The Structure of the Pulvillus and Its Taxonomic Value in the Land () '

S. C. GOEL AND CARL W. SCHAEFER Department of Zoology, R. K. College, Shamli, U. P., India; and Systematics and Environmental Biology Section, Biological Sciences Group, University of Connecticut, Storrs 06268 ABSTRACT The following terminology is proposed for the heter- of which consists of a basal basipzdvillus and a lamel- opteran pretarsus : The median unguitractor plate, bear- lated distal distipulvillus. The presence or absence of the ing basally the claws and terminally a small extension, parempodia and pulvilli, and the structure of the latter, the empodium, from which arises a pair of bristles, the in general confirm current ideas as to the relationships parempodia. Basal and ventral to the claws (and lateral of the major groups of land Heteroptera (Gcocorisae). to the parempodia) is a pair of pads, the pulvilli, each

~ Few attempts appear to have been made to put of the pretarsus, but they did not consider homologies pretarsal nomenclature upon a firm footing. Cramp- from order to order and, like Crampton, they ignored ton (1923) described and in part homologized most some parts (e.g., parempodium). Dashman's attempt of the structures, but his conclusions have unfortun- (1935a) was marred by misprints, a few contradic- ately been largely ignored. MacGillivray (1923) tions (e.g., his Fig. 2 and his definition of pulvillus), gave a set of terms many of which were new, some and a failure in some cases to distinguish among of which were poorly defined, and few of which have different structures (e.g., pulvillus, pseudarolia (as been adopted by morphologists. Snodgrass (1927, a singular), and parempodium). Perhaps the most 1935) and Levereault (1935) also discussed the parts successful attempt was that of IIolway (1935), who nevertheless did little to determine homologies. How-

1 Reccived for publication May 16, 1969. ever, his terminology seems the most consistent. 308 ANNALSOF THE ENTOMOLOGICALSOCIETY OF AMERICA [Vol. 63, no. 1 Despite a recent revival of interest in the compara- gest pulvilli arose from a division of a median aro- tive morphology and relationships of the major heter- lium. However, pulvilli und an arolium are possessed opteran groups, characters of the pretarsus have by many , including some studied by Cramp- been used but sparingly. This reluctance may well ton; and Levereault’s evidence consists of the phrase be a result of the nomenclatorial confusion, and this “most probably.” confusion is manifest in the names applied to the 2 The basal part of the pulvillus is a small sclerotized paired structures found beneath the claws. plate, in many orders (Snodgrass 1935), including In this paper we hope to establish a consistent the JIeteroptera. This plate is the basipulvillws terminology, based somewhat on Holway’s (1935) (Crampton 1923, IIolway 1935), or auxiliu. With and Dashman’s (1953b) definitions and Crampton’s Dashman (1953a), we prefer the former, more de- (1923) arguments (buttressed by our own) ; and we scriptive, term. The larger part of the pulvillus is hope also to stimulate interest in the comparative distal to the basipulvillus and appears to have no study of the heteropteran pretarsus by presenting name. We dislike coining new names in a field where descriptions of the most complex pretarsal structure, so many already exist, but this distal part of the in the major groups of land bugs. The structures pulvillus is of taxonomic importance: we propose are easy to work with and can be removed with a the name distipulvillus for it. The basipulvillus (BP) minimum of damage to a pinned specimen. forms a hollow, flattened (anteroposteriorly), well- Our descriptions are based largely on material sclerotized, basal region attached to the claw’s base collected in India at lighttraps by one of us (S.C.G.) , by a membrane (Fig. 1). The distipulvillus (DP) is with the principal exception of a synoptic series of flattened and somewhat cuplike, concave ventrally, supplied by Ih. J. A. Slater, University and with a smooth margin. It often bears numerous of Connecticut. The pretarsi were studied by disec- lamellae (L) (as de Meijere 1901 [e.g., his Fig. 161 tion and as whole mounts after KOH treatment. In and Tullgren 1918 also noted), bound together by a some cases pretarsi were removed from dried speci- membrane. The amount of lamellation, and the di- mens, heated in Na,P0,.12H20 (tribasic), to remove rection in which the lamellae run, are useful taxon- air bubbles, and studied as wet mounts in glycerine omic characters, as are other features of the pulvillus. at 100 and 430x. The unguitractor place has been described and dia- grammed by Dashman (1953b) for several hetero- STRUCTURE AND TERMINOLOGY pteran families. The presence and shape of the “aro- The heteropteran pretarsus consists of a median lia” and “pseudarolia” have used widely in separating plate, the unguitractor plate, from which arise later- the major niirid groups (Knight 1941, 1968). Stys ally the claws (or ungues). Ventrobasal to the (1959) separated the subfamilies of in claws is a pair of complex pads; medially, arising part on the presence or absence of these structures, from a small median extension (empodium) of the and a similar separation may be made of the sub- unguitractor plate, is a pair of bristles. families of the Thaumastocoridae (Drake and Slater The 2 pairs of structures (pads and bristles) have 1957; and see following). Wygodzinsky (1966) used been called “arolia” and “pseudarolia,” but there has some pretarsal characters in separating tribes of the been little consistency among heteropterists as to Eniesinae (). Reuter (1910, 1912) divid- which term applies to what. A few examples taken at ed the Heteroptera into major groups based partly randoin are: the bristlelike median pair is “arolia” on the presence or absence of the “arolia,” and Tull- to Knight (1941, 1968) and to Stys (1967), but gren (1918) has discussed “arolia” and “pseudarolia” “median bristles” to Usinger and Matsuda ( 1959) ; briefly. Otherwise, however, pretarsal characters they are not mentioned by Reuter (1910, 1912) or seem to have been little used in studies of heterop- by Tullgren (1918). The fleshy outer pad is “pseu- teran higher systematics, although occasional de- darolia” to Knight and Stys, but “arolia” to the scriptions have been made of these structures in in- others. dividual species. The median, paired structures (not named by In the rest of this paper we shall describe the pul- Crampton or Snodgrass) appear identical with the villus in several heteropterans from most of the fam- pareinpodia, as defined by Holway (1935) : “bristle- ilies of the pentatomomorphan land bugs (Geocori- like appendages of the empodium.” We prefer this sae) and a few species of the Cimiconiorpha. We descriptive term to either “arolium” or “median recognize that 1 or 2 species are an inadequate rep- bristles.” resentation of a family; the following data are frag- The paired pads lateral to the parempodia and bas- mentary, and conclusions are tentative. al to the claws are not arolia. An arolium is a medial single structure (Crampton 1923 ; Snodgrass 1927, DESCRIPTIONS 1935) and is probably absent in Heteroptera. These (Fig. 1) .-The basipulvillus is hollow, pads are clearly pulvilli (singular, pulvillus). The flattened, and with a distal groove. The distipulvil- pulvilli in Heteroptera appear to be homologous to Ius is oval, lamellate, and lacks a sclerotized core. homonymous structures in Diptera, although they may Species studied : Chrysocoris stockerzts ( L.), C. pur- not be if Martin (1968) is right and primitive dip- pureus (Westwood), Poecilocaris latus Dallas, and terans lacked them. Crampton ( 1923) very tenta- Canto ocellatus (Thumberg). tively and Levereault (1935) more forthrightly, sug- (Fig. 2-4) .-The basipulvillus is January 19703 GOEL.9XD SCH.4EFER : PC1,VLLLT:S CF 1-IETEROPTERA 309

4

FIG.1.-Pretarsus of Chrgsocoris pivpureus (Scutelleridae). FIG. 2-8.-Pulvilli. 2, Andrallus spinidens (Pentatomidae) ; 3, Dorpius indiciis (Pentatomidae) ; 4, Dalpada versicolor (Pentatomidae) ; 5, l'essavatoma jazianica () ; 6, Coridius jarzits () ; 7, Coptosoiiza contecta () ; 8, dethus nigritrcs (). RP, basipulvillus ; C, claw ; DP, distipulvillus ; EMP, empodium ; L, lamellar area ; S, sclerotized area ; U, unguitractor plate.

grooved distally and the distipulvillus is oval in with a sclerotized core at its junction with the basi- Andrallus spinidem (F.) and Cairtheconidea furcel- pulvillus, in Dalpada versicolor (Herrich-Schaeffer) lata (Wolff) (both Asopinae). The following spec- and D. tenzpoketsaensis Cachan, of the pentatomine ies, except where noted, are of the pentatomine tribe tribe Halyini. Pentatomini. The basipulvillus is grooved distally In general, pentatomids resemble scutellerids in and the distipulvillus is triangular in Bngruda cruci- lacking the sclerotized core (except Dalpada) and in ferarum Kirkaldy, whereas the distipulvillus is rec- having some breaks at the tip of the lamellae. tangular in Nezarn antemata Scott, Agonoscelis Tessaratomidae (Fig. 5);-The basipulvillus is flat- rutila (F.) and Eusarcocoris montivagus (Distant). tened. The distipulvillus is large and rectangular, its Thc basipulvilliis is straight distally and the distipul- lamellae originating directly from the junction with villus is triangular in Carbula insocin (Walker) and the basipulvillus in Agapophyta bipztnctata Boisduval, Dorpius indicus Distant (Aeptini), and the distipul- but from a sclerotized core at this junction in I'essar- villus is rectangular in Adria parvula (Dallas). The atoma javanica (Thunberg) and Lyranzorpha diluta distipulvillus is lamellate and lacks the sclerotized Stil. In A. bipunctata the lamellae show breaks at core in all of the above-named species. The basi- their distal end. pulvillus is straight distally, and the distipulvillus is Dinidoridae (Fig. 6) .-The basipulvillus is elon- rectangular but with many breaks in the lamellae and gate and transparent, and the distipulvillus is large, 310 ANNALSOF TIIE ENTOMOLOGICALSOCIETY OF AMERICA [Vol. 63, no. 1 ovate, and with only a few short lamellated setae (Fig. IO).-The basipulvillus is indis- medially. Species studied : Coridius jams (F.) (de- tinguishable, apparently not differentiated. The dis- scribed in more detail in Goel 1967). tipulvillus is triangular, transparent, and lacks lamel- The Dinidoridae seem, with respect to the pul- lae. Species studied: Glochocoris sp. There is much villus, transitional between those pentatomoids with variation in the Aradoidea (Usinger and Matsuda lamellae and those without them. 1959). Plataspidae (Fig. 7) .-The basipulvillus is lightly (Fig. 11, 12) .-The basipulvillus sclerotized, hollow, and lacks a distal groove. The is well sclerotized, flattened, hollow, and sickle-shaped distipulvillus is rectangular and lacks both the scler- with the pointed end dorsal. The distipulvillus is lo- otized core and the lamellated setae. Species studied : bate. Lamellae are basal ; the margins of the pul- Coptosowta contecta Montandon, Coptosoma sp., and villus are transparent and lack lamellae in Dysder- subaeneus (Westwood). cus koenigii (F.), Scantius clavimafius (F.) (Pyr- Cydnidae (Fig. 8) .-The basipvlvillus is crescentic rhocoridae), and Macroceroea grandis Gray (Larg- and narrow distally. The distipulvillus is elongate idae) ; but the lamellae are more distal in Odontopus (straplike) and transparent, lacking lamellae. The aigricovnis Stll () . distal end of the pulvillus is blunt, probably a fos- The pyrrhocoroid pulvillus appears generally inter- sorial adaptation as the cydnid tarsus is vestigial. mediate between that of the and that Species studied : Macroscytus expansus Signoret, of the and . Aethus indicus Westwood, A. nigritus F., and Stib- Lygaeidae (Fig. 13, 14; Table l).-The lygaeid aropiis callidus Schiodte. pulvillus is rather consistent in structure (Table 1) Urostylidae (Fig. 9).-The basipulvillus is cres- and rather like that of related groups. The basipul- ceiitic distally, and the distipulvillus is flattened, with villus is more or less sickle shaped, except in Myodo- numerous lamellae, and rectangular in Urolabida cha () and Drymus () where nilgirica Yang, but triangular in I/. histrionica West- it is nearly straight, and in (Stygno- wood (Urostylinae) . corini) where the distal third is bent at a right

FIG.9-l7.-Pulvilli. 9, Urolabida nilgirica (Urostylidae) ; 10, GZochocoris sp. (Aradidae) ; 11, Dysderczts koenigii (Pyrrhocoridae) ; 12, Odontopus nigricornis (Pyrrhocoridae) ; 13, SpiZoJ tethus parsdzhr?ls (Tygaeidae) ; 14, (Lygaeidae) ; 15, Howkoeocerus lacertosus () ; 16, Akharatm fasciatus, (Alp didae) ; 17, CarvaZhoia arecae (). BP, basipulvillus ; DP, distipulvillus. January 19701 GOELAND SCHAEFER: PULVILI.US OF ~IETEROPTERA 311 Table 1.--The Pulvillus in the Lygaeidae. ill-__- -_ -. 111111 --- Rasipulvillar Subfamily or tribe Species shape Distipulvillar lamellae _II-. .. II ______-_ Spilostethus pandurus militaris sickle-shaped distal %-% indistinct (F.), S. p. elegans Wolff, S. (Fig. 13) hosfes (J?.), Graptostethus ser- vus Scopoli Orsillinae Nysius raphanus Howard ,, ,, to tip Chauliopinae Chauliops rutherfordi Distant sickle-shaped, slightly almost to tip incised ventrally RIalcinae Malcus flavipes StHl sickle-shaped not to tip Geocorinae Geocoris aiiaabilis Stal sickle-shaped, but to tip, becoming less dis- somewhat more tinct and more irregu- straight lar distally Oxycareninae Crophius bohenzani (StHl) as Geocoris indistinct, absent from dis- tal third Pachygronthitiae Pachygrontha bipunctata Stil sickle-shaped to tip, those of distal % weak, not continuous with those proximal Hete rogas t rinae Heterogaster longirostris Wag- ,, ,, not to tip ner Blissinae Extaredemus macer Van Ihee not to tip, lamellae well sclerotized Rhyparochrorninae Plintltisus lougisetosus Barber 11 1, not to tip, lamellae few Plinthisini and weak Lethaeus gzctfulatus Stal 1, 9, not to tip, medial lamellae approaching tip more closely than lateral Drymini Drynaus unus (Say) almost straight, turned like Pachygrontha up at tip Myodochini serripes Olivier tapering slightly, bent not to tip at tip Megalonotini Megalonotus chiragrus (F.) sickle-shaped like Mjiodocha Gonionotini Elnblethis zirarius Horvath 11 11 not to tip, well sclerotized Rhyparochromirii Dieuches leucocerns (Walker) , 1, iestricted to base Stygnocorini Stygnocoris sabulosus Schilling tip sharply bent (Fig. reduced, perpendicular to 14) long axis (Fig. 14)

I-___I ___ I_ ____ --- _____ -- 11111 __

angle (Fig. 14). The end of the basipulvillus points and Leptocoris tagalica Hurmeister (Serinethinae) . dorsally. l'he distipulvillus bears many flattened lam- A fuller account of representatives of all the rhopa- ellae which rarely extend all the way to the tip, line tribes will be published elsewhere by one of us and when they do, are often weaker distally. The (C.W.S.). lamellae are much reduced in Plinthisus (Plinthisini) Stenocephalidae.-As in 1,ygaeidae. Species studied : and in Stygnocoris. In the latter genus those lamellae lateralis Signoret. present are perpendicular to the long axis of the dis- Miridae (Fig. 17) .-The basipulvillus is sclero- tipulvillus, which is not circular like those of other tized basally and short, and the distipulvillus is lo- lygaeoids, but narrow (Fig. 14). These differences bulate and large in Helopeltis theivorn Waterhouse, assume added significance in the light of Sweet's leafy and reduced with delicate transparent lamellae (1967) suggestion that the Stygnocorini may be quite in Carvalhoia arecae Miller & China, and absent in primitive in the Lygaeidae with respect to several Sokenus uvarovi Ballard and Nicosfratus princeps other characters. Distant. Coreidae (Fig. 15) .-The basipulvillus is like that --Neither pulvillus nor parempodium is in the Lygaeidae. The distipulvillus is cup shaped, present in Leptopharsa heidemanni Osborn & Drake its concavity ventral. The lamellae are flattened and and this absence is general in the family (Drake and embedded in a transparent chitinous membrane. The Davis 1960). species studied, all coreines, were Hornoeocerus lacer- Thau1nastocoridae.-The pulvillus in the subfamily tosids Distant, 11. variubilis Dallas, and Cletus bi- Xylastodorinae was described elsewhere (Schaefer f~zctntusWestwood. These species, of course, are 1969). Both pulvillus and parempodium are absent a narrow sampling of the family. in the other subfamily, 'l'haumastocorinae (Drake and (Fig. 16).-The pulvillus here is like Slater 1957). that in the Lygaeidae, but the other margins of the Reduviidae. -The pulvillus is absent. Species lamellae bear many delicate setae in Akbaratus fasci- studied : Reduvius ciliatus Reuter, Acanthaspis pav- atus Distant, and the lamellae are restricted to the ipes StHl, Triatoma rubrofasciatus (De Geer), Ecto- base in Leptocorisa varicornis I;. wtocoris biguttulus StHI, E. cordiger Stil,, Oncoce- .---In general, similar to Lygaeidae. phalus annulipes Stll, Oiecocephalus sp., and Tribel- Species studied : CortLus hyoscyami Id. ( Rhopalinae ocephala indica Walker. 312 ANKALSOF THE ENTOMOLOGICALSOCIETY OF AMEKICA [Vol. 63, no. 1 .---Roth the pulvillus and the parempho- groups included here, should yield useful taxonomic dium are apparently absent (Usinger 1966), but information, particularly at the tribal and subfamilial there are some fusions and reductions basal to the levels. claws in Cimex henzipterus (F.). These modifica- tions may be adaptations to a (partly) ectoparasitic ACKNOWLEDGMENTS life. One of us (S.C.G.) is deeply indebted to Dr. A. -There is no pulvillus, and the parempo- K. Dattagupta, Professor of Zoology, Rirla Institute dia are indistinguishable, except by position, from the of Technology and Science (Pilani), for guidance setae of the tarsus. Species studied : ferus L. and supervision ; and to Dr. S. C. Rastogi for aid in preparing an early draft of this manuscript. We DISCUSSION AND CONCLUSIONS both thank the following for specimens : the Director, The land Heteroptera have been divided into 2 Zoological Survey of India (Calcutta), Dr- H- L. complexes on the basis of several characters (Leston Kudu (Pilani), Dr. K. Kumar (Accra, Ghana), Dr. et al. 1954), not including the pulvillus. It appears M. R. G. K. Nair (VellaYani), Dr. A- K. Sen from our study that 1 of these complexes, the Cim- (Ranchi), Mr. S. Kumar (Saharanpur), Mrs. Sudha icomorpha, is characterized by absence or reduction Goel (Tinsukhia), and Dr. J. A. Slater (University of the pulvillus, which is well developed in the other of Connecticut). We are grateful also for aid ill complex, the , In 1 family, the Re- identification to the Entomologist, Forest Research duviidae, the loss of the pulvillus may be secondary, Institute (Dehradun) , the Director, ~0”onwealth its place being taken by the “fossa spongiosa tibia- Institute of Ento”h3’ (London), and Dr. w- J- rum”; the fossa may be an aid in climbing (Gillette K11ight, British Museum (Natural IIistorY). and Wiggleswortli 1932) or in grasping prey (Miller Finally, we thank StePhatli Shefer for help in 1942). the preparation of the figures, and Dr. J. A. Slater Too few cin1icomorphs have been included in our for reading the final manuscript. The junior author’s study to characterize the cimicomorph superfamilies, work was supported by grant GB-5985 from the Na- and the scant representation of certain important tlonal %ience Foundation- pentatomoniorphan superfamilies ( Coreoidea, Arad- oidea) makes conclusions there only tentative. REFERENCES CITED There are 5 superfamilies in the Pentatomomor- China, w. E., and J. G. Myers. 1929. A reconsidera- pha : the Aradoidea, Lygaeoidea, Pyrrhocoroidea, tion of the classification of the ciniicoid families Coreoidea, and Pentatomoidea. The structure of the (Heteroptera), with the descriptions of two new spider-web bugs. Ann. Mag. Natur. Hist. (10)3: pulvillus, as described here, supports these groupings. 97-125. The reduction of the pulvillus and the sickle-shaped Crampton, G. C. 1923. Preliminary note on the term- distal end of its basipulvillus place the Pyrrhocoroi- inology applied to the parts of an ’s leg. Can. dea near both the Pentatomoidea and the lygaeoid- Entornol. 55: 126-32. Dashman, T. 1953a. Terminology of the pretarsus. coreoid complex. To place the Pyrrhocoroidea be- Ann. Entomol. Soc. Amer. 46: 5M2. tween the last 2 groups is in agreement with South- 1953b. The unguitractor plate as a taxonomic tool in wood (1956), but other work (Stys 1964; work in the Hemiptera. Ibid. 46: 561-78. progress of Schaefer) strongly suggests a wholly in- Drake, C. J., and N. T. Davis. 1960. The morphology, phylogeny, and higher classification of the family dependent origin of the pentatomoidea from a proto- Tingidae, including the description of a new genus lygaeoid stock. To the extent one can generalize and species of the subfamily Vianaidinae (Hemip- from the few specimens studied, the superfam- tera:Heteroptera). Entomol. Amcr. 39 (n. s.) : 1- ilies Lygaeoidea and Coreoidea appear closely re- 100. lated, both groups having the cup-shaped pulvillus DraE,’dc;~;.e~~~c~ 2 t~~&m;i”,j7+2~~2~$l~22 and other pulvillar similarities. The close relation- (Hemiptera : Heteroptera). Ann. Entomol. Soc. ships of these 2 superfamilies, and of them to the Amer. 50: 353-70. Pyrrhocoroidea, has been confirmed (Schacfer 1964). GiIlett, J. DV and v. B- Wiggleswomh. 1932. The c1imbing Organ Of an insect, Rhodnius firozixzts The absence of lamellae in the pulvillus appears to (Hemiptera: Reduviidae). Proc. Roy. Soc. (B) 111 : ally the Aradidae with the Pentatomoidea, if the 1 364-76. aradid species studied is representative. However, Goel, S. C. 1967. The thorax of the adult Coridizrs just as the Aradoidea is distinct from all other penta- ianus Fabr. (Heteroptera : Dinidoridae). Ann. Soc. Entomol. Quebec 12: 69-96. tomomorph ’uperfamilies because Of the absence Of Holway, R. T. 1935. Preliminary note on the structure abdominal trichobothria, so may it be distinct by the of the pretarsus and its possible pllylogenetic sig- absence of a differentiated basipulvillus. Many more nificance. Psyche 42 : 1-24. examples of the Aradoidea should be studied. Knight, H. H. 1941. The plant bugs, or Miridae, of Reuter (lglO) indicated the Pulvillus (“arolia’’) Illinois. Ill. Natur. Hist. Sum. Bull. 22: 1-234. 1968. Taxonomic review: Mirjdae of the Nevada Test and claws are god taxonomic characters, but the site and the western United States. Brigham Young value of these characters was doubted by China and Univ. Sci. Bull. (BioI. Ser.) 9: 1-282. Myers (1929). Although the claws are nearly ident- Leston, D., J. G. Pendergrast, aad T. R. E. Southwood. ical in the families studied here, it is clear that the 1954. Classification of the terrestrial Heteroptera (Geocorisae). Nature (London) 174 : 91-92. pulvillus is indeed a useful character. Study of ad- Levereault, p. 1~35. The insect tarsus. univ. K~~~. ditioiial groups, and of more representatives in the Sci. B:111. 22: 521-5. January 19701 GOEL AND SCHAEFER : PULVILLUS OF KETEKOPTEKA 313 MacGillivray, A. D. 1923. External Insect Anatomy. the classification of the group. Trans. Roy. Entomol. Scarab Co., Urbana, Ill. p. i-viii, 1-388. Soc. London. 108: 163-221. Martin, C. H. 1968. The new family Leptogastridae Stys, P. 1959. The 5th stage nymph of Cerutacordms (the grass flies) compared with the Asilidae (rob- (Cerutocombus) coleoptrutus (Zetterstedt, 1819) and ber flies) (Diptera). J. Kans. Entomol. Soc. 41: notes on the morphology and systematics of Dipso- 70-100. coridae (Heteroptera). Acta Entomol. hlus. Nat. de Meijere, J. C. H. 1901. Uber das letzte Glied der Pragae 33: 377-88. Bcine bei der Arthropoden. 2001. Jahrb. (Anat. Sweet, M. H. 1967. The tribal classification of the Ontog. Tiere). 14: 417-76. (Heteroptera : Lygaeidae) . Ann. Miller, N. C. E. 1942. On the structure of the Icgs in Entomol. Soc. Amer. 60 : 208-26. Reduviidae (Rhynchota). Proc. Roy. Entomol. Soc. 1964. Thaumastellidae-a new family of pentatomoid London. 17: 49-58. Heteroptera. Acta. Soc. Entomol. Cechoslov. 61 : Reuter, 0. M. 1910. Neue Beitrage zur Phylogenie und 236-53. Systematik der Miriden, nebst einleitenden Bemer- 1967. Monograph of Malcinae, with reconsideration kungen iiber die Phylogenie der Heteropteren Fam- of morphology and phylogeny of related groups ilien. Acta. Soc. Sci. Feiin. 37: 1-171. (Heteroptera, ). Acta. Entomol. Mus. Nat. 1912. Bemerkungen iiber mein neues Heteropteren- Pragae. 37 : 351-516. System. Ofv. Fin. Vet-Soc. Forh. (A) 54: 1-62. Tullgren, A. 1918. Zur Morphologie und Systematik Schaefer, C. W. 1%4. The morphology and higher der Ihmipera. I. Ueber das Vorkominen von s.g. classification of the Coreoidea (Hemiptera : Heterop- Trichobothrieii bei Hemiptera-Heteroptera und ihre tera) : Parts I and TI. Ann. Entomol. Soc. Amer. 57: mutmassliche Bedcutung fur das Heteropterensysteni. 1670-84. Entomol. Tidskr. 39: 113-33. 1%9. Morphological and phylogenetic notes on the Usinger, R. L. 1966. Monograph of Cimicidae (IIemip- Thaumastocoridae (Hemiptera-Heteroptera) . J. Kans. tera-IIeteroptera) . Thomas Say Foundation, College Rntoniol. Soc. 42: 251-6. Park, Md. p. i-xi, 1-585. Snodgrass, R. E. 1927. Morphology and mechanism of Usinger, R. L., and R. Matsuda. 1959. Classification of thc insect thorax. Smithson. Misc. Colt. 80: 1-108. the Aradidae ( Hemiptera-Heteroptera). British Mus- 1935. Principles of Insect Morphology. McGraw-Hill, eiim (Natur. Hist.), London. p. i-vii, 1-410. New York. p. i-ix, 1-667. Wygodzinsky, P. W. 1%. A monograph of the Emcs- Southwood, T. R. E. 1956. The structure of the eggs inae (Reduviidae, Hemiptera). Bull. Amer. Mus. of the terrestrial Heteroptera and its relationship in Natur. Hist. 133: 1-614.

Reprillted from the ANNALSOF THE ENTOMOLOGICALSOCIETY OF AMERICA Volume 63, Number 1, pp. 307-313, January 1970