Insects in Amber

17 Pages·2016·0.66 MB·English

AnnuR.e v.E ntomo1l9. 93.46:145-59
Copyrig©ht 1 99by3A nnuaRle vieIwnsc A.l lr ighriess erved
INSECTSI NA MBER
GeorOg.Pe o inJarr.,
DepartmoefEn ntt omological UnSicvieernoscfieC sta,yl ifoBrneirak,e lCeayl,i fornia
94720
g
ory.
s.nl KEY WORDS:f ossiinls ecttiss,s puree servaptailoeno,e nto,m olpoaglyeosymbiosis, paleoen­
wo
ee vironments
ww.annualrevior personal us Introduction
m w16. F Asidfer otmh ebiera uitnys,e icnat msb erre pretshefeni tn feosstrs eimla ins
aded fron 12/22/ boifto hgIeen osgermacaintpmdaohi ffyceb,rnre yu h,ma evreioonoupvrspi, or rotnumrtneeoisnc tttoiuanedmlssyi tr cuercxotteiivononcl,tu itoino,n ,
nloy o paleosymabnimdoo sliesc,pu hlyalro geny.
Dowersit Amberiza(ticohna nignevso livnte hdpe r ocoeffs osr maimnbge frr ofmr esh
9. niv resiang )e nitpslr eo ciensdsu ctiotv hepe r eserovfai tnisoetnch taastr se m all,
5U
3.38:145-1gon State dfmeioilnrnui dstcaeeeatnac tadsntieo obdls,ffts y, pt h re-eexb soIoestdrk ittvehiehtimledseshnoe . urgst ccef tohoeo n m-s,pds cliitaemlynbtfesp eoeoe ne f rfas cmosio,sicol slonpoiymaarl plt kia tnzwraeoaienrnwttdd,nih o n
199Ore theeixrt adnets cendants.
ol. by Thes tuodfya mbeirn sewcatsis n itioavte2er0d 0y eaargswo i tthh fIer st
v. Entomprovided idtnehpvesa ecss5rtt0iy i opoegftfaata i mrhtobsBeine,ai osr lnn twsha ioeamtcrcvbob kteefdees rapag r ueyutsnar noe nat xtd flapo oBil ranao(al dr1 tdde0ieIi) cpnt. oadi semoHimebnotodaedwss,lreeo.t pvf toei hsrnei, t s.
nnu. ReAccess ThTehsaeoer ll edi eassmittbneT ed dare b p1lo.esc iotnst aiinnisneagcr tfesr otmh Mei ddElaes atn d
A
arec ommonklnyo wans L ebaneasmeb eTr.h ey date fErarolmy the
Cretacaenoedu xst ebnadc1 k3 5m illyieoanrI snt. h pea s2t0y earast,t ention
hasc enteornet dh eh ighfloys siliTfeerrtoiauamsrb yed re posiintt sh e
DominiRceapnu bwlhiicca,hr f ea ierxltye nasnipdvr eo vais dtee aidnyc ome
foDro miniwcoarnk eprrmisa rlyit hrosuaglfheo urs aes j ewelBreyc.a uosfe
thsec ienitnitfiecor fse psetc imfernosm tdheepsoest ihteasiv,ra ilaabnidl ity,
threa ngofei nsercetpsr es[esnotme2ed3 5f amilaitpe rse s(e1n3t) t]h,i s
revieemwp hasDiozmeisn iRceapnu b(loiDrco miniacmabne)ir n sects.
Ambefro rgecrainse usd deanplpye aanrdl oovke rayu thenFtoircg.e ries
invotlhvpeel acemoefpn rte senitn-sdeaicynt tpsol asotrsi ecm i-fossilized
145
0066-4170/93/0101-0145$02.00
146 POINAR
Tabl1e M ajowro rladm bedre posciotnst ainfionsgsi inls ectsa
Approximaagtee
Amberd eposits Location (millyieoanr s) Plansto urce
Tertiary
Baltic NortheErnu rope 40 Notk nownp,o ssibly
araucarian
Burmese Burma 40( somyeo unger) Notk nown
Claiborne Arkansas 45 Shor(eDai pterocarpaceae)
Dominican DominicRaenp ublic 25-40( somyeo ungerH,y menaperao teanrda
g
ory. somep ossioblldye r) possisbolmye o ther
ws.onl Hymenasepap( .L egum­
ee
ww.annualrevior personal us RSMFoieumxcsaiihcnluainina a nn CCRShhoiiimcnaaiapnl ayiMs ae, x ico 43230200-- -524360( som7e0 ) THNNaooyixttmnkko oenndsonoiewwaa(annecLe eae)ag eu?m inoseae)
m w16. F Cretaceous
aded fron 12/22/ AAtllaasnkCatonia cs tPalla inNA olSrattsakhtCaeeonasa s stteParnll a Uinni 78t50e-?d1 42 AArraauuccaarriiaapccleeuoaastee h ers
oo
nly Canadian CedaLra keM,a nitoba7 0-80 Araucariaceae
Dowersit French NortheFrnr ance 95-100 Notk nown
9. niv Middle East Lebanon, IJsorradealn1, 2 0-135 Araucariaceae
3.38:145-15gon State U Taa Diamyttakrae nf roRme f1.3S .ov iAertc tic 78-115 Notk nown
9e
19Or res(icno paSlc)i.e ntimfaidceaf lolryg ewriileplsa stsh es tandtaersdt s
mol. d by employbeygd e ma ndm ineirnaslt itbuusttpe es,c itfeistcth saw .ti ldle tect
v. Entoprovide moTshftepo rregseaaercrnicaeteov soua fnia tlm( ab2bei0lrn)e s. e ecmtpsh asviazreisao ruesoa fps a leobio­
Ress logGye.n earnisdcp ecliifsaitncasd c couonfat msb eirn seacrtaesl saov ailable
nnu. Acce (51,0 1,3 3,2 -38).
A
DatiAnmgb er
Ambehra tsr aditbieoendnaa ltlbeyyde xaminiinndgfe oxs s(iflosr aminifers,
coccoliintt hhbsee )d roccokn taitnhiaenm gb eTrh.i mse thoodfc, o urse,
proviad meisn imudma tbee cauistde o enso tta kien taoc cotuhnett ime
involfvoetrdh fi en adle posiotfti hoaenm beirnt hbee drock.
Alalm befrr oamp artigceuolgarra pshiimtcaeay nl o bte i dentiinac gael.
Nuclmeaagrn erteisco n(aNnMcRes) p ecsthroaw tehdaa tm befrr odmi fferent
mineisnt hDeo miniRceapnu bvlaircii enad g aen do thcehra racte(r9i)s.t ics
Ageosfa mbeirnt hneo rthaenrned a stpeornr tiooftn hsDeo miniRceapnu blic
wergei vaesn2 5-40m illyieoanrw si,ta hm befrr otmh wee ll-estLaab lished
AMBER INSECTS 147
Tocmai nbee intgho el de(s4tm0 i llyieoanr Tsh)e.sd ea tceosrr elfaatierl y
welwlit ht hceo ccoalgidete ht erminparteisoennstC eedp e(bk2y 9w )h,og ave
agersa ngfirnog3m 0 t o4 5m illyieoanfr ostr h Lea T oemai ndee posits.
FirOsctc urroefIn ncseTesac xtia nA mber
Iti si mporttoka nnotww h einn segcrotu fpisra sptp eairnte hdfe o ssil record,
anda mbedre posciotnst maainnyfi rsrte coorfdi sn sefcatm ilainedes v en
somoer derIsn.s taonfct ehgsee ologiecaarllrlieyec sotor fdi sn sefcatm ilies
occurirnai mnbge arrt eo nou mertooum se ntihoenra en tdh eea rlrieecsotr ds
orgy. ofg enearraee v emno reex tensive.
ws.onl Alolf t hteh rmeaei gnr ouopfss ociianls e(cbtesea sn,t asn,dt ermites)
ee
ww.annualrevior personal us aCd(pro2peu7Aetb) airata.bneCrsc o apr etumueohtabttaue neeasasi trdcsn nLa , eid om og wbounfeeC(fs1Tiurrir r 2nr nei1,gtvgt3 hoea4p,enscr1 Lartie) eh sio;boi(cgu awantas netthota evohelimRse fa dobaeroteknews ,s snsronm sto uniai whl tbbyblnadese t e wes ey th ax )eoeUrp a prrrpaelensirtse eesddt
m w16. F knowrne coorftd h igrso uipfi ndetehdse p eciimsde ent ermtiobn eea dna nt
aded fron 12/22/ (tAhToA.eerc redrtrpaTisae,ha mr yrbssyecoa cronno( naum3tlrm0aMt au)iah n,nenei sat c roald(tkie1inea3oZos )nowt,u)rn n.a d pi(ts6e1p)rrEu,aemt pberideo spe(tn2eot8rfa)a t, i ves
oo
Downlersity SiFprhaogininal(spe1et t3ceah)trna,nsadSt o t rrmeapalsrln(eiyo8 p ft)to .eu rniadn f osfsoirlmm a ys ometimes
59. Univ tumu pi na mbeBru.t terafrloein eees x ampalp er;o bambeltea lmaadrukl t
45-1ate (RiodinLiedpaied:o pitnDe ormai)n iacmabne( rF ig1u)ri esp resedrovwend
3.38:1gon St toAt mhfibene aerlds setcot o aniitnlact,ilh nfuessid c riaasnplatgpen esdca orloapfomna ract netby el rnoso.d -suflcikeiisnn g
9e
mol. 19d by Or tChafeno assdirailema cbnoe(i rr1sbd 3 yfA ) ar.tre h coeefl nidtnekb dsnyT to e wfdPno i skooseffait m lhoC esu qluiiicnC(ti rFodei atge2au )cr.ee o us
v. Entoprovide IndiErveicdteo nfPc leaO nctc urrence
Ress Indireevcitd efnoctreh eex istoefnc ceer tpaliangn rto udpesr ifvreostm h e
nnu. Acce appearoafsn pceec iailniszeeicdnat msb er.D oTmhien iacmabnef riw ga sp
A (AgaoniHdymaeen:o pstheroawi)nnF igu3ra el rehaadisyt a sd aptaftoiro ns
enterfiifngrg u iatnsid,t p sr eseinsec vei detnhcaFeit c utsr etehsr iivnte hde
vicidnuirtiytn hgae tp och.
Biogeography
Dominiacmabnec ro ntaiinntse reesxtaimnpsglh eosw itnhgat thd ei stribution
ofs oemi nsewcatsms u cmho reex tenisnit vhpeea stth aantp reseTnhte.
abiltiort eyl aamtbee irn setcomt osd em-fdoarym asl louwsts o m ake some
interesting oonbt sheepr avsdatit sitornisob fuf taimoinl ies,a ngde nera,
specTiheess t.u odfay m bearn tisos n eex ampRleep.r esenotfta hteeix vteisn ct
angte nuSpsh ecomyromfat heex tinct subfamily (fSoprhmesc omyrminae
148 POINAR
intermebdeitawtemeeo nd eamn tasn dn onsowcaisaplhs a)v bee efno unidn
CretacNeeowJu esr saenyCd a nadaimabne( r4 1T)h.e sfeo ramrsne o we xtinct
att hseu bfalmeivlAeyll t.h omuegmhb eorfts h aen gte nLuesp tomyrmeexxi st
inD ominiacmabne trh,ag te nuisse xtifnrcottm h Nee wW orladn di sn ow
confitnoet dh eA ustrarleigaino( n2s)M .e mberosft hea nts ubfamily
Dolichodearriern eaper esebnyt seedv ergaeln e(rian dudAizntgec a,
DolichodeHryupso,C /inaenaMd,o naciisnD) ominican( 4a2mH)bo.ew re ver,
noe xtarnetp resenotfta hteigsveeen seo rcac iunrt hGer eaAtnetri lles.
Verfye ws awflioecsc uirnt hWee sItn diaensdn onaer ken owfnr om
g HispanDioomlian.i acmabnec ro ntasienvses rpaelc oifDe isd ymi(aA rgidae:
s.ornly. Hymenoptwehriacb)he, l otnoag s ubfanmoiwld yi strifbruoPtmae nda mtao
wo
ee southeaBsrtaez(rni3 l1 ).
ww.annualrevior personal us lgieOvntMiurhanseebsg prit r ooetgesereoimongneDftrst o aaahmntpigioihsenmvi niaaecuicnl iasnatMdinl nhe ec w exslo ti urhcdalideaemdsci n b os e(ncr6fo7 ,oivt) fneTto.t eroe rdydro amptyiofih,tfoocte e nrah seele s t
m w16. F arFeuiarnNst o dhriestrjh Auedunrnisc stttr railbiiunatv.io olnavsmi bnegir n sewcetrsde e monstrated
aded fron 12/22/ waimtbrhee( rp4 r)Oe.fs t ehneott fhaw rtoegioeevd negesno(r afAatw n soi osdogpnoaDdtiispd taereior:ennDa p eo)oo m frti thneeidmc, a n
oo
nly (Mycetobnioal )o ngeexri sitnt sh Ger eaAtnetri lalneost,(h Meers ochriisa )
Dowersit nowf ouonndl iynt hOel Wdo rld traontpdhi tech si,(rV dal esegiuyska n)o wn
9. niv froams ingle sfpreocAmiu msetnr aliat.h eCaslnemed aa nrmylo yr eex amples
5U
45-1ate shohwo wa mbeirn clusciaobnen u ss etdod eterpmaisndtei ffereinntc hees
3.38:1gon St diTshtear bioboveufexit animsopeonlcfsOet lssWd .o rladn Adu strarleilaant iownistihhni spesc ts
9e
19Or froDmo miniacmabne prr ovifduer tehveird eonfca e f aunraell ationship
ol. by between ASmoeurtiahcn aAd u straaswl eilaal so thpearr otfts h weo rladn d
md
v. Entoprovide 2sh5o-4hw0mo iwtl hlyeie oarnorTa fschn .eigr rseet diaunicsnhteh aicbasotbes nsee e endsn rt aasbtlniiocsotahn lelflidyoy nti r hn rpesea edscuttc se,d
Ress buatl sfoo sro mpel anitnDs o miniacmabne Fro.re xamptlhete,r eteh at
nnu. Acce produmcuecdoh f t hDeo miniacmabneH ry,me naepar oteriasm, o sctl osely
A relattoea dne xtasnpte cfioeusn idnE asAtf ri(c1a5 S)u.c ehx ampolfe s
disjudnicstt ribouctciuiornon tsh aemrb edre posaiswt esl els,p ecitahlel y
BaltaincdC anadian.
Behavior Intfreormp rAemtbeedrI nsects
Thes udddeena tohfi nsetchtafsta liln troe soifnt aelnl otwhse tmo b e
preseirnvb eedh aviaocrtTashl.u msa tiinngs eacrtoesft epnr esebrevfeodr e
thecya dni sassoMcaitaiptnaegi. or fss c atoflpiseiasdsw, e lalsc hironomids
andg alfll i(eFsi g4u)rh ea vbee enno tiendD ominiacmabne r.
Figu5r seh owasr arfei nodfm atgeu ardiinan p ga iorfw atesrtr iders,
Electrobsaptiensi p(e)1sf roDmo miniacmabne Irn.m atgeu ardwihnigc,h
AMBER INSECTS 149
g
ory.
s.nl
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ee
ww.annualrevior personal us
m w16. F
aded fron 12/22/
oo
nly
Dowersit
9. niv
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3.38:1gon St
9e
19Or
ol. by
md
v. Entoprovide
Ress
nnu. Acce
A Figu1r eA butterpfrloyb,a bolfty h fea miRliyo dini(dLaeep idopitnDe ormai)n icaamnb eNro.t
onlayr es ucfho ssrialrsbe u tt hearye q uiteex pensoinvt eh oep emnar kets,e llfiontrgh ousands
ofd olla(rpsr esleoncta toifos npe cimeunn known-unoltehsesr wniostee, ald ls pecimenisn
figuraersei nt hPeo incaorl lecmtaiionnt aiantte hdUe n iverosfiC tayl ifornBiear,k eley).
Figu2r eA femalmeo squi(tCou liciDdiapet:e trah)aw ta sr ecendtilsyc ovebryeT de dP ike
(UniverosfiC tayl garAyl,be rtai)nC anadiaamnb e(rp hotcoo urteosfTy e dP ikes;pe cimen
deposiitnte hd eRo yaTly rreMluls eumo fP aleontology).
FigureA f3em afliew ga sp( AgaoniHdyamee:n opteiranD) o minicaamnbe ri ndirecttellyol sf
thpere senocfefi gtr ee(sspe cimeni nt hBerodz inskTy opez-cPeonlhlae Scmtiitohns,o nian
collec.t ion)
Figu4r eA matinpga iorfg almli dge(sC ecidiomyDiiipdtaeeir:naD )o minicaamnbe r.
150 POINAR
occuirnss o meex tasnpte coifew sa tesrt ridtehrmesa ,li en sercetm aiinns
contwaictth maittbese twepeenr ioofdcs o pulaatpipoanr,e tnopt rleyv ent
othmearl efsr oamt tempttomi antgwe i tthh sea mfee maTlhee.a bovcea se
appeatorb set hfei rrsetc oorfmd a tgeu ardiinnvgo livnisnegic ntt hsfe o ssil
record.
Afteexrt ensivelyt hrfeeo vsirseeiwclio fnrogder v ideonfbc eeh avainodr
coevoluBtoiuocno(,t3 )s howetdh atth eb ehavoifoo rr ganirsemmsa ins
constaanndtt h aotn ec ana ssutmhea tth bee havoifoe rx tionrcgta niassm s,
determbiynf eudn ctimoonraplh oliossg iym,i tloat rh aotfm odemd escen­
g dantTsh.u st,h bee havdiiosrc,u isnst ehdfe o llopwairnagg roafpth hste,w o
ory.
s.nl insefcrtosDr.n o miniacnaMdne xicaamnb esrh owisnpge ciamloirzpehdo log­
wo
ww.annualrevieor personal use fitihcTfseaxheou liefafb t fit fyoreuac ssxmraoHstabineroi me lsgrllp y iaeo l stncpe hiot aesnion mslfbtstit uo .rnhgnP ufaet, rcae h abtm eeaei cRsdlooei dyrftdsoi hum spiiv nrsiii (icnFda(ciano1igeaf4tpu 6a )har)m b,eWle.ee i mhtbathevoh rife o ral
aded from wn 12/22/16. F vbeoaoxntrct ahiehmtempo it ootudrdfisolai o anoicfnnsefntti pdp eihca ev afearcebo tpai drirlmao oyeeltmnozynloiasfoiit, nn nnhn aatk gegWnlsn ht .oOesTs wnnaa.htnem nne esa bo t menud aictgipta rsfpere iOrsetfcloi. iradWasa ootg oc tt nmilrhhhs obaleeig aonunds slnp gd , ahde ana nrrdviae eni gs es
oo
nly itbso dayn edx poisteassb domisntaelm iStoeosan.ft eirm bibtihnsege cretions,
Dowersit thaen bte comleest haragnidtc h,et nh beu gs triiknesse, rittibsne ga akn d
9. niv withdratwhipenr ge yh'esm olymTphhel. o ng hsatiiorffnst haen tennae and
5U
45-1ate legosft hbeu ga rfeo prr otecatgiaoinn st bniotytie nptga raanltyAszn eodt.h er
3.38:1gon St laanbttuf tgoeP,srras onipbttliu lgao rcdteeowr luoodss ro uicsbe codudi rinBose af asl n t iacDno tla,imc bFheoorduet nwre(duri3 stt9t ih)ha .rii su s,
9e
19Or proviaddidnigt ieovniadleo nfbc eeh avifoirxabilet cya utsheeex taannbttu g,
mol. d by Ptiloceorcuhsr acefuese odnsm embeorfst hsea maen gte nuDso,li choderus
v. Entoprovide bailtshuoab evbreec uedFlneao stsucssori.ffDil o rbsloe Dimdco hmoidn(eirgacuemsanbn nueo srwe . x tiinntc htWe e sItn dies)
Ress Termibtueg bse lotnogt heg enuTse rmitaraodfut sh eT ermitaphidae
nnu. Acce (Hemiptaenrdoa c)c uirnt hceo lonoifte esr mibteelso ngtiotn hgfe a mily
A RhinotermTihteisdfeao er.ma sr hei ghaldya ptainvdte h eeinrt ibroed iys
stronfgllayt tdeonresdo ventrtahluclsoy n,c eatlhienhiger a da ndb ody
appendaIgnae dsd.i tion,b utghesar vmleio tsteth eciorm pouenyde osc,e lli,
andw ingcsl,e aarsfl uyr tmhoedri ficfaotsriu ornvsi ivnit negr mciotleo nies.
A termbiutg(e T ermitarpardoutse rianM) e xicaamnb e(r2 1c)l eahralsy
thmeo rpholomgoidciafli ctaytpiioocnfta hlee x tafnotr m(sF ig7u)rT.eh ese
morpholoagdiacpatla ttioognestw,hi ettrhh aed jacent orfte wmtoa eirnmsi tes,
strongly tshucepo pnotretntsth iatothn fe o ssil aslpseloci ivieensdt ermite
colonies.
Functimoonraplh olaolgscyoa nb eu seads e videfnoccrea terpillar-ant
symbioAs icsa.t erpciolnltaaric nliunsgto efbr lsa ddesreltipakreeo jecting
AMBER INSECTS 151
g
ory.
s.nl
wo
ee
ww.annualrevior personal us
m w16. F
aded fron 12/22/
oo
nly
Dowersit
9. niv
5U
45-1ate
3.38:1gon St
9e
19Or
ol. by
md
v. Entoprovide
Ress
nnu. Acce
A
Figu5r eA paiorf w aterst rid(eGresr riHdeamei:p teirnDa o)m inicaamnb erw,i tthh em ale
(arroshwo)w itnygp imcaatle -guarbdeihnagv ior.
Figur6e An antb ugP, raecoris do(RmeidnuivciaiHndeaame i:p terianD) o,m inicaamnb er.
Notet heh eavsye taoent hea ppendafgoeprsr otecftriooamnn tb ites.
Figu7r eA termibtuegT ,e rmitaradus(T eprrmoittearphuHised miidpatee:ir nMa e)x icaamnb er.
Notea f latteonfit nhgeb odya ppendaagsea sna daptatfioolrni viinntg e nnictoel onies.
Figu8r eC aterpiplolsasri,ob flt yhf ea miRliyo dini(dLaeep idopitnDe ormai)n ican amber. Insert
showpsa irgelda nodnse ighth abdomina(la rsreogtwmhesacn)tot u lbed a ttracttoai nvtes .
152 POINAR
forwaarbdo vieth se acdo ulbdea metalmbaurtkt elrafrl(vyRa i odinidae:
PapilionLoeipdiedao:p( tFeirga8u))r Ie.tp osseasp saeisor fc onspicuous
glanodnst heei ghatbhd omisneaglm einntc,o nspidcourossuaeslt aea but
frinogfle o nlga tesreatlaa en,da p aiorfv ibraptaopriylo lnta hete h oracic
shieTlhde.sc eh aracatrteeyr psi cfaolulniydntr opicriaold iansisdosc iated
witahn ttsh aotb tasienc reftriootmnh spe a iraebdd omignlaalna dnsdp rotect
thcea terpfirloleman resm iTehsil.sa rmvaaya lshoa vbee eans sociwaittehd
antisni tosr ighianbailt at.
Mimicry
g
ory.
ws.onl Fosseivli deonfcm ei micirsry a rheo;w evaenre ,x ampolfae n mti micry
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ww.annualrevior personal us Ta(bynmepdyeair trgmcmleeoeae ndc luTiolisfilmy ileo inosdrtnDc t ph olrheMyamuyte p iucm)mansti bioi.aumerfwc morir habefcstine, i sht r n aieg vbcnse oeod bmnlal eaautovrofh rsene reeak n tsamividto tatbin oo smilheogfd tiioee rs ihdmn ro cueei, faern c nra etata(llhcsml1yet by1t iy )rvc.aei , d
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tob ef aiardlayp tianbr leeg atrogd r owicnogn ditAinoontshs.eu rg gesitsi on
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