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Rev. 6yrgh y vw rgh rrv

THE CHEMICAL ECOLOGY OFDFNSE IN ARTHROPODS

M asteels and rgoire

Laboraore de Bologe ne e Celuare, Brue Unvery0 . Roevet Avene 00 Bre Bem

M owelahier

Zooogice Intitut er nvert Bae Reinprng 0 Baewitzerland

nroduconThc webs e nence sn checa snas [sechecas(10) ecnes sens Pasees (139 140)] an whch efensvessances a a nen e Snce he can wen eas a fe s eew n ceca efense n as (4) a cnseaean f wk has een eve he encan f new efensvecns We w n ae an exhasve evew f hs ae f eae. e exceen an ecen s f en (10) an (19)

s nee e a f s evew s scse aes awn easn

ae ecns as wc as w e ceca efene aansw wen wee an w Defensve cns as fequen fncn as nasecc snas an ae scsse n hs cnex

e vs excens aes f checa efense ae e c u sace ans exce an exhasve evew f hs caesan c s e cee Mean c s exane e as a fac ha a nence he ves f efensve secen Mc n aho ha een eewe esewhee (e 58 0)

Defene n ne aec f he checa neacns exn eweena}hs an e enees Hs- eseachn ehav s aseae checa ces an se eas e ve secc anes cae k e e hs asec s cvee n sevea evews(e 133 201)

64170/83/0101023$02

Annu.Rev.Entomol.1983.28:263289.

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2 PASTEELS, GRGOIRE & ROWELL-RAHIER

Chemica iteractio betwee pat ad herbivore are already illumiated by theorie tat permit predictio (78 79 1 1) I thi paperthe theory of pat cemca defee i ued a a aalogy

The umber of poibly igicat ecological parameter i obvioulyhigh ad teir iteractio are complex Their reative importace imotly uow ad oe are till urecogized. The exiig data aretherefore complemeted by ome pecuatio ad i may cae quetioare raied rater tha awered

The Pah Dbu Chemal DeeeWh he Ahd

Oly a a fractio of arhropod have bee eamied for their abity to

ecrete defeive compoud ad ome group have bee tudied moreiteivey ta other. Our preet view of te ditributio of ceicaldefee i te phyum i terefore udoubtedy ditorted everthee ti clear tat chemica defee i very uevely ditrbuted For exape thedefeive metathoracic glad of the Heteroptera are o commoly preettat tey are coidered caractetic o e axo 5 e oter adchemical defee eem to be abet or rare i the Ephemeroptera orDiptera. Witi the Coleoptera ome familie are regulary chemicaly

defeded eg Carabidae (123) or Teebrioidae (9 19) wherea iome other familie eg Elaterdae Curculioidae cemica defee ha toour kowledge ever bee repored Furter the ditrbutio of chemicaldefee i oe patchy withi a give family eg amog the Coccielidae(144) Chryoelidae (47) Staphyiidae (3 138) or Tethrediidae (139) i whic cheica defee ha evoved everal time ad ha bee lotecodaly oe taa. lealy pylogey aoe doe ot deteie teditrbutio of ceical defee.

Ecologca coditio ay be a ore imporat factor Chemical defee

hould be poitivey correlated with the probability of dicovery by preda-tor [apparecy eu Feey (78)] ad egatively wit te exitece ofateative defee eca Tu witi the Cryomelidae te eabeele (Alticiae) wic poe a quic ecape jump have reduced oreve lot the defeive glad tha are copicuou i their lowmovgrelative the Chryomeiae The Hipiae alo Cryomeidae havetrog cuticular pie coverg their dorum but lack defeive glad (47)Cemica defee copicuou i may termite pecie (49) i abet ithe aate the trategy adopted by warmig alate termite i ecape i time

(4) I ay pecie warmig i trictly eaoa ad retrcted to partic-ular hour of the day or ight (134) Durg warmig the capacity of thepredator may alo be aturated by the udde appearace of touad ofiect. O a loger tie cale ecape i te ad predator aturatio are


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CEMCAL DEFENE N ARTHROPOD 2

coo sraeges epoyed by oherwse reavey udefeded orga-ss cudg secs (eg 7year ccada ad pas (eg as frug Dperocarps Soca secs acvey proec her sybos so ha

yrecophy ad erophy ca occur as a aeave o checadefese boh secs ad pas The erga defese gas of aeochareSaphydae are srogy reduced ad soees os hghy specazederophous speces (37 The coces are ofe shor or udeary yrecophous aphds (9. Aacacas have os he ceca defese of oyrecophye speces (.

The apparecy of a poea prey s deered pa by he aure ofhe predaor. For exape aposeac secs vg o herbs are appareo brds bu oaposeac secs vg o he so surface are probaby

ore appare ha he former o predaceous as as hey expore hegroud search of food Severa facors uece he aarecy of reyogevy sze spaa dsrbuo haba ad feedg habs ad we shaassess her eec o he dsrbuo of defesve aooes

Logevy creases apparecy ad herefore favors defese Ideedshorved secs eg adu Epheeropera are usuay o checaydeeded h he bueres he daads hecods hoe dacraeds are kow o be checay proeced ad a eas he rs hree areuusuay ogved for her group (7 1 Bes (2 suggesed haposrproducve ogevy such upaaabe aposac secs gh beadapve because coud prove he dvdua's ess by educagpredaors a o he coor appearce of s hs has o o our owedge bee subsaaed O he corary ogevy s assocaed whcoued ovposo H whose poefeed habs rovdehe wh a couous source o deary roe usuay houh o reproduco ad ogevy Ledopera (

Mos ow checay defeded secs are arge or edu szed Ths

gh be ae o dcae a reduced predao pressure or seeco pres-sure a dere dreco o sa secs Sa arhropods oe behaveas sraegss reyg upo hgh reproducve raes ad usg spaa oepora refuges (189 They oer e food reward o verebrae predaorsfurher hey ca easy hde ad are ofe crypc. They are however subjeco heavy predao by oher ahropods For exape oeboa are hefavore prey of severa speces of as (33 The boogca coro eraureabouds wh exapes of such predaos e.g. upo aphds ad es byoeds syphds heperas or chysopds Moreover checa

defese has fac bee reored Coeboa (2 aphds ( 1hrps (9 ad 3 og saphyds (3. The fac ha osdeed secreos are fro arge or eduszed secs ay prc-pay reec he araco ha her copous secreos have for chess



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266 PAEEL GRGORE & ROWELL-RAHER

Chema deeses are very key to our aggregated arthropods orat east two reasos Frst brds as we as some arthropod predators tedto expore oger the surroudgs o paes where they have aready oud

ood () eod gregarous sets requety ome rom the same batho eggs ad are thus osey reated The mehasm or the evouto oaposematsm may requre k seeto avorg sgas potetay detrmeta to the dvdua but beea to the k (8) Wag sgasassoated wth toxty oud thereore evove easy umped sets.Gregarousess aposemat sets aso reases the eetveess owg sgas Aposemat beetes (69) ad Heteroptera (1) emt aggregato pheromoes hymaus grasshoppers are vsuay attrated to eahother (1)

Habtats ad eedg abts uee the dstrbuto o artropods adosequety ther apprey d deesve strategy Ths s best see byomparg aposemat ad rypt spees o sets. Predator are usuaymore mobe ad dspersed tha say speast herbvores. More apose-mat spees are thereore to be expeted amog the atter ome apparetxs b x t s xm m eous Europea adybugs are ommoy oud grouped o pts sum-mer ths aggregato s however seodary to that o ther aphd preywhose ooes sometmes aggregate o pats o pathy dstrbuto.

At ast the temperate rgos aposemat sts seem to be oudmore oe o herbs or srubs ope habtats tha o orest trees; orexampe hemay eded hrysomeds eed prmary o erbs (11)ad are aposemat Brght oors the Heteroptera are repored to orre-ate wth herbaeous habtats (18). Looper aterpars (Geometrdae)eedg o rees deduous temperate orest are rypt sets par exe-ee resembg twgs; but a obvous exepto s the aposemat ooperaterpar o the magpe moth Abraas rssularaa eedg o urrats

goosbry ushs a aa. Amg prus sets t svspot adybug beete Cla uaa preemety aposemat amogthe Europea adybugs s ommoy oud o herbs spees suh asAha obraa Calva guaa or Myrrha guaa whh hutaphds o trees ar ar more rypt.

Amog pytophagous sets aposematsm seems to be assoated wthmoophagy or ogophagy Aposemat hrysomeds are ess poyphagoustha rypt speces (). May aposemat aterpars are speasts e.g.Tyra jaba a maha rs brassa ad may other apose

mat sets are speasts o tox pats. e oa ood pat preeree a be arrower tha the tota spetrum o pats eate over te whoedstrbuto area o the spees ths tred s dut to assess rom pubshedormato Wkud (13) made a detaed ed esus o the host pats

Annu.R

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HEMIA EENE IN ARHROPO 267

o hree oigophagous caerpiars in weden ao maao Aoaaam and a a Ineresingy he ony aposemaic species mahao was aso he ony one ha behaed as a monophage

ahough nine o is poenia hos pans were presen in he aea sudiedSuch behaio appears o be chaacerisic of mahao 19 dieren hosUmbeierae ae ised as occurring in Fennoscandinaia bu popuaionsof P mahao speciaie on dieren pan species in his amiy a dieenocaiies. n apparen ecepion o his associaion beween aposemaismand monophag is ound in he widerange feeding omaeid gasshoppesha are known o sequeser pan oins (18 6) and are aposemaic Thismay howeer be consideed as oigophagy on oic pans rahe han ruepoyphagy owe in peparaion)

Seera facors appear o be inoed in hese rends. isua aposemaismcoud be moe eecie in open habias and hee are pobaby oe hidingpaces in foess o on he gound han on hebaceous oiage. he associa-ion beween aposemaism and herbs may eis because according o pandeense heoy ess appaen hebs ae usuay more oic han perenniaees (8) Hebioe insecs can nd in hose pans oins ha hey cansequese ad use o heir own deense (2 166) he cose associaionbeween aposemaic insecs and oic pans is ampy documened (32, 166)

The aaiaby of pan oins ahough faoing he eouion of aposema-ism, is howee no in ise sucien o epain i eea aposemaicspecies boh predaos and heboes manuacure hei own oxins egmeoids (188) coccineids (199) chysomeids (12) and he saphyinidau (36) Insecs ha soe pan oins may addiionay pro-duce hei own oins (168). The cyanogenic gucosides inamarn and 10ausain fund in he Zygaenidae and in Araa and Hous buees ae anuacued by he insecs hemsees ahough hey oeneed on cyanogenic pans 5 126). The abiiy o synhesize cyanogenic

compounds and o be immune o hem may hae peadaped hese insecso cyanogenic pans (166). simia echanis is aso suggesed oraciid ohs (168).

igophay inceases he pobabiiy of an aggegaie disbuion orhe insecs een oe when he ood pan is an epheea herb and hasa pachy disibuion An addiiona cue is aaiabe o predaors o oigo-phages i hey are abe o recognie specic pans ome parasioids usecues deed o he ood pan o hei hos (20). I oigophagy endesinsecs moe predicabe o pedaors as compared o poyphagy, cas for

einforced defense Pedaory adybugs feeding on seasona aggegaed andood speciais aphids ae aso hesees highy predicabe (see aboe)and heir own chemica deenses may in par be an adapaion o hischaracerisic



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26 PAT GRGOR & ROW-RAR

There are ners exeptns t the abve generalzatns abt thehaas a f has f hally f ss. h lak r saphys r spp as wel as h lak a rae ri

mr ve he grn srfae Severa f the apseaaterplars ste by Marsh thshl (112) are frest tree feerssetes lttle spealze n ther hst plants. A better knwlege f thegy f apsat spes an f hr prears s rsa hr l a sr r ap h frssawes Dr a Nr feeg nfers regarsess -reases the srvva f yng arae y fatatng the rs attak n tghneeles (1) Eet heal efense s prve by tree terpees seqes-rate n spa fregt verta (6) Gregars lfe assae wthapseas a heal fs s ths a re e srategy fr hsfres ses ha rypss a aleary sri

Tage ecc Deee AmeMs efesve aes appear t be eetve aganst a large array fptental enes r exaple he hpsrp Magorou gganeurepels preatrs as verse as ants slpgs brs lzars an aalsby ejetng a xtre f aet an apryl as (73) Salylalehye

serete by se hrysel larvae (1) s h tx t nsets abaterstat (5) The eensve seretn f the fssral yn a-rs rs a re f alehyes frans an qs repels atsan s fnsta (163) May ther atprar ps av anbt atvty (5) Hst terpees regrgtate by say lavae are eetveaganst parass an brs (53) as ell as aganst ants an sprs (6).he seqestratn f plant txns by herbvres s well knn as a efenseaganst brs (3 66) bt an als have eleters eets n the herbr's parass (52) h ak f spey f fesv ales ay

slee fr by he rsy f prears Larsa parae ses the avty f efesve ps verse rrestral prars arergetly reqre It reas lt t assess the sp atn f efensve pns n preatrs

Se efensv allnes appear t be rete at spe preatrs Afew ses ses ha h ps s fr heal efs aqases were sele fr her eay aast sh whh are hht t behe a preatrs fresh aer evres Mler Ma (11)pare the txty an aesthet atvty f a large varety sters,

lng thse serte by ytss t the sh mehale rmela Thest ely rrng pents the yts seretns pregnen3nes a relat ervatves were ag he ps st ae bassays The athrs nle that "the efensve arsenal f ytss



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HMA DFN N AHD 26

has bn so slcd to ncud onl thos srods hhl anshtc andtoc o sh Hbu t al () sd 2 knon ahrood dnsvscrons blonn to varous chmcal classs (acds srs, carbonl, or

hnolc comounds) on h omnvorous sh laa arma s a ru,dnsv scrons roucd b auac arthroos r mor rlnt toh sh han thos o thr crozoc or tstra athroods

Som cass o narro scc o dnsv allomons ar ound narthrood vnoms. Th vnoms o buhnn scoons conan slctvmamma ons that ar nacv on hr arthroo r (219221) Thvnom o th harvstr ant myrmx a s mor toc to mammas than an othr knon nsct vnom, an t aars to hav vovas a dtrrnt or vrtbra rdaors (1) houh t s a ont nuro

ton or mammals, s onl modral toc or nscts and s dvod oscc nura toct (5) Morovr nrct arumns ar avalablor susn ossbl as o scc.

Dnsv scrons can b rouhl class n thr cators accorn o hr mod o acon. Frst, stck, slm or ntann scrons,actn mchancall rathr than chmcall (1 6) mar th movmno nms or lu hr mouhars or sns orans. k lant ums,rsns or muclas th ar chmcal vrs: ronacous as n thml mr maraa (6 12) an robab n srh larva (1)

trn rsns as n nasu trm soldrs (70 1); murs o 2227hdrocarbons and mucoosacchards as n th a o Crmsoldrs (12); or as n varous Homotra, . ahds (60).

Scond nonscc rrtants actn as rllns rt thr acton va th"common chmca sns (63 65). Th nclu unons aromac comouns, small ahatc ahs, kons, acs, or monotrns and arusual ut volat and rahr racv comounds Esnr (6) and Ha(5) obsr hat th acs and alhds naur rotns, as o hstolo

calatvs, an ar thror nonscc. Svral authors sust hat

th coul also ac on chmorcors b nactvatn or ovrsmulatnth rcos (16 720 5 5).

Th thrd caory nclus tru osons actn a scc sts or ntrfrn th scc hsolocal rocsss. Thr ct s on dlad ..mss or vsaton (63); bu th can also ha mor mma csbcaus o h "bad as on assocat h osons. Th ar usuallhavr comouns than nonscc ants an ar uvaln and somms dncal to h comouns classd as ualtatv dnss n ants

(7 16). Eamls o such tons ar h cardnolds ound n vaoussecs (3 13 167 7) buadnolds roducd b rs (7) cantharidin characteristic omelids (4 9 naolnon alkaoids secretedb som lomr mllds (63 1) and n all robablt man o th



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70 PA GGOI & OWAI

varus alkals sequestere r pruce by arthrps (revee n 1)The pharaclgcal prpetes st the are unrtnately stllunknn.

These categres are nt necessarly eclusve yrgen cyane s aellknn tn but prbably acts als as an ecent lactry repellentan nteeres th the unctnng the chereceptrs (107). Cantha-n s a ptent eeng eterrent t nsects, th pssble rrtant prpertes(7)

These erent unctnal categres eensve cpuns can be -recte paly at specc grups preatrs, ther apparent bra spectu actn ntthstanng Entanglng secretns are anly ecentaganst ahp preatrs (, 63), an setes agans parasts

(60). Vetebrates are prbably nt very aecte by these secretns unlessthey are enrche th re tc cpuns (6). Ths type eensvecpun sees t have evlve st requently n slvng arthrps, unable t escape qucky ater havng release ther secretn anhghly epse t ant preatn, e.g. sl arthrps such as nychphrans(, 6), gle lpees (63, 1) gephlh an lthbrhcentees (10), an sps (C De Vrey, unpublshe bservatns)Vscus secretns are als ell nn n ther terrestral r sl anals,e.g. eahs an snas (, ). pheatng syph avae laybugs have a hgh prbablty encunterng ants an they t setesapt ths e eense (). In cccnells ree bleeng eters ants,even n thse speces that nt secret alkals, e.g. Aha obraar Rhobs ra J. M. Pasteels an C. De Vrey, unpublshe bserva-tns) he bl s very vscus n these beetles an clttng s quck heecency ths echancal eense aganst ants has been escbe netal r the secnarly phytphagus cccnell aha arss(6). Tertes are hancappe n ther ghts aganst ants, hch are prba-

bly ther an enees (6), by the relave clusness ther veentsan the slness ther reactns (70) It s therere nt suprsng thatbeses the essn vlatle rrtants lke nterpenes, ketnes, rqunnes (6, 1), stcky secretns parng the veent ants haveevlve at least s erent tes. Nasrms Trrrms an re-late genera (Terae, asuttertnae) rcbly eject r ther rntalglan a vscus slutn tepenes n nterpenes, rng a stckyresn (1) Coorms (Rntertae) pruces n hlgus glansa ture ucplysacchares an hyrcarbns (11) lers

oorms bas (etae, Macrtetnae) use ther salva reense an secrete a ture prtens an qunnes, hch, hen yng,raply beces stcky an rubbery (17). A sehat slar eenseechans has evlve nepenently n the prtve terte Masrms

Annu.R

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HMAL S AHOPOS 21

aw (Masoermiidae) (121). The soldiers of Glbrm lhr Termiinae) burs open when molesed, releasing opious, siksava sereons rom he rpured abdomnal wall (132). upure o he

abdomen is also observed in he workers of some soldierless ermies (Termiidae, Apoermiinae) ha release visous feal maerial aumulaedin large pouhes of he hindgu (177). Siky sereions are someimes usedby ans agains larger ans (1 1 4)

Volaile, nonspei irrans appear o be aimed mainl a arhropods,and possibl a mammalian predaors, and seem o be less of a deerren obirds. There are man repors of ans being repelled b hese ompounds(13, 21, 56, 207, 22). Several hings sugges ha nonspe volales areaimed pariularl a ans or oher arhropods Firs beause o heir smallsie he ans anno avoid having heir anenna bahed in noious vaporwhen he are lose o defended arhropods. Volaile deerren are ideallysuied o eep small bu numerous predaors a a disane Seond, henonspei rrians are oen produed in admixure wih more lipophii

ompounds, suh as fay aids, esers, or hydroarbons, whih onsiderablenhane boh peneraion hrough and spreading over he uile of hepredaor (see below). Third Blum (18) poined ou he simlari beweendefensive volailes and he alam pheromones of ans. He suggesed ha

some ompounds ould mimi he alarm pheromones, allowing he pre oesape while ausing pani in he ans. As wih enangling sereions,nonspei rrians are imporan defensive ompounds of ground arhro-pods, e.g. ground beeles, enebronids, and saphylinids (wih he exepionof ar spp.) (reviewed in 2 1 1) he sink glands of Heeopera alsoseree volaile irrians reviewed in 210), espeiall in speies ving on hesoil surfae (18). Oher arhropods living on he ground, e.g. millipedesand enipedes, seree nonspei irrians, inluding hdrogen anidewhih an also be a poen bu nonspei oxin (53, 65, 1 16). Ans hem

selves are major produers of his lass of ompounds (reviewed in 22, 23).Mashwi 1 14) ommened ha hese hemial weapons supplemen hesing apparaus and are more eien agains oher ans, heir maor enemies

Birds are someimes deeed by volaile irrians when hey are produedin large amouns or when he are spraed and reah he bird's ees (62,73, 7). In many insanes, however, he irrans are no spraed and heireeiveness may be far less The volailes produed by many Heeroperado no repel inseivorous birds reviewed in 190). Sawy lavae wih well

developed sen glands, like Nma spp, probably produe benzaldehde(139) bu an be heavily preyed upon b birds 6). The noodonid aerpillar rara = Crra) vla serees a 0% soluion of formi aid(183) bu is readily eaen b he blakbird Tr mrla The sereion



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272 PA, GGO & OWA

appeas howeve to dete paasitoid nsets (186) The aid seetion of thegound beetle yhr arab epels small mammals but not the ommnay Gar /aa (9);owee te bd was nee t nea te

eyes. It seems that most of the volates ae ost befoe eahing any sensitivetssue when the bds manpulate te pey Subsequent posonng aftengestion is unlely beause the eatve moleules would be quly alteedn te gut as suggested fo benzoqunones (65).

Spe toxns appea to be mu moe eetve tan nonspe tantsaganst bds and pobably othe vetebate pedatos Indeed vetebatesae moe senstve tan nsets to toxns su as alalods adenoldes ohydogen yande (8) althoug mpedes podung phenos and hydogen

yande ae pefeentay eaten by babets (Pfomes) and quals aeghly nsenstve to adenoldes (69) At least n Lepdoptea aposema-tsm s stongly oelated wth toxty (12) Most of te nset om-pounds atng as potent posons when ngestd ave been solated fomaposemati speies: eg adenoides n the monah buttey (56, 57),the pygomophid gasshoppe Poekilocerus buJonius (03) Aphis neri(7) ygaeds (87, 05) the tenuhid moth Syntomeida epilais (73)eambyds (54), and hysomelid beetles (43) bufadenoldes n ies(75) aistoloh ads n som swalowtal buttees (7, 04), py

olzdine alalods n atiid moths (68) and te gasshoppe Zonocerusvariegaus () st ohe aaods n oneds (44), some sode beees(4) and te staphyinid Paederus Juscipes (46) yanogeni gyosides zygaends (4599), cea and Heconius butteies (6), and the opald bug Lpocois iso/aa ( C Baeman et a submitted) and antha-dn in meloid beetles (49) Tese toxns ae often sequesteed fom plantsbut some ae manufatued by te nsets temseves (see above) We aenot awae o ases of nonaposemati nsets stoing sgniant amounts ofplant toxins fo selfdefense exept possbly aganst paastoid lavae (4)

The podution of toxins s pobably a peequste fo the evoluton ofaposematism (66) Waing sgnals whh an be leaned ae liely to bepatulay eetve aganst pedatos lie vetebates sne te onditoning o pedatos s ewading ony when the pedatos ae ew ongvedand fequently enounte pey Aposemat ooation is eognized aseient aganst bids and othe waing signals suh as sounds (39 7)odos (65) o taste (29) have been epoted Poisoning assoated wtwanng signals su as olos taste o odos is vey eetve fo theleaning of food avoidane by pedatos (80 36 25) Colo is a bette

onditionng stimuus than taste is fo bids but the evese is tue fomammas (24) moneys being a possible exepton (98). At least in atsodo avesions an be ndued f odo s st assoiated wth taste (36).Avesve ondtioning ous oten ae a sngle tia een when thee isa onsdeabe deay between ingestion and punishment (80).



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HEMAL DEENE IN ARTHROPOD 23

The loose age specici of defensive allomones could help o explainappaen edundances in soe defensve mechanss of ahopods oexample Ons fasas and possib ohe gaeids secee boh vola

e defensve adehdes (191) and cadenoides () Al ladbugs emisic blood doples bu on aposemaic species conain alaloids (144)The pohoacic gands of discids poduce seoids ha ae acive againssh and amphbian pedaos ( 1 19 180) hei pgdial glands secee hdoxbenzadehde and benzoic acid and vaious derivaives ha ae be-lieved o be dieced main agains micooganisms (19) enoic acid ishoweve an ecien sh epellen (88) and hese compounds have no beenesed on auaic insec pedaos Specializaion in esponse o dieenpedaion pessues could also explain some of he dieences in he defen

sive seceions of dieen insas of he same species (see below).

Blgal gae MxueWih ver few excepions defensive seceions appea o be emaablcoplex xues Ths s even oe evden when mno consuens aeseached fo ssemacal The defensve seceon of he saphlinid sla canalclaa conans no ess han 4 compounds 4 adehdes hdoquinones 1 quinone and 4 hdocabons (28) Gilb Waehouse(83) idenied 18 componens in he seceion of he sin bug Nza and Tschinel (19) idenied 13 compounds in he defensiveseceion of he enebonid Es bami and ised 10 addiional unidenied consiuens

This complexi is poorl undesood Snegism beween consiuenshas been demonsaed man voaie ans ae pola and hei eecive-ness is gea enhanced b moe lipophilic consiuens which faciliae hespeading of he compounds on he cuicle of arhopod pedaos and heipeneaion hough i (2 3 18) The nonl aceae pesen in admix-

ue wh fomic acid in he defensive spa of he aabid genus Hd geal inceases he sensaion of buing pain because ienhances he peneaion of formic acid hough human sin suggesing asnegisic eec on vetebaes as well (2) Soe consiuens could alsodela evapoaion of moe volaile ones polonging he acivi of heseceion (see 2) is hough ha he main funcion of onl sighl oxiccompounds such as hdocabons which ae commonl found in defensiveseceions is o ac as surfacans peneaos o evapoaor eadansbu some auhos have suggesed ha hese compounds could have a less

obvious bu ipotan defensive ole b inefeing wh cheoeepion(16 20 1)Defensive compounds ae ofen poisons fo living issues incudng hose

ha secee hem Consequenl he nal seps of biosnhesis ae ofenperformed in exacellula spaces o esevois (66 84 181) Some of he



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27 PEE GRGRE & RWERHER

cnsiuens f e defensive seceins may be pecuss bypducs fese nal eacins An bvius example glucse may ccur wen epecuss f e acive cmpunds f e secein ae gucsides (26 )

Te enymes caaying ese las seps ae emselves smeimes deecedin e secein and i as been suggesed a is culd be easn f epesence f e pein facins deeced in erwise nnpeinaceusseceins (2).

In many cases e divesiy f defensive cnsiuens wiin e samesecein emains uneplained and appeas unnecessaily edundanTscinkel (6) suggesed a "wiin limis e cemical naue f ecmpunds is f n gea impance as lng as e secein mees ceainpysical equiemens (lippiliciy vlailiy meling pin irrian

value). Tis culd allw muc divesiy a is n necessaiy f adapivesignicance Te cmpleiy f defensive blends culd als be a way avid cuneadapain by pedas bu ee is n evidence suppis ypesis

Intraspecc Variabty in Defensive SecretionsIndividuals f e same insa and se may pduce seceins diering inquaniy and qualiy [efeence in ( )] Sme vaain nly eecs sagesin bisynesis ( ) egeneain afe deplein (208). Seasnalvaiain is descibed in wae beeles and can be a leas paly elaed age dieences and e fequency f uiliain (8 28 2)

Geneic plympism in defensive cemisy is ply dcumened bumuans die in ei bisynesis f defensive cmpunds and in eisequeseng f plan ins (76 68) and cnsideable vaain can ccubeween vaus ms fD fm e same lcaliy (7)Inaspecic vaiain in cemical defense culd educe cuneadapain(see belw) as in plans ( 50).

Inepulain vaiain pbably eecs geneic dieences Te defen-sive seceins f dieen ppulains f enebnids (6) cysmeldaduls (3) and lavae () ans (7) and emies (50) die quaiaivelyand quaniaively. Te sage f cadenlides als appeas be dieeninvaius ppuains f e buery nus chsius even wen eaedn e same fd pan (7) n e cysmelid beele Chsoin opplains de in e TC paes f cadenides e deences e-man cnsan in successive labay geneains eaed n vaus spe-es f mn (3) N saisfacy adapaive eplanain as been ppsed

suc geneic dieences. Tey may be due a lack f specic selecivepessue peaps exemplied by e dieen pprins f dierpenefund in eee eu (150 e deenes fm a scyesin wen disslved in mnepenes (70). and eswic () suggesed



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N N RR 2

that "a hgher veral cncentratn f gle lecles can be btaned ftere a tre f trctral type A tre s elected f rather tanany ne prece cbnatn

exual eence n Chemcal eeneFeale aseatc edtera have been rerted t be re tc reetc than ales (30 112) These seal derences ay erely reect thefact tat a lage at f tn ncprated wt te egg eg na aja (10 and yaa a . Untl t yptes dsprven re cple evtnay elanatn ee neceay

In several nstances te abty f ales t yntee ns cndssees t be eater than n feales r eaple n the adlt spansh y

ya vsaora nly the ale are able t anfactre canthardn whch tred n te accesry gland f te ae seal rgans and njected ntte feale dring clatn (1) Te etatracc repgnatral gand better develed n se ae Hetertera tan n feae revewed n10) bt te reasn reans nclear Strngly drs sbstances aretanfeed by ale t feales dng atng n te bttery Hoao and rbably act a antaprdsac perne () It a al beenggeted tat the dr cld have a re general defenve fnctn 4)Scaled ahrdsac hernes n Ledtera are ften dentcal twellknwn nsect defensve cnds Ther re n cshp has beenestabshed n nly a few nstances 5) and they cld well have a defensvefnctn (11) Te prdctn f defensve cpnds by ales and tertranfer t feale cld pare te feale te ct f ter ynte anday rereent ale nvetent n te brd Mrever ter reeae dingatng cld rtect nsects n cla

eences n Chemcal eense eween Insas and Cases

In te chryseld tribe Paednn arvae secrete fr egental abdnal gland atgens vlatle cycc nterenes r sacyaldehydedeved fr the fd ant (141, 16) In cntrast, the adlt secreteanne glcsdes d crresndng ntrrnate derivatve(141) Bt the larvae and te adlts eat te sae ant arvae are lesscnspcs tan te apseatc adlt Thee derences n defenvecesty ay reect a derence n te natre f te an predatr (eeabve) The cptn f te setea ecretn f e wallwtalcaterllars changes a the frh ecdyss (3) fr a cle tre f

n and sesterends n vars carbylc acds and rlated estersThese checa changes are syncrns wth a clete change n appearance Yng larvae are crptc te ast ntar s aseatc The rle fcarbylc acds and esters s bscre Tey are eectve agant ant (1)



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276 PASTEES GRGORE & ROWERAHER

bu a leas in apo machaon hey are n necessary induce reecinf he caerillars by birds larae deried f heir smeerum are alsrejeced 97 Te dr f e smeeal secren s mre nense n e las

nsar 93). may ac as an addnal wang sgnal fr eebrae redars s remans ure seculan hweerMre dcul nerre are he derences bsered beween he la-

iles rduced by laral and adul eerera reference in 190)]. Freamle n Cx enes ans4c2nl cnsues ca 92% fhe laral screin bu 4he2enal accuns fr 95% f hesecrein f aduls 92). N eclgical inerreain has been gien frsuch a small bu sriing dierence.

he defensie deices f insec eggs hae been reiewed by Hinn 90)

Tns resen n e females may be ransferred e eggs smemesdurng sn. Tey aear be lnglasng feedng deerrens rcnac reellens n e surface f e eggs. The female meld beelecers he eggs wh canhardn receed during culan frm e male). Similarly Cule and Culesta msuies desi defensie uid neach egg 9 90).

emarkable dierences are smemes bsered n he cmsn f hedefense secrens f dieren cases f e same clny f scal insecs.

Thus in he ermie Ancstotemes cavthoa he labial gland secrein fe small sldiers diers frm ha f he large sldiers. The minr sldiersrduce he sesuierpene ancisrdial he majr sldiers secree he ismec ancisrfurn n admiure wh a bcyclc unsauraed daldeydesmall amuns f luene and a- and cyclgeranlenes 77) Thesesubcases ms rbably gae frm dere female sars dung s-embrync deelmen 3) Small slders usually accmany he fraging aries while he majr sldiers say in he nes 00) bu hislyehsm as n ye been relaed e cemcal cmsn f e

defense seren.

Cngn gn f Chmal fnn Symparc Spcsefense allmnes are usually cnsdered a he secies leel and cm-aisns are made n a anmic basis They hae rarely been cnsidereda he cmmuniy leel hs is necessary hweer if ne is undersandheir funcin and eluin.

Rhschild 6) suggesed ha he srng dr characeing many

asemaic nsecs culd ac as an addiinal waing signal fr redarsand ha Mllerian mimcr f drs culd be freuen Ths wuld redca cnergene f defense drs n symarc seces The drs f deren

Annu.R

ev.Entomol.1983.28:263289.


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CHEMICAL DEFENS IN ARTHROPOS 277

apoema beele e.g. ly eamby a oell ae hmcally ieical (125) We sugges ha coegece o oos woul be aoe whe he oo is isic om he mai oxi as is appaely uei may aposemaic species (125 165)

Paallel eoluio o eical eesie allomoes has bee obsee ompleely uelae pee (eeee 1). Coegee howee oeo appea o ou bewee oapoema pee. o eample heeesie seceios o closely elae chysomeli laae show isiclyee mes o mooepees ( 141) These ieeces ae o he sameoe o magiue o boh sympaic a alopaic species. Chemicaley wh a commuiy coul lim he abily o peao o oecome eeie allomoes. Habiuaio osh peaos o a gie hmial

has bee epoe a his shoul ao iegece o seceio amogsympaic specie (88) oeoe he peaceous sawy Tenthedo olivaceaca e coioe o seec as pey sympaic chysomei aae ieigpicpally hei eesie eceios bu o physial appeaae J. Paeels a C. Ggoie upublishe eul).

Phooal Fuco o Dv ScoDeesie alomoes ca easily u io alam pheomoes i socia ogegaiou iec I eeal gegaious eeopea (5 108 1 1 5) bu o

ohe (1 he alam pheomoe ha aue he peo o he ewhe aae ae he mao eee aomoe I aou eme healam pheomoes ae eesie mooepee (122 206). I eealaphis he waxy seceio om he coicle coais alam pheomoeseihe asaesee o gemacee A (7, 130) Aam pheomoes iocial Hymeopea ae usually pouce i eesie glas oe i cloeoecio wh he ma weapo he mable a he ig (15 1 1 3).

Wlo oe (216) peulae ha alam i be accomplihe byahe simple olaile compous ee a he expese o speciciy whichis olile impoace i his coex As see aboe he emissio o olailes also a eece chemcal eee aga ahopo peao a equely ecouee ahopo. Thu boh alam a eese haeimila physicochemical equieme

I highe Hymeopea eesie allomoes oe ale ecuis a aeemie i ohe coexs ha he eesie omic aci ecuis as iseeal omicies o oly uig eese bu also uig peaio (113).I he "leag pheomoe ee by ecue o amoo o

o gue emae owa oo (2). I eeal melpoe igle beehe maibula seceio is use o buil aeial ecuime ails leaigo oo (10) The epeic maibula seceio o he meipone bee



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PASEELS GRGORE & ROWELL-RAHER

ia la is an inraspei atraan bu it is used as an oen-sive deterrent during raids against Tiona bees (). Ants similarly utiliedefensive secretins frm the Dufur r isn glands while sacing therant nests ( 5)

Bergstrm Lfqvis () suggested tha ants culd specically labelforeigners or inuders with heir defensive seretions his ould palyeplain the compleity of some secretions like those of the Dufour glandsin whih speic minr cmpunds are ften obseved () his attrativehypthesis remains t be tested

Defensive sereins are also reprted at as assembling sens ingregarious insets he sereion from the pygidial gland of he tenebrionidB/a onaa ats as an alarm pheromone when produed in large

quantity tuing int an aggregative phermne after t (). Inthe stinkbug a rosa 2hexenal is an alarm phermne butacts as an aggregain phermne fr the rst instar when emitted graduallyin very small quantities (6) he dorsal abdominal glands of Dyini produe a sereion ha is suspeted o inuene aggregain() However evidene fr the artiipation of defensive substanes in theaggregaion f Heteropera has been quesioned () he soure andnaure of the assembling sens remain unknown in Ci llai (6)and in apsemati oreids () n he aanhominid bug laa ia

he ohesion of lavae and moher is assured by a rail pheromne releasedby the larvae (5)

ur knwledge in nly tw cases were defensive secretins reprtedt reel individuals f the same secies favring disersins he urbeetle Tiboli asan utiies its defensive quinnes t repel con-sei adults () Adult leaeeles Gaya iila are repelled byhe defensive seretions of he lavae his avids verrowding of inseson single plants and uld be highly adaptative in speiast erbivres() Many epideii pheromones are ontinuously deposied by her-bivores while feeding (referene in 5) or by some parasitids duringeploraion (eg 5). Cmmuniatin by pheromnes requires onlyminue amounts of he ompounds n onas high nenraion of hem-icals are discharged suddenly in defense. his ould limi the verlap between eidectic and defensive functins

Atopods Vesus Plant emical DefenseArhroods and plans ofen use the same hemial weapons for defense

(6). he bilogial satus f plans and tha of ahropods are so distinthwever that divergenes mus be epeed a least in the relative fre-queny of ocurrene f partiular mdes of protetion r in the ways both



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CHEMICAL DEFENSE N ARTHROPODS 9

group ue chemica for defene Plant can regenerate and urvive evenafter large par have been bitten o. A ingle wound can be fatal forarhropod but the are uuall able to ee and to control the emiion of

their defenive ecretion. Selective preure i much tronger in arthropodfor the development of a chemical defene eective at the earliet intantafter an encounter with a predator or even better before epellent ecre-tion (bet eved b volatile and waing ignal dominate arhropoddefenive trategie, wherea feeding deterrent [enu Dethier et al (48]and antibiotic are more frequent in plant. Ahropod defenive chemitri dominated b mall molecule tored a highl concentrated miture ineteal reevoir invaginated in the bod. Of the 30 terretrial Coleoptera lited b Weatherton Perc (21 1 283 produce compound with a

molecular weight le than 10Some rearable adaptation have been obeved i ahropd that

allow the predator to reject toic inect b tate onl. n the monarchbutterie cardenolide are more concentrated in the wing, which are thepat rt bitten b bird (30 Mot of the cardenolide preent in chro-melid beetle are concentrated in the ecretion that cover the pronotumand eltra oon a the are attacked The concentration of cardenolidein the ecretion ha been etimated to be a high a 2 M for Clrb (143 The bug OlJ emit doral droplet of uidin which the cardenolide are concentrated at the level of more than 10g/ l ( The rejection of unharmed apoematic inect b bird i vercommon and it ha been potulated that individual election alone ma beucient to eplain the evolution of apoematic coloration (7 2

Arhropod, with the poible eception of ome ocial inect are a farle predictable ource of food than apparent plant Quantitative defeneimilar to thoe of tree are onl epected in ome ocial inect egperennial termite and ant ocietie living in denel ditibuted apparent

net In termite like Nrm or Trrrm chemical defene ienured b terile oldier. Defene i their ole function; the are evenunable to feed themelve Their propotion vaie from 10 to 30% of theactive popuation of the ociet 6 About 1% of the annua productionof Trrrm m i devoted to the productionof oldier (101.The oldier produce a miture ofd and monoterpene a chemical defenequite imilar to that of coniferou tree (148. Each major oldier of Trrm contain about 3 1 g of terpene ie 8% of the frehweight (17 thi would repreent 4% ofthe energ equivalent (in joule

of the oldier a etimated b Joen for a ver imilar pecie (101 hiquantitative defene i eective againt verebrate predator. The earh wolfl r i peciazed for feedng on Trrrm b



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280 ASEES GGOE OWEAE

which fage in the pen in Suth Afica 10) Thi entmphagu haenid i n abe t captue ma numbe f temite at the ta fit attack,when the wke ti age utnumbe the die he we quick

withdaw, but die ae ecuted t the pace f ditubance The eahwf eem unabe t feed n die n, and i appaent epeed bthe tng cncentatin f tepene.

Quantitatie chemica defene i a pectacua in me ant pecieWke in the ubfami Fmicinae ae we knwn f the age amuntf fmic acid the te in the eei f thei pin gand. In EupeanFrmic pp. the cncentatin f fmic acid in the enm eache 61 t65% 135) Accding t the gue f Oman Bande [Tabe 1 in 135)],the amunt f enm pe we f Formic polycen ae fm 07 t

mg which eeent fm 95 t 3 f thei feh weight Unfunate, pecie data that wud aw a cmpan f inetment in chemi-ca defene cia and nncia inect ae acing

Chemica defene ha been ecndai t in man temitid, pehapbecaue it i ct efene i then tanfeed t inticate net tuctuefm which the inhabitant ae e icut t dientange (J eigne anJ M Patee, in epaatin) Peha the net mateia in tee peciei anthe anag t the quantitatie defene f appaent tee

ConluonBum 19) it 316 empiica fmuae f cmpund ued b ahpdf defene The tta numbe f knwn amne i een geate becauef the exitence fime Amng the ma mecua weight cmpundthee amne ange fm chemica a impe a hdgen canide fmic acd cadac gcide aad a cmpex a peden

In thi eiew, we ugget that, depite thei geat dieit, patte fccuence f defenie cmpund can be ecgnied. The ditbutin f

chemica defene depend n the "appaenc enu Feen 8) fthe pet thei maj pedat Mnphagu igphagu hebie ma beme pedictabe f pedat if the a ue pant cue in thei eachnect feedng n heb ma equete txin fm thei fd pant, andcumped ditibutin euting fm the patchine f the he inceaethei appaenc and fa the eutin f apematim b kn eectin

Ntwithtanding thei appaent bad pectum f actin, defenie cmpund ae nt equa eectie againt the aiu tpe f pedatEntanging ecetin ae eecte againt ma athpd pedat Va-

tie nnpecic iitant ae eectie againt athpd pedat, andmabe againt mamma, but e againt bid Entanging ecetin aeexpected in wming anima and atie itant in fate ne pe



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CHEMICAL DEFENE IN ARTHROOD 28

cic xins pbb h ms ci chmicl dns ins bs which cn ddiinll b cndiind b wnin sinls Appndundnc in chmicl dns dincs bwn inss m b

pl xpind b his spcicimpxi dnsi mixus is pl ccund b h nd snism bwn cmpunds nd pl b h psnc bisnhiclbpducs pcuss In sm insncs mixu cmpunds wihcin phsicchmicl ppis is scd ; his lws ibili in h pcis cmpsiin h dnsi scins Sxul dincs in chmicl dns cn b inpd in p s din d pducin insmn bwn h sxs

nnc in h win sinls smpic spcis cn ccu, bu

cnnc in h xins hmsls is impbb inc dnsiscins, impin pssibl dpin b pds culd pl plinh cmpx sm scins nd hi inspcic in in smp-ic spcis

Sm dnsi mpunds h ddiin bili mins, sphmns hs uncins mus ls b cnsidd whn uin hcic snicnc chmicl dns

unit dnss nus hs und in pnnil s und in sm scil inscs in which h insmn in nnspcc xinsis cnsidb Whil cmpn h chmicl dns pns nd thpds, w sus h h mphsis n insnnus dns is much h l This xplins wh lil ins nd winsinls, sscid wih xins, dmin h chmicl dns h-pds

Th nizins ppsd in his iw smims bsd nindic scc inmin h shul b cnsidd s huisicpdicins ndin cnmin mndmn A b knwd h

pcis md cin dnsi cmpunds is unl ndd, p-b wihin n hli mw in which pms suh s hppnc nd pdicbili p spcis und nul cndins pln imptn

AK

W hnk D. Mc nd . Rwl hi inub iw nd

ciiu h mnuscip M S Bum, M Rhschild B W Sddn simulin discussin nd usu inmin nd mn uhs -lwin us qu hi suls, unpublishd sil in pss



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CHEMCA DEENSE N AHPDS 283

ion of ha so i th monarch buttry Scence 889

3 Browr, L. P, v Zandt Bowr Covo, M. 96 Plan poisons in a

trrtra fo cha Nat cadSc U 893983 Brow, K S. Jr 98 Insos

aposmtos dcador naturas dpant micas. Cenc. Cul SaPaul 3289 (Supp.)

33 Brown W L. Wlon E. O 99 hvouton of h dtn n. Q RB 389

3 Campl, B. C., Dy S. S. 99Tomatn and piic waps: Potntalinompabili of planibiosis wihbolocal conrol. ence

3 Carayon, 9 Not t dumntssu 'appal orant metahoraciqd Hempr. Ann. S Entm (NS 3

36 Cardani, C. Fugant C. GhirghlD., Grass, P. 93 h biosythsof r. Tethedn et 88

3 Crr E Enr, T 9 Cnthardn Pot fng dtrn o nsts Scence 83

38 Carrl, J E. Thompson, W, Macaghln, M 9 Parnta rnsmis

sion of a dfiv chmica (canhadn) in bsr t. So 38

39 Clardg M 9 Sdlation anddfnsv bhavour n h gond t Ch cade (L). EntmS A 9

4 Cran 9 Imagnal havior of aTrindad buy Helcnu eathaa wison with sia rfrnc to th sal u of color. lgcaNY 696

uot L 896 Su a sai rx

l moyn d dfns d qlqun h 68 Cro E. 96 e Ethl da

tn. Br Spingr-Vrlag. pp.3. aoz . Ps J. M 99 Pro

dcion of adac gycosids by chrysom bt and lv Che c63

Davi, R. H., Nahrst, A. 99Lnamarin ad lotausan hsourc of cyand ygaena lendulae L. (pdoptra) Cm Bche Ph B 6399

Davis R. Nahrs A 98 Occurnc d variation of h cyaognic glcosid lin and loasrain s of h yganida (Ina Lpdopra). Cm BochePhy/ B 393

6 Dgn, Quny, A., Bum, M. S98 h mi nd dfn mhasms of tmt. I Scl Inect, . R. mann. Nw Yok Acadmic.

9 pp Dr C., Ps J M. 98 Dribon of adul dfns gds chysomds (Cooptra Chrysomlda) d is scanc th voluonof dfns mhansms wth h fy. Che c 868

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Forpersonaluseonly.

past eels 1983

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