(C) 2011 Daniil S. Aristov. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
For reference, use of the paginated PDF or printed version of this article is recommended.
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.
Grylloblattida, Permulidae, Aliculidae, Ideliidae, Eoblattida, taxonomy, key, new taxa, Permian, Europe, North America
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, Alicula acra (Kukalová 1964) from the Obora locality in Czech Republic, the Upper Permian Alicula 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, Sojanopermula 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.
MaterialAll 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 forked into branched first and simple second veins), CuA2 second cubitus anterior, CuP cubitus posterior, A1 first anal, A2 second anal vein.
TaxonomyOrder Grylloblattida Walker, 1914
Suborder Protoperlina Brongniart, 1885
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).
| 1(10) | The basal portion of M distinctly separated from CuA. | |
| 2(5) | Costal area distinctly broader than subcostal one. | |
| 3(4) | The basal portion of costal area broadly rounded, without an abrupt narrowing | Permula |
| 4(3) | Costal area distinctly narrowing in its basal portion | Sojanopermula |
| 5(2) | Costal area narrower than or as broad as subcostal one. | |
| 6(7) | CuA simple | Kazanalicula gen. n. |
| 7(6) | CuA with two branches. | |
| 8(9) | The base of anterior branch of CuA distinctly sclerotized | Elmopterum |
| 9(8) | The base of anterior branch of CuA desclerotized | Tshepanichoptera |
| 10(1) | The basal portion of M fused with CuA completely or at certain distance. | |
| 11(12) | First fork of CuA situated distinctly proximal to main fork of M. Costal area with a row of simple or bifurcated anterior branches of SC | Mezenalicula gen. n. |
| 12(11) | First fork of CuA situated near to main fork of M. Anterior branches of SC form double row of cells near the middle of costal area | Neprotembia |
| 1(10) | The basal portion of M distinctly separated from CuA. | |
| 2(5) | Costal area distinctly broader than subcostal one. | |
| 3(4) | The basal portion of costal area broadly rounded, without an abrupt narrowing | Permula |
| 4(3) | Costal area distinctly narrowing in its basal portion | Sojanopermula |
| 5(2) | Costal area narrower than or as broad as subcostal one. | |
| 6(7) | CuA simple | Kazanalicula gen. n. |
| 7(6) | CuA with two branches. | |
| 8(9) | The base of anterior branch of CuA distinctly sclerotized | Elmopterum |
| 9(8) | The base of anterior branch of CuA desclerotized | Tshepanichoptera |
| 10(1) | The basal portion of M fused with CuA completely or at certain distance. | |
| 11(12) | First fork of CuA situated distinctly proximal to main fork of M. Costal area with a row of simple or bifurcated anterior branches of SC | Mezenalicula gen. n. |
| 12(11) | First fork of CuA situated near to main fork of M. Anterior branches of SC form double row of cells near the middle of costal area | Neprotembia |
http://species-id.net/wiki/Permula
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).
| 1(4) | CuA with three or four branches. | |
| 2(3) | Anterior margin of wing straight; CuA with four branches | Permula lebachensis |
| 3(2) | Anterior margin of wing convex; CuA with three branches | Permula tshekardensis comb. n. |
| 4(1) | CuA with two branches. | |
| 5(6) | Cross-veins simple or forming a double row of cells in the areas between RA and RS, between MP and CuA, and between CuA and CuP | Permula minor comb. n. |
| 6(5) | Cross-veins forming numerous rows of cells in all areas | Permula edemskii sp. n. |
| 1(4) | CuA with three or four branches. | |
| 2(3) | Anterior margin of wing straight; CuA with four branches | Permula lebachensis |
| 3(2) | Anterior margin of wing convex; CuA with three branches | Permula tshekardensis comb. n. |
| 4(1) | CuA with two branches. | |
| 5(6) | Cross-veins simple or forming a double row of cells in the areas between RA and RS, between MP and CuA, and between CuA and CuP | Permula minor comb. n. |
| 6(5) | Cross-veins forming numerous rows of cells in all areas | Permula edemskii sp. n. |
http://species-id.net/wiki/Permula_lebachensis
Fig. 1Asselian 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.
Forewings of Permulidae 1 Permula lebachensis (Schlechtendal) (after Handlirsch 1906–1908) 2 Permula tshekardensis Aristov (holotype PIN 1700/1151, after Aristov 2004) 3 Permula minor Aristov (holotype PIN 1700/4934, after Aristov 2004) 4 Permula edemskii sp. n. (holotype PIN 94/1027, orig.) 5 Sojanopermula lucida Storozhenko (holotype PIN 94/117, after Storozhenko 1992) 6 Sojanopermula rasnitsyni sp. n. (holotype PIN 117/301, orig.) 7 Sojanopermula rasnitsyni sp. n. (paratype PIN 117/195, orig.). Scale bars 2 mm.
Forewings of Permulidae 1 Permula lebachensis (Schlechtendal) (after Handlirsch 1906–1908) 2 Permula tshekardensis Aristov (holotype PIN 1700/1151, after Aristov 2004) 3 Permula minor Aristov (holotype PIN 1700/4934, after Aristov 2004) 4 Permula edemskii sp. n. (holotype PIN 94/1027, orig.) 5 Sojanopermula lucida Storozhenko (holotype PIN 94/117, after Storozhenko 1992) 6 Sojanopermula rasnitsyni sp. n. (holotype PIN 117/301, orig.) 7 Sojanopermula rasnitsyni sp. n. (paratype PIN 117/195, orig.). Scale bars 2 mm.
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.
http://species-id.net/wiki/Permula_tshekardensis
Fig. 2Holotype 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.
http://species-id.net/wiki/Permula_minor
Fig. 3Holotype 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.
urn:lsid:zoobank.org:act:7728E3EC-1FCE-409B-BF16-1C6A84D372AE
http://species-id.net/wiki/Permula_edemskii
Figs 4, 15Holotype 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.
http://species-id.net/wiki/Sojanopermula
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.
| 1(2) | CuA with three branches. Costal area as broad as intercubital area | Sojanopermula lucida |
| 2(1) | CuA with four branches. Costal area distinctly broader than intercubital one | Sojanopermula rasnitsyni sp. n. |
| 1(2) | CuA with three branches. Costal area as broad as intercubital area | Sojanopermula lucida |
| 2(1) | CuA with four branches. Costal area distinctly broader than intercubital one | Sojanopermula rasnitsyni sp. n. |
http://species-id.net/wiki/Sojanopermula_lucida
Fig. 5Holotype 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.
Forewings of Permulidae and Eoblattida incertae sedis 8 Kazanalicula reducta sp. n. (holotype PIN 117/467, orig.) 9 Tshepanichoptera lacera Aristov (holotype PIN 3286/14, after Aristov and Bashkuev 2008) 10 Mezenalicula connata sp. n. (holotype PIN 94/883, orig.) 11 Mezenalicula conjuncta sp. n. (holotype PIN 3840/1243, orig.) 12 Neprotembia truncata Aristov (holotype PIN 1700/1026, after Aristov 2004) 13 Neprotembia complicata Aristov (holotype PIN 1631/314, after Rasnitsyn et al. 2005) 14 Neraphidia mitis Novokshonov & Novokshonova (orig. reconstruction based on holotype PIN 1700/643 and spec. PIN 4987/10). Scale bars 2 mm.
Forewings of Permulidae and Eoblattida incertae sedis 8 Kazanalicula reducta sp. n. (holotype PIN 117/467, orig.) 9 Tshepanichoptera lacera Aristov (holotype PIN 3286/14, after Aristov and Bashkuev 2008) 10 Mezenalicula connata sp. n. (holotype PIN 94/883, orig.) 11 Mezenalicula conjuncta sp. n. (holotype PIN 3840/1243, orig.) 12 Neprotembia truncata Aristov (holotype PIN 1700/1026, after Aristov 2004) 13 Neprotembia complicata Aristov (holotype PIN 1631/314, after Rasnitsyn et al. 2005) 14 Neraphidia mitis Novokshonov & Novokshonova (orig. reconstruction based on holotype PIN 1700/643 and spec. PIN 4987/10). Scale bars 2 mm.
Length of forewing about 34.6 mm.
urn:lsid:zoobank.org:act:7D76CEE0-2646-45F2-B738-072281C9CD4E
http://species-id.net/wiki/Sojanopermula_rasnitsyni
Figs 6, 7, 16, 17Holotype 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.
Forewings of Permulidae 15 Permula edemskii sp. n. (holotype PIN 94/1027) 16 Sojanopermula rasnitsyni sp. n. (holotype PIN 117/301) 17 Sojanopermula rasnitsyni sp. n. (paratype PIN 117/195) 18 Kazanalicula reducta sp. n. (holotype PIN 117/467) 19 Mezenalicula connata sp. n. (holotype PIN 94/883) 20 Mezenalicula conjuncta sp. n. (holotype PIN 3840/1243). Scale bars 2 mm.
Forewings of Permulidae 15 Permula edemskii sp. n. (holotype PIN 94/1027) 16 Sojanopermula rasnitsyni sp. n. (holotype PIN 117/301) 17 Sojanopermula rasnitsyni sp. n. (paratype PIN 117/195) 18 Kazanalicula reducta sp. n. (holotype PIN 117/467) 19 Mezenalicula connata sp. n. (holotype PIN 94/883) 20 Mezenalicula conjuncta sp. n. (holotype PIN 3840/1243). Scale bars 2 mm.
Length of forewing about 30 mm.
The new species is named in honor of the Russian paleoentomologist Prof. A.P. Rasnitsyn.
urn:lsid:zoobank.org:act:2444F483-EB8B-4A2A-AD84-EA98E5B6E872
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.
urn:lsid:zoobank.org:act:5225F245-8B20-4FA3-93EE-5F39F7A019AB
http://species-id.net/wiki/Kazanalicula_reducta
Figs 8, 18Holotype 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).
http://species-id.net/wiki/Elmopterum
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.
http://species-id.net/wiki/Elmopterum_rotundum
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.
http://species-id.net/wiki/Tshepanichoptera
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.
http://species-id.net/wiki/Tshepanichoptera_lacera
Fig. 9Holotype 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.
urn:lsid:zoobank.org:act:B823B199-72D0-499E-A595-68097693F7E4
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.
| 1(2) | Cross-veins forming a double row or numerous rows of cells. RS distinctly separated from MA | Mezenalicula connata sp. n. |
| 2(1) | Cross-veins simple. RS fused with MA at short distance near the base | Mezenalicula conjuncta sp. n. |
| 1(2) | Cross-veins forming a double row or numerous rows of cells. RS distinctly separated from MA | Mezenalicula connata sp. n. |
| 2(1) | Cross-veins simple. RS fused with MA at short distance near the base | Mezenalicula conjuncta sp. n. |
urn:lsid:zoobank.org:act:71A86E11-B0EC-4E75-AD48-255EF52A6030
http://species-id.net/wiki/Mezenalicula_connata
Figs 10, 19Holotype 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).
urn:lsid:zoobank.org:act:4549C5ED-EB14-4526-A1F3-DA4DAB596428
http://species-id.net/wiki/Mezenalicula_conjuncta
Figs 11, 20Holotype 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).
http://species-id.net/wiki/Neprotembia
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.
| 1(2) | Anterior margin of forewing distinctly convex; the basal part of wing broader | Neprotembia truncata |
| 2(1) | Anterior margin of forewing gently convex; the basal part of wing narrower | Neprotembia complicata |
| 1(2) | Anterior margin of forewing distinctly convex; the basal part of wing broader | Neprotembia truncata |
| 2(1) | Anterior margin of forewing gently convex; the basal part of wing narrower | Neprotembia complicata |
http://species-id.net/wiki/Neprotembia_truncata
Fig. 12Holotype 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.
http://species-id.net/wiki/Neprotembia_complicata
Fig. 13Holotype 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.
Taxa erroneously included in PermulidaeOrder Grylloblattida Walker, 1914
Suborder Grylloblattina Walker, 1914
Family Ideliidae Zalessky, 1928
urn:lsid:zoobank.org:act:CA6C68E9-C9E6-4DC2-8A44-FF87AE4C48D2
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.
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.
The family Permulidae consists of thirteen species in seven genera. The oldset Lower Permian representative of the family, Permula lebachensis, is known from the Asselian of Central Europe. In the Leonardian Elmo locality (North America) Permulidae is represented by Elmopterum rotundum only. The Kungurian Chekarda locality (East Europe) is characterized by very rich fauna of Grylloblattida, but only three species of the family are described from here (Permula tshekardensis, Permula minor and Neprotembia truncata). Another, probably younger Kungurian representative of Permulidae, Neprotembia complicata, is known from the Pechora Basin (East Europe). Five Middle Permian species (Permula edemskii sp. n., Sojanopermula lucida, Sojanopermula rasnitsyni sp. n., Kazanalicula reducta gen. et sp. n. and Mezenalicula connata gen. et sp. n.) are described from the Kazanian Soyana locality in East Europe. A later Middle Permian species, Tshepanichoptera lacera, is known from Urzhumian Chepanikha locality (East Europe). The only Upper Permian representative of family, Mezenalicula conjuncta sp. n., is described from Severodvinian Isady locality in East Europe. Almost all species are known solely by their holotypes, and only Sojanopermula rasnitsyni sp. n. is known by two specimens. Thus, Permulidae was a widespread but not abundant and diverse Permian family of the order Grylloblattida.
The authors are grateful to Drs A.V. Gorochov (Zoological Institute, Russian Academy of Sciences, St Petersburg, Russia) and P. Vršanský (Geological Institute, Slovak Academy of Sciences, Bratislava, Slovakia) for important comments, and to Dr R. Beckemeyer (University of Kansas, Lawrence, Kansas) for checking the English of the MS. This study is supported by the Program ‘Biosphere Origin and Evolution of Geo-Biological Systems’ of the Presidium of the Russian Academy of Sciences and the Russian Foundation for Basic Research projects 09-04-01241, 10-04-01713, and 10-04-10032.