Permula Handlirsch 1919.

Aliculidae Storozhenko, 1997: 8 (type genus: Alicula Schlechtendal, 1913); Storozhenko 1998: 97. Invalid name according to Article 39 of the Code (ICZN, 1999).

In Permulidae CuA is either simple or branching distal of its middle and CuA2 not individualized. In other families of the order Grylloblattida either the first fork of CuA is situated near the base, or CuA1 and CuA2 are well individualized.

Medium-sized insects. Forewing uncolored, without spots and stripes. Costal area narrow to broad. SC ending near the apical third of wing or apically. RS originates in basal third or quarter of wing. The base of M and CuA distinctly separated or fused. M forked distinctly before or near the base of RS. MA simple or branched. MP desclerotized near the middle of wing. M5 not individualized. CuA not divided into CuA1 and CuA2, CuA is branched distal of its middle, with 2–4 branches, distal branches of CuA not parallel to posterior margin of wing, rare CuA simple. CuP simple, straight or gently curved. A1 simple. Cross-veins either simple or forming two or more rows of cells in the majority of areas. Hind wing similar to forewing but anal area enlarged. Body unknown.

Seven genera are known from the Permian of Europe and North America, two of them are described as new below.

The generic name Alicula Schlechtendal, 1913 (Insecta) is a junior homonym of Alicula Eichwald, 1830 (Mollusca). The name Alicula Schlechtendal, 1913 must be replaced by Permula Handlirsch 1919 (the next oldest available name from among its synonyms). Thus, the family name Aliculidae Storozhenko, 1997 must be replaced by a new replacement name Permulidae based on the valid name of the former type genus according to Article 39 of the Code (ICZN, 1999).

The base of costal area in forewing broadly rounded; costal area is considerably broader than subcostal one, crossed with simple or furcated anterior branches of SC; RA simple; RS with more than two branches; the base of M distinctly separated from CuA; M forked before the base of RS; CuA with 2–4 branches, first fork of CuA is situated proximal to fork of M; the base of anterior branch of CuA sclerotized; CuP curved or straight; A1 and A2 simple.

Four species from the Lower and Middle Permian of Europe.

Apparently being unaware of Schlechtendal’s paper, Handlirsch (1919) described Permula lebachensis as a new genus and species based on the same specimen as Alicula lebachensis Schlechtendal, 1913. Later Carpenter (1992) synonymized Permula lebachensis Handlirsch, 1919 with Alicula lebachensis Schlechtendal, 1913 and included Permula acra Kukalová, 1964 in the genus Alicula. But the generic name Alicula Schlechtendal, 1913 is a junior homonym of Alicula Eichwald, 1830 and must be replaced by Permula Handlirsch 1919. Herein two species are transferred from Sojanopermula to Permula based on the shape of costal area and one new species is described. Moreover, Permula acra Kukalová, 1964 is not congeneric with other species of Permula and is placed in a new genus of the family Ideliidae (see below).

Asselian Stage, Lebacher Schichten (the Lower Permian).

Lebach locality, Saar-Nahe Basin, Saarland, Germany.

Anterior margin of forewing gently convex; costal area as broad as intercubital one; RS distinctly separated from MA; MP forked; CuA with four branches; CuP curved; cross-veins in almost all areas forming a row of two cells, cross-veins simple in the basal portion of areas between SC and R, and between M and CuA, cross-veins forming rows of three cells in the basal portion of intercubital area.

Length of forewing about 25 mm.

Permula lebachensis Handlirsch 1919 is an objective synonym of Alicula lebachensis Schlechtendal 1913 because the descriptions of both species are based on the same specimen.

Holotype only.

Kungurian Stage, Koshelevka Formation (the Lower Permian).

Tshekarda locality, Krasnaya Gora Mountain on the left bank of Sylva River near the Tshekarda village, Suksun District, Perm Region, Russia.

Anterior margin of forewing convex; SC reaching the apical quarter of wing; costal area distinctly broader than intercubital one; RS with more than two branches, distinctly separated from MA; MA simple or bifurcate near the apex of forewing; MP with three or four branches directed to the posterior margin of wing; CuA with three branches; CuP curved; A1 and A2 straight and simple; cross-veins are simple or forming a double row of cells in the areas between MP and CuA, and between CuA and CuP.

Length of forewing 30 mm.

Holotype only.

Kungurian Stage, Koshelevka Formation (the Lower Permian).

Tshekarda locality, Krasnaya Gora Mountain on the left bank of Sylva River near the Tshekarda village, Suksun District, Perm Region, Russia.

RS with more than two branches, distinctly separated from MA; MA simple or probably with short terminal fork; MP with three branches sharply turning towards the posterior margin of wing; CuA with two branches; CuP straight; A1 and A2 simple and straight; cross-veins are simple or forming a double row of cells.

Length of forewing about 23 mm.

Holotype PIN 94/1027, positive and negative imprints of an incomplete forewing.

Kazanian Stage, Lower Kazanian Substage, Iva-Gora Beds (the Middle Permian).

Soyana locality, right bank of the Soyana River 56–60 km upstream of the mouth, Mezen District, Arkhangelsk Region, Russia.

Anterior margin of forewing distinctly convex; costal area considerably broader than intercubital one; R distinctly separated from M; CuA with two branches; CuP straight; cross-veins forming numerous rows of cells in all areas.

Length of forewing about 30 mm.

The new species is named in honor of the Russian geologist M.B. Edemsky, who collected the type specimen.

The base of costal area in forewing distinctly narrowing; costal area is broader than subcostal one, crossed with simple or furcated anterior branches of SC; RA simple; RS originated in the basal quarter of the wing, with three branches; the base of M distinctly separated from CuA; M forked before the origin of RS; CuA with three or four branches, first fork of CuA is situated distinctly proximal to fork of M; the base of anterior branch of CuA sclerotized; CuP straight.

Two species from the Middle Permian of Europe.

Holotype only.

Kazanian Stage, Lower Kazanian Substage, Iva-Gora Beds (the Middle Permian).

Soyana locality, right bank of the Soyana River 56–60 km upstream of the mouth, Mezen District, Arkhangelsk Region, Russia.

Anterior margin of forewing straight; SC reaching apical one fifth of the wing; costal area as broad as intercubital one; RS distinctly separated from MA; MA with fork; MP with three branches; CuA with three branches; cross-veins in the almost all areas simple, but forming a row of two cells in the area between RA and RS, in the basal portion of areas between MA, MP and CuA, and between branches of CuA; in the intercubital area cross-veins forming numerous rows of cells.

Length of forewing about 34.6 mm.

Holotype PIN 117/301, positive imprint of incomplete forewing. Paratype PIN 117/195, positive imprint of forewing without anal area.

Kazanian Stage, Lower Kazanian Substage, Iva-Gora Beds (the Middle Permian).

Soyana locality, right bank of the Soyana River 56–60 km upstream of the mouth, Mezen District, Arkhangelsk Region, Russia.

Anterior margin of forewing straight; SC reaching apical quarter or one fifth of wing; costal area distinctly broader than intercubital area; RS separated from MA; MA simple; MP with two branches; CuA with four branches; cross-veins in the almost all areas simple, but forming a row of two cells in the basal portion of area between MP and CuA; in the intercubital area cross-veins forming two or three rows of cells.

Length of forewing about 30 mm.

The new species is named in honor of the Russian paleoentomologist Prof. A.P. Rasnitsyn.

Kazanalicula reducta Aristov & Storozhenko, sp. n.

Similar to Elmopterum and Tshepanichoptera by the narrow costal area, but distinct from these in the simple CuA.

The base of costal area in forewing gentle rounded; costal area is narrower than subcostal one, crossed with simple anterior branches of SC; RA simple; RS originated in the basal third of the wing, with two branches; the base of M distinctly separated from CuA; M forked slightly before the base of RS; CuA simple; CuP straight; A1 simple, A2 furcated.

One species from the Middle Permian of Europe.

After the name of the Kazanian Stage and genus Alicula; gender feminine.

Holotype PIN 117/467, positive imprint of forewing without apical portion.

Kazanian Stage, Lower Kazanian Substage, Iva-Gora Beds (the Middle Permian).

Soyana locality, right bank of the Soyana River 56–60 km upstream of the mouth, Mezen District, Arkhangelsk Region, Russia.

Anterior margin of forewing weakly convex; costal area 2.5 times narrower than intercubital one; RS distinctly separated from MA; MA simple; MP with three branches directed to the posterior margin of wing; CuA simple; cross-veins in almost all areas forming a double row of cells.

Length of forewing about 20 mm.

From Latin reductus (distant).

In forewing costal area as broad as subcostal one, crossed with simple anterior branches of SC; RA simple; RS originated in the basal third of the wing, with three or four branches; M forked distinctly before the base of RS; CuA with two branches, fork of CuA is situated distinctly proximal to fork of M; the base of anterior branch of CuA sclerotized; CuP straight; several branches of anal veins reaching posterior wing margin. In hind wing RA simple, RS dichotomously branched, with four branches; M branched distal of the first fork of RS.

One species from the Lower Permian of North America.

Leonardian Stage, Wellington Formation (the Lower Permian).

Elmo locality, Kansas, USA.

Anterior margin of forewing weakly convex; SC reaching the apical one fifth of the wing; costal area as broad as intercubital one; RS distinctly separated from MA; MA simple or with two branches directed to the apex of wing; MP with three branches directed to the posterior margin of wing; CuA with two branches; cross-veins in almost all areas forming a double row of cells.

Length of forewing about 18 mm.

In forewing costal area narrow, as broad as subcostal one; anterior branches of SC absent; RA simple; RS originated in the basal third of the wing, simple; M forked distinctly basally to main fork of R; CuA with two branches, the fork of CuA is situated distinctly proximal to fork of M; the base of anterior branch of CuA desclerotized; CuP straight; A1 simple, A2 reduced.

One species from the Middle Permian of Europe.

Holotype only.

Urzhumian Stage (the Middle Permian).

Chepanikha locality, Rossokha River 1.8 km north of the Chepanikha village, Zavyalovsk District, Udmurtia, Russia.

Anterior margin of forewing straight; SC reaching the apical quarter of wing; costal area considerably narrower than intercubital one; RS distinctly separated from MA; MA forked; MP with two branches directed to the posterior margin of forewing; CuA with two branches; cross-veins in almost all areas forming a double row of cells, but in the areas between MP and CuA, between branches of CuA and in anal area, cross-veins forming numerous rows of cells.

Length of forewing about 18 mm.

Mezenalicula connata Aristov & Storozhenko, sp. n.

Similar to Neprotembia by the base of M fused with CuA, but distinct from it in the more proximal first fork of CuA, and anterior branches of SC not forming double row of cells.

The base of costal area in forewing broadly rounded; costal area broader than subcostal one, crossed with simple or furcated anterior branches of SC; RA simple; RS originated in the basal third or near the mid wing, with four or fewer branches; the base of M completely fused with CuA; M forked before the base of RS; CuA with two or three branches, first fork of CuA is situated distinctly proximal to fork of M; the base of anterior branch of CuA sclerotized; CuP straight; A1 and A2 simple.

Two species from the Middle and Upper Permian of Europe.

After the name of the Mezen District in the Arkhangelsk Region of Russia and genus Alicula; gender feminine.

Holotype PIN 94/883, positive imprint of incomplete forewing.

Kazanian Stage, Lower Kazanian Substage, Iva-Gora Beds (the Middle Permian).

Soyana locality, right bank of the Soyana River 56–60 km upstream of the mouth, Mezen District, Arkhangelsk Region, Russia.

Anterior margin of forewing convex in basal portion of wing; SC reaching the apical third of wing; costal area considerably broader than intercubital one; RS distinctly separated from MA; MA simple; MP with three branches directed to the posterior margin of forewing; CuA with three branches; cross-veins forming numerous rows of cells in all areas.

Length of forewing about 21 mm.

From Latin connatus (connate).

Holotype PIN 3840/1243, positive imprint of incomplete forewing.

Severodvinian Stage, Upper Severodvinian Substage, Poldars Formation, Kichuga Bundle (the Upper Permian).

Isady locality, left bank of the Sukhona River 1.8 km downstream of the Mutovino near Isady village, Veliky Ustyug District, Vologda Region, Russia.

Anterior margin of forewing convex in the mid wing; SC reaching the apical third of wing; costal area as broad as the intercubital one; RS fused with MA at short distance near the base; MA and MP simple; CuA with two branches; cross-veins in all areas simple.

Length of forewing about 20 mm.

From Latin conjunctus (connected).

The base of costal area in forewing broadly rounded; costal area about two times broader than subcostal one; the anterior branches of SC form double row of cells near the mid of costal area; RA simple; RS originated in the basal third of the wing, with two branches; the base of M fused with CuA at certain distance; M forked before the base of RS; CuA with three branches, first fork of CuA is situated near to fork of M; the base of anterior branch of CuA sclerotized; CuP straight; A1 and A2 simple.

Two species from the Lower Permian of Europe.

Holotype only.

Kungurian Stage, Koshelevka Formation (the Lower Permian).

Tshekarda locality, Krasnaya Gora Mountain on the left bank of Sylva River near the Tshekarda village, Suksun District, Perm Region, Russia.

Anterior margin of forewing convex; SC reaching the apical quarter of wing; costal area as broad as intercubital area; RS distinctly separated from MA; MA and MP simple; CuA probably with three branches; cross-veins are simple in the subcostal area and forming double row of cells in the areas between RA and RS, and between CuA and CuP.

Length of forewing about 15 mm.

Holotype only.

Kungurian Stage, Lek-Vorkuta Formation, Rudnik Subformation (the Lower Permian).

Khalmer-Yu coalfield, borehole KhK-371, deep 88 m, Pechora Basin, Perm Region, Russia.

Anterior margin of forewing weakly convex; SC almost reaching the wing tip; costal area as broad as intercubital one; RS distinctly separated from MA; MA and possibly MP are simple; CuA with three branches; cross-veins simple and forming double row of cells in the apical portion of the areas between RA, RS, MA, MP and CuA (in the subcostal area cross-veins absent).

Length of forewing 13 mm.

Order Grylloblattida Walker, 1914

Suborder Grylloblattina Walker, 1914

Family Ideliidae Zalessky, 1928

Permula acra Kukalová, 1964, Lower Permian of Czech Republic.

By shape of forewing CuA, the new genus undoubtedly belongs to family Ideliidae. It is easily distinguished from all known genera of Ideliidae in excised base of costal area.

Anterior margin of forewing strongly convex; costal area broad, crossed with mostly furcated anterior branches of SC, the base of costal area distinctly excised; subcostal area very narrow; RA simple; radial area broad, with numerous slightly backwardly slanted cross-veins; RS with several branches; M dividing before the origin of RS; CuA concavely bent, with a long pectinate series of branches. CuA2 not individualized; intercubital area with single intercalary branch of CuA; cross-veins forming numerous rows of cells in almost all areas.

A type species only.

After the species name acra and genus Permula; feminine in gender.

The description of a new genus is based on illustrations and diagnosis of Permula acra (Kukalová 1964).

Sakmarian Stage, Boskovice Graben, Letovice Formation (the Lower Permian).

Obora locality, 1 km north-west from the Obora village near Boskovice, Moravia, Czech Republic.

Metidelia Martynov, 1937

Metidelia asiatica (Storozhenko, 1997)

Notes. This species was described in the genus Alicula (Srorozhenko 1997). After examination of holotype it was transferred to the genus Metidelia of family Ideliidae by Aristov (2004b).

Order Eoblattida Hanlirsch, 1906

Incertae familiae

Neraphidia Novokshonov & Novokshonova, 1997

Notes. The genus Neraphidia was described in the grylloblattid family Protembiidae (Novokshonov and Novokshonova 1997), but later it was placed in family Aliculidae (Aristov 2004a). The examination of holotype of Neraphidia mitis Novokshonov & Novokshonova, 1997 (Fig. 14) allows us to eliminate this taxon from order Grylloblattida. Neraphidia is characterized by SC terminating on RA, and by M5 stronger than the main stems of M and CuA, both characters are typical for the order Eoblattida (in order Grylloblattida SC terminating on C or disappearing in the area between C and RA; M5 usually not individualized, if present in some families, when M5 is as broad as, or narrower than stems of M and CuA). Undoubtedly Neraphidia is similar to genera of the family Tillyardembiidae in the shape of SC and CuA, but distinguished from the latter by most basal fork of M (in Tillyardembiidae first fork of M is situated distinctly proximal). Herein Neraphidia is transferred to the order Eoblattida (sensu Aristov and Rasnitsyn 2009) as a genus of uncertain taxonomic position inside this order.