Table Of Content©EntomologicaFennica.11October2006
Agonistic behaviour of Scarites buparius (Forster, 1771)
(Coleoptera: Carabidae) in relation to body size
TeresaBonacci,PietroBrandmayr,AnitaGiglio,AlessandroMassolo,AntonioMazzei,
RosalbaOdoguardi,MariastellaRomeo,FedericaF.Talarico&TulliaZettoBrandmayr*
Bonacci,T.,Brandmayr,P.,Giglio,A.,Massolo,A.,Mazzei,A.,Odoguardi,R.,
Romeo,M.,Talarico,F.F.&Brandmayr,T.Z.2006:AgonisticbehaviourofSca-
ritesbuparius(Forster,1771)(Coleoptera:Carabidae)inrelationtobodysize.—
Entomol.Fennica17:340–344.
Intra-maleaggressivebehaviourofScaritesbupariuswasanalysed;agonisticin-
teractionbetweenmalesconsistedofarepeatedseriesoffightingevents.Wede-
finedthisbehaviouras“agonistic”becauseadominance/submissionstatuswas
established.Wemeasuredthemalesandfoundthattheattackbehaviourpersis-
tenceiscorrelatedwiththebodylength.
T.Bonacci,P.Brandmayr,A.Giglio,A.Mazzei,R.Odoguardi,M.Romeo,F.F.
TalaricoandT.Z.Brandmayr,DipartimentodiEcologia,UniversitàdegliStudi
della Calabria, 87036 Rende (CS), Italy; *correspondent author’s e-mail:
[email protected]
A.Massolo,DipartimentodiBiologiaAnimaleeGenetica,UniversitàdiFirenze,
viaRomana,17,I-50125Firenze,Italy
Received12December2005,accepted22May2006
1.Introduction Agonisticbehaviourisaninherentcapacityof
mostanimals.Forinvertebratesithasbeenstud-
In the sense of King (1973), “agonistic behav- iedinspiders(Austad1982,1983,Leimaretal.
iour”includesallbehavioursassociatedwiththe 1991), cuttlefish and squid (Adamo & Hanlon
strugglebetweenindividualsofthesamespecies. 1996,Kingetal.2003),scorpionflies(Thornill
Themotororactionpatternsinthepredator-prey 1984) and most exhaustively in crustaceans
encounterorininter-specificcompetitionareof- (Hazlett 1981, 1999, Bruski & Dunham 1987,
ten the same as those exhibited between con- Copp 1986, Daws et al. 2002, Gherardi & Pie-
specificrivals.Thebehaviourissimilar,butthe raccini2004,Goessmannetal.2000,Guiasu&
functionoftheagonisticbehaviourchangesfrom Dunham1997,Karavanich&Atema1998).Usu-
inter-specificcompetitionandpredator-preyrela- ally,agonisticbehaviouristriggeredbythecom-
tionshipstointra-specificsociality.Usually,los- petitionforresources(reproduction,feeding,ter-
ers in the agonistic encounter are variously op- ritory ormany amongthem)anditseemstoin-
posed,dependingonthespecies;converselywin- crease with the population density (Hazlett
nersgainsocialstatusoraccesstoresources.The 1968), and the dominants might hypothetically
evolution of restraints to high aggressiveness occupymostofthenichespaceifresourcesarein
may buffer a strong negative outcome for the lowdensity(Levins1968).Morse(1974)found
loser. that the dominant individuals have larger body
ENTOMOL.FENNICAVol.17 (cid:127) Intra-specificaggressivenessinScaritesbuparius 341
sizethansubordinatesinmosthierarchies.Inin- Kiel,Germany).Dataanalysis(frequencyandse-
sects,sizeisanimportantcharacteristicdetermin- quence of events) was performed with the soft-
ing reproductive success in males (McLain wareTheObserver.
1982).Insoldierbeetleshighersuccessinobtain-
ing mates is positively correlated with antennal
scape diameter (Mason 1980). Positively varia- 2.2Morphometricanalysis
tioninmaledryweightanddiameterofantennae
andpalpiinMeloidae(Coleoptera)maycausean The sample consisted of 16 individuals of
increasing of the courtship duration (McLain Scaritesbuparius.Magnifiedimagesweretaken
1982). by mean of a stereoscope (Zeiss Stemi SV
AgonisticbehaviourinColeopteranshasbeen 11Apo),andimagecapturewasperformedusing
poorlyinvestigated;particularlyforcarabidbee- thesoftwareMatroxPC-VCR(Windows2000).
tles. S. buparius is one of the first species for For each individual, we measured body length
whichintramalefightinghasbeenreportedboth (mm),mandiblelength(mm),headlength(mm),
in the field and in the laboratory (Alicata et al. head width (mm), pronotum length (mm),
1980). Recently, Mossakowski (2003) studied pronotum width (mm) and elytral length (mm).
the fighting behaviour of Broscus cephalotes MeasurementsweretakenusingSigmaScanPro
(L.). In this paper we describe the intramale 5software(SPSS®Inc.).
agonisticbehaviouranditsrepertoireofinterac-
tions.
2.3.Statisticalanalysis
The relationship between morphometric mea-
2.Materialandmethods surements and agonistic behaviour events was
analysed by logistic regression (Hosmer & Le-
Sixteen specimens (4 females and 12 males) of meshow1989).
S.buparius,collectedinsanddunehabitatsonthe WeusedtheSpearmancorrelationcoefficient
Ionianseacoast,Squillace,Italy,wereseparately totestforassociationbetweenthedurationofag-
kept in glass containers (diameter 5.5 cm, 8 cm gressive behaviour and body size (Siegel &
tall),with2cmofsandysoil,inaclimaticcham- Castellan 1998), and then we used nonlinear
beratday/nightT24°C/18°C,underL/D:12/12 curveestimationtocomparethefittingofdiffer-
photoperiod and fed on Theba pisana (Gastro- entequations(Linear,power,exponential,loga-
poda)collectedinthesamehabitats. rithmic,inverse)(SPSS®Base12.0User’sGuide
2003).
All statistical analyses were performed with
2.1.Behaviouralanalysis theStatistical Packagefor Social Sciences, ver-
sion12.01(SPSS®).
EachdyadofS.bupariuswastestedinthelabora-
tory(35recordingsessionsintotal)for30min-
utes.Animalswerefairly fedbeforetheexperi-
ment.Ineachtestonespecimenwasmarkedwith 3.Results
non-toxic paint. The experimental arena was a
glasscontainerwithaplastersubstrate(8.5cm× After the introduction into the experimental
5.5 cm); the test started when both specimens arena,thebeetlesexhibitedaggressivebehaviour
wereplacedinside. with a typical repertoire of eleven interactions
Thebehaviourwasrecordedbydigitalvideo (events).
equipment (Sony digital video camera and a
DVD recorder) and behavioural analyses were (1) Latencyofattack:thetimepassingfromthe
carried out with software for behavioural re- introduction into the arena to the attack of
search(TheObserver®version2.0,Universityof one of the two rivals (Frequency of occur-
342 Bonaccietal. (cid:127) ENTOMOL.FENNICAVol.17
Fig.1.Agonisticrepertoirein
Scaritesbuparius(observedbe-
haviouralevents).–a.Mandible
grip:oneindividualgraspsthe
otherbyitsmandibles.Thisgrip
involvesthecontactofanten-
nae.–b.Head-pronotumgrip:
oneindividualgraspstheother
bytheneck(left)orbythe
pronotum(right).–c.Pronotum-
abdomengrip:oneindividual
graspstheotherbythe
mesonotum.Thesubordinatein-
dividualmayberaisedhead-up
(left)orhead-down(right).–d.
Defensivegrip:thesubordinate
individualgraspswithmandibles
theotheronthelegorbythe
mandibles.
rence(F)=34;x=293.34±370.67sec). behaviourwascarriedoutbymeansofthree
o
(2) Attack:oneindividualadvancestowardsthe items (mandible grip, head/pronotum grip,
other with opened mandibles gripping the pronotum/abdomengrip).
opponent’sbody (F =648;x=8.24±8.31 (3) Mandible grip: one individual grasps the
o
sec). The event “grip” during the agonistic other by its mandibles. This behavioural
ENTOMOL.FENNICAVol.17 (cid:127) Intra-specificaggressivenessinScaritesbuparius 343
eventinvolvesthecontactofantennae(F =
o
142;x=21.92±72.48)(Fig.1a).
(4) Head-pronotum grip: most frequently the
dominant grasps the subordinate individual
by thehead/pronotumarticulation (a) or by
thepronotum(b)(F =149;x=21.71±26.69)
o
(Fig.1b).
(5) Pronotum – abdomen grip: one individual
graspstheotherbythemesonotum.Thesub-
ordinateindividualmayberaisedhead-up(a)
or head-down (b); (F = 133; x = 36.05 ±
o
98.62sec)(Fig.1c).
(6) Grip-raising:oneindividualgraspstheother
withthemandiblesandraisesitfromthesub-
strate(F =256;x=26.26±16.52sec).
o
(7) Defensive grip: the subordinate individual
graspswithmandiblestheotheronthelegor
onthemandibles(F =82;x=5.65±11.75 Fig.2.Relationshipbetweenmalebodylengthof
sec)(Fig.1d). o Scaritesbupariusanddurationofmaleaggressivebe-
(8) Skipping: this event was displayed only by haviourinthelaboratory.Blackdots=behaviouraltest
the dominant individual; it skips without events;curve=exponentialcurve.
movingfromitsposition(F =138;x=9.99±
o
24.03sec).
(9) Immobility: the subordinate during the grip tack”, one individual approaches the other with
remains motionless (F = 219; x = 42.97 ± openedmandiblestogriptheantagonist’sbody;
o
96.08sec). “mandiblegrip”,oneindividualgraspstheother
(10)Antennal contact: during the fight the indi- by themandibles,whilearapidantennationoc-
vidualsinteractviatheirantennae(F =150; cursbetweenthetwoindividuals;“grip”,onein-
o
x=24.55±36.67sec). dividual grasps the other by various parts of its
(11)Escapemovement:oneindividual,duringan body.
aggressivebehaviourepisode,escapes(F = Throughouttheagonisticbehaviour,S.bupa-
o
590;x=8.2±4.22sec). rius specimens establish the dominant/subordi-
nate status. Dominants show superior body
Our data showed a significant relation between strength; in particular, they have larger body
dominantandsubordinatespecimens.Pronotum length and a wider and longer pronotum. The
length(B=28.390;p=0.041),pronotumwidth dominant individual attacks and raises subordi-
(B=35.839;p=0.021)andtotalbodylength(B= natesmorefrequentlybutitneverinjurestherival
60.803; p = 0.027) influenced significantly the severely.Conversely,subordinatessuffertheat-
acquisitionofthedominance-status.Durationof tack without an aggressive reaction. It is likely
aggressive behaviour was positively correlated thatmaleaggressivebehaviourinS.bupariusin-
withtotalbodylength(r =0.589,p<0.001),con- volvesreproductivefitness,asvigorousmalesin-
s
firmedby asignificantexponentialfitting(R2= creasetheprobabilitytofertilizealargenumber
0.334,b =0.594,p<0.001;Fig.2). offemales.
1
Thebodysizeofarthropodsmayevendeter-
minetheresultofagonisticencountersbetween
4.Discussion males(Austad1983,Suter1990)and,asaconse-
quence,thefecundityoftheinteractingmales.
Inthispaperweanalysedthefightingbehaviour Significantcorrelationsbetweensizeofmale
ofS.buparius.Inthiscarabidbeetletheaggres- anditsreproductivesuccesswerealsofoundby
siveness involves some important phases: “at- McLain(1982)inEpicautapennsylvanica(Cole-
344 Bonaccietal. (cid:127) ENTOMOL.FENNICAVol.17
optera,Meloidae);largermalesizehasbeenasso- iouralEcologyandSociobiology48:418–428.
ciatedwithhighercompetitiveabilityorsuccess Guiasu,R.C.&Dunham,D.W.1997:Initiationandout-
comeofagonisticcontestsinmaleforminCambarus
inobtainingmatesinavarietyofbeetles(Brown
robustus Girard, 1852 crayfish (Decapoda,
1980,Otte&Joern1975).
Cambaridae).–Crustaceana70:720–736.
During the different phases of the agonistic Hazlett,B.A.1968:Effectsofcrowdingontheagonistic
behaviour, the beetles continuously interact via behaviourofthehermitcrabPagurusbernhardus.—
theirantennae.Ourhypothesisisthatthisbehav- Ecology49(3):573–575.
Hazlett, B. A. 1981: The behavioral ecology of hermit
iourmayplayaroleintheinhibitionofaggres-
crabs.—AnnualReviewofEcologyandSystematics
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12:1–22.
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abouttheconspecificrecognition. bythecrayfishOrconectesvirilis.—Behaviour136:
161–177.
Hosmer,D.W.&Lemeshow,S.1989:AppliedLogistic
Regression.—JohnWiley&Sons,NewYork,USA.
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