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Zoological Journal of the Linnean Society, 2008, 152, 17–24. With 4 figures New evolutionary evidence of Grylloblattida from the Middle Jurassic of Inner Mongolia, north-east China (Insecta, Polyneoptera) DI-YING HUANG1*, ANDRÉ NEL2* and JULIAN F. PETRULEVIČIUS3 1 Nanjing Institute of Geology and Palaeontology, State Key Laboratory of Palaeobiology and Stratigraphy, Chinese Academy of Sciences, Nanjing, 210008, China 2 CNRS UMR 5202, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 45 Rue Buffon, F-75005, Paris, France 3 CONICET, Argentina & CNRS UMR 5202, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 45 Rue Buffon, F-75005, Paris, France Received 1 March 2006; accepted for publication 13 April 2007 The small modern insect order Grylloblattida has an abundant fossil record during the Late Palaeozoic and the Mesozoicirca. The relationships between these fossil taxa and the modern grylloblattids remain unclear because most of them are based on isolated wings or have poorly preserved body features. Modern grylloblattids are wingless insects. The new grylloblattid family Plesioblattogryllidae fam. nov. is erected for the new genus and species Plesioblattogryllus magnificus gen. nov., sp. nov., from the Middle Jurassic of north-eastern China. The well-preserved specimen provides further evidence that could support its close relationships with the modern grylloblattids: (1) several very similar head structures, e.g. developed laciniae with inner row of setae, maxillary palps segmented into five, labial palps segmented into three, large labrum, and morphology of antenna; (2) paired eoplantulae on tarsomeres 1–4; (3) long ovipositor and large eggs comparable with those of modern taxa. The new genus has strongly developed mandibles with sharp pointed apical teeth and strong marginal teeth, and strong hook-like fore claws with basal teeth, suggesting it was carnivorous. The major differences between the extinct and extant Grylloblattida, such as the lack of wings, the eyes and ocelli either degenerated or absent, and the thorax degenerated in the modern forms, are probably related to their adaptation to their life under rocks and rock-crawler habits. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 17–24. ADDITIONAL KEYWORDS: evolution – Grylloblattodea – Plesioblattogryllidae fam. nov. INTRODUCTION The small order Grylloblattida (rock crawlers), currently considered as ‘living fossils’, comprises 26 living species within four genera (Vrsansky et al., 2001). Its fossil record is distinctly larger with more than 44 families, even extending to the Late Carboniferous (Storozhenko & Vranský, 1995; Storozhenko, 1997). However, many of these taxa are based on isolated wings, with their body features unknown. *Corresponding author. E-mail: [email protected]; [email protected] Extant grylloblattids are wingless insects. The definition of the Grylloblattida sensu Storozhenko (1997, 2002: 279) is far from being clear. This author indicated that the synapomorphies of the Grylloblattida are ‘absent because of paraphyletic state of the order in respect to other perlideans’. Storozhenko (1998, 2002) considered that the Embioptera are ‘derived’ from the Permian grylloblattid Sheimia Martynova, 1958. There is still not consensus on the relationships between Plecoptera, modern Grylloblattidae, Embioptera, and Dermaptera. Wheeler et al. (2001) considered that the modern Grylloblattidae are the sister group of the Dermaptera, being together the © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 17–24 17 18 D.-Y. HUANG ET AL. sister group of (Phasmida + Orthoptera), and that the (Plecoptera + Embioptera) are not directly related to this clade. Beutel & Gorb (2001) considered that the Grylloblattidae are the sister group of the clade {Phasmatodea + [Mantodea + (Isoptera + Blattodea)]}. Grimaldi (2001) supposed that the extant and fossil ‘Grylloblattodea’ falls in an unresolved polytomy with the majority of the other ‘polyneopteran’ orders. Terry & Whiting (2005) and Cameron, Barker & Whiting (2006) considered that the Grylloblattida could be the sister group of the recently discovered order Mantophasmatodea, and altogether could be the sister group of the Dictyoptera. Grimaldi & Engel (2005: 223) also supposed that the Palaeozoic and Mesozoic alate ‘grylloblattids’ could ‘represent a stem group of both apterous Grylloblattida and Mantophasmatodea. Lastly, even if Rasnitsyn (1976, 1980) listed several similarities between Blattogryllus and the extant Grylloblattidae, the potential synapomorphies of this last group with the fossil ‘Grylloblattida’ are not clear. Thus, the present discovery of a new Middle Jurassic fossil insect attributable to the ‘Grylloblattida’, with well-preserved body structures, is potentially of great interest to solve the problem of the relationships between the extant Grylloblattidae and the fossil ‘Grylloblattida’. We follow the terminology of the wing venation of Storozhenko (1998) in order to allow a more simple comparison of our fossil with those already described, even if we reserve our opinion on its real phylogenetic value, especially concerning the problem of the exact nature of the vein ‘M5’ (see Kukalová-Peck, 1991). SYSTEMATIC PALAEONTOLOGY ORDER ‘GRYLLOBLATTIDA’ WALKER, 1914 (SENSU STOROZHENKO, 2002) PLESIOBLATTOGRYLLIDAE FAM. NOV. Type genus: Plesioblattogryllus gen. nov. Diagnosis: Wing venation nearly identical to that of Blattogryllus karatavicus Rasnitsyn, 1976; mandibles very strong with a sharp pointed apical tooth and a few marginal teeth with a broad base; eyes and ocelli present; antenna short with segments 2–7 shorter than others; pronotum with broad lateral expansions; tarsomeres 1–4 with a pair of rather large euplantulae; tarsi with strong claws and no arolia; fore tarsal claw with a large base and strong basal tooth. Remark: Some new material of grylloblattids was discovered in the same locality after recent fieldwork. Two new forms seem to be closely related or attributable to the new family. Thus, a further cladistic analysis of the Plesioblattogryllidae will be possible in the near future. PLESIOBLATTOGRYLLUS GEN. NOV. Derivation of the name: Named after the Greek ‘plesios’ meaning close or near, and Blattogryllus. Type species. Plesioblattogryllus magnificus sp. nov. Diagnosis. That of the family. PLESIOBLATTOGRYLLUS MAGNIFICUS GEN. NOV., SP. NOV. FIGURES 1–3 Derivation of the name: Named after the wonderful state of preservation of the type specimen. Material: Only the holotype included (NIGP 133701). The holotype is an almost complete female specimen with brownish colour, dorsally compressed, and without counterpart. The material is deposited in the Nanjing Institute of Geology and Palaeontology. Type locality: Daohugou Village, Ningcheng County, Inner Mongolia, north-east China; Middle Jurassic, Jiulongshan Formation. Description: Head 10.5-mm long, 7.7-mm wide, rather developed, upper surface covered with small tubercles and tiny setae; eyes 3.4-mm long, 1.9-mm wide (left one), 4.0-mm apart, in lateral position, subelliptic in shape, contracted posteriorly; three ocelli 0.6 mm in diameter at middle, olive shaped, disposed in a triangle but nearly aligned between eyes; left mandible 4.8-mm long (without tip), 2.8-mm wide at base, right mandible 5.2-mm long and 2.6-mm wide at base, both very robust, asymmetrical (problem of preservation?), left mandible with a long, curved, and sharp pointed apical tooth and three marginal teeth, first and second being strong and pointed, and third marginal tooth located near base and very blunt; lacinia c. 4-mm long and very developed, pointed at apex, with a row of dense setae in middle inner margin; preserved part of maxillary palp 4.5-mm long, developed, with five segments of distinctly increasing width forwards, covered with fine setae; labial palp 4.3-mm long, developed, with three elongated segments of nearly the same width, covered with fine setae, second segment being shorter than the other two, the last one distinctly swollen and rounded at apex; clypeus 1.2-mm long, 2.8-mm wide, strong, with the shape of a transverse ellipse; labrum 2.5-mm long, developed, slightly projected in median part of anterior edge, extending to two-thirds of the © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 17–24 NEW MESOZOIC GRYLLOBLATTID FAMILY 19 Figure 1. Plesioblattogryllus magnificus gen. nov., sp. nov., holotype NIGP 133701. A. Photograph of general habitus (scale bar represents 10 mm). B. Photograph of foreleg (scale bar represents 2 mm). C. Photograph of head (scale bar represents 5 mm). D. Photograph of hind leg (scale bar represents 2 mm). E. Photograph of eggs (scale bar represents 2 mm). mandibles; antenna c. 15-mm long, as long as head + pronotum, its base located close to the mandible; about 24 antennomeres, first one large, second shortest, third rather long, segments 4–7 distinctly shorter than the others, segments 8–24 with a notable decreasing width, last one with rounded end, all antennomeres covered with fine setae; ecdysial distinctly visible, Y-shaped. Pronotum 5.2-mm long medially, 5.7-mm long laterally, 9.3-mm wide, broader than head, covered with tiny setae, with middle line and large rounded lateral expansions, anterior margin slightly projected, with pronounced angles, posterior margin rounded; mesonotum 4.2-mm long, about 4.7-mm wide, contracted backwards, distinctly narrower than pronotum. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 17–24 20 D.-Y. HUANG ET AL. Figure 2. Plesioblattogryllus magnificus gen. nov., sp. nov., holotype NIGP 133701. Drawing of general habitus. Legs all similar; femora, tibiae, and tarsi densely covered with distinct setae; fore and hind coxae visible, and rather elongate; forelegs rather sturdy, femur 6.8-mm long, 1.8-mm wide, tibia 5.7-mm long, 0.8-mm wide; median legs rather short, femur 9.2-mm long, 1.7-mm wide, tibia 6.5-mm long, 1.1-mm wide; posterior legs slender, femur 9.6-mm long, 1.9-mm wide, tibia 9.7-mm long, 0.8-mm wide, tarsi 5.0-mm long; inner margin of median tibia with three strong aligned spines; inner margin of posterior tibia with four strong aligned spines, apical end with three strong spines; all tarsi with five segments, tarsomeres 1–4 with a pair of rather large euplantulae, fifth segments rather elongate with increased width; no arolia but two claws, hind tarsal claws 0.7-mm long, strong, and rather straight, fore tarsal claws 0.9-mm long, with a large base and a strong basal tooth. Wings partly overlapping and difficult to interpret, in particular for the anal areas; wings dark infuscate with darker spots in distal halves; preserved parts of Figure 3. Plesioblattogryllus magnificus gen. nov., sp. nov., holotype NIGP 133701. Drawing of head. fore and hindwings similar, but the possible differences in the anal areas are not preserved; wings hyaline; forewing 43-mm long, about 11-mm wide; costal area between subcosta (Sc) and anterior wing margin narrow, 1.0-mm wide, in all wings, with about 20 more or less sigmoidal cross-veins between Sc and anterior wing margin; costal vein reaching wing apex; vein Sc ending on anterior wing margin 3.7 mm to wing apex; radius (R) ending very close to wing apex; Sc and R closely parallel; several cross-veins in area between Sc and R, the most distal ones being oblique; base of Rs 22 mm from wing base, in basal half of wing; area between R and radial sector (Rs) rather narrow, 1.2-mm wide in its broadest part, with one or two rows of cells; Rs posteriorly pectinate, with five main branches; cubitus apex (CuA) emerging from Cu at wing base and basally fused with common stem R + media (M); R and M + CuA separating about 4 mm from wing base; M + CuA anteriorly pectinate, with first branch media anterior (MA) emerging 7 mm © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 17–24 NEW MESOZOIC GRYLLOBLATTID FAMILY from its base; MA posteriorly pectinate, with three branches in its distal fourth; media posterior (MP) emerging 10 mm from base of M + CuA and apparently simple; CuA posteriorly pectinate with three long and simple branches; cubital area between M + CuA and cubitus posterior (CuP) broad, with long sigmoidal cross-veins; CuP simple; anal vein 1 (A1) posteriorly pectinate with three simple branches; A2 forked; hindwing about 40-mm long. Abdomen 15.8-mm long, segmentation not clearly visible, first visible abdominal segment shorter than half of the second, and other visible segments with distinct lateral lobes; last segment very small with slight lateral angles; two long, parallel-sided, and straight structures corresponding to the ovipositor, c. 8.0-mm long; right cercus (?) with three visible elongate segments, second segment distinctly longer than the first one; at least 11 visible large oliveshaped eggs, c. 3-mm long, 1.5-mm wide, with several strongly developed longitudinal ridges, first egg at level of second abdominal segment, but smaller than the more distal eggs. COMPARISON WITH FOSSIL GRYLLOBLATTIDA After the available phylogenetic analysis of the fossil taxa currently attributed to the Grylloblattida, P. magnificus gen. nov., sp. nov., would fall in the Blattogryllidae Rasnitsyn, 1976 because of the following characters, after Storozhenko (1998, 2002): (1) forewing with sigmoidal cross-veins in area between CuP and M + CuA/CuA. Storozhenko (2002) interpreted them as branches of CuA terminating on CuP; (2) costal area of fore and hindwings narrow; (3) cross-veins of forewing area between Sc and C simple, not H- or Y-shaped. Storozhenko (2002: 284) indicated that the Blattogryllidae have ‘Sc veinlets H- or Y-shaped’, which is absolutely not the case in all the taxa currently included in the family; (4) forewing veins CuA and M basally fused (no veinlet ‘M5’ between them); (5) forewing cubital area with a series of branches of CuA directed to posterior wing margin; (6) forewing CuA with a strong anterior flexure at base. In fact, its forewing venation is closely similar to that of B. karatavicus Rasnitsyn 1976. However, Storozhenko (1988a, b, 2002: 284) indicated that both B. karatavicus and Blattogryllulus mongolicus have tarsi with a large arolia and lacking claws. Plesioblattogryllus has strong claws and no arolia. These two characters are sufficient to separate Plesioblattogryllus from the Blattogryllidae, as the type species of this last family is B. karatavicus. Another possible difference is the thorax as broad as pronotum in the Blattogryllidae, unlike the thorax 21 with broad lateral expansions of Plesioblattogryllus. But it is necessary to be prudent with this character because of the taphonomic deformations of the body structures in these insects with large thoraces and heads. Interestingly, the thorax of Parakhosara nasuta Storozhenko, 1993 (Megakhosaridae Sharov, 1961; sister group of Blattogryllidae + Mesoblattinidae, after Storozhenko, 2002) is similar to that of Plesioblattogryllus. We also have to compare Plesioblattogryllus with the taxa currently attributed to the Blattogryllidae because some of these taxa are based on isolated wings: (i) Baharellus Storozhenko, 1988a (two species: Baharellus lineatus Storozhenko, 1988a, Middle Jurassic, and Baharellus madygensis Storozhenko, 1992; Triassic). Both have the forewing Rs with only two main branches and with a corresponding area relatively reduced (Storozhenko, 1988a, 1992, 1998). (ii) Blattogryllulus Storozhenko, 1988a (two species: B. mongolicus Storozhenko, 1988a, and Blattogryllulus lucidus Storozhenko, 1988a, both Upper Jurassic). Both have a forewing MA and CuA simple, and an Rs not pectinate, with a double fork (Storozhenko, 1988a). (iii) Parablattogryllus Storozhenko, 1988a (one species: Parablattogryllus obscurus Storozhenko, 1988a, Lower Cretaceous). This taxon is based on a hindwing, even if it was labelled as a forewing in Storozhenko (1988a: fig. 1.i). Thus, it is difficult to compare with our specimen but its Rs is not pectinate. (iv) Dorniella Bode, 1953 (two species: Dorniella pulchra Bode, 1953; Lower Jurassic, Dorniella primitiva Storozhenko, 1992, Middle or Upper Triassic). Both have Rs with only two branches (Storozhenko, 1988a, 1992). D. primitiva is based on a hindwing, which is very difficult to compare with the known forewings of D. pulchra. Thus the generic identity of these two species is questionable. (v) Costatoviblatta Storozhenko, 1992 (one species: Costatoviblatta aenigmatosa Storozhenko, 1992; Middle or Upper Triassic). (vi) Mesoblattogryllus Storozhenko, 1990 (three species: Mesoblattogryllus intermedius Storozhenko, 1990; Mesoblattogryllus conjuctus Storozhenko, 1992; both from Middle or Upper Triassic, Mesoblattogryllus longipennis Storozhenko, 1992; Lower or Middle Jurassic). These two genera have a short Sc and its first branch of Rs is pectinate, instead of being simple (Storozhenko, 1992, 1998). (vii) Baharellinus Storozhenko, 1992 (two species: Baharellinus dimidiatus Storozhenko, 1992; Baharellinus pectinatus Storozhenko, 1992; both from Middle or Upper Triassic). They have very different wing shapes, rounded in B. dimidiatus and very elongate © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 17–24 22 D.-Y. HUANG ET AL. in B. pectinatus. Nevertheless, their Sc is short and Rs has three branches (Storozhenko, 1992, 1998). (viii) Protoblattogryllus Storozhenko, 1990 (three species: Protoblattogryllus zajsanicus Storozhenko, 1990; Protoblattogryllus asiaticus Storozhenko, 1990; Protoblattogryllus abruptus Storozhenko, 1990; Middle or Upper Triassic). They have a short Sc and an Rs with two branches (Storozhenko, 1990). (ix) Microblattogryllus Storozhenko (1990) (one species: Microblattogryllus variabilis Storozhenko, 1990; Middle or Upper Triassic). It has a short Sc and a simple Rs. (x) Anoblattogryllus Storozhenko, 1990 (one species: Anoblattogryllus fundatus Storozhenko, 1990; Middle or Upper Triassic). Its Sc is short. (xi) Gryphopteron Handlirsch, 1939 (one species: Gryphopteron molle Handlirsch, 1939; Lower Jurassic). Its MA is fused with Rs, and its Rs has two distal branches (Ansorge, 1996). (xii) Megablattogryllus Storozhenko, 1990 (two species: Megablattogryllus magister Storozhenko, 1990; Megablattogryllus austerus Storozhenko, 1990; Megablattogryllus pinguis Storozhenko, 1990; Middle or Upper Triassic). Their CuA emerges first from M + CuA, then MP, and lastly MA, unlike our fossil. (xii) Blattogryllus Rasnitsyn, 1976 (two species: B. karatavicus Rasnitsyn, 1976; Upper Jurassic, Blattogryllus rasnitsyni Storozhenko, 1990; Lower Jurassic). The only difference of Plesioblattogryllus with Blattogryllus in the wing venation is its MP being apparently simple. All other characters are identical (Rasnitsyn, 1976; Storozhenko, 1990). Remark: All the species currently attributed to the Blattogryllidae based on isolated wings now fall into uncertain position between the Blattogryllidae and the Plesioblattogryllidae because these two families can be separated only on the basis of the body characters. EVOLUTIONARY RELATIONSHIP The present discovery and characters listed in Rasnitsyn (1976) show that there are numerous structures of at least some fossil ‘Grylloblattida’ (Blattogryllidae and Plesioblattogryllidae) similar to those of extant Grylloblattidae. But these characters have never been polarized in a phylogenetic analysis. Thus, the establishment of a direct phylogenetic relationship between these groups remains questionable. A well-supported phylogenetic analysis of all polyneopterous orders, including the fossil taxa, is necessary. Re-analyses of the characters available in fossils (wing venation) could help to solve this problem, as Béthoux & Nel (2002a, b) already showed for the orthopteroid taxa. Although more than 44 extinct families have been attributed to Grylloblattida, very few fossils have well-preserved body structures (Storozhenko & Vranský, 1995; Storozhenko, 1997). The new fossil material from the Middle Jurassic of north-east China provides some new similarities with the modern grylloblattids. 1. Some head structures of P. magnificus gen. nov., sp. nov. correspond to those of the living grylloblattids, namely: laciniae developed with an inner row of marginal setae; maxillary palp segmented into five; labial palp segmented into three; labrum developed; antenna with a large first segment, smallest second segment, and segments 2–7 distinctly shorter than the others. The structure of the antennomeres, viz. antennomere 2 smaller than 3, and antennomeres 4–7 distinctly shorter than the other distal segments, visible in Plesioblattogryllus, is a rather typical feature present in extant grylloblattids, but also in the modern Embioptera Embia major (Crampton, 1917: fig. 9). The rather developed antennomere 3 and its adjacent segments are especially similar to Galloisiana yezoensis (Asahina, 1961), and the third antennomere is even more developed in some other species such as Galloisiana biryongensis (Namkung, 1974). The mandibles and eyes in the new genus are remarkably larger than those of the extant forms. Moreover, the large ocelli of our fossil, absent in all modern grylloblattids, is an important difference. In addition, no evidence indicates that the clypeus of Plesioblattogryllus is divided into an anteclypeus and a postclypeus, as in extant forms. 2. The five-segmented tarsi and the tarsal euplantulae of P. magnificus gen. nov., sp. nov. are strongly similar to those of some modern grylloblattids (Walker, 1938: pl. 10; Caudell & King, 1924: plate 3: 2–3; Namkung, 1974: figs 10,11; Storozhenko, 1998; Beutel & Gorb, 2006). However, Crampton (1927: 125) indicated that ‘some Mantids have flange-like tarsal pads very suggestive of those occurring in Galloisiana and in certain Gryllacrids’ (see also Crampton, 1933). Thus, this type of euplantulae seems to be not uniquely present in Grylloblattida, but also occurs in some Mantodea and Orthoptera. Thus, it is not sufficient alone to attribute Plesioblattogryllus to the same clade as the extant Grylloblattidae. Nevertheless, the wing venation and body structures of Plesioblattogryllus strongly differ from those of the Mantodea and Orthoptera, suggesting that it belongs to a very different order. The marginal and apical strong spines of posterior tibia in Plesioblattogryllus also display similarities with the modern forms. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 17–24 NEW MESOZOIC GRYLLOBLATTID FAMILY 3. The ovipositor and eggs are similar to those of modern forms. We interpret two long and parallelsided straight structures as the ovipositor bases. Such structures are not only comparable with the ovipositor of extant female grylloblattids, but also to the similar structures described in some other extinct grylloblattid taxa (Storozhenko & Novokshonov, 1994). The large elliptic eggs are similar to those of modern grylloblattids (Crampton, 1927, 1933). They occupy a broad space in the abdomen, suggesting the adult female of Plesioblattogryllus had large ovaries. Similar size and arrangement of eggs in the abdomen are also found in some other Mesozoic grylloblattids (family Tshekardominidae) (Novokshonov & Aristov, 2002). In addition, the body of P. magnificus is covered with tiny setae as are those of the modern grylloblattids. The well-developed wings of this large insect suggest it was probably an active flyer. Its fore tarsal claws differ from those of hind legs (the middle claws are not visible) in their hook-like shape with a large base and bearing a strong basal tooth. Moreover, the fore tarsal claws are also distinctly longer than the rather straight hind claws. After preservation, the fore tarsal claws and the most apical tarsomeres were probably movable structures. Such structures could have been associated with the capture of small prey. The very strong mandibles with a sharp pointed apical tooth could help to kill the prey, and the strong marginal teeth would help to destroy it. The developed eyes and ocelli suggest it was an active hunter. Plesioblattogryllus was probably a predator. The fossil evidence indicates that the Permian grylloblattid Tillyardembia was probably a pollen eater (Afonin, 2000). Therefore the food habits of the early grylloblattids were very diverse. The modern grylloblattids probably became adapted to live under rocks or hidden in moss liver through the degeneration of their sensitive organs and wings. Even the eyes are absent in some cave-living species (Namkung, 1974). Their thorax trends to be very developed in relation to their habit of being a strong rock crawler. Their mandibles degenerated in relation to their habit of omnivory. Nevertheless, the modern Grylloblatta can also attack a living insect if it is very hungry (Ford, 1926). ACKNOWLEDGEMENTS We are pleased to acknowledge the joint support provided by the NSFC (grant no. 40672013, 40632010), the Major Basic Research Projects of MST of China (2006CB806400), and the State Key Laboratory of Palaeobiology and Stratigraphy (No. 053106, CAS). We thank Prof. Shu-Yong Wang (Institute of Zoology, Chinese Academy of Sciences) for kind bibliographic help. 23 REFERENCES Afonin SA. 2000. Pollen grains of the genus Cladaitina extracted from the gut of the Early Permian insect Tillyardembia (Grylloblattida). Paleontological Journal 34: 105–109. Ansorge J. 1996. Insekten aus dem Oberen Lias von Grimmen (Vorpommern, Norddeutschland). Neue Paläontologische Abhandlungen 2: 1–132. Asahina S. 1961. A new Galloisiana from Hokkaido (Grylloblattoidae). Kontyû 29: 85–87. Béthoux O, Nel A. 2002a. Venation pattern of Orthoptera. Journal of Orthoptera Research 10: 195–198. Béthoux O, Nel A. 2002b. Venation pattern and revision of Orthoptera sensu nov. and sister groups. Phylogeny of Palaeozoic and Mesozoic Orthoptera sensu nov. Zootaxa 96: 1–88. Beutel RG, Gorb SN. 2001. Ultrastructure of attachment specializations of hexapods (Arthropoda): evolutionary patterns inferred from revised ordinal phylogeny. Zeitschrift für Zoologische Systematik und Evolutionforschung 39: 177–207. Beutel RG, Gorb SN. 2006. A revised interpretation of the evolution of attachment structures in Hexapoda with special emphasis on Mantophasmatodea. Arthropod Systematics and Phylogeny 64: 3–25. Bode A. 1953. Die Insektenfauna des Ostniedersächsischen Oberen Lias. Palaeontographica, (A) 103: 1–375. Cameron SL, Barker SC, Whiting MF. 2006. Mitochondrial genomics and the new insect order Mantophasmatodea. Molecular Phylogenetics and Evolution 38: 274–279. Caudell AN, King JL. 1924. A new genus and species of the notopterous family Grylloblattidae from Japan. Proceedings of the Entomological Society of Washington 26: 53–60. Crampton GC. 1917. A comparison of the antennae of the Grylloblattidae and Embiidae to demonstrate the relationship of these two groups of insects. Canadian Entomologist 49: 213–217. Crampton GC. 1927. The abdominal structures of the orthopteroid family Grylloblattidae and the relationships of the group. The Pan-Pacific Entomologist 3: 115–135. Crampton GC. 1933. The affinities of the archaic orthopteroid family Grylloblattidae, and its position in the general phylogenetic scheme. Journal of the New York Entomological Society 41: 127–166. Ford N. 1926. On the behaviour of Grylloblatta. Canadian Entomologist 58: 66–70. Grimaldi D. 2001. Insect evolutionary history from Handlirsch to Hennig, and beyond. Journal of Paleontology 75: 1152–1160. Grimaldi D, Engel MS. 2005. Evolution of the insects. Cambridge: Cambridge University Press. Handlirsch A. 1939. Neue Untersuchungen über die fossilen Insekten mit Ergänzungen und Nachträgen sowie Ausblicken auf phylogenetische, palaeongraphische und allgemeine biologische Probleme. Teil 2. Annalen des Naturhistorischen Museums in Wien 49: 1–240. Kukalová-Peck J. 1991. Chapter 6: Fossil history and the evolution of hexapod structures. In: Naumann ID, ed. The © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 17–24 24 D.-Y. HUANG ET AL. insects of australia, a textbook for students and research workers. 2nd ed. vol. 1. Melbourne: Melbourne University Press, 141–179. Namkung J. 1974. A new species of cave dwelling Grylloblattoidea (Grylloblattidae) from Korea. Korean Journal of Entomology 4: 1–7. Novokshonov VG, Aristov DS. 2002. New and little-known Permian insects (Insecta: Grylloblattida; Orthoptera) from the Chekarda locality, central Ural mountains. Paleontological Journal 6: 73–77. Rasnitsyn AP. 1976. [Grylloblattidae – living representatives of order Protoblattodea (Insecta).] Doklady Akademii nauk SSSR, Moscow 228: 502–504. [in Russian, translated into English in Doklady Biological Sciences 228 (1–6): 273– 275.] Rasnitsyn AP. 1980. [Order Grylloblattida.]. In: Rohdendorf BB, Rasnitsyn AP, eds. [Istoricheskoe razvitie klassa nasekomykh [Historical development of the class Insecta.]. Trudy Paleontologicheskogo Instituta Akademii Nauk SSSR 175: 150–153 + 166–171. [in Russian]. Sharov AG. 1961. [Order Protoblattodea.]. In: Rohdendorf BB, Becker-Migdisova EE, Martynova OM, Sharov AG, eds. [Paleozoic insects of the Kuznetsk basin.]. Trudy Paleontologicheskogo Instituta Akademii Nauk SSSR 85: 157–164. Storozhenko SY. 1988a. New Mesozoic Grylloblattida. Paleontological Journal 22: 48–54. Storozhenko SY. 1988b. A review of the family Grylloblattidae (Insecta). Articulata 3: 167–181. Storozhenko SY. 1990. New Permian and Mesozoic insects (Insecta, Grylloblattida: Blattogryllidae, Geinitziidae) from Asia. Paleontological Journal 24: 53–61. Storozhenko SY. 1992. New Mesozoic grylloblattid insects from Soviet Central Asia. Paleontological Journal 26: 67–75. Storozhenko SY. 1993. [Megakhosaridae.] pp. 100–112. In: Ponomarenko AG, ed. [Mesozoic insects and ostracods of Asia.]. Trudy Paleontologicheskogo Instituta Akademii Nauk SSSR 252: 1–160. [in Russian]. Storozhenko SY. 1997. Classification of order Grylloblattida (Insecta), with description of new taxa. Far Eastern Entomologist 42: 1–20. Storozhenko SY. 1998. Sistematika, filogeniya i evolyutsiya grilloblattidovykh nasekomykh (Insecta: Grylloblattida) [Systematics, phylogeny and evolution of the grylloblattids (Insecta: Grylloblattida).] Vladivostok: Dal’nauka, 1–207. Storozhenko SY. 2002. Chapter 2.2.2.2.1. Order Grylloblattida Walker, 1914 (= Notoptera Crampton, 1915, = Grylloblattodea Brues et Melander, 1932, +Protorthoptera Handlirsch, 1906, = Paraplecoptera Martynov, 1925, +Protoperlaria Tillyard, 1928). In: Rasnitsyn AP, Quicke DLJ, eds. History of insects. Dordrecht, Boston, London: Kluwer Academic Publishers, 278–284. Storozhenko SY, Novokshonov VG. 1994. Revision of the Permian family Sojanoraphidiidae (Grylloblattida). Russian Entomological Journal 3: 37–39. Storozhenko SY, Vranský P. 1995. New fossil family of the order Grylloblattida (Insecta: Plecopteroidea) from Asia. Far Eastern Entomologist 19: 1–4. Terry MD, Whiting MF. 2005. Mantophasmatodea and phylogeny of the lower neopterous insects. Cladistics 21: 240– 257. Vrsansky P, Storozhenko SY, Labandeira CC, Ihringova P. 2001. Galloisiana olgae sp. nov. (Grylloblattodea: Grylloblattidae) and the paleobiology of a relict order of insects. Annals of the Entomological Society of America 94: 179– 184. Walker EM. 1914. A new species of Orthoptera forming a new genus and family. Canadian Entomologist 46: 93– 99. Walker EM. 1938. On the anatomy of Grylloblatta campodeiformis Walker. 3. Exoskeletton and musculature of the neck and thorax. Annals of the Entomological Society of America 31: 588–640. Wheeler WC, Whiting M, Wheeler QD, Carpenter JM. 2001. The phylogeny of the extant hexapod orders. Cladistics 17: 113–169 + 403–404 (erratum). © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 17–24