ISSN 0031�0301, Paleontological Journal, 2011, Vol. 45, No. 2, pp. 159–166. © Pleiades Publishing, Ltd., 2011Original Russian Text © D.S. Aristov, L. Grauvogel�Stamm, F. Marchal�Papier, 2011, published in Paleontologicheskii Zhurnal, 2011, No. 2, pp. 39–45.

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INTRODUCTION

The earliest grylloblattid insects are known fromthe beginning of the Late Carboniferous; by the end ofthe Early Permian and the beginning of the MiddlePermian they reach their heyday (Aristov, 2005b).At the end of the Middle Permian and in the Late Per�mian the diversity of grylloblattids decreases. Four ofthe 11 Late Permian families (Probnidae, Tshekardo�minidae, Chaulioditidae, Geinitziidae, Liomop�teridae, Ideliidae, Permotermopsidae, Megakhosa�ridae, Blattogryllidae, Tunguskapteridae, andSylvaphlebiidae) become extinct during the Late Per�mian: Liomopteridae, Tshekardominidae, Permoter�mopsidae, and Sylvaphlebiidae (Aristov, 2009). At thePermian–Triassic boundary, most of the Upper Per�mian families disappear from the Lower Triassicdeposits. In the largest Induan grylloblattid localityBabii Kamen’ (Kemerovo Region, Russia), the orderis represented by two families, which make up 20% ofall insect fossils. The dominant family is Chaulioditi�dae; Geinitziidae are scarce (Aristov, 2003, 2004;Shcherbakov, 2008). Scarce Chaulioditidae are knownfrom Induan deposits of Russia (Nedubrovo inVologda Region, Anakit and Tura in KrasnoyarskRegion; Aristov, 2003) and Mongolia (Yamaan Uslocality, Gobi�Altai Province; Aristov, 2005). The sin�gle specimen from Entala locality in Vologda Regionprobably belongs to Chaulioditidae (Aristov, 2005a).From the Olenekian deposits of Russia (Tikhvinskoelocality in Yaroslavl Region) only Chaulioditidae areknown, accounting for up to 50% of all insects (Aris�tov, 2003). From the Middle Buntsandstein of Ger�many (Olenekian Stage of Saxony�Anhalt), onlyscarce Chaulioditidae from the locality of Gödevitz

are known (Ansorge and Braucmann, 2008; Aristovet al., 2009). Numerous undescribed Chaulioditidaewere found in the locality of Gambach in Lower Fran�conia (Upper Buntsandstein, lower layers of the Ani�sian Stage of Germany) (Ansorge and Braucmann,2008). Undescribed members of Chaulioditidae(=Tomiidae; Aristov, 2004) and Blattogryllidae fromthe Lower Anisian Grès à Voltzia localities in Alsace,making up about 2% of all insects, have been men�tioned in the literature (Marchal�Papier, 1998; Aristov,2003, 2004; Gall et al., 2006). In this study, membersof four grylloblattid families (Chaulioditidae, Mesor�thopteridae, Megakhosaridae, and Blattogryllidae)are described from these deposits. A single record ofBlattogryllidae is known from the Buntsandstein ofSpain (Induan–Anisian Stage) on the Islandof Majorca, locality Port de Estellencs (Aristov andZessin, 2009). In the Middle Triassic locality of Nan�shenghu in China (Guizhou Province) scarceChaulioditidae were found (Lin, 1978; Aristov, 2003).The Ladinian fauna of Madygen locality in Kyrgyzstan(Osh Region) includes members of the dominantMesorthopteridae and the less abundant Megakhosa�ridae, Blattogryllidae, Geinitziidae, Ideliidae, Tun�guskapteridae, Gorokhoviidae, and Madygenophlebi�idae (Storozhenko, 1998). The family Mesojabloni�idae, described from this locality as a grylloblattid, is amember of the order Homoptera (D.E. Shcherbakov,pers. comm.). Grylloblattids make up about 4% of allinsects collected in this locality. Taking into accountthe fact that some insect groups (beetles, cockroaches)were selectively collected in Madygen, the actual per�centage of grylloblattids is considerably lower andunequal at different levels of the locality (Shcherbakov,

New Grylloblattid Insects (Insecta: Grylloblattida) from the Grès à Voltzia of the Vosges (Middle Triassic of France)

D. S. Aristova, L. Grauvogel�Stamm, and F. Marchal�PapieraBorissiak Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya ul. 123, Moscow, 117997 Russia

e�mail: danil_aristov@mail.ru; lea.stamm@gmail.comReceived May 20, 2009

Abstract—New grylloblattid insects (Grylloblattida) are described from the Grès à Voltzia of the Vosges (Ani�sian of Alsace and Lorraine, France): Chauliodites anisicus, sp. nov. (Chaulioditidae), Palaeomesorthopteron pul�lus gen. et sp. nov. (Mesorthopteridae), Megakhosarodes vosgesicus sp. nov. (Megakhosaridae), Dorniella elca�noides sp. nov., D. apectinata sp. nov., D. diluta sp. nov., D. ovalis sp. nov., Baharellinus umbrosus sp. nov. Embig�ryllus shcherbakovi gen. et sp. nov., and Vosgesopterum arzvillerensis gen. et sp. nov. (Blattogryllidae).

Keywords: Insecta, Grylloblattida, Anisian, France, Bas�Rhin, Moselle, new taxa.

DOI: 10.1134/S0031030111020043

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2008). A Grylloblattida incertae sedis was describedfrom the locality of Cacheuta in Argentina (MendozaProvince), which has the same age as Madygen (Aris�tov et al., 2006; Spalletti et al., 2008).

In localities of the Vosges, representatives of fourfamilies are known. Members of the family Blattogryl�lidae, which make up 72% of grylloblattid fossils, aredominant. This family is represented here by the fol�lowing genera: Dorniella, characteristic of the Middleor Upper Triassic of Kyrgyzstan and Jurassic of Ger�many, including four species: D. elcanoides sp. nov.,D. apectinata sp. nov., D. diluta sp. nov., and D. ovalissp. nov.; by Baharellinus, characteristic of the MiddleTriassic of Kyrgyzstan, including one species:B. umbrosus sp. nov.; and by two monotypic endemicgenera: Embigryllus (E. shcherbakovi gen. et sp. nov.)and Vosgesopterum (V. arzvillerensis gen. et sp. nov.).The family Chaulioditidae, which makes up 20% ofgrylloblattid fossils, is represented here by the genusChauliodites, characteristic of the Upper Permian ofRussia, Lower Triassic of Germany, Russia, and Mon�golia, and Triassic of China, including one species:C. anisicus sp. nov. Members of each of the familiesMesorthopteridae and Megakhosaridae make up 4%of grylloblattids fossils. Mesorthopteridae are repre�sented by the endemic genus Palaeomesorthopterongen. nov., including one species: P. pullus sp. nov. Thefamily Megakhosaridae is represented by one genus,Megakhosarodes, characteristic of the Upper Permianof Russia and Middle Triassic of Kyrgyzstan, includingone species: M. vosgesicus sp. nov.

Thus, the grylloblattid fauna of the Vosges, in spiteof the presence of Chaulioditidae, typical of the UpperPermian and Lower Triassic, more closely resemblesthe fauna of Madygen than Lower Triassic faunas,because of its high diversity and the small proportionof grylloblattids among insects. The composition ofthe Vosges faunas is similar to that of Madygen in thepresence of Mesorthopteridae, Blattogryllidae, andMegakhosaridae and the genera of the two latter fam�ilies known from the Middle or Upper Triassic. Thedifferences between the Vosges and Madygen faunasare in the smaller proportion of grylloblattids amonginsects (1 and 4%, respectively), dominance of Blat�togryllidae, which in Madygen make up about 20%,and presence of Chaulioditidae. In addition, the fam�ilies Geinitziidae, Ideliidae, Gorokhoviidae, andMadygenophlebiidae, known from Madygen, havenot been found in the Vosges. The small proportion ofgrylloblattids among insects in combination with thedominance of Blattogryllidae is only characteristic ofthe much younger locality of Karatau (Upper Jurassicof Kazakhstan).

To date, Grylloblattids are known from five Grès àVoltzia localities of the Vosges: Arzviller (Moselle),Bust (Bas�Rhin), Gottenhouse (Bas�Rhin), Adam�swiller (Bas�Rhin), and Vilsberg (Moselle). From Arz�

viller locality, the families Megakhosaridae (M. vosge�sicus sp. nov.) and Blattogryllidae (D. elcanoidessp. nov., D. apectinata sp. nov., D. diluta sp. nov.,B. umbrosus sp. nov., E. shcherbakovi gen. et sp. nov.,and V. arzvillerensis gen. et sp. nov.) are known. FromBust locality, the families Chaulioditidae (C. anisicussp. nov.) and Blattogryllidae (D. elcanoides sp. nov.,D. apectinata sp. nov., and B. umbrosus sp. nov.) areknown. From Gottenhouse locality, the familiesChaulioditidae (C. anisicus sp. nov.), Mesorthop�teridae (P. pullus gen. et sp. nov.), and Blattogryllidae(D. elcanoides sp. nov. and D. ovalis sp. nov.) areknown. The localities Adamswiller and Vilsberg haveyielded only Blattogryllidae (D. elcanoides sp. nov.)and Chaulioditidae (C. anisicus sp. nov.), respectively.The differences between the original faunas of theselocalities are apparently random.

MATERIAL

The material described, including the types, isstored in the private coll. L. Grauvogel�Stamm(G.�S.) (Strasbourg, France).

SYSTEMATIC PALEONTOLOGY

Order Grylloblattida

Subrder Grylloblattina

Family Chaulioditidae Handlirsch, 1906

Genus Chauliodites Heer, 1864

Chauliodites anisicus Aristov, Grauvogel�Stamm et Marchal�Papier, sp. nov.

Plate 7, fig. 1

E t y m o l o g y. From the Anisian Stage.H o l o t y p e. G.�S. no. 5541/5542, part and coun�

terpart of forewing; Vilsberg locality; Middle Triassic,Anisian Stage.

D e s c r i p t i o n (Fig. 1a). The anterior margin ofthe wing is weakly convex; the costal field basally ismore than twice as wide as the subcostal field. SC issharply bent at the base, ending in the distal third ofthe wing, anterior branches of SC are simple andY�shaped. The base of RS is positioned in the basalthird of the wing; RS, MA, and MP are two�branched.The branches of RS and MA are S�shaped. CuAbefore its division into branches is simple, dividinginto simple CuA1 and CuA2 near the basal quarter ofthe wing; CuP is simple. The transverse veins are sim�ple; the coloration pattern consists of large and smallspots.

M e a s u r e m e n t s, mm. Forewing length, 10–12.5.

C o m p a r i s o n. The new species differs from theother species of the genus in the sharp basal bend of SCand the S�shaped branches of RS and MA. In theother species of the genus, SC is not sharply bent

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NEW GRYLLOBLATTID INSECTS (INSECTA: GRYLLOBLATTIDA) 161

basally, and the branches of RS and MA are notS�curved (Aristov, 2003, 2004, 2005, 2008, 2009).

M a t e r i a l. In addition to the holotype, paratypesG.�S. nos. 5832/5833, 5821/5822, and 5774/5775from Gottenhouse locality and no. 5551/5552 fromBust locality.

Family Mesorthopteridae Tillyard, 1916

Genus Palaeomesorthopteron Aristov, Grauvogel�Stamm et Marchal�Papier, gen. nov.

E t y m o l o g y. From the Greek palaios (ancient)and the generic name Mesorthopteron.

Ty p e s p e c i e s. P. pullus, sp. nov.D i a g n o s i s. Small insects. Costal field at level of

RS base slightly narrower than subcostal field. Interra�dial field narrower than radial. RS, MA, and MP sim�ple. Distal branches of CuA ending in distal third ofwing. Apices of RS, MA, and MP and distal branchesof CuA forming anastomoses along wing margin.

S p e c i e s c o m p o s i t i o n. Type species.C o m p a r i s o n. The new genus differs from all

known genera of the family Mesorthopteridae in thecombination of the narrow costal field, simple RS,MA, and MP with anastomoses at apices of RS, MA,MP, and of distal CuA branches. In the other genera of

(c)

R

SC

CuP

CuA MP

MA

RS

RS

MA

MPCuP CuPCuA

A

R

(e) (f)

(d)

(a) (b) SC

CuP CuP

CuPCuP

CuA

CuA

MP

MP

A

AA

A

R

RS + MA

(g)

RS + MA

(h)

MP

MP

MP

R

R

CuA

CuA

CuA

RS + MA

RS + MA

RS + MA

MP

MA

RS

CuA

SCR

R

Fig. 1. New Grylloblattida from the Grès à Voltzia of the Vosges, forewings: (a) Chauliodites anisicus sp. nov., holotype 5541/5542,Vilsberg locality; (b) Palaeomesorthopteron pullus gen. et sp. nov., holotype 5711/5712, Gottenhouse locality; (c) Megakhosarodesvosgesicus sp. nov., holotype 5844/5845, Arzviller locality; (d, e) Dorniella elcanoides sp. nov.: (d) holotype 5791/5792, Gottenhouselocality, (e) paratype no. 5829, Arzviller locality; (f) D. diluta sp. nov., holotype 9352, Arzviller locality; (g) D. apectinata sp. nov., holo�type 5539/5540, Arzviller locality; (h) D. ovalis sp. nov., holotype 5895, Gottenhouse locality. Scale bar, 1 mm.

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Plate 7

1 2

3 4

5 6

87

9 10

E x p l a n a t i o n o f P l a t e 7

Figs. 1–10. New Grylloblattida from the Grès à Voltzia of the Vosges, forewings: (1) Chauliodites anisicus sp. nov., holotype5541/5542, Wilsberg locality; (2) Palaeomesorthopteron pullus gen. et sp. nov., holotype 5711/5712, Gottenhouse locality;(3) Megakhosarodes vosgesicus sp. nov., holotype 5844/5845, Arzviller locality; (4) Dorniella elcanoides sp. nov., holotype5791/5792, Gottenhouse locality; (5) D. diluta sp. nov., holotype 9352, Arzviller locality; (6) D. apectinata sp. nov., holotype5539/5540, Arzviller locality; (7) D. ovalis sp. nov., holotype 5895, Gottenhouse locality; (8) Baharellinus umbrosus sp. nov., holo�type 5537/5538, Bust locality; (9) Vosgesopterum arzvillerensis gen. et sp. nov., holotype 5787/5788, Arzviller locality;(10) Embigryllus shcherbakovi gen. et sp. nov., holotype 6643, Arzviller locality. Scale bar, 1 mm.

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NEW GRYLLOBLATTID INSECTS (INSECTA: GRYLLOBLATTIDA) 163

the family, the costal field is wider than or equal inwidth to the subcostal field; RS, MA, and MP arebranching, without anastomoses apically (Storozhenko,1998).

Palaeomesorthopteron pullus Aristov, Grauvogel�Stamm et Marchal�Papier, sp. nov.

Plate 7, fig. 2

E t y m o l o g y. From the Latin pullus (dark).H o l o t y p e. G.�S. no. 5711/5712, part and coun�

terpart of forewing; Gottenhouse locality; Middle Tri�assic, Anisian Stage.

D e s c r i p t i o n (Fig. 1b). The anterior margin ofthe wing is straight; the wing apex is weakly pointed.RS begins in the basal third of the wing. CuA has fouror more branches. The transverse veins are simple; thewing membrane is dark.

M e a s u r e m e n t s, mm. Forewing length,about 10.

M a t e r i a l. Holotype.

Family Megakhosaridae Sharov, 1961

Genus Megakhosarodes Storozhenko, 1993Megakhosarodes vosgesicus Aristov, Grauvogel�Stamm

et Marchal�Papier, sp. nov.

Plate 7, fig. 3

E t y m o l o g y. From the Vosges.H o l o t y p e. G.�S. no. 5844/5845, part and coun�

terpart of forewing; Arzviller locality; Middle Triassic,Anisian Stage.

D e s c r i p t i o n (Fig. 1c). RS begins dividing nearthe middle of the wing to form five branches. MA andMP divide near the posterior margin of the wing, eachhas two branches. CuA1 has three branches; CuA2 issimple and straight, posterior branches of CuA areweakly curved. The cubital field and the field betweenА1 and А2 are narrow. The transverse veins are simple.

M e a s u r e m e n t s, mm. Forewing length,about 13.

C o m p a r i s o n. The new species is especiallyclose to M. zajsanicus Storozhenko, 1993 from theUpper Permian locality of Karaungir II in Kazakhstanand differs from it in the branching MP and narrowfields between CuA and CuP and between А1 and А2.In M. zajsanicus, MP is simple, and fields betweenCuA and CuP and between А1 and А2 are wide (Storo�zhenko, 1993).

M a t e r i a l. Holotype.

Family Blattogryllidae Rasnitsyn, 1976

Genus Dorniella Bode, 1953Dorniella elcanoides Aristov,

Grauvogel�Stamm et Marchal�Papier, sp. nov.

Plate 7, fig. 4

E t y m o l o g y. From the generic name Elcana.

H o l o t y p e. G.�S. no. 5791/5792, part and coun�terpart of forewing; Gottenhouse locality; Middle Tri�assic, Anisian Stage.

D e s c r i p t i o n (Figs. 1d, 1e). The anterior marginof the wing is weakly convex; the wing apex is pointed;the costal field is 1.5 times narrower than the subcostalfield. RS begins in the basal third of the wing, forms ananastomosis with MA; RS+MA is pectinate, with fouror five branches. M does not have a free base; MP issimple. CuA divides into CuA1 and CuA2 proximal of itsmiddle; CuA1 and CuA2 are simple or with short bifur�cations. Some of the posterior branches of CuA areS�shaped. The wing membrane is dark.

M e a s u r e m e n t s, mm. Forewing length, 8.3–10.C o m p a r i s o n. The new species is especially

similar to D. primitiva Storozhenko, 1992 from Mady�gen locality (Ladinian of Kyrgyzstan), from which itdiffers in the pectinate RS+MA and CuA dividingrather proximally. In D. primitiva, RS+MA is dichoto�mizing, and CuA divides distal of its middle (Storo�zhenko, 1992).

M a t e r i a l. In addition to the holotype, paratypesG.�S. nos. 5701/5702, 5739/5740, 5754, 5829, and9146 from Arzviller locality, specimens G.�S.nos. 6656/6657, 6656/6657, and 9350 from Adam�swiller locality, specimen G.�S. no. 5735/5736 fromBust locality and specimens G.�S. nos. 9351,5719/5720, and 5694/5695 from Gottenhouse locality.

Dorniella diluta Aristov, Grauvogel�Stamm et Marchal�Papier, sp. nov.

Plate 7, fig. 5

E t y m o l o g y. From the Latin diluta (light).H o l o t y p e. G.�S. no. 9352, part and counterpart

of forewing; Arzviller locality; Middle Triassic, Ani�sian Stage.

D e s c r i p t i o n (Fig. 1f). RS begins in the basalthird of the wing and forms an anastomosis with MA.RS+MA is pectinate, with four or more branches.M does not have a free base; MP is simple. CuAdivides into CuA1 and CuA2 proximal of its middle;CuA1 divides in its basal third and has two branches;CuA2 is simple. The posterior branches of CuA are notS�shaped. The wing membrane is light.

M e a s u r e m e n t s, mm. Forewing length,about 16.

C o m p a r i s o n. The new species differs from theespecially similar species D. elcanoides sp. nov. in theproximally branching CuA1.

M a t e r i a l. Holotype.

Dorniella apectinata Aristov, Grauvogel�Stamm et Marchal�Papier, sp. nov.

Plate 7, fig. 6

E t y m o l o g y. From the Latin apectinata (non�pectinate).

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ARISTOV et al.

H o l o t y p e. G.�S. no. 5539/5540, part and coun�terpart of forewing; Arzviller locality; Middle Triassic,Anisian Stage.

D e s c r i p t i o n (Fig. 1g). SC does not reachbeyond the middle of the wing. RS begins in the basalquarter of the wing, forms an anastomosis with MA;RS+MA is dichotomizing, with four branches. M hasa free base; MP is simple. CuA divides into CuA1 andCuA2 distal of its middle; CuA1 and CuA2 are simple.The posterior branches of CuA are straight. The color�ation pattern consists of a dark area in the distal half ofthe wing along the apex and the posterior margin ofthe wing.

M e a s u r e m e n t s, mm. Forewing length, 9–10.C o m p a r i s o n. The new species is especially

similar to D. primitiva Storozhenko, 1992 from Mady�gen locality (Ladinian of Kyrgyzstan) and differs fromit in the relatively proximal beginning of RS and thefree base of M. In D. primitiva, RS begins near thebasal third of the wing, and M does not have a free base(Storozhenko, 1992).

M a t e r i a l. In addition to the holotype, paratypesG.�S. nos. 6654/6655 from Arzviller locality and5737/5738 from Bust locality.

Dorniella ovalis Aristov, Grauvogel�Stamm et Marchal�Papier, sp. nov.

Plate 7, fig. 7

E t y m o l o g y. From the Latin ovalis (oval).H o l o t y p e. G.�S. no. 5895, part and counterpart

of forewing; Gottenhouse locality; Middle Triassic,Anisian Stage.

D e s c r i p t i o n (Fig. 1h). RS begins in the basalthird of the wing, forms an anastomosis with MA.RS+MA is not pectinate, has four branches; MP hastwo branches. CuA divides into CuA1 and CuA2 proxi�mal of its middle; CuA1 divides in its basal third andhas three branches; CuA2 is simple. The posteriorbranches of CuA are S�shaped. The wing membrane islightly colored.

M e a s u r e m e n t s, mm. Forewing length,about 7.5.

C o m p a r i s o n. The new species is especiallysimilar to D. diluta sp. nov., from which it differs in thetwo�branched MP and three�branched CuA1.

M a t e r i a l. Holotype.

Genus Baharellinus Storozhenko, 1992

Baharellinus umbrosus Aristov, Grauvogel�Stamm et Marchal�Papier, sp. nov.

Plate 7, fig. 8

E t y m o l o g y. From the Latin umbrosus (dark�ened).

H o l o t y p e. G.�S. no. 5537/5538, part and coun�terpart of forewing; Bust locality; Middle Triassic,Anisian Stage.

D e s c r i p t i o n (Figs. 2a, 2b). The wing apex isweakly pointed. RS begins in the basal quarter of thewing, does not form an anastomosis with MA, has fivebranches. M does not have a free base. MA and MPbranch off CuA in separate stems; MA has three�fivebranches; MP is simple. CuA divides into CuA1 andCuA2 proximal of the middle; CuA1 divides into two

(a)

CuACuP

CuP CuP

CuPCuA

CuA CuA

SC

MPMA

RS

A

AAMP

MP

R

SC R (b)

(c) (d)

RS + MA

MPMA

RS

RS + MA

R

Fig. 2. New Grylloblattida from the Grès à Voltzia of the Vosges, forewings: (a, b) Baharellinus umbrosus sp. nov.: (a) holo�type 5537/5538, Bust locality, (b) paratype 5854/5855, Arzviller locality; (c) Vosgesopterum arzvillerensis gen. et sp. nov.,holotype 5787/5788, Arzviller locality; (d) Embigryllus shcherbakovi gen. et sp. nov., holotype 6643, Arzviller locality. Scale bar,1 mm.

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NEW GRYLLOBLATTID INSECTS (INSECTA: GRYLLOBLATTIDA) 165

branches in itsbasal quarter; CuA2 is simple. The pos�terior branches of CuA are not S�shaped. The wingmembrane is dark.

M e a s u r e m e n t s, mm. Forewing length,about 16.

C o m p a r i s o n. The new species is especiallysimilar to B. dimidiatus Storozhenko, 1992 fromMadygen locality (Ladinian of Kyrgyzstan) and differsfrom it in the relatively proximal branching of CuA1. InB. dimidiatus, CuA1 divides near its basal third (Storo�zhenko, 1992).

M a t e r i a l. In addition to the holotype, paratypeG.�S. no. 5854/5855 from Arzviller locality.

Genus Vosgesopterum Aristov, Grauvogel�Stamm et Marchal�Papier, gen. nov.

E t y m o l o g y. From the Vosges and the Greekpteron (wing).

Ty p e s p e c i e s. V. arzvillerensis sp. nov.

D i a g n o s i s. Small insects. Anterior margin ofwing convex; wing apex weakly pointed. RS beginningin basal quarter of wing, forming anasotmosis withMA. M with free base; MA and MP branching off CuAin separate stems. CuA dividing into CuA1 and CuA2proximal of MA base. CuP in its basal third notapproaching А1.

S p e c i e s c o m p o s i t i o n. Type species.

C o m p a r i s o n. The new genus is similar to thegenera Griphopteron Handlirsch, 1939 from Mecklen�burg locality (Toarcian of Germany) and Embigryllusgen. nov. from Arzviller locality and differs from themin CuA dividing into CuA1 and CuA2 proximal of thebase of MA. In Griphopteron and Embigryllus, CuAdivides into CuA1 and CuA2 distal of the base of MA.

Vosgesopterum arzvillerensis Aristov, Grauvogel�Stamm et Marchal�Papier, sp. nov.

Plate 7, fig. 9

E t y m o l o g y. From Arzviller locality.

H o l o t y p e. G.�S. no. 5787/5788, part and coun�terpart of forewing; Arzviller locality; Middle Triassic,Anisian Stage.

D e s c r i p t i o n (Fig. 2c). The costal field at thebase of RS is equal in width to the subcostal field; SCreaches beyond the middle of the wing. The apex of Ris S�shaped. The anastomosis of RS+MA has fivebranches, MP is simple. CuA1 has one short bifurca�tion; CuA2 is sharply bent at points where the trans�verse veins enter it. А1 is simple. The transverse veinsare simple.

M e a s u r e m e n t s, mm. Forewing length, 11.

M a t e r i a l. Holotype.

Genus Embigryllus Aristov, Grauvogel�Stamm et Marchal�Papier, gen. nov.

E t y m o l o g y. From the order Embioptera andthe generic name Blattogryllus.

Ty p e s p e c i e s. E. shcherbakovi sp. nov.D i a g n o s i s. Small insects. Wing apex rounded.

RS beginning in basal quarter of wing and forminganastomosis with MA. M without free base, MA andMP branching off CuA in separate stems. CuA withone short bifurcation at posterior margin of wing. CuPin basal third closely approaching А1.

S p e c i e s c o m p o s i t i o n. Type species.C o m p a r i s o n. The new species is similar to the

genera Griphopteron Handlirsch, 1939 from Mecklen�burg locality (Toarcian of Germany) and Vosge�sopterum gen. nov. from Arzviller locality and differsfrom them in M without free base, CuA dividing rela�tively distally, and the small size. In Griphopteron andVosgesopterum the base of M is free, CuA branchesproximal of the middle, and the length of the forewingis 11–20 mm.

Embigryllus shcherbakovi Aristov, Grauvogel�Stamm et Marchal�Papier, sp. nov.

Plate 7, fig. 10

E t y m o l o g y. In honor of the paleoentomologistD.E. Shcherbakov.

H o l o t y p e. G.�S. no. 6643, impression of forew�ing; Arzviller locality; Middle Triassic, Anisian Stage.

D e s c r i p t i o n (Fig. 2d). The anterior margin ofthe wing is straight. RS+MA divides near the middleof the wing and has two branches; MP has one shortbifurcation. CuA is curved parallel to the posteriormargin of the wing. А1 and А2 are simple. The trans�verse veins are simple.

M e a s u r e m e n t s, mm. Forewing length, 4.M a t e r i a l. Holotype.

ACKNOWLEDGMENTS

The authors are grateful to A.P. Rasnitsyn (Boris�siak Paleontological Institute, Russian Academy ofSciences) and S.Yu. Storozhenko (Institute of Biologyand Pedology, Far East Branch, Russian Academy ofSciences) for important comments and toD.E. Shcherbakov (Borissiak Paleontological Insti�tute) for assistance.

This study was supported by the Russian Founda�tion for Basic Research, project nos. 07�04�01158 and09�04�01241, and by the Program “Origin of Bio�sphere and Evolution of Geo�Biological Systems” ofthe Presidium of Russian Academy of Sciences.

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