Blattula langfeldti ( Geinitz, 1880 )

Vrsansky, Peter & Ansorge, Jörg, 2007, Lower Jurassic cockroaches (Insecta: Blattaria) from Germany and England, African Invertebrates 48 (1), pp. 103-126 : 115-122

Blattula langfeldti ( Geinitz, 1880)

Figs 8–15

Blattina langfeldti: Geinitz 1880: 521 , fig. 3 [Fw, holotype FGWG 117 /20, examined].

Blattina nana: Geinitz 1883: 30 , pl. 22, fig. 2 [Hw, holotype FGWG 118 /3, examined]. Syn. n.

Blattina (Mesoblattina) dobbertinensis: Geinitz 1884: 570 , pl. 13, fig. 1 [Fw, holotype FGWG 119 /2, examined]. Syn. n.

Blattina incerta: Geinitz 1884: 571 , pl. 13, fig. 2 [Hw, holotype FGWG 119 /2, examined]. Syn. n.

Dipluroblattina scudderi: Geinitz 1887: 200 , pl. 5, fig. 7 [Fw, holotype FGWG 120/2, examined]. Syn. n. Ctenoblattina langfeldti (Geinitz) : Scudder 1886: 443. Mesoblattina dobbertinensis (Geinitz) : Scudder 1886: 455. Mesoblattula dobbertiniana: Handlirsch 1906 –08: 430, pl. 40, fig. 23 [Fw, holotype FGWG 122/103,

examined]. Syn. n. Mesoblattula geinitziana: Handlirsch 1906 –08: 430, pl. 40, fig. 24 [Fw, holotype FGWG 122 /10-11,

examined]. Syn. n. Blattula dobbertinensis (Geinitz) : Handlirsch 1906 –08: 431, pl. 40, fig. 25. Blattula langfeldti (Geinitz) : Handlirsch 1906 –08: 431, pl. 40, figs 26, 27. Blattula ancilla: Handlirsch 1906 –08: 431, pl. 40, fig. 28 [Fw, holotype FGWG 122/12, examined]. Syn. n. Blattula geinitzi: Handlirsch 1906 –08: 432, pl. 40, fig. 29 [Fw, probably lost]. Syn. n. Battula scudderi (Geinitz) : Handlirsch 1906 –08: 432, pl. 40, fig. 30.? Blattula incerta (Geinitz) : Handlirsch 1906 –08: 432, pl. 40, fig. 31.? Blattula debilis: Handlirsch 1906 –08: 433, pl. 40, fig. 32. Syn. n.? Blattula pusillima: Handlirsch 1906 –08: 433, pl. 40, fig. 33 [Hw, holotype FGWG 122/250, examined].

Syn. n. (? Mesoblattina ) nana (Geinitz) : Handlirsch 1906 –08: 435, pl. 40, fig. 38. Parablattula reticulata: Handlirsch 1920: 191 , fig. 153 [Fw, holotype FGWG 123 /26, examined]; 1939: 61 ,

pl. 5, fig. 83. Syn. n.? Mesoblattula ala: Handlirsch 1939: 58 , pl. 4, fig. 74 [Hw, holotype FGWG 123/45, examined]. Syn. n. Blattula riparia: Handlirsch 1939: 58 , pl. 4, fig. 75 [Fw, holotype FGWG 123/20, examined]. Syn. n. Blattula intercalata: Handlirsch 1939: 58 , pl. 4, fig. 76 [Fw, holotype FGWG 123/21, examined]. Syn. n. Blattula brunneri: Handlirsch 1939: 59 , pl. 5, fig. 77 [Fw, holotype FGWG 123/22, examined]. Syn. n.? Blattula vicina: Handlirsch 1939: 59 , pl. 5, fig. 79 [Fw, holotype FGWG 123/23, examined]. Syn. n.? Blattula acutipennis: Handlirsch 1939: 59 , pl. 5, fig. 80 [Hw, NHMW 1984/33/11]. Syn. n. Peloblattula oligoneura: Handlirsch 1939: 60 , pl. 5, fig. 81 [Fw, holotype FGWG 123/24, examined]. Syn. n. Parablattula simplicissima: Handlirsch 1939: 60 , pl. 5 fig. 82 [Fw, holotype FGWG 123/25, examined;

counterpart NHMW 1984/33/12]. Syn. n. Metablattula lipomena: Handlirsch 1939: 61 , pl. 5, fig. 84 [Fw, holotype FGWG 123/27, examined]. Syn. n. Chiloblattula simplex: Handlirsch 1939: 61 , pl. 5, fig. 85 [Fw, holotype FGWG 123/28, examined]. Syn. n.? Chiloblattula subcostalis: Handlirsch 1939: 62 [Hw, holotype FGWG 123/29, examined]. Syn. n.? Chiloblattula longipennis: Handlirsch 1939: 62 , pl. 5, fig. 86 [Hw, holotype FGWG 123/30, examined]. Syn. n.

Ectinoblattula medialis: Handlirsch 1939: 63 , pl. 5, fig. 90 [Fw, holotype FGWG 123 /57, examined]. Syn. n. Blattula hattorfensis: Bode 1953: 114 , pl. 5, fig. 93 [Fw, SMN W 13 ]. Syn. n.

Englyptoblatta elegantula: Bode 1953: 120 , pl. 6, fig. 100 [Fw, SNM W 9/11]. Syn. n.

Elisama langfeldti (Geinitz) : Ansorge 2003: 296, fig. 2B.

Blattula langfeldti (Geinitz) : Ansorge 2004: 782, fig. 2.

Diagnosis: Forewing: length 6.5– 9 mm; Sc 1, R 8–14, M 2–7, Cu 4–10, A 5–12. Hindwing: length 6.5–7.5 mm; Sc 1, R 3–4+3–5, M 2–3, Cu 6–8.

Description: Pronotum ovoid in shape, transverse, with distinct coloration. Forewing length 6.5– 9 mm, width about 2.3 mm; Sc simple, rarely with some weak lateral branchlets; R with 8–14 veins; M with 2–7 branches; Cu with 4–10 branches.A reticulate, with 5–12 veins reaching margin, veins in clavus may be reticulate. Hindwing length 6.5–7.5 mm with Sc simple; R divided near base into R with 3–4 veins and RS with 3– 5 veins; M with 2 or 3 branches; CuA with 6–8 veins. Legs thin, with sparse, long and thin spines.

Holotype (examined): FGWG 117 /20, isolated forewing. GERMANY: Mecklenburg : former clay pit of Schwinz , near Dobbertin; carbonate concretions of the marine “Green Series” clay, Lower Toarcian, exaratum Subzone.

Additional material examined: Besides the re-studied holotypes described by Geinitz and Handlirsch from the Lower Toarcian of Dobbertin and listed in the synonymy, the following additional material is in Ansorge’s collection. Hindwings : LDA 40; 76; 85; 86; 109; 455; 485; 594/3; 660/1, 2; 593/3; LGA 86; 96; 275; 331/1, 2; 380; 425; 579; 594; 888; 893; 1077; 1201; 1275; 1372; 1529; 1578; 1739; 1820; 2169. Forewings : LDA 7, 66; 169; 240; 145; 254; 289; 326; 348; 581; 586; 629; 611; 798; 869; 910; 1017; 2196; LGA 13; 118; 154; 880; 1328; 1435; 1519; 1842; 1887; 1926; 2006; 2015; 2196; 2356, S 137; 139. Complete specimens: LDA 96; LGA 888; 1820. Two species described by Bode (1953) from the Lower Toarcian of Hattorf (Lower Saxony) have not been re-examined .

Note: A few complete specimens with associated fore- and hindwings allow us to determine isolated fore- and hindwings.

Variability (Tables 2, 3): Both complete specimens from Grimmen show considerable variation between the left and right wings. Sc is stable, simple in all wings. Other veins vary. Hindwing variation, considered as more stable, is expressed in the varying number of RS branches (4 in left wing; 5 in right wing), while R is stable in this case (3). M is 3-branched in the left wing, and 2-branched in the right wing. Unfortunately, the wings are not complete, so it is uncertain whether the Cu will compensate differences in the total vein number (6 in right wing, and evidently more than 3 in the left wing).

The variability studies of B. langfeldti revealed some general principles regarding the control of the cockroach wing venation development. With the increasing number of veins in a competent venational complex, the variability falls (coefficient of variation (CV) is lower). Therefore we have also analysed the dependence of the CV according to the number of veins. Results show that the CV is comparatively low in the competent venational systems (R, M, Cu), higher in neighbouring venational complexes and the highest between complexes that are not neighbouring (in the forewing of B. langfeldti it is low in (R+M) complex, high in (R) and (Cu); in the hindwing, the most apparent is the relatively high variability of (R+Cu) complex). That means that the control mechanisms are developed autonomically, “working best” in each venational complex (R, M, Cu) (each major branch has its own control mechanism) but these separate systems communicate with their surroundings, and influence each other (see Schneider 1977, 1978 a).

Remarks: There is no gap in the variability of the venation which would suggest dimorphism in the flight abilities of both sexes.

Figures

Fig. 8. Blattula langfeldti (Geinitz, 1880): (A–C) LGA 1820, Lower Toarcian of Grimmen: (A) total before preparation of legs, (B) after preparation of legs and antennae, (C) details of legs; (D) LDA 96, Lower Toarcian of Dobbertin.

Fig. 9. Blattula langfeldti (Geinitz, 1880), LGA 1820, Lower Toarcian of Grimmen: (A) details of specimen before preparation of legs, (B) details of legs after preparation, (C) details of antennae.

Fig. 10. Blattula langfeldti (Geinitz, 1880), LGA 888/1-2, Lower Toarcian of Grimmen: (A, B) photographs of part and counterpart, (C, D) details of body structures.

Fig. 11. Blattula langfeldti (Geinitz, 1880): (A) FGWG 123/26, forewing, holotype of Parablattula reticulata Handlirsch, 1939, Dobbertin; (B) LDA 7, forewing, Dobbertin; (C) LDA 66, forewing, Dobbertin; (D) LGA 2356, forewing, Grimmen; (E) LDA 586, forewing, Dobbertin; (F) LGA 96, hindwing, Grimmen; (G) LGA 1275, hindwing, Grimmen; (H) LGA 2507, hindwing, Grimmen; (I) LGA 380, hindwing, Grimmen; (J) LDA 85, hindwing, Dobbertin.

Fig. 12. Blattula langfeldti (Geinitz, 1880), details of venation of forewings, Lower Toarcian of Dobbertin and Grimmen: (A) FGWG 122/12, holotype of Blattula ancilla Handlirsch, 1906; (B) LGA 880; (C) FGWG 123/26, holotype of Parablattula reticulata Handlirsch, 1920; (D) FGWG 123/25, holotype of Parablattula simplicissima Handlirsch 1939; (E) FGWG 123/28, holotype of Chiloblattula simplex Handlirsch, 1939; (F) FGWG 123/27, holotype of Metablattula lipomena Handlirsch, 1939; (G) FGWG 119/2, holotype of Blattula dobbertinensis (Geinitz, 1884); (H) FGWG 122/103, holotype of Mesoblattula dobbertiniana Handlirsch, 1906.

Fig. 13. Blattula langfeldti (Geinitz, 1880), details of venation of forewings, Lower Toarcian of Dobbertin, Grimmen (LGA 2356) and Schandelah (S 139): (A) FGWG 123/21, holotype of Blattula intercalata Handlirsch, 1939; (B) FGWG 120/2, holotype of Dipluroblattina scudderi Geinitz, 1887; (C) FGWG 123/20, holotype of Blattula riparia Handlirsch, 1939; (D) FGWG 117/20, holotype of Blattina langfeldti Geinitz, 1880; (E) FGWG 123/57, holotype of Ectinoblattula medialis Handlirsch, 1939; (F) FGWG 122/10-11, holotype of Mesoblattula geinitziana Handlirsch, 1906; (G) LGA 2356; (H) S 139; (I) FGWG 123/24, holotype of Peloblattula oligoneura Handlirsch, 1939; (J) FGWG 123/22, holotype of Blattula brunneri Handlirsch, 1939.

Fig. 14. Blattula langfeldti (Geinitz, 1880), details of venation of hindwings, Lower Toarcian of Dobbertin (FGWG) and Grimmen (LGA): (A) FGWG 123/23, holotype of?Blattula vicina Handlirsch, 1939; (B) LGA 579; (C) LGA 96; (D) LGA 893; (E) FGWG 123/30, holotype of?Chiloblattula longipennis Handlirsch, 1939; (F) FGWG 122/250, holotype of?Blattula pusillima Handlirsch, 1906.

Fig. 15. Blattula langfeldti (Geinitz, 1880), details of venation of hindwings, Lower Toarcian of Dobbertin and Grimmen (LGA 380): (A) FGWG 123/45, holotype of?Mesoblattula ala Handlirsch, 1939; (B) LGA 380; (C) FGWG 118/3, holotype of Blattina nana Geinitz, 1883; (D) FGWG 119/2, holotype of Blattina incerta Geinitz, 1884; (E) LDA 85; (F) FGWG 123/29, holotype of?Chiloblattula subcostalis Handlirsch, 1939.