New and Poorly Known Grylloblattids (Insecta: Grylloblattida) from the Lopingian of the Lebombo Basin, South Africa

ABSTRACT A new genus and species of the grylloblattid family Chaulioditidae, Iphikozulu kwayayaensis Aristov & Mostovski, gen. et sp. n., is described from the Lopingian locality of KwaYaya in KwaZulu-Natal, South Africa. This is the first record of the Chaulioditidae in Gondwanan deposits. The genera Triadosialis Handlirsch, 1906, Protomia Aristov, 2004 and Miralioma Aristov, 2004 are transferred to the family Chaulioditidae, and the genus Kargalella Martynov, 1937 is transferred to the Liomopteridae. Neoliomopterum picturatum Riek, 1976 from the Lopingian locality at Emakwezini Railway Station in KwaZulu-Natal is redescribed and transferred from Grylloblattida incertae sedis to the family Liomopteridae. The subfamily Kargalellinae Aristov, 2004 is synonymised under Liomopteridae Sellards, 1909.

Thus, the grylloblattid fauna of the South African Lopingian is represented by two families: the dominant Liomopteridae (63 % of all grylloblattids) and the considerably rarer Megakhosaridae (22 %). The family Chaulioditidae, being widely distributed in the Tatarian (= Lopingian, or Upper Permian) of Russia, has so far been unknown in Gondwanan deposits.
A new representative of the Chaulioditidae, Iphikozulu kwayayaensis gen. et sp. n., is herewith described from the locality of KwaYaya in KwaZulu-Natal. In addition, Neoliomopterum picturatum from the contemporaneous Emakwezini Railway Station locality is transferred from Grylloblattida incertae sedis to the family Liomopteridae.
To summarise, the composition of the South African grylloblattid fauna is typically Lopingian, with dominating Liomopteridae, and rare Megakhosaridae and Chaulioditidae. The only difference is a high ratio (35 %) of grylloblattids to other insect taxa.

Locality information
The holotype of Iphikozulu kwayayaensis gen. et sp. n. was found in fine-grained, olive-grey mudstone in a railway cutting approx. 25 km west of Empangeni along the R34 and approx. 200 m west of the KwaYaya railway siding (28°41.915'S:31°40.730'E) (Fig. 1). The holotype is housed at the Natal Museum (NMSA).

Geological setting and palaeoenvironment
The fossiliferous deposits at KwaYaya lie within the upper part of the Emakwezini Formation, Adelaide Subgroup, Beaufort Group, Karoo Supergroup. The Emakwezini Formation was deposited in the Lebombo Basin, and crops out in a narrow strip just inland of, and parallel to, the north-eastern KwaZulu-Natal coastline. The formation is considered to be lithostratigraphically equivalent to the Normandien Formation (incorporating the previously recognised Estcourt Formation) in the north-eastern and eastern parts of the main Karoo Basin (e.g., Johnson et al. 2006). A review of the Emakwezini Formation is provided elsewhere (Bordy & Prevec 2008). Bordy and Prevec (2008) interpreted the autochthonous nature of the deposits at KwaYaya, together with various associated sedimentary features, as indicative of deposition in a floodplain setting in the immediate vicinity of a broad, shallow, gently meandering channel in the overbank area of a large fluvial system. According to general climatic reconstructions (Chumakov & Zharkov 2003), this locality was situated within the southern temperate semiarid belt of middle latitudes. Locally, the Emakwezini Formation appears to have been deposited under cool, humid, perennially moist conditions. The latter assumption is supported by records of grylloblattids, which are known to be associated with cold and moist environments from their early days until the present (e.g., Shcherbakov 2008) contrary to the statement that they are indicative of warmer and drier conditions (Geertsema et al. 2002).

Associated flora
The KwaYaya locality has yielded a glossopterid-dominated palaeoflora, containing multiple elements typical of Late Permian floras from the main Karoo Basin of South Africa (e.g., Lacey et al. 1975;Anderson & Anderson 1985). These include at least five morphotypes of Glossopteris leaves and the glossopterid ovuliferous organs Lidgettonia africana, Rigbya arberioides and Plumsteadia gibbosa, and the glossopterid polleniferous organ Eretmonia natalensis. Also present are the sphenopsids Trizygia speciosa, Phyllotheca australis and Benlightfootia sp. (Bordy & Prevec 2008). TAXONOMY Order Grylloblattida Walker, 1914Suborder Grylloblattina Walker, 1914Family Chaulioditidae Handlirsch, 1906 Type genus: Chauliodites Heer, 1864 with 11 species from the Severodvinian-Olenekian of Russia, Induan of Mongolia, Olenekian of Germany, Anisian of France and Middle Triassic of China (Aristov, 2003(Aristov, , 2004b(Aristov, , 2008. Diagnosis: Costal space broader than subcostal one; M not fused with CuA, branching beyond RS base; CuA divided into CuA 1 and CuA 2 . Description: Small and middle-sized insects. Anterior margin of forewing convex. Costal space broader than subcostal one. SC ends beyond midlength or in distal third of wing. RS starts in basal third of wing, with 2-4 branches. Base of M free, M 5 absent, M bifurcates into MA and MP beyond RS base. MA with 2-4 branches, sometimes forming regular comb; MP with 1 or 2 branches. CuA bifurcates into simple CuA 1 and CuA 2 , and may change its state from concave to convex almost immediately after branching. Intercubital space narrow, traversed by simple crossveins. Anal area short and broad.  Handlirsch, 1906, Protomia Aristov, 2004and Miralioma Aristov, 2004. The genus Triadosialis was described by Handlirsch (1906) for Chauliodites zinkeni Heer, 1864 from the Lower Triassic (Middle Bundsandstein) locality of Gödevitz in Germany (Heer 1864). Handlirsch assigned this genus to the family Chaulioditidae in the order Megaloptera. The family Chaulioditidae, with the only species Ch. picteti Heer, 1864, was transferred to the order Grylloblattida and considered a senior synonym of the family Tomiidae Martynov, 1936 (Aristov 2004b). The genera Protomia and Miralioma were described in the family Liomopteridae (Aristov 2004c). Both of them differ from other liomopterids in the M that branches late, and from chaulioditids in the branched CuA 1 . The position of the M in relation to the RS base is a more reliable character than the number of CuA 1 branches. Thus, the genera Triadosialis, Protomia, Miralioma and Iphikozulu gen. n. are herewith transferred to the family Chaulioditidae, the type genus of which is characterised by the M that branches beyond the RS base. The description of the new genus and the transfer of the genera Triadosialis, Protomia and Miralioma to Chaulioditidae have necessitated changes to the familial diagnosis. Previously, one of the diagnostic features was simple CuA 1 . Currently, a broad costal space and M branching beyond the RS base remain diagnostic for chaulioditids.
The genus Kargalella Martynov, 1937 from the Urzhumian of Russia was placed in a separate subfamily of Chaulioditidae on the basis of a broad costal space and simple CuA 1 . The M in Kargalella bifurcates proximal to the RS base (Aristov 2004b). The combination of these characters precludes us from keeping this genus in Chaulioditidae, and it is transferred to the family Liomopteridae, with the subfamily Kargalellinae Aristov, 2004 being considered as a new junior subjective synonym of Liomopteridae Sellards, 1909. Kargalella differs from other liomopterids in the combination of fused RS+MA and simple CuA 1 .
An incomplete grylloblattid forewing, which has been recently found at a locality near Colenso (Prevec et al. 2009: 486, pl. XIV, figs 1, 2), also probably belongs to Chaulioditidae on the grounds of its wing venation. Regrettably, its further identification is difficult without a closer examination of the specimen.
Genus Iphikozulu Aristov & Mostovski, gen. n. Etymology: From Zulu iPhiko (wing) and Zulu (Zulu Kingdom). Gender masculine. Type and only species: I. kwayayaensis Aristov & Mostovski, sp. n. Diagnosis: Middle-sized insects. Costal space near RS base 3 times as broad as subcostal one. SC with simple anterior branches, ends before distal third of wing. RS starts in basal quarter of wing. CuA 1 starts branching before midlength, with four branches. Comparison: The new genus is closest to Protomia from the Kazanian deposits of Russia, from which it differs in simple branches of SC, as well as in CuA 2 and CuP ending on the posterior margin of the wing. In Protomia, CuA 2 anastomoses with CuA 1 , and CuP terminates on CuA 2 (Aristov 2004c). Family Liomopteridae Sellards, 1909Genus Neoliomopterum Riek, 1976 Type and only species: N. picturatum Riek, 1976, by monotypy. Diagnosis: Small insects. Anterior margin of forewing weakly convex, costal space slightly broader than subcostal one. SC with simple anterior branches, enters C near wing tip. RS base near wing midlength, at level of separation of M into MA and MP. Branches of CuA simple. Riek, 1976 Figs 2C, 2D
Remarks: N. picturatum had originally been described as a forewing fragment of a representative of the family Liomopteridae (Riek 1976b). It remained in this family until some time ago (Carpenter 1992), but later was transferred to Grylloblattida incertae sedis on the basis of its incomplete preservation (Storozhenko 1998). Re-examination of the holotype illustrated by a photograph in the original description has shown that the preservation of the wing is almost complete. This species is characterized by a shortened wing, which is twice as long as it is broad (wings of the majority of grylloblattids are three times as long as broad), and by the stems of RS, MA, MP and CuA thickened basally and at the wing midlength. Such characters are typical of shortened wings, which can be divided into brachypterous and micropterous forms. In brachypterous individuals, the wing is longer than half of a fully developed wing, and does not have considerably modified venation. In micropterous individuals, the wing is shorter than half of a normal wing, and both its shape and venation are heavily modified (Sinitshenkova 1987). Shortened wings are recorded in some grylloblattids. Micropterous wings are known in Protoblattiniella minutissima Meinier, 1912 from the Upper Carbo-niferous locality of Commentry in France and Sylvamicropteron harpax Aristov, 2004 from the Kungurian locality of Tshekarda in Russia. The genus Protoblattiniella belongs to the family Protoperlidae (Storozhenko 2002), and Sylvamicropteron has been described as Grylloblattida incertae sedis (Aristov 2004a). Despite the unusual proportions of the wing, N. picturatum still displays venation typical of grylloblattids, suggesting that we are dealing with a brachypterous wing. A rather wide costal space that is traversed by simple branches of SC, and a simple CuA 1 are features typical of two grylloblattid families, viz. Liomopteridae and Chaulioditidae. The latter family is characterized by M being divided into MA and MP beyond the RS stem (Aristov 2004b). This character is missing in Neoliomopterum, which favours its assignment to the Liomopteridae. N. picturatum differs from other liomopterids in having both a long SC, which reaches the wing apex, and oligomerized venation of RS, MA, MP and CuA 1 . These differences are possibly caused by changed proportions of the wing, but the comparison with other known liomopterids does not offer an opportunity to synonymise this genus.