University of Connecticut OpenCommons@UConn
ANSC Articles Department of Animal Science
1988 The food plants of some 'primitive' Pentatomoidea (Hemiptera: Heteroptera). Carl W. Schaefer University of Connecticut, [email protected]
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Recommended Citation Schaefer, Carl W., "The food lp ants of some 'primitive' Pentatomoidea (Hemiptera: Heteroptera)." (1988). ANSC Articles. 9. https://opencommons.uconn.edu/ansc_articles/9 9!E
THE FOOD PLANTS OF SOME "PRIWtrTTIVE" PENTATOMOIDEA(HEMIPTERA: HETEROPTERA)
CARL W. SCHAEFER
Department of Ecotogy and Evolutionar.t Biolog.r, Unit,ersity of Connecticut, Storrs, CT 06268 U.S.A.
,ABSTR.ACT
The iood plants of the Cydnidae (Cydninae,Sehirinae, Scaptocorinae, Amnestinae, Garsauriinae, Thau- mastellinae,Parastrachiinae, Corimelaeninae), Plataspidae, Megarididae, Cyrtocoridae, and Lestoniidae, compiled {iom the literature, are discussed.So too are the habitats ofthese insects,most ofwhich live on or are associatedwith the ground. This associationsupports an earlier assertionihat life on the ground was the way of lile o{ the early hemipterans. Most of these groups are polyphagous. However, the Plataspidae feed largely upon legumes, the Scaptocorinaeupon the roots ol Gramineae,some Cydninae also upon legumes,and many Sehirinaeupon members of the advanced dicot subclassAsteridae. Fallen seedsand roots are the preferred plant parts.
A group ol mostly small drab shieldbugsappears to be primitive in the Pentatomoi- dea: "primitive" in the senseof possessingseveral plesiomorphic features, and'oa group" becausethey also sharesome apomorphies; neither the autoplesiomorphies nor thc autapomorphiesare sharedwith other pentatomoids(Schaefer, in prep.).I havealso suggested (1981) that the earlyheteropterans lived on the ground,not up on plants (seealso Schaef'er,in press).Accordingly, some knowledge of thesepentato- moids' biology may be use{ul in working out their phylogenetic relationships, and interestingof course in its own right. 'lhese lamilies are the Cydnidae (sensulato'. seebelow), Plataspidae,Lestoniidae, Cyrtocoridae,Megarididae, and Canopidae.Their food plants and habitats are listed and brielly discussedhere. Almost all recordsare liom the literature, and I have been careful to eschew"sitting on" or probing records;I have always prel'erredrecords of nymphs'feeding. Therefore I hope any errors are ones of omission, not commission.
RESULTS Cydnidae
I {ollowhere the classilicationofthis lamily of Dolling(1981), who includedthe
Phytophaga:2 (1988) 19 45 g 20 CARLw. ScHAEFER
sublamiliesCydninae. Sehirinae,Amnestinae. Gu.ruuriinu., and Scaptocorinae I (groups usually included) as well as the Corimelaeninaeand Thaumastellinae(groups previously treated as separatefamilies). There is some evidencethe Corimelaeninae should be raised to {amily level, with two subf'amilies;my resuitshere argue neither for nor against such a view, and I follow Dolling's placement.The Parastrachiinae,a subfamily recently erectedand placed in the cydnidae (Schaet'eret al., in press)may also some day require Ibmily rank. Cydninae.- Members of this sublamily I'eedvery widely indeed (Table l), The {bod plant lamilies representsixteen disot and two monocot orders and all dicot subclassesexcept one (classiiicationo1'Cronquist, 1968). only two species{'eed on members of the Magnoliidae (on Papaveraceae),the most primitive ofthe dicot subclasses(cronquist, 1968),and none on anotherprimitive subclass,Hamamelidae. Six speciesin six genera{'eed on membersol the most advancedsubelass, Astericiae. This is a low number, comparedto the number o{'speciesin this subclass;and only three species(and three genera)I'eed on composites,an asterid lamily that is one of the largest plant lamilies. By lar the most cydninesleed on Rosidae,members of sevenorders being ied upon. The most popular lamily is Leguminosae(Rosales), particuiariy among memberso1'the genusGeotoruus, where Crucil'eraeand Gramineaeare also used. Grassesare also commonly I'edupon by other.cydnines,particularly by speciesof Aethus. Grassesand legumesoccur most fiequently in Table l, but in no consistent way; they olten occur as but part o1'a more generalizeddiet. No clear pattern emergesfiom these data. Cydnines feed very wideiy, and legumes,crucil'ers, and grassesare perhapslavored" No real specilicity is evident, at the sublamily level or at the generic(except perhaps Aethus on grasses).To some extent,the variety ofplants in Table I reflectsthe ditiiculty ol'knowing il'the bugs actually f'eed on the roots where they are fbund. It may be signilicantthat thesebugs, probably primitive within the Pentatomoi- dea, do not {'eedon the most primitive subclasses;nor do they f'eedto any great extent on the most advancedsubclass, the Asteridae. According to most authors, they I'eedon the roots of plants.But Stoner (1920) several times {bund Cydnus obliquus carrying a small seed with the middle legs. "when the insect was lieed liom the sand and allowed to walk lieely" (p. 340), it retained the seed and walked with the other legs. other cydninesare also known to I'eedon seeds.Concerning Adrisa, Van den Berg ( 1980,p. 225)writes, "The Adrisa spp. live in the debris underneathwattle trees. They t'eedon the seedsol both Alcacial cyclops and A. salignaas well as many other wattle species.All the seedsof A. saligna collected about 2 months after seedibll showed the characteristic I'eedingtubes of seed I'eeding Hemiptera and were non viable [sic]. As Adrisa spp. are the most dominant f'eedersunderneath the trees,most ,.i of the inviability can be attributed to them". t,.\ Many cydnineslive in dry areasand severalare actually in sand.This is often .1i aeoliansand (e.g., several species ol'Aethus 1983]),but Georomuspunctula- ,l [stehlik, 'j tus and Aethuspilosulus live in shoredunes. A great majority ofcydnines {'eedson roots in the sand or sandysoil, but Stehlik (I983) reportsol one, C),dnusaterrimus, ,,i that the llrst instarsl'eed on r,ri Euphorbiaroots but that "older nymphsand adultscreep Food Plants oJ' Pentatomoidea 2l
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io be E.$ (t .}i s 3 f;E €a P iE s t" Eg .:tr f SA $ 50o € Eg E> F!f; € E FH .l .a $s \: v vv C. impressus Horvath Thesiunt Santalaceae Heiss1977
C. maculipes (Mulsant & Rey) Centronlhus Valerianaceae de la Fuente 1972
Adomerus biguttatus Melamp.t'rum Scrophulariaceae on plants and among de la Fuente 1972, (L.) dry leaves Stelhik1983
Tritomegas sexamaculalus Ballota Labiatae Boselli1932, Heiss 1977, (Rambur) Daucas Urnbelliferae de la Fuente1972
"prefers humid Bakuni& Vasarhelyil98l conditicns"
T. bicolor (L.) Lamium, Stachys Labiatae immatures on Labiatae de la Fuente l!172
Crataegus Rosaceae adults on inflorescences de la Fuente 1972 Prunus, Rubus of these plants Clematis R.anunculaceae as above 40 found at Southwood& Hine 1950 \ Lamium Labiatae [stage?] o- roots fhibernating?] 6 in April adults feed on stems
Cydonia Rosaceae Lodos & Onder 1980 A
Ligustrum Oleaceae Lodos & Onder 1980 o- early instars in Paul 1953 s calyx of Lamium Ballota Labiatae nymphs& adults on Woodward 1949 young seedheads *
\oN) CARL W. SCHAEFER
on the ground' ffi japonensis adults live up in low treesbut do not feed;the nymphs I'eed and Schae1er,1985)' * or, ,rr. ripe seedsol Scioep/iaiasminodcra(Table 3; Tachikawa (Table 3) has been f The Garsauriinae too contains only one genus. One species specimensof G' ara- lbund under bark (a groundlike habitat) in Aliica. And five tree;the tree was cloideswere collectedin Japan below the loosebark of a large fallen 25, i983). The llattened moist but not rotted ilrt. sat
Plataspidae
Carayon (1949) writes of several Plataspis species,"inl'eodes aux Leguminosae' (in on comme la majorit6 des Plataspidae";those he studied laid their eggs ocithecae) quoted the bark of their arboreal hosts. T'able 5 attests to the accuracy of the and statement. Other plant families t'ed upon are the Rhamnaceae,Santalaceae, Acaly- Sapindaceae,all of the subclassRosidae. Linnavuori ( 1977)gives a number ol pha recordsfor A{iican plataspids(Table 5), but I am not certain that theserepresent legumes. irue leedingrecords and that the plataspidsdo indeedpre{'er this euphorbto Table 3. Food p-lgnts and habitats ol the Cydnidae: Scaptocorinae, Garsauriinae.Amnestinae, and Parastrachiinae
Insect species Plant genus Plant family Remarks Reference
Scaptocorinae
Scaptocoris divergens Musc Musaceae (monocot) Timonin 196l Froeschner feeds on roots Ruth 196[, Willis & Roth 1962 1' Scaptocoris sp. Solanwn Solanaceae Basso,et 1L 1974
S. castaneaPefty Saccharum Gramineae (rnonocot) in soil Roth 198'l Amophila Gramineae (rnonocot) amnng roots (from Froeschns & Uniola ballast?) Steiner 1983
S. talpa Champion Saccharum Gramineae (monocot) anNng roots Champiom1900
Stibaropus Saecharum Gramineae (monocot) on roots Wilbrink l9l2
molginw 6chiddt) !l Hasegawa1960, S. formosaizus (Takano Oryza, Eleusine, Gramineae (monocot) in soil and around & Yanagihara) Digitaria, roots Miyaanoto1965 Saccharum s
4 Saccharum Gramineae (monocot) a pest in Taiwan Esaki& Ishiharal95l
(!- Garsauriitrae s *under $ Garsamiahaglund bark of huge Linnawori 1977 Bergroth fallen tree" s A * Amnestinae a
Amnestus lenkoi "associatedwith Froeschner1975 Froeschner ant nests' tt r-
"associatedwith Froeschner1975 S A. pequinus ant nests" Froeschner occasionally Blatchley1926 g A. pallidus Zimmer Antmnaria Compositae understones ? (PanhleY1923)
ground,moist Froeschner1960 H A. spinifrons(SaY) (summary of literature) situations F irl F Parastrachiinae eggS' Tachikawa & parastrachiajaponensis SchoepJia Olacaceae ripe seeds; (Scott)nymphsongrourrdSchaeferl9Ss ,rqw i ,iI Table 4. Food plants and habitats of the Cydnidae: Corimelaeninae I
Reference lnsect species Plant Senus Plant family Remarks 1978 Corimelaeno obscura Sanicula Umbelliferae McPherson& Sailer McPherson & Sailer
C. pulicaria (Germar) Daucus, Chaerophyllum Umbelliferae McPherson& Mohlenbrock 1976 Apocynum Apocynaceae Rhus Anacardiaceae Asclepiu Asclepiadacae Verbascutn Scrophulariaceae Erigeron, Compositae Antmnaria
1974, C. agrella McAtee Phlox Polemoniaceae leaves McPherson Chaerophyllum Umbelliferae McPhers Nicotiana Solanaceae ripening fruit Lung & Goeden1982 4 A 1923 C. cognata Van Duzee Eucnide Loasaceae Van Duzee li o- 1972 C. l. lateralk (F.\ Daucus Umbelliferae nymphs:umbel McPherson s adults: cup Chaerophyllum Umbelliferae McPherson& 1976, 6' Elymus Gramineae (monocot) Mohlenbrock s Erigeron Compositae Van Duzee 1923 Geranium Geraniaceae (,(t rr: I flower spikes Van Duzee 1904 C. nitiduloides (Say) Planrago Plantaginaceae 5 seeds McPherson1979 C. nigra Dallas Ranuncalus Ranunculaceae o Stoner1922 Thyreocoris ater Agrotis Gramineae (monocot) t. (Amyot & Serville) = w Stoner 1922 Plantago Plantaginaceae o 1922 ..blue grass" Gramineae (monocot) stoner seedsot Castilleia Stoner 1922 lrt Castilieia Scrophulariaceae a'l rrt de la Fuente 1972, F T. scarabaeoides (L.) Ranunculus Ranunculaceae Lodos & Onder 1980 Singer 1952 Sambucus Caprifoliaceae in plant litter on drY ground Wagner 1966 adults under mats of McPherson & Galgupha arerrima Veronica Scrophulariaceae Euphorbia maculata; Mohlenbrock 1976' McAtee & Malloch Carex Cyp.ra"ea" (monocot) Wheeler l98l Euphorbia Euphorbiaceae nymphs on inflorescences and developing caPsules (Wheeler l98l) see G. aterrima McPherson & G. atra Arnyot & Plantago Plantaginaceae Mohlenbrock 1976, Serville Rubus Rosaceae Wheelerl98I Helianthus Compositae Euphorbia Euphorbiaceae $ailer l94l G. loboprostethia Medicago Leguminosae Sailer klcPherson& McAtee & Oxalis Oxalidaceae on caPsules G. carinata Mohlenbrock1976 Malloch on flowers Callan 1948 G. schulzii(F.) Boerreria Rubiaceae '..'...*:'iffi ,",",,.,',, :'',':{,;'6:dii4iii$ii{t;.:a-.ir],i,';r:t,i:ii, iii "'i ''i$a*i#i':l'" 1:-ieuliatt'i G. t'intulatu Germar Boerreria Rubiaceae on flowers Callan 1948 (i. olalr'.r Hussey Plantago Plantaginaceae on llowers Biehler & McPherson I 982 Allocoris gilletti Galium Rubiaceae damages fruits Batra 1984 Van Duzee Cl,dnoides ciliatus in loose sand around Hart l9l9 orientis McAIee & grassroots Malloch ^J t .o \ ts' 6 * ffi 36 CARLW. SCHAEFER Thelamilymaybecharacterizedasspecializingonlegumes,sharingthisdistinc- (Schaefer, 1980; Schaei'erand o'Shea, 1979; ,'iT tion with several coreoid groups (Sweet and Schae{er' Schael'erand Mitchell, 198J), the Malcidae: Chauliopinae i groups' The t] 1gg5)and perhaps a {'ew-but certainly not many-other heteropteran part o{ the rrt t'ew statementsavailable indicate plataepl'Jsf'eed less oh reproductive .',X plants, than on vegetative parts (Table 5)' with the li{ These insects feed ,rp on plants. But plataspids too are associated host plants in l ground. The nymphs ol European Coptosoma spp. "live on and below p' Maxwell-Lefioy g iu.g" ugg..gatio.ts" (Davidov6-Vilimov6 and Stys, 1982: 144)' ""' it is probable ii anJ Howt"tt (1909)confirms ttris observation,lor indian species: o{ the soil and at $ that nymphs o1'this group live wholly in concealmentat the surlace sp'' "a '! the roots of plants" (p. 6'72).Finally, Green (1900) reports a CoptosomQ (p' 185)' colony o1which was Leing attendedby a specieso1Crematogasler" ,] i ',1 Megarididae, Canopidae, Cyrtocoridae, Lestoniidae haustorifera Almost nothing is known about the biology ol these{amilies. Lestonia but not on a China occurs on the gymnosperm Callitris preissi (Cupressaceae)' related species(McDonald, 1970)' (F.) has been The other three lamilies are Neotropical. Cyrtocoris gibbus ol cyrtocoris recorded from the legume Mimosa (costa Lima, 1940),and a species with thirdlitth neartrigonus (Germar) has been collectedin large numbers(together (J. personal instars)on the ground below someunidentified low dried plants Eger, Barner and M' communication). Two speciesof Megaridid ae,Megaris puertoricensis Wol- semiamicta McAtee & Malloch, l'eed on Eugenia (Myrtaceae) (Barber' 1939; not cotr, 1936,respectively), the latter on the plant'sllowers (Wolcott, 1936).I have {bund any mention ofthe biology ol the Canopidae' GENERAL f'eedon Of some interest are the plant groups that thesebugs do not leed upon. Few the monocots, gymnosperms,or (except for the Plataspidaeand some cydninae) on primitive of Iargest of tfie Oicoilamilies, Leguminosae;and I'ew {'eedupon the most o{' the dicot subclasses,Magnoliidae and Hamamelidae. Almost all other members the Pentatomoidea, as well as of the related super{'amiliesLygaeoidea, Coreoidea, and Pyrrhocoroidea, also shun these plants (Schael'er,various papers' and unpub- lished), although there are exceptions.Plants of the most advanced dicot subclass, Asteridae, u." ulro little used, although the Sehirinae I'eedon rather more of them than do members o{' the other heteropteran groups listed' Legumes are perhaps protected liom these bugs by chemical def'enses(Janzen, 1969).The lack of {'eedingon the others, however-all relatively primitive plants,- suggeststhe major groups of thesebugs arose coeval with the plants they f'eedupon. UaO ttt.y arisen early and been t'eedingon sarly llowering plants, it is unlikely they had switched to the later plants when these became available (if the early ones Eq F'ood Plants o./ Pentatomoidea 37 ;i l produced'adequateseed). For thesebugs are-as family-gro$ps-quite polyphagous' no on. group beingtied to a particulargroup ofplants (alwaysexcepting the Platas- pidae).being gene.alleeders, it is unlikelythese bugs would havefbrsworne the early ilo*..ing plants;rather, they would havecontinued to {'eedon them aswell as on the later plants.That this has noi uiCi.ifrcd, rir.lt tht:carlitr plantsare almoststudiously ignored,suggests the bugs arose later than these early planrs' On the other hand, it may be that the earlyllowering plants(principally gymno- sperms,magnoliids, hamamelids) produced seeds not suitedto the earlyseed-I'eeding pentatomoids.The early bugs may have been I'ew, and their radiation may have awaitedthe arising ofthe large angiosperrnsrrbclassts, especially the Rosidae.This would placesomewhat more recentlythe radiation ofthese bugs,and placelurther into the pasttheir origin. Certainly thesebugs are old: they with their severalautoplesiomorphies I believe olderthan many other pentatomoids:and they are alsoold enoughthat considerable host-switchinghas occurred:this at leastpartly explainsthe variety ol hostswithin lamily-groupsand evenwithin genera.But it is not at all certainthat host-switching hasoccurred in a largeway by the ancestralbugs {rom early{lowering plants to more recentones. For now the questionremains open. Answers may be suggestedby fbssil evidence(of the bugs)ond strrdvnf'fhe early lloweringplants'seeds'suitability. Ihe Plataspidaespecialize on legumes,as do severalother land heteropteran groups (seeabove). lt is likely that thesegroups' ancestorsbroached the legumes' deterrentdelenses and the groupsradiated on this abundantand nitrogen-richf'ood. Although root-l'eedingis olten assumed,particularly in the Cydnidae,many of theseinsects I'eed upon seeds(see above, under Cydninae).Only the Scaptocorinae {'eedexclusively on roots. The othersprobabiy leed on what is available.lt is likely that seed{'eeding is primitive: seed{'eeding is widespreadin the relativesofthese groups(Schael'er, various publications); and seed{'eeding occurs so liequently in the literatureon cydnidsand their relatives(see Tables) that it is quite possiblythe way of leedingo{ thesebugs' ancestc'rs"It is lessclear that membersof the other pentato- moid lamilies {'eedon seeds. l-iving on the ground too is widespread,correlated ofcourse with {'eedingon tallenseeds and on roots (althoughI maintain the on-groundhabit camebefore the seed-and root-leedinghabit [Schae{'er,I 98 I ]). Eventhe Plataspidae,which appearto leed on above-groundvegetative parts (Table 5), have some associationswith the ground. Such associationsoccur liequently throughout the major groups(Schaefer, 198l); they are frequent too in the Homoptera (Schael'er,in press,in prep.). ACKNOWLEDGEMENTS I am deeplygrateful to Dr. M. Tomokuni lbr unpublishedobservations, and to him and Dr. S. Tachikawafbr openingto me the Japaneseliterature on thesebugs. Ms R. (iovern expertly renderedtyped order fiom the scribbleclchaos given her. ' d*sfij;fu;;;;ad'&{isiieb# *;;'jh;i*r;,*#3,w r @ Table 5. Food plants and habitats of the Plataspidae o Remarks Relerence Plant genus Plant lamily lnsect sPectes - Yang 1934 Desmodium Legumrnosae Coptosoma varregalum Yang 1934 1 Dodonaea Sapindaceae (t Montandon Chatterjee 1934 Santalaceae o Santalum Chatterjee 1934 Rhammaceae Zizyphus H Tl Chatterjee 1934 F Santalum Santalaceae C. siamicum Walket Chatterjee1934 Sapindaceae Dodonaea Chatterjee l9-14 Rhammaceae Zizyphus Chatterjee 1934 Rubiaceae Canthium Chatterjee 1934' Leguminosae Pterolobium, Singh et al. 1978 grain legumes Chatterjee 1934 Bulea Leguminosae C- ostensum Distant Chatterjee 1934 Santafum Santalaceae C. Juscomaculatum Distant Chatterjee 193'4 Zizyphus Rhamnaceae Heiss 1977,Voigt Astragalus, Leguminosae 1974 C. scutellatum 1978,Dobsik Coronilla" (GeollieY) OnobrYchis Davidov6-Viliruovi Onionis, Leguminosae C. mucronatum & StYs1982 Medicago Seidenstr'icker Singh et al. 1918 grain legumes Leguminosae C. angularis (StLl) Singh et al. 1978 grain legumes Leguminosae C. atomaria (Stti) Singh et al. 1978 grain legumes Leguminosae C. cincta (KirkaldY) C. noualhieri Xanthiurtt Compositae Moizuddin&Ahmad 1975 Montandon C. xanihogramma Canavalia, Leguminosae Beardsley& Fluker 1967 (white) Strongylodon C. nubilum (Germar) grain Iegum€s Leguminosae Singhet al. 1978, Acalypha Euphorbiaceae Linnavuori 1977 C. marginella Dallas grain legumes, kguminosae stem & fruit Golding 1927 Desmodiwn, stem Cajanus, stem Dolichos stem Gliricidio, stem Crotalaria stem C. punctissimum Glycine [,aguminosae stem, petiole, Kobayashil$l . Montandon leaf, pod C. biguttula Glycine Leguminosae stem, p€fiole, Kobayashil98l i, Motschulsky leal, pod A C. japonicum Glycine Leguminosae stem, petiole, Kobayashil98l ts Matsumura leal, pod 4 C. hirsutum Chaetacme Leguminosae Linnavuori 1977 Montandon * o- C. nigropunctatum Acalypha Euphorbiaceae Linnawori 1977 s Stel 1977 C. p y gmaeum Jensen-Haarup unidentified Leguminosae I-innavuori R. s C. rt4lt (Montandon) grain legu.mes Leguminosae Singh et aL 1978 {^} C. costale StLl Psoralea Leguminosae Seidenstricker19{i3, Lodos \o & tinder 1978 40 CARL W. SCHAEFER K.s€ ii i E Eh€ i:6 '! 5 6r ?ats E 5 EE E 5E$ ;;:H Etl c gEE i5s; ! E ::qdBtE EEp'q.66f g €3SsETsi € E :5 s E A e 3 gEgiEeeaEgg FF BE p a a f E E B e ssE "s E€ E$s*S i i i i _E E6 *t s $ $ S SBi':. i i i s a s g $ss sFgs- :uFe'i: $ $€i i t iE? g sao tF et s:aa €F EE s E e; Es i : E b-E :? SE * 3 SE I E tg s t so s= n i + Es-E SO i. !z qi qi 'd cti i Lj qj Li E cd ti { d#&i&{"".,c,e*#ic;tri,.:,...,ix6*.s&i**kfuii*riiji \{iias il5M&*e4'r#{&diliil#;tus*Ills ry' P. cottinellokles aS atrove Carayon 1949 (Laporte) Carayon 1949 P. Jlavosparsa Albizia Leguminosae Montandon Megatopto lobata Glochidion Euphorbiaceae Young buds and Ren 1984 (Walker) leaves fu|. lusrvathi Vigna, Leguminosae pest on tender Ren 1984 (Montandon) Puerorio, shoots other Iegumes 1958,Singh M. cribraria (F.) Acacia, [,eguminosae shoots Banerjee Pueroria, 1978,Kershaw 1910, Sesbania, Yang l9$4, Chatterjee Cajanus, 1934, lull:mad & Dolirhos Moizudidin 1976 "important pest ol Ren 1984 .il Iegrumesin China" a % .o o 6 G 6" s ffil /1'' CARL W. SCHAEFER ',fr'-q{ g REFERENCES j plataspids(Heteroptera: and M. Moizuddin (1976) Biological control measuresol bean li Ahnrad, l. Pentatomoidea)in Pakistan. Prot' Etilomal' Sac' Karachi 6: 85 86' i Hortob6gyNational G. and T. Vasarhelyi(1981) Contribution to the Heteropteranlauna ol the t Bakonyi, Park, l. The I'auna o./ the llortobhgl' Natirtnal Park, l98l:55 63' speciesol Hemiptera ,i Banerjee,M.R. (1958)A study ol the chromosomesduring meiosisin twenty-eight '1 (Heteroptera,Honoptera). Prot'. Zool. Sot Calcutta ll:9-31 ' Barber,H.G. (lg3g) lnsectsol PuertoRico and lhevirgin islands lleniiptera-Heteroptera(exceptingthe l4(3): 263 441' Miridac and corixidae). stieniilit survq' o./ Puerto RiLrt and the virgin Islands rf em santa Maria' RS' Basso,L.V., D. Link, and o.J. Lopes(1974) Entomolauna de algunassolanaceas Ret,.Centro Cient. Rur 4:263 2'70' $ Eurasiaand N orth Amertca' Batra,S. W.l'. ( I 984) Phytophagesand pollinatorsol Galium ( Rubiaceae)in i Environ l:nrontol. 13: I I l3 I 124. (Hemiptera:Plataspidae) a Beardsley,J.W., Jr., and S. Fhrker( 1967\CoPttt,tt;tvtu xilittlxo,t'atllmd (White)' new pest ol legumes in Hawaii. Pror'. [lau'- Entontrsl S'ot l9: 367-'312' o|a/rs(Hemtp- Biehler.J.A. and J.E. McPherson(1982) l-i{e historyand laboratoryrearing ol Galgupha Amet. 75:.465 4'70' tera: Corimlaenidae), with descriptionsol immature stages.Ann. Entontol. Soc. Nature Publ' Blatchley.w.s. (1926)Heteroptera or True Bugsol EasternNorth America.lndianapolis. Co. ll16 pP. (Heteroptera: Bol. Lab' Boselli.F.ts. (1932)lnstinti materni del Sehirussexmaculatus Rhr. Cydnidae). Zool. Portiti 26; 2 7. Verii.l/.Natur- Burghardt,G. and C. Rieger(1978) Die Wanzenlaunader sandhausenerFlugsanddinen. s('hutz. L.an(ls(ha/rsp/legeBad -Wurtt 47 148: 393 411' B.'N.1. Prot Rtt.t'al Callan. E.M. (1948)l'he Pcntatomidae,Cydnidae and Scutelleridaeof Trinidad, $ Entomol. Sot. I'ontlon 17: ll5 124. gull. Sot Entontol. ,,t Carayon, J. (1949) L'oothdque d'H6miptbresPlataspides de I'A{iique tropicale. :! France 54:66-69. Perty,lound at the roots ol sugar-cane.Entonol. Mon. -;i Champion,G.C. (1900)A speciesol Scuptotoris, Mag 36:.235 237. (24). Pentatomidae Chatteriee,N.C. (1934)Entomological investigations on the spikedisease o{ sandal ,$ (Hemipt.). Indian Foresr Ret. 20: | 31. genera China, W.E. (1931)Morphological parallelism in the structureol the labium in the hemipterous Coptosomoides, gen. nov., and Bozius, Dist. Ann. Mag. Nat. //i.rl.7(10):281 286. da CostaLima, A. (1940)ln setosdo Brasil.2' Tomo, Capitulo XXll. Hemipteros.Rio de Janeiro.Escola Nacional de Agronomia. 351 PP. Cronquist,A. (1968)The Evolution and Classilicationol Flowering Plants.Boston, Houghton-Mifllin. 396 pp. Davidova-Vilimov6,J. and P. Stys (1982) Bionomics ol European Coptosomaspecies (Heteroptera, Plataspidae).A(ta [Jniv. Carolin., Biol. l9E0t 463-484' Dethier, M. (1980) H6miptbresdes pelousesalpines au Parc National Suisse.Rev. Suisse7'ool.87t 975 990. Dolling, W.R. (l93l) A rationalizedclassilication ol the burrower bugs(Cydnidae). S...'st. Entontol.6t 413 431. Ehanno. ts. fi961) Contribution ir la connaissancedes H6teroptbres Pentatomides armoricaines. Bull. Sot. Sti. Bretagne 36: 47-58. Esaki,T. and T. Ishihara(1951) Hemiptera of Shansi,North China. ll. Pentatomoidea.Mu.rhi22:29 44. Froeschnner,R.C. (1960)Cydnidae ol the Western Hemisphere.Prot. U.S. Natl. Mus. lll:337 680. Froeschner,R.C. (1975)Three new speciesol burrowing bugs lbund in associationwith ants in Brazil (Hemiptera:Cydnidae). J. Kans. Entomol. Sor. 48: 105 ll0' Froeschner,R.C. (1980)Notes on a collectionof Cydnidae(Hemiptera) Irom Ecuadorand the Gallpitgos Islandsmadeinlg64lg65bytheExpeditionN.etJ.L,eleup.Bull. lnst.Ro.v. Sci.Nal. Belg.(Enromol.) 52" | 4. Froeschner.R.C. and W.E. Steiner,Jr. (1983)Second record ol South Americanburrowing bug, Scapto- coris castaneusPerty (Hemiptera: Cydniclae),in the United Slates. Etttttntrtl. Nen's 94: ll6. F orttl P!unts o,f Pentatontoidea 43 . I*.t1. Il. de]aFuentc..l.A.(1972)ll.cr'isiorrdclosPentat6midr:sib6ricos.FanriliaCydnidaeBillberg,l820 Sttt. I:..:;1tuit.llirr' \ttr' i/inrl') 70: -3378' wagner (Hemiptera:Heteroptera : Cydni- ( 1975)(). ,, ,,cu,srrbspecies ol Aathttslatircllis chauri, ivl.S.K. Sot. 72: dire)asaserioilspcsroll'?,nniserunlt.vphoities(Euirl1.)in'lnrlia.'l.llomha-t'Nol'Hisl' 226 229. Golding'F.D.(1927)N()lcsonthcl'()od-plantsandhahitsol.somesouthernNigerianinsects.B,/1. (.)5 Entotnol. /ics. lll: 99. Green,E.E,(1900)NotcoI]thcatt|activcpropertiesol'certainlarvalHemiptera.Entomol'Mon'Mag.36]!' l 85. Die wanzentaunades l-aturschutzgebietes"Mainzer Grinther,H.. C. Ricgcr.and (i. llurghardt (1982) Mainzer Naturv'..Arth' 20: l-36' Sand" und benachbartcrSandgciicte (lnsecta: Heteroptera)- keysro the nearcticgenera. Bull. lllinoisNatur' Hist' Hart, C.A. (lglg) T.hcl,eutatomoiJca o{ lllinois with Surv. 13; 157 223. Hasegawa,H.(1960)LlctcI()ptclitolNiigataPref.ecture,Japan.BuIl.NagaokaMunit'ipalSd'Mus.(|): l9-65. (in JaPanesc) Hasegawa,H.(1966)[lemrptcra.itrYKurosawaandtHidakaed'lconographiainsectorum'Sh6gaku- kan, Tokyo, 476 PP.(in .laPancse) (lnsecta:Heteroptera). vI: PentatomoideaVerijf|'. Heiss,E. (1917) Zur Hctct.optcrenllurraNordtirols Mus. Ferdinantleurtt(ltttt:hru4 )57:53 77' Hertzel.G(1983)llcitragczutlnsektcnl;tuniiderDDR:Heteroptera-PlataspidaeundCydnidaeFcanli' Abh. Staatt.MLr's. lit'rl;. l)rtvlt'rt l0: l1l-i24' llight activity ol the burrowing bug' Highland, H. B. and P.F . l .u nr rn is ( 1986) u sc o{ light traps to monitor field inlestations in peanuts' J' Econ' Pangaeusbilinearus (Hcntiptcra: Cydnidae), and associated Entomol. 79:523 526. (in Japanese) Hiura, l. (1958)The topics ol bugs. Naltrre Stud! 4:100-102 (Hemiptera,Pentatomoidea)' Hiura, l. ct al. (1959)l,cntatonlid bugs in Tokushima Pre{ecture.Shikoku Konchff-Kagaku.(9) 9 46. (in 'lapancse) Hoebeke,E.R.andA.(i.whcctcr..lr.(l9tii4)Aethusnigritus(F.),aPalearcticburrowerbugestablishedin soc. wash' 862138-744' easternNorth Amcrict (llcmiprcra-Heteroptera:cydnidae). Prot. Entomol. Entomol lS:82 84' Horvath, G. (1899)Notc srrr lc Schirusmaculipes et especesaflines. Rev Dept. Agric. Farmers Bull' Howard, L.c. (19(n) lhe principal inscctszrflecting the lobacco plant. u.s. 120: | 12. Georgia.J. N.Y. Entomol' t Hussey,R.F. (f 925) Sonrc ncu or little-knownHemiptera liom Florida and i Sot. 33:' 61 6'1. 55-69.(in Japanese) $ lshihara,T. (1917)n n rr)rro(lucrionto Pcntatomidaeliom Japan. Mushi-shizenl7t I Evolution23" | 27' Janzen,D.H. (1969)Secd-caters versus seed size, number, toxicity and dispersal. Instituts' Josiiov. M. (1970)I:rgcbnissc dcr Albanien-Expeditionl96l des DeutscheriEntomologischen 82. Beitrag. Hetcroptcra. Ileitr' Iintomol. 20: 825 956. $ Macao Ann' Soc Kershaw,J.C.W. (1910)on thc mctamorphosesof two coptosomineHemipteraliom 3 j' Entornol. llelg 5l: 69 7 i (Hemiptera). Kobayashi,T. ( 1953) 1-lre tlevcltrpmental stages ol six speciesof the JapanesePentatomoidea 'l.qrit. St'i. Rep. Mtttsu.ttrttttt Coll' (ll): 73 89 $ Kobayashi,T. (1954a)lhc hiology of some pentatomoids(2). Shin Kont'htr7(10\:22-24.(inJapanese) l 'l i Kobayashi,T.(1954b)hclcaringmethodolthepentatomoids. ShinKont'htt7(ll):ll 14.(inJapanese) j Kobayshi, T. (1964) t)$clopmcntal stagesol Geotomus pygmaeus (Dallas and Sehirus niveintarginatus (Scott)(Cydnidrc). A'orrr.tir 32:21 27. * a Kobayashi,l . ( 1974)A note on Acthus indius (Westwood) and GeotomuspNgnlaeus (Dallas) in lshigaki * l2l. (in { Island. /lo,rrlro2.1: .lapanese) Kobayashi,l.(l9lll)lnsectpcstsof soybeansinJapan. Mist:.Publ.TohokuNatl.Agric.Exp.Sta.2:l39. g Lindberg,FL (19.12)lnvcnta cntomologicaitineria Hispaniciet Maroccaniquod ab 1926{ecerunt Harald et Hakarrl.indbcrg. Xlll. tlcmiptera Heteroptera(excl.Capsidaeet Hydrobiotica). Comment. Biol.3: I 55. l.innavuori. I{. (1960) llcmiptera ol lsrael. L Ann. Zool. Soc. Zool. Bot. Fenn. "Vanamo" 22: l-71. l.innavrrirri.lt. (197.1)Studics on Aliican Heteroptera.Arq. Mus. Bot'age(Ser. 2) 4: 26-69. I-innavuori.I{. ( 1977)Hcnriptera ol the Sudan,with remarkson somespecies ol the adjacentcountries. 5. ffi 44 CARL W. SCHAEFER l,* Acta zool. Fenn l4T: l-81. Tingidae, piesmidae, cydnidae, Thaumastelliiae and Plataspidae. (Heteroptera) rist lamilyasi r'izerindearastirmalar' L"d*,"N and F. 6nder 1tlaOl fUrnye Pentatomoidea Yavin 38l: 81 pp' III. Familya Cydnidae Billberg. Ege Univ' Ziraat Fakillt extensaot tree tobacco' Nocotiana glauca' Lung, K.y. and R.D. CoeOen(19J2) Biology oi Corimelaena Ann. Entomol. Sot. Amer.75: 177-180 Lile. London' Thacker, Spink & co' 786 pp' Maxwell-Lelioy, H. and F.M. Howlett (1909) lndian Insect (Palisot de Beauvois) (Heteroptera: McDonald, F.J.D. (196g) Some observations on Sehirus cinctus Cy (Heteroptera' Plataspididae)' Reichenbachia20z Seidenstricker'G. (1963) Anatoliens Coptosoma-Arten 155,160. Singer'K.(1952)Diewant-ei1(Hemiptere.Hetgroptera)desunterenMaingebietesvonHanaubis(n F 5: Naturwissensch.fufus. stadt AschaJJbnbarg ) wrirzburg mit EinschlussJ., sp"rruur. Mitt. I 128. Singh,S.R.,H.F.vanBmden,andT.A.Taylor(1978)Checklistofinsectandmitepestsofgrainlegumes' London' Academic Press' of Crain Legumes: Ecology and Control Pp 399 417lr Pests cmctus and laboratory rearing ol sehirus cinctus Sites, R.w. and .1.E. Mcpherson (lgg2) Life.history l'?n Enro.mol'Soc' Amer'75t210 215' (Hemiptera: Cydnidae),*i* i.,"'iptions of imniature stagcs' smith,J.w.andJ.l-.pitts(1g74)peststatusolpangaeusbilineatusattackingpeanutsinTexas'J'Econ' Entomol.6T:lll-113. (Hem'' on the biology of sehirus bicolor (L') Southwood, T.R.E. and D.J. Hine (1950) Further notes Cydnidae). Entomol. Mon. Mag' 86: 299-301' qnc Bugs o{ the British Isles London' F' Warne & southwood, T.R.E. and i). l-eston( 1959)Land water Co., Ltd. 436 pp. Stehlik,J.L.(1983)Resultsol.theinvestigationsonHemipterainMoraviamadebytheMoravian 172' Museum (Pentatomoidea ll) Atta Mus' Moraviae 682 153 Stud' Nat' /lsl' 8: l-140' ,tStoner, D. (1920) The Scutelleroideaol lowa' Llniv' Iowa third European speciesof Plataspidae Stys, P. and J. Davidova-v'itimova( 1979)Coptosoma sandahli-a (Heteroptera). Acto Etltottt lloheittosi T6: 140-142' bioiogy al Chauliops.fal/at scott, with a Sweet, M.H. and C.w. Schaeler(19g5) Systematicstatus ancl (Hemiptera: Lygaeidae). Ann. discussion of the phylogenetic relationships of the Chauliopinae Entomol. Soc' Anter. 78: 526 536' 1l(10):220-223. Tachikawa, S. (1974) on parental care in Heterop|era. Inse1tariunr japonensis (Hemiptera: Pentatomoidea: Tachikawa, S. and C.w. Schaeler( 1985)Biology ol Parasttachia '!-\dae).Ann. Entontol. Sot. An*r' 7Ei 387-397' and shieldbugs Thomas, D.C. (1954)Notes on the biology ol someHemiptera Heteropteral.-Introduction (Pentatomoidea\.Entonutlttgist 87: 25 30' # Can'J'Bot'392 Timonin,M.I.(1961)Theinteractionolplant,pathogen,andscaptocoristalpaChamp $&l 695-703. H Don and Acacia saligna(Labill') Van den Berg (1980)Natural enemiesol Atutia t'yclopsA. cunn. ex G. Wendl. in western Australia lll. Hemiptera, Phytophylactica l2i 223-226' ffi ( 11emiptera aitacking Acacia dealbata Ltnk., Acacia decurrensWllld. Acacia Van den Berg, M.A. 1982) melanoxylon R. Br. in Australia. tongifolii 1A,nar.)Willd., ,4r'atia mearnsil De Wold. and Acacia PhYtoPhYlactiru14: 47 50' north of Mexico, with uun Dur.", E.p. (1904)Annotatcd list ol the Pentatomidaerecorded liom America descriptionsol somc new species.Trans. Amer. Entornol Soc' 30: I 80' Gulf of California in van Duz"e, E.P. (1921) Ilxpedition ol the Calilbrnia Academy ol sciencesto the 1921.Proc. Calil. Atad 'Sr'i (4) l2: 123-200' Bad.' Voigt, K. (1978)Dic Wanzcn dur RussheimerAltreheingebiets. Natur. u. Landschaftsschutz.-gebiete Wurtt', l0: 401 444 Voigt, W. (1980) Ein urlcr\r'artctcsVorkommen von Canthophorusimpressus (Horv.) (Heteroptera, Cydnidae) im llolimischcn Mittelgebirge (Cesk6 Stredohofr). Acta Faun. Entomol. Mus. Nat. Pragae 16: 129-130. 'l Wagner. E. ( 1966)Die ierrvclt L)eutschlands.54. Teil. Wanzen oder HeteropterenL Pentatomorpha [sic]. Jena, G. FischerVcrlag 235 PP. Wheeler. A.G., Jr. (l9lll) lnsect associatesol spurges,mainly Euphorbia Maculata L., in easternUnited States. Prrr'. Iitttttntol. ,St\'. lf ash. 83: 631 641. Wilbrink, G (1912)[)c kcdirischewortelwants.Meded. ProeJstationJava-Suikerindustrie.22:llll-1123. Willis. E.R. and 1..M. Itoth (1962) Soil and moisture relations ol Scaptocoris divergens Froeschner (Hemiptcra:('ldnidae). Ann. Entontol. Soc. Anter. 55: 2l-33. Woodward,l'.E. (1949)Noles on the biology ol some Hemiptera-Heteroptera.Entomol. Mon. Mag.85: I93 206. Yang, W. (1934)Rcvision of ChinesePlataspidae. Bull. Fan. Mem. Inst. Biol (Peiping)5:137 235.