Review of the Permian family Permulidae nomen novum pro Aliculidae Storozhenko, 1997 (Grylloblattida)

Abstract A new replacement name Permulidae nom. n. is proposed for the Permian family Aliculidae Storozhenko, 1997 (Insecta: Grylloblattida). A review of Permulidae is given. All genera and species are redescribed and illustrated. A key to genera of Permulidae is given. New taxa of Permulidae are described: Permula edemskii sp. n., Sojanopermula rasnitsyni sp. n., Kazanalicula reducta gen. et sp. n., Mezenalicula connata gen. et sp. n., all from the Soyana locality (Middle Permian, Kazanian Stage; Arkhangelsk Region, Russia), and Mezenalicula conjuncta sp. n. from the Isady locality (Upper Permian, Severodvinian Stage; Vologda Region, Russia). New combinations are proposed: Permula minor (Aristov, 2004), comb. n. for Sojanopermula minor Aristov, 2004; Permula tshekardensis (Aristov, 2004), comb. n. for Sojanopermula tshekardensis Aristov, 2004. A new genus Acropermula gen. n. (type species: Permula acra Kukalová, 1964, from the Lower Permian of Czech Republic) is established in the family Ideliidae. Neraphidia Novokshonov & Novokshonova, 1997 is transferred from Aliculidae to the order Eoblattida as a genus of uncertain taxonomic position inside this order.


introduction
Family Aliculidae was established for two genera of the Permian Grylloblattida characterized by distinct narrowing of the costal area in the forewing base and by CuA not dividing into CuA1 and CuA2 (Storozhenko 1997). Originally it included the Lower Permian Alicula lebachensis Schlechtendal, 1913 from the Saar-Nahe Basin in Germany, A. acra (Kukalová 1964) from the Obora locality in Czech Republic, the Upper Permian A. asiatica Storozhenko, 1997 from the Karaungir-II locality in Kazakhstan, and the Middle Permian Sojanopermula lucida Storozhenko, 1992 from the Soyana locality in Russia. Aristov (2004a) described in Aliculidae two new species (Sojanopermula minor Aristov, 2004, S. tshekardensis Aristov, 2004 and one monotypic genus (Neprotembia truncata Aristov, 2004) from the Lower Permian Tshekarda locality in Russia, and transferred to family Aliculidae the genus Neraphidia Novokshonov & Novokshonova, 1997, originally described in the family Protembiidae (Novokshonov and Novokshonova 1997). At the same time he re-examined the holotype of Alicula asiatica and placed this species in the genus Metidelia Martynov, 1937 of the family Ideliidae (Aristov 2004b). Neprotembia complicata Aristov, 2005 was described from the Lower Permian Vorkuta locality in Russia (Rasnitsyn et al, 2005). The genus Elmopterum with a single species Elmopterum rotundum Béthoux & Beckemeyer, 2007 from the Lower Permian Elmo locality in the USA, originally described as Grylloblattida incertae sedis (Béthoux and Beckemeyer 2007), was placed in Aliculidae by Aristov (2009). Finally Tshepanichoptera lacera Aristov, 2008 was described from the Middle Permian Chepanikha locality in Russia (Aristov and Bashkuev 2008). Thus, the composition and conception of family Aliculidae are changed considerably during last ten years and a critical examination of this family is necessary. Moreover, the family name Aliculidae Storozhenko, 1997 was based on Alicula Schlechtendal, 1913, a junior homonym of Alicula Eichwald, 1830 in Mollusca. So, this generic name must be replaced by the synonymic name Permula Handlirsch, 1919, and the family name Aliculidae by Permulidae nom. n.
A review of the family Permulidae nom. n. is given below. Two new genera and five new species are described. The keys to genera and to species of each genus are given based on forewing characters. In addition the taxa previously erroneously included in Permulidae are listed, and a new genus of family Ideliidae is established.

Material
All studied material including the holotypes of new species is deposited in the Borissiak Paleontological Institute of the Russian Academy of Sciences, Moscow (PIN).
The vein symbols are as follows: C costa, SC subcosta, R stem of radius, RA radius anterior, RS radius sector (= radius posterior), M stem of media, MA media anterior, MP media posterior, M5 strong oblique vein between stems of media and cubitus, Cu stem of cubitus, CuA cubitus anterior, CuA1 first cubitus anterior (if CuA distinctly 1(10) The basal portion of M distinctly separated from CuA.

3(4)
The Description. 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. Composition. Four species from the Lower and Middle Permian of Europe. Notes. 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 andincluded 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). Horizon. Asselian Stage, Lebacher Schichten (the Lower Permian). Locality. Lebach locality, Saar-Nahe Basin, Saarland, Germany. Description. 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, crossveins forming rows of three cells in the basal portion of intercubital area.
Measurements Description. 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.
Measurements. Length of forewing 30 mm. Locality. Soyana locality, right bank of the Soyana River 56-60 km upstream of the mouth, Mezen District, Arkhangelsk Region, Russia.
Description. 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.
Measurements. Length of forewing about 34.6 mm. Description. 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.

Sojanopermula rasnitsyni
Measurements Diagnosis. Similar to Elmopterum and Tshepanichoptera by the narrow costal area, but distinct from these in the simple CuA.
Description. 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.
Composition. One species from the Middle Permian of Europe. Etymology. After the name of the Kazanian Stage and genus Alicula; gender feminine.

Horizon. Leonardian Stage, Wellington Formation (the Lower Permian).
Locality. Elmo locality, Kansas, USA. Description. 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; crossveins in almost all areas forming a double row of cells.
Measurements. Length of forewing about 18 mm. Description. 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. Composition. One species from the Middle Permian of Europe.
Description. 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.
Measurements. Length of forewing about 18 mm. Description. 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.

Mezenalicula
Composition. Two species from the Middle and Upper Permian of Europe. Etymology. After the name of the Mezen District in the Arkhangelsk Region of Russia and genus Alicula; gender feminine.  Kukalová, 1964, Lower Permian of Czech Republic. Diagnosis. 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.
Description. 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.
Composition. A type species only. Ethymology. After the species name acra and genus Permula; feminine in gender. Notes. The description of a new genus is based on illustrations and diagnosis of Permula acra (Kukalová 1964).