SYSTEMATICS Ga?loisiana olgae sp. nov. (Grylloblattodea: Grylloblattidae) and the Paleobiology of a Relict Order of Insects PETER VRSANSKY,^' ^ SERGEI Y. STOROZHENKO,' CONRAD C. LARANDEIRA,^' ^ AND PETRA IHRINGOVA'' Ann. Entomol. Soc. Am. 94(2): 179-184 (2001) ABSTRACT An extant species of the relict insect order Grylloblattodea is described from the Ussuri River Basin of southeastern Russia. This species, Galloisiana olgae, is a member of the family Grylloblattidae that probably originated as a lineage during the mid-Genozoic and experienced subsequent range constriction associated with Pleistocene glaciation. The Grylloblattodea have a richer fossil history in warm-temperate habitats during the Late Paleozoic than the four confamilial genera of today would suggest. These modern taxa represent a speciahzed Genozoic lineage that adapted to cool-temperate habitats in northwestern North America and northeastern Asia, and parallel other similar distributions of seed plants and insects. KEY WORDS Galloisiana olgae, Grylloblattodea, rock crawlers, new species, paleobiology, Ussuri River Basin THE GRYLLOBLATTIDS OR "rock-crawlers" are the least diverse of modern insect orders, consisting of 26 spe- cies, including a new species described below. All of the known extant species, which belong to the family Grylloblattidae, can be considered "living fossils" with presently relict distributions. The single extant family can be contrasted with >44 families described from the fossil record, which extend to the Late Carbonif- erous (Storozhenko 1992, 1997). Grylloblattodeans occupied diverse habitats during the Paleozoic and Mesozoic but currently are restricted to several spe- cialized populations in northern, cool-temperate re- gions. The newly described species bears some of the most primitive features currently within the order and survives as a relict lineage from the mid-Cenozoic Turgai Forest of northern Eurasia. Modern grylloblattids are wingless insects that live on and in soil, in caves, and beneath stones and in crevices of mountainous regions. They are principally carrion feeders on other insects though they will con- sume plant material (Pritchard and Scholefield 1978). They are extremely cryophilic, having a temperature optimum as low as 0?C (Henson 1957, Edwards 1982) and are negatively phototrophic and nocturnal when active (Rentz 1982). The order contains four extant genera, all occurring poleward of =35? latitude in cool-temperate areas of the Northern Hemisphere Department of Zoology, Comenius University, MlynskaDolinaBl, 842 15, Bratislava, Slovakia. Palaeontological Institute, Profsoyuznaya 123, Moscow, Russia 117647. ?' Institute of Biology and Pedology, Vladivostok 22, Russia 690022. "^ Department of Paleobiology, National Museum of Natural His- tory, Smithsonian Institution, Washington, DC 20560-0121. ^ Department of Entomology, University of Maiyland, College Park, MD 20742. " AMBA Projects, Ticha 4, 811 02, Bratislava Slovakia. (Fig. 1). Although living species are wingless, fossil representatives are fully winged and capable of flight (Rasnitsyn 1980). The correlated loss of wings, a sub- terranean life, and adaptation to cold temperature is a departure from the major evolutionary trend in Gryllo- blattodea that was established during the Mesozoic in warmer environments. Mesozoic species ancestral to the modern taxa were probably similar to Grylloblat- tina, the representative harboring the greatest number of primitive features, and a taxon that inhabits a moun- tainous region of the Khakaskaya Oblast in south- central Siberia (Fig. 1). Galloisiana also lives in old- growth, cool-temperate forest, in the Sikhote-Alin Mountains in the Ussuri River Basin, which also sup- ports some plant species that are relicts of the mid- Cenozoic Turgai Forest. It was this locality in which the new species Galloisiana olgae sp. nov. was found, representing the northernmost occurrence of the or- der in the Far East of Russia. The Ussuri River Basin encompasses the greatest species-level diversity of this order and is the only region in which more than one genus of this order has been found. Galloisiana olgae Vrsansky & Storozhenko, new species (Figs. 2 a-f and 3 b-e) Material Examined. HOLOTYPE: female: Russia, Primorskii Region, Olga Mts., the basin of Vasilkovka River, 19.VIII.1997; collected by Vrsansky and Ihrin- gova; repository in Zoological Institute, St. Petersburg. Paratypes. Two adult females, 16-21.VIII.1997; im- mature males and females from the same locality not collected. Adult (Female). Medium-sized for genus; dorsal body covered with numerous short hairs. Head broader than pronotum. Eyes 2.2-2.4 times longer than 0013-8746/01/0179-0184$02.00/0 ? 2001 Entomological Society of America 180 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 94, no. 2 Fig. 1. Contemporary distribution of Grylloblattida. 1 Grylloblatta (11 species), 2 Galloisiana (12 species), 3 GryUoblattina (one species), 4 Grylloblattella (2 species). wide, well pigmented. Parietal suture reaching eyes; frontal and clypeal sutures well differentiated. Anten- nae with 37-38 segments; third segment =3 times longer than second. Pronotum as long as wide, with broadly rounded posterior margin. Cervical sclerite with 7-8 setae (macrotrichiae) on external margin. Basisternum oviform. Legs long; femur ratios (length divided by width) as follows: fore femur 2.8 times, hind femur 4.3 times; tarsal pulvilli relatively short. Hind margin of I tergite with 4 (2+2) setae, II-V tergites with 8 (4+4), VI with 10 (5+5), VII-IX with 6 (3+3); anterolateral part of II-VII tergites with 1+1 setae. Sternites with two regular rows of setae. Cerci 9-seg- mented (including weakly separated first and second segments). Ovipositor reaching fifth segment of cerci. Fig. 2. (a) Galloisiana olgae sp. nov., female, (b) head, (c) fore leg, (d) cervical sclerites and basisternum, (e) 4-6 sternites, (f) apex of abdomen, lateral view. Body brown, legs and ventral surface of abdomen light brown, ovipositor brown. Body Dimensions (Holotype). Length: body 17.2 mm; head 4.5 mm; pronotum 3.6 mm; fore femur 3.7 mm; hind femur 4.7 mm; cerci 9.3 mm; ovipositor 3.7 mm. Width: head 4.0 mm; pronotum 3.6 mm. Adult (male) : Unknown. Immature male typically similar to female, including well pigmented eyes and number and placement of setae on body, but easily distinguished by structure of abdominal apex. Comparison. This new species is closely related to Galloisiana iissuriensis Storozhenko, 1988 from Russia, G. sofiae Szeptycki, 1997 from North Korea, and G. nipponensis (Caudell and King 1924) from Japan. G. olgae sp. nov. is distinguished from the first two species by the number of tergital setae and by presence of two regular rows of setae on sternites. G. nipponensis dif- fers from the new species by its elongated pronotum and shorter ovipositor. Etymology. Found in Olginskij Chrebet, near the village of Olga, but named in the honor of Taiga god- dess Olga, an inspiration to S.Y.S. for many years. Biology and Biogeography. In addition to reduced eyes, the sensorial apparatus of grylloblattids is highly specialized. These features are associated with a highly localized, soil-based, and often subterranean existence. Thirteen distinct types of sensilla have been observed in modern grylloblattids (Mclver and Sut- cliffe 1982). The most noticeable are ocular setae, which serve as tactile sensors. Unlike all other species, G. olgae has heavily pigmented eyes from its earliest instar, which may indicate that this taxon is less sub- terranean than its congeners. Other externally evident microstructures of G. olgae, with an exception of the distribution of sensilla, are identical to those of G. nipponensis of Japan, indicating a close relationship between these species. Five populations of G. olgae have been identified in the Ussuri River region, all occupying banks of the Vasilkovka River. Each of these populations is distributed along a 50- to 150-m swath paralleling both river banks; each contains min- imally several hundred individuals. The entire geo- graphic range of the species is small, ^8 km^. Typi- cally, this species inhabits wet soil and can be found under rocks covered with moss. The annual average temperature of this habitat is under 8?C, and humid- ities approach 100% (unpublished data; also see Namkung 1982). The habitat of G. olgae contains veg- etation that is considerably denser from that occupied by other grylloblattodean species (Fig. 2a). Notably, G. olgae always is associated with two species of an- giosperms? one phylogenetically basal and the other advanced?namely, the magnoliid dicot Schisandra si- nensis (lUiciales: Schisandraceae) and asterid dicot Eleutherococciis senticoccus (Apiales: Apiaceae) (Plunkett et al. 1997, Qiu et al. 1999). A relictual distributional pattern of closely related genera in northwestern North America and northeast- ern Asia typifies members of the Grylloblattidae, but also characterizes the distribution of many insects (Linsley 1963) and seed plant taxa, including species of Metasequoia (Cupressaceae), Ginkgo (Gink- March 2001 VRSANSKY ET AL.: GRYLLOBLATTODEAN PALEOBIOLOGY 181 Fig. 3. Galloisiana olgae sp. nov. (a) habitat, (b) sixth immature stage, (c) head of second immature, (d) ocular seta, large and middle macrotrichia and microtrichia, (e) tarsus. 182 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 94, no. 2 goaceae), Cercidiphyllum (Cercidiphyllaceae), and many angiosperms (Manchester 1999). The only ex- ception is that the North American plant occurrences are extinct and occur commonly as fossils of Pacific Northwest floras, whereas the grylloblattids have sur- vived to the present day in both regions. An additional, more intriguing, biogeographic parallel is the distri- bution of the wingless woodroach Cryptocercus relic- tus Bey-Bienko(Blattodea: Cryptocercidae), which inhabits rotting logs and occurs in the same general habitat as the Grylloblattidae. Cryptocercus probably represents a Cenozoic lineage of cockroaches that harbors wood-digesting intestinal symbionts (Nalepa et al. 1997, Nalepa and Bandi 1999), although this interpretation has been contested (Grandcolas 1999). Populations of C. relictus have been documented in the Ussuri Region near Slavyanka and Vladivostok (Bei-Benko 1935,1950), and apparently its congeners, including C. punctulatus of the southern Appalachian Mountains, C. clevelandi of the Pacific Northwest and C. primarius of Sichuan Province in China, are relic- tual populations confined to mountainous regions (Mamaev 1973, Asahina 1991, Nalepa et al. 1997). Nevertheless, new species have been observed in South Korea, North Korea, and in Yunnan Province of China (P.V. Vidlicka, and S.Y.S., unpublished data). Grylloblattodeans, like Cryptocercus, possess an ele- vated number of distinct species in this region, but the co-occurrence of two grylloblattodean genera that are restricted to southeastern Russia, the adjacent Korean Peninsula, and northern Japan, supports the hypoth- esis that this region was the center of origin for the extant genera. Discussion Because of the close taxonomic relationship be- tween G. olgae and G. nipponensis, species similar to G olgae presumably inhabited Japan during the mid- Cenozoic, when modern genera of East Asian Gryl- loblattidae originated. Their subsequent evolution in Japan and Korea reflects isolation and adaptation to a life primarily in caves (Nagashima et al. 1982, Namkung 1982). Undoubtedly, additional speciation took place in northeastern Asia during the Pleisto- cene, when species such as G. olgae presumably orig- inated, as well as other related species in Japan, North Korea, and (sympatrically?) in the Ussuri River re- gion. The distinctions among these species are mor- phologically subtle and involve differences of G. olgae from G. ussuriensis, G nipponensis and G. sofiae that are based variously on chaetotaxy, ovipositor length, and pronotal size. A major influence in grylloblattid evolution was the ecological partitioning of their hab- itats, which in the case of the Russian taxa, probably was associated with renewed volcanic activity from the Sikhote-Alin Mountains, during the mid-Cenozoic (Bersenev and Vasiliev 1969). Later, during the Pleis- tocene or perhaps earlier in the Pliocene, some taxa became adapted to conditions at the edges of glaciers, of which modern species oi Gryllohlatta in northwest- ern North America are an example. These species evidently migrated from refugia south of receding Cordilleran and Keewatin ice sheets during the latest Pleistocene, reaching their current distribution in the Holocene, where there has been subspecific differen- tiation (Kamp 1979). Speciation within the Gryllo- blattodea is a rare example of how a taxon can adapt to extreme conditions, given changing paleoenviron- mental conditions and perhaps given competitive pressure by more advanced groups. The order Grylloblattodea is believed to be ances- tral, perphaps as a stem group, to several insect orders, including the Dermaptera, Plecoptera, and extinct Protelytroptera (Walker 1937; Rasnitsyn 1976, 1980; Rentz 1982; Storozhenko 1998). Nevertheless, the his- torical distribution of the order Grylloblattodea is enigmatic. During the Late Carboniferous, the group was present as two rare families, Protoperlidae and Daldubidae, and occurred in tropical to temperate Euramerican and Gondwanan localities (Storozhenko 1996a). Representatives from as many as 30 families are known from many fossil deposits of the Permian, albeit at the beginning of the Permian Period gryllo- blattids are absent, when the climate became more xeric and cooler, particularly for the Southern Hemi- sphere. Although largely based on negative evidence, this indicates that the group did not tolerate the cooler climates at that time. During the later Permian, the group was abundant and diverse (Storozhenko 1991, 1992); in several Permian sites the order comprises 30-50% of all insects. In particular, one member of the Grylloblattodea, Sojanidelia florea Rasnitsyn (Ideli- idae) from the mid-Permian Cherkada locality of the central Ural Mountains, consumed pollen of Lunatis- porites, Protohaploxypinus, and two species oiVittatina (Rasnitsyn and Krassilov 1996, Krassilov and Rasnitsyn 1999). These pollen genera have been identified with conifer, glossopterid, peltasperm, and probably gne- tophyte seed plants ( Clement-Westerhof 1974, Tra- verse 1988, Zavada 1991, Balme 1995). From the same deposit, another grylloblattodean, Tillyardemhia an- tennaeplana Zalessky (Tillyardembiidae), contained gut pollen oiCladaitina (Krassilov et al. 2000), whose source was the conifer-like male cones of rufloriacean cordaites (Meyen 1987, Taylor and Taylor 1993). A third species, Tschekardaenigma pollinivorum Rasnitsyn, contained gut pollen similar to Protohap- loxypinus (Rasnitsyn and Krassilov 1996). These three grylloblattodean taxa were representatives of a world- wide Late Paleozoic assemblage of spore- and pollen consuming insects (Labandeira 1998) that occupied the paleocontinents of Euramerica and Angara (the latter currently approximating Siberia), and indicate that there has been a dramatic dietary shift between Permian and recent members of the order (Krassilov and Rasnitsyn 1999). After a hiatus during the earlier Triassic, the fossil record of the Grylloblattodea re- sumes with the presence of 11 families in Middle and Upper Triassic deposits (Storozhenko 1996b), of which the Madygen locality of Kirghizistan is most notable (Storozhenko 1991, 1994). Successively younger occurrences are known from the Lower Ju- rassic localities at Soguty in Kirghizistan (Storozhenko March 2001 VRSANSKY ET AL.: GRYLLOBLATTODEAN PALEOBIOLOGY 183 1991), Braunschweig in Germany (Storozhenko 1989), and the Upper Jurassic localities of Khoutiyn- Khotgor in Mongolia (Storozhenko 1989) andKaratau in Kazakhstan (Rasnitsyn 1976, Doludenko et al. 1990), which contain rare material representing five families. From Lower Cretaceous deposits, particu- larly Baissa in the Transbaikalian region of Russia, three rare species from three families are known (Storozhenko 1989). The fossil record of the Gryllo- blattodea is absent from the mid-Cretaceous to the present. Apparently the extant taxa are a relatively late, specialized lineage adapted to cryophilic condi- tions where possibly competition with other ecolog- ically equivalent groups was minimal. Acknowledgments We are grateful to Alexandr P. 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