Pollen Feeding and Reproductive Biology of Heliconius Butterflies Lawrence E.

Gilbert Proceedings of the National Academy of Sciences of the United States of America, Vol. 69, No. 6. (Jun., 1972), pp. 1403-1407.
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Proc. Nu(. Acacl. Sci. ( J S A Vol. 69, No. 0, pp. 1403-1407, Jttnc 1972

Pollen Feedirig and Reproductive Biology of Heliconius Butterflies

(fat I)otlj/nitroge~lorrsconlpor~ rltls/rrprodr~c*t straLegj) ive

LAZ\\iltEN(>li:i:. C:ILl<ElbT l
L)ei).~rtnrent r Zoology, Univeri~tyo l Texas, Auit113, Texa5 78712 o

('omm~~nzcutetl Vc rnc Cranl, 1tfarc.h22, 197'2 bit

I3r1t ~erflies ~ I l e OF neo~ropical Crnrrs llrlihUSTI{ AC'I' c n n i ~ ~ eet1 on pollrrl. 'l'kris is ~ I l e Fs first linown ins~arlre ill
r
7

:LS long :is 10 min, ahereas nec~tar viiiti typic1:tlly 1:ast 3 ~ no Inore t i r : ~ ~see. 1)1ir1r1gpollen viiits, the butterfly reI)r~t~rrflirs a IlahiL ~kraL wrll krrowrl for othrr irrsrcls. OF is 1)e:ttetlly icrlapei its prohoiclis tlp over tire lantircrs with 1hr hrr~trrflirs remove a r l ~ i ~acids arrd prolrirrs from pollo illort jerky t h r ~ i i t s :ar~dr:irely thrusts deep into the corolla len; his frrclirlg irlrrova~iorl plays a rolr in the reprodr~rLive and pop~tla~iorr lriologj of these inserts. 11 is sr~gfor n e c t ~ r . gestrd t l l a ~o ~ l i e r animals may rrsr pollrn in a similar (2) Other species of butterfly do not ~ac~c~urnul~ate poller~ Fashion. 1o:itls even wllcr~visiti~rgtllosc flo\vers t l ~ r)rovlde pollc~r t for :a rlunlber of Ifeliconi7~sspecicls. For eaamplc, during :L It i i gei1er:illy tho~~g'lrt : ~ d ~ i1epidopter:tni :ire url:~hle tl1:~t lt stutly (1)ec~cnlber 1!)60-1)ec1cmber 1070) :it Arima, l':tss, :tnd protei~r, :tlrd tllcrefore tlmt the to :issim~l:~te amino : L C I ~ S I Tri~ritl:td,MT.I.,Ieliconius elkilln frequently clollected elaiily rlitrogenoui conlpourltls ol tlrcir eggs lare tlerivetl only from viiiblc polle~r 1o:ttls from I'nlicorea crocea (Ibubi:tce:~e),cvirere:ts reiervcs (fat body) laid tlolvrr :is a rcsult of 1:trv:~l feeding Parzdes spp., tire only otllcr cLommon butterfly viritori to (1, 2). Eggs :~rc,tllerefo~e,protlucled :it tire eypensc ol f:tt thescl flo~vcri,tl~tl rot collecd, pollen. I n arl 1r1sect:try a t Tire furtirer eggs :ire protluc~etl. body, :t~rtlcvl1c.11 it d~s:tppe:iis ~ r o A U~rivcrsity01 '1'ex:as : ~ t ustin, 1)ryns julia la~rtl Eueides Itecently, ho\vevcr, la hl:~l:~y:an oth h:ti been described t h : ~ t m isabella (hotir nor~pollen c c ~ h ~ I~clic~oniines), ~ rHeliconius f rg :~ tl takes hlootl nleals nlrrc~l1:is nloicl~iitoci(lo (3). Tllc prcre~rt erato all visit /,anlanu cnmara h u t orlly I f . erato builds pollen tire report d i s ( ~ ~ ~ i s c ic~videncelor :in :tn:tlogot~stlevelopment 1o:tds. butterfly gerlus Ilelini c~crt~alil members of tllc r~eotroplc~~al I'ollcn 1o:ttis simillar to tir:tt in Fig. I ir:~vcbeen ohicrved conius (Nympir:ill~r~ac), ~vllichthe source ol mino no acids in 111 tire follocvirlg sptlcies of Ileliconius: ernto, nzelpomene, proteins for tire 2ad11lthuttei fly is pollen r:tther than blood. :~r~tl ckaritonia, clysoni?tzus, sara, hecale, ethilla, doris, isncenius, tire field Studies performctl in the l:ib, 111 insect:tries, :inti i ~ r sapho, wallacei, rydno, pachinus, la~rtlheu~itsoni. Eueides, a iilthcirte hocv tire nitroqeirous c~ompourltlsof polle~r :ire ~ c gc~rus ften lumpctl ~vitllIleliconius, tloes not contlair~polleno ~novetl y I~elicolii~is Iron tlrcly ~r~fluc~r~csc,reprotluc~tlvc b the inrtl g:~tirerir~g specLies. 'l'he slanlc is true lor tire renllaining six it1ategy of tllesc b~itteifiic~> :tnd genera ol hclico~rii~res :~llotllcr New TVorltl groups cx:imir~etl. (3) 'I'llere exists : ~ n el2ahor:ite pollen procsessilrg bel1:tvior N o s t fieltl obierv:~t~orrs were nl:ttle i r l locve~ molrt:~~rc wet tire tlmt hegirls \vitir tire form la ti or^ of :i dry m:Lss o ~ r verltr:~l forest [Ci l 0 In (2000 ft) elev:itio~r] rie:Lr Brimla I':~si, Tri1lid:ad iide of the 1)robos(sis 11c:trtire 1le:itl. N e ~ t:i c1e:tr liquid (prob, :ir~tlin two lo~vl:ir~d r:tin-lorcit sit('s 111 ('entral Bn1enc:t [O:tbly nect:tr) 1s e\utl(,tl from the proboscis tip :i~rdis miyeti 122 m (0-400 ft) elev~ation]:Ihtrro (>olor:ttlo Isl:tr~tl,C:t~rlal tvit'lr tlle polle~r.Subsequently, tllc \vet pollen load i i lagltlated Zone, : L I OS:LI'eilinsul:t, ('ost:~Iti(1:t. F ~ ~ r t l lobscrv:~tion:~ild I~ cr ur~coilingof the profor several llours throligll :i (soiling :~ild noi it eq)criment:il \vork \v:~ipc>rformed on butterflies livi~rg boscsis. citller in outdoor sclreened ii1sect:trics (:it tire Necv York (4) hIorpirologlc~al fc:ttureiof the probos(is corre1:tte cvitll the Zoolog~c~~al Socsic~ty's 'I'ropic:~l Resc:ar(~I~ 8t:~Iion : ~ t Siml:~,
functioni dcscrihed hove. P2apill:ie on the prohoicir tip (Fig. lrinlla Valley, 'l'rir~id:ad) or in 6.4 X 4 nl (2 1 X 13 ft) g1:iss
2ac.t rcc~eptors ~atldltion in to 2A) prohz~hly :is cllrerno-rnech:~~ro grerrllrousci a t Stz~nlordIrliivcrsity : ~ r ~Tire IJnivc~rs~ty tl of
r , ts f~mctionillg togetllcr :LS :L pollen brusll. Some ~ ~ r s e ctir:it do
1 eu:ti, d u s t i r ~ 'l'hc gree~r'lro~~se . pop11l:itioi1~
beer1 nl:tilrhave not fcctl or1 pollen sucs'lr :LS Papilio :irltl Danaus 1:ick these
hrertling r)opul:itior~s over 2 ye:~rs, for t:airlcd :is (~1rti1111o11sly
in s 1)al11llac~ i r t i ~ c (Fig. 2C), w'lrll(~ otlrc~ (such :LS nume1 ous
ly :~ntl nrlult h c h i ~ v i o irl 511('11 err(~101111cii it1 m o ~ t s l ~ t ~ ( ~ l i ~ i a :L ~ tliffereiltly and :ire
nymp1~:ilinc~c,nc,r:~) ~ : L ~ ) I I ~:are( o r i ~ n t ~ d
g to t id~nt~(>:aIh a t of \vild i~rdtvidu:~li. (L. gro~ipetltlifferc~~rtly :is to scwe :L thffcrcl~rtfur~cstio~r
io 1~;videnrror pollen Feeding f Gilbert, ~ l r ~ p l l b l ~ sobs~rv:itioni).Large mc~c~h~ar1o-rc~c~(~1)tor ir~d
h:tirs ne:tr tire I1c:ttl nl:ty provitle lcc,db:~ck cwr~cserr~ir~g the Nunlerous lillcs ol eviderlce irrtliclatc that pollerl is c:ollcctctl pollrrl 1o:td sl1:tpc :ilrtl size (lj'ig. 2B). Tllclsc I1:iirs :irc lcrs for its r~utritivc valuc :allti lot ~)ic:kctl 11) :LS :a11 ir~tlirect 1 result tlcvelopetl in :ill spcc1ic.s c\:tminc~l t h a t do riot lcctl or1 pollcn. of r1ect:ir visits. Rlood-lcc,ding mothi hlavc c~volvc~l simi1:ir nlorpl1ologic~:~l (1) Thcrc is :L tlistirictive pollc~r-collccti~lg be11:ivior.
fc:tturc,s tllat cwrrelatc cv1tl.r tire fcctlilrg I1:ibit (4). Ileliconius erato g:ttllerilig pollcn in the ir~scct:iry11:~vcbcerl
(5) E\pc>rimcrlts with a~tifici:~l o ~ v ? ~ ~ r:an ini(,ct:~rya t fl 111 i t obscrvcd to remair~: ~ a siilgle floret of Lanfana camam for

1404

Zoology : (:ilbc~ t

I'roia. ,Vat. .tcad. Aci. ITALL (lMT2) iili

Frc:. I . IIr[iconiz~sctkillu I1c::~rirrg1:irgc (no. : 3 ) poller~ load collec(ce1 i l l less 111:u13 nrir~f r o r r r

:I.

sil~glrnr:rIc flowt:~.ol' G'iiortr~itr spin,,iiosn

rlenr Arinru I'ass, 'l'1,irlidad. St;~nford slioa- t11:lt I/eiiconi/ts eralo Irns cxqnal ~)rcfc~i.rncc lor glass i pollen a l ~ t l~)olle~l-siz~(l 1)c.atls. 111 :c:lroic:e i:s])erirnc~lt, Hclir:oniu,s in ;all ilrsec:t,nry wcre given ti : ~ r r : ~ y 25 :~l.tifi(:i:~l of flo.c~ersiri sr:t,s ol 5 coritairlirrg tlry l)ollcir, gl:~ss br:~tls, :L sucrose estritct 01' l)ollen, 15% s~~c:i.osc, wiilel.. A li~ilrant1 1:~l)sc ideo ti~l)e v recot.tler itllo\vecl 20,000 si:c ol' visit tirlle to be tlle :~vcl.iigcvisit i l l itriitlyxed. 011 first (lay, tlie Ierigtli of : ~ n sccoritls wits -10, 40, 2 1, 34, itntl 9, \vhilc t h percSerit:~ge tot:~l ~ of visiting t i ~ n e 15,:~s32, 2 1, 17, 17, itird :j for l)oll(:n, g1:~ssk)e:~(ls, 1.cs11cctively. On sul)1)ollen cwtr:~ct, sucrose, :~rrd .c~:~tci,, scclucirt tlitys tlie butl,ei.flics seerlletl lo tlifCereirliitte I~etwecrr ~ ~ o l l :LII(I~ glass I~catls, el I~rlt tlic I:~t,l,er still 1,cceivcd i i gr(::~t(:r ~)ercerit,:~gc tot:~lvisit,irig tirnc thiirl tlitl tlie otlicr "flowers" ol' thitl (lo riot Ilossess pollclr. l\'Io~.eover,1o:~cfsof g1:~ssbcatls itre collecstctl :inti tl.c:~trtl ex:lrtly :LS iS Ihcy were ~ ~ o l l r n loittls. l'llis clirrrirrates tlie possil)ilil,y tl1:l.t but,t;crAirs irrvolur~litrilj~ 1)ick 1111 l)ollcli hcc::n~seoC sorncbchcrllie::~I11lirllie:ry of rlec:t,:~t.. will elscwlicre. I)ei,:tils of tlicsc cxl)c~rirlletrt,s :~l)pe:~r (6) Esl)(:rirnctit,s by I,i~rskerrsi~rltlcoworkers liavc slrowtr tliitt cert:lirl ~ ~ o l l c will :~ctivelyrcbleitsc rllost ol' tlrcir 1)roteiris ns : L I I ( ~ i'rec, :~rnino:~c:itlswil,liin rnirn~tesof bci~ig I I C I I ~ ) : L ~ ,ill ~ ~ C( :1k)o11lr 50% suc:rose solutiolr (5, 6). 1)uritlg tlic, first niilrl~tc, ol' t,Ii(. i'tw arnino :~(;i(ls II:LVC IICCI~ r(:l(:;tse(l frorn living 1)oll~lr sc gr:~irrswllcn incltb:rtctl : ~ 25' irr LOO& s ~ ~ c r osolution. Pollen t pregcrn~itiatio~i concentr:ttion oi' mitli :L total :~n~ino-:~citl 2.56 @nlol/nig of ~ ~ o l l e n , rcblc:tsc~l5fi0 rlrr~oloi' :~niino:tcitls ir~t,otlxr sl~crosc~ sol~~tiorr aftel. 2 hr of iriclil):ttion. l)uririg ger~rii~r:it,io~r t,Ilc'i.c~is :I ~i(tt, ~~-1ti11('sih :itnil~oil(*i(ls ill(' of 1)y ]I~)II(~II. of gy, ilritl 1ic.c:;ausc of Illis fe:~t,rtr.c, 11ollelr~ ~ l ~ y s i o l oirlg:.c>st,io~~ ' or cnxym:~tictligcstiori :LIT irot 1.cr1uirrtl Sol 1,11(,I)11tlcl.f1yto I~itlcrtl, wlirlr ~ v c 1)olle11 t k)e :ll)le to remove i,hesr c~oml)orl~ltls. ilici 1o:~tlswcre ~.crriovctl fro111insectary If rIic.o,?ius :rl~tl r 1)rotc,iris \VPI.C iinitlyze(l I)y ~ l ~ c t r o ~ ) J ~ ~ ril,' W;IS fot111(1 ( SiH, 1,lr;tt t h ~ y ; g ~ t colrtailrctl 1)ollcir clrxyines o~rly tro h~ll,tcl.fly ~cllayrrlcs wet.(: ill tlct(~c:tctl.IE'ollcl~ loirtls \vcl,e irnrnctliit1,ely srlsl)c~~tletl c.lrilliltl 15% sucrose solutiolr itii(1 c*~rltt.ii'ttg~d. I'r.ot(~ir~s s111)rl.of n:~t:~lit \trcrc s-;cl):~ritt,~d clecl,rol)horc,sis or1 tliscolll,iri~lo~~i: by gels (EC Al)l)iirittt~~ itcr~~liirni(l(: (:orl)., l ' l ~ i l : ~ ( l e l ~ ~ I ~ i : ~ ~ I'ii., 'I'eclrlrical ISrllletilr no. 140), wlriclr wct.c st:~ilt~(l i ( ~ t ~ e i ~ ~ o l'or itrniuo 1)eplitlase (E(!3.4.1.1) :~ntl~lo~rsl)ccific esl,ct.:lsclh.1,ilic.of W \ Sot, wisr crop :iti(l gul, (;o~ltrill~ Llr(>k)utt,erfIi(~s ( ~ I Yitss:t~,r(l tlie s:lrnc, C I I B ~ I I II ( ~ S . (7) To sc~:i S itniiiro :LC~(IHrrg(~st(%(l :~iIull, i I)!. lIr:li(.onitis ~ ~ ( ' r r tlil.rct,ly il~volvctlilr cgg l)rotlrlctioti, I ~.;trr tlrc Followiiig cxt)crirlic~it: '::lcli (lay for 8 tl:iys, 15 11. crnto i'ern:rlcs \vc%~.c\ c ~ i 1 S :1 silificic~rt: ~ m o ~ ~ r r t ol' ll"(:j:~rrri~roitcitls,o :1.1,1)t.osirn:rtc.t i ~ v t irit,:llic of tllosc :~mirlo:~(;i(ls l l ~rl:~.\inlt~rn i (1:lily 1)0111~11 1o:reis IIIII(IC~I.fi(:l(l (;o~~(litior~s trig) 1; :~ssuririirg 2-111,i~~c~uI):~t,ie)r~ (1 .5 :L 1)olle11 pcriotl. h f t r r 2 1ir oi' iric:uh:rtion, 1 rng of gcrrni~r:~ti~rg 3.3, )\-ill h : ~ v (i~~le:~s(:(l :3.3j 4.4> : I L I ( ~ 6.0 rirne~lof :11,girii11(1> ~ le~ici~~c., ant1 valirl(%, Iysilre, r~s~)(:(:ti~.ely (6). Eggs wcre c-ollectetl i'ol 6 tl:tys I)ci'ore, : L I I ( ~% d:~ys:~ft(,r. li;:~cl~ the 8-(lay perio(1 ( I ~ i i i ~ ~vI~i(:lx ig l:~k)(:l w:is it(l~riirrisi(~~~(~(I.

(1;ry's (!gg p~.ocl~ioLio~r t~,cl:~t,(~l t v ( ~ \ k~ l ~ l o so111tioi1 tvns \vi011 ( ~~ox :rcscsossoi,y I II t,o ~.c~rl~ovc. gl:~lrtil ~ r o t o :iL~ I~~ (:11loi.i011,O I I I O ~ C I I ~ Z C ~ i l l sci~~i,ill:~t,io~~A \ ( ~ ~ ~ :S s o l ,P;~igl:u~(l tlui(l ( iew Nrl(sIe:i~, arl).)> C itritl (.OIIII tetl. 'l'lrc rc~s-;lilts j~lotlrtlits :tvei,:Lgc c-ounts I)clr rriirr ~ W I ygg :ig:ti~rstd:tys, i ~ ~ ( l i c ~ : ~ t v , LIl:tt, 111)t:~k(>f i , e ~ :: i n ~ i ~:t(,i(l,q ol' ro eggs :III(I (Pig. 3). 'I'lris i b ~loi, i1l1,o (iov(~lol)i~ig is 1,:11)id striki~rg I I I I ( ~ X ~ W ( ~ , ~l(l Yiris(~c>t,s, ( -as 1 i ~ ~ ~ 1 rrially l l ~ ~ t i r o n isl~(~ci(:s, t~s tvl~icll ~ ~ ~ ~ o tll:il,tll.e ,eggs i'ronl t,irly oocytes ill just, :t few rl:~ys..lpor thic~ :it :LILY givcr~ l,itl~e, eitcli ov:~riol(: I$. <:r(l/oC O I I ~ , : L ~ I I S of i~lst,:~~lrc, t i 7 17isii)lc oocaytc:s, tlic l:trgt,ht of wliic~h\\rill rnatr~l.~. ilito :i 1:1,1,gr0.50- t,o 0.G-rng egg witlrirr :I.t h y , l,li(:srrlallest of I\-liic-lr ih :d)o~it, pt11ill rli:tni(:t(:r. 80 I t is 0 S i~lt,ei.est o ~)oilil, t out Irere t l i : ~ tI(:l)itlol)l,el.:i~rs :LI.(> \<now11 to tr:~sSc:r i~lje(:te(lSorc:igri i)rot,eirr u11e11:~~ig~l iroln tlre 1rc~rriolytrrl)ll ir11,otlcvrlol)iirg eggs 1)y pi~loc.ytosis (,7), sug:~caitls wot~ltl rc~novctl l l ~)l.ol)ortio~r be i gesti~ig t11:tt S I Y ~:trni~ro i i r tlic Ilctllolyrrlj)li. lo t,lrrii.c~o~rc~c~rl,r:tt~io~l (8) Irl :trr c:sl)c:l.irric~~rl iri :nl ou1,tloor i~lsectui.g rP~.iiiirulr irr ethilla Srrri:~leh tl:~tl,c1:tily egg lwocluctiorl by five Ue1ic:o~~ius d fctl :I dirt, of irect,:~l.: L I I ( ~ 1)ollrrl fro111llreir rtrlergc>lr(a(+: ~ y (Groul) I ) 1 ~ : c:orrij):t~~c~rl t,li:~L five frrriules ferl ilrc:t,:~i. ~s with of 6.25 olrly ((:roul) I I). (:rorlp 1 :~vclr:~gctl eggs per 0:1g tlurirlg 1)urilrg t,llc first, 5 tiilys 1,lic firs(, 5 rlayh of cgg ~)l.od~~ct,iorr. :iftcr cr:isirrg pollcrl Scctli~lgllrc:ir irrrlivitlu:~Icgg l)rotluctio~r s \\-ere 28, 32, 24, 23, :rild 21 for B tot:~iof 128 ~ g g s (:1~)111) 11 . (5 i~~s(:(>t,s) ~ tt,li(:s:~r~rr in :~~rotli(:r $\-it llout polle~r; (::~g(: \V:LS , tirrie

I\dllr:r~ f c e t l i r ~ ~ 1,iolcrgy ol: N < ~ i i c . c ~ r ~ i r r . s ancl

Y:irio~is: ~ s l ~ ( ~oft ,t,lit>life liisl,o~,yL I I C ~ ~ ~ ( ~ l ) ~ ~ o ( biology , i v e a s : l~i(~l ol' ~~ollri~-c~ollcc~li~ig He1ic~oni.i~~ cor~sistc~rrl, :LIV wi1,li 1,lir Iiy1)ot,li(:sis(11:1t t l ~ ~ r u I , ~ ~ i1)1,ovi(l(j(l po1lr11:rrc 01' III:LJOI. : ( ~ i i t ~ 11y irr~llo~.t,:~~ice to :~dr~Il, nl:~irit,ril:~~lce ~ ~ :IS1 i~epr~o(l~i(~tive :is 1 :tcalivil,y. 1Svitlerrer fl.otrr t>lircc sl~t,cicxsirrtlic:~t,c:stlrat Olicsr nrny i,ossehs the: lo~igrstuclivr :~tlultliSr hll:irrs for 1111ttnflits, mi1,ki i~~(livid~i:iIh livi~lg6 nic~~itirs oSl,(:r~ 11otlr in i~rscc~ s( ilic ) titries ((.;ilk)(:rl,, ~ ~ i i ~ ~ ~ ~ ~:~ lr i~iirll r e r lbviltl (rcl'. 8, l~;li~~li(~li oS ~g :ti~rl(.;ill~r~,t., r111l)11i)Iislr(~cl). ~)rok)1(:111 o l l t : ~ i ~ ~:Ii rregu'l'lie I:ir sourc:e of <:llrl'g)' for fliglrt rtl:ry 11c. ~);rr.t,ly solvetl fil,s(,. 1)y oi' ~~l~olirlc, 50% 01' t l i ~ :rk)~l~itl:~~rc.r wliirli caollstitutrs :~I)olll (ti) elrc.rgyTr.c:e urrli~ro:tc*itlsirr ~ ~ o l l c ~ r[l)roliire is :11iiml)o~,t,:t~rl il fliglrt, muscle stoi,:~gcc~onll)o~lrlrll irrhrcal lic:rriolyrri[)li :i11(3. ($1, 10) 1, :~rl(l s(:co~~rl, :~\::~il:~k)ilit,y ~)oIle~r, by 01' ~vliiclris hu11y lr oS ~)lit,il u st,e:~tl r,:tt,c: l l r l ~ ~ u g t,lic lif~-hl):~il :t I I ~ l i ~ o t z , i ~ ~ s at, thir to special I'e:itur.cs of Olrr l)larrts tlrut, ~)rovitle l)olle~l t,lic (8). l'olle~if'cetlirlg also 1rc:ll)s t,o csl)l:ri l l &liel):rt,t,c:rrr of rc:l)rotluct,io~lscc~iill nlull?. Ilcliconius sl)ccaies. 'I'lii~ougl~out, lollg i1,s Scrrl:~lec~o~rli~rucs lil'c, t,lrc tl:~ilyc.gg l~rotluc:tio~r a Ilclico~~iics oS 10~1' , :1,11d a t :L re1:~tivt'ly c o ~ ~ s t , : ~ ~ l l~.:~te', v:~ri(:sS~.onl1-4 eggs ()I.rnto, ?r~el~)o.n~este) 10 cygs (11. c//I~:/~(x). ex:~.r~i[)I(~, e to i 1Cor in of f~ t,li(; rirulik)(r of ~ r l : i L ~t3ggs ft~~iri(l :i s(~ric~s i~is(,cl:~ry l . ~r(~ ~ 0, errcto m:is 0, 0, 4, 1 , -1, 3, 5, 4, :li~tl4 , f o fenl:~los:Lgc~d I , 14, 39, 53, 74, 84, 100, :nlti I82 ct:rys, resl~rc~l.ivc~ly. elltiiicr /I. I'ern:rl(~s wllc~~tc:d c i'l.olli 1111. fit,lti tlr:~t:ire, Itrroti-11(hy ni:~t.ltirrg) 4 2i.t L t,o be ovt,~. rrrorit~lisoltl, 1:~y~ g g s :L I.:ILC ~ : ~ I I :to I(.hat of is in l:rbor;ttot.y ferii:rl(~sl(1ss t1i:nr 20 tl:r~,soltl. 'l'kris ~):ttt,(,~.i~ L t11:tt o i(~s striking (~o~ll,~~:l,qt, for s ~ ) c ( ~ sllrh 21s t l ~ c ~ y r ~ i l ) l i : r l i ~ ~ ( ~ ~ cclitirtr, w11ic:li 1:tys sc~\.c~r:~I Irl~~ltlrc~ti dur.irlg c,ggs I<r*;uh~d7yu.s liC(, I 3 the first tl:tys of : ~ ( l r ~ l t :111d l i v ~ sI O I ~ I O I . Ct,ll:~r~ IT(\C~<Si l l the wild ( I I). I{cc::tusa of l)oll(,rl I'c,c>tling,Lllc total ti-i:tss of pggs ~)~.o(Irfi':eti Ileliconius l)~~)l)ill)ly by eclu:tls :allti rli:Ly cs-

P I G 2. ( A )Prok)oscis t i p of HclLc.oni7rs~ h i / l ( sho\viirg tc~rmi~~:ll . t x p:rl~ill:~cx. S(>slc: .J rrrrrl. X 160. (13) l'ollel~ lo:ttl 0 1 1 pro1)osds of 0 11. ~i~clpoitcc'r~r'. R'Lech:t~ro-~~ec~(~l~lor visik)l(x rrc:lr lh(x top o h:tirs :II.(> C t k r c x plrologral)h (trrrow). Ii11ttcl.fly';i hcx:~tl i;i just, off t.hc, iippcxr riglrl of the llic:t~lr.c. Scale: 0.1 rrlnl: X 165. ( C ) 1'rok)osc:is lip of 1)nnc~ci.s p1c.r.ippct.s hlrowi~lg :L~scII(:(: p i ~ l ~ i l l S~ ,i~ . l0.1 111111: of : c ~ ~: X
200.

DAY

Iprc:. 2. I~rco~~l~or:tl.io~~[14(:]:rrr~itro:tc.idh of free illlo flcIicor~i,t~s eggs; c:l]rrl pet cgg :we plo(led sgaitrst d:tys. L:ll)eled c:orrr~:oi~rrds \ V ( ~ I . ( :~dmi~ristcxred ~ frorn 1:tk iri day Y i 1,o d:ty 14 (tioltt;ti 1inc.s).

c ~ o l l ~ l ) o ~ r ~ ~ t l s11ollc11 f'l.orll l)~,ovi(ic:s:l r.c;~,soirf o r ~.(:cxi~rllillillg t11c 1ll~~:llls ~)ollcrl 0 1 (ll~c5lloll ot11cr lll5cct5 sllcll :Is llolrc) 1)) 1 ) ( ~nrltl s\ 1.1)111(1 flrcs i,l1:1( :IVC 1~110\511 1(~11111(, 5 to l)olIr~r111 tllc :~thrlttl1c.t lor 1 ; ~ I ~ o t l(18) :ltltl c,gg p~othrctiorl(]!I), t ~ rcil~cc--

c.:lsio~r:llirlsc1c.l i'cc~tlil~g, frrlly ccllri1)l)c~tl llscl l)ollri~ :u.c to rllllc.11 :ts llrc I)llllclflics (lo. 'l'lris 1)ossil)ility is sLr~c~rgOlrc:~~etl 1))- tlrc~sc, 11:1ily ~ ~ ( : ~ ~ ~ r ~1)1~otl1r(~lio11 ~ ccr~crgcOi(~( ~ ~ ~ i tv : ~ l c ~ ~ : ~1(\:1slsor11(\ s :~to~)l~o~ c 11c &Ire Illl~lr~lrillgl~i~.(ls I ~ O \ Y I I t,o illgcst l)ollc~r i~,r(,< J I':rc~ts: l fi tl~(>ss. ygg oS :I. I'c~~rr:~l(:'s l,l.othlctiorl To1 ccjlr:~l I ):~l.~villi:~l~ (I:,. I<. (~oI\v(~ll*, 1)cl,st111:11 corr11111111ic:1t,io11), I I C ~ & : I I , I):rOs ;1,1r(1 r I Iris strggoslion is slrpl)o~~Oc:tl Ilrc: i':~cat,tlr:~ISc:rllnl(,s c 4 0 r l by arc ol't,crr t:~kcirnlitll girts stlrfctl witll ~ ) o l l c ~ l . sistc~rt,ly c;1,1~yt,lrc, I:~~.gc:sl ) o l l lontls (see Fig. 1) i ~ tlrc: f i ~ l t l , ~ c~~ r coll(:t~t(:tll)y I,ll(: r ~ ~ o l ) i:ltltrlt is i n l(~ lk:(::~trs(::L ~ ~ : s o t r ~ ~ : ( :
3

i,:lt,cs of' rcl)l.otluctiorr by IIcliconi?t.s, Illc :lnlomlL : L I I ( ~(list,l.il)t~l,ioll of tllis rcsot~l~cc sl):lcc :~11tl ill tilrlc is : L I ~ irllpo~.l:~llt key lllc :~ ~ 1,o t~~rtlcrst:~rrtli~rg s t r l l ~ t l l r L I I ( ~ ( ~ ~ T I : L I ~ Iof C lSl e l i ~ o ~ t ~ i i ~ ~

11:lr.c bcclr Illc tlcc~isivcstel) ilr Lllc tlivcl.gcrlcc of tkris gr.ou1) of' 'l'l~isis OII(> i~rll)ort,:lrrl, lnct.l~o(l (lisj)(;lxio~r oSfs1)1,irrgi l l t,ilrr(>. f'ronl l)rinlil,ivcllclicorlii~lc stock. oS (,S(~:I~)(: l):l~,:~sit.(:s l)r(~l:ll,ors, v l ~ i c l ~ \ fro111 \)lc-ir~l:lgi~~:~I :I,II(~ 1,llc of :i~.cvil,lloul tlot~k)L rn:ljo~. \ lllc rllor1,:llily 1':lctol.s for 1r10~1,1~111,1,(~1~- RIol~corc~., tlcrclol)r~rcrlt l)ollcll fcctlillg ill : L I ~ irrscc~t, flivs, cs1)cc:i:llly 1,rol)ic:~I sl)c,cics. Alol~covcr,Lllc :~1Io~:ltio11 to :ltlull Ileliconi~rsof' wkr:lt I'or nlost b~rt,t,cl.flics tyl)ic:~Ily is L:~,rv:~l of I'(:(>(lirrg Sor:~ging:~clivity(i.(;., t,l~:lt. fr:~(atio~~ (I(~vo1,ctl 1,o cgg 1)l.otlucliorr) worrltl 1,cvltl to s l r o r l c ~fcctlirrg tilrlc :nltl ~ /It lit onilts (1, (:ilbcrl, ~rlll)t1k)li5lrc~tl), i t l l r r nl:~~~lzcd c,. :LIT Ilcr I(:SS(>II I:LIY:II x \ ) o s t ~1,o~ ) ~ ~ ~ l : ~ t'I'l~isis . ~ T I O W I IL e ~ l, ,io~l o t.lr(, l nqtLrzrr trip11yllo, l ut~ibroso, Curtrnzn spint~loso, (:dl ( 'rr(::IS(: i l l rllosclt~itoc~s s~rc:l~ . I c ~ ~ n l r o p ( ~ l p ~Str:~ills of .I. :LS cs ~r~. caull ) l t : ~ ( ~ ~ : iI'(~lico~rret~ (6, CPJ)/I~I( fort~it (,), croct ntosa, (bot,li 11's trtropalpt~s vllllicall :L blootl 1rrc:ll is ~.ccluii.ctl egg l)rotlt~cfor Tor l,io~l sl)c~rtl less l,irrlc, i l l tlrc 1:lrr:r.l s t : ~ g c 1,11:1,rr st.r:~i~rs \vllic:ll ill nntr. Ccplitrcl~s lon~~cr~losa. i t : ~ ttic.:~; I. fuarcefuiczii, ('o , :r.tlrrll,s(lo rlol 1,cclllirc :l l)lootl IIIC:LI y(:L 1:~y:LII (:(111:1111r11nl)e~. I':~.TI:L~II:I; I I ~ I ~ I I ( ~ I ~r(:I:~tc~I1)1:1~11 :LII(I O~IS sl)(:ei(~st,h~.o~rghollt of (:gas (12, 13). :\t 111(, S:LIII~: tirllc:, I l o ~ v c v ctllis c s t r : ~ ~, :i.tl~rll, 1,llc trollies of' lllc Ken; \\'o~,ltl. 'I'llcsc pl:i~rLs, a11 of n.llit~lr fo~.:~gillg :ic~l,ivit,y wot~ltilc~rtl1,o ~ I I C ~ ( : ::i(Irrlt (:x~)oslll.(> LSC I,o for I j)~.tx(l:itio~r.~ r I , c : ~ ~ o s l i ~ r g l ~riglll~.tlisl,:lst,cft~l i~rscclslilw I flelic.onit~s k)rrtl(>lflics(14), ('orlsl)i(~rro~rs :i.(:livil,y i h 1):11,t :1(11111, o I,lr(: ~ : I I , I I ~ Isig11:11,o v o ~ ~ l ( ; l ~ r : ~ lc ~ i : ~ l o ~ , s . S I ~ 1 l)~ o 01,lrcl trol)ic::~l :r.lrtl t,c~rl11)c3~.:lt,c~ I~trttcrflics Iri:iy I1:irc. i l l crc,:rsc:tl : ~ t l t ~Iorrg(>vity:LII(I : ~ ( l t ~ l t lt 111~ fl.:~(:liollof 1,01,:1.1 1.c1)l.otlt~clivc cfi'o~~t, c>\-l)loitilrg 113. otllcr soln.c3cs ~rill,ogc,l~ous of c:olrl1 I O ~ I T I ( ~ S . cx:~~rrl)l(,, Yor iI,kror~~ii~~(: k)t~tt,crfli(ls t'(~:(l n t ' ~ ~ ;k)i1,(1 o sl~ th,ol)1)iilgsill :~tltlitiorr1,o ~rc,c~t:~r : ~ ~ rnl:~rkirrgsl,~rtlics (15), tl ill 'l'l.i~ritl:~tl i~ltlic~:i.tc :11, 1r:lst l\vo sl)ccics (Itlion~itr Ikr:~t. pelkccid(x FI.~c:II(~s :ri,St,:~~f'or(i,p:~,~,t,i(:~il:i,r ill IT. l~;tlc~~i)c~~g, i l t ) ,J. 13.l , (>. ~ ~ ( c~~ , J O ~ ~ I I S O A. , hlo~,row,I). h I ~ ~ ( : h i ~ l :~11tl, 1. (J. Si~~gf:], 1'. I I o r t ~1 p1.oa ~ l t lIJypotliyris clrclcn) lire, : ~ I(,:~st, lrlo~~tlrs Gi lbcrt,, t 4 (1,. help \vitrh :~.spcc(s this wo1.1~. .J. 1!Iit,c:hcll ol' 1'1. vitletl :~tlvic:c :~iltl 11rr~)rrk)Iiskr(!(l). i'kr:~r:lxi~~c I I ~ ~ I I I ) ~ spct-ics ~s11t-11:IS :LII(I I:I~~ I(, ol' Kcllt C:~.mk)ridgt~ S(:itsi~t,ific: l c . 1':~lo Alto, (>:rlif.provitlctl fal~ , Cliaraxes, .I r~nca, IJ,isioris, l'ry)or~tr, :I r/clp/~a, d/larpesin, (,ililics : I , I I ( ~ liclp with tli(~S ( : : I , I ~ I I ~ I I ~ n~i(:ros(:opf: t~lf:(:tro~~ worlc. :I slerocm~pn,Ctrliqo, cl,c. :LIT klrow~rl o l'cc~lO I I l,ot,l,i~~g I'l.t~il,, (Jost:r I:.~(::LII wo~,lc pcrl'orrrlctl i l l c:orrj~u~ct,iorr t,c:rchfield was \vit,h S ( ; I , I I I ( ; T I ~ ~ TS:II), ~ r r i r ~:111(l I~ ( : ~ (lt111g. t rn:i,y :i,lso 1x1 1)ossil)lcl11:1t, I ir~g ics i l l the Org:rl~ix:rtiolrfor 'l'ropic::rl St,utiitss fu~rd:trr~r:~~l,:r.Is tlul ,J~tly l!)i I (:ol~l.st~, - A I I ~ I I S ~ ., It,. 11. l3:~1.1,k1, K. s . 13rowr1,Jr., I ) . 11. ~ ~ ~ sl>(>(;i:~l ( i c r (:ir(*t~~r~st,:l~~(:(,s t:~ills ~ r n i :u;i(ls, : ~~o II(Y:~,:LI, it,s(,lI' (:OII I~'et%irr:r,I I ( ~ (:. 'r. , J ~ I . ~ : I . I Ik it~(ily :L rf::~d : L I I ~ (:ril,i(:izf:d tllt~nl:rl111all,l.rot~gll:L s y s l , ( ~ ~ n :(;I~e(:k ~ l , i ~ 1'01. ~ ~ i I , r o g ~COIIII)OIIII~IS ill ~~o~rs sc:l.ipt. Tlris st,~tdy :I.II ol'l'sllool, of worli c:o~~d~~c:ted is i l l 'l'ri~~i(l:~d I I ( Y : ~ , : L ~ 11:~s 1101 I)CCJI l)(~~.I'ol.r~l~tl\\!lli lc Illcsc. s o u ~ ~ c c ~ s (1 6). of wii,ll 1'. 1:. 1~;lrrIit:li , I I ( ~ spo11so1~1 his IV.S.JP. (:r:r111,s(;13-I!)ti8ti :I k)y ~lil,l.og(l~~ 11~ ~ I 1 L :i(It~Il, (lief, irligllt S I ~ ~ ) I ) I C ~ I L C I I ~ l,h(, eor~ll)oull(Is (: a l ~ d 15-2285::~. si,or~c(Ik)y tkr(> I:LI.V:L~sl,:lg(,, i r o ~ ~ofl l l ~ ( ~ :lrv~ likely 1,o s~rl)l)ly ( rl i r lr~sects(W. I I. I . I ):rvtsy, K. (:. (!!)ti:) i l l l~'cprorfi~rctio.r~l~t ~ c
1,11(. Iliglr ly coi~c~crrl,~~:~t,ctI of Lllc I0 : ~ r ~ l:~citls soul.c:c i~~o rcclt~irctl F ~ ( Y , ~ ~ I : L(Ih., F~.:i,~~(;is(;o),
:r11(iI S:rr~ 1). 2 I . l)y ir~sc~c~ts :~l.c~)roritl(,tl y 1 ) o l l a ~ 1,lr:ll b (:ll,gi~rilrc, I~ist,itli~lc, F. (1070) 2. I~:l~gclrr~:~r~r~, irk 7'11,~ /'lr!/,?io/og!/ oj' I?rscct /~c.~~rotlirc.l'rt~ss,I!~lt~rsf'o~~(l, N.Y.) i s o l ~ i ~ ~I(~uci11c1, lysirrc, r11111io11i1 l ) l ~ c ~ r y l : ~ I : ~ ~ r i l l ( ~ , L;OTL (L'crg:rir~or~ :, /i(;,s. I~~~Lort~ol.58, 1 2 - 16:;. :3. 13:1,rlxigcr,1 I . (I 968) 13~111. t,llrx:o~ri~rc:, I,r>.l)l,ol )kra~l,:~ntl v:ilirrc,). li'or ( ~ s : L I ~ I ~ I ct11(, 1)t,1., 4. I%:l.~~xigt%~,, 11 rttc 7'rop. 27, 54--88. I I. ( I!)'iO) t ~ ~ r r t , :oI'g~ '~ Y Y , I ~ I ~ T I irr ~ ) : I , ~ I : ~ I I : ~ S : ~ l ~ o lIl/ I20 ol ~ ( I :I :~ritls~ (17) is l 1,ll:~L 01111tl i l l 1)ollcll (O), :LII(I I I I I I I I C ~ O I I S ilrll)o~.t:~llt 1' :11rri11o :~citls lrlissillg I'r.olrl tlrc, II:LII:LII:L~. :rr.c, r , I llc (,:IS(>mitll wkric:l~ Ilcliconi?ts c:;lrr cstr.:rd, ~litl,ogcrrorrs

1 I I P 1):~t,l,c~.1r of ovil~ositioll :~,llo\vctl l)oll(:i~ 1))fcctlillg nr:lsinlizcs

3

S[,;i,~~cly, ( i . & l,i~~sl<c!i~s,V. (I!)(;>) I'1~3:~j.xio/. I<. I I. lilur~,L. 8, 1 47-5:;. I,i~>sl<e~is, LC. & S ( ~ ~ ~ I ~ : L,J. (I!)(;!))I,lc11~ 11. I~\V(:I , 1101. ;Y(,r1,.18, (iO.3-614. T(:lf(k~,. IT. (IH6.i) '4 I / I ~ ~ L . I+r~Lo1?1o/. I ( j l 1x4. S(V: 1). W. 11)r11. 10,

~: ~~I , 12. ~ ' ~ \ I C : L(i.I11'., , & K I ~ : I , S I I(-;. .J,< (1!)70) .I. ;lit(/, ICIILOI~LOL. 7, ::2S :::14.
I .I. 1.1. Ilrowe~,,,. I'., I31.owe1., V. %. & (:ollills, (:. 'I'. (106::) Zoo-
lo{/iccx. 8, 65-84. 4 1.5. ( iilk)e~,t,,,. 11;. ( l!)ti!l 1 Or!/(x.r~ I ;~11;ori ,[or 'I'~,opi(nl~SL~i(/ic~s ~1li1r1,oo 11'(jporl, ,I (/I,(LI~. /{;01. (:i ~~(i:i(i /~(,,s, I'op. lJt~iv(:r,si I:il,i:~,, OCi!l). Cost,:t I < ic:i ( , J I L ~ ~ - II~ I I , ~ . f'h,?jLoI. 60, 2:;3-281. 16. l'(!t,civ:il, ILr. S.(1071 ) :\F(;I~> F'rc~ila csrril Llwii L?;?;]I~, r ~ 17. I<ol.~,or I ,. I<'.( 1 970) i ~ 7 ' 1 Ilinclsc.,rr.isll.!/t,~rslij ~ ~, I'i~ot.i1rcLs, etl. I liillr~c:,A. (>. (Ac::~dertlio ' I v sN(:\v Yorli). 1:. .J I~~. 18. I,ot lrl:~.~., (I!)::!)) I,trrrt.imi~~l. ( L / L Brtswciz. 53, :;4 70. I<. I ! St,r~rltetl, (1!l(i4) %. .l ttgctu. Zool. 51, :;X3 4 1'7.