Tachysphex austriacus Kohl, 1892 and T. pompiliformis (Panzer, 1804) (Hymenoptera, Crabronidae) are a complex of fourteen species in Europe and Turkey

Abstract Tachysphex pompiliformis (Panzer, 1804) and Tachysphex austriacus Kohl, 1892 species subgroups belong to the Tachysphex pompiliformis species group, and both subgroups consist of morphologically similar species. The Tachysphex austriacus Kohl species subgroup comprises four species in Europe and Turkey. For this subgroup, differential diagnoses of Tachysphex austriacus and Tachysphex prismaticus Straka, 2005 are presented, and Tachysphex hungaricus sp. n. from Hungary and Tachysphex smissenae sp. n. from Spain, France and Turkey are described. The Tachysphex pompiliformis species subgroup consists of ten species from Europe and Turkey. For this subgroup, differential diagnoses of Tachysphex ferrugineus Pulawski, 1967 and Tachysphex opacus F. Morawitz, 1893, as well as the differential diagnosis and redescription of Tachysphex pompiliformis, are presented. Tachysphex dimidiatus (Panzer, 1809), Tachysphex jokischianus (Panzer, 1809) and Tachysphex nigripennis (Spinola, 1808) are resurrected from synonymy with Tachysphex pompiliformis and redescribed. Neotypes of Tachysphex dimidiatus (Panzer) and Tachysphex nigripennis (Spinola) are designed. Tachysphex bohemicus sp. n. from the Czech Republic; Tachysphex cretensis sp. n. from Crete, Greece; Tachysphex nobilis sp. n. from Bulgaria, the Czech Republic, Hungary, Poland, Slovakia and Turkey; and Tachysphex punctipleuris sp. n. from Bulgaria, Germany, Hungary, Italy, Slovenia and Turkey are described. Identification keys to all species from Tachysphex pompiliformis and Tachysphex austriacus species subgroups known from Europe and Turkey are presented.


Introduction
Tachysphex pompiliformis (Panzer, 1804) and T. austriacus Kohl, 1892 species subgroups belong to the T. pompiliformis species group. Tachysphex austriacus Kohl has been considered to be a sister species to T. pompiliformis (Panzer) (Straka 2004). Later, T. austriacus Kohl, 1892 was recognized to be a cluster of closely related species different from T. pompiliformis (Panzer). Both species are now considered to belong to separate lineages and form two different species subgroups (Straka 2008). Only three species that were placed in the T. pompiliformis subgroup were recognized in the previous revision of Tachysphex in the Palearctic region (Pulawski 1971). Species from the T. austriacus subgroup were thought to be conspecific with T. pompiliformis. Most of the species from the T. austriacus subgroup were recently described or restored from synonymy. First, T. austriacus Kohl was resurrected and redescribed by Straka (2004). Subsequently, two other species were described and were considered to be related to T. austriacus Kohl: T. prismaticus Straka, 2005 and T. stysi Straka, 2008(Straka 2005, 2008. Two new species that belong to the T. austriacus subgroup are described in the present study. In a revision of the Palearctic species (Pulawski 1971), only T. pompiliformis (Panzer), T. ferrugineus Pulawski, 1967, and T. opacus F. Morawitz, 1893 were recognized as valid; all other names from respective groups were considered to be junior synonyms. Subsequently, one additional species, T. kaszabi Tsuneki, 1972 from Mongolia, was described (Tsuneki 1972). Pulawski (1971) presented information on the extensive variability within T. pompiliformis (Panzer) species; however, no species recognition or change in taxonomy was made within this variable species until this time, except for the separation of species of the T. austriacus subgroup. In this study, ten species that belong to this subgroup are recognized in Europe and Turkey. Three species names are restored from synonymy, and four additional new species are described. Morphological terms are used according to Bohart and Menke (1976) and Krombein and Pulawski (1994), except morphological description of volsella. There is volsellar corpus -the main part with setae on outer margin and a depression on inner side, which connecting volsella with gonocoxites; apical volsellar process -a thin process which bears setae on its outer margin, a row of setae continues from the corpus; dorsal volsellar process -the process, which can have various shapes and extending dorsally from the corpus. The following abbreviations are used in the morphological descriptions:

PTLC
WML clypeus median lobe width LCL clypeus maximum length WCL clypeus width LA3 length of antennal article III, dorsally WA3 width of antennal article III, dorsoapically LA5 length of antennal article V, dorsally WA5 width of antennal article V, dorsoapically WV vertex width LV vertex length MOD diameter of median ocellus LF1 forefemur length WF1 forefemur width, laterally All of the newly described species were labeled in the following manner: "HOLO-TYPUS ♂ or ♀, name of taxon sp. nov., J. Straka det. 2015" on a red card; neotypes were designated on red cards, and paratypes were designated on yellow cards. Exact label data are cited only for the holotypes. Separate lines on the labels are indicated with a slash "/", and separate labels are indicated with double slashes "//".

Tachysphex austriacus species subgroup
Diagnosis of subgroup. The Tachysphex austriacus species subgroup belongs to the T. pompiliformis group, which is characterized by the following character combination: La-brum flat; frons uniformly convex; scutum and mesopleuron finely reticulate or punctate, microsculptured or unsculptured among punctures; episternal ridge developed; tarsomeres V of mid and hind legs unmodified, straight ventrally and mildly convex dorsally; tarsomeres IV of the mid and hind legs distinctly emarginated and longer than wide; anal lobe of the hindwing small; sternum I without a longitudinal carina; sterna without distinct setal patches; pygidium triangular, narrowly truncate at the apex, with indistinct transverse carina, integument sparsely punctate, or without punctures in some species, interspaces between punctures finely sculptured to unsculptured. In males, the volsellar corpus is not separated from the dorsal process by constriction, and thus the corpus is extended to a process; aedeagus with distinct well-developed teeth, usually not more than 10 in number.
From the other species of the T. pompiliformis group, the T. austriacus species subgroup (species complex sensu Straka 2005) differs in having the following characters: glossa and galea short, galea as wide as long, apex densely setose; glossa shorter than galea; mesopleuron microsculptured, dull or slightly shiny, with or without punctures, punctures ill-defined when present, and with distinct interspaces; fore and midfemora uniformly sculptured throughout, with only small punctures and uniform interspaces; lateral parts of tergum II with variably sparse punctuation and shiny interspaces; and terga I-III with silvery apical fasciae (rarely absent). In the female, clypeus with lateral incisions and with shallow or absent median emargination. In the male, clypeus arcuate with conspicuous lip corner and forebasitarsus without rake. The T. austriacus species subgroup differs from the T. pompiliformis species subgroup by two consistent male characters. In the T. austriacus species subgroup, the forefemoral notch, slightly shiny to shiny, has no elevated plate and carinae on the margins and has distinct small setae on the surface; the setae on the volsellar apical process are uniformly directed ventrally (or nearly so). In the T. pompiliformis species subgroup, an elevated plate is present in the forefemoral notch with more or less distinct carinae on the margins; its surface appears nearly glabrous, finely or coarsely microsculptured; the ventral setae on the volsellar apical process are randomly directed. Characters for distinguishing females of T. pompiliformis and T. austriacus subgroups have not been identified so far.
Metasoma. Terga I-III with slightly developed, but distinct silvery apical fasciae, densely, very finely micropunctate, punctures ill-defined, evanescent in microsculpture, interspaces aciculate, slightly shiny; apical depressions shallow with indistinct micropunctures; sculpture of terga VI-V distinctly coarser than on previous terga, punctures denser, ill-defined, but large, one to less than half diameter apart, interspaces microsculptured, apical depression of tergum V coarsely microsculptured, with few scattered punctures. Pygidium of usual size, sparsely punctate, large and minute punctures intermixed, ill defined, but large punctures deep, interspaces nearly unsculptured, shiny. Central part of sternum II with several larger punctures, interspaces microsculptured, shiny; lateral part slightly shiny, densely micropunctate; remaining sterna with uniform sculpture similar to that on sternum II, but more or less reduced laterally.
Coloration. Large area in center of mandibles, terga and sterna I-III and base of tergum and sternum IV red. Distal tarsomeres, apex of pronotal lobe, ventral part of distal antennal segments, apex of clypeal bevel and tip of pygidium dark red. Tegulae reddish translucent. Apical parts of terga I-III slightly translucent. Remaining body parts all black.
Geographic distribution. Known only from Hungary. Note. Male unknown. Name derivation. The species is named after the country of origin. Straka, 2005 Type material. Holotype: ♂, Kazakhstan: "Kazakhstan 20km / SE Aksay env / 16. -19. 6. 1992 / leg.K.Denes", printed label. Holotype in OLML, examined. Diagnosis. Tachysphex prismaticus Straka resembles T. austriacus Kohl, T. hungaricus sp. n., and T. smissenae sp. n. in the shape of the clypeus and well developed punctures on the ventral part of the mesopleuron. It differs from the other species of the subgroup in the following character combination: ♂, ♀: mesopleuron rugose to densely punctate; venter of all trochanters with punctures several diameters apart, their interspaces unsculptured; all femora and tibiae black; ♂: volsella with large dorsal process; ventral volsellar setae in one line; ♀: clypeal lip with distinct irregular median emargination; clypeal bevel forming flat plate, separated from basomedian clypeal area by relatively sharp, more or less angulated transition; scutum and scutellum sparsely punctate, punctures up to two diameters apart at scutum center; forebasitarsal rake with four apical spines.

Tachysphex prismaticus
Geographic distribution. Kazakhstan, Kyrgyzstan and Turkey. Note. Full description and all material is given in Straka (2005).
Mesosoma. Scutum without distinct anterior impression; scutum and scutellum densely punctate, punctures well defined, most punctures less than half diameter apart, interspaces distinct, unsculptured, slightly shiny, setae about 1.0 × MOD or less. Mesopleuron coarsely punctate to rugose, punctures well defined in lower part and ventrally, but indistinct dorsally, interspaces small to absent, slightly shiny; hypoepimeral area densely punctate to uniformly rugose, dull. Propodeal dorsum coarsely sculptured with irregular longitudinal ridges, uniformly rugose to areolate between ridges; propodeal side longitudinally ridged, ridges well developed, interspaces slightly shiny; posteromedial margin of propodeal dorsum slightly elevated, marginal ridges nearly parallel above groove on posterior side. Legs densely punctate, punctures small; forefemoral notch small, but deep, nearly semicircular, shorter than distance that separates it from forefemoral base, central part of notch without microsculpture, with distinct small setae, shiny. Wings not infumated, with brown veins.
Coloration. Apical part of mandibles and terga I and II red. Distal tarsomeres, forefemoral notch, sternum I and base of sternum II dark red. Tegulae reddish translucent. Apical parts of all terga slightly translucent. Remaining body parts black.
Variation of males. Body length: 6.5-7.0 mm. Clypeus slightly to conspicuously convex medially. Forefemoral notch and all metasoma dark in some specimens.
Mesosoma. Scutum without distinct anterior impression; scutum and scutellum densely punctate, punctures well defined, uniform in size, about half diameter apart (up to one diameter apart in some specimens), interspaces uniform in size, slightly shiny, setae about 0.5 × MOD or less. Mesopleuron coarsely punctate to rugose, punctures well defined in lower parts and ventrally, but indistinct dorsally, interspaces small to absent, slightly shiny; hypoepimeral area densely punctate, with punctures ill-defined, appearing uniformly rugose, dull; ventral part of mesopleuron with punctures well defined, interspaces shiny. Propodeal dorsum coarsely sculptured, uniformly rugose to areolate, with irregular longitudinal ridges; side longitudinally ridged, ridges well developed, interspaces slightly shiny; posteromedial margin of dorsum slightly elevated, marginal ridges nearly parallel, but turning ventrally and directed toward groove on posterior surface. Legs, including trochanters, densely punctate, punctures small; forebasitarsal rake pale yellowish, with reddish iridescence, with three apical spines, one preapical spin and two or three additional spines. Wings slightly yellowish, veins brown.
Metasoma. Terga I-III with silvery apical fasciae; densely and very finely micropunctate, punctures ill defined, evanescent in microsculpture, interspaces very finelly microsculptured, slightly shiny; apical depressions shallow, with indistinct micropunctures; sculpture of terga IV-V distinctly coarser than on previous terga, also punctures denser; terga IV-V coarsely punctate, punctures ill-defined, but large, one to less than half diameter apart, interspaces microsculptured, apical depression of terga IV and V coarsely microsculptured, apical depression of tergum IV very finely punctate, apical depression of tergum V with a few scattered punctures, slightly shiny, but coarsely microsculptured and dull in some specimens. Pygidium relatively wide, sparsely punctate, with large and minute punctures intermixed, large punctures well defined, deep, interspaces nearly unsculptured, shiny. Central part of sternum II with several larger punctures, interspaces microsculptured, shiny; lateral part slightly shiny, densely micropunctate; remaining sterna with uniform sculpture similar to that on sternum II, but more or less reduced laterally.
Coloration. Central part of mandibles and terga and sterna I-II red. Distal tarsomeres dark reddish. Tegulae brown translucent. Apical parts of terga I-III slightly translucent. Remaining body parts black.
Geographic distribution. France, Spain, Turkey. Name derivation. The species is named in honor of the outstanding entomologist and collector of most of the type specimens, Jane van der Smissen.

Tachysphex pompiliformis species subgroup
Diagnosis of subgroup. The Tachysphex pompiliformis species subgroup differs from the T. austriacus species subgroup by two male characters. In the T. pompiliformis species subgroup, the surface of the forefemoral notch is microsculptured (very finely in some species), slightly elevated, with a distinct carina on the inner margin, sometimes on the outer margin as well; and the setae on the lower surface of the forefemoral notch are hardly visible (magnification 50 ×). The ventral setae on the volsellar apical process are randomly directed. In the T. austriacus species subgroup, the forefemoral notch is slightly shiny to shiny, without an elevated plate and carinae on the margins and with distinct small setae on the surface; the setae on the volsellar apical process are uniformly directed ventrally (or nearly so). Characters for distinguishing females of T. pompiliformis and T. austriacus subgroups have not been identified so far. See the characterization under the diagnosis of the T. austriacus species subgroup.
Species included. Tachysphex bohemicus sp. n.; T. cretensis sp. n.; T. dimidiatus (Panzer, 1809); T. ferrugineus Pulawski, 1967;T. jokischianus (Panzer, 1809); T. kaszabi Tsuneki, 1972  Diagnosis. Tachysphex bohemicus sp. n. is most similar to T. jokischianus in having nearly impunctate apex of tergum V and wide pigydium with shiny surface. It also resamble T. dimidiatus in the overall sparsely micropunctate metasoma, which makes the appearance of metasomal sculptures more shiny than in other compared species from the T. pompiliformis species subgroup. The apex of the male volsella of the new species is sparsely setose and may resemble volsella of males from the T. austriacus species subgroup. Both sexes of T. bohemicus sp. n. are easily recognizable by a slightly prominent supraantennal tubercles, which are separated by punctate area; the tubercle is distinctly punctate on the top and nearly at all sides. This character is unique in the T. pompiliformis species subgroup. Like T. jokischianus, the apical depression of tergum V is nearly impunctate medially, but the terga are finely and sparsely punctate and the pygidium is basally microsculptured; the metasoma thus resembles that of T. dimidiatus, except for the apical depressions of all terga, which are very sparsely punctate and well impressed. Tachysphex bohemicus sp. n. possess an unusual combination of characters: the propodeum lack longitudinal ridges dorsally and laterally, the female clypeus is distinctly arcuate at apex and uniformly convex dorsally, and the male forefemoral notch is black, positioned more anteriorly than in the other species of the subgroup and its proximal margin is rounded and elevated over the distal margin.
Metasoma. Silvery apical fasciae developed faintly on terga I-IV. Apical depressions of terga distinct, sparsely micropunctate, nearly like more anterior parts. Terga I-IV sparsely micropunctate, punctures ill defined, interspaces coarsely microsculptured, slightly shiny to dull; sculpture of terga IV-VII slightly more conspicuous than that of other terga, also densely punctate. Sterna uniformly punctate, nearly like terga. Volsella light brown, ventral setae pointing in various directions in medial part, but almost in one direction at apical process; apical process relatively sparsely setose; dorsal process large, wider than corpus; corpus lined with distinct ledge along base of setae on inner side of corpus.
Coloration. Apical part of mandibles, distal tarsomeres and terga I-III red. Forefemoral notch black. Tegulae reddish translucent. Apical parts of all terga slightly translucent. Remaining body parts black.
Description of female. Body length: 10.3 mm.
Metasoma. Terga I-III with slightly developed but distinct silvery apical fasciae. Apical depressions of all terga well developed, very sparsely micropunctate. Terga I-III sparsely micropunctate, punctures ill-defined but distinct, interspaces microsculptured, slightly shiny; sculpture of tergum IV-V distinctly coarser than on previous terga, also slightly more densely punctate. Pygidium sparsely punctate, punctures variable, some ill defined, some well defined, but shallow, interspaces microsculptured medially, unsculptured laterally, slightly shiny to shiny. Central part of sternum II with several larger punctures, interspaces microsculptured, shiny; lateral part slightly shiny, densely micropunctate; remaining sterna with uniform sculpture similar to that on sternum II, but more or less reduced laterally. Diagnosis. Tachysphex cretensis sp. n. is most similar to T. pompiliformis and T. opacus. They share a coarsely sculptured mesopleuron and the shape of the mandible is as in the latter species. The species is well recognizable by a coarsely sculptured dark body, four metasomal segments with silvery apical fasciae, the clypeal lip produced medially, the mandibular inner margin shallowly emarginated distally from the inner tooth, lacking any furrow, and a densely microsculptured forefemoral notch. The new species is probably endemic to the island of Crete, Greece.
Mesosoma. Scutum without anterior impression; scutum and scutellum densely punctate, punctures well defined, most punctures less than half diameter apart, interspaces distinct, microsculptured, slightly shiny, setae about 1.0 × MOD or less. Mesopleuron rugose to densely punctate, most punctures in ventral half well defined, interspaces coarsely microsculptured to rugose, slightly shiny to dull; hypoepimeral area coarsely rugose, without distinct punctures, with irregular longitudinal ridges, dull; ventral part of mesopleuron with punctures large, relatively well defined, interspaces finely microsculptured, slightly shiny. Propodeal dorsum coarsely sculptured, with longitudinal ridges; side longitudinally ridged, ridges well developed, interspaces microsculptured, slightly shiny; posteromedial margin of dorsum insignificantly elevated, marginal ridges evanescent, directed ventromedially. Legs densely punctate, punctures small; forefemoral notch small but relatively deep, semicircular, shorter than distance that separates it from forefemoral base, proximal margin relatively sharp, but not distinctly elevated over distal margin; central part of notch slightly elevated, anterior and posterior margin lined by faint ledge, notch surface without distinct setae, coarsely microsculptured, dull. Wings moderately infumate, with brown veins.
Metasoma. Silvery apical faciae of terga I-IV distinctly developed. Apical depressions of all terga shallow, nearly as densely micropunctate as more anterior parts. Terga I-III densely and distinctly micropunctate, punctures ill defined, interspaces microsculptured, slightly shiny; sculpture of terga IV-VII coarser than on previous terga, also puncture denser. Sterna uniformly punctate, nearly like terga. Volsella brown, ventral setae pointing in various directions; apex with numerous setae; dorsal process about as wide as corpus.
Coloration. All body black, except central part of mandibles and terminal tarsomeres dark red. Tegulae brown translucent. Apical parts of all terga slightly translucent.

Tachysphex dimidiatus (Panzer, 1809), restored from synonymy
The gena is robust. The dorsolateral cuticular projection of the propodeal spiracle is slightly arcuate to nearly straight, with the apex reddish transparent. The posteromedial margin of the propodeal dorsum is elevated and produced between the marginal ridges, the marginal ridges directed ventromedially toward the groove on the posterior surface. The propodeal side is longitudinally ridged; ridges anteriorly inconspicuous or absent. The propodeal dorsum often has inconspicuous, irregular longitudinal ridges. The punctures of the mesopleuron are exceptionally variable but usually small and inconspicuous. The terga are sparsely punctate, the punctures are ill defined. Apical depressions of tergum V densely microsculptured with micropunctures, densely punctate latereally. The pygidium is of usual size and distinctly microsculptured in most specimens. ♂: The clypeus is convex, conspicuously elevated in the basal half, continuously declining apically, the top at the clypeal midlength or in the basal half of the clypeus. The clypeal bevel is well developed, shiny. The gena is robust. The supraclypeal area is flat, the supraantennal tubercles are connected, slightly elevated ventromedially. The mesopleuron is distinctly punctate laterally and ventrally, the punctures are ill defined. When the interspaces between the punctures are distinct, then they are microsculptured, slightly shiny to dull. The forefemoral notch is relatively shallow, as long as the distance that separates it from the forefemoral base, central part slightly elevated and lined by a ledge on the anterior as well as the posterior margin. The propodeal side is variable but without basal ridges in most specimens. The posteromedial margin of the propodeal dorsum is elevated and produced between marginal ridges, which are directed ventromedially toward the groove on the posterior surface. The terga are sparsely micropunctate with variable interspaces between punctures.
Mesosoma. Scutum without distinct anterior impression; scutum and scutellum densely punctate, punctures well defined, most punctures less than one diameter apart, interspaces distinct, unsculptured, shiny, setae about 1.0 × MOD. Mesopleuron coarsely microsculptured, with or without distinct punctures; hypoepimeral area irregularly microsculptured, impunctate; ventral part of mesopleuron with punctures ill defined, but deep in some specimens, less than one diameter apart, interspaces shiny. Propodeal dorsum relatively finely, irregularly rugose, with irregular longitudinal ridges basally in some specimens; propodeal side irregularly longitudinally ridged, ridges variable, usually faint, or absent basally, microsculptured, dull; posteromedial margin of propodeal dorsum elevated and produced between marginal ridges, marginal ridges directed ventromedially toward groove on posterior surface. Legs densely punctate, punctures small; forefemoral notch relatively shallow, as long as distance that separates it from forefemoral base to proximal margin of notch, central part slightly elevated and lined by ledge on anterior as well as posterior margin, surface without distinct setae, microsculptured, dull. Wings slightly infumate with brown veins.
Coloration. Apical part of mandibles, tarsi, forefemoral notch, tegulae, terga and sterna I, II, and partly or all tergum III red. Apical parts of all terga and tegula translucent. Remaining body parts all black. Forefemoral notch very rarely black in specimens from high altitude mountains.
Metasoma. Terga I-III with silvery apical fasciae. Apical depressions of all terga shallow, with micropunctures evanescent in microsculpture. Terga I-III sparsely and finely micropunctate, punctures ill defined, interspaces microsculptured, shiny to slightly shiny; sculpture of tergum IV-V slightly coarser than on previous terga, also slightly more densely punctate. Pygidium sparsely punctate, punctures ill defined, interspaces microsculptured, slightly shiny. Central part of sternum II with several larger punctures, interspaces microsculptured, shiny; lateral part slightly shiny, densely micropunctate; remaining sterna with uniform sculpture similar to that on sternum II, but sculpture more or less reduced laterally.
Coloration. Central part of mandibles, three distal tarsomeres, terga and sterna I-III red. Tegulae brown translucent. Apical parts of terga I-III slightly translucent. Remaining body parts all black.
Variation of females. Extremely variable in most of characters. The clypeus is typically conspicuously elevated basally, with the top in the basal third; the bevel can be well separated from the clypeal base in some specimens, althouth infrequently. Vertex wider than long in large specimen and slightly wider than long in small specimens, WV:LV = 1.2-1.5. Antenna short, LA3:WA3 = 2.2-2.4, LA5:WA5 = 2.6-2.7.
Geographic distribution. Austria, Czech Republic, Bulgaria, France, Germany, Italy, Sweden and Switzerland. Specimens from mountains of Greece, Morocco and Spain probably also belong to this species. Diagnosis. Tachysphex ferrugineus Pulawski is easily recognizable by the body coloration, but in morphological characters and body sculpture it resembles T. dimidiatus (Panzer), T. jokischianus (Panzer), T. pompiliformis (Panzer) and T. punctipleuris sp. n. The species is similar to T. stysi Straka from central Asia in coloration, but T. stysi belongs to the T. austriacus species subgroup. Tachysphex ferrugineus Pulawski is largely light red colored. All tibiae, tarsi and nearly all metasoma, except basal part of terga IV-VI in males and IV-V in females, are light red. The species is similar to T. dimidiatus in body morphology and sculpture but differs in the following characters: ♀: Supraantennal tubercle distinctly elevated, but supraclypeal area nearly flat. Gena inconspicuously developed but not short. Pygidium densely punctate, all coarsely microsculptured, dull. ♂: Gena dorsally inconspicuous. Vertex finely and densely punctate, and postocellar impression shallow. Forefemoral notch large, usually longer than the space that separates it from forefemoral base.
Mesosoma. Scutum without distinct anterior impression; scutum and scutellum densely punctate, punctures well defined, less than half to one diameter apart, interspaces well developed, variable in size, microsculptured, slightly shiny, setae about 1.0 × MOD or less. Mesopleuron coarsely microsculptured, without distinct punctures; hypoepimeral area irregularly microsculptured without punctures; ventral part of mesopleuron with punctures ill defined, less than one diameter apart, interspaces small, slightly shiny. Propodeal dorsum relatively finely, regularly rugose, with few indistinct irregular, longitudinal ridges basally, dull, but slightly shiny posteromedially; propodeal side regularly longitudinally ridged, ridges inconspicuous basally, interspaces microsculptured, slightly shiny; posteromedial margin of propodeal dorsum elevated, but not produced between posterior marginal ridges, marginal ridges positioned nearly horizontally above groove on posterior surface. Legs densely punctate, punctures small; forebasitarsal rake light yellow with reddish opalescence at distal half, with three apical spines and two to three additional spines. Wings slightly infumate with brown veins.
Metasoma. Terga I-III with ill defined silvery apical fasciae. Terga I-III densely, finely micropunctate, punctures ill defined, interspaces microsculptured, slightly shiny; apical depressions shallow, with distinct micropunctures; sculpture of terga IV-V distinctly coarser than on previous terga, also slightly more densely punctate; tergum V coarsely punctate, punctures ill-defined, but large, half to one diameter apart, interspaces microsculptured, apical depression of tergum V very sparsely and finely punctate. Pygidium large, sparsely punctate, punctures large, ill defined, interspaces nearly unsculptured, shiny. Central part of sternum II with several larger punctures, interspaces microsculptured, shiny; lateral part slightly shiny, densely micropunctate; remaining sterna with uniform sculpture similar to that on sternum II, but more or less reduced laterally.
Coloration. Central part of mandibles, terga and sterna I-III red. Distal tarsomeres dark red. Tegulae reddish translucent. Apical parts of terga I-III slightly translucent. Remaining body parts black.
Variation of females. Very variable in sculpture, but finely sculptured over all body in most specimens except apical terga (in comparison to other species). Clypeal bevel reaching base of clypeus in some specimens, but well separated from the base in most.
Antenna short, LA3:WA3 = 2.2-2.4, LA5:WA5 = 2.6-2.7. Pygidium distinctly microsculptured, of usual width and dull in some specimens, but shiny and wide in most of specimens. Some specimens appear dark in coloration, with only terga I and II dark red and nearly all sternum II black. The most characteristic coloration of metasoma is as follows: segments I and II red, tergum III red basally and laterally, with larger or smaller black area apicomedially and sternum III basally red and apically dark.
Mesosoma. Scutum without distinct anterior impression; scutum and scutellum densely punctate, punctures well defined, most punctures less than half diameter apart, interspaces distinct, microsculptured, slightly shiny to dull, setae about 1.0 × MOD or less. Mesopleuron coarsely microsculptured with distinct or indistinct punctures in lower areas, most punctures ill defined, interspaces dull; hypoepimeral area coarsely microsculptured to finely rugose, dull; ventral part of mesopleuron with small, ill defined punctures, interspaces slightly shiny. Propodeal dorsum variably sculptured, irregularly ridged, areolate, or coarsely irregularly rugose, without distinct longitudinal ridges, short longitudinal ridges developed basally; propodeal side distinctly ridged, more densely and finely ridged basally than in the center in most specimens; posteromedial margin of propodeal dorsum elevated, but not produced between marginal ridges, marginal ridges positioned nearly horizontally above groove on posterior side. Legs densely punctate, punctures small; forefemoral notch light red and large, its diameter usually longer than space that separates it from forefemoral base, central part of notch slightly elevated, anterior and posterior margins lined by small distinct ledge, notch surface without distinct setae, microsculptured, dull. Wings slightly infumate, veins brown.
Metasoma. Terga I-III with silvery, apical fasciae. Apical depressions of all terga distinct, with micropunctures similar as on central part of terga. Terga I-III densely and distinctly micropunctate, punctures ill defined, interspaces microsculptured, slightly shiny; sculpture of terga IV-VII slightly coarser and slightly more densely punctate than other terga. Sterna uniformly punctate nearly like terga.
Coloration. Apical part of mandibles, terga and sterna I and II and forefemoral notch red. Metasoma very variable in coloration; some specimens with only tergum I dark red, or terga I and II and sternum II basally red, or terga and sterna I-III equally red. Distal tarsomeres dark reddish. Tegulae brown to reddish. Apical parts of all terga and tegulae translucent. Remaining body parts black.
Tachysphex nigripennis (Spinola, 1808), restored from synonymy Figs 6,18,32,46,66,89,98,103 = Tachytes nigripennis Spinola, 1808 (p. 260), ♀. Holotype or syntypes: Italy: "prope Genuam". Type material lost (de Beaumont 1952, 44). Diagnosis. Tachysphex nigripennis (Spinola) is easily recognizable from the other species of the T. pompiliformis and T. austriacus species subgroups. It is generally dark and coarsely punctate, and thus resembles T. opacus Morawitz and T. pompiliformis (Panzer). Tachysphex nigripennis (Spinola) is relatively well characterized species. It possesses an inconspicuous gena. The vertex is distinctly wider than long. The metasoma is coarsely punctate with densely micropunctate apical depressions. The mesopleuron is coarsely sculptured, rugose, or coarsely punctate, and the hypoepimeral area has more or less distinct longitudinal ridges. The metasoma has terga I-II dark red and a black sternum II. In females, the basomedian area of the clypeus is sparsely punctate; the pygidium is exceptionaly narrow, bright shiny and with inconspicuous lateral marginal carina. In males, the mandibular inner margin possesses one distinct rectangular tooth and a distinct furrow next to the tooth distally; the clypeal lip is distinctly arcuate, longest medially; the forefemoral notch is dark red but black in some specimens, with a shiny, nearly unsculptured surface; the dorsal process of volsella is larger than in species from the T. pompiliformis species subgroup.
Metasoma. Terga I-III with distinct but sparse silvery, apical faciae. Terga I-III densely micropunctate, with very large, but superficial, sparse punctures, micropunctures ill-defined, evanescent in micosculpture, faint, interspaces microsculptured, slightly shiny; apical depressions well developed, with distinct micropunctures; sculpture of terga III-V distinctly coarser than other terga, also more densely punctate; tergum V coarsely punctate, punctures ill defined, but large, half to one diameter apart, interspaces microsculptured, apical depression of tergum V distinctly, densely finely punctate. Pygidium distinctly narrower than in other related species, integument sparsely punctate, large and small punctures intermixed, punctures ill defined, interspaces nearly unsculptured, bright shiny. Central part of sternum II with several large punctures, interspaces microsculptured, shiny; lateral part slightly shiny, densely micropunctate; remaining sterna with uniform sculpture similar to that on sternum II, but more or less reduced laterally.
Coloration. Central part of mandibles, terga I-II and distal tarsomeres dark red. Tegulae brown translucent. Apical parts of terga I-III slightly translucent. Remaining body parts all black.
Mesosoma. Scutum with shallow anterior impression, densely punctate, punctures well defined, most punctures less than half diameter apart, interspaces distinct, microsculptured, slightly shiny to dull, setae about 1.0 × MOD or less; scutellum slightly more sparsely punctate than scutum, punctures well defined, most punctures half to less than one diameter apart, interspaces shiny to slightly shiny. Mesopleuron rugose to densely punctate, most punctures in ventral half well defined, interspaces coarsely microsculptured to rugose, dull; hypoepimeral area coarsely rugose, without distinct punctures, with distinct longitudinal ridges in most specimens, dull; ventral part of mesopleuron with small ill-defined punctures, interspaces finely microsculptured, slightly shiny. Propodeal dorsum coarsely sculptured, irregularly areolate to irregularly coarsely ridged, without longitudinal ridges except at very base; propodeal side longitudinally ridged, ridges well developed, ill defined in small specimens, interspaces microsculptured, dull; posteromedial margin of propodeal dorsum slightly elevated or not elevated, marginal ridges on posterior surface well developed, horizontal, or slightly directed toward propodeal groove medially. Legs micropunctate, punctures small; forefemoral notch large, deep, semicircular, about as long as distance that separates it from forefemoral base or slightly longer, proximal margin obtuse, not elevated over distal margin, central part of notch inconspicuously elevated, anterior and posterior margin lined by nearly indistinct ledge, notch surface without distinct setae, unsculptured, or very finely microsculptured, shiny. Wings infumate, with brown veins.
Metasoma. Silvery apical fasciae of terga I-III distinctly developed, but fasciae brown in specimens from eastern Turkey. Apical depressions of all terga shallow, densely micropunctate, nearly as densely as on tergal disk. Terga I-III densely and distinctly micropunctate, punctures ill defined, interspaces microsculptured, slightly shiny; terga IV-VII more coarsely and more densely punctate than previous terga. Sterna uniformly punctate nearly like terga.
Coloration. Apical part of mandibles red; terga I and II dark red; apex of sternum I and base of sternum II dark red in some specimens; distal tarsomeres dark red on ventral side; forefemoral notch dark red in most specimens, but black in some. Tegulae brown translucent. Apical parts of terga slightly translucent. Remaining body parts all black.
Mesosoma. Scutum without distinct anterior impression; scutum and scutellum densely punctate, punctures well defined, less than half diameter apart, interspaces well developed and relatively uniform in size, microsculptured, slightly shiny, setae about 1.0 × MOD or less. Mesopleuron coarsely microsculptured, without distinct punctures; hypoepimeral area finely rugose with indistinct, longitudinal, irregular ridges, without punctures; ventral part of mesopleuron finely, densely punctate, with punctures ill defined, less than one diameter apart, interspaces small, slightly shiny. Propodeal dorsum relatively finely, regularly rugose, with short irregular longitudinal ridges basally, dull, posteromedially with transverse ridges on small longitudinal slightly shiny area; propodeal side regularly longitudinally ridged, ridges inconspicuous basally, microsculptured, dull; posteromedial margin of propodeal dorsum elevated, slightly produced between marginal ridges, marginal ridges positioned nearly horizontally above groove on posterior surface. Legs densely punctate, punctures small; forebasitarsal rake light reddish, with three apical spines, one preapical and one or two additional ones. Wings slightly infumate, yellowish, with brown veins.
Metasoma. Terga I-III with distinct silvery, apical faciae; terga I-II very densely and finely micropunctate, punctures ill-defined but distinct, interspaces microsculptured, slightly shiny; apical depressions well developed, with distinct micropunctures; sculpture of terga III-V distinctly coarser than on previous terga, also more densely punctate; tergum V coarsely punctate, punctures ill-defined, but large, half to one diameter apart, interspaces microsculptured, apical depression of tergum V very sparsely, finely punctate. Pygidium of usual size, sparsely punctate, punctures large, ill defined, interspaces microsculptured basally, but unsculptured in apical half, slightly shiny to shiny. Central part of sternum II with several large punctures, interspaces microsculptured, shiny; lateral part slightly shiny, densely micropunctate; remaining sterna with uniform sculpture similar to that on sternum II, but more or less reduced laterally.
Coloration. Central part of mandibles, terga and sterna I-III red. Distal tarsomeres are dark red. Tegulae brown translucent. Apical parts of terga I-III slightly translucent. Remaining body parts all black.
Variation of females: Body length: 7.2-9.0 mm. Antenna relatively short, LA3:-WA3 = 1.9-2.1, LA5:WA5 = 2.9-3.0. In general, the species is not significantly variable in key characters. There is the usual variation in size dependent characters like the number of spins on foretarsal rake. The shape of the lateral emargination of the clypeal lip is one of a few variable characters; it can be well developed to nearly absent, but never angulated. The pygidium is slightly variable in width and sculpture; it is of usual shape in most specimens with distinct microsculpture, slightly shiny in basal half, but also wider and bright shiny in some specimens.
Mesosoma. Scutum without distinct anterior impression; scutum and scutellum densely punctate, punctures well defined, most punctures less than half diameter apart, interspaces distinct, microsculptured, slightly shiny to dull, setae about 1.0 × MOD or less. Mesopleuron coarsely microsculptured, with distinct or indistinct punctures in lower areas, most punctures ill defined, interspaces dull; hypoepimeral area coarsely microsculptured to finely rugose, dull; ventral part of mesopleuron densely punctate, punctures small ill defined, interspaces slightly shiny. Propodeal dorsum finely irregularly areolate to rugose, without longitudinal ridges, posteromedially with finely sculptured longitudinal area, which appears slightly shinier than adjacent areas; lateral surface of propodeum coarsely microsculptured, with or without ill defined longitudinal ridges, dull; posteromedial margin of propodeal dorsum elevated, but not produced between marginal ridges, marginal ridges positioned nearly horizontally above groove on posterior side. Legs densely punctate, punctures small; forefemoral notch light red and large, semicircular, its diameter is usually longer than distance that separates it from forefemoral base, central part of notch slightly elevated, anterior and posterior margins lined by small ledge, nearly indistinct in some specimens, notch surface without distinct setae, microsculptured, dull. Wings slightly infumate, veins brown.
Coloration. Apical part of mandible, terga and sterna I-III and forefemoral notch red. Distal tarsomeres reddish. Tegulae brown to reddish. Apical parts of all terga and tegulae translucent. Remaining body parts all black.
Geographic distribution. Bulgaria, Czech Republic, Greece, Hungary, Poland and Turkey.
Geographic distribution. China, Iran, Kazakhstan, Kyrgyzstan, Tajikistan, Turkey, and Uzbekistan. It is probably also distributed in Jordan, Lebanon and Syria.
Note. Holotype came from China (Uighur region). Type material not examined. The species was redescribed by Pulawski (1971).
Type material. Holotype: ♀, no specific locality at the label, but probably from Germany. Holotype by monotypy. "Coll. Sturm Diagnosis. Tachysphex pompiliformis (Panzer) is difficult to distinguish from T. dimidiatus (Panzer), T. jokischianus (Panzer) and T. punctipleuris sp. n. It is exceptionally variable in most characters and no single character alone can differentiate it from morphologically similar species. This species is generally darker and more coarsely punctate than similar species. The metasoma is densely micropunctate, including apical depressions. The mesopleuron is coarsely sculptured, rugose, or coarsely punctate, and the hypoepimeral area has more or less distinct longitudinal ridges. In males, sternum II is often dark red to partly black, but tergum II is red; however, specimens with red metasomal segments I-III also occur. The terga are darker than in other species in some females, but this occurs less often. The forefemoral notch of the male is dark in most specimens. The gena is robust. The following combination of additional characters is also characteristic of this species: ♀: The clypeus is slightly convex, and the most elevated point is not well defined but located at about the clypeal midlength. The vertex setae are short and uniformly semierect both medially and laterally. The dorsolateral cuticular projection of the propodeal spiracle is arcuate to semicircular and usually dark. The propodeal dorsum and side are longitudinally ridged; the dorsum is coarsely sculptured, but the ridges are indistinct in some specimens. The forebasitarsal rake is reddish in fresh individuals, with two, well separated apical spines and one additional preapical spine. The wings are often but not always dark. ♂: The clypeus is coarsely punctate, the interspaces between the punctures large, approximately one diameter apart. The forefemoral notch is markedly carinated on the anterior and posterior margin. The volsella is dark, the dorsal process narrow, about as wide as the volsellar corpus basally.
Mesosoma. Scutum without distinct anterior impression; scutum and scutellum densely punctate, punctures well defined, most punctures less than half diameter apart, interspaces distinct, microsculptured, slightly shiny to dull, setae about 1.0 × MOD or less. Mesopleuron rugose to densely punctate, most punctures in ventral half well defined, interspaces coarsely microsculptured to rugose, slightly shiny to dull; hypoepimeral area coarsely rugose, without distinct punctures, with distinct longitudinal ridges in most specimens, dull; ventral part of mesopleuron with punctures large, well defined (ill defined in small specimens), interspaces finely microsculptured, slightly shiny. Propodeal dorsum coarsely sculptured, longitudinally ridged, irregularly ridged, or areolate; side longitudinally ridged, ridges well developed, interspaces microsculptured, dull; posteromedial margin of dorsum slightly elevated or not elevated, marginal ridges fine, variably sinuous, horizontal, with ventromedially directed medially. Legs densely punctate, punctures small; forefemoral notch small, but relatively deep, semicircular, shorter than distance that separates it from forefemoral base, proximal margin relatively sharp, elevated over distal margin, central part of notch slightly elevated, anterior and posterior margins lined by faint ledge, notch surface without distinct setae, coarsely microsculptured, dull. Wings moderately infumate, but relatively pale in some specimens, veins brown.
Coloration. Apical part of mandible red; terga I and II dark red, base of tergum II dark red in some specimens; apex of sternum I and base of sternum II dark red, infrequently specimens with three red metasomal segments also occur; distal tarsomeres dark red on ventral side; forefemoral notch black in most specimens, but dark red in a few. Tegulae reddish or brown translucent. Apical parts of all terga slightly translucent. Remaining body parts all black.
Head. Clypeus distinctly convex, top at clypeal midlength; basomedian area relatively large, more sparsely punctate and punctures larger than on lateral section, interspaces between punctures basolaterally small, but getting sparser in mesoventral direction, interspaces well developed, but punctures mostly less than one diameter apart; bevel large, distinctly convex, as long as basomedian area both laterally and medially, not triangular, with sparse large punctures, not reaching base of clypeus, shiny; lip slightly arcuate, with lateral incision, separate from bevel by punctate groove, WML:LCL = 1.6, WCL:WML = 1.8. Supraclypeal area flat, distinctly punctate, punctures ill-defined, interspaces between punctures shiny to slightly shiny. Supraanten-nal tubercle slightly elevated on inner side. Antenna relatively short. Frons uniformly punctate, punctures well defined, about half diameter apart or less, interspaces slightly variable in size; frontal median line distinct, narrow, finely impressed. Vertex punctate, punctures well defined, less than half to one diameter apart, laterally sparser than medially, interspaces microsculptured, dull. Vertex setae short semierect, slightly longer laterally than medially, about 0.5 × MOD; postocellar impression distinct, shallow, Vshaped; vertex slightly wider than long; WV:LV = 1.3. Gena dorsally well developed.
Metasoma. Terga I-III with silvery apical fasciae inconspicuously developed. Apical depressions of all terga shallow, with distinct micropunctures, on terga I-II evanescent in microsculpture. Terga I-III densely, finely micropunctate, punctures ill defined but distinct, interspaces microsculptured, slightly shiny; sculpture of tergum III-V distinctly coarser than on previous terga, also punctures slightly denser. Pygidium sparsely punctate, punctures ill defined, interspaces microsculptured, dull. Central part of sternum II with several large punctures, interspaces microsculptured, shiny; lateral part slightly shiny, densely micropunctate; remaining sterna with uniform sculpture similar to that of sternum II, but more or less reduced laterally.
Coloration. Central part of mandible, three distal tarsomeres, terga and sterna I-III red. Tegulae brown translucent. Apical parts of terga I-III slightly translucent. Remaining body parts all black.
Variation of females: Body length: 6.7-9.3 mm. WV:LV = 1.3-1.4. Antenna, LA3:WA3 = 2.2-2.4, LA5:WA5 = 2.6-2.7. Very variable in coloration and sculpture. The holotype female is among the specimens with the finest sculpture. The mesopleuron is coarsely punctate to coarsely rugose; the hypoepimeral area is longitudinaly ridged, but also coarsely and uniformly rugose in some specimens. The propodeal dorsum is longitudinaly ridged, but also coarsely and uniformly rugose in some specimens (usually the hypoepimeral area and the propodeal dorsum with a comparable sculpture). The abdominal segments I-III equally red or dark red, but also two, or just one tergite red, and then sterna darker than terga.
Geographic distribution. Austria, Czech Republic, Bulgaria, Germany, France, Greece, Hungary, Italy, Kyrgyzstan, Portugal, Slovakia, Switzerland and Turkey. from the similar species. It possesses the following combination of characters: ♀: Clypeus slightly convex, the most elevated point not well defined, located approximately at clypeal midlength. Clypeal bevel variable, reaching the clypeal base in some specimens but well separated from it in others. Clypeal lip in fresh specimens sinuate, with ill-defined emargination medially. Vertex setae short, uniformly semierect medially as well as laterally. Postocellar impression well developed, deep, an open widely Y-shaped, or the posterior margin of the impression sinuous. Head nearly round, with uniformly, densely punctate frons. Gena robust. Punctures of mesopleuron variable but distinct and well defined in some specimens. Dorsolateral cuticular projection of propodeal spiracle arcuate to semicircular and usually all dark. Propodeal side uniformly longitudinally ridged, except in small specimens. Punctures of central part of tergum IV and V ill defined, interspaces between the punctures large, distinct, slightly shiny to shiny. Pygidium wide and shiny in most specimens, similar to T. jokischianus (Panzer). ♂: Clypeus slightly convex. Clypeal bevel large, densely punctate, interspaces between punctures small, punctures less than one diameter apart. Vertex approximately as wide as long. Gena robust. Supraclypeal area flat, supraantennal tubercle slightly elevated ventromedially. Mesopleuron laterally and ventrally distinctly punctate (punctures distinct also on hypoepimeral area in some specimens), punctures well defined, interspaces between the punctures distinct, slightly shiny to shiny on most parts. Forefemoral notch small, inconspicuously carinated, anterior carina distinct, black to dark red in coloration. Terga sparsely micropunctate, with variable interspaces between punctures. Volsella light brown, dorsal process basally wider than volsellar corpus.
Mesosoma. Scutum without distinct anterior impression; scutum and scutellum densely punctate, punctures well defined, half to one diameter apart, interspaces slightly microsculptured, shiny, setae about 1.0 × MOD. Mesopleuron variable, dis-tinctly, densely punctate, punctures well developed, interspaces microsculptured, slightly shiny to dull; in some specimens mesopleuron with ill-defined punctures, or with indistinct punctures in posterior and dorsal part; hypoepimeral area dull, with distinct punctures, or coarsely microsculptured, or finely longitudinally ridged, dull; ventral part of mesopleuron with punctures well developed in most specimens, less than one diameter apart, interspaces shiny. Propodeal dorsum relatively finely, irregularly rugose, but with distinct longitudinal ridges in some specimens; propodeal side irregularly, incompletely longitudinally ridged, ridges well developed in large specimens, completely absent in some small specimens, dull; posteromedial margin of propodeal dorsum slightly elevated, slightly produced between marginal ridges, marginal ridges directed ventromedially toward groove on posterior side. Legs densely punctate, punctures small; forebasitarsal rake pale reddish to yellowish, with three apical spines, one preapical spine, and two additional spines. Wings slightly yellowish, veins brown.
Coloration. Central part of mandibles, three distal tarsomeres, terga and sterna I-III red. Tegulae brown translucent. Apical parts of terga I-III slightly translucent. Remaining body parts all black.
Coloration. Apical part of mandibles, tegulae, terga and sterna I-III red; distal tarsomeres and forefemoral notch in center dark reddish. Apical parts of terga and tegula translucent. Remaining body parts all black.
Variation of males: Mandibular inner margin with one distinct rectangular tooth and distinct furrow next to the tooth distally. Antenna relatively short, LA3:WA3 = 1.5-1.6, LA5:WA5 = 1.9-2.1. Volsella light brown, ventral setae pointing in various directions; dorsal process slightly wider than corpus in most specimens. Sculpture of mesopleuron very variable, densely to sparsely punctate, punctures mostly well defined, but indistinct in some specimens. Propodeum coarsely to finely sculptured, with or without longitudinal ridges. Forefemoral notch often black.

Discussion
Morphological variation in T. pompiliformis sensu Pulawski (1971) was found to be extensive (Pulawski 1971). Some species of this group have already been separated by Pulawski himself (Pulawski 1971) or later recognized as valid (Straka 2004(Straka , 2005. Present recognition of fourteen species and two species subgroups may appear surprising, but this variation is correlated with ecological and biogeographic data. Not all species are ubiquitous. Only two reach Scandinavia (T. jokischianus and T. dimidiatus). Some species occur predominantly on sandy or loess habitats (T. jokischianus, T. nigripennis, and T. nobilis). This uneven distribution have suggested the presence of different species within a group for a long time, validated in the present study. These species delimitations are supported by morphological characters.