Wandering males are smaller than territorial males in the damselfly Calopteryx virgo (L.) (Zygoptera: Calopterygidae)

2009·1 MB·English

by  Koskimäki, J

Odonatologica38(2): 159-165 June I.2009
Wanderingmalesare smaller thanterritorial
males inthe damselfly Calopteryx virgo (L.)
(Zygoptera: Calopterygidae)
J. Koskimäk¹i,M.J.Rantala²andJ.Suhonen²*
1SectionofEcology,DepartmentofBiology,UniversityofOulu,POBox 3000,
FIN-90014University ofOulu,Finland
2Section ofEcology,DepartmentofBiology,UniversityofTurku,FIN-20014 Turku,Finland
Received September17,R2008/ evisedandAcceptedJanuary15,2009
Interritorial Odonata,adult 63mayuse2matingtacticsthatmaybe genetical-
lyorenvironmentallydetermined:territoriality and non-territoriality. The non-ter-
ritorial tactichasbeensometimesfound toinclude 2additional males: sneakingand
wandering.Thenon-territorial 6 3,however,oftenhave lowerreproductivesuccess
than territorial 33.Studiesonvarious Calopteryxspp. haverepeatedlyshownthat
territorial andnon-territorial behaviours areconditional matingtacticsandthatbody
sizedoesnotpredict3resource-holdingpotentialandterritorialbehaviour. Instead,
theresourceholdingpotentialseemstodependonthe amountof 3 fat resources.
Here,bothterritorial andwanderingC. virgo33werecollected fromacreek incen-
tralFinland. Itwasfoundthatterritorial 33werelargerand heavierthanwandering
33.Thedatashowthatthesizeoftheindividual maypredictthereproductivetactic
ofsomeodon. 33 toagreaterdegreethanpreviouslythought.
INTRODUCTION
Alternativemating strategies and tacticsevolvebecause ecologicaland social
environmentsoftenfavourmorethanonephenotype (GROSS, 1996;SUHONEN
etah, 2008). However,not all malesare abletoobtaina territory.Interritorial
odonatespecies, adultmalesmayusetwoorthreematingtacticsorstrategies that
maybe genetically(SIVA-JOTHY&TSUBAKI, 1989a, 1989b;WATANABE &
TAGUCHI, 1990;TSUBAKI, 2003)orenvironmentally determined, forexample
being territorialor not(FORSYTH & MONTGOMERIE, 1987; PLAISTOW
&SIVA-JOTHY, 1996;TYNKKYNENetah,2006;SERRANO-MENESES et
*Correspondingauthor:[email protected]
160 J.Koskimaki, M.J.Rantala&J.Suhonen
al.,2007; RAIHANI etal.,2008), SomeCalopterygidae malesmayalso actively
moveandsearchforfemales, withoutdefending anyterritoryduringtheirlifetime
(RAIHANI etal.,2008). Thisnon-territorialtactichasbeen occasionally men-
tionedtotaketwoadditionalmodes:sneaking andwandering (WATANABE &
TAGUCHI,1990;CORDOBA-AGUILAR, 1995). Intheformer,non-territorial
malesconsistently stay in the same places while in the latter, malescross many
territoriesanddo not showany sortofsitefidelity. Malesmayadoptalternative
mating tacticsandbehave as sneakers, trying to intercept femalesarriving atat-
tractiveterritorialsites beforebeingdetectedby theresident males(PAJUNEN,
1966).InHetaerinaamericanamaleswere foundto usethreematingtactics: ter-
ritorial, non-territorialand switcher (individuals withbothterritorialandnon-
territorialperiods during theirlife-span) andallthreetacticshavesimilarmating
success (RAIHANI etal., 2008). Thenon-territorialtactic, however, is usually
not as successful as the territorialone (SUHONEN et al., 2008). For example,
it was shown that the mating success of territorial Calopteryx splendens xan-
thostomamalesmaybethousandtimeshigherthanthatofnon-territorialmales
(PLAISTOW&S1VA-JOTHY, 1996). Malefitness is alsoaffected by territory
quality. For example, C. s. xanthostomafemalesprefer males withaterritory in
whichtheriver flowrateenablestheireggs havehighhatching success (GIBBONS
& PAIN. 1992,SIVA-JOTHY etal., 1995).
Ouraimwasto determinetheeffectofbody size(which variesconsiderably be-
tween individuals) on territorialbehaviourin C. virgo males(PAJUNEN, 1966;
RANTALA etal., 2001; KOSKIMAKI etal., 2004). They havehigh survival
rates. Thedailysurvival probabilityofanadultmaleisabout86%,whichmeans
thattheirexpected longevity is6-19 days,withthemaximumlife-spanbeingover
40 days (CORDOBA-AGUILAR & CORDERO-RIVERA,2005). Mostof C.
virgo malesdefendriparian territories,which includepatchesoffloating vegeta-
tion, used by femalesas oviposition sites (PAJUNEN, 1966; KOSKIMAKI et
ah,2004;TYNK.K.YNEN etah, 2006). In this study, we compared wing length
andfreshbody weight measurements ofterritorial andwandering males during
early andadvancedflyingseason. Thus,wetestedwhethertherewereany differ-
ences inthe body sizebetweenterritorialandnon-territorialmalesandwhether
therewas any seasonalvariationinthisrespect.
METHODS
Thestudy wasconductedontheMustajokicreek,closetothecityofJyvaskyla (62°I6'N,25°30’E)
in central Finland. The creek isca400m long,has agravelly/sandy bottomand abundant aquatic
vegetation.
InordertofacilitatetherecordingoftheCalopteryxpositions,fiber-tapeswereputoutonthetrees
at5mintervals.Several dayspriortotheactualstudy, allC, virgomales werecollected,theirleft and
right hindwingsmeasured (0.01 mm)with a digitalcalliper,dragonfliesweremarked with asilver
marking pen,and subsequentlyreleased back into thehabitat (RANTALAetal.,2001).The same
Sizein Calopleryxvirgowanderingandterritorial males 161
procedurewasfollowed with thenewmalesthatturnedup inthestudy sectionduringthestudy.
Onthe 18thand28th ofJuly1998,thebehaviourandthepositions ofmaleswererecorded hourly.
BasedontheobservationsofPAJUNEN(1966),WAAGE(1973),FORSYTH&MONTGOMERIE
(1987),TYNKKYNEN etal.(2006)males thatdefendedaterritorywithin adistancenotexceeding
2m and duringatleast 3consecutive hours wereconsidered territorial. Sinceterritorial males are
knowntoperchclosetothewatersurface(PAJUNEN, 1966),amalewasacceptedasterritorial only
ifitwasalways observed within 50cm ofthewatersurface. Males that were notdefendingaterri-
toryduringthedays ofourstudyweredesignatedasnon-territorial. Thenon-territorial males that
movedatleast 100m duringadaywereclassified aswanderers.
Inthisstudy, onlymaturemales,with unflexible wingedge,wereconsidered. Inall,wecollected
11wanderers and28territorial males. Theywereplacedindividuallyinacooler,transferredtothe
laboratorywhere theirwetbody mass wasmeasured with anaccuracy of0,1 mg,whereuponthey
werereturned back tothestream.
Wetestedproportionofwanderer males onstudy dayswith chisquaretest,and analyzedthedif-
ferencesinmeanwinglengthsand bodymassofterritorial andwanderer males withtwo-wayANO-
VAinwhichthemalestatus(territorialorwanderer)andstudy days(18 and28July)arefactors.All
theanalyseswereperformedwith SPSSforWindows(Version14.0).
RESULTS
On the18thofJuly thereweresevenwanderersoutofatotalof 18males(39%)
andonthe28lhofJulyfourwanderersoutofatotalof21 males(19%).Therewas
nosignificant differencebetweenthetwo daysinproportion ofwanderermalesin
theMustajoki creek(x2= 1.88,df=\,p =0.17).Themeanwinglengths(Two-way
ANOVA,F{ =0.10,/?=0.76)andwetbodymass(Two-way ANOVA,F] =2.02,
M 35
p=0.16)didnotdiffer
between days. Wan-
derers were smaller
thanterritorialmales
as estimatedby their
wing length (F, =
34
15.35,< 0.001) and
wet body mass (F,
35
= 15.25, p < 0.001,
Fig. 1).Themeanwet
bodymassofwander-
ers was 99 mg(SD -
11 mg)andofterrito-
rialmaleswas 112mg
(SD = 9.4 mg). The
mean length of left
hindwing ofterritori-
Fig. 1.Winglengthand wetweightofwanderers (opensymbols)and
almaleswas 30.9mm
territorialmales(blacksymbols).Squaresindicatethefirstdayofcap-
(SD = 1.0mm) while
ture(18July)and dotsindicate theseconddayofcapture(28July)at
thatofwandererswas theMustajokicreek.
162 J.Koskimaki, M.J.Rantala&J.Suhonen
29.4mm(SD = 1.0mm).Theinteractionbetweenthesetwomaineffectsdidnot
accountforsignificant variationinwinglength (F, =0.03,p=0.86)orwetbody
J4
mass (F,J5 - 0.56, p= 0.45).
DISCUSSION
Wefoundthatterritorial C. virgomalesare largerthan wandering males;they
havelongerhindwingsandareheavierthanwanderers.However,several previous
studiesin various Calopteryx species have shownthat territorialand non-terri-
torialbehavioursare conditionalmating tactics(MARDEN & WAAGE, 1990;
PLAISTOW&S1VA-JOTHY, 1996;KOSKIMAKI etah,2004;CONTRERAS-
-GARDUNO et ah, 2006). For example, C. virgomales thatwere winners of
stagedterritorialcontests hadmorefatthan losersbuttherewas no differencein
body size (KOSKIMAKIetal., 2004). Our datacontradictprevious studiesin
calopterygids inwhichtherewas nosize differencebetween territorialand non-
territorialmales(FORSYTH & MONTGOMERIE, 1987;CORDOBA-AGUI-
LAR, 1995; PLAISTOW& SIVA-JOTHY, 1996; BECK & PRUETT-JONES,
2002;TYNKKYNENetal.,2006;SUHONEN etal.,2008). However, in C.mac-
ulata,largermalesareableto holdterritoriesforlonger timethansmallermales
(BECK&PRUETT-JONES,2002),similarlyas inthepseudostigmatid, Megalo-
prepuscoerulatus(FINCKE, 1984).Moreover,recentworks onHetearinaameri-
canahavealso shownthatterritorial malesare larger thannon-territorialmales,
withlarge territorialmaleshaving highenergyreserves andacapabilityto hold
territorieslongerthansmaller, non-territorialmales(SERRANO-MENESES et
ah,2007;RA1HANI etah, 2008).
Mating success is likely tobe higher interritorialmalesthan innon-territorial
males(PLAISTOW&SIVA-JOTHY, 1996;SERRANO-MENESESetah,2007;
SUHONEN etah,2008), becausefemalespreferterritorialmales(WAAGE, 1973;
PLAISTOW&SIVA-JOTHY,1996;CORDOBA-AGUILAR,2000;SERRANO-
-MENESES etah, 2007). However, non-territorialmales are still likely to have
mating opportunities. For example, non-territorialC. virgoand C. maculatecan
sometimesclasp femaleswithoutcourtship (PAJUNEN, 1966;WAAGE, 1973)
but only 20% of non-courtedclasped females allowed copulation (WAAGE,
1973).Inarecent studyonHetaerinaamericana,thematingsuccess ofterritorial
males, switchermales(thosethatdemonstratebothterritorialandnon-territorial
behaviour) and non-territorialmalesdidnot differsignificantly (RAIHANI et
ah,2008). Itis possible thatwanderersandterritorialmalesmaybetwo morphs
withdifferentlife-historiesbutequal estimatedreproductive success as hasbeen
shown inthe Mnaisdamselflies(WATANABE & TAGUCHI, 1990;WATAN-
ABE, 1991;NOMAKUCHI, 1992;TSUBAK1etah, 1997).For example, male
M. costalisoccur asterritorialorange-winged “fighter”males ornon-territorial
clear-winged “sneaker”males,and thetwomorphs haveequal reproductive sue-
Sizein Calopleryxvirgowanderingandterritorial males 163
cess (TSUBAKI et al„ 1997; PLAISTOW& TSUBAKI, 2000). In M. costalis,
orange-winged fightermalesare largerthanclear-winged sneakermales(TSUB-
AKIetah, 1997).Studiesonthisspecies haveshownthattheclear-winged, non-
territorialmorph liveslonger(TSUBAKI etah, 1997; TSUBAKI& HOOPER,
2004). Femalesinthissystem matepreferentially withterritorialmalesbutsince
non-territorialmaleshavea larger sperm displacement ability, they are ableto
fertilisemoreeggs asthey can transfermorespermto thefemale(SIVA-JOTHY
& TSUBAKI, 1989a).Theestimatesoflifetimereproductive success indicatethat
bothmorphs achieveasimilarfertilizationsuccess, whichexplains thecoexistance
ofthe twomorphs inthe samepopulation (TSUBAKI etah, 1997).
Inconclusion, ourdatashowthatwandererC. virgomalesaresmallerthanter-
ritorial males. However, inour study only individuals fromasingle population
wereexamined.Inadditionto thebody size ofwanderersandterritorialmales,
future studiesshouldalso measure thebodysizeofsneakersandpossible switch-
ers (RAIHANI etal.,2008) andshouldinvestigate whethersimilar patterns are
apparentin other C. virgo populations andin otherspecies. Measuring thelife
timereproductive success inthe differentbehaviouralphenotypes will also bea
fruitfulavenue ofinvestigation.
ACKNOWLEDGEMENTS
We thank DEREKW.DUNN,KEN FEDORKA,KALEVI SALONEN andKATJATYNK-
KYNEN forvaluable commentsonanearlier versionofthemanuscript.Specialthanksaredue to
SEPPOKUUKASJARVI forassistance inthefield.Thelanguagewaschecked by DerekW.Dunn.
Thestudywasfundedby theFinnish Biodiversity Research Programmeand the AcademyofFin-
landundertheFinnish CentreofExcellence ProgrammetoJukkaSuhonen,AcademyofFinlandto
JukkaSuhonenandMarkusJ.Rantala,byEllen andArtturiNyyssonenFoundation andbySocietas
proFaunaetFloraFennicato JaniKoskimaki.
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