalesor those
with insufficientsperm reserves shouldbeexpected withoutPCR. However, the
lackofPCR couldalsobearesultofcontact-mate guarding. Thiswas thecase in
thepresentstudy where2outof 13observedcopulations were withoutPCR dueto
contact-guarding. Thus,the behaviourpattern describedabovecan alsobe seenas
a consequenceofa sexualconflictbetweenmaleandfemale.Ifthemalecontrols
theterminationofcopulation heshouldflyin wheelposition with hismate tothe
oviposition site.Inthecaseoffemalecontrolshe shouldhavethechoiceofwhether
toinitiatePCR(for whateverreason) or toescapeandlaterreturntotheoviposition
site.Inbothcases, however,themale’scontrolofpaternity decreases.Inthis situ-
ationit wouldbe advantageous fora maleto beready forcopulation with other
femalesandtoperformSTwhile“waiting” forthefemale’sdecision(to leaveorto
oviposit)orby stimulatinghertooviposite.g.by fluttering infrontofher(MILLER
& MILLER, 1989;this study).
The frequent male-femaleencounters, the partial ability of males to control
oviposition sites,theoccurrence ofguarded andunguarded ovipositions, andmul-
tiple matings per day by the two sexes leadto the acceptance ofone ofthe two
resource-based mating systems (sensu CONRAD & PRITCHARD, 1992) for L.
albifrons. Outside themain mating activity, females are able to oviposit uninter-
rupted, clearly indicating thatmalescannot completely control oviposition sites.
Thus, for L. albifrons the resource-limitation mating system (CONRAD &
PRITCHARD, 1992)applies. Thelong copulation durationin L.albifrons (up to
30 min,mean 11 min inthisstudy) andthegenusLeucorrhiniaingeneral (15-40
min inL dubia,PAJUNEN, 1962a;20-30min inL. caudalis,PAJUNEN, 1964b;
15-25 min in L. pectoralis, KIAUTA, 1964;approx. 15 min in L. hudsonica,
HILTON, 1984),theabsenceofcourtship display, andtheoccurrence ofcontact-
-guarding favours the assumption ofa non-resource-basedmating system (see
CONRAD & PRITCHARD, 1992).Thus, themating system couldat bestbe de-
scribedas acombinationoftheresource-limitationandfemale-controlsystems. A
combinationoftwo systems ratherthanonepurestrategywas previously described
forLestes sponsa(STOKS etal., 1996).However, thelackofreliableinformation
on the mating behaviourof L. albifrons at lower density remains a problem in
identifying the mating system ofL.albifrons.
Reproductivebehaviour in Leucorrhinia albifrons 209
ACKNOWLEDGEMENTS
Iacknowledgethe commentsbyS.GORB,Kiev, A.MARTENS,Braunschweig,S.HALLE, Jena,
and ananonymous refereeon apreviousversion,which greatlyimprovedthemanuscript.
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