Mating Behavior of Caribbean Reef Squid, Sepioteuthis ...
Transcript of Mating Behavior of Caribbean Reef Squid, Sepioteuthis ...
Mating Behavior of
Caribbean Reef Squid, Sepioteuthis sepioidea
Jamie Morris June 6, 2010
Dominica2010Dr.HeymanDr.Lacher
2
Abstract:
Sepioteuthissepioidea,CaribbeanReefSquid,usechromatophoresand
iridiphorestochangetheircolorandpatterns.Thesechangesareusedaspartofa
complexcommunicationsystem.Thecomplexityofthismatingsystemhasbeen
mentionedinpreviousstudiesbutrarelyexplainedinmoredetail.Thispaper
attemptstoidentifyanddescribethecolorandpatternchangesseenineveryday
behavior,butmostimportantlyfocusesondescribingthecomplexcourting
behaviorandmale‐malecompetition.
Introduction
ThefocusofmystudywasSepioteuthissepioidea,CaribbeanReefSquid.Reef
squidareafavoriteamongsnorkelersandscubadiversbecausetheyarecommonly
foundamongshallowreefsandcandisplayavarietyorcolorsandpatternsthatcan
changeinlessthanasecond(Byrneetal.2003).Notonlyarethesecolorand
patternchangesfascinatingtoobserve,butalsotheyplayagreatrolein
reproductivebehavior.Reefsquidhaveaveryshortlifespanofabout1to2years,
andthemajorityoftheirmaturelivesarespentcompetingtomaximizetheir
reproductivesuccess(Hanlonetal.2002).Sincereefsquidaresemelparous;thatis,
theydieafterreproducing,theyarenotmonogamous.Malescanfertilizemany
femalesintheshorttimebeforetheydie,whilefemaleslaytheireggsanddie
immediatelyafter(MarineBio2008).Becauseofthedifficultyofstudyingreefsquid
bydirectunderwaterobservation,thereisnotmuchinformationontheircomplex
matingsystems.PreviousstudiesbyMoynihanandRodaniche(1982)and
Moynihan(1985)(ascitedbyHanlonetal.2002)offerbasicdescriptionsofthe
3
complexityofthematingsystemforS.sepioidea,butlittlesubsequentinformation
hasbeenpublished.Sincethesestudiesareabout25yearsold,Iwasinterestedto
seeifmyobservationsareconsistentwiththeseearlydescriptionsandifIcan
describeinmoredetailthecomponentsofthematingsystem.
MaterialsandMethods
Studyorganism
Reefsquidareamemberofthe10‐armcephalopodsandcanbeidentifiedby
theirtorpedoshapeandfinsthatspanalmosttheentirelengthoftheirbody.Allten
oftheirarmsareattachedtothehead,witheightshortarmsclosetothemouthand
twolargetentaclesforcapturingprey(MarineBio2008).Thecolorationand
patternchangeisduetopigmentedorganscalledchromatophoresandreflecting
cellscallediridiphores.Thechromatophorescellsareinthedermallayeroftheskin
anddisplayreds,yellows,brownandblackandupclosearesmallpatchesanddots.
Theiridiphoresproducethegreenandbluecolorations(Figure1)(MarineBiological
Laboratory2008).Thesecellsallowthesquidtocommunicateonemessagetothe
squidtotheirright,whiledisplayingacompletelydifferentmessagetothesquidto
theirleft(Byrneetal.2003).Reefsquidarecommonlyfoundinsmallschoolsof
about4‐30individualsandrangefromabout8‐20cminlength(MarineBio2008).
Studyarea
TheCaribbeanSeaisatropicalsealocatedadjacenttotheAtlanticOceanat
9‐22°Nand89‐60°W.TheCaribbeanSeaishometomanycoralreefs,whichallow
forgreatecologicaldiversity.DominicaispartoftheLesserAntillesbetweenthe
4
islandsofMartiniqueandGuadeloupe,andhelpsmakeuptheeasternboundaryof
theCaribbeanSea.This“natureisland”,hascoralreefssurroundingitmakingita
greatplacetodomystudy.Itookdataattwodifferentlocations,ChampagneBay
andScott’shead.BothlocationsarelocatedatthesouthwesternendofDominica
andareseparatedbyapproximately4.9km(Figure2).
StudyMaterials
TodomyobservationalstudyIusedsnorkelinggearconsistingofamask,
snorkel,fins,andawetsuit.Ialsotookstillphotosandvideoofthereefsquidwith
anOlympusStylistTough8010camera.
Figure1:FemaleCaribbeanreefsquidinbasicbrowncolorphaseshowingthedifferencebetweenchromatophoreandiridiphorecoloration.PhotographbyJamieMorris
5
Figure2:ThismapshowsthesouthwesternendofDominica,pointingoutthetwo
areas,ChampagneBayandScott’sHead,weremyobservationstookplace.Mapby
KinnieEijsink
Results
Reefsquidcanbefoundinabasicpale,basicintermediate,andbasicbrown
colorationinthewild.Thebasicbrownphaseistheonlyphasewhereitispossible
todifferentiatebetweensexes(Figure3).Thefemaleshaveagradualtransition
frombrowntopale,whilethemalesaresolidbrownwithdistinctfindots.Basic
paleisseenoftenatdawnorwhenapredatorswimsby(Figure4).Thiscoloration
6
consistsofapalebackground,withreflectivegreenontheeyebrowsandmantle
midline.Thebasicintermediateisthein‐betweenstageofpaleandbasicbrown
(Byrneetal.2003).
A)
B)
Figure3:A)AfemaleCaribbeanreefsquidinthebasicbrownphaseB)AmaleinthebasicbrownphasePhotographbyJamieMorris
7
A)
B)
Figure4:AandB)Caribbeanreefsquidintheirbasicpalephaseofcoloration.Photographsby:JamieMorrisandDr.Heyman
8
Reefsquidspendthemajorityofthedaycasuallyswimminginshoals
(groups)andchangingtheircolorperiodicallyforcamouflagepurposes.Theadult
reefsquidchangedtopaleorwouldblendinwiththeirenvironment,whilethe
juvenilereefsquiddisplayaplaidpatterntohidethemselves(Figure5).Theshoals
Iobservedwereverysmall,only2‐4squidpershoal.Interestingly,Iobservedthe
squidswimmingwithonlytheirownsexwhennotcourtingeachother.
CourtshipbehaviorwasobservedatChampagneBayon5/30/2010from
about2:15PM‐2:25PM.Courtshipbetweenmalesandfemalesinvolvedaslow
parallelswimmingtypeofdancebetweenamaleandfemalesquidwithsome
interruptionbycompetingmales.Theprimarymaleflickeredbrightlyusuallyon
thesidefacingawayfromthefemale,whilethesidetowardthefemaleshoweda
stripedisplay(Figure6aand6b).Themaleperiodicallyexhibitedafullbodyflicker.
Thefemaleoccasionally“answered”backwithapartialsaddledisplay(herbottom
Figure5:JuvenileCaribbeanreefsquidshowingtypicalplaidcolorationdisplay.Photographby:ArielSklar
9
endturnedlighterleavingtherestofthemantledark)(Figure7).Thecourtship
dancelastedabout1.5minutesbeforeanothermalecompetingforthefemale
interruptedthecourtshipbehavior.Themale‐malecompetitionquicklyheatedup
withdynamiczebradisplaysfrombothmales.Themalesroseupcompletely
verticalinthewatercolumntochallengeeachother(Figure8).Tointensifytheir
displaysthemalesspreadtheirtentaclesandspreadtheirfinsaroundthemantleto
makethemselveslooklarger(Figure9).
A)
Figure6:A)Amalereefsquidisshownhoveringoverandtotheleftofafemaleand
heisshowingapartialstripedisplayandapartialflicker.
B)Amaleshowsaclearflickeronthesideofhisbodythatisawayfromthefemale.
StillshotfromavideobyArielSklar
10
B)
Figure7:Amalesquidshowsafullbodyflicker,whilethefemale(below)responds
withasaddledisplay.StillshotextractedfromvideobyArielSklar
11
Figure9:Themaleexhibitselaboratezebrapatternaspartofanintensedisplayof
male‐malecompetition.Thespreadingoftheirtentaclesintensifiesthedisplay.Still
shotfromavideobyDr.Heyman
Discussion:
Figure8:Twomalesquidareswimverticallyinanelaboratedisplayofmale‐malecompetition.Notealsothespreadoftheirtentaclestointensifythedisplay.StillshotfromavideobyDr.Heyman
12
Myobservationsofthereefsquidcourtshipandnon‐courtshipdisplays
followverycloselytothoseofBurkeetal.(2003).Thoseauthorsdescribedwellthe
differentdisplaysseenduringcourtshipand,male‐malecompetition.Whiletaking
videoitwashardtotellifwhatwewereseeingwascourtshipbehaviorormale‐
malecompetition.ItwasnotuntilIwatchedthevideoseveraltimesthatIfound
thatthevideowehadcapturedwasindeedmostlymale‐malecompetition.The
Marinebio.orgwebsitestatedthattherecouldbecompetitionbetween“2‐5other
males”forthefemale,buttheintensityofthecompetitionwasnotclear.Themale‐
malecompetitionthatIobservedwasabouttwofullminutesofnon‐stopdisplays
andcompetition.
Throughdirectobservation,Iwasabletonotonlyseethedynamicdisplays
thataccompanycourtshipbehaviorbutalsothelengthoftimeandeffortspenton
eachcomponentofthematingsystem.Theactualactofmatingissoquickthat“if
youblinkyoumissedit”butwiththetimespentcourtingandcompeting,itiseasyto
seehowtheyspendthemajorityoftheirshortlivestryingtomaximizetheir
reproductivesuccess(McKay2008;Hanlonetal.2002).
IfIweretodothisprojectagain,Iwouldhaveamuchbetterideaofwhatthe
male‐malecompetitionversesthecourtshipbehavior,sothatIwouldknowwhatI
wasfilming.Becauseofthislackofknowledge,Ididnotknowtostayandkeep
recordingtohopefullycatchthematingofthesquid.Also,becausesquidtendto
comebacktothesamematingarea,IwouldplantoreturntotheChampagneBayat
about15.24°Nand‐61.38°Wwherethecourtshiptookplaceinsteadofchanging
13
locations.Extrapracticewithanunderwatercamerawouldalsobebeneficialto
captureclearervideoofthecourtshipprocess.
Acknowledgments:
MostimportantlyIwouldliketothankDr.HeymanandDr.Lacherfortheir
helpandsuggestionsformyproject.IwouldalsoliketothankDr.HeymanandAriel
SklarforhelpingmecapturethevideoIneededtomakethisprojectpossible.
ThankyouKinnieEijsinkforprovidingmewiththemapofDominca.Ialsothank
theSoufriereandScott’sHeadMarineReserveandtheDepartmentofForestry,
ParksandWildlifeofDominicaforallowingmeaccesstoChampagneBayandScott’s
Headtodomyresearch.AndfinallytheArchboldTropicalResearchandEducation
Centerforlettingmestayhereandusetheirfacilities,andfortheirhospitality.
References:
Byrne,RA,GriebelU,Wood,JBandMatherA.2003.Squidsayitwithskin:a
graphicmodelforskindisplaysinCaribbeanReefSquid(Sepioteuthis
sepioidea).BerlinerParabiol.Abh03:29‐35.
Hanlon,R.T.,Smale,M.J.,Sauer,W.H.H.,2002.ThematingsystemofthesquidLoligo
Vulgarisreynaudii(Cephalopoda,Mollusca)offSouthAfrica:fighting,
guarding,sneaking,matingandegglayingbehavior.Bull.Mar.Sci.71,331–
345.
MarineBio.2008.Sepioteuthissepioidea,ReefSquid.
<http://marinebio.org/species.asp?id=286>.
14
MarineBiologicalLaboratory.2008.Loligopealei:Thelong‐finnedSquid.How
SquidChangeColor.
<http://www.mbl.edu/publications/pub_archive/Loligo/squid>.
McKay,KellyM.2008.MarineInvertebratesofBermuda.CaribbeanReefSquid
(Sepioteuthissepioidea).<http://www.thecephalopodpage.org/
MarineInvertebrateZoology/Sepioteuthissepioidea.html>
Moyihan,M.1985.Communicationandnoncommunicationbycephalopods.Indiana
Univ.Press,Bloomington.141p.
Moynihan,MandA.F.Rodaniche.1982.Thebehaviorandnaturalhistoryofthe
CaribbeanreefsquidSepioteuthissepioideawithaconsiderationofsocial,
signalanddefensivepatternsfordifficultanddangerousenvironments.
Advancesinethologyno.25;Supplement25toJ.Comp.Ethology.VerlagPaul
Parey,BerlinandHamburg.151p.