New Saldoid Bug of the Family Archegocimicidae (Hemiptera: Heteroptera: Leptopodomorpha) from the Middle Jurassic of Eastern Siberia

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1 ISSN , Paleontological Journal, 2013, Vol. 47, No. 2, pp Pleiades Publishing, Ltd., Original Russian Text O.V. Ryzhkova, 2013, published in Paleontologicheskii Zhurnal, 2013, No. 2, pp New Saldoid Bug of the Family Archegocimicidae (Hemiptera: Heteroptera: Leptopodomorpha) from the Middle Jurassic of Eastern Siberia O. V. Ryzhkova Borissiak Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya ul. 123, Moscow, Russia Received June 28, 2012 Abstract A new genus and species of saldoid bugs, Kubecora ignamica gen. et sp. nov. (Heteroptera, Archegocimicidae), is described from the Middle Jurassic (Itat Formation) locality Kubekovo, East Siberia. Keywords: Jurassic, Transbaikalia, bugs, Archegocimicidae DOI: /S INTRODUCTION The extinct Mesozoic bug family Archegocimicidae occupies a basal position in the infraorder Leptopodomorpha and is probably ancestral to the recent family Saldidae (Popov, 1985, 1988, 1989; Shcherbakov and Popov, 2002). The morphological similarity of these two families has also been repeatedly mentioned in the literature (Handlirsch, 1925; Popov, 1985, 1988, 1989; Popov, Dolling, and Whalley, 1994). The family Archegocimicidae was originally described by Handlirsch (1906) and included the genus Archegocimex Handlirsch, 1906, which contained a single species, A. geinitzi Handlirsch, 1906 from the Lower Jurassic deposits of Germany (Dobbertin municipality, Mecklenburg-Vorpommern). The families Eonabidae and Diatillidae, described by Handlirsch in 1925, were subsequently synonymized under Archegocimicidae (Popov and Wootton, 1977). Such genera from the same deposits as Archegocoris Handlirsch, 1906, Progonocoris Handlirsch, 1939, Anosmus Handlirsch, 1939, Eonabis Handlirsch, 1925, Diatillus Handlirsch, 1925, Somatocoris Bode, 1953, Eurynotis Bode, 1953, Entomecoris Bode, 1953, Corynecoris Bode, 1953, Macropterocoris Bode, 1953, and Ensphingocoris Bode, 1953 were also included in the family; some of them were previously placed in their own monotypic families (Handlirsch, 1939; Bode, 1953). However, detailed study of the type materials of these genera, as well as of the genera Ophthalmocoris Bode, 1953, Deraiocoris Bode, 1953, and Euraspidium Bode, 1953 revealed that they were all synonymous to the genus Archegocimex (Popov and Wootton, 1977; Popov, 1985). The systematic position of the family Archegocimicidae long remained unclear. It was either considered as family of unclear systematic placement (Handlirsch, 1939; Bode, 1953; Becker-Migdisova, 1962; Popov and Wootton, 1977), or placed close to Pentatomidae (Handlirsch, 1906), or Pentatomomorpha, Cimicomorpha, and Dipsocoromorpha (Popov and Wootton, 1977), or tentatively included in Pentatomomorpha (Popov, 1980, 1981), or in Cimicomorpha (Whalley, 1985). However, it was noted already by Handlirsch (1925) that the families Archegocimicidae and Saldidae were similar in the structure of the head. Popov (1973) described the new genus Saldonia from the Early Jurassic of Transbaikalia in the family Saldidae, erecting for this genus the monotypic subfamily Saldoniinae. Subsequently he transferred the genus to Archegocimicidae and included this family in the superfamily Leptopodoidea, placing it close to Saldidae, and simultaneously abolishing the subfamily Saldoniinae (Popov, 1973, 1985, 1989). The Middle Jurassic locality of Kubekovo, Krasnoyarsk Region, is formed by deposits of a very large lake (Zherikhin, 1985). The insect assemblage of Kubekovo is especially close to the Toarcian entomofaunas of Germany and to some Early to Middle Jurassic entomofaunas of Siberia (Zherikhin, 1985). Examination of materials collected in this locality revealed an impression of the elytron of a bug of the family Archegocimicidae with a small number of apical cells (five), which is atypical of this family. This number of apical cells is normally characteristic of the Early Cretaceous saldoid family Enicocoridae, which is currently included by some specialists in the family Saldidae as a subfamily (Shcherbakov and Popov, 2002, Zhang et al., 2005). The author has placed the bug from Kubekovo in a new genus, rather similar in the shape of the radial cell and position of radial branches to members of the genus Archegocimex, especially A. geinitzi (Fig. 1). Nevertheless, the new genus is 180

NEW SALDOID BUG OF THE FAMILY ARCHEGOCIMICIDAE 181 erected here, since the number of apical cells in members of the genus Archegocimex is no less than six (Popov and Wootton, 1977).

2 NEW SALDOID BUG OF THE FAMILY ARCHEGOCIMICIDAE 181 erected here, since the number of apical cells in members of the genus Archegocimex is no less than six (Popov and Wootton, 1977). It has to be noted that there are rather strong differences in venation between species of the genus Archegocimex, and this genus is probably composite. Unfortunately, at present the author can study only the figures of these bugs drawn by R. Wootton (Popov and Wootton, 1977), and photographs of the holotypes. The likely division of Archegocimex into several genera will be possible only after direct re-examination of the type material. The material studied is stored in the Borissiak Paleontological Institute, Russian Academy of Sciences (PIN). SYSTEMATIC PALEONTOLOGY I n f r a o r d e r Leptopodomorpha Family Archegocimicidae Handlirsch, 1906 Genus Kubecora Ryzhkova, gen. nov. E t y m o l o g y. From Kubekovo locality and latinization of the Greek kore (daughter). Type species. Kubecora ignamica, sp. nov. D i a g n o s i s. Medium-sized bug. Costal area relatively wide (length to width ratio approximately 7 : 1), strongly chitinized, occupying 2/3 of wing length. Base of anterior carina 1.25 times as close to base of posterior carina as to base of hindwing. Arculus long; R and M diverging at angle smaller than 45 ; M without bend at point of divergence of arculus, making radial cell triangular. Median cell pentagonal. Radial cell subequal in length to median cell. Most distal one of additional crossveins joining one of branches of most distal fork. Number of apical cells 5. R with two branches parallel at base, but weakly diverging distally. R1 reaching anterior margin of wing. Species composition. Type species. C o m p a r i s o n. The new genus is distinguished from all other genera of the family by the smaller number of apical cells. The triangular shape of the radial cell makes it similar to the genera Archegocimex and Shartegocimex; the new genus differs from the latter in the longer basal portion of the radial cell and the divergence of R1 and R2. It differs from the genus Britannicola in the longer basal area of the radial cell. It differs from the genera Britannicola, Saldonia, and Sondalia in the triangular shape of the radial cell; it also differs from the latter genus in the longer costal area. It differs from the genera Saldonia and Shartegocorpus in the closely set bases of the carinae. R e m a r k s. The absence of a distinct membrane, long costal fracture reaching the point of divergence of R and M, presence of two well-developed carinae in the precostal area, wide costal area, long M5, triangular radial cell, and pentagonal median cells are conclusive evidence that the new genus belongs to the family Archegocimicidae. Nevertheless, the presence of five apical cells and the divergence of R branches (although Fig. 1. Archegocimex geinitzi Hand., 1906, Lower Jurassic of Germany (from Popov and Wootton, 1977). Scale bar in Figs. 1, 2, 3, 1 mm. much more strongly pronounced) are more typical of the family Enicocoridae. Moreover, the additional crossvein between the branches of M, although it is more typical of Archegocimicidae, sometimes occurs also in enicorids (Ryzhkova, 2012), providing yet another evidence of their close relatedness. Kubecora ignamica Ryzhkova, sp. nov. Plate 11, fig. 1 E t y m o l o g y. From the specific epithet of Saldonia ignota and the Latin amica (female friend). H o l o t y p e. PIN, no. 1255/447, adult, sex unknown; positive and negative impressions of elytron without clavus; Kubekovo village; Middle Jurassic, Itat Formation. D e s c r i p t i o n (Fig. 2). M has two branches, slightly diverging towards the wing apex. M1+2 and M3 divide distal to the crossvein m-cu. The crossvein r-m is present. The segment of M1+2 between the bifurcation of the branches of M and the divergence of r-m is S-shaped. The ratio of the length of the radial cell and the length of internal apical cell is 0.43; the ratio of the length of the external apical cell and the internal apical cell is 0.6. Two additional crossveins are present: ms1, diverging from the S-shaped segment of M1+2 and joining the radial stem proximal to the divergence of R1, and ms2, located between M1+2 and M3 and joining M1+2 at 2/3 of its length from r-

3 182 RYZHKOVA Plate 11 1 mm 1a 1 mm 1b Explanation of Plate 11 Fig. 1. Kubecora ignamica gen. et sp. nov., holotype PIN, no. 1255/447: (1a) positive impression; (1b) negative impression. PALEONTOLOGICAL JOURNAL Vol. 47 No

4 NEW SALDOID BUG OF THE FAMILY ARCHEGOCIMICIDAE 183 ca cp R R+M Cu R1 R2 eac ms1 rc M M1 M2 ms2 CuA1 CuA2 iac Fig. 2. Kubecora ignamica gen. et sp. nov., holotype PIN, no. 1255/447. Notation: (ca) anterior carina; (cp) posterior carina; (rc) radial cell; (eac) external apical cell; (iac) internal apical cell. m. The lightly colored spot is pronounced in the distal part of the basal cell and under this cell. Measurements, mm. Length of elytron, 4.5; width of preserved part of elytron, R e m a r k s. The relative width of the costal area and the venation pattern of the membrane (presence and location of two additional crossveins) make Kubecora ignamica gen. et sp. nov. especially similar to Saldonia ignota Popov, 1988 (Fig. 3), but the new species clearly differs from that species in the triangular shape of the radial cell and in the number of apical cells. Material. Holotype. ACKNOWLEDGMENTS The author is grateful to Yu.A. Popov, A.P. Rasnitsyn, and D.E. Shcherbakov (PIN) for useful comments and help in manuscript preparation. Fig. 3. Saldonia ignota Yu. Popov, 1988, holotype PIN, no. 4098/3; Upper Jurassic to Lower Cretaceous of the Chita Region. REFERENCES Becker-Migdisova, E.E., Order Heteroptera: Heteropterans, or True Bugs, in Osnovy paleontologii. Tom 9. Chlenistonogie. Trakheinye i khelitserovye (Fundamentals of Paleontology: Vol. 9. Arthropoda: Tracheata and Chelicerata), Rohdendorf, B.B., Ed., Moscow: Akad. Nauk SSSR, 1962, pp Bode, A., Die Insektenfauna des ostniedersächsischen Oberen Lias, Palaeontographica, 1953, vol. 103, sect. A, nos. 1/4, 1953, pp

5 184 RYZHKOVA Handlirsch, A., Die fossilen Insekten und die Phylogenie der rezenten Formen: ein Handbuch für Paläontologen und Zoologen, Leipzig: W. Engelman, Handlirsch, A., Palaeontologie, in Handbuch der Entomologie, Schröder, C.W.M, Ed., Leipzig: G. Fischer, 1925, vol. 3, pp Handlirsch, A., Neue Untersuchungen über die fossilen Insecten, Ann. Naturhist. Mus. Wien, 1939, vol. 49, pp Popov, Yu.A., The First Discovery of the Hemiptera Family Saldidae (Heteroptera) in the Mesozoic of Siberia, Dokl. Akad. Nauk SSSR, Nov. Ser., 1973, vol. 209, no. 3, pp Popov, Yu.A. and Wootton, R.J., The Upper Liassic Heteroptera of Mecklenburg and Saxony, Syst. Entomol., 1977, vol. 2, pp Popov, Yu.A., Order Cimicida Laicharting, 1781, in Istoricheskoe razvitie klassa nasekomykh: Tr. Paleontol. Inst. Akad. Nauk SSSR, T. 175 (Historical Development of the Class Insecta: Proc. Paleontol. Inst. Acad. Sci. USSR, vol. 175), Moscow: Nauka, 1980, pp Popov, Yu.A., Historical Development and Some Questions on the General Classification of Hemiptera, Rostria, 1981, vol. 33 (Suppl.), pp Popov, Yu.A., Jurassic Bugs and Coleorrhyncha of Southern Siberia and Western Mongolia, in Yurskie nasekomye Sibiri i Mongolii: Tr. Paleontol. Inst. Akad. Nauk SSSR. T. 211 (Jurassic Insects of Siberia and Mongolia: Proc. Paleontol. Inst. Acad. Sci. USSR, Vol. 211), Rasnitsyn, A.P., Ed., Moscow: Nauka, 1985, pp Popov, Yu.A., New Mesozoic Coleorrhyncha and Heteroptera from Eastern Transbaikalia, Paleontol. Zh., 1988, no. 4, pp Popov, Yu.A., Some Aspects of the Systematics of Leptopodoidea, Acta Biol. Siles., 1989, vol. 13, no. 30, pp Popov, Yu.A., Dolling, W.R., and Whalley, P.E.S., British Upper Triassic and Lower Jurassic Heteroptera and Coleorrhyncha (Insecta: Hemiptera), Genus, 1994, vol. 5, no. 4, pp Ryzhkova, O.V., New Saldoid Bugs of the Family Enicocoridae (Hemiptera: Heteroptera: Leptopodomorpha) from the Lower Cretaceous of Mongolia, Paleontol. J., 2012, vol. 46, no. 5, pp Shcherbakov, D.E. and Popov, Y.A., Superorder Cimicida Laicharting, Order Hemiptera Linné, The Bugs, Cicadas, Plantlice, Scale Insects, etc., in History of Insects, Rasnitsyn, A.P. and Quicke, D.L.J., Eds., Dordrecht: Kluwer Acad. Publ., Whalley, P.E.S., The Systematics and Palaeogeography of the Lower Jurassic Insects of Dorset, England, Bull. Brit. Mus. (Nat. Hist.). Geol. Ser., 1985, vol. 39, no. 5, pp Zhang, J., Golub, V.B., Popov, Y.A., and Shcherbakov, D.E., Ignotingidae fam. nov. (Insecta: Heteroptera: Tingoidea), the Earliest Lace Bugs from the Upper Mesozoic of Eastern China, Cret. Res., 2005, vol. 26, no. 5, pp Zherikhin, V.V., Insects, in Yurskie kontinental nye biotsenozy Yuzhnoi Sibiri i sopredel nykh territorii: Tr. Paleontol. Inst. Akad. Nauk SSSR. T. 213 (Jurassic Continental Biocenoses of Southern Siberia and Adjacent Areas: Proc. Paleontol. Inst. Acad. Sci. USSR, Vol. 213), Moscow: Nauka, 1985, pp

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