Tillyardembiidae G. Zalessky, 1938

Aristov, D. S., 2020, Revision of the genera Heteroptilon, Nugonioneura and Opistocladus from the Lower Permian of USA (Insecta: Cnemidolestida: Tillyardembiidae, Nugonioneuridae and Parmapteridae), Far Eastern Entomologist 401, pp. 1-9 : 2-3

Family Tillyardembiidae G. Zalessky, 1938

Tillyardembiidae G. Zalessky, 1938: 64 (type genus: Tillyardembia G. Zalessky, 1937 );

Sharov, 1962: 124; Rasnitsyn, 1980: 152; Carpenter, 1992: 123; Storozhenko, 1997: 8;

1998: 91; 2002: 279; Aristov & Rasnitsyn, 2009: 263; 2010: 17; 2012: 48; 2014: 411;

Aristov, 2014: 20; Aristov, 2018: 1382.

Permocapniidae Martynov, 1940: 20 (type genus: Permocapnia Martynov, 1940 ); synonimized by G. Zalessky, 1950: 41–60.

Heteroptilidae Carpenter, 1976: 346 (type genus: Heteroptilon Carpenter, 1976 ); Carpenter,

1992: 126; syn. n.

Type genus: Tillyardembia G. Zalessky, 1937 .

EMENDED DIAGNOSIS. Small and medium sized insects. SC ends in forking on C and RS. Starting in wings basal third, RS branching abundantly, with branches occupy whole wing apex. M with prominent М 5 or М merged with CuA on a short section near the base. M with few branches, not split into MA and MP, start branching past base of RS. CuA start branching in its distal third, forms posterior branching comb. Posterior branches of CuA are lacking. Anal area small, with two or three anal veins.

COMPARISON. The family is most similar to Emphylopteridae from Carboni-

ferous of France, but differs in branching CuA. Emphylopteridae have simple CuA

(Aristov, 2014).

COMPOSITION. Five genera: Tillyardembia G. Zalessky, 1937 from Chekarda,

Barda, Kishert’ ( Russia: Perm Region; Lower Permian, Kungurian Stage), and

Zalazna ( Russia: Kirov Region; Upper Permian, Vyatkian Stage), Heteroptilon

Carpenter, 1976 ( USA: Kansas; Lower Permian, Leonardian Stage), Kungurembia

Aristov, 2004 from Chekarda and Soyana ( Russia: Arkhangelsk Region; Middle

Permian, Kazanian Stage), Kamamica Aristov et Rasnitsyn, 2014 from Tyul’kino

( Russia: Perm Region; Lower Permian, Kungurian Stage), Paralongzhua Prokop,

Szwedo, Lapeyrie, Garrouste et Nel, 2015 from Lodève ( France: Languedoc Region;

Middle Permian, Salagou Formation), and Udembia Aristov, 2018 from Kostovaty

( Russia: Udmurtia; Middle Permian, Urzhumian Stage).

NOTE. The genus Paralongzhua was described within Archaeorthoptera nec

Panorthoptera as family undetermined taxon (Prokop et al., 2015). Comparison with the rest of tillyardembiids shows attribution of Paralongzhua to this family

(see key below).

Key to genera of the family Tillyardembiidae based on forewing characters

1(6) Costal field near base of RS is equal in width with subcostal one.

2(3) SC ends in wings distal third .............................................................................................

.................................... Paralongzhua Prokop, Szwedo, Lapeyrie, Garrouste et Nel, 2015

3(2) SC ends near wings mid length.

4(2) Preradial field narrow ................................................................... Udembia Aristov, 2018

5(7) Preradial field wide ............................................................. Heteroptilon Carpenter, 1976

6(1) Costal field near base of RS wider that subcostal one.

7(10) Forewing with costal fan at the base of subcostal field.

8(9) Base of M not merged with CuA, M and CuA connected with short М 5 .........................

........................................................................................ Tillyardembia G. Zalessky, 1937

9(8) Base of M merged with CuA ................................ Kamamica Aristov et Rasnitsyn, 2014

10(7) Costal fan is lacking ........................................................... Kungurembia Aristov, 2004