Performance consequences of food mixing in two passion vine leaf-footed bugs, Holymenia clavigera (Herbst, 1784) and Anisoscelis foliacea marginella

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1 Performnce consequences of food mixing in two pssion vine lef-footed ugs, Holymeni clviger (Herst, 1784) nd Anisoscelis folice mrginell (Dlls, 1852) (Hemipter; Coreide) Rodrigues, D. *, Durte, LS. nd Moreir, GRP., Progrm de Pós-Grdução em Biologi Animl, Deprtmento de Zoologi, Instituto de Biociêncis, Universidde Federl do Rio Grnde do Sul UFRGS, Av. Bento Gonçlves, 95, Bloco IV, Prédio 43435, CEP , Porto Alegre, RS, Brzil Progrm de Pós-Grdução em Ecologi, Instituto de Biociêncis, Universidde Federl do Rio Grnde do Sul UFRGS, Av. Bento Gonçlves, 95, Bloco IV, Prédio 43435, CEP , Porto Alegre, RS, Brzil *e-mil: Received Septemer 27, 25 Accepted Octoer, 25 Distriuted Ferury 28, 27 (With 4 figures) Astrct Holymeni clviger (Herst) nd Anisoscelis folice mrginell (Dlls) (Hemipter: Coreide: Anisoscelini) re distriuted in southern Brzil nd use vrious pssion vine species (Pssiflorcee) s host-plnts. Preliminry oservtions indicte high coexistence of these species in terms of host-plnt use; in ddition, there is strong similrity regrding egg nd nymph morphology. In this study, the most suitle feeding sites for nymph performnce on wild (Pssiflor sueros Linneus nd Pssiflor miser Humold, Bonplnd et Kunth) nd cultivted (Pssiflor edulis Sims) hosts were determined y rering them on ech host nd on the comintion of hosts. Performnce ws determined y evluting nymph development nd survivorship, nd dult size t emergence. Plnt prts used were lso recorded. For oth species, P. sueros ws the most suitle host plnt. First instr nymphs of oth species fed on terminl uds more frequently when compred to other plnt prts. Second instr nymphs switched to green fruits, whose ehvior ws more pronounced for H. clviger. Thus, H. clviger nd A. folice mrginell immtures re extremely similr in terms of host-plnt use nd consequences for performnce, in ddition to their morphologicl similrity. We suggest tht these coreids my hve evolved through severl processes, including prsimony etween the immture stges fter specition, evolutionry convergence, mimicry or genetic drift. Keywords: Coreide, pssion vines, host-plnt selection, immture stges, nymph performnce. Conseqüêncis d diet mist n performnce de dois coreídeos do mrcujá, Holymeni clviger Herst, 1784 e Anisoscelis folice mrginell Dlls, 1852 (Hemipter; Coreide) Resumo Holymeni clviger (Herst) e Anisoscelis folice mrginell (Dlls) (Hemipter: Coreide: Anisoscelini) ocorrem no sul do Brsil e são hóspedes de diverss espécies de Pssiflorcee. Oservções preliminres indicm um lt coexistênci dests espécies em termos de uso d plnt hospedeir; em dição, ovos e ninfs são extremmente semelhntes. Neste estudo comprou-se performnce ds ninfs ds dus espécies em pssifloráces ntivs silvestres (Pssiflor sueros Linneus e Pssiflor miser Humold, Bonplnd et Kunth) e cultivds (Pssiflor edulis Sims), oferecids em seprdo e conjuntmente. Form determinds s txs de desenvolvimento e mortlidde ninfl, em como o tmnho do dulto à emergênci. As prtes ds hospedeirs mis utilizds pr limentção form registrds. Pr mos os coreídeos, P. sueros conferiu melhor performnce. Ninfs de primeiro instr de ms s espécies limentrm-se com mior intensidde d região picl dos rmos qundo comprdo com s demis prtes ds plnts. Ams s espécies mudrm pr os frutos verdes no segundo instr, sendo este comportmento mis pronuncido em H. clviger. Deste modo, lém d semelhnç morfológic, os imturos de H. clviger nd A. folice mrginell são extremmente similres no tngente à interção com sus plnts hospedeirs. Sugere-se que estes coreídeos possm ter evoluído trvés de vários processos, entre os quis prcimôni entre s fses imturs pós especição, convergênci evolutiv, mimetismo ou deriv genétic. Plvrs-chve: Coreide, mrcujás, seleção d plnt hospedeir, estágios imturos, performnce ninfl. Brz. J. Biol., 67(1): 91-99, 27 91

2 Rodrigues, D., Durte, LS. nd Moreir, GRP. 1. Introduction Performnce studies re importnt for understnding life history trits nd their dynmics. For phytophgous insects, this pproch provides informtion relted to different trophic levels (see Arhmson nd Weis, 1997; Singer et l., 24). The outcome of the performnce of given insect my result from physicl nd/or chemicl plnt properties, or my reflect selective pressures imposed y oviposition ptterns, nturl enemies (see Bernys nd Grhm, 1988; Thompson, 1988; Arhmson nd Weis, 1997), or the evolution of insect speciliztion (Joshi nd Thompson, 1995). Phytophgous hemipterns re commonly not restricted to vegettive or reproductive prts of their hostplnts, nd sometimes studies of performnce hve fced difficulties to mintin the nutritionl qulity of vegettive prts tht they require. As consequence, most informtion ville refers to seed sucking species, mny of them of economic importnce. Studies with coreids include nymph development time of nymphs feeding on plnts of no economic importnce [Leptoglossus gongr (Amrl-Filho nd Storti-Filho, 1976), Venez stigm (Amrl-Filho nd Cjueiro, 1977), nd Crinocerus snctus (Amrl-Filho, 1986)], or on host-plnts of economic importnce [Phthi pict (Amrl-Filho, 1981) Leptoglossus zontus (Pnizzi, 1989), nd Ans tristis (Bonjour nd Frgo, 1989)]. Holymeni clviger nd Anisocelis folice mrginell (Anisoscelini) re distriuted in southern Brzil nd use the sme host-plnts s feeding resources (Pssiflorcee) (Schefer nd Mitchell, 1983). Some individuls were found feeding on oth wild (e.g., Pssiflor sueros Linneus) nd cultivted species (e.g., Pssiflor edulis Sims) (Lim, 194). Although nymphs of these species re morphologiclly very similr (Rodrigues nd Moreir, 25), their ontogenetic trjectories nd dult morphology differ significntly (Rodrigues et l., 25). Few dt re found in the literture, which include notes on nymph mortlity of Anisoscelis (Amrl-Filho, 1981), nd of Holymeni histrio, showing high nymph mortlity on different pssion vine fruits (Bldin nd Boiç-Júnior, 1999). Therefore, we determined nymph performnce of H. clviger nd A. folice mrginell on three pssion vine species ntive to southern Brzil (Rio Grnde do Sul Stte - RS): P. sueros, Pssiflor miser Humold, Bonplnd et Kunth nd P. edulis. The wild form of P. edulis is rrely found in RS Stte, nd it is restricted to Mt Atlântic ptches (northestern RS). Both forms re, however, similr in size nd shpe (C. Mondin nd G. R. P. Moreir, UFRGS, unpulished dt). The plnt prts used during the ontogeny were lso recorded, since the high moility of these species mkes the use of severl prts possile within given host plnt. 2. Mteril nd Methods 2.1. Insects nd plnts H. clviger nd A. folice mrginell dults were field-collected from the Porto Alegre (3 5 S nd 51 1 W) nd Eldordo do Sul (3 8 S nd W) Municiplities, RS. Adults were kept in cges (2 x 2 x 93 cm; n = 12/cge) under controlled iotic conditions (14 hours of light: 1 hours of drkness; 25 ± 2 C). P. sueros nd P. miser shoots were otined from pssion vine grdens t the Deprtmento de Zoologi/ UFRGS, Porto Alegre Municiplity. P. edulis ws otined from cultivtion locted either t the Centro de Biotecnologi/UFRGS, Porto Alegre Municiplity or Estção Agronômic/UFRGS (Eldordo do Sul Municiplity). In order to void inducing responses nd/or feeding preference, dults were fed d liitum with shoots ering oth vegettive (stems, terminl uds nd leves) nd reproductive (flower uds, open flowers, green nd purple fruits) prts of P. sueros, P. miser, nd P. edulis. Shoots were plced in plstic ottles provided with 5 cm high, wooden frme supports (n = 3/ottle), nd replced when wilted. Tp wter plced on Petri dishes with wet cotton ws lso offered to the dults. Cges were checked dily, nd eggs were collected nd plced on Petri dishes, lined with moist filter pper Experimentl design Freshly-htched nymphs were rndomly distriuted on ech host plnt (= tretments: P. sueros, P. miser, nd P. edulis) nd on the three hosts comined (mixed tretment). Immtures were fed s descried for the dults, nd ll nymphs of given tretment were plced in single cge. Nymphs (n = 2) were mrked dorslly (thorx or domen) with dots y using enmel pint (Fer Cstell ) of ten different colors: white, yellow, ornge, pink, red, green, ple lue, lue, rown nd lck. As result, nymphs hd the following identifiction: nymph #1 = mrked with white dot on the thorx; nymph #2 = mrked with white dot on the domen, nd so on. Body structures such s wing pds nd repugntory glnds were voided in terms of pinting. We knew from previous studies tht nymph mortlity would e high, ut pproximtely equl mong tretments during ontogeny. Thus, to mke possile comprisons regrding development rte nd size mong tretments, nymphs were replced in cse of deth; s consequence, ech cge (= tretment) hd pproximtely twenty nymphs throughout the experiment. Nymphs were oserved dily. Nymph development nd survivorship, nd dult size t emergence (the ody length, from tylus to the lst dominl segment) were recorded. Plnt prts used during the ontogeny were lso recorded twice dy (9 nd 15 hours), nd ech oservtion lsted 3 minutes. When nymphs were rered in the mixed tretment, it ws recorded whether the sme individul hd used different plnt prts Sttisticl nlysis First, development time nd dult size were nlyzed seprtely. The normlity nd homocedsticity of vri- 92 Brz. J. Biol., 67(1): 91-99, 27

3 Food mixing effects on coreids nces were evluted using Kolmogorov-Smirnov s nd Brtlett s tests, respectively (Sokl nd Rohlf, 1995). One-wy ANOVAs were used nd, in cse of significnt difference, tretments were sumitted to Tukey- Krmer s (in cse of homocedsticity) or Dunn s (in cse of heterocedsticity) multiple comprison tests. All tests were performed using n lph =.5, on GrphPd Prism softwre (Motulsky, 1999). Secondly, performnce ws treted s multidimensionl fctor tht considers ll life history trits here recorded - nymph survivorship, development time, nd size t emergence (sensus Thompson, 1988). The effects of tretment nd coreid species on nymph survivorship ( qulittive vrile) using ANOVA with Rndomiztion Testing (1 iterctions) (Mnly, 1991) were evluted. Then, the effects of tretment, coreid species nd sex on development time nd size (quntittive vriles) were evluted only for the individuls tht reched the dult stge (ANOVA with Rndomiztion Testing; 1 interctions). Both tests were run fter clculting Eucliden distnce mtrix etween groups of smpling units, using MULTIV softwre (Pillr, 21). Develolpment (dys) Survivorship (%) Results 3.1. Nymph survivorship Less thn 5% of the nymphs of H. clviger nd A. folice mrginell survived until the dult stge in ll tretments (Figure 1). Effects on nymph survivorship were not significnt regrding either tretment or coreid species (Tle 1) Nymph development time H. clviger nymphs grew significntly fster when rered on P. sueros thn on P. miser (Dunn s multiple comprison tests, p <.5), nd on P. edulis (Dunn s multiple comprison tests, p <.1). Nymphs rered on P. edulis grew significntly slower thn those rered on ll pssion vines together (Dunn s multiple comprison tests, p <.1). For A. folice, P. sueros provided significnt fster development time thn P. edulis (Tukey-Krmer s multiple comprison tests, p <.5). Nymphs rered on P. edulis lso hd significnt longer development time thn ll pssion vines together (Tukey-Krmer s multiple comprison tests, c <.5) (Figure 1) Adult size For H. clviger, nymphs were significntly lrger when they fed on P. sueros thn on P. miser (Dunn s multiple comprison tests, p <.1), nd on ll pssion vines together (Dunn s multiple comprison tests, p <.5). For A. folice mrginell, dult size ws not significntly different mong tretments (ANOVA, p =.8892) (Figure 1c) Performnce For oth coreids, there ws tretment effect on nymph performnce (Tle 2). P. sueros provided su- Body length (mm) c P. sueros P. miser P. edulis mixed 7 Tretment Figure 1. Nymph survivorship ), development time ), nd dult size c) of H. clviger (open rs) nd A. folice mrginell (closed rs) (men + stndrd error) rered on different pssion vines lone nd in comintion (= tretments). Aric numers ove rs indicte the totl numer of dults otined per tretment. perior performnce thn P. miser nd P. edulis. Nymphs rered on P. edulis hd significnt poorer performnces when compred to the other tretments. There ws no effect of coreid species nd sex on nymph performnce (Tle 2) Use of plnt prts For oth coreids, progressive use of reproductive structures of ll pssion vines s nymphs developed ws detected (Figures 2, 3 nd 4). Fruit feeding ws less pro- 6 1 Brz. J. Biol., 67(1): 91-99, 27 93

4 Rodrigues, D., Durte, LS. nd Moreir, GRP. Tle 1. ANOVA with Rndomiztion Testing (1 iterctions) of the effects of tretment nd coreid species on nymph survivorship of H. clviger nd A. folice mrginell. Source of vrition Comprison Sum of squres P Tretment etween groups P. sueros P. miser P. sueros P. edulis P. sueros mixed P. miser P. edulis P. miser mixed P. edulis mixed Coreid species etween groups H. clviger A. folice Interction coreid pecies x tretment Tle 2. ANOVA with Rndomiztion Testing (1 interctions) of the effects of tretment, coreid species nd sex on nymph performnce of H. clviger nd A. folice mrginell. Asterisks indicte significnt differences etween sources of vrition nd their corresponding comprisons. Source of vrition Comprison Sum of squres P Tretment etween groups * P. sueros P. miser * P. sueros P. edulis.175.1* P. sueros mixed P. miser P. edulis * P. miser mixed P. edulis mixed * Coreid species etween groups H. clviger A. folice Sex etween groups mle femle Intertion tretment x coreid species tretment x sex coreid pecies x sex ll vriles nounced for A. folice mrginell thn for H. clviger. The ltter used fruits of P. sueros more intensively, followed y P. miser. Both coreids fed on the vegettive prts of P. edulis more frequently thn on P. sueros or P. miser (Figure 4). Focl oservtions on feeding ctivity for the mixed tretment indicted tht oth coreid species used severl plnt species nd prts (Tles 3 nd 4). This pttern ws more significnt for the lst instr nymphs, whose feeding ctivity nd instr durtion lsted longer. 4. Discussion P. sueros provides the superior performnce for oth H. clviger nd A. folice mrginell thn P. miser, P. edulis, or ll pssion vines in comintion. This my e explined ecuse P. sueros is ntive plnt whose wild form is mintined, nd is not modified for economic purposes. Moreover, P. sueros is the most undnt pssion vine in the re (C. Mondin nd GRP. Moreir, UFRGS, unpulished dt), nd presents oth vegettive nd reproductive structures throughout the yer. In contrst, P. miser, the second est host, is hevily defolited, nd presents no reproductive structures during fll nd winter (Rodrigues nd Moreir, 24). Field nd lortory oservtions indicte tht these insects do not undergo dipuse, remining ctive throughout fll nd winter. In ddition, preliminry dt indicte tht H. clviger immtures do not rech the forth instr when fed with only one structure of P. sueros, either veget- 94 Brz. J. Biol., 67(1): 91-99, 27

5 Food mixing effects on coreids = 4 = = 6 = = 16 = = 22 = 12 Reltive frequency (%) c d e TBU LEA STE FBU FLR FRU Plnt structure = = 17 = 16 = 23 = 22 = 16 Reltive frequency (%) c d e TBU LEA STE FBU FLR FRU Plnt structure = 14 = 21 = 26 = 21 = 26 = 21 Figure 2. Feeding ctivity of H. clviger (open rs) nd A. folice mrginell (closed rs) immtures (Figures to e - first to fifth instr, respectively) in reltion to P. sueros plnt prts. Aric numers following the corresponding squres represent numer of individuls oserved in given instr. TBU, terminl ud; LEA, lef; STE, stem; FBU, flower ud; FLR, flower; FRU, fruit. Figure 3. Feeding ctivity of H. clviger (open rs) nd A. folice mrginell (closed rs) immtures (Figures to e - first to fifth instr, respectively) in reltion to P. miser plnt prts. Aric numers following the corresponding squres represent numer of individuls oserved in given instr. TBU, terminl ud; LEA, lef; STE, stem; FBU, flower ud; FLR, flower; FRU, fruit. tive or reproductive (D. Rodrigues nd GRP. Moreir, UFRGS, unpulished dt). Therefore, the presence of oth vegettive nd reproductive structures, in ll sesons, is n importnt ttriute of potentil host-plnt for oth coreids. H. clviger nd A. folice mrginell femles nd mles show similr performnces, indicting tht they re ffected nd respond eqully to ll pssion vines tested. Although femles hd wider domens thn mles, they re similr in length. Thus, our findings give support to the hypothesis for the existence of tesin mimicry for oth sexes of H. clviger in reltion to ichneumonids (Rodrigues, 23), despite the common size dimorphism for coreids (Schuh nd Slter, 1995). Survivorship ws not ffected y tretment nd coreid species, eing less thn 5% in ll cses. This vlue is considered lower thn tht found in other studies regrding heteropterns (Pnizzi et l., 1996). Adopted methodologicl procedures, such s the sence of soil nd/or shoot sectioning my hve led either to nutrition- Brz. J. Biol., 67(1): 91-99, 27 95

6 Rodrigues, D., Durte, LS. nd Moreir, GRP. Reltive frequency (%) TBU LEA STE FBU FLR FRU c d e Plnt structure = 3 = 3 = 2 = = 8 = 16 = 24 = = 23 = 13 Figure 4. Feeding ctivity of H. clviger (open rs) nd A. folice mrginell (closed rs) immtures (Figures to e - first to fifth instr, respectively) in reltion to P. edulis plnt prts. Aric numers following the corresponding squres represent numer of individuls oserved in given instr. TBU, terminl ud; LEA, lef; STE, stem; FBU, flower ud; FLR, flower; FRU, fruit. l losses or the increse of induced defenses to herivory, which my explin the high mortlity. Such hypotheses, however, need further investigtion. Both H. clviger nd A. folice mrginell feed on reproductive structures of the wild ntive species (P. sueros nd P. miser) more often thn on the cultivted one, P. edulis. This fct my e relted to physi- cl nd chemicl chrcteristics of the pssion vines. For instnce, the fruit is less coriceous in the wild species. Moreover, ll fruit prts of P. sueros nd P. miser my e used, including the seeds, due to the length of the stylets (totl rnge: 1.7 to.5 mm; Rodrigues nd Moreir, 25) tht rech them. According to Scco (198) nd Acioli (1999), P. sueros nd P. miser fruit dimeters rnge from.4 to 1.5 cm. This is not the cse of P. edulis, whose fruits hve dimeter from 5 to 6 cm (Scco, 198), which mkes the seed use limited. The fct tht seeds re the most nutritive resource for seedsucking insects (Slnsky-Júnior nd Pnizzi, 1987) explins the poorest performnce of oth species on this host-plnt. The reltion etween stylet size nd seed loction in Heteropter is poorly investigted, lthough host rce rdition cses relted to tht hve een reported for some hemipterns (Crrol nd Boyd, 1992). In reltion to host-plnt chemicl defenses, cinogenic glycosides hve een descried nd isolted in purple fruits of P. edulis, nd lredy detected in fruits of other cultivted pssion vines species (Chssgne nd Crouzet, 1998). H. clviger nd A. folice mrginell my use the vegettive tissues of P. edulis more frequently thn the reproductive ones to sequester, synthesize or trnsform secondry metolites (see Bowers, 199). Such compounds my e more concentrted in vegettive structures of cultivted plnts due to their high pltility nd strong susceptiility (MS. Singer, Wesleyn Universtiy, personl communiction). This hypothesis, however, must e tested, since these hemipterns pprently use repugntory glnds for defense, whose compound is minly formed from lcohols, ldehyds, nd esters (see Whitmn et l., 199). The individul feeding pttern oserved in the mixed tretment suggests tht nymphs use different tissues nd different host-plnts throughout ontogeny, even within given instr. This corroortes food mixing condition (Singer nd Bernys, 23) of these immtures, since severl heteropterns re polyphgous. The nisocelins studied here re considered oligophgous (use of host-plnts tht elong to the sme fmily; Bernys nd Chpmn, 1994). However, this should e tken with cution, ecuse this clssifiction does not tke into ccount the vriety of plnt prts explored y them. Besides, H. clviger nd A. folice mrginell showed good performnces when rered in the mixed tretment. The high moility of nymphs tht cn esily rech severl prts of their host-plnts, which enles them to lnce nutritionl needs s well s contct with toxins, my explin the dvntges of hving mixed diet. In ddition, lthough not tested, we did not identify ny evidence for the existence of the oviposition site selection regrding the different plnt prts. Both in the field nd in the lortory, eggs re rrely found on plnts, eing lid either isolted or ggregted on severl structures such s dry stems nd tendrils, cge wlls, nd others. It is known tht the correltion etween the oviposition preference nd offspring performnce is medited y 96 Brz. J. Biol., 67(1): 91-99, 27

7 Food mixing effects on coreids Tle 3. Focl oservtions on the feeding ctivity of H. clviger immtures in the mixed tretment. Ech line represents oservtions of given individul t different occsions. FBU. flower ud; GFR. green fruit; LEA. lef; NEC. extr-florl nectry; PFR. purple fruit; TBU. terminl ud. Instr I II III IV V P. miser LEA P. edulis GFR P. miser PFR P. miser LEA P. edulis NEC P. miser LEA P. edulis LEA P. miser TBU P. miser PFR P. edulis FBU P. edulis LEA P. miser FBU P. miser LEA P. miser LEA Tle 4. Focl oservtions on the feeding ctivity of A. folice mrginell immtures in the mixed tretment. Ech line represents oservtions of given individul t different occsions. FBU. flower ud; FLR. flower; GFR. green fruit; LEA. lef; PFR. purple fruit; TEN. tendril. Instr I II III IV V P. edulis TEN P. edulis TEN P. miser FBU P. miser FBU P. miser FBU P. sueros LEA P. miser FBU P. miser FBU P. sueros GRF P. miser FBU P. miser FBU P. miser BFU P. miser PFR P. edulis FLR P. edulis FBU oth genetic nd environmentl fctors, nd my explin spects relted to the evolution of insects plnt interction (Arhmson nd Weis, 1997). However, individuls rered in the mixed tretment did not hve superior performnce thn other tretments, pttern lredy chrcterized for some generlist insects, including heteropterns (Bernys nd Minkenerg, 1997). In generl, H. clviger nd A. folice mrginell use the green fruits more intensively thn other plnt prts. In fct, they prefer green fruits (Rodrigues, Sores nd Moreir, in prep.), pttern lredy descried for severl sucking species (see Pnizzi, 2). When in contct with leves, Brz. J. Biol., 67(1): 91-99, 27 97

8 Rodrigues, D., Durte, LS. nd Moreir, GRP. they use xylem (Rodrigues et l., sumitted), nutritionlly poor, ut importnt resource. A generl sttement out hemiptern feeding, not corroorted in this study, points out tht first instr nymphs do not feed (e.g., Simmons nd Yergn, 1988), including Holymeni histrio (Bldin nd Boiç-Júnior, 1999). Feeding ctivity on vegettive structures t this instr ws only recorded for Leptoglossus fulvicornis, genus closely relted to Anisoscelis (Wheeler-Júnior nd Miller, 199), s well s for Ans tristis, coreid tht feeds on cucurits (Bonjour et l., 1991). There is not cler reltion regrding the hemiptern use of ntive, non-ntive, wild or cultivted plnts nd performnce. For exmple, Neomeglotomus prvus nymphs nd dults tht use legumes show etter performnces when fed on cultivted, non-ntive plnts (Sntos nd Pnizzi, 1998). However, the etter performnce on wild ntive plnts hs een recorded for some penttomid species (Pnizzi, 1997). This study indictes superior performnce on wild, ntive plnts thn on cultivted ones. In ddition, preliminry dt showed tht H. clviger nd A. folice mrginell young nymphs hd high mortlity when rered on P. lt shoots, whose reproductive structures hd high scission rtes fter cutting. Field studies crried out on southestern Brzil detected smll density of H. histrio nd A. folice mrginell individuls on the cultivted pssion vines P. lt nd P. edulis (Cetno et l., 2). Our conclusions regrding the poorest performnces of these coreids on these species, in ssocition to multiple use of plnt prts, my question their pest sttus. In conclusion, this study suggests n ecologicl similrity of these sucking species with respect to the interction with their host-plnts, in ddition to the strong morphologicl similrity of the immture stges (Rodrigues nd Moreir, 25). Although elonging to different genus, these coreids my hve evolved s sister species due to the resons mentioned ove. Alterntively, the immture stges my present evolutionry convergence or e mimics, in order to mintin the coexistence nd the common use of their host plnts. Finlly, genetic drift is n lterntive scenrio for explining H. clviger nd A. folice mrginell evolution. Acknowledgments We would like to thnk Michel S. Singer (Wesleyn University) for discussions concerning food mixing ehvior. We re especilly grteful to Antônio R. Pnizzi (EMBRAPA/CNPSoj) for criticlly reding the mnuscript. Finncil support cme from CAPES Scholrship (Ph.D. Disserttion) grnted to Dniel Rodrigues. This is contriution numer 48 of the Deprtmento de Zoologi of Universidde Federl do Rio Grnde do Sul. References ABRAHAMSON, WG. nd WEIS, AE., Evolutionry ecology cross three trophic levels: goldenrods, gllmkers, nd nturl enemies. Princeton University Press, Princeton, 456p. ACIOLI, MF., Estudo preliminr d iologi florl de Pssiflor sueros Linneus (Pssiflorcee). Honors Thesis, UFRGS, Porto Alegre, 23p. AMARAL-FILHO, BF., Efeito ds diets nturis e de ftores mientis n iologi de Phthi pict (Drury, 177) so condições de lortório (Hemipter, Coreide). Rev. Brsil. Biol., vol. 41, p AMARAL-FILHO, BF., Oservções sore o ciclo iológico de Crinocerus snctus (Fricius, 1775) (Hemipter; Coreide) so condições de lortório. An. Soc. Entomol. Brsil, vol. 15, p AMARAL-FILHO, BF. nd CAJUEIRO, IVM., Oservções sore o ciclo iológico de Venez stigm (Herst, 1784) Osun, 1975 (Hemipter, Coreide) em lortório. An. Soc. Entomol. Brsil, vol. 6, p AMARAL-FILHO, BF. nd STORTI-FILHO, A., Estudos iológicos sore Leptoglossus gongr (Fricius, 1975), (Coreide, Hemipter) em lortório. An. Soc. Entomol. Brsil, vol 5, p BALDIN, ELL. nd BOIÇA-JÚNIOR, AL., Desenvolvimento de Holhymeni histrio (Fr.) (Hemipter: Coreide) em frutos de cinco genótipos de mrcujzeiro (Pssiflor spp.). An. Soc. Entomol. Brsil, vol. 28, p BERNAYS, EA. nd CHAPMAN, RF., Host-plnt selection y phytophgous insects. Chpmn nd Hll, New York, 312p. BERNAYS, EA. nd GRAHAM, M., On the evolution of host specificity in phytophgous rthropods. Ecology, vol. 69, p BERNAYS, EA. nd MINKENBERG, PJM., Insect herivores: different resons for eing generlist. Ecology, vol. 78, p BONJOUR, EL. nd FARGO, WS., Host effects on the survivl nd development of Ans tristis (Heteropter: Coreide). Environ. Entomol., vol. 18, p BONJOUR, EL., FARGO, WS., WEBSTER, JA., RICHARDSON, PE. nd BRUSEWITZ, GH., Proing ehvior comprisons of sqush ugs (Heteropter: Coreide) on cucurit hosts. Environ. Entomol., vol. 2, p BOWERS, MD., 199. Recycling plnt nturl products for insect defense. In EVANS, DL. nd SCHMIDT, JO. (eds.): Insect Defenses. Stte University of New York Press, Alny, p CAETANO, AC., BOIÇA-JÚNIOR., AL. nd RUGGIERO, C., 2. Avlição d ocorrênci szonl de percevejos em cinco espécies de mrcujzeiro, utilizndo dois métodos de mostrgem. Brgnti, vol. 59, p CARROLL, SP. nd BOYD, C., Host rce rdition in the soperry ug: nturl history with the history. Evolution, vol. 46, p CHASSAGNE, D. nd CROUZET, JA., cynogenic glycoside from Pssiflor edulis fruits. Phytochemistry, vol. 49, p JOSHI, A. nd THOMPSON, JN., Trde-offs nd the evolution of host speciliztion. Evol. Ecol., vol. 9, p LIMA, AC., 194. Insetos do Brsil: Hemípteros. Tomo II. Escol Ncionl de Agronomi, Rio de Jneiro, 351p. MANLY, BFJ., Rndomiztion nd Monte Crlo methods in iology. Chpmn nd Hll, New York, 281 p. MOTULSKY, H., Anlyzing dt with Grph Pd Prism softwre. Grph Pd Softwre, Sn Diego, 397p. 98 Brz. J. Biol., 67(1): 91-99, 27

9 Food mixing effects on coreids PANIZZI, AR., Desempenho de ninfs e dultos de Leptoglossus zontus (Dlls, 1852) (Hemipter: Coreide) em diferentes limentos. An. Soc. Entomol. Brsil, vol. 18, p PANIZZI, AR., Wild hosts of penttomids: ecologicl significnce nd role in their pest sttus on crops. Annu. Rev. Entomol., vol. 42, p PANIZZI, AR., 2. Suoptiml nutrition nd feeding ehvior of hemipterns on less preferred plnt food sources. An. Soc. Entomol. Brsil, vol. 29, p PANIZZI, AR, VIVAN, LM., CORRÊA-FERREIRA, BS. nd FOERSTER, LA., Performnce of southern green stink ug (Heteropter: Penttomide) nymphs nd dults on novel food plnt (jpnese privet) nd other hosts. Ann. Entomol. Soc. Am., vol. 89, p PILLAR, VD., 21. MULTIV Multivrite Explortory Anlysis nd Rndomiztion Testing. User s Guide, v RODRIGUES, D., 23. Descrição, trjetóris ontogenétics de Holymeni clviger (Herst, 1784) e Anisoscelis folice mrginell (Dlls, 1852) (Hemipter: Coreide) e interção com três espécies de Pssiflorcee. Unpulished Thesis, UFRGS, Porto Alegre, 127p. RODRIGUES, D. nd MOREIRA, GRP., 24. Sesonl vrition in lrvl host-plnts nd consequences for Heliconius erto (Lepidopter: Nymphide) dult ody size. Austrl Ecol., vol. 29, p RODRIGUES, D. nd MOREIRA, GRP., 25. Comprtive description of the immture stges of two very similr leffooted ugs, Holymeni clviger (Herst, 1784) nd Anisoscelis folice mrginell (Dlls, 1852) (Hemipter: Coreide). Rev. Brs. Entomol., vol. 49, p RODRIGUES, D., SANFELICE, D., MONTEIRO, LR. nd MOREIRA, GRP., 25. Ontogenetic trjectories nd hind tii geometric morphometrics of two very similr lef footed ugs, Holymeni clviger (Herst) nd Anisoscelis folice mrginell (Dlls) (Hemipter: Coreide). Neotrop. Entomol., vol. 34, no. 5. p RODRIGUES, D., SAMPAIO, DS., ISAIAS, RMS. nd MOREIRA, GRP. Xylem nd seed feeding y two pssion vine lef-footed ugs, Holymeni clviger (Herst) nd Anisoscelis folice mrginell (Dlls) (Hemipter: Coreide), with notes on mouthprt morphology. Ann. Entomol. Soc. Am. sumitted. SACCO, JC., 198. Pssifloráces. In Reitz, R. (ed.): Flor ilustrd ctrinense. Herário Bros Rodrigues, Itjí, 13p. SANTOS, CH. nd PANIZZI, AR., Nymph nd dult performnce of Neomeglotomus prvus (Hemipter: Alydide) on wild nd cultivted legumes. Ann. Entomol. Soc. Am., vol. 91, p SCHAEFER, CW. nd MITCHELL, PL., Food plnts of the Coreidoide (Hemipter: Heteropter). Ann. Entomol. Soc. Am., vol. 76, p SCHUH, RT. nd SLATER, JA., True ugs of the world (Hemipter: Heteropter): clssifiction nd nturl history. Cornell University Press, Ithc, 336p. SIMMONS, AL. nd YEARGAN, KV., Feeding frequency nd feeding durtion of the green stink ug (Hemipter: Penttomide) on soyen. J. Econ. Entomol., vol. 81. p SINGER, MS. nd BERNAYS, EA., 23. Understnding omnivory needs ehviorl perspective. Ecology, vol. 84, p SINGER, MS., RODRIGUES, D., STIREMAN III, JO. nd CARRIÈRE, Y., 24. Roles of food qulity nd enemy freespce in host use y generlist insect herivore. Ecology, vol. 85, p SLANSKY-JÚNIOR, F. nd PANIZZI, AR., Nutritionl ecology of seed-sucking insects. In: F. Slnsky Jr. nd J. G. Rodriguez (eds.), Nutritionl ecology of insects, mites, spiders, nd relted invertertes. John Wiley & Sons, New York, p SOKAL, RR. nd ROHLF, FJ., Biometry. 3 rd ed. Freemn, New York, 887p. THOMPSON, JN., Evolutionry ecology of the reltionship etween oviposition preference nd performnce of offspring in phytophgous insects. Ent. Exp. Appl., vol. 47, p WHEELER-JÚNIOR, AG. nd MILLER, GL., 199. Leptoglossus fulvicornis (Heteropter: Coreide), specilist on mgnoli fruits: sesonl history, hitts, nd descriptions of immtures stges. Ann. Entomol. Soc. Am., vol. 83, p WHITMAN, DW., BLUM, WS. nd ALSOP, DW., 199. Allomones: chemicls for defense. In: EVANS DL. nd SCHMIDT JO. (eds.), Insect defenses. Stte University of New York Press, Alny, p Brz. J. Biol., 67(1): 91-99, 27 99

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