Annals of The Entomological Society of America ‘Volume 72 ‘SEPTEMBER 15, 1979 Numbers The Annals ofthe Entomological Society of Ameria introduces with ths issue a new featare, The Forum. Like The Forum in Environmental Entomology, this one will publish interpretative and spec ‘lative articles and, indeed, ftom time to time, controversial ones. The Forum will appear as often as suitable manuscripts are accepted under the usual review process. We urge anyone who wishes to spec: ‘ulate, to argue, to ponder i do $0 the Annals; and we look forward eagerly to these manuscript, and to the excitement they will bring tothe Annals Forum ‘This section of the Annals of the Entomological Society of America is reserved for in- texpretive articles dealing with the broader manifestations of entomological activities, and the basic scientific approach to these activites. Dissenting viewpoints may be published subsequently if they meet the editorial criteria of the Annas. On the Original Feeding Habits of the Hemiptera (Insect we svar stn nS 78 Pagers pean br ly ae ue teal rahe than cations and yropow es proposed oy Cate wRE sce ot Seperate caret fedora eet St ne Cpe tc Stn Cage a See ar tft eee ated Ris recreates 9 een Mabe mec eee er nage peter toate re SS Cement eee ete eee See Recreate en ae ES ein ecatne en ab tana SEES es cnt ee eee nh epee SRE cca ofa cea econ apr pan cee nt near Ae eet Sioa eel sah station nee eke Bers Re ini aes CCobten (1978, in his reset compechensive worth suborder Heteroptera ad perhaps also ofthe subor- Evolutionary tends in Heteroptera: Pat Meutipurt. der Homoptera, He bas Ke conclsion on the wide structures and feeding satis," argues that can. spread presence of camivory among the heterplerous ‘ory not herbivory wa the original feoding suatgy of iacrder,ineloing the Digsocormorph, the Falco (©1979 Emiomological Society of America (0013-8746/79/0S05-7505$00.75/0516 (Amphibian) which he boise oe the pi {ive sek ofthe Heteroptern Many Cinicomoré, But {ew Penatomomorpt re carvers. Coben as a ‘cs dat the Heteroptera evolved in a semantic Shorcine type habitat sce hss where the more rims |tve Garmorpha now oct, especialy the Heb. Thave no qual bere with Cobben's dete and ex: celle sues; however, it should te noted that his Se {Sons about prinive al svanced sates often hinge fn is concen tht the Oxo ae primi. “Moreover the lage amocnt of prt angen ad convrgsat evolution tat Cobben i willing to scope takes ful to aceopt hs phylogenetic ani tring fee of bia, post ho, ergo proper hoe. ‘This bias in favor ofthe Geromorpa ss being the ast plesionnphic ofthe Hetroper seems cider a Cobbens ating ofthe rate Conon othe het ‘oper archeype (Caen 1978, p. 250) OF hese, the folowing appear f be ster coadaons of th Ge. romorpa per sear ae cold wih the cao. fous habit ad semi aquate babi ofthe Gemomor pba: (1) ygroptlus, ing inthe ite ae: 2 al Revie female wih secondary econ cana @) ferale witht a iacniate vip (3) epg epoited ‘supricial: (6) sid chorion withou pores for srg stmowphric ar) presence in egg of posterior ser ‘al Ryopyle; @)lavee with ooe tama segment, ‘dls wid 2 taal segments; (head wih 3 pas of Tcobeas (10) song bated non ‘rere (or order) are suicly homelogous, i follows fen ta the sheath vas lot inthe agua and sem ‘aquatic lineages probaly as a consequence of thee fend ing mechanisms the mani ze isrted rapily with ‘he mandibles fonctoning t hold the prey (Coton 1978, p13, 33. “Another argument fora herbivorous ain of the Het- croptera concerns the evolution of the compe hemp ‘eroas moutpars. Cunivory is «commen phenomenon ‘mong insects bat when i evolves, as seen a Odonata, Plecoptera, Newropira, and Coleoptera, eg, st i volves mainly the modification of chewing mouthparts forlaeral biting sharpmaniblesanor mails athe ‘than sleader proboscis fo unidirectional preing. Pr lel evolution of uniecuonal pring sucking moat pats is seen csewhere among parasite insect feeding fn 8 lage hos (Anopora, Siphonapea) a, as espe cially seen among the Diptera, are modifications of ower-feeding mouthpas. is ot clear to me how the complex pering sucking mautparts of the Hemipiers ould have gradually developed through earivory. Sim {any fhe Homoptere had ino evalved, ss Coben ‘sugges, from carnivorous ances, how ould these lncestrs have been effective carivares without «gus? eis aking oi too much to evolve and thea Tose the ula especialy a pytophagn Heep bve re envision the evoltion of feeding habits among the ‘Hemiptera much s seen by Goodehid (1966 nd Miles (1972), except that place much weight on x poytoph: fous ancestry. As is well known, the Hemiptera ae Strongly relied othe order Pacopera and Thyeoop- tera by the lak of cer, los of juve oell, cosa. ‘ion of tie nervous system, preseace of few map Swarr: Oucinat Hisaprena Feanine Hanne sn ian tubules, reduced tral segmentation, and 2 well ‘developed ciara sacking pump. While some members fae caaivorous, both ores ate predominalybebiv- ‘tous. Procheriterans much like most sysanopterans could use thr mouthparts to cut ato plas ster and ‘he the eibaral pump fo sock up lige food. From this bepinaing the complex hemiperoos mouthparts could evolve gradually sep by step. perhaps conomitanly ‘withthe evolution of coraceous epidermal plant asbes ‘moog the gymnosperms plants nthe ltr Paes ‘The slvay sheath id evolved at this ine Ta this scheme the protbemipteran ely divided into 2 main ncages representing alspation to? diferet ‘ypes oF plant feeding. The fst neage, which went oo toform the Hemopter,pesTected driling ino he phloem effectively and asa resi the beak rises early between ‘he frcleps to fctate downward dling io vat ‘lar plant sues. This taps ch food our Bu ne Inter low in pretcin, hence st homopterane must fi tera large amount of nivogen-por tap to obtain su ‘ent amin acide and ther nuttonal neds (Goodchild 1966). The 2ad lineage, which evolved into the Heter copter, developed a quite diferent feeding sty. Rather than sit directly upon the plant sues and dl somewhat lindly into sues Benet, the prose foptras searched for tsves already sch in aval nivogen-spical meristems, bods, fruits, seeds— ‘xplting a basically erent fod from that of hom ‘opterans. Searching and prcbing for atropen ich fod ‘2 up Selection which brought the mouthparts forward by elongating the bead, thus foming the gla. From his positon the fsrum could be extended forward ud wed 5 see organ in conjunction wth the eyes and ate ‘ae fo cuamine the plat meri oc ikely feeding ‘prs, much as seen now among modes Penttomomer ghePeoaonie. Care, Loess Phar. ‘After these morphological changes occured, the an- esl hetropteran coal feed ot only on mien lant material bat could also explo itogen rich seasile Sima foed such as eggs. Digestion of proteinaceous Plant issue readily lend self to the digestion of imal Protin. At fist, mach as fequenly occu now, sy {imal food sequred was nitogen heh supplement ‘Such opportunistic carver is very common aman te Peaatesomorph toy, aswell at among the Mia, ‘he plan bags (Wheeler 1978). ‘Once this evolutionary stage is reached, the hetero ‘eras ould aow readily eater the ew adaptive zoe of ‘ariory by evolving the necessary speciation. In aly, as Coben (1978) seses was the tse among the shoreline Genomoepis, timid camiores eval, ‘which could cape inactive peey. Later hetcroperans ‘would evolve the comples morphological nd behavior Adaptations necesary to cape active prey, bah on Plants and onthe ground ‘Several early inages which specialized for cara- vor, the Enicocepalomxpha and Dipsocoromexph, Survive today a very ld carnivorous shoots whore Feationhips wih other Hetroptea are highly probe iti. Net, one or moe major incages valved i d- ‘pation to searching for grey in the wash-p habia and ‘averse tothe Lepepodemorph andthe Geromer,sm ‘which imately adapted tothe surface fi, and tothe [Nepomorpha, which adapted ultimately ino swimming forms. The plesiomorphous phytophagous sister groups ofthe shove carnivorous groupe were eventually clin ‘ued by competition though te evolution f more ad ‘anced tna of phytophagous insect [a this view the, ‘he Geromorpha form nt the peslomorpie stem group ofthe Hemiptera buts highly specialized although ar ‘hae lnege. ‘Within the landbugs 2 main phytophagou lineages sve. One is the hiphly advanced Penatomomorpts, ‘which are neverelese the diece descendants of the lant feding tm. The dificly of elang the existing Pewatomompan groups, the Aradciea and the Te ‘hophora to each oer and together othe other Het ‘pte ead understandable: the ogal sock of ie Penttomomorph, at well asthe egal stock of the Heteropers became extinct trough competion ith later Hneages. This appreat extensive extinction i Sup ported by 2 line of evidence the complete lick of any ‘Shmupomorphous comectns beeweea the Penatoa- ‘morph apd any oh subgroup of Heteropters; and Po- povrand Wooton (1977) have desebed fom the Lisle (Of Europe 2 large umber of unusual petatomomor- halite bugs, which resemble o ext famlis and Indicate the anquty of tis lineage aswell as that ex- tensive extinctions had ocsred ‘The 2ad phytophagous lineage is ho miPiaingid ciicomerpban ince. Tis lineage shows evidence of being derived from carnivorous ancestors because Fst, ‘the slivary uid is absent second. the acrate-ash feoting on mesophyltasses highly toxic and way leaves nerai pot on leaves Miles 1969, Tingey and Pilemer 1977 and hed, mos cinicomorphan amis ‘ae camvorous. As Wheeler (1978) noes, he Mirdae themselves are fequenly carnivorous, andthe Isome- topiae,evidely tie plesomorp sister group of te Mirae, ae predaceoor on scales (Wheclr and Henry 1978), whieh adds support to the crivoros oigin of the Mirae andthe Tnghe. OF couse, ii equally conceivable that cimicomorphans were originally lace fe and fash feeders om plans and the salivary sheath ‘id was an early loss. Io, then this feeding method really gee ise tothe many caivoroos fies of the CGmicomorpha wile being retained in the Mids, Ti tae and the Thamastocrda, In any cas, te lrgely Dytophagous Mindae comprises the largest family moog te Heteroprea, which atests to the success of this feeding meted "The scenario proposed hres evluionaly sting Jn hat it permis changes in head suuctures to occu ‘small increments and doesnot require salary vols tion. The preence ofthe salivary sheath fad in te Pentatomomerpha and Homoptera would be a sag ev. ‘luonary ever which accounts forthe close chemical Similarity of the sheath Nid in the wo txa (Miles 1968) Coben (1978, p. 251) in contrast, expais the ‘heath fads being a consequence of anagenetic con ‘ergent evan. That, the seat was ether lost and revolved in te Heteropiera or it evolved by pull ANAL of THE ERTOMOLOGICAL SOCIETY OF AMERICA Val. 72, no. 5 ‘the amino acids ofthe sath proti. Iti, eveutionally speaking, easier to love complesatrbutes tha to git ‘hem, and complex charaters, ocelot re rarely re ‘sined inthe same erg frm. The absence of shea, $id from many Hinagesof Heteroptera would this be best explained by apomarphic losses related 0 adapt tions 1 canivary; the posence, inthe Penatomenter: pa, a symplesiomorphy withthe Homoptera. The re feation of the salivary sheath in the carnivorous Pentcomomorpha may be a consequence ofthe feeding swexbann of he insects In bat theIygacld Gescar- Shae and the peistomoic Atopimae (Sect 1968) the ‘axillae and mandibles eter togeer as inthe phiyto- fagons lincagee and the sheath eves suppt poi fer penevation, ain sc! feeding, where the shut is laid down only at the vrfer (Miles 1968, Saxons 1963), and doesnot sound the set paths as occurs ‘with vascular tse edi. Finally it may be poised out that this seqoene of evolution ins is parallel mong the Cooper. The ‘Adephaga ae cleat mor peimisve tian the Plyphags which are largely phytophagous although many fais Ue seconduly caravoros. Hecase of ts, i as Log ‘hovght tthe Coleoptera is oigas in carnivorous Invests. Cromnon (1987) demonstrated thatthe Myx0- ‘haga andthe Archeostnma are considersby more Primitive and tht these frm were phyophagoss or ‘iycetophagous. The scess ofthe Adephaga owes i Seif to an early arquston of effective predton which ‘ermited tes insets nt only o survive sucessfully in competion with te Polyphaga but also allowed the ‘Asepnga to saccesfly Invade the agease abit ‘much af did he Geomorphn and the ‘Nepomompa sanong the Heteroptera [REFERENCES CITED ‘Cobbes, RH. 197. Evduscery Trend in Heeroper Patil Metatarsal Feoing Soin. ADE es. Repos, Wages 1-5 1-0 ‘Cromon, He A. i967 The Natal Cantino te Fa "Jes fCacopen- BW. Cay La Hanon, Me Goodall A. 1. P. 196, Bvt of the iment carl ‘nthe Henge, il Re. #1 6-10, ‘es, PW 186k Sai ome alvaypyslogy of un bas: Th hey of frat of fe eh {ei Ins, Pysal 10 147-60, 196, Tse secretin ia plane. Ase. Rev. Phytpabl 199 ren lt es a rye ihe fenslpbeves pn and HempursEstnal yp. Appl 2 Test 97h Thea of Heit. Adv, Inet, Phys. 9 fre Popor, Ys Ay and R. J. Wontn. 197. he upper Lise Racopre of Mae aed Sasy. Sit Zc ‘ek EF i971 The gine es, Pro. 13 i. Cons ral, Mozow 162993. ‘Sanen, KN 196. The oe of ingen ina etropcus Inst, Dysart hoe (2) Pyne). Ts Pay 87Sweet: Omnia Hesarréea Fosbive Haars oo Sweet M.H. 1964 The iog and csogy of he Raya. Whee, A. A og ee sigh wee ee nsotae E ne ep a ee Tesmsce sa peopel me fog aay Ball ‘Ema. Soe‘Am. 2327-7 EE A ty ee eae dat Lieve rate See Skee Saree ae Reprinted from the ‘Awxas op ie Ensowaocteal Sorry o Axeatca