Ecology, 84(10), 2003, pp. 2549±2556 ᭧ 2003 by the Ecological Society of America

THE EVOLUTION OF OMNIVORY IN HETEROPTERAN

MICKY D. EUBANKS,1,3 JOHN D. STYRSKY,1 AND ROBERT F. D ENNO2 1Department of Entomology and Plant Pathology, Auburn University, Auburn, Alabama 36849 USA 2Department of Entomology, University of Maryland, College Park, Maryland 20742 USA

Abstract. Although omnivory is common and widespread across many taxa, the evolutionary origin of omnivory, the selective forces that promote or constrain omni- vory, and the morphological, physiological, and behavioral hurdles that have to overcome to become omnivores have not been studied. The goal of this paper is to stimulate the development of ideas concerning the evolution of omnivory. We focus on the terrestrial lineages of the order and use published life history data and recent phylogenies to test two hypotheses concerning the evolutionary origin of feeding on both plants and prey: (1) that the propensity to feed on seeds and pollen is correlated with the evolution of omnivory, and (2) that broad host range (polyphagy) is correlated with the evolution of omnivory. In order to test these hypotheses, we mapped the plant part consumed and host plant range of insect species in two heteropteran suborders onto their respective phylogenies and used phylogenetically independent contrasts to test for correlations of these traits with omnivory. We found evidence that seed and pollen feeding and broad host ranges are correlated with the evolution of omnivory within both ancestrally herbivorous and ancestrally predaceous lineages of terrestrial heteropterans. Key words: feeding habits; herbivory; Heteroptera; omnivory; predation; seed and pollen feeding; sister-group comparisons. pca Feature Special

INTRODUCTION holding prey) (Faucheux 1975, Cobben 1979, Cohen 1996). Feeding on both prey and plant food is a widespread The ecological signi®cance of omnivory has histor- feeding habit in insects (Whitman et al. 1994, Alomar ically received little attention. Recent studies, however, and Wiedenmann 1996, Coll and Guershon 2002). suggest that omnivory can deny prey density-related Thousands of omnivorous insect species occur through- refugia from predation, dictate the strength of top-down out a broad range of insect taxa, including grasshop- control and resulting trophic cascades, alter the stabil- pers, earwigs, thrips, true bugs, beetles, and ants. These insects represent a unique blend of morphological, ity of food webs, and profoundly in¯uence the move- physiological, and behavioral adaptations found in ment of energy and nutrients through ecosystems (Polis their predaceous and herbivorous relatives. Omnivo- et al. 1989, Polis 1991, Polis and Strong 1996, Fagan rous insects in the order Heteroptera (true bugs), for 1997, Holt and Polis 1997, Ostrum et al. 1997, Holyoak example, possess digestive tracts and accessory sali- and Sachdev 1998, McCann et al. 1998, Rosenheim vary glands that are intermediate in length, size, and 1998, Eubanks and Denno 1999, 2000). placement of those found in their herbivorous and pre- Studies of the evolution of omnivory, in contrast, are daceous relatives (Slater and Carayon 1963, Goodchild almost nonexistent in the literature (but see Cooper 1966). In addition, many species of omnivorous het- 2002, Denno and Fagan 2003, Diehl 2003). We know eropterans produce protein-digesting enzymes (pro- relatively little about the adaptive advantages of om- teinases and phospholipases) and plant-digesting en- nivory and the selective forces that favor or constrain zymes (amylases and pectinases) whereas their strictly the evolution of omnivory. The evolution of the mor- herbivorous and predaceous cousins produce only a phological, physiological, and behavioral traits asso- subset (Baptist 1941, Goodchild 1966, Kahn and Ford ciated with omnivory have not been studied and we do 1967, Miles 1972, Varis et al. 1983, Cohen 1990, 1996, not know if these traits evolve as a suite of correlated Schaefer and Panizzi 2000, Wheeler 2001). Omnivo- characters. Further, we know very little about the evo- rous heteropterans also have piercing-sucking mouth- lutionary consequences of evolving the ability to feed parts (stylets) with characteristics of both herbivorous on both plants and prey. For example, the evolution of heteropterans (smooth stylets to penetrate plants) and omnivory might in¯uence the diversi®cation rate of predaceous heteropterans (toothed or curved stylets for omnivorous taxa, but questions such as this have not been raised in the literature. The goal of this paper is to stimulate the devel- Manuscript received 5 July 2002; revised and accepted 28 opment of ideas concerning the evolution of omni- October 2002. Corresponding Editor: A. A. Agrawal. For reprints of this Special Feature, see footnote 1, p. 2521. vory. Insight into the evolutionary origin of omnivory 3 E-mail: [email protected] will help us understand the adaptive signi®cance of 2549 Special Feature loafc hi rpniyt vleteadaptations 1979). Cobben the 1979, (Sweet evolve feeding prey to for necessary propensity their affect also both from omnivory of predation. evolution and transi- herbivory a the be in to state seeds hypothesized tional on therefore, Feeding is, to 1979). pollen similar and Cobben parts more 1979, plant nutritionally (Sweet nitrogen are prey pollen) high and because (seeds foliage feed that plant omnivores than on pollen omnivores and to seeds rise on hy- give feed to that are likely more insects be to of be- pothesized lineages match nutritional predaceous a Conversely, prey. nitrogen-rich of and parts basis plant nitrogen-rich the tween on This predicted 1979). Cobben is foliage- 1979, to (Sweet of rise herbivores lineages feeding give than frequently will more herbivores of omnivores pollen-feeding lineages or that consequently, seed and, ni- prey consume to trogen-rich preadapted 2001). be may pollen- Wheeler herbivores and feeding seed 2000, that hypothesized Cohen have Panizzi authors 1985, Some and Kirk Schaefer 1984, al. 1996, pollen-feeding et and en- (Houseman seed these their relatives but by pollen, produced and are seeds zymes in found digest to proteins them the allow would produce that enzymes not digestive do the species 1985). Kirk foliage-feeding 1979, example, (Janzen plants For or part plant particular a host on their feed of al. to herbivorous adapted result, et usually a are Strong As insects 1988). 1979, me- al. et al. antiherbivore Thomison et of 1984, (Tamas degree defenses and type chanical the of and concentrations allelochem- icals, the defensive carbohydrates, in nutrients, vary Weaver other also and Douglas parts 1984, Plant Cordova-Ed- al. 1989). and et Strong Murray 1984, 1982, wards Evans 1935, (An- respectively) drews 0.0002%, nitrogen and less (0.005% even leaves contain nitrogen than tissues 10% xylem and nitrogen. to 0.7% Phloem as little up as contain Seeds often contain leaves 1984). whereas frequently al. et pollen (Strong and insects most limiting important for an is nutrient concentra- nitrogen the and in nitrogen tremendously of of vary tions parts can species plant Different the host 1979). by same Cobben consumed part 1979, plant (Sweet the may to prey consume related and be capture to ability the species evolve herbivorous to an of propensity the herbivory, of with omnivory. associated of is evolution (polyphagy) (2) the range and host omnivory, of broad evolution that pollen the and seeds with on correlated feed is to and tendency plants the both that on (1) feeding prey: of origin and concerning evolutionary hypotheses data the two order test history insect to life phylogenies the published recent of use We lineages and food. terrestrial Heteroptera plant must the and on animals prey focus both that consume hurdles to overcome behavioral physio- morphological, and the logical, elucidate and omnivory 2550 h otrneo ebvru netseismay species insect herbivorous of range host The state ancestral an from omnivory of evolution the In IK .EBNSE AL. ET EUBANKS D. MICKY ots o orltosbtenteetat n omni- and traits these between vory. correlations contrasts for independent test to phylogenetically used phyloge- two and respective in nies their insects onto of range suborders plant heteropteran host plant and the consumed mapped parts we omnivory, of origin evolutionary om- of evolution the nivory. regarding historical ideas and test biological to rich framework Heteropter- a 1998). provide Coll therefore, Schuh 1995, ans, 1979, Slater Sweet and Schuh 1979, 1986, (Cobben herbivores giving to lineages rise predaceous and predators lineages to herbivorous rise with giving order, mul- this evolved within have times habits tiple feeding three the addition, relationships. In evolutionary well- and relatively taxonomic have established and biologies, di- well-known have have species, omnivorous they many because contain habits, studying habits feeding verse for feeding group of Ter- evolution ideal the Heteroptera. an order are ter- insect heteropterans several restrial the of of history lineages evolutionary restrial and histories life they plants host the about speci®c consume. are very omnivores be these to taxa, unlikely plant spe- for speci®c selection on to undergo cialization likely they Until also recently polyphagous. are be have lineages predaceous that within evolved species omnivorous om- relatives. evolve monophagous Conversely, their to are likely than her- species more nivorous polyphagous be may of consume therefore, Lineages and bivores, attack 2001). to (Wheeler predisposition prey their as well requirements monophagous nutrient as their their affect may than this often and relatives move more oli- far herbivores or plants Polyphagous among monophagous their relatives. than gophagous omnivorous they that be likely more will it make that (Sweet traits behavioral also species have may herbivores monophagous Polyphagous 1979). than Cobben 1979, prey consume likely more to species polyphagous detox- make as enzymes such ifying adaptations that Some suggested have 1997). authors Minkenberg and (Bernays allelochemical-based defenses have of plant and variety a species detoxify plant to of ability enzymes the variety a the digest produce to For necessary lifestyle. insects omnivorous polyphagous an families) example, to plant themselves more lend or may two representing di- plants on (feeding verse polyphagy with taxa associated adaptations behavioral plant and of physiological, morphological, spe- diversity The insect cies. omnivorous the or herbivorous by an by de®ned consumed is range Host 2fmle,adtosbresfo 4 published 146 terrestrial the from on focused suborders We Appendices). two (see sources genera, and 232 families, in 22 distributed heteropterans terrestrial of nodrt etortohptee ocrigthe concerning hypotheses two our test to order In the on focusing by hypotheses these evaluated We eetatdlf itr nomto o 9 species 398 for information history life extracted We M ETHODS clg,Vl 4 o 10 No. 84, Vol. Ecology, October 2003 WHY OMNIVORY? 2551 heteropterans in the suborders Cimicomomorpha and TABLE 1. Numbers of omnivores and herbivores used in eight independent contrasts to test for correlations of feed- because they include large numbers ing habit with plant part consumed (foliage vs. reproductive of herbivores, omnivores, and predators, and because parts) and with host range (monophagy or oligophagy vs. strict herbivory, omnivory, and strict predation have polyphagy). evolved independently multiple times within these groups (Cobben 1979, Sweet 1979, Schuh 1986, Schuh No. No. omnivores herbivores and Slater 1995, Alomar and Wiedenmann 1996). We Taxa in contrast in contrast used the most recent phylogenies for each suborder, Miridae 54 66 Schuh and Stys (1991) for the Cimicomomorpha (Ap- Pentatomoidea² 28 64 pendix A) and Henry (1997) for the Pentatomomorpha Alydidae ϩ Coreidae 1 14 (Appendix B), for our analyses. Both of these phylog- 1 3 Pyrrhocoridae 2 7 enies are based on morphological characters. These two Berytidae 7 4 lineages are particularly useful for this study because Ishnorhynchinae ϩ Orsillinae 1 8 the Cimicomomorpha is ancestrally predaceous and the Lygaeinae 3 7 Pentatomomorpha is ancestrally herbivorous (Sweet ² Includes families Acanthosomatidae, Cydnidae, Penta- 1979, Cobben 1979, Schuh 1986, Schuh and Stys 1991, tomidae, Scutelleridae, and Thyreocoridae. Henry 1997). The correlation of seed and pollen feed- ing and broad host range with the evolution of omni- vory in both lineages would suggest that these traits orous; Table 1). This procedure reduced our data to play a pivotal role in the evolutionary transition from eight family-level groups (Table 1). We then conducted herbivory to omnivory and from predation to omni- a logistic regression analysis with family group, plant vory. part consumed (foliage or reproductive), and host range For each heteropteran species surveyed, we scored (restricted or polyphagous) as predictor variables and the family and subfamily classi®cation, the plant part

feeding habit (herbivory or omnivory) as the dependent Feature Special consumed by the species, and whether or not the spe- variable (SAS version 8.2, Proc Logistic with class cies consumed prey. From this information, we char- statement; Stokes et al. 2000). acterized the species' feeding habit as strictly herbiv- This analysis corrects for phylogenetic noninde- orous (consumes only plants), omnivorous (consumes pendence among family-level groups, but treats spe- plants and prey), or strictly predaceous (consumes only cies within these groups as independent and does not prey). We also characterized the species host range as control for phylogenetic non-independence at levels monophagous (consumes plants belonging to one ge- above family. The second analysis controls for phy- nus), oligophagous (consumes plants in two or more logenetic nonindependence at all levels and is con- genera within the same plant family), or polyphagous sidered a conservative test of correction for similarity (consumes plants in two or more families). due to common ancestry. In this analysis, we reduced We used two analyses to control for possible phy- logenetic nonindependence among heteropteran spe- the data set further to include only phylogenetically cies. Our approach follows that of Fagan et al. (2002) independent contrasts among sister taxa (Ridley 1983, and all methods are based on the principal of phylo- Felsenstein 1985, Harvey and Pagel 1991). This re- genetically independent contrasts (Felsenstein 1985). quired that we assign a feeding habit to each family- The two tests represent each end of the continuum be- level group. If all species within a group were her- tween strictness of correction for similarity due to com- bivorous, then we scored that group as herbivorous. mon ancestry and potential statistical power (Mazer If some or all species within the group were omniv- 1998, Ackerly and Reich 1999). We used both liberal orous, then we scored that group as omnivorous. We and conservative techniques to control for phylogenetic identi®ed four meaningful phylogenetically indepen- constraints because omnivory has evolved multiple dent contrasts using the phylogenies of Schuh and times within multiple heteropteran lineages and is un- Stys (1991) and Henry (1997): Miridae (omnivorous) likely to be highly conserved (i.e., is phylogenetically vs. Tingidae (herbivorous), Coreidae (herbivorous) labile). vs. Alydidae (omnivorous), Berytidae (omnivorous) The ®rst analysis partitioned species into a set of vs. Colobathristidae (herbivorous), and Ischnorhyn- family-level groups each containing at least one phy- chinae (omnivorous) vs. Orsillinae (herbivorous). We logenetically independent contrast between herbivores used two 2 ϫ 2 contingency table analyses to test the and omnivores (sensu Fagan et al. 2002). These group- hypotheses that evolutionary changes in feeding habit ings corresponded to single, monophyletic taxa in the were independent of plant part consumed and host case of the superfamily Pentatomoidea, the families range. A signi®cant G is evidence that pairs of char- Berytidae, Miridae, Pyrrhocoridae, and Rhopalidae, acter states are not independent, but that some com- and the subfamily Lygaeinae. In other cases, these fam- binations of characters are more or less common than ily-level groups consisted of pairs of sister taxa with expected by chance (Ridley 1983, Harvey and Pagel different feeding life histories (omnivorous vs. herbiv- 1991). Special Feature ␹ ϭ 2552 nflaewr mioos(i.1.Teefc of (Wald effect signi®cant The not was 1). analysis this (Fig. in group omnivorous fed family that were species pollen the foliage and of seeds on 25% Bery- on only the fed and that in omnivorous species were striking the more of even 85% (Fig. tidae: was omnivorous were pattern foliage This on 1). fed of that 37% only species reproduc- whereas on the omnivorous fed were that parts species Miridae, plant the tive the of In omnivorous 53% 1). be example, for (Fig. to species likely foliage-feeding more than were pollen and seeds nivory fteegtfml-ee rusue norlgsi ersinaayi.Msigbr niae`zr' values. ``zero'' indicate bars Missing analysis. regression logistic our in used groups family-level eight the of h ee ffml ru (Wald group at family nonindependence of level phylogenetic the for analysis the controlled in signif- habit feeding that a on part was the plant There of with effect pollen. icant correlated and is seeds of omnivory consumption of evolution the that in®atymr ieyt eonvru hntheir than omnivorous ( taxa be were sister pollen foliage-feeding to and likely seeds on more fed signi®cantly that taxa analysis, levels. this all In at nonindependence phylogenetic for trolled 2 pce fet t rpniyt eonvru.There heteropteran omnivorous. a be to of propensity range its host affects species the that evidence found aooopa eg,Brtdei i.1). Fig. in (Pen- Berytidae lineage (e.g., herbivorous ancestrally tatomomorpha) the ancestrally and 1) the Miridae Fig. in both (e.g., (Cimicomomorpha) to in lineage appeared omnivory predaceous pollen with and correlated seeds be on feeding addition, In F oyhg screae ihomnivory with correlated is Polyphagy om- with correlated is pollen and seeds on Feeding .4) vrl,htrpea pce htconsumed that species heteropteran Overall, 0.045). ϭ efudtesm atr nteaayi htcon- that analysis the in pattern same the found We IG .Pretgso pce htaeonvru mn pce htfe nflaeo nsesadple,i each in pollen, and seeds on or foliage on feed that species among omnivorous are that species of Percentages 1. . 11,df 11.10, Ðefudsrn upr o h hypothesis the for support strong found .ÐWe ϭ 7, P R ϭ ESULTS 0.134). G ϭ .,df 6.1, ␹ 2 ϭ ϭ .1 df 4.01, 1, IK .EBNSE AL. ET EUBANKS D. MICKY P Ðealso .ÐWe Ͻ 0.025). ϭ 1, P edn ai nteaayi htcnrle o phy- for (Wald level controlled family-group on that the range analysis at logeny host the of in affect habit signi®cant feeding statistically a was anonvru pce hntermnpaosor monophagous ( their taxa sister than oligophagous species omnivorous con- to tain likely more signi®cantly levels: were taxa all polyphagous at nonindependence phylogenetic for trolled omnivores. of distribution phylogeny of the effect on an was there that indicating 0.038), ϭ ihteeouino mioyi ersra heterop- terrestrial in correlated omnivory are of range evolution plant the host with broad and feeding len ettmie,teAlydidae the Pentatomoidea, sssaitcly hr a ttsial signi®cant (Wald group statistically family a of hypoth- was effect this There neither test was statistically. to set enough esis data balanced more nor our are enough but large species omnivorous, be monophagous to groups likely other omnivorous in be to and likely more groups are some in effect species that polyphagous such interactive range an host and be group family may These of there 2). that (Fig. in suggest Pyrrhocoridae evident results the was and pattern Rhopalidae opposite the The 2). (Fig. gaeinae a,teIschnorhynchinae the dae, a nFg 2). ancestrally Fig. the in Beryti- dae and (e.g., 2) (Pentatomomorpha) Fig. lineage herbivorous in (Cimicomo- Miridae lineage (e.g., predaceous in morpha) ancestrally correlated be the to appeared both omnivory and range Host eswtesm atr nteaayi htcon- that analysis the in pattern same the saw We u eut upr h yohssta edadpol- and seed that hypotheses the support results Our 1, P Ͻ .0) hswseieti h iia,the Miridae, the in evident was This 0.001). D ISCUSSION G ϩ ϭ ␹ ϩ .,df 6.0, rilne n h Ly- the and Orsillinae, 2 clg,Vl 4 o 10 No. 84, Vol. Ecology, ϭ oede h Beryti- the Coreidae, 48,df 14.83, ϭ ␹ 1, 2 P ϭ Ͻ ϭ 65,df 26.52, 0.025). 7, P ϭ October 2003 WHY OMNIVORY? 2553

FIG. 2. Percentage of species that are omnivorous among species with restricted host ranges (monophagous ϩ oligoph- agous) or broad host ranges (polyphagous) in each of the eight family-level groups used in our logistic regression analysis. Missing bars indicate ``zero'' values. pca Feature Special

terans. Heteropteran species that fed on seeds and pol- be addressed by comparative studies of closely related len were signi®cantly more likely to be omnivorous herbivores, omnivores, and predators. than their foliage-feeding relatives (Fig. 1). Likewise, polyphagous species were signi®cantly more likely to Ecological consequences be omnivorous than their relatives with restricted host ranges (monophagous or oligophagous), although this The consumption of seeds and pollen as well as po- effect may vary to some extent among family-level lyphagy could dramatically affect the spatial and tem- groups (Fig. 2). These patterns were apparent when we poral abundance of omnivorous insects. Omnivores used both liberal and conservative methods to control that consume seeds and pollen may be forced to track for the effects of phylogeny on the evolution of om- changes in the abundance of these plant resources in nivory and were apparent in both ancestrally preda- space and time (Eubanks and Denno 1999, 2000). Om- ceous (Cimicomomorpha) and ancestrally herbivorous nivorous insects are noted for their attraction to pollen (Pentatomomorpha) lineages. and seeds (Kiman and Yeargan 1985, Read and Lamp- We suggest that seed and pollen-feeding insects are man 1989, Coll and Bottrell 1991, 1992) and some preadapted to consume prey and that predaceous insects studies even suggest that omnivorous ``predators'' ac- are similarly predisposed to consume nitrogen-rich tually track variation in the production of seeds and plant parts. Preadapation may be associated with phys- pollen more than they track variation in the population iological traits. For example, the digestive enzymes size of prey (Cottrell and Yeargan 1998, Eubanks and produced by seed and pollen-feeding herbivores may Denno 2000). The impact of these omnivores on prey be able to digest prey. Similarly, the digestive enzymes is largely dictated by variation in the abundance of produced by predators may be able to digest seeds and seeds and pollen because the presence of high quality pollen. Further, some morphologies or behaviors of plant food not only affects omnivore abundance, but seed and pollen-feeding herbivores and predators may also alters the per capita consumption of prey by om- be functionally interchangeable. For example, how nivores (Cottrell and Yeargan 1998, Eubanks and Den- similar do the mouthparts of omnivores have to be to no 2000). Likewise, the abundance of polyphagous their herbivorous and predaceous relatives to function plant feeders is affected by spatial and temporal vari- adequately in both prey and plant consumption? Do ation in host plants, and polyphagous herbivores are host-plant identi®cation behaviors and mobility asso- frequently more widespread and abundant than mo- ciated with polyphagous heteropterans allow them to nophagous relatives, especially if the monophagous in- track prey? To our knowledge, these hypotheses have sects specialize on relatively rare plants (Ehrlich and never been tested; however, questions like these could Raven 1964). Special Feature lmr . n .N idnan dtr.19.Zooph- 1996. editors. Wiedenmann, N. R. and O., Alomar, and Convergence 1999. Reich. B. P. and D., D. Ackerly, 2554 930 oR .Dno n E-434 oR .Denno F. R. to Eubanks. DEB- DEB-9423543 D. grant and M. Denno, Foundation and F. Science R. to National 9903601 to Eubanks, Agriculture of D. Department in M. States supported United was the and from research DEB-0074556 funding grant This Foundation Science project. National by this part of var- stages at encouragement Al ious and and discussion Raupp, thoughtful Mike for Singer, Wheeler Mike Mitter, Charlie Lamp, Bill im- this of habit. consequences feeding portant and origin evolutionary concerning the ideas as- of heteropteran our development in the hope stimulates omnivory We insects pre- of range. evolution be host the may of broad and sessment a pollen have and to seeds pre- adapted on be or feed may single to predators a adapted Conversely, on species. specialize plant that few relatives their in their prey include than to diet ability are the evolve species to likely polyphagous more Likewise, foliage- relatives. their than feeding diet their adaptations in the prey include evolve to to necessary likely more far are parts reproductive herbivorous plant nitrogen-rich, heter- that these consume within suggest that range results species of host Our consumption broad insects. the and opteran with pollen and correlated seeds is omnivory of biodiversity. of generation the for consequenc- es profound have may omnivory the of that evolution suggests Co- result vs. interesting Berytidae This and lobathristidae). Tingidae vs. herbivorous (Miridae taxa taxa their omnivorous sister than the diverse contrasts, more three dramatically the were of out (herbivorous). two Colobathristidae In vs. Beryti- (omnivorous) and Coreidae dae (omnivorous), (herbivorous), Alydidae vs. Miridae Tingidae (herbivorous) species: extant vs. of number (omnivorous) the of and/ species estimates described we or of number which the for ®nd of readily taxa could contrasts sister herbivorous independent hy- and are three omnivorous this results found test preliminary We to our intriguing. data but enough statistically, have pothesis not rel- predaceous did We or herbivorous atives. ``success- strictly more their be than to ful'' plant species and omnivorous prey both allows consume food to ¯exibility ability the ecological by the provided if diversi®cation taxonomic ayad USA. 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APPENDIX A The phylogeny of family relationships in the heteropteran suborder Cimicimomorpha is available in ESA's Electronic Data Archive: Ecological Archives E084-063-A1. Special Feature E084-063-A3. 2556 nsbresCmcmmrh n ettmmrh saalbei S' lcrncDt Archive: Data Electronic ESA's in available is Pentatomomorpha and Cimicimomorpha suborders in aooi lcmn,hs ag,patpr edn rfrne n edn ai f38tretilhtrpea insects heteropteran terrestrial 398 of habit feeding and preference, feeding part plant range, host placement, Taxonomic Archive: h hlgn ffml eainhp ntehtrpea uodrPnaooopai vial nEAsEetoi Data ESA's Electronic in available is Pentatomomorpha suborder heteropteran the in relationships family of phylogeny The clgclArchives Ecological E084-063-A2. IK .EBNSE AL. ET EUBANKS D. MICKY PEDXC APPENDIX PEDXB APPENDIX clg,Vl 4 o 10 No. 84, Vol. Ecology, clgclArchives Ecological