Dziriblatta (Dziriblatta) undulata, Bohn, 2021

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 : 214-215

10. Dziriblatta (Dziriblatta) undulata , spec. nov.

Figs. 23A–I , 24 , 25A–I , 26E, G , 27 , 28 , 29

Etymology. The species name refers to the undulate shape of the posterior border of T6 having three concavities (unda, Latin word for wave).

Diagnosis. In the male sex easily recognized by the strikingly whitish area posterior to the glandular pit of T7; further characterized by the well-produced latero-posterior corners of T5, the undulate shape of the posterior border of T6 with deep lateral concavities, the huge dome-shaped glandular pit of T7 without larger internal elevations, and the dense layers of spatular bristles on the lateral parts of T5-7.

Material studied. Type material. MOROCCO. Holotype, 1♂, Rif, Bab-Besen (15 km W Ketama ), 1600 m, leg. B. & H. Bohn (completely on two slides, Ma 6b/8). (Coll. Bohn, ZSM).

Additional material. MOROCCO. 1♂, 1♀, Tidiguin [= J. Tidirhine], Ketama, Rif, VI .1930, Exp. C. Bolívar (slides: ♂, Bo 229, without abdomen; ♀, Bo 137) [Ma 18]. ( MNHN) .— 7♂, 3♀, 3L, Tidiguin [= J. Tidirhine], Ketama, Rif, VI .1930, Exp. C. Bolívar (slide: 1♂, Bo 197) [Ma 18] ; 2♀, Bab Ruadi , Beni Siyyel, VI .1932, C. Bolívar [Ma 19]; 1♂, Hauta Kasdir , Beni Seyel, Gomara, V .1941, E. Morales (slide: MNCN _Ent 233303) [Ma 19] ; 5♀, Imasinen [= Imassinne], Beni Seddat, Rif, VI .1930, Exp. C. Bolívar [Ma 20]; 3♂, 2♀, 2L, Tizi [n’] Taka , Beni Seddat, Rif, VI .1930, Exp. C. Bolívar (slides: 1♂, MNCN _144929; 1♀, Bo 138) [Ma 21] ; 3♀, Zoco Telata , Ketama [= Tleta Ketamen], Rif, VI .1930, Exp. C. Bolívar [Ma 22]; 1♀, Zoco Telata , Ketama, Rif, VIII.1932, F. Escalera [Ma 22] ; 6♀, Iguermalen [= Iguermalet], Beni Mesdui, VI .1932, M. Escalera (slide: 1♀, Bo 138) [Ma 23] ; 1♀, Iguermalen , Targuist, Rif, VI .1930, Exp. C. Bolívar [Ma 23]; 3L, Bab [-] Taza (El Ajmas, Yebala), VI .1941, E. Morales (Ma 53); 1♂, 10♀, 1L, Marruecos español, I[s]saguen, VI .1941, Morales (slide: 1♂, MNCN _233293) ( Ma 105). ( MNMS).— Same locality as holotype : 11♀, 14.VIII.1984 (Ma 6) / exL: 13♂, 9♀, 15L, 1.IV.1988 (Ma 6a) / 5♂, 18♀, 18O, 6. VI .1989 (slides: 1♂, Ma 6b/13; 2♀, Ma 6b/9,10) ; 22♂, 40♀, Rif , 3 km S Ketama, 1500 m, 5./ 6. VI .1989, leg. B. & H. Bohn (slides: 7♂, Ma 105/3–8,11; 1♀, Ma 105/10) ; 8♂, 5♀, ex L: 4♀, Rif , E slope of J. Kouine, 1700-1800 m, 12./ 13.IV.1998, leg. B. & H. Bohn (slide: 2♂, Ma 225/2,7). (Coll. Bohn, ZSM) .

Description. Size. Length of pronotum in male 1.95–2.18 mm (mean 2.06 mm), in female 2.19–2.40 mm (mean 2.25 mm); N 8/8.

Male structures. Latero-posterior corners of T2–4 weakly produced, those of T5, 6 rather strongly produced; posterior border of T2–5 weakly convex or straight, of T6 undulate, with three concavities; in contrast to the other species of the lobososacculata -species group lateral concavities much deeper than median one, intermediate parts strongly convex, transversal ridge with a mesal excurvation to the anterior ( Figs. 23A–C, F, G ); in T5,6 length of preglandular margin (pm) distinctly increasing towards the median notch ( Fig. 23B,C ); surfaces of T3–6 mesally deepened to a common shallow sagittal trough, deepest and broadest on T6, fading away in both dimensions up to T3 (the trough not visible in the figures). T7 ( Figs. 23D, H, I , 24 ) with a deeply concave posterior border, producing broadly rounded triangular latero-posterior corners; surface of the tergite appearing tripartite by areas with whitish cuticle encircling the opening of the glandular pit; consisting of two narrow, towards posteriorly converging areas with longitudinal folds (upper arrows in Fig. 24 ), each starting in a membraneous area at the latero-anterior corners of the tergite and posteriorly merging into a broader whitish area with wrinkled surface extending between the posterior borders of pit opening and tergite (between lower arrows of Fig. 24 ). Opening of the glandular pit large, broadly transversely oval, occupying more than half of the breadth and about two thirds of the length of the tergite (between ab and pb). Pit rather deep, dome-shaped, with steeply descending walls; bottom small, bowl-shaped, adjacent on the anterior wall with a membraneous window (w) containing the openings of the two pouch lobes. Posterior wall with a low, narrow, partly unsclerotized sagittal ridge (pr) reaching down to the bottom and up to the posterior border of the tergite. Pouch lobes ( Fig. 24 ) narrowly cylindrical, of more than abdominal length, wriggled, opening into the pit near together but separately within the membraneous window; interiorly near the opening with numerous bristles reaching with their ends into the pit hole; cuticular lining of the lobes with bristle-like microtrichial structures, shorter than in the preceding three species and radially aligned ( Figs. 25H, I ). T10 with median part of posterior border shallowly, but distinctly concave ( Figs. 23E ). Spatular bristles laterally on T5–7: densely arranged in all three tergites ( Figs. 25E–G ). No glandular pores on T2.

Colouration. Very similar to Dz. (Dz.) lobososacculata . Tegmina ( Figs. 25A–D ) transparent; in the male along the anterior margin infuscated, along the posterior margin with variously sized patches, at the base partly uniting to a larger patch; female tegmina only basally at the posterior margin slightly infuscated, but disc often with several dark spots of various size. Male. Mainly brown to dark brown. Head dark, with yellowish post-interocular stripe; discs of thoracic nota uniformly dark, margins transparent ( Fig. 25C ); legs mostly dark except for the lighter coloured coxatrochanter and femur-tibia joints, rarely with more extended yellowish areas; tergites mainly dark, but often variably lightened, especially laterally on T2–4, with a maculose appearence ( Fig. 23A ); lightenings in the specimens from the most eastern locality (Ma 225, Fig. 23G ) more extensive than in those from the more western localities; T7 medially posteriorly to the pit opening strikingly whitish ( Figs. 23D, H, I , 24 ); sternites almost completely dark, only lateral margins slightly lightened. Female. Head mainly dark, with yellowish post-interocular stripe; discs of thoracic nota dark, at most with very slight lightenings, margins transparent ( Fig. 25D ); legs almost completely yellowish; tergites on a yellowish ground colour extended maculose; sternites mainly dark, laterally with a relatively broad yellowish margin.

Distribution. Distributed in the Rif mountains between Chefchaouen in the West and Aknoul (about longitude of Al Hoceima) in the East, at elevations of 1500–1800 m ( Figs. 27 , 28 ).

It was not possible to exactly identify the localities Bab Ruadi and Hauta Kasdir; both should be situated in the region Beni Esjjel [in older literature called Beni Sayel, B. Seyel, or B. Siyyel] in the NE of Chefchaouen [= El Aaiún]. The position of the corresponding locality [Ma 19] in the distribution map, therefore, only indicates the approximate position.

Remarks. The distribution of this species partially overlaps with the distribution of Phyllodromica vignai Failla & Messina, 1989 ; both species, which are very similarly coloured, were found together at locality Ma 6 ( Bohn 1993). While the males of the two species can easily be distinguished solely by external inspection ( Ph. vignai has longer tegmina considerably surpassing the posterior border of the mesonotum), a reliable distinction of the females requires the study of the genital sclerites: dorsal basivalvular sclerites (bd) of both sides in Ph. vignai separate ( Fig. 26F ), in Dz. (Dz.) undulata (as in most of the subgenera of the genus Dziriblatta ) fused to an ringlike structure ( Fig. 26E ); laterosternal shelf (ls) in Ph. vignai fairly rounded with long lateral arms (a) ( Fig. 26H ), in Dz. (Dz.) undulata (as in most subgenera of Dziriblatta ) transversal with very short arms ( Fig. 26G ).

Figures

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).

FIGURE 26. A. Male of Dziriblatta (Dz.) lobososacculata, comblike cuticular scales covering the inside of the pouch lobes, the arrows point to the margin of the pouch.— B–D. Anterior part of T7 showing the evolutionary steps in the development of pouch and pit: B: Dz. (Pauciscleroblatta) haffidi, only with a bilobed pouch, no pit present; C: Dz. (Discleroblatta) merrakescha, pouch posteriorly opening into a trough-shaped pit with membraneous bottom; D: Dz. (Autumnoblatta) spec., with bipartite pit, and a relatively long bilobed pouch.—E–H. Female genital sclerites of Dz. (Dz.) lobososacculata (E, G) and Phyllodromica vignai (F, H), ventral view, posteriorly on top. E, F: terminalia and dorsal genital complex; G, H: ventral genital complex.—Abbreviations: c cercus; dv,bv dorsal,ventral basivalvulae; gu gutter ascending from the membraneous bottom of the pit (m) towards the surface of the tergite; if intersternal folds; ls laterosternal shelf and its arms (a); pt8,9 paratergites 8,9; p pouch; pm pre-glandular margin of T7; pp paraprocts; r sagittal ridge; tl trail, sclerotized bottom of pit; v ovipositor valves (in F distorted).—Enlargements: Same scale for B–D.—Identifications:A (Ma 89/1), B (Ma 87b/1), C (Ma 151/1), D (Ma 165a/1), E, G (Ma 6b/9), F, H (Ma 6b/11).

FIGURE 27. Distribution of the species of subgenus Dziriblatta in Morocco and southern Spain (for mountains and locality numbers see Fig. 28). The lobososacculata-species group is distributed along the main mountain ranges of the Middle and High Atlas and the Rif, the bolivari-species group mainly further North: along the western Tell Atlas in Algeria (see fig. 32 in Bohn 2019), in northern Morocco including the northern part of the Middle Atlas, and in southern Spain along the Cordillera Penibetica, West of the Sierra Nevada up to Algeciras. The names of the species and their symbols are listed in the sequence of their distribution, which in the bolivari-species group is from East to West, in the lobososacculata-species group from North to South.The numbered arrows indicate localities with the common occurrence of two different species of subgenus Dziriblatta. In most cases Dz. (Dz.) algerica is found associated with another species, at locality 1 with Dz. (Dz.) bolivari, at 2 with Dz. (Dz.) ramososacculata, and at 4 with Dz. (Dz.) lobososacculata; at locality 3 Dz. (Dz.) ramososacculata is occurring together with Dz. (Dz.) planotuberculata.—Abbreviation: indet. females = indetermined females, only in Spain.

FIGURE 28. Locality numbers of all species of subgenus Dziriblatta found in Morocco (Ma) and Spain (Sp.).

FIGURE 29. Cladogram of the species of subgenus Dziriblatta reconstructed via the states of 9 male characters as listed below. Due to the complexity of the structures the “characters” are mostly character complexes rather than single characters. The presumed character states of the last common ancestor are listed at the root of the cladogram.—The alphabetic sequence of the states usually follows the assumed sequence in the evolution. But after a branching two different evolutionary lines may develop; the branching point is marked by an arrow (k) indicating where the line branching off is continued. The end of a line is marked by a slash.—For the species of the bolivari subgroup a separate character (10) was introduced in order to allow a differentiation between the four very similar species.