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Ent18 1 007 016 Shcherbakov for Inet.Pmd Russian Entomol. J. 18(1): 716 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2009 Permian insects from the Russky Island, South Primorye Ïåðìñêèå íàñåêîìûå ñ îñòðîâà Ðóññêèé â Þæíîì Ïðèìîðüå Dmitry E. Shcherbakov1, Vladimir N. Makarkin2, Daniil S. Aristov3 & Dmitry V. Vasilenko4 Ä.Å. Ùåðáàêîâ1, Â.Í. Ìàêàðêèí2, Ä.Ñ. Àðèñòîâ3, Ä.Â. Âàñèëåíêî4 1, 3, 4 Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya ul. 123, Moscow 117647, Russia. 1 E-mail: [email protected] 3 E-mail: [email protected] 4 E-mail: [email protected] 2 Institute of Biology and Soil Sciences, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok 690022, Russia. E- mail: [email protected] Ïàëåîíòîëîãè÷åñêèé èíñòèòóò ÐÀÍ, Ïðîôñîþçíàÿ óë. 123, Ìîñêâà 117647, Ðîññèÿ. Áèîëîãî-ïî÷âåííûé èíñòèòóò ÄÂÎ ÐÀÍ, Âëàäèâîñòîê 690022, Ðîññèÿ. KEY WORDS: fossil insects, Megasecoptera, Titanoptera, Grylloblattida, Homoptera, Neuroptera, Coleoptera, Mecoptera, entomofauna, South Primorye, Permian, Kungurian. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: èñêîïàåìûå íàñåêîìûå, Megasecoptera, Titanoptera, Grylloblattida, Homoptera, Neuroptera, Coleoptera, Mecoptera, ýíòîìîôàóíà, Þæíîå Ïðèìîðüå, ïåðìñêèé ïåðèîä, êóíãóðñêèé âåê. ABSTRACT. The first Permian insect assemblage ÐÅÇÞÌÅ. Âïåðâûå ñ Äàëüíåãî Âîñòîêà îïèñàí from the Russian Far East, collected in the lower ïåðìñêèé êîìïëåêñ íàñåêîìûõ èç íèæíåé ÷àñòè Pospelovo Formation (Kungurian, Lower Permian) of ïîñïåëîâñêîé ñâèòû (êóíãóðñêèé ÿðóñ, íèæíÿÿ the Russky Island near Vladivostok, is described. It is ïåðìü) îñòðîâà Ðóññêèé áëèç Âëàäèâîñòîêà.  í¸ì dominated by Grylloblattida (Ideliidae and Lio- äîìèíèðóþò Grylloblattida (Ideliidae è Liomopteridae: mopteridae: Liomopterites svetlanae Aristov, sp.n.) Liomopterites svetlanae Aristov, sp.n.), ïðèñóòñòâóþò and includes Megasecoptera (Scytohymenidae: Ocean- Megasecoptera (Scytohymenidae: Oceanoptera elenae optera elenae Shcherbakov, gen. et sp.n.), Homoptera Shcherbakov, gen. et sp.n.), Homoptera Aucheno- Auchenorrhyncha (Prosbolopseidae Ivaiinae: ?Cicad- rrhyncha (Prosbolopseidae Ivaiinae: ?Cicadopsis sp.), opsis sp.), Plecoptera, Neuroptera (Permithonidae: Plecoptera, Neuroptera (Permithonidae: Okolpania Okolpania eugeniae Makarkin et Shcherbakov, sp.n.), eugeniae Makarkin et Shcherbakov, sp.n.), Coleoptera Coleoptera Archostemata (Permocupedidae: Mari- Archostemata (Permocupedidae: Maricoleus valentinae coleus valentinae Shcherbakov, gen. et sp.n.), Mecop- Shcherbakov, gen. et sp.n.), Mecoptera (Permo- tera (Permochoristidae: Protopanorpa sp., ?Petroman- choristidae: Protopanorpa sp., ?Petromantis sp.), è, tis sp.), and presumed Titanoptera s.l. (Gerarina), but âîçìîæíî, Titanoptera s.l. (Gerarina), íî íå îáíàðó- no Blattida or Orthoptera. Traces of insect-plant inter- æåíû Blattida è Orthoptera. Íàéäåíû ñëåäû âçàèìî- actions include cordaite seeds with punctures made by äåéñòâèÿ íàñåêîìûõ è ðàñòåíèé ïðîêîëîòûå ïà- Palaeodictyopteroidea and clutches of relatively large, ëåîäèêòèîïòåðîèäàìè ñåìåíà êîðäàèòîâ è äîâîëü- lense-shaped eggs attached to leaves and stems (possi- íî êðóïíûå ëèíçîâèäíûå ÿéöà, îòëîæåííûå íà ëèñ- bly produced by megasecopterans; the earliest record òüÿ è ñòåáëè (ñàìûå äðåâíèå ýêçîôèòíûå êëàäêè, of exophytic oviposition). In the regular occurrence of âîçìîæíî, ïðèíàäëåæàùèå ìåãàñåêîïòåðàì). Ïî palaeodictyopteroids and their feeding traces the ðåãóëÿðíûì íàõîäêàì ïàëåîäèêòèîïòåðîèäîâ è ñëå- Russky Island assemblage is similar to the Vorkuta äîâ èõ ïèòàíèÿ êîìïëåêñ îñòðîâà Ðóññêèé ñõîäåí ñ Group (Pechora Basin) entomofauna, while in the ýíòîìîôàóíîé âîðêóòñêîé ñåðèè Ïå÷îðñêîãî áàñ- absence of cockroaches it is closer to the Siberian ñåéíà, à ïî îòñóòñòâèþ òàðàêàíîâ ñ êîìïëåêñàìè Middle Permian assemblages. The systematic position èç ñðåäíåé ïåðìè Ñèáèðè. Îáñóæäàåòñÿ ñèñòåìàòè- of Nemuropsis Martynov, 1925 is discussed. The new ÷åñêîå ïîëîæåíèå Nemuropsis Martynov, 1925. Íî- insect species are named after the paleobotanists from âûå âèäû íàñåêîìûõ íàçâàíû â ÷åñòü Å.Á. Âîëûíåö, Vladivostok, Elena B. Volynets, Valentina I. Burago, Â.È. Áóðàãî, Ñ.À. Øîðîõîâîé è Å.Â. Áóãäàåâîé Svetlana A. Shorokhova and Eugenia V. Bugdaeva, ïàëåîáîòàíèêîâ èç Âëàäèâîñòîêà, îêàçàâøèõ íàì who greatly facilitated our fieldwork. íåîöåíèìóþ ïîìîùü â îðãàíèçàöèè ïîëåâûõ ðàáîò. 8 Dmitry E. Shcherbakov, Vladimir N. Makarkin, Daniil S. Aristov & Dmitry V. Vasilenko The first Permian insect from Primorye was collected pteridosperms and Cathaysian elements [Tashchi & in 1923 near Vladivostok (the valley of Obyasnenie Burago, 1974; Zimina, 1977, 1997; Zakharov et al., River, southern slope of Komarov Mt. now within 2009]. This plant assemblage is similar to those from the city boundaries) by Pavel V. Vittenburg, an eminent the Rudnik Subformation (Lek-Vorkuta Formation, Russian geologist, from the Taonurus horizon. These Vorkuta Group, Kungurian or Ufimian) of the Pechora beds are now assigned to the upper subformation of the Basin and from the Usa and Starokuznetsk Formations Pospelovo Formation (V.I. Burago, pers.comm.), dated (Kuznetsk Subgroup) of the Kuznetsk Basin [Kotlyar et Late Kungurian according to the International Strati- al., 2006]. Abundant mosses (Intia sp. and Uskatia sp.: graphic Scale [Zakharov et al., 2009] (Ufimian of the [Burago in Meyen, 1982]) and liverworts are recorded. East European Stratigraphic Scale). The fossil was de- The insect remains are usually wing fragments or scribed by Andrey V. Martynov [1925] as Nemuropsis isolated body parts (nymphal wing pads, body seg- tenuis Martynov, 1925, constituting a new family Nem- ments, legs). The insect assemblage is quite diverse: 8 uropsidae, which he placed, along with three other, orders per 42 identifiable specimens. It is dominated by Carboniferous families, in a new suborder (or even Grylloblattida (24 specimens, or more than half of all order) Paraplecoptera within Protorthoptera, on account the identified insects). The palaeodictyopteroid Megas- of several similarities to Plecoptera. ecoptera are represented with two wings, about 40 mm The classification of Paleozoic orthopteroids is still long, and probably also with several nymphal frag- disputable; most protorthopterans are now assigned to ments. Homoptera (suborder Auchenorrhyncha) and the orders Grylloblattida and Eoblattida. Nemuropsis is Mecoptera are each known from two small wings. The a poorly preserved, large insect (forewing length ca. 60 remaining orders are represented each with a single mm) possessing a nearly quadrate pronotum without specimen: one nymphal exuvium of Plecoptera, an in- paranota, and wings completely overlapping in repose. complete forewing of Neuroptera, an elytron of Co- Although the two latter features are characteristic of leoptera (suborder Archostemata), and a large femur stoneflies, the largest Permian Plecoptera are only half with two spine rows along the ventral surface, possibly as large as Nemuropsis. The grylloblattids lacking prono- belonging to a primitive Titanoptera s.l. (suborder Ger- tal paranota (part of suborder Protoperlina) are also arina). More than thirty insect fragments were impossi- smaller (forewing length less than 25 mm), and their ble to identify to order. wings only partly overlap in repose. So far as known, The insect groups recorded in this assemblage dis- large eoblattids did not survive to the mid-Permian, but play a variety of diets: specialized sucking herbivory on such a possibility cannot be excluded. Before re-exam- vascular plants (Megasecoptera and Auchenorrhyncha), ination of the holotype (the location of which is un- xylomycetophagy (i.e., feeding on rotting wood: larvae known) it is not possible to place Nemuropsis in one of of Archostemata), saprophagy (most probable for Gryl- these orders. loblattida, larvae of Mecoptera, and nymphs of Plecop- In the 1970s, four insect specimens were collected tera), and zoophagy (Gerarina, larvae of Neuroptera). by V.I. Burago (Primorye Prospecting and Mapping The only doubtless aquatic insect is the nymph of Ple- Geological Expedition, Vladivostok) from three differ- coptera. ent Permian outcrops in Primorye (two of them from the The most significant traces of insect feeding are lower Pospelovo Formation of Russky Island, outcrop cordaite seeds with punctures made by beaks of Megas- 429), and one additional specimen by a field party of the ecoptera (represented in this assemblage by isolated Arthropoda Lab, Paleontological Institute (PIN), but wings) or some other palaeodictyopteroids (Figs 12), these finds remain undescribed. In 2008, our field party indicating that these insects were more abundant in the of the Arthropoda Lab, PIN, joined by Elena B. Voly- lower Pospelovo Formation than one may assume based nets and Valentina I. Burago, visited this outcrop on the only on their body fossils. Such feeding damage of Russky Island for several days and collected some cordaite seeds was first described from the Late Car- seventy insects. The first Permian insect assemblage boniferous of Tunguska Basin and attributed to Palaeo- recorded from the Russian Far East is of high interest. dictyoptera by Sharov [1973]. Recently, it was reported All the specimens mentioned below are deposited at also from the Early Permian Vorkuta Group (Kozhim- PIN. They were collected from black coaly shales (con- rudnik Formation, correlated to the Lek-Vorkuta For- taining abundant plant fossils, Bivalvia and few Con- mation, Kungurian or Ufimian) of the Pechora Basin chostraca) of the lower sub-formation of Pospelovo [Shcherbakov, 2008]. Relatively large eggs, attached to Formation,
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