Doratura, J. Sahlberg, 1871, J. Sahlberg. A, 1871

Bückle, Christoph & Guglielmino, Adalgisa, 2022, Revision of the genus Doratura Sahlberg (Hemiptera, Cicadellidae, Deltocephalinae) with particular regard to its distribution in Italy and description of four new species, Zootaxa 5112 (1), pp. 1-116 : 61-65

Key to males (in part additionally with characters concerning the corresponding females)

1. Aedeagus ( Figs 41A–C ) ventrally with two carinae divided by a deep furrow; margin of each carina with row of distinct teeth; styles ( Fig. 41D ) evenly bent with denticle apically of mid-length (female pregenital sternite ( Fig. 34H ) with concave hind margin and triangular median notch)......................................................... D. salina Horváth

-. Aedeagus never with deep ventral furrow dividing two dentate carinae; shaft provided with distinct spinules ( Fig. 23B ) or robust spines ( Fig. 31A ), or ventrally slightly scabrous ( Fig. 10C ) or smooth and shiny ( Fig. 10O )..................... 2

2. Aedeagus dark, ventrally with few (ca. 5–10), but particularly robust and long, black, thorn-shaped spines ( Figs 31A, B ); genital plates laterally strongly sinuate ( Fig. 33A )................................................ D. concors group 3

-. Aedeagus with tooth-shaped spines ( Figs 37A, B ), testaceous, or with small spinules ( Fig. 23B ), or ventrally slightly scabrous ( Fig. 10C ), or completely smooth ( Fig. 10O )............................................................... 5

3. Aedeagus ( Figs 31M–R ) with 1–2 smaller apical spines and ca. 4–7 robust spines in the apical half, shaft in ventral view very narrow, strongly compressed, proximally with large shield-shaped ventro-proximal plate; styles ( Figs 32F, G ) stout, portion apically of denticle short (female pregenital sternite ( Fig. 34D ) strongly protruding caudad with very deep narrow median incision)........................................................................... D. medvedevi Logvinenko

-. Aedeagus shaft ( Figs 31D, G ) less strongly compressed, without shield-shaped ventro-proximal plate, robust spines also in the central part of shaft (female pregenital sternite ( Figs 34A, C ) without median incision).............................. 4

4. Aedeagus ( Figs 31E, G ) with proximal portion of shaft without spines, not widening proximad in ventral view; styles ( Fig. 32C ) stout, portion apically of denticle short (female genital sternite ( Fig. 34C ) abruptly narrowing in basal part, then caudad protruding as tongue-shaped lobe)....................................................... D. rusaevi Kusnezov

-. Aedeagus shaft ( Figs 31C, D ) proximally abruptly widening, proximal portion with spines; styles ( Fig. 32B ) slenderer, portion apically of denticle long and thin (female pregenital sternite ( Fig. 34A ) shorter, hind margin ± semicircular).................................................................................................. D. concors Horváth

5. Aedeagus ( Figs 37A, F ) testaceous, without robust thorn-shaped spines, but ventrally with tooth-shaped spines, arranged in two rows, above all in the apical portion, basal socle very low, fused with shaft (female pregenital sternite ( Figs 34E, F ) very long with protruding rounded hind margin).................................................... D. homophyla group 6

-. Aedeagus with small spinules ( Fig. 23B ), or ventrally slightly scabrous ( Fig. 10C ), or completely smooth ( Fig. 10O )...... 7

6. Aedeagus shaft in lateral view ( Fig. 37C ) basally and medially straight, ventrally slightly concave, apicad abruptly bent dorsad; genital plates ( Fig. 39A ) with rather sharp latero-caudal angle................................. D. homophyla (Flor)

-. Aedeagus shaft ( Figs 37F, G ) ventrally not concave, in its apical half evenly curved; genital plates ( Fig. 39C ) with obtuse laterocaudal angle........................................................................ D. caucasica Melichar

7. Aedeagus ( Fig. 23B ) with spinules of ± equal size distributed evenly on ventral and lateral parts of aedeagus; styles ( Fig. 24A ) ± evenly curved with denticle near mid-length (female pregenital sternite ( Figs 20B, D ) with straight or concave hind margin, with or without slight rounded notch...................................................... D. paludosa group 8

-. Aedeagus with spinules only along its ventral margin ( Fig. 14A ), or ventrally slightly scabrous ( Fig. 10C ), or completely smooth ( Fig. 10O )................................................................................... 10

8. Aedeagus shaft ( Fig. 23G ) proximally with abruptly elevated ventral crest (female pregenital sternite ( Fig. 20E ) with ± straight hind margin without median notch)........................................................... D. iblea D’Urso

-. Aedeagus shaft ( Figs 23B, D ) without abrupt crest, ventrally ± evenly curved (female pregenital sternite ( Figs 20B, D ) with straight hind margin with slight, rarely completely lacking rounded notch)........................................ 9

9. Aedeagus shaft ( Fig. 23B ) slender, dorsal margin almost straight, spinules present also on the apex.... D. paludosa Melichar

-. Aedeagus shaft ( Fig. 23D ) stout, dorsal margin ± curved, apex smooth and shiny........ D. butzele Guglielmino & Bückle

10. Aedeagus shaft ( Figs 14A, E ) with distinct spinules along its ventral margin, arranged in two rows ( Figs 14B, F ); styles ( Figs 14C, G ) slender with long thin apical portion, denticle basally of mid-length (female pregenital sternite ( Figs 19D, E ) with straight hind margin, in the middle slightly convex)...................................... D. marandica subgroup 11

-. Aedeagus shaft without spinules or spines, smooth ( Fig. 10O ) or ventrally slightly scabrous ( Fig 10C )................ 12

11. Aedeagus ( Fig. 14A ) with ventral margin at mid-length abruptly protruding ventrad, apical portion only with few tiny spines.................................................................................. D. marandica Dlabola

-. Aedeagus ( Fig. 14E ) with ventral margin evenly curved, not protruding ventrad in the middle, spines distinctly present until apex.................................................................................... D. jole sp. nov.

12. Styles ( Figs 28H, K, M ) with robust apical portion, towards apex not or only slightly narrowing; medio-apical margin of genital plates ( Figs 29A, C, D ) rounded (female pregenital sternite ( Figs 20I–L ) with ± deep caudal notch)...................................................................................................... D. heterophyla group 13

-. Styles ( Figs 11A, E ) with thin, slightly curved apical portion; (female pregenital sternite ( Fig. 19A ) straight, often in the middle of its hind margin slightly convex, rarely with a ± distinct notch).............................................. 15

13. Genital plates ( Fig. 29A ) very long, latero-apically completely rounded, lateral margin sinuate; styles very long, in dorsal view ( Fig. 28H ) slightly bent, in median view ( Fig. 28G ) straight from basal apodeme to hook-shaped tip; large species of ca. 3.8–4.2 mm (females ca. 4.8–5.6 mm) with long fore wings ( Figs 27A, B ) (length from shoulder to hind margin ca. 1.7–2.1 mm; ratio sutural length of fore wings/head width 0.75–0.85) (female pregenital sternite ( Fig. 20I ) with deep, acuminate or narrowly rounded notch)..................................................................... D. heterophyla Horváth

-. Genital plates short, latero-apically rounded ( Fig. 29D ), or long with distinct latero-apical angle ( Fig. 29C ); styles in dorsal view ( Figs 28K, M ) distinctly curved; smaller species of ca. 3.2–3.9 mm (females ca. 4.1–4.4 mm) with shorter fore wings ( Figs 27C–E ) (length from shoulder to hind margin less than 1.3 mm; ratio sutural length of fore wings/head width 0.46–0.68)... .................................................................................................. 14

14. Styles in dorsal view ( Fig. 28K ) rather abruptly bent in their median portion, apex not hook-shaped in median view ( Fig. 28I ); aedeagus ( Fig. 28C ) in lateral view apicad distinctly narrowed; genital plates ( Fig. 29C ) long (portion protruding from genital valve longer than wide) with distinct latero-apical angle, lateral margin almost straight; (female pregenital sternite ( Fig. 20K ) with distinct, acuminate notch).......................................................... D. kusnezovi Vilbaste

-. Styles in dorsal view ( Fig. 28M ) evenly curved, apex hook-shaped in median view ( Fig. 28L ); aedeagus ( Fig. 28E ) in lateral view ± of equal width; genital plates ( Fig. 29D ) short (portion protruding from genital valve about as long as wide), latero-apically rounded, lateral margin sinuate; (female pregenital sternite ( Fig. 20L ) with shallow, rounded notch........................................................................................... D. rikele Guglielmino & Bückle

15. Genital plates ( Fig. 41K ) very long, apically rounded, protruding beyond genital segment; styles ( Fig. 41I ) slender, evenly curved (female pregenital sternite ( Fig. 34I ) with hind margin straight or very slightly convex in the median portion)........................................................................................ D. horvathi W. Wagner

-. Genital plates ( Fig. 12B ) not protruding beyond genital segment, apex with (sometimes indistinct) median and/or latero-caudal angles............................................................................................. 16

16. Aedeagus ( Figs 26A, D ) in ventral view proximad evenly widening, not forming an abruptly widening laterally carinate ventroproximal shield; denticle of styles directed more ventrad, almost invisible in dorsal view, styles not or indistinctly geniculate in position of denticle ( Figs 26F–H ) (see also the remarks concerning the vefele -complex at the end of this key)............. .................................................................................................. 17

-. Aedeagus ( Figs 10A, B ) forming an abruptly widening laterally carinate ventro-proximal shield; styles ( Fig. 11A ) generally ± geniculate near denticle, or evenly curved, but then denticle directed more mediad, distinctly visible in dorsal view (see remarks at the end of this key)................................................................................. 18

17. Styles ( Figs 26F, G ) with basal portion distinctly more robust than apical one, lateral apodeme securiform; aedeagus ( Figs 26B, C ) evenly but distinctly narrowing towards apex (female pregenital sternite ( Figs 20F, G ) with ± distinct rounded notch in the middle of hind margin)................................................................. D. gravis Emeljanov

-. Styles ( Fig. 26H ) very slender, evenly narrowing from base to apex, lateral apodeme acuminate; aedeagus, in lateral view ( Fig. 26E ), also in its basal portion quite narrow, apical portion only slightly narrower, in ventral view ( Fig. 26D ) very narrow until basal widening (female pregenital sternite ( Fig. 20H ) with ± straight hind margin)............. D. lukjanovitshi Kusnezov

18. Styles ( Figs 11A, E ) with very long and thin apical portion, central denticle at mid-length or basally of mid-length of style, in most cases directed mediad (hind margin of female pregenital sternite ( Fig. 19A ) medially straight or slightly convex)......................................................................................... D. stylata group 19

-. Styles ( Fig. 41O ) with denticle distinctly apically of mid-length and with short apical portion; proximal shield-shaped portion of aedeagus ( Fig. 41N ) in ventral view with distinct angular latero-distal margin (female pregenital sternite ( Fig. 34L ) with distinct notch).......................................................................... D. lobele sp. nov.

19. Vertex long, roughly triangular ( Figs 7C, D ; 15A, B ), ratio length to width including eyes 0.43–0.5; aedeagus ( Figs 10K, L ; 17B, D ) in ventral view narrow, basal shield rather small, ventral margin of shaft smooth, in lateral view not sinuate (see also the remarks concerning the vefele -complex at the end of this key); fore wings without green tinge.................... 20

-. Vertex proportionally shorter, roughly semi-circular, ratio length to width including eyes 0.37–0.43 ( Fig. 6A ); aedeagus ( Figs 10A, D ) in ventral view wider, basal shield well developed, ventral margin of shaft sometimes scabrous, in lateral view often sinuate (if aedeagus narrow and with straight ventral margin, then body size small and forewings proportionally short).... 21

20. Fore wings long ( Figs 15A, B ), ratio wing length from shoulder to hind margin/width of head ≥ 1.03; large species with body size ≥ 3.85 mm (see also the remarks concerning the vefele -complex at the end of this key)............. D. vefele sp. nov.

-. Fore wings shorter ( Figs 7C, D ), ratio wing length from shoulder to hind margin/width of head ≤ 0.91; small species with body size ≤ 3.25 mm ......................................................................... D. roesle sp. nov.

21. Large species ( Figs 8A, B ) of ca. 3.9–4.5 mm (females ca. 5.3–6.0 mm) with long legs (hind tibia ˃ 2.1 mm), ratio length of hind tibia to head width 1.64–1.78; (length of ovipositor from hind margin of pregenital sternite to ovipositor tip ca. 1.9 mm; nymph V ( Fig. 9I ) brownish with diffuse light parts, abdomen with dark sprinkles)................. D. impudica Horváth

-. Smaller species, ratio length of hind tibia to head width 1.43–1.61; (length of ovipositor from hind margin of pregenital sternite to ovipositor tip ˂ 1.35 mm)............................................................................ 22

22. Species of medium size ( Figs 6A–H ) of ca. 2.9–3.65 mm (females ca. 3.55–4.3 mm) with longer fore wings: Length from shoulder to hind margin ≥ 1.25 mm, ratio length from shoulder to hind margin to width of head 0.95–1.24; ventral margin of aedeagus with or without minute spinules, medio-ventral aedeagus carina often doubled, black ( Fig. 10A ); fore wings generally with green tinge; (nymph V ( Fig. 9A ) with distinct light areas on vertex, pronotum and abdomen; lateral parts of tergites V-VI and median part of tergites VIII-IX light)................................................. D. stylata (Boheman)

-. Small species ( Figs 7A, B ) of ca. 2.5–3.1 mm (females ca. 3.35–4.0 mm) with short fore wings: Length from shoulder to hind margin ≤ 1.05 mm, ratio length from shoulder to hind margin to width of head 0.76–1.02 (specimens from Central Italy and the Peloponnese Peninsula may display somewhat higher values); ventral margin of aedeagus never with spinules, medio-ventral aedeagus carina simple, sharp, ± testaceous ( Fig. 10I ); fore wings without green tinge; (nymph V ( Figs 9B, C ) brownish with vertex, median part of pronotum and median and lateral bands on abdomen light); (see also “Remark 1” in the paragraph concerning D. exilis in “Results”).............................................................. D. exilis Horváth

Remarks. In some cases, specimens belonging to the vefele -complex (in Spain, Morocco, Tunisia, see above) may share a part of the characters presented in the key (guide numbers 16 and 19) as exclusive of other taxa (especially of the D. gravis -group or some members of the D. stylata -group).

Key to females

Preliminary remarks. Only a few characters are available for the identification of Doratura females. Besides several morphometric characters (body length, relative and absolute length of wings, length of tibiae, length of the protruding portion of ovipositor), the morphology of the pregenital sternite furnishes particularly helpful characters. Unfortunately, morphometric characters are quite variable, and as to the pregenital sternites, only some Doratura species display a highly derived shape of the pregenital sternite (e.g., D. medvedevi , Fig. 34D ), whereas in others such specific morphological features are missing. Moreover, for some species the number of specimens we could examine was too low for an estimation of the variation range of their characters.

In view of this situation, the key to females presented below is surely insufficient in many regards, as some taxa cannot be identified, and others only by means of rather indistinct differences, which may turn out completely unsuitable after examination of more material. Nevertheless, it may constitute a first attempt, which in the future hopefully will be improved, corrected and elaborated. It may be useful not only for the identification of isolated females, but for the safe assignment of females to the corresponding males, particularly where two or more species occur in the same area.

1. Hind margin of pregenital sternite straight or slightly concave, always with median, sometimes very shallow notch ( Figs 20B, F, I )................................................................................................ 2

-. Hind margin of pregenital sternite strongly protruding caudad ( Figs 34A, B, D ) or slightly convex in the middle ( Fig. 19A ), or straight ( Fig. 19C ), in the latter case without median notch.................................................... 9

2. Hind margin of pregenital sternite with distinct, V-shaped notch ( Fig. 20I )........................................ 3

- Hind margin of pregenital sternite ( Figs 20B, D ) with shallow or semicircular, often indistinct notch................... 6

3. Large species ( Fig. 27A ) (body length ca. 5.3–5.6 mm; width of head ca. 1.35–1.49 mm) with long forewings (length from shoulder to hind margin ca. 1.9 mm; ratio wing length to width of head ca. 1.3–1.4).............. D. heterophyla Horváth

-. Smaller species (body length ca. 4–5 mm; width of head <1.25 mm) with in proportion shorter forewings (length from shoulder to hind margin 0.86–1.34 mm; ratio wing length to width of head generally ˂1.2)................................... 4

4. Hind margin of pregenital sternite ( Fig. 20K ) with distinct lateral angles, between these angles and median notch slightly concave; ovipositor in lateral view protruding from pygofer ca. 0.53 mm ........................... D. kusnezovi Vilbaste

-. Hind margin of pregenital sternite ( Figs 34H, L ) with rounded lateral angles, between these angles and median notch slightly convex or straight; ovipositor in lateral view protruding from pygofer at most ca. 0.48 mm ........................... 5

5. Hind margin of pregenital sternite ( Fig. 34H ) from lateral angles to median notch obliquely directed medio-rostrad; ovipositor in lateral view protruding from pygofer ca. 0.43–0.48 mm; fore wings proportionally longer: ratio wing length from shoulder to hind margin to width of head generally ˃1; fore wings normally with green tinge................... D. salina Horváth

-. Hind margin of pregenital sternite ( Fig. 34L ) from lateral angles to median notch at most very slightly obliquely directed medio-rostrad; ovipositor in lateral view protruding from pygofer only ca. 0.29–0.38 mm; fore wings proportionally short: ratio wing length from shoulder to hind margin to width of head generally 0.8—0.9; fore wings without green tinge................................................................................................. D. lobele sp. nov.

6. Fore wings longer, ratio length from shoulder to hind margin/wing width rarely ˂ 1.4, generally 1.45–1.55; fore wings normally with distinct green tinge, black bands on face and genae very distinct, dark markings on abdomen less pronounced ( Fig. 25A )..................................................................................... gravis Emeljanov

-. Fore wings shorter, ratio length from shoulder to hind margin/wing width generally 1.1–1.4; fore wings pale, without green tinge, but often with dark longitudinal markings; black bands on face and genae in light specimens indistinct, dark markings on abdomen more pronounced............................................................................. 7

7. Pregenital sternite ( Fig. 20L ) proportionally wider (ratio width/length measured on severed flattened sternite ca. 1.9), less protruding caudad, with latero-caudal margin rounded................................. D. rikele Guglielmino & Bückle

-. Pregenital sternite ( Figs 20B, D ) proportionally narrower (ratio width/length measured on severed flattened sternite ca. 1.6), distinctly protruding caudad, with latero-caudal margin more angular............................................ 8

8. Pregenital sternite ( Fig. 20B ) with median notch shallow, sometimes indistinct, rarely completely lacking (see below, remarks)............................................................................ D. paludosa Melichar

-. Pregenital sternite ( Fig. 20D ) with median notch distinct, ± semicircular................ D. butzele Guglielmino & Bückle

9. Hind margin of pregenital sternite straight ( Figs 19C ), or slightly convex in its median portion ( Fig. 19A ).............. 10

-. Hind margin of pregenital sternite ( Figs 34A, B, D ) strongly protruding caudad.................................. 16

10. Vertex long ( Figs 7C , 15A ), roughly triangular, ratio length to width including eyes 0.43–0.5; fore wings without green tinge. .................................................................................................. 11

-. Vertex proportionally shorter ( Fig. 6E ), roughly semi-circular, ratio length to width including eyes 0.37–0.43........... 12

11. Fore wings long ( Fig. 15A ), ratio wing length from shoulder to hind margin/width of head ≥ 1.03; large species with body size ca. 6 mm .............................................................................. D. vefele sp. nov.

-. Fore wings shorter ( Fig. 7C ), ratio wing length from shoulder to hind margin/width of head ≤ 0.91; small species with body size ≤ 4.25 mm ............................................................................ D. roesle sp. nov.

12. Large species ( Figs 8A , 13A, D ) (body length ca. 5.3–5.8 mm) with long legs (hind tibia 2.1–2.65 mm, ratio hind tibia to width of head generally ≥ 1.6); ovipositor length ventrally from hind margin of pregenital sternite to ovipositor tip ca. 1.9 mm, in lateral view protruding from pygofer generally 0.58–0.67 mm; fore wings with green or blue tinge........................................................................ D. impudica Horváth , D. marandica Dlabola , D. jole sp. nov.

-. Smaller species (body length ˂ 4.4 mm); legs shorter (hind tibia ˂ 1.9 mm, ratio hind tibia to width of head generally ≤ 1.6); ovipositor length ventrally from hind margin of pregenital sternite to ovipositor tip ≤ 1.6 mm, in lateral view protruding from pygofer generally 0.3–0.6 mm .......................................................................... 13

13. Small species ( Fig. 7A ) (body length ≤ 4.0 mm) with short fore wings (length from shoulder to hind margin ≤ 1.05 mm; ratio length from shoulder to hind margin/width of fore wings ca. 1.15–1.35) (specimens from Central Italy and the Peloponnese Peninsula may display somewhat higher values); fore wings without green tinge (see also “Remark 1” in the paragraph concerning D. exilis in “Results”)................................................................. D. exilis Horváth

-. Larger species (body length ca. 3.75—4.4 mm) with longer fore wings (length from shoulder to hind margin ˃ 1.05 mm)... .................................................................................................. 14

14. Slender species (width of head ca. 1.10—1.15 mm) with rather short fore wings (ratio length from shoulder to hind margin/ width ca. 1.1—1.2), hind margin of pregenital sternite straight, never convex in its median portion; fore wings without green tinge, light specimens without markings on median and hind area of vertex, prothorax and scutellum, darker specimens with dark ± distinct marking in sagittal line on vertex and/or prothorax and scutellum, but without dark spots on lateral part of central and posterior portion of vertex; dark marking on anterior part of pregenital tergite often line-shaped........................................................................................................... D. iblea D‘Urso

-. More robust species (width of head rarely ˂ 1.15 mm, generally 1.15—1.25 mm, sometimes ˃ 1.25 mm) with longer fore wings (ratio length from shoulder to hind margin/width ca. 1.25—1.5); fore wings often with green tinge, only very dark specimens with dark marking in sagittal line on vertex and/or prothorax and scutellum...................................... 15

15. Hind margin of pregenital sternite ( Fig. 20H ) straight, rarely with indistinct median notch; ovipositor protruding beyond pygofer tip in lateral view ca. 0.7 mm; length of pygofer in dorsal view ca. 0.4 mm; coloration pale without markings on the middle and posterior portion of vertex.................................................. D. lukjanovitshi Kusnezov

-. Hind margin of pregenital sternite ( Figs 19A , 34I ) with median portion generally distinctly convex, more rarely straight; ovipositor protruding beyond pygofer tip in lateral view generally ˂ 0.6 mm; length of pygofer in dorsal view ca. 0.3–0.35 mm; vertex in dark specimens with distinct spots on its middle and posterior portion... D. stylata (Boheman) , D. horvathi Wagner

16. Pregenital sternite ( Fig. 34D ) with deep incision in the middle of its hind margin.............. D. medvedevi Logvinenko

-. Pregenital sternite without incision on its hind margin....................................................... 17

17. Pregenital sternite ( Figs 34B, C ) in its basal quarter with almost parallel margins, posteriorly narrowing in a truncate triangle with slightly concave lateral margins..................................................... D. rusaevi Kusnezov

-. Pregenital sternite in its posterior portion lobe-shaped with convex lateral margins................................ 18

18. Pregenital sternite ( Fig. 34E ) laterally with angular transition to posterior free portion, posterior portion sub-quadrangular, at its base abruptly narrowing, then protruding posteriad with almost parallel margins, widely rounded at its apex; forewings without green tinge........................................................................ D. homophyla Horváth

-. Pregenital sternite ( Figs 34A, F ) laterally with rounded transition to posterior free portion, posterior portion protruding posteriad with narrowing margins, more sub-triangular, apical margin often somewhat truncated with shallow, indistinct or without median notch....................................................................................... 19

19. Larger species, length of vertex ca. 0.53–0.58 mm, ovipositor protruding from tip of pygofer in lateral view 0.53–0.67 mm; forewings pale, straw-coloured........................................................ D. caucasica Melichar

-. Smaller species, length of vertex ca. 0.43–0.48 mm, ovipositor protruding from tip of pygofer in lateral view ca. 0.4–0.5 mm; forewings with distinct green tinge........................................................ D. concors Horváth

Remarks. The abdomen of the female syntype of D. paludosa as presented in Fig. 20A seems to have a pregenital sternite with strongly concave hind margin and semi-circular median notch. In reality, this is largely due to the position of the sternite still integrated in the ventral part of the abdomen, with strong lateral and slight longitudinal curvature, and in view somewhat obliquely from behind (in order to give the maximum extension of the ovipositor). The isolated sternite (after dissection and under a cover slip) would give a different picture, more similar to the specimen from Manziana ( Fig. 20B ).

Figures

FIGURE 6. Doratura stylata (Boheman). A: male. Czech Republic, Bohemia, Srdov, (MNHN)EH 23391; B: male. Mongolia, Baga-Tenger, 26.7.1965, (MNHN)EH 23452; C: male. Spain, Palencia, Embalse de Requejada, 1100 m, 11.08.1990; D: male. Spain, Palencia, Embalse de Requejada, 1100 m, 11.08.1990 (other specimen); E: female. Spain, Segovia, Vegas de Matute, 950–1050 m, 22.08.1990; F: male. Spain, Cuenca, Laguna de Marquesado, ~1300 m, 25.08.1990; G: female. Spain, Teruel, Gudar, 1400–1500 m, 27.08.1990; H: male. Spain, Teruel, Gudar, 1400–1500 m, 27.08.1990. (Photos Christoph Allgaier).

FIGURE 7. Doratura exilis Horváth, D. roesle sp. nov. A, B: D. exilis. A: female. Germany, Baden-Württemberg, Heidenheim, Iggenhausen, 12.08.2013; B: male. Germany, Baden-Württemberg, Schwarzwald-Baar-Kreis, Mühlhausen, 15.07.2013.—C–E: D. roesle. C: female. Greece, Achaia, Erymanthos, Elliniko, 943 m, 31.07.2017; D: male. Greece, Achaia, Kalavryta, Lousiko, 960 m, 02.08.2017; E: male. Greece, Achaia, Erymanthos, Tsapournia, 1206 m, 31.07.2017. (Photos Christoph Allgaier).

FIGURE 8. Doratura impudica Horváth. A: female. Bulgaria, Sofia, Dragomansko blato, 720 m, 23.06.2012; B: male. Bulgaria, Vlahina Mountains, Boboshevo, 730 m, 30.06.2012; C: male. Syntype D. ivanovi Kusnezov. Ukraine, Kiev surroundings, Holosiivskyi forest; D: female. Syntype D. ivanovi Kusnezov. Ukraine, Kiev surroundings, Holosiivskyi forest. (Photos Christoph Allgaier).

FIGURE 9. Doratura stylata group. Nymphs. A: D. stylata (Boheman). Italy, Friuli-Venezia Giulia, Basovizza, 432 m, 21.07.2016.—B, C: D. exilis Horváth. B: Greece, Thessalia, Larissa, Tempi, Kallipefki, 1120 m, 10.08.2017; C: Greece, Thessalia, Larissa, Tempi, Kallipefki, 1120 m, 10.08.2017 (other specimen).—D–G: D. roesle sp. nov. D: Greece, Achaia, Erymanthos, Elliniko, 943 m, 31.07.2017; E: Greece, Achaia, Kalavryta, Lousiko, 960 m, 02.08.2017; F: Greece, Achaia, Erymanthos, Tsapournia, 1206 m, 31.07.2017; G: Greece, Achaia, Kalavryta, Lousiko, 960 m, 02.08.2017.—H: D. stylata group?. Greece, Corinthia, Killini, Ziria, 1525 m, 04.08.2017.—I: D. impudica Horváth. Italy, Friuli-Venezia Giulia (Udine), Torrente But c/o Sutrio, 526 m, 05.07.2017.—K: D. jole sp. nov. Turkey, Tunceli prov., Ovacik.—L: D. vefele sp. nov. Morocco, Marrakech, Tahanaout, 1000–1050 m, 25.05.1981.—M: D. vefele complex. Tunisia, Beja, 170–220 m, 20.09.1986. (Photos Christoph Allgaier).

FIGURE 10. Doratura stylata group. Aedeagus. A–G: D. stylata (Boheman). A, B: Germany, Bayern, Memmingen; C: Italy, Basilicata, Monte Pollino; D, E: Mongolia, Bulgan, (MNHN)EH 23459; F, G: Spain, Aragón, Huesca, Ansó.—H, I: D. exilis Horváth. Germany, Baden-Württemberg, Gültlingen.—K, L: D. roesle sp. nov. Greece, Achaia, Kalavryta, Lousiko.—M–Q: D. impudica Horváth. M, N: Bulgaria, Sofia, Beledie Han; O: Italy, Basilicata, Matera, Fiume Sinni; P: Germany, Mecklenburg- Vorpommern, Insel Poel; Q: syntype D. ivanovi Kusnezov. Ukraine, Kiev. (A, D, G, I, K, N ventral view; B, C, E, F, H, L, M, O–Q lateral view).

FIGURE 11. Doratura stylata group. Right style, dorsal view. A–C: D. stylata (Boheman). A: Germany, Bayern, Memmingen; B: Mongolia, Bulgan, (MNHN)EH 23459; C: Spain, Aragón, Huesca, Ansó.—D: D. exilis Horváth. Germany, Baden-Württemberg, Gültlingen.—E: D. roesle sp.nov. Greece, Achaia, Kalavryta, Lousiko.—F–I: D. impudica Horváth. F: Bulgaria, Sofia, Beledie Han; G: Italy, Basilicata, Matera, Fiume Sinni; H: Germany, Mecklenburg-Vorpommern, Insel Poel; I: syntype D. ivanovi Kusnezov. Ukraine, Kiev.

FIGURE 12. Doratura stylata group. Genital plate. A–D: D. stylata (Boheman). A, B: Germany, Bayern, Memmingen; C: Mongolia, Bulgan, (MNHN)EH 23459; D: Spain, Aragón, Huesca, Ansó.—E, F: D. exilis Horváth. E: Germany, Baden-Württemberg, Gültlingen; F. Italy, Friuli-Venezia Giulia, Basovizza.—G: D. roesle sp. nov. Greece, Achaia, Kalavryta, Lousiko.—H, I: D. impudica Horváth. H: Bulgaria, Sofia, Beledie Han; I: syntype D. ivanovi Kusnezov. Ukraine, Kiev.—K, L: D. marandica Dlabola. K: Iran, Kandavan-Pass; L: Holotype. Iran, Marand.—M, N: D. jole sp. nov. Turkey, Tunceli, Ovacik. (B–I, L, M: right genital plate, ventral view; A, K, N: left genital plate, dorsal view).

FIGURE 13. Doratura marandica Dlabola, D. jole sp. nov. A, B: D. marandica. A: female. N-Iran, Ziaran, 2400 m, 10.– 16.7.1977; B: male. N-Iran, Kandavan-Pass, 2700–2900 m, 4.–9.7.1977.—C, D: D. jole sp. nov.; C: male. Paratype. Turkey, Tunceli prov., Ovacik; D: female. Paratype. Turkey, Tunceli prov., Ovacik. (Photos Christoph Allgaier).

FIGURE 14. Doratura marandica Dlabola, D. jole sp. nov. A–D: D. marandica. A, B: aedeagus, lateral and ventral view. Holotype. Iran, Marand; C: right style, dorsal view. Iran, Kandavan-Pass; D: right style, dorsal view. Paratype. Iran, Marand.—E–G: D. jole. Turkey, Tunceli, Ovacik. E, F: aedeagus, lateral and ventral view; G. right style, dorsal view.

FIGURE 15. Doratura vefele sp. nov. A: female. Paratype. Morocco, Marrakech, Tahanaout, 1000–1050 m, 25.05.1981; B: male. Paratype. Morocco, Marrakech, Tahanaout, 1000–1050 m, 25.05.1981; C: male. Paratype. Spain, Granada, Huescar, 850–950 m, 16.08.1972; D: female. Paratype. Spain, Granada, Huescar, 850–950 m, 16.08.1972. (Photos Christoph Allgaier).

FIGURE 17. Doratura vefele sp. nov., D. vefele complex. A–L: Aedeagus. A–D: D. vefele. A, B: Morocco, Marrakech, Tahanaout; C, D: Spain, Andalusia, Granada, Huescar.—E–L: D. vefele complex. E, F: Morocco, Fès, Sefrou; G–I: Tunisia, Beja (two different specimens); K, L: Spain,Aragón, Huesca, Bailo. (A, C, F, H, I, L lateral view; B, D, E, G, K ventral view).—M–S: right style. M, N: D. vefele. M: Morocco, Marrakech, Tahanaout; N: Spain,Andalusia, Granada, Huescar.—O–S: D. vefele complex. O, P: Morocco, Fès, Sefrou; Q, R: Tunisia, Beja (two different specimens); S: Spain, Aragón, Huesca, Bailo. (M–O, Q–S dorsal view; P median view).

FIGURE 19. Doratura stylata group. Female pregenital sternite. A: D. stylata (Boheman). Germany, Bayern, Memmingen; B: D. exilis Horváth. Germany, Baden-Württemberg, Gültlingen; C: D. impudica Horváth. Italy, Basilicata, Matera, Fiume Sinni; D: D. marandica Dlabola. Iran, Kandavan-Pass; E: D. jole sp. nov. Turkey, Tunceli, Ovacik; F: D. vefele sp. nov. Spain, Andalusia, Granada, Huescar; G: D. vefele complex. Tunisia, Beja; H: D. roesle sp. nov. Greece, Achaia, Kalavryta, Lousiko.

FIGURE 20. Doratura paludosa group, D. gravis group, D. heterophyla group. Female. A–E: D. paludosa group. A, B: D. paludosa Melichar. A: apical part of abdomen, ventral view. Syntype. Italy, Friuli-Venezia Giulia, Monfalcone; B: pregenital sternite. Italy, Latium, Roma, Manziana. C, D: D. butzele Guglielmino & Bückle. C: apical part of abdomen, ventral view. Paratype. Italy, Apulia, Foggia, Gargano; D: pregenital sternite. Paratype. Italy, Apulia, Foggia, Gargano. E: D. iblea D’Urso. Pregenital sternite. Paratype. Italy, Sicily, Siracusa, Buccheri.—F–H: D. gravis group. F, G: D. gravis Emeljanov. Pregenital sternite. F: Mongolia, Ara-Changaj-district, Ugij Nur; G: Russia, Primorskij Territory, Platonovka. H: D. lukjanovitshi Kusnezov. Pregenital sternite. Mongolia, Bajanchongor, Bogdo.—I–L: D. heterophyla group. I: D. heterophyla Horváth. Pregenital sternite. Bulgaria, North Black Sea Coast, Chernevo; K: D. kusnezovi Vilbaste. Apical part of abdomen, ventral view. Russia, Astrachan, Ulan Chol; L: D. rikele Guglielmino & Bückle. Pregenital sternite. Paratype. Italy, Calabria, Crotone, Caccuri.

FIGURE 23. Doratura paludosa group. Aedeagus. A–C: D. paludosa Melichar. A: holotype Doratura veneta Dlabola. Italy, Veneto, Laguna Veneta; B, C: syntype. Italy, Friuli-Venezia Giulia, Monfalcone.—D–F: D. butzele Guglielmino & Bückle. D, E: paratype. Italy, Apulia, Foggia, Gargano; F: Herzegovina, Trebinje.—G: D. iblea D’Urso. Paratype. Italy, Sicily, Siracusa, Buccheri.—H–L: D. cf. paludosa Melichar. H: Italy, Sicily, Messina, Cesarò; I: Italy, Sicily, Messina, Cesarò (same specimen as in Fig. H); K: Italy, Sicily, Messina, Cesarò (other specimen); L: Italy, Sicily, Messina, Cesarò (3rd specimen). (A, B, D, F, G, I–L lateral view; C, E, H ventral view).

FIGURE 24. Doratura paludosa group. A–H: right style, dorsal view. A–C: D. paludosa Melichar. A: Italy, Latium, Roma, Manziana; B: syntype. Italy, Friuli-Venezia Giulia, Monfalcone; C: holotype Doratura veneta Dlabola. Italy, Veneto, Laguna Veneta. D–G: D. butzele Guglielmino & Bückle. D: paratype. Italy, Apulia, Foggia, Gargano; E: Herzegovina, Trebinje; F: Italy, Veneto, Lago di Garda; G: Italy, Sicily, Etna, Citelli. H: D. iblea D’Urso. Paratype. Italy, Sicily, Siracusa, Buccheri. I–M: right genital plate, ventral view. I, K: D. paludosa Melichar. I: syntype. Italy, Friuli-Venezia Giulia, Monfalcone; K: holotype Doratura veneta Dlabola. Italy, Veneto, Laguna Veneta. L, M: D. butzele Guglielmino & Bückle. L: paratype. Italy, Apulia, Foggia, Gargano; M: Herzegovina, Trebinje.—N: left genital plate, dorsal view. D. butzele Guglielmino & Bückle. Italy, Sicily, Etna, Citelli.

FIGURE 25. Doratura gravis group, D. heterophyla group.A, B: D. gravis group.A: D. gravis Emeljanov. Male. Russia, Maritime Territory, 14.07.1966, (MNHN)EH 23462; B: D. lukjanovitshi Kusnezov. Male. Syntype. Russia, Transbaikalia, Dzhida, Enchor, 20.07.1928.—C, D: D. heterophyla group. Nymph; C: D. heterophyla Horváth. Bulgaria, Strouma valley, Kozhuh Hill, Rupite, 100 m, 01.07.2012; D: D. rikele Guglielmino & Bückle. Italy, Calabria (Crotone), Petilia, 509 m, 06.07.2011. (Photos Christoph Allgaier).

FIGURE 26. Doratura gravis group. A–C: D. gravis Emeljanov. A, B: aedeagus, ventral and lateral view. North Korea, Pukchŏng, Hamgyŏng-Namdo; C: aedeagus, lateral view. Paratype D. jurtica Dlabola. Mongolia, Bajan-Agt.—D, E: D. lukjanovitshi Kusnezov. Aedeagus, ventral and lateral view. Syntype. Russia, Transbaikalia, Dzhida, Enchor.—F, G: D. gravis. Right style, dorsal view. F: North Korea, Pukchŏng, Hamgyŏng-Namdo; G: paratype D. jurtica Dlabola. Mongolia, Bajan-Agt.—H: D. lukjanovitshi. Right style, dorsal view. Syntype. Russia, Transbaikalia, Dzhida, Enchor.—I–L: D. gravis. I, K: right genital plate, ventral view. I: North Korea, Pukchŏng, Hamgyŏng-Namdo; K: paratype D. jurtica Dlabola. Mongolia, Bajan-Agt. L: left genital plate, dorsal view. Paratype D. jurtica Dlabola. Mongolia, Bajan-Agt.—M: D. lukjanovitshi. Right genital plate, ventral view. Syntype. Russia, Transbaikalia, Dzhida, Enchor.

FIGURE 27. Doratura heterophyla group. A, B: D. heterophyla Horváth. Bulgaria, Strouma valley, Kozhuh Hill, Rupite, 100 m, 01.07.2012. A: female; B: male.—C: D. kusnezovi Vilbaste. Male. Kazakhstan, Karagand, 10.06.1961.—D, E: D. rikele Guglielmino & Bückle. Paratypes. Italy, Calabria (Crotone), Caccuri, 581 m, 05.07.2011. D: female; E: male. (Photos Christoph Allgaier).

FIGURE 28. Doratura heterophyla group. A, B: D. heterophyla Horváth. Aedeagus, ventral and lateral view. Bulgaria, Strouma valley, Kozhuh Hill, Rupite.—C, D: D. kusnezovi Vilbaste. Aedeagus, lateral and ventral view. Russia, Astrachan, Ulan Chol.—E, F: D. rikele Guglielmino & Bückle. Aedeagus, lateral and ventral view. Paratype. Calabria, Crotone, Caccuri.—G, H: D. heterophyla Horváth. Right style, median and dorsal view. Bulgaria, Strouma valley, Kozhuh Hill, Rupite.—I, K: D. kusnezovi Vilbaste. Right style, median and dorsal view. Russia, Astrachan, Ulan Chol.—L–N: D. rikele Guglielmino & Bückle. L, M: right style, median and dorsal view. Paratype. Italy, Calabria, Crotone, Caccuri; N: right style, dorsal view. Paratype. Italy, Calabria, Crotone, Caccuri (other specimen).

FIGURE 29. Doratura heterophyla group.A, B: D. heterophyla Horváth.A: right genital plate, ventral view. Bulgaria, Strouma valley, Kozhuh Hill, Rupite; B: left genital plate, dorsal view. Bulgaria, Strouma valley, Kozhuh Hill, Rupite.—C: D. kusnezovi Vilbaste. Right genital plate, ventral view. Russia, Astrachan, Ulan Chol.—D: D. rikele Guglielmino & Bückle. Right genital plate, ventral view. Paratype. Italy, Calabria, Crotone, Caccuri.

FIGURE 31. Doratura concors group.Aedeagus.A–D: D. concors Horváth.A, B: Russia, Altai, Čibit, Čuja; C, D: Montenegro, Nikšić, Vidrovan.—E–L: D. rusaevi Kusnezov. E–G: Kazakhstan, Karaganda, Zhana-Arka; H, I: Russia, Astrachan, Sör-Kata; K, L: Kazakhstan, Akmolinsk.—M–R: D. medvedevi Logvinenko. M–O: West-Kazakhstan, Saykhin; P, Q: Russia (?), Caspian Sea coast; R: Ukraine, Crimea, Kertsh Peninsula. (A, C, E, H, K, M, Q, R lateral view; B, D, G, I, L, O, P ventral view; F: Apex in apical view in direction of arrow in Fig. E; N: Apex in apical view in direction of arrow in Fig. M).

FIGURE 32. Doratura concors group. Right style, dorsal view. A, B: D. concors Horváth. A: Russia, Altai, Čibit, Čuja; B: Montenegro, Nikšić, Vidrovan.—C–E: D. rusaevi Kusnezov. C: Kazakhstan, Karaganda, Zhana-Arka; D: Kazakhstan, Akmolinsk; E: Russia, Astrachan, Sör-Kata.—F, G: D. medvedevi Logvinenko. F: Russia (?), Caspian Sea coast. G: West-Kazakhstan, Saykhin.

FIGURE 33. Doratura concors group. Genital plate.A–C: D. concors Horváth.A: Russia,Altai, Čibit, Čuja; B, C: Montenegro, Nikšić, Vidrovan.—D–G: D. rusaevi Kusnezov. D, E: Russia, Astrachan, Sör-Kata; F: Kazakhstan, Akmolinsk; G: Kazakhstan, Karaganda, Zhana-Arka.—H, I: D. medvedevi Logvinenko. H: West-Kazakhstan, Saykhin; I: Russia (?), Caspian Sea coast. (A, C, D, F–H: Right genital plate, ventral view; B, E, I: Left genital plate, dorsal view).

FIGURE 34. Doratura concors group, D. homophyla group, D. salina, D. horvathi, D. lobele. Females.A–D: D. concors group. A: D. concors Horváth. Pregenital sternite. Montenegro, Nikšić, Vidrovan; B, C: D. rusaevi Kusnezov.Apical part of abdomen, ventral view. B: Russia, Astrachan, Sör-Kata; C: Kazakhstan, Karaganda, Zhana-Arka; D: D. medvedevi Logvinenko. Apical part of abdomen, ventral view.—E, F: D. homophyla group. E: D. homophyla (Flor). Pregenital sternite. Germany, Baden-Württemberg, Baden-Baden, Sandweier; F: D. caucasica Melichar. Pregenital sternite. Azerbajan, Talysch, Alekseevka.—G, H: D. salina Horváth. G: Apical part of abdomen, ventral view. Romania, Cluj, Someşeni; H: pregenital sternite. Romania, Constanţa, Techirghiol.—I: D. horvathi W. Wagner. Pregenital sternite. Germany, Niedersachsen, Lengder Burg.—K, L: D. lobele sp. nov. Greece, Arcadia, Tripoli, Kandalos. K: apical part of abdomen, ventral view; L: pregenital sternite.

FIGURE 37. Doratura homophyla group. Aedeagus. A–E: D. homophyla (Flor). A: Germany, Mecklenburg-Vorpommern, Insel Poel; B, C: Germany, Baden-Württemberg, Baden-Baden, Sandweier; D: Spain, Andalusia, Jaén, Cazorla; E: Irak, Bagdad.—F–I: D. caucasica Melichar. F, G: Paratype (?) D. arenicola Logvinenko. Azerbajan, Lenkoran, Alekseevka, 01.06.1967; H, I: Georgia, Tbilisi, Lac Lisi. (A, C, D, E, F, H lateral view; B, G, I ventral view).

FIGURE 39. Doratura homophyla group. Genital plate. A, B: D. homophyla (Flor). Germany, Baden-Württemberg, Baden- Baden, Sandweier.—C–F: D. caucasica Melichar. C: paratype (?) D. arenicola Logvinenko. Azerbajan, Lenkoran, Alekseevka, 01.06.1967; D: paratype D. arenicola Logvinenko. Azerbajan, Lenkoran, Talysh, 12.07.1970; E, F: Georgia, Tbilisi, Lac Lisi. (A, C, E: right genital plate, ventral view; B, D, F: left genital plate, dorsal view).

FIGURE 41. Doratura salina Horváth, D. horvathi W. Wagner, D. lobele sp. nov. A–F: D. salina. A, B: aedeagus, ventral and lateral view. Romania, Cluj, Cojozna; C: aedeagus, rostral view (in direction of arrow in Fig. B). Ukraine, Charkow, Zelenino; D: right style, dorsal view. Romania, Cluj, Cojozna; E: right genital plate, ventral view. Romania, Cluj, Someşeni; F: left geni- tal plate, dorsal view. Romania, Constanţa, Techirghiol.—G–L: D. horvathi. Germany, Niedersachsen, Lengder Burg. G, H: aedeagus, ventral and lateral view; I: right style, dorsal view; K: right genital plate, ventral view; L: left genital plate, dorsal view.—M–Q: D. lobele. M, N: aedeagus, lateral and ventral view. Greece, Arcadia, Tripoli, Kandalos; O: right style, dorsal view. Greece, Arcadia, Tripoli, Kandalos; P: left genital plate, dorsal view. Greece, Messenia, Kalamata, Dyrrachio; Q: right genital plate, ventral view. Greece, Arcadia, Tripoli, Kandalos.