Paleontological Journal, Vol. 38, Suppl. 2, 2004, pp. S173–S184. Original Russian Text Copyright © 2004 by Novokshonov, Zhuzhgova. English Translation Copyright © 2004 by MAIK “Nauka /Interperiodica” (Russia). Discussion of the System and Phylogeny of the Order Palaeomanteida (= Miomoptera) with Description of New Representatives of the Genus Permosialis Mart. from the Late Permian of Kirov Region and Triassic of Kyrgyzstan V. G. Novokshonov† and L. V. Zhuzhgova Perm State University, ul. Genkelya 1, Perm, 614005 Russia Received June 19, 2002 Abstract—Taxonomic and phylogenetic relationships within the order Palaeomanteida are discussed. Morpho- logical characteristics of little-known miomopteran taxa are supplemented. An idea of sexual dimorphism in the species of Permosialis Mart. is advanced. Permosialis ualentovae sp. nov. from the Late Permian of the Kirov region and Permosialis triassica sp. nov. from the Triassic of Kyrgyzstan are described. INTRODUCTION any other holometabolous insect order, they are consid- ered to be a separate superorder. The adult morphology The order Palaeomanteida (= Miomoptera), which of miomopterans suggests that they lived in open rather occurred from the Middle Carboniferous to the Early– than cryptozoic habitats and, probably, fed on gymno- Middle Jurassic, is considered to be a common ancestor sperm strobili (Rasnitsyn, 1980a). For a long time, of all holometabolous insects (Rasnitsyn, 1980a). This there was no direct evidence of such trophic relation- order was transferred into Scarabaeiformes (= Holom- ships; they have been confirmed, however, by pollen etabola) from Gryllones (= Polyneoptera) by Rohden- found in the gut of miomopterans belonging to Palaeo- dorf (1977), but without any comment (Rasnitsyn, mantiscidae and Palaeomanteidae from the Lower Per- 1980a). Rasnitsyn (1980a) analyzed the problem in mian of Tshekarda (Rasnitsyn, pers. comm.; the detail and noted that the nymphs described as being authors’ personal observations). Taking into account miomopteran (Sharov, 1957) most probably belong to the morphology of the ovipositor in Palaeomanteida, it Gryllones and that the segmented cerci of Palaeo- appears that their larvae fed on pollen in more or less manteida do not preclude assigning them to holometab- ripe gymnosperm cones, moving between the scales olans, because three-segmented cerci are retained in from one microsporangium to another. Hymenopterans living scorpionflies and, therefore, should also be form a direct continuation of this lineage; another present in primitive Scarabaeiformes. In addition, Ras- group of miomopteran descendants consists of beetles, nitsyn (1980a) listed those features of Palaeomanteida neuropteroids, and mecopteroids (Rasnitsyn, 1980a). that are characteristic of Scarabaeones (Palaeoptera, The beetles represent a separate lineage, which is Para- and Oligoneoptera), but the full set of said fea- related to almost equal extents to both mecopteroids tures never occurs in Gryllones. These are wings that and neuropteroids; the larvae of the first beetles were are folded rooflike over the abdomen in repose (except apparently xylomycetophagous; the neuropteran larvae for Permonka Riek), a hindwing anal area not tucked were predatory, whereas the first mecopteroids proba- under in repose, cryptosterny, the third valvulae of the bly became detritivorous with a tendency toward being ovipositor forming its sheath, and the male gonocoxae necrozoophagous (Ponomarenko, 1983). None of the and gonostyles forming a clasping organ. One more known Carboniferous groups of insects has been iden- character listed is generally specific for holometab- tified as ancestral to Palaeomanteida. olous insects: additional medial articulation of pterothoracic coxae. Therefore, in adult morphology, MATERIAL miomopterans turned out to be, on the one hand, suffi- ciently advanced to be included in Scarabaeiformes Collections of the Paleontological Institute of Russian and, on the other hand, so primitive and ancient that Academy of Sciences (PIN) and Perm State University they might be ancestral to the remaining Scarabaei- (PGU) were studied. All specimens are stored at PIN. formes, all the more so as no other group is known to meet the required conditions. Since no smooth transi- SYSTEM OF THE ORDER tion has been uncovered between Palaeomanteida and Data on the taxonomy of miomopterans have been †Deceased. summarized by many authors (Martynova, 1962; Riek, S173 S174 NOVOKSHONOV, ZHUZHGOVA 1976; Kukalová, 1963; Rasnitsyn, 1980a, 1980b; Car- characteristic of most miomopterans (Palaeomanteidae penter, 1992; Storozhenko, Novokshonov, 1999; and Palaeomantiscidae). If so, miomopterans of the Novokshonov, 2000; Rasnitsyn, 2002). Currently, family Permosialidae should, perhaps, be excluded Palaeomanteida are thought to comprise the family from the supposed ancestors of hymenopterans and Palaeomanteidae Handl., which includes Palaeomantis may only be considered to be ancestral to archaic Handl. (Late Carboniferous–Permian of Eurasia), mecopteroids and neuropteroids. On the basis of its Delopterum Sell. (Late Carboniferous–Permian of Eur- morphology, the genus Palaeomantis (see below), asia and North America) and Epimastax Mart. (Late which is apomorphous only in having four-segmented Permian of eastern and northeastern Europe); Permo- tarsi, could be nearest to the origin of hymenopterans. sialidae Mart., which includes Permosialis Mart. (Per- In the Permian, Palaeomantis and Delopterum mian–Triassic of Eurasia and Australia) and Permonka became especially diverse and widespread. Their mor- Riek (Late Permian–Early and Middle Jurassic of Eur- phology is quite well known today, mainly due to asia and South Africa): and Palaeomantiscidae A. Rasn., numerous finds in the Lower Permian of the Urals which includes Sellardsiopsis G. Zal. (Early Permian of (Novokshonov, 2000). The representatives of Palaeo- the Urals) and Palaeomantina A. Rasn. (Early Permian mantis (body structure being known for P. aestiva of the Urals). The reasons for the taxonomic changes Novokshonov, 2000) are medium-sized insects and the status of those genera that were previously (Figs. 1b–1f). The antennae are setaceous and about as known but are not listed here have been discussed in long as the forewing; the eyes are ovoid and weakly detail by Rasnitsyn (2002). convex; the ocelli form a single group; the maxillary Miomopterans from the Middle Carboniferous of palps are long, and the labial palps are much shorter. the Mazon Creek locality have not yet been described The pronotum is large, rounded quadrangular, but (see below). Late Carboniferous miomopterans have slightly shorter than it is wide; the meso- and metano- been discovered in several localities within France, tum are subdivided as usual; the pleural region is of Germany, and the United States, where they are repre- generalized structure. The tarsi are four-segmented. sented by two genera, Delopterum and Palaeomantis The male abdomen bears long, multisegmented cerci, (Rasnitsyn, 2002). Thus, the earliest known mio- forming a forceps-shaped figure (probably, in P. aes- mopterans belong to the family Palaeomanteidae. This tiva, cerci aided in fixing the female abdomen during suggests that future studies of their morphology will copulation); the gonocoxae are large, apparently firmly apparently be most useful in resolving the problems of fused to each other by their upper and lower processes; both the phylogeny of the order and the origin of the gonostyles are small and curved; nearly all speci- Holometabola. The remaining miomopteran families mens show a central baculiform sclerotized process, the appeared only in the Early Permian, virtually synchro- tip of which could extend far beyond the gonostyle api- nously with the first scorpionflies, caddisflies, neu- ces (presumably, this is an evaginable penis); a similar ropterans, and beetles. Hymenopterans (order Vespida) structure of the male genitalia is characteristic of both are known only from the Triassic onward. some mecopteroids (caddisflies) and hymenopterans The family Palaeomanteidae comprises most of (Rasnitsyn, 1969, p. 138, text-figs. 244–250). The the representatives of the order and is characterized by female abdomen bears a powerful, straight, and wide the following main wing features: wing folded rooflike; ovipositor reaching up to the apices of the folded forewings with a slightly thickened membrane, rela- wings; no armature is traceable on the outer valves, but tively narrow costal area, and with several anterior SC often a narrow pair of the inner valves reaching the ovi- branches (up to seven in Epimastax) or, rarely, no SC positor tip is clearly visible due to their distinct trans- branches (Delopterum); in forewings, M and CuA verse striations (very similar structures are present in anastomose subbasally for a greater or lesser but still the ovipositors of some extant hymenopterans: Ras- relatively long distance, i.e., the base of M5 is absent as nitsyn, 1969, p. 132, text-fig. 236). In the forewings, SC a separate vein. This last feature is supposedly a sec- terminates slightly distal to the wing midlength, with ondary development in Palaeomanteida, the original four or five anterior branches; the costal area is about as state having been a retention of the free M5 base that wide as the subcostal one; R has a more or less distinct was fused to CuA to form a Y-shaped figure, as in mio- scooplike bend distally; RS has normally three or four mopterans Permosialidae and in all archaic mecop- terminations