Male behaviour in the male dimorphic damselfly Paraphlebia quinta Calvert (Zygoptera: Megapodagrionidae)

2005·1.1 MB·English

by  González-Soriano, E

Odonatologica34(4):379-385 December 1,2005
Malebehaviourinthemaledimorphic
damselflyParaphlebiaquinta Calvert
(Zygoptera: Megapodagrionidae)
E.González-Soriano¹ andA.Córdoba-Aguilar²
1DepartamentodeZoologia,InstitutodeBiologia,Universidad Nacional Autonoma deMexico,
Apdo.Postal70-153,MX-04510 Mexico, D.E,Mexico
[email protected]
2Instituto deEcologia,Universidad Nacional Autonoma deMexico,Apdo.Postal70-275,
Circuitoexterior,Ciudad Universitaria,MX-04510 Mexico,D.E,Mexico
[email protected]
ReceivedMarch 19,2004/RevisedandAcceptedMarch8, 2005
P.quintaisatropicalsp.with 23morphs:theblack-winged(BW) 3and thehya-
line-winged(HW) 3;here theirsexual behaviour isdescribed.Ingeneral,63seem
tospendrelatively little timein flyingactivities.This maybeexplainedeitherby the
inabilitytorecogniseconspecificsand,hence,engagein socialinteractions,orbythe
reduced energeticreservesthatpreventthem fromengaginginexpensive activities.
BW<J3weremoreaggressive and site-faithful than HW <J6.BW defended spaces
containingdebris(plantandwood)againstconspecificswhile HWdid not.BW-BW,
BW-HWand HW-HWaggressiveencounters werecommon.Despitetheirnon-ag-
gressivenaturetowardBW <J<J, HW 33behaved aggressively when facedby HW
3 3.Thedistanceflown byeachmorphfrom 3 graspingofthe ? untilshe started
ovipositionwasmeasured:HWflewlongerdistancesthanBW.Thesedifferencesbe-
tween3 morphsarecomparedtothose found in Mnaisp.pruinosa,another 3 di-
morphiczygopteran.Similartowhathappensinthatsp.,bothtacticsinP. quintaare
possiblymaintained duetothe similar reproductiveand energeticcosts accrued by
andbenefitspaidtoeachmorph.
INTRODUCTION
Maleterritoriality representsan ubiquitous phenomenon inmany dragonfly
taxa(reviewed by CORBET, 1999).IntheZygoptera, territorialityis particularly
prevalent amongseveraltaxaofthesuperfamily Calopterygoidea, such asinthe
generaCalopteryx (e.g. WAAGE, 1973), Cora (e.g. GONZALEZ-SORIANO
& VERDUGO-GARZA, 1984; FRASER & HERMAN, 1993), Heliocypha
380 E.Gonzalez-Soriano &A.Cordoba-Aguilar
(ORR, 1996),Hetaerina(JOHNSON, 1962),Libellago(ORR, 1996),Mnais(e.g.
HIGASHI, 1981), Platycypha (ROBERTSON, 1982), and Rhynocypha (ORR,
1996). Inotherzygopteran families, territorialbehaviouris a commonfeature
amongsomegeneraofMegapodagrionidae, such asHeteragrion (GONZALEZ-
-SORIANO & VERDUGO-GARZA, 1982) and Paraphlebia (GONZALEZ-
-SOR1ANO & CORDOBA-AGUILAR, 2003); inProtoneuridae,such as Chlo-
roneura (SRIVASTAVA & BABU, 1985) andNososticta(THOMPSON, 1990);
in Platystictidae, such as Palaemnema(GONZALEZ-SORIANOet al., 1982);
in Pseudostigmatidae, such as Megaloprepus (FINCKE, 1984);in Platycnemidi-
dae,such as Platycnemis (HEYMER, 1966);andinsomeCoenagrionidae, such
as Megalagrion (MOORE, 1983)andPseudagrion (MESKIN, 1986).
Malesofthe orientalgenusMnaisare dimorphic. One morph bearspigment-
ed wings and shows territorial behaviour, while the other morph has hyaline
wings andbehaves assubordinate, non-territorialindividuals(WATANABE &
TAGUCHI, 1990;SIVA-JOTHY & TSUBAKI, 1989a, 1989b). Thisphenome-
nonisnotconfinedtothistaxon. Malesinsomespecies oftheNewWorldgenus
Paraphlebia, such as P. quintaand P. zoe,arealsodimorphic(GONZALEZ-SO-
RIANO, 1997;GONZALEZ-SORIANO &CORDOBA-AGUILAR,2003). In
P. quinta,forexample, one morph is theblack-winged (BW)malewhiletheother
is thehyaline-winged(HW) male.BWmalesare usually largerindividuals, bear-
ing ablacktransverse band onthewing tips, whileHW males are smaller, with
entirely clear wings, resembling thoseof females(GONZALEZ-SORIANO &
CORDOBA-AGUILAR,2003). How thetwo morphs are maintainedis uncer-
tain.InMnais, themaintenanceofbothmorphs ispossibly dueto thebalanced
reproductive benefitsandcostsofeachphenotype (SIVA-JOTHY &TSUBAKI,
1989a;PLAISTOW& TSUBAKI,2000).
Herewepresentasummaryofthebehaviouralcomponentsofthemalesexual
activitiesofP. quinta. Our firstaimis to provide a descriptiveframeworkofthe
behaviourofeach morph.Additionally, wepresent apreliminary analysisofthe
cost ofaccompanying afemaleby each morph, whichis discussedintheframe-
workofthe possible fertilisationsuccess achievedby bothtypesofmales.
MATERIALAND METHODS
Field work wascarriedoutatasmallseepage, dividedinto36 quadrants(2x2m),attheEstacion
deBiologiaTropical Los Tuxtlas in the state ofVeracruz, Mexico (95°04-95°09’Wand 18°34’-
18°36’N).Mostobservations wereperformedon23-26August 1984 and 19 July-15August 1986. In
bothyears, malesand females wereindividuallymarked ontheirwings, usinganindelible inkpen,
allocatingdistinctive numbers. Inboth years, observations of 30-60 min duration offocal males
werecarriedout.In 1986,dailycensusesweremadeandthepositionofeachmarkedindividual was
located onascalemap. Bythisprocedure,dailymovementsofeachindividual weremeasured and
trackedwith relativeaccuracy.
On those datesand alsoon8-11 July 1988 and 19-23 August 1990,werecorded the trajectories
followed byeach morphfromthemalegraspingofthefemaleprothorax(attheonset ofcopulation)
Malebehaviour inParaphlebiaquinta 381
tothebeginningofthefirstoviposition bout.
For theuseofparametric tests,wetransformed thosedata that wereamenable forsuchmanipu-
lation. Mann Whitneytestsarereferred toasUtests. Resultsarepresentedasmeans± STDunless
indicated otherwise.
RESULTS
Dailycensuses were carriedout during 18consecutive days(19 July-5 August
1986).Thenumberofindividualsofbothmalemorphs founddailyfluctuatedbe-
tween33and41individuals(36.5 ± 2.59). However, malesremainedinthestudy
sites for variableperiods (in days;BWmales, 11.5 ± 6.5, N =35; HW males, 4.8
± 4.8, N = 45).For BW males, theminimumnumberof days anindividualwas
foundinthesame site(during amaximumof a 17-day period) was 1,whilethe
maximum(butnot necessarily consecutive days)was 17(N =35). Morethan50%
ofBWmaleswerefoundin thesamesitefor16-17days,andonly 14%were seen
onthe dayofmarking. For HW males,theminimumof dayspresentinthesame
sitewas 1,whilethe maximumwas 16(N =45). Only4.6%oftheHWmalesre-
mainedbetween 15-16days,65% were concentratedwithina greaterintervalof
1-15days,andalargenumber(almost30%)werepresentonly onthedayofmark-
ing. ThesedifferencesindicatethatBWmalesstayedforlongerperiodsthanHW
malesintheplaces ofmarking(Utest= 1878.5, P<0.0001). Asforfemales, they
werefound fewerdays (0.5 ± 0.8)compared to BW (U test= 3019, P < 0.0001)
andHW males(Utest=3653.3,P<0.0001).Theminimumnumberofdaysthey
returnedto thegeneral areawas 1 day, whilethemaximumwas 4.
Thefourmainflying activities, performed by the twomalemorphs at the re-
productive site were;changes ofperching site, aggressive flights(mainly directed
atsame-sexconspecifics), investigatory flights(short-duration flightsofafewcm
withnoclearexplanation), and patrolling flights(in whichtheanimalcovers the
areaarounditsperching site)(Tab. I). Flying activity wasfairlylowincompari-
son withtotaltime inactivity(0.87% forBW and0.38%forHW). Theonly sig-
nificantdifferenceinflyingactivitiesbetweenmorphs wasintheaggressive flight:
Table I
Accountofthedifferentflyingactivitiesatthe reproductivesite
byP.quintablack-winged(BW)andhyaline-winged(HW)males
Perching site Aggressive Investigatory Patrolling
change flight flight flight
BW HW BW HW BW HW BW HW
Occurrence 20 17 15 1 55 20 20 1
Meanduration 2.0 2.2 4.3 5 1.6 1.2 0.8 0.4
%occurrenceinrelation to all flights 18.2 43.5 13.6 2.6 50 51.2 18.2 2.6
%durationin relation toall flights 22.9 25 49.4 56.8 18.3 13.6 9.3 4.5
MannWhitneyUtest (P) 57.5 (>0.05) 69.5 (<0.05) 66.0 (>0.05) 66.0(>0.05)
382 E.Gonzalez-Soriano &A.Cordoba-Aguilar
BW males made more of
theseflightsthan HW males
(U test = 69.5, P < 0.05).
No further differenceswere
foundinchanges of perch-
ing site (U test = 57.5, P >
0.05)orinvestigatoryandpa-
trolling flights (both U test
= 66.0,P> 0.05).Inperched
males,threemainbehaviours
wereobserved:wingclapping
(BW: 14.2± 12.8;HW; 12.60
± 6.8), wing warning (BW:
0.2±0.5; HW;0.4±0.9)and
abdominalcleaning(BW:6.9
± 9.2; HW; 4.2± 3.5).Sam-
ple size forallcomparisons:
Fig. 1.ExamplesofdistancesflownbyeachP. quintamorph
whilewith afemale(measuredfromthe onsetofcopulation N =8forBWmalesand5for
tothefirstovipositionbout):(a-b)twoBWmales;1(c-d)two HW males. Themorphs dif-
F1Wmales; — ®onsetofflight;—xfirstovipositionbout;— feredinnoneofthesebehav-
dotsrepresentquadrats(2x2m). iours(wingclapping,Utest=
57.0; wingwarning, Utest=56.0;abdominalcleaning,U test=53.0;allP>0.05).
Agonistic interactionsoccurredbothbetween(BW-HW)andwithin(BW-BW
and HW-HW) morphs. Most inter-morph interactionsconsisted basically ofa
chasing flightofHW malesby BW malesinwhich invariably theHW male left
theterritory(25 outof25 cases). Inthefewoccasions thatescalatedflights were
observed, they tookplace mainlybetweenneighbouring BW males. Thesecon-
testsconsistedofafrontal,slowly-ascending interaction.Inmostcases,theowner
remainedin theterritory(3 outof 5 times). As forHW-HW encounters,the in-
truderwaschasedandexcluded fromtheterritory(5 outof6 times). Inthis type
ofencounters,physical contact was observed.
Interactionswerenotonlyintraspecific. P. quintamalesalsochasedotherdam-
selflies, especially Palaemnemapaulitaba and Hetaerinasempronia. BW males
had9 interactions withP.paulitabaandonewithH.sempronia, whileHWmales
had 12withP.paulitaba andone withH. sempronia. P.paulitabahas clearwings
withblack tips, H. sempronia clearwings withred bases and smallbrownspots
atthe hindwingtips.
TRAJECTORIESOFPAIRS. — Therewas adifferenceinrelationto malemorph
in thedistance travelledby thepair to thetimeatwhichthe femalefirst ovipos-
ited: HWmalesflewalongerdistance(N=4; 10.65±7.1 m)thanBWmalesdid
(N =5; 1.28 ± 1.1 m)(t on log transformeddata=-3.36, P= 0.015). Someex-
7
amples ofthis differenceare shownin Figure 1.
Malebehaviour inParaphlehiaquinta 383
DISCUSSION
P. quinta malemorphs exhibit clearbehaviouraldifferences, whichpresuma-
blyrelateto theirdifferentmate-seeking tactics. TheBW morph ismoreaggres-
sive (reflected in the numberof aggressive flights andtheir chasing HW males
butnot being chased by them)and site-attachedthanthe HW morph. An in-
teresting convergence to this dimorphism is thesituation inthe Japanese Mnais
(SIVA-JOTHY &TSUBAK1,1989a, 1989b;WATANABE& TAGUCHI,1990;
PLAISTOW&TSUBAKI,2000), Wheretheterritorialmorph isalsoaggressive,
whilethenon-territorialmorph seems toavoidagonistic encounters. Addition-
ally, theterritorialmorph of M. pruinosa requires moremass than non-territo-
rialmales, anindicationthatthe territorial tactic is morecostly. Our results in-
dicatethattheBW malesalsoseemto incurhighercosts, as they haveto defend
aterritory. Thiscost maybebalanced bythe reproductive benefits. In P. quinta,
copulations by the non-territorialmorph are fewer, they are more frequently
genitalicallyinterrupted and, during these interruptions,they are likely toresult
in moresperm-ejection events by females (GONZALEZ-SORIANO & COR-
DOBA-AGUILAR,2003). Giventheseresults, someinteresting avenues forre-
search in thesespecies are:(a)toelucidatethegeneticbasisofmaledimorphism
(to determinetheenvironmentalroleingiving risetoeachmorph); — and(b) to
measurethelong-termreproductive success ofeach morph.Theseaimswillshed
lightonhowboth morphs are undergoing selection.
BEHAVIOURAL REPERTOIREOFP.QUINTA
Territorialbehaviourin P. quinta shows two unusual featuresin comparison
withotherterritorialzygopterans,viz. malesarerelativelyinactive,whilethenon-
territorialmorph isfairlyaggressive towardnon-territorialconspecifics.
BothP quintamorphs showedextremely reducedflightactivity: nomorethan
0.9%oftotaltime.Incomparison, malesofthecoenagrionid Pseudagrion hage-
niinvest 6.7% oftheir territorialoccupation in flight (MESKIN, 1986), while
HIGASHI (1981) pointed out that “territorialmales of M. pruinosa seemed
to havealmost notimeperching on theirperch sites for pursuing theintruding
males”. Also unusual for otherterritorial zygopterans butconcurrent with the
above:P quinta engaged inrelatively fewmale-maleinteractions, withmost of
themlasting only afewseconds(foracomparison seeWAAGE, 1988). Atleast
two explanations canbeputforwardforthis;(a) thereis notmuch lightintheir
habitats, whichpossiblyaffects visualrecognition ofconspecifics; — and/or (b)
malesmaypossess areducedamountofenergetic reserves which preventsthem
frommovingtoomuch.
Itisalsointerestingtonoticethatalthough HW malesalwaysshowedasubmis-
sivebehaviourin thepresenceofa BW male, they behavedaggressively against
384 E.Gonzalez-Soriano &A.Cordoba-Aguilar
malesoftheirownform.Possibly, thesubmissivebehaviouronlyemergesinthe
faceofaBWmale. Actually, according to theunpublished observationsonare-
latedspecies, P. zoe, attheendoftheflightseason,whennoBWmalesarepresent,
HW malesaggressively defendterritories(A. Romo-Beltran, pers.comm.) This
indicatesthatthe submissivebehaviourof HW males occurs only towards BW
males. Perhaps the valueofa territoryis “appreciated” by HW males, butthey
do notdispute itwith BWmalesbecausethey are notableto undertakeacostly
fight (forasimilarrationaleseePLAISTOW& TSUBAKI,2000). Thiscost can
beaffordedonly against HW males.
Pairs formed by HW males flew longer thanthoseformedby BW malesdur-
ing andaftercopulation. Thisexpenditure isnot meaningless. Thedelay incop-
ulationand/oroviposition couldaffectfitnessofbothpartners in at leastthree
non-exclusiveways:(1) agreaterrisk of predation; — (2) anincreasednumber
ofattacks fromconspecific males whichmay cause tandemdisruption; - and
(3) areduced dispositionto oviposit byfemales.Wehaveindirectevidenceof the
lasttwo. Although not measured, during theincreasedHW male tandemdura-
tion, thecouple was morefrequentlyapproached by conspecific maleswhichoc-
casionally induced tandembreakage. Theincreased attack may be“perceived”
by the femaleandaffects her disposition to oviposit. This mayexplain why fe-
malesspend lesstimein oviposition aftermating withan HW malethanwitha
BWmale(GONZALEZ-SOR1ANO &CORDOBA-AGUILAR.2003), though
thereisalso the possibility that HW malesmaynot be good mate guarders, as
showninsomeotherspecies. Thereducedoviposition timemaybeactuallytrans-
latedto the numberof eggslaid. In M. pruinosa there is a direct relationship
betweenthenumberofeggslaidand timespentin oviposition (WATANABE
& TAGUCHI, 1990). Inodonates, the lastmaleinseminatesusually mosteggs
(CORDOBA-AGUILAR etal., 2003). Ifthese relationships holdforP. quinta,
thenHW malesmay seetheirfertilisation success reduced as a consequenceof
reducedoviposition bouts.
ACKNOWLEDGEMENTS
EG-SthanksROLANDOMENDOZA,JOSELUISVILLAREAL,and LIZBETHHERAS for
helpduringthefieldwork.
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