Dziriblatta (Bohn, 2019)

Bohn, Horst, 2021, Revision of the genus Dziriblatta Chopard, 1936 (Blattodea, Ectobiidae, Ectobiinae) III. The species of the subgenus Dziriblatta, Zootaxa 4964 (2), pp. 201-250 : 215-216

Key for the determination of the males of subgenus Dziriblatta

1. T6 unusually long, length/breadth ≥39% ( Figs. 2C–F ); T5–7 laterally with sparsely distributed spatular bristles ( Figs. 1B–D ); pouch lobes of T7 gland at least basally sclerotized, shortly before opening into the pit united to a large common orifice almost of the breadth of the pit (ep, Figs. 1A , 8 ); transversal mesal part of the posterior border of T10 between weakly convex to weakly concave ( Figs. 1E, F , 3G–I )....................................................................... 2

- T6 not unusually long, length/breadth ≤36% ( Figs. 10F–H ); at least T7 laterally with densely arranged spatular bristles ( Figs. 12D, E ); pouch lobes of T7 gland unsclerotized, each separately opening with a relatively small orifice into the pit ( Fig. 11C ); transversal part of the posterior border of T10 distinctly concave ( Fig. 10E )....................................... 6

2. Pouch lobes much longer than the length of T7, only in the basal part and there only weakly sclerotized, throughout their length flat; only with a large bipartite pit ( Fig. 8 ); discs of thoracic nota usually completely dark......... Dziriblatta (Dz.) algerica

- Pouch lobes not or only scarcely longer than the length of T7, conical, in full length three-dimensional and sclerotized; apart from the large bipartite pit additionally on each side with a smaller circular pit (hole) ( Fig. 1A ); discs of thoracic nota never completely dark, either almost completely lightly coloured or dark maculose...................................... 3

3. Pouch lobes of about the length of T7 (87–126%, mean 102%), conical, with narrowly rounded tip ( Figs. 1A , 3C ); long pit bristles strictly straight, forming on each side a slender and dense bundle ( Figs. 3A, B )........... Dziriblatta (Dz.) bolivari

- Pouch lobes shorter, measuring less than 84% (mean 70% or less) of the length of T7, tip usually broadly rounded; long pit bristles less straight, forming usually less dense bundles....................................................... 4

4. Posterior border of T6 with a deep sinusoidal excavation ( Fig. 7C ), T7 with strongly produced latero-posterior corners and a wide concavity in between ( Fig 7D ); pouch lobes very short, caplike, with more or less transversely cut tip, length 37% ( Fig. 7E ); all pit bristles strongly curved or curled ( Figs. 7F, G )......................... Dziriblatta (Dz.) pilleata spec. nov.

- T6,7 similarly shaped as in the other members of the species group: posterior border of T6 only with a week excavation ( Fig. 4C ), latero-posterior corners of T7 moderately produced ( Fig. 4D ); pouch lobes longer, obtusely conical, with broadly rounded tip ( Fig. 3D ), at least long pit bristles less strongly bent....................................................... 5

5. Length of the pouch lobes 58-83b% (mean 70%) ( Figs. 3D–F , 4D , 5A–C); long pit bristles almost straight, only tips may be slightly bent, arranged in rather compact bundles ( Figs. 4F–H )..................... Dz. (Dz.) brevisacculata spec. nov.

- Length of pouch lobes 45-59% (mean 53%) ( Figs. 5D–F , 6D ), long pit bristles more strongly bent, loosely distributed over the full width of the pit hole ( Fig. 6F–H )....................................... Dziriblatta (Dz.) curvisetosa spec. nov.

6. Pouch lobes very flat, elongated lappet-shaped, basally broad, about as broad as half of the glandular pit of T7, apically pointed, often unregulartly lobated and folded ( Figs. 12A , 13 )............................. Dz. (Dz.) lobososacculata spec. nov.

- Pouch lobes narrowly cylindrical, tubelike, longer than the abdomen, wriggled ( Fig. 15 )............................. 7

7. Latero-posterior corners of T5 triangularly produced ( Fig. 23B ); T5‒7 laterally densely covered with spatular bristles ( Figs. 25E–G ); posterior wall of the pit apart from a low and narrow sagittal ridge without elevations ( Fig. 24 ); surface posteriorly of the pit opening more or less membraneous and rather conspicuous by its whitish appearance... Dz. (Dz.) undulata spec. nov.

- Latero-posterior corners of T5 not produced ( Figs. 10B , 14B ); only T7 laterally densely covered with spatular bristles ( Figs. 14E–G ); posterior wall of the pit with an elevation covering a large part of its breadth ( Figs. 15 , 18 , 22 ); surface posteriorly of the pit opening not membraneous......................................................................... 8

8. Pre-glandular margin of T7 only slightly produced anteriorly; elevations on the posterior wall of the glandular pit relatively flat ( Figs. 15 , 16 )............................................................ Dz. (Dz.) planotuberculata spec. nov.

- Pre-glandular margin of T7 strongly produced anteriorly; elevations on the posterior wall of the glandular pit higher ( Figs. 18 , 22 )................................................................................................. 9

9. Pouch lobes with numerous short side branches; latero-posterior corners of T7 scarcely produced, broadly rounded ( Fig. 22 )...................................................................... Dz. (Dz.) ramososacculata spec. nov.

- Pouch lobes unbranched or with only few, very short side branches; latero-posterior corners of T7 strongly produced, narrowly rounded ( Fig. 18 )......................................................... Dz. (Dz.) altotuberculata spec. nov.

Figures

FIGURE 1.A. T7 of the male type of Dziriblatta (Dz.) bolivari, dorsal view. Abbreviations: ep common end piece of the pouch lobes; gu gutter; h lateral hole; lbr bundle of long bristles protruding from the pouch into the pit hole; sbr short bristles arranged in short ribbon on the pit bottom; p pouch lobe with long bristles shining through; pm pre-glandular margin of T7 covering dorsally the basal parts of the pouch and forming together with the roof of the pouch the edge of the pouch opening po; pw posterior wall of the pit; r sagittal ridge; tl trail, sclerotized posterior part of the pit bottom.—B–D. Dz. (Dz.) algerica, lateral parts of T5 (B), T6 (C) and T7 (D) showing sparsely distributed spatular bristles on all three tergites, as is true for all species of the bolivari-species group.—E, F. Shape of the posterior border pt of T10 in Dz. (Dz.) algerica (E) and Dz. (Dz) bolivari (F), c cercus.—Enlargements: Same scale for B–D and for E,F.—Identifications:A (Type of Dz. (Dz) bolivari), B–D (Ma 118/1), E (Ma 57/3), F (Sp 209/1).

FIGURE 2. A–C. Tergites of a rather darkly coloured specimen of Dz. (Dz.) bolivari. A: T2–4; B: T5; C: T6.—D–F. T6 of lightly coloured specimens from D: Dz. (Dz.) curvisetosa; E: Dz. (Dz.) brevisacculata; F: Dz. (Dz.) bolivari.—G–J. Lateral pre-ridge area pa of T2 with glandular pores (arrows), tr transversal ridge. G: Dz. (Dz.) bolivari; H: Dz. (Dz.) curvisetosa; I: Dz. (Dz.) brevisacculata; J: Dz. (Dz.) pilleata.—Enlargements: Same scale in A–C and in G–J.—Identifications: A–C (Ma 188/1), D (Sp 219/5), E (Sp 193a/4), F (Ma 5a/4), G (Ma 222/1), H (Sp 219/6), I (holotype, Sp 193a/2), J (holotype, Sp 187/13).

FIGURE 3. A–C. Male structures of Dziriblatta (Dz.) bolivari. A, B: dorsal view into the interior of the central pit extending between the dorsal edge of the pouch opening po and the posterior wall of the pit pw. In A bases of bristles (light circles) focused to show how the layer of bristles on the ventral wall of the pouch p is continued outside the lobes in two ribbons on the membraneous part of the pit bottom m; tl trail, slerotized part of the pit bottom; in B bristles in full length focused, notice the curving of the shorter mesal bristles sbr and the straight appearance of the long bristles lbr originating in the pouch lobes, and their common ending near the sagittal ridge r. C: male T7 with pouch lobes; abbreviations not yet mentioned: gu gutter; h lateral hole.—D–F. Male of Dz. (Dz.) brevisacculata, T7 with pouch lobes, in D one of the three specimens with relatively narrowly rounded lobe tips.—G–I. Male terminalia showing the posterior border pt of T10 of the species Dz. (Dz.) brevisacculata (G), curvisetosa (H), and pilleata (I).—Enlargements: Same scale for A,B, for D–F, and for G–I.—Identifications:A (Sp 123/1), B,C (Sp 209/1,5), D,E (Sp 193a/4,3), F (Sp 457/2), G (holotype, Sp 193a/2), H (Sp 214/4), I (holotype, Sp 187/13).

FIGURE 4. A–J. Male structures of Dziriblatta (Dz.) brevisacculata. A: T2–4; B: T5; C: T6; D: T7; E: T7, central region in more detail; F–H: dorsal view into the interior of the central pit; I, J: thoracic nota with tegmina.—Abbreviations: gu gutter; h lateral hole; lbr, sbr long, short bristles; n notch; p pouch; po dorsal edge of pouch opening; pa pre-ridge area; pw posterior wall of pit; r sagittal ridge; tl trail; tr transversal ridge.—Enlargements: Same enlargements for A–C, for F–H, and for I,J.—Identifications:A–F,I (holotype, Sp 193a/2), G (Sp 192a/3), H (Sp 457/6), J (Sp 186a/5).

FIGURE 5. A–F. Male T7 of species of the bolivari-species group. A–C: Dziriblatta (Dz.) brevisacculata, in A specimen with exceptional shape and length of the pouch lobes; D–F: Dz. (Dz.) curvisetosa.—Abbreviations: br bristles; gu gutter; h lateral hole; p pouch lobe; r sagittal ridge; tl trail.—Identifications: A (Sp 193a/1), B,C (Sp 458/2,3), D,E (Sp 214/5,7), F (Sp 219/2).

FIGURE 6. A–I. Male structures of Dziriblatta (Dz.) curvisetosa. A: T2–4; B: T5; C: T6; D: T7; E: T7, central region in more detail; F–H: dorsal view into the interior of the central pit. I: thoracic nota with tegmina.—Abbreviations: br bristles; gu gutter; h lateral hole; m membraneous part of the pit bottom; p pouch lobe; po dorsal edge of pouch opening; pw posterior wall of pit; r sagittal ridge; tl trail.—Enlargements: Same scale for A–C and for F–H.—Identifications: A–F,I (holotype, Sp 214/4), E,G (Sp 214/5), H (Sp 214/6).

FIGURE 7. A–I. Male structures of Dziriblatta (Dz.) pilleata. A: T2–4; B: T5; C: T6; D: T7; E: T7, central region in more detail; F–H: dorsal view into the interior of the central pit, F,G with focus on bristles, H focus on the trail, sievelike due to numerous glandular pores; I: thoracic nota with tegmina.—Abbreviations: arrow in D indicates the missing shoulders; br bristles; gu gutter; h lateral hole; m membraneous part of the pit bottom; p pouch lobe; po dorsal edge of pouch opening; pw posterior wall of pit; r sagittal ridge; tl trail.—Enlargements: Same scale for A–C and for F–H.—Identifications: All images from the holotype, Sp 187/13.

FIGURE 8. Dziriblatta (Dz.) algerica, tergite 7 of male with glandular pit and the two pouch lobes, from the locality Ma 24/4 in Morocco.—Abbreviations: ep common end piece of the pouch lobes; gu gutter; lbr long bristles within the pouch lobes; m membraneous anterior part of the pit bottom; pm pre-glandular margin of T7; po dorsal edge of pouch opening; pw posterior wall of pit; s sclerotized part of pouch lobes; r sagittal ridge; tl trail.

FIGURE 10. A–N. Structures of Dziriblatta (Dz.) lobososacculata.—A–H. Male tergite structures. A: T1–4; B: T5; C: T6; D: T7, gl gandular pit, p pouch lobe; E: terminalia, showing the excavated posterior border pt of T10, c cercus; F–H: T6 of three specimens showing the variability in the shape of the posterior border; arrow heads in H point to three excavations, a deeper mesal and two very shallow lateral ones, in F,G only the mesal one present; length of pre-ridge area pa slightly increasing towards the median notch n, tr transversal ridge.—I, J. Tegmina of male (I) and female (J).—K–N. Thoracic nota with tegmen of male (K) and female (L–N).— Enlargements: Same scale for A–D, for I,J, and for K–N.— Identifications: A–D,J,K (holotype, Ma 14a/2), E (Ma 22a/3), F,G (Ma 14a/5,6), H (Ma 89/2), I (Ma 92/3), L (Ma 158/2), M (Ma 84/2), N (Ma 63/3).

FIGURE 11. A–D. T7 glandular pit of four male specimens of Dziriblatta (Dz.) lobososacculata with the openings of the pouch lobes into the pit. The arrow in B points to a dark contour indicating a transition between the inclined plane of the posterior wall of the pit and the right lateral gutter. Notice the great variability of the pit structures; profile of posterior ridge and neighbouring grooves considerably strengthened from A to C.—Abbreviations: aw anterior wall of the pit; gu gutter; lbr, sbr long, short bristles; n nose, basal end of the posterior ridge pr; pm pre-glandular margin; po opening of the pouch lobe via the tail pipe tp, between arrows in C; tl trail; w membraneous window.— Enlargements: Same scale for A–D.—Identifications:A (Ma 22a/3), B (Ma 166/1), C (Ma 27a/3), D (Ma 251/1).

FIGURE 12. A–E. Male structures of Dziriblatta (Dz.) lobososacculata. A: T7 with pouch lobes. Abbreviations: gu gutter; lbr long bristles, only few present in the basal region of the pouch lobes; sbr ribbon of short bristles; pb line indicates approximate level of posterior border of the pit opening; pm pre-glandular margin of T7, mesally rather transparent, forming the anterior border of the pit opening; po dorsal edge of the fairly semi-circular opening of a pouch lobe into the pit; pr posterior ridge, accompanied on both sides by grooves gr; tl trail, bottom of the pit; B–E: distribution of bristles on the lateral parts of T5–7, sparse on T5 (B), moderately dense on T6 (C), rather dense on T7 (D,E).—Enlargements: Same scale for A–E.—Identifications: A (Ma 89/1), B–D (Ma 14/4), E (Ma 89/2).

FIGURE 13. Dziriblatta (Dz.) lobososacculata, T7 of the holotype (Ma 14a/2), with rather unregularly shaped pouch lobes. For the identification of structures see Figs.10, 11.

FIGURE 14. A–N. Structures of Dziriblatta (Dz.) planotuberculata.—A–K. Male tergite structures. A: T1–4; B: T5; C: T6; D: T7, with latero-posterior corners only slightly produced; p tubular pouch lobes; gl glandular pit; tl trail; ce central elevation on the posterior wall of the pit, weakly developed; bu sagittal bulge posteriorly of the pit, in this preparation hardly visible (compare with Fig. 16); E–G: spatular bristles on the lateral parts of T5 (E), T6 (F), and T7 (G), only T7 with densely arranged spatular bristles; H: terminalia showing the excavated posterior border pt of T10, c cercus; I–K: T6 of three specimens showing the variabilty in the shape of the posterior border of the tergite; n median notch; pa pre-ridge-area; tr transversal ridge.—L–N. Thoracic nota with tegmina of male (L) and female (M,N).—Enlargements: Same scale for A–D, for E–G, and for L–N.—Identifications: A–G,I,L (holotype, Ma 13a/6), H,N,M (Ma 81/1,2,3), J (Ma 80/1), K (Ma 178/1).

FIGURE 15. Dziriblatta (Dz.) planotuberculata, T7 of male with pouch lobes, holotype (Ma 13a/6).Abbreviations: bu bulge, scarcely set off (compare with Fig 16); ce central elevation; gu gutter; p pouch lobes; pb approximate level of the posterior border of the pit opening; pm pre-glandular margin of the tergite; pr posterior ridge; tl trail; w membraneous window in the anterior wall of the pit, bearing the pouch openings.

FIGURE 18. Dziriblatta (Dz.) altotuberculata, T7 of male, holotype (Ma 11a/5). Abbreviations: arrowheads point to slight signs of side branches; bu bulge, angularly set off (arrow); ce central elevation; gu gutter; p pouch lobes; pb approximate level of the posterior border of the pit opening; pm pre-glandular margin of the tergite; pr posterior ridge; tl trail.

FIGURE 22. Dziriblatta (Dz.) ramososacculata, T7 of male with pouch lobes (Ma 184/1).Abbreviation:pm pre-glandular margin. For the identification of other structures see preceding figure.

FIGURE 23. A–I. Male tergites of Dziriblatta (Dz.) undulata. A: T1-4; B: T5; C: T6; D: T7; gl glandular pit; p pouch lobes; E: terminalia, showing the excavated posterior border pt of T10, c cercus, pp paraproct structures surpassing pt; F,G: T6 of two specimens with differently deep median excavations at the posterior border; n median notch, pa pre-glandular margin; tr transversal ridge; H, I: T7 of specimens from two other localities with slightly differing pit structures; aw, pw anterior, posterior wall of pit; ab, pb anterior, posterior border of the pit opening (line).—Enlargements: Same scale for A–D, for F,G, and for H,I.—Identifications: A–F (holotype, Ma 6b/8), G,I (Ma 225/2), H (Ma 105/3).

FIGURE 24. Dziriblatta (Dz.) undulata, T7 of male with pouch lobes, holotype (Ma 6b/8). Abbreviations: ab, pb anterior, posterior border of the pit opening; aw, pw anterior, posterior wall of pit; pm pre-glandular margin of T7; pr posterior ridge; w membraneous window in the anterior wall of the pit; arrows mark the membraneous areas surrounding the pit opening: the upper oblique arrows point to long diagonal folds, the upright arrows below define the lateral extension of the membraneous area posteriorly of the pit opening.

FIGURE 25. A–G. Structures of Dziriblatta (Dz.) undulata. A, B: left tegmen of male (A) and female (B); C, D: thoracic nota with tegmina, male (C) and female (D); E–G: lateral parts of male T5 (E), T6 (F), T7(G), all tergites densely covered with spatular bristles.—H–K. Phase contrast images of bristlelike microtrichial processes arising from short indentations (arrows) of the cuticular lining of the tube wall; H, I: Dz. (Dz.) undulata, processes relatively short, radially directed; sample in H from the tip of a pouch lobe, in I from an intermediate region; J, K: Dz. (Dz.) ramososacculata, processes longer, obliquely downward directed; sample from side branches, in J with part of the main axis (a apical, b basal site); s remnants of soft tissues, in K removed by image editing.—Enlargements: Same scale for A,B, for C,D, for E–G, and for H–K.—Identifications:A,B (Ma 105/5,9), C,E–I (holotype of Dz. (Dz.) undulata, Ma 6b/8), D (Ma 6b/10), J,K (Ma 184/1).