Description of Three New Cleptes Species from Cyprus and Lebanon with Updates and Revisions to the Genus (Hymenoptera, Chrysididae)  ^†

Abstract

Three new species of Cleptes Latreille, 1802, in the C. nitidulus species group are described: C. sfenthourakisi Rosa & Makris, sp. nov. (from Cyprus), C. christi Rosa & Makris, sp. nov. (Cyprus), and C. adonis Rosa & Boustani, sp. nov. (Lebanon). A new synonymy is proposed: C. ignitus var. chevrieri Frey-Gessner, 1887, syn. nov. of C. splendidus (Fabricius, 1794). The two members of the C. morawitzi species group, and the single member of the C. rugulosus group are transferred to the C. nitidulus group. The Cleptes semenovi species group is transferred from the subgenus Chrysocleptes Móczár, 1962 to the subgenus Leiocleptes Móczár, 1962, and synonymised with the C. nitidulus species group. Cleptes viridis Gravenhorst, 1806, is considered nomen dubium. An updated checklist of the C. nitidulus group is given, including all the species described and synonymised after the latest revision by Móczár.

1. Introduction

Cleptes Latreille, 1802 is the largest genus in the subfamily Cleptinae, with 95 recognized worldwide species [1,2,3,4,5,6,7,8,9,10,11,12,13,14,15,16], 72 of which occur in the Palaearctic realm only. The subfamily includes two other small genera, Cleptidea Mocsáry, 1904 (20 species) and Lustrina Kurian, 1955 (1 species), which are respectively distributed only in the Neotropical and in the Oriental realms [17,18,19,20]. Cleptes are parasitoids of the prepupae of sawflies (Hymenoptera, Symphyta) in the families Tenthredinidae and Diprionidae [1,4,21].

Móczár [2,3,4,5,6,7,8,9,10] provided extensive data for the establishment of all Cleptes species groups, starting from the revision of the C. nitidulus species group [2]. He later reintroduced the use of subgenera [3,4], which were synonymised with Cleptes by Kimsey and Bohart [1], and included the C. nitidulus species group in the subgenus Leiocleptes Móczár, 1962. We follow Móczár’s taxonomic arrangement, although we propose to synonymise some species groups, namely C. morawitzi, C. rugulosus and C. semenovi groups, with the C. nitidulus species group based on the examination of type material and other specimens that were not available to the Hungarian author. We here describe three new species of the C. nitidulus species group from Cyprus and Lebanon and provide further taxonomic information on other members of this group. Finally, we provide an updated checklist of the members of the C. nitidulus species group.

2. Materials and Methods

Holotypes of the newly described species are deposited in the Linsenmaier collection housed at the Natur-Musem (Luzern, Switzerland, NMLU). Paratypes are deposited in the private collections of P. Rosa (Bernareggio, Italy, PRC) and C. Makris (Cyprus, CMC). Other specimens studied in this paper are deposited at: Institute of Biology and Pedology of the National Academy of Sciences of Kyrgyzstan, Bishkek (IBPB); Muséum Histoire Naturelle, Geneva (Switzerland, MHNG) and Zoological Institute of St. Petersburg (Russia, ZIN). Species group is abbreviated into “group” in the following text.

Morphological terminology follows partly Kimsey and Bohart [1] and partly Lanes et al. [22]. Abbreviations used in the descriptions are as follows: F1, F2, F3, etc. = flagellomeres 1, 2, 3, etc., respectively; l/w = length/width ratio; MOD = median ocellus diameter (measured in frontal view); MS = malar space, the shortest distance between base of mandible and lowest margin of compound eye; OOL = oculo-ocellar line, the shortest distance between posterior ocellus and compound eye; P = pedicel; PD = puncture diameter; POL = posterior ocellar line, the shortest distance between posterior ocelli; T1, T2, T3, etc. = metasomal terga 1, 2, 3, etc. Length measure of metascutellum is taken from the antero-median margin to the postero-median margin, and the width is given by the shortest distance between the mesoscutellar troughs.

Type specimens of Cleptes sfenthourakisi sp. nov. and C. christi sp. nov. were photographed by C. Makris with a Camera Canon EOS 5D Mark III (Canon Inc., Ōta, Tokyo, Japan), with Canon MP-E 65 mm f/2.8 1-5x Macro Photo lens (Canon Inc., Ōta, Tokyo, Japan); images were stacked with software Zerene Stacker (Zerene Systems LLC, Rich-land, WA, USA) then enhanced with Adobe Photoshop (Adobe Inc., San Jose, CA, USA). Images of C. semenovi and C. adonis were taken by P. Rosa with a Camera Olympus E-M1 Mark II (Olympus Corporation, Shinjuku, Tokyo, Japan), with Olympus Zuiko 60 mm(Olympus Corporation, Shinjuku, Tokyo, Japan); images were stacked with Helicon Focus 7.7.5 (Helicon Soft Ltd., Oakland, CA, USA) software then enhanced with Adobe Photoshop. Images of C. morawitzi were taken with a Camera Nikon D-80 (Nikon Corporation, Minato, Tokyo, Japan) connected to the stereo microscope Togal SCZ (Togal, Thailand) and stacked with the software Combine ZP (Alan Hadley, GPL Software, UK).

3. Results

3.1. Taxonomy

Cleptes Latreille, 1802

Cleptes Latreille, 1802: 316. Type species: Sphex semiaurata Linnaeus, 1761, by monotypy.

Cleptes (Leiocleptes) Móczár, 1962

Leiocleptes Móczár, 1962: 118 (as subgenus of Cleptes Latreille, 1802). Type species: Cleptes nitidulus Fabricius, 1793, by original designation.

Leiocleptes Móczár: Junior subjective synonym of Cleptes Latreille, 1802 according to Kimsey and Bohart, 1991: 53.

Leiocleptes Móczár: Revalidated as subgenus by Móczár, 1997b: 89.

Diagnosis of Cleptes (Leiocleptes). Pronotum simple (Figure 1, Figure 2 and Figure 3A–C), without median, longitudinal row of pits; without posterior groove, irregular punctures and/or a basal, longitudinal keel; pronotal disc convex, posteriorly connected to mesonotum without any depression; lateral margins of first two metasomal segments, although less evidently sharp than in the subgenus Oxycleptes Móczár, 1962; mesopleuron smooth below alar foveae, punctures on anterior margin usually aligned in striae [3].

Móczár [3,4] considered five species groups in Leiocleptes: C. alienus, C. nitidulus, C. morawitzi, and C. rugulosus groups. Here, we consider only two species groups C. nitidulus and C. alienus (Nearctic, not evaluated in this article, to be revised).

Diagnosis of the Cleptes (Leiocleptes) nitidulus species group. The species included in the C. nitidulus group are recognized by the simple pronotum without posterior pit row, longitudinal median sulcus or median basal keel. The metasomal colour is variable; usually, the metasoma is basally yellowish, brownish or reddish and apically blackish with or without metallic segments, but some Asian species have entirely black metasoma (such as C. morawitzi Radoszkowski, 1877 [23], C. helanshanus Wei, Rosa & Xu, 2013, C. mareki Rosa, 2003 and C. sinensis Wei, Rosa & Xu, 2013 [14]), or black metasoma laterally and apically with violet or green highlights (C. transoxianus Rosa, 2018 and C. semenovi Kuznetzov-Ugamszkii, 1927).

Until Móczár [3], the distinction with the Nearctic C. alienus group was based only on the colour of the metasoma, given as reddish and apically blackish in the C. nitidulus group and:

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with basal segments often metallic laterally, and apical segments almost entirely metallic (from Kimsey and Bohart [1])

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with metallic green, blue or purple highlights along lateral margins and (or) also last tergites (except in some specimens, cf. Cleptes speciosus). Head and thorax usually blue to purple or bright green to bluish green (male), bronze to copper (in Cleptes speciosus, female). T-III rarely with double punctures (from Móczár [3]).

The diagnosis of the Cleptes asianus group should therefore be re-evaluated based on morphological characters and not only on body colour. New findings in the last decade have broadened the spectrum of the metasomal colour in members of the C. nitidulus group. The analysis of members of the C. asianus group is postponed to future research since not all the North American species were available for this study, and this analysis is beyond the scope of this article.

3.2. New Species Description

3.2.1. Cleptes (Leiocleptes) sfenthourakisi Rosa & Makris, sp. nov.

LSID: urn:lsid:zoobank.org:act:FB2261EA-6A9C-4DF6-9258-6462B32CFBA0

Figure 1A,B, Figure 2A,B,D,E, Figure 3A,B,D, Figure 4A,B and Figure 5A–D.

Type Material

Holotype: female, Cyprus: Larnaka Salt Lake, near Hala Sultan Tekke, 34°53′17″ N 33°36′15″ E, 26.v.2012, leg. C. Makris (NMLU). Paratypes: one male, five females, same locality, 27.v.2012 (NMLU, PRC, CMC).

Diagnosis. The female is recognised by an androchrome colour pattern, with blue head and mesosoma, and black face; deep and dense punctation on mesosoma and metasoma; double punctation on T3. The male is recognised by a combination of diagnostic features, such as shape of genitalia with elongate apex of gonostylus, metascutellum subsquare, propodeal projections with straight lateral side, metallic tibiae.

Description. Female. Body length 7.3–7.5 mm (holotype 7.3 mm, Figure 1B).

Head. Vertex with small, deep, dense punctures, with even interspaces (1–3 PD); face medially with sparser punctures (2–3 PD), laterally closer; clypeus apically emarginate, with straight, protruding medial truncation, with right angle and without teeth at corners; frontal sulcus broad and deep, almost complete from anterior ocellus to clypeal margin (Figure 2B), deeper and broadened below anterior ocellus and before clypeal margin; ocellar triangle isosceles, without distinct post-ocellar sulcus, but with slight depression (Figure 2E) more or less visible in the paratypes; lateral pits of posterior ocelli deep, elongated (1.0 × MOD), as large as 0.5 × MOD; MS = 1.6 × MOD; POL = 1.6 × MOD; OOL = 2.2 × MOD. P:F1:F2:F3 = 1.0:1.5:0.6:0.6.

Mesosoma. Pronotum unmodified; with deep punctures larger than those on the vertex; dorsally with relatively large, polished interspaces, becoming narrower at the sides; punctures smaller along the posterior margin; mesoscutum and mesoscutellum with shallow and relatively small punctures, with polished, large interspaces (1–3 PD) (Figure 3B); notauli and parapsidal lines deep and complete; upper mesopleuron with wide polished interspaces (Figure 3D), and small, shallow punctures; deep, short scrobal pit (Figure 3D); large and transversally confluent punctures postero-medially; metanotum with large axillary trough irregularly and longitudinally wrinkled; metascutellum subsquare; metapleuron transversely wrinkled; dorsal surface of propodeum short, as long as the mesoscutellum, irregularly reticulate, with sharp ridges (Figure 3B); posterior propodeal projections subparallel, with straight angle.

Metasoma. T1 almost impunctate, with shallow, sparse, and tiny punctures medially; T2 basally densely micropunctate (Figure 4B); T3 with double punctation with small dots among punctures in the first two-thirds, with shallow and sparse punctures in the apical third; T4 with large punctures and polished interspaces; all terga with impunctate stripe along posterior margin.

Colouration. Face and large part of frons medially dull black; rest of the head metallic blue dorsally (Figure 2B); mesosoma blue with blackish areas on median area of mesoscutum and declivitous part of propodeum (Figure 1B); some specimens show small discoloured brown areas on pronotum and mesonotum; metasoma orange on T1–T2 and basally on T3; apical half of T3 and T4 black without metallic reflection. Pedicel and F1 foxy, other antennal articles brown; tegulae non-metallic brown; legs dark brown to black without metallic reflections on femora; fore tibia light brown; fore tarsi testaceous.

Vestiture. On head with long (1.5 × MOD) and thick black setae, clearer to yellowish on mandibles; mesosoma laterally with relatively long (up to 2.0 × MOD), dark grey setae on pronotum and mesonotum; with dense, long and whitish setae on lower mesopleuron; femora with elongate, erect and appressed, greyish setae; metasoma laterally and posteriorly, with long (up to 3 × MOD) sparse, erect, greyish setae on T3 mixed with short, whitish and appressed setae.

Male. Body length: 7.0 mm. Similar to female in size and body punctation, with deep and dense punctures on mesosoma and metasoma (Figure 1A); metascutellum, metapectal-propodeal complex and posterior propodeal projections similar to female with lateral side straight. Sexual dimorphic characters: fore body uniformly blue and largely black T3 (Figure 1A); face, seen in frontal view (Figure 2A), narrowed medially, and convergent apically, with shorter malar spaces (0.8 × MOD); fully black flagellomeres, distinctly elongate, with F2–F11 l/w = 2 (Figure 1A).

Variability. Two female paratypes show two slight sub-basal pronotal depressions, thus generating a weak postero-median keel, which has to be considered as a visual effect and not a sort of keel as in the Cleptes satoi group.

Distribution. Cyprus.

Etymology. The specific name is dedicated to Spyros Sfenthourakis (Professor of Ecology and Biodiversity, Dept. of Biological Sciences, University of Cyprus) who discovered the new species.

Habitat. The collecting site is 5–7 m above sea level (Figure 6A). The specimens were collected on flowers of the Apiaceae Opopanax hispidus in May (Figure 6B). The habitat is a field border with Crataegus azarolus, Convolvulus dorycnium and Prosopis farcta; in this site, flowering plants which attract cuckoo wasps are Ferula communis, flowering in February–April and Opopanax hispidus flowering in May–June.

Comparative diagnosis. Cleptes sfenthourakisi sp. nov., C. christi Rosa & Makris, sp. nov. and C. adonis Rosa & Boustani, sp. nov. females show an androchrome colour pattern, with blue head and mesosoma. Nevertheless, the fully black face without metallic reflections is apparently a distinctive character of the female, because the male face is usually concolour with the rest of the head. The androchrome colouration separates these three species from all the other females of Leiocleptes from the Palaearctic Region. Other females show large part of mesosoma red, violet, coppery, non-metallic orange or black. Androchrome females are rarely observed in the Palaearctic area (see below, in the C. aerosus and C. dahlbomi groups) and are more frequent in the Oriental groups (e.g., C. asianus and C. fudzi groups), yet these groups belong to different subgenera, distinct from Leiocleptes by the pronotal structure [4].

Cleptes sfenthourakisi sp. nov. female can be separated from C. christi Rosa & Makris, sp. nov. female by the following combination of characteristics: larger dimensions (7.3 mm vs. 4.5 mm); deeper and denser punctation on mesosoma (Figure 3B) (vs. shallower and sparser (Figure 3C)); denser punctation on T2 and T3 (Figure 4B) (vs. sparser (Figure 4C)); face with a deep medial sulcus extended from anterior ocellus to clypeus (Figure 2B) (vs. barely visible only before anterior ocellus (Figure 2C)); posterior propodeal projections straight, expanded at base, right angled (vs. narrow, subparallel). Cleptes sfenthourakisi sp. nov. can be separated from Cleptes adonis Rosa & Boustani, sp. nov. by the antennae with pedicel and F1 light brown, and the rest of flagellomeres brownish (vs. all flagellum dark black); metascutellum subsquare, l/w = 1.2 (vs. metascutellum subrectangular, l/w = 1.4); posterior propodeal angles with straight sides, subparallel, apically right angled (vs. laterally sinuous, with angles apically pointed, divergent).

Males of C. christi sp. nov. and C. adonis sp. nov. are unknown. The male of Cleptes sfenthourakisi sp. nov. is similar to other two species distributed in the East Mediterranean and Middle East: Cleptes splendidus Fabricius, 1794 and C. femoralis Mocsáry, 1890 for the general habitus, colouration and even, simple punctation of the third tergum; the females of these two species can be easily separated from the female of C. sfenthourakisi sp. nov. by the forebody colouration, red with golden or violet areas. The male of C. sfenthourakisi sp. nov. can be separated from the male of C. femoralis by the metallic femora and tibiae (vs. yellowish to brownish meso-, meta-femora, and tibiae); and it can be separated from the male of C. splendidus by subsquare metascutellum (vs. subrectangular) and propodeal ridges less elaborated and sharp. The male of C. sfenthourakisi sp. nov. can be separated from the males of both species by the shape of the genital capsule: compare Figure 5C,D with Figure 5C,E,G,H, in which the apex of the gonostylus is elongate and curved and the upper inner margin of volsellae is distinctly angled and not gently curved.

3.2.2. Cleptes (Leiocleptes) christi Rosa & Makris, sp. nov.

LSID: urn:lsid:zoobank.org:act:3A65320B-1C2C-44C8-8C85-567799D43C9C

Figure 2C,F, Figure 3C,E, Figure 4C and Figure 7

Type material

Holotype: female, Cyprus: Pafos District, near Eledio, 34°48′41″ N 32°34′32″ E, 10.v.2020, leg. C. Makris (NMLU). Paratype: 1 female, same locality, 30.v.2020, leg. C. Makris (CMC).

Diagnosis. Cleptes christi Rosa & Makris, sp. nov. is recognisable by dark blue colour pattern of head and mesosoma; small-sized body (4.5 mm); shallow and sparse punctation pattern over body. The male is unknown.

Description. Female. Body length 4.5 mm (Figure 7A).

Head. Vertex with small, shallow, sparse punctures and wide interspaces (2–4 PD); face with sparse punctures; clypeus apically emarginate, with straight, protruding medial truncation, with right angle and without teeth at corners; frontal sulcus as fine line on frons and with deep fovea at base of toruli; ocellar triangle isosceles, without post-ocellar sulcus; lateral pits of posterior ocelli deep, rounded, small, as large as 0.3 × MOD (Figure 2F); MS = 2.5 × MOD; POL = 2.0 × MOD; OOL = 2.4 × MOD. P:F1:F2:F3 = 1.0:1.2:0.6:0.5.

Mesosoma. Pronotum unmodified; with shallow, small punctures, similar to those on vertex, with wider polished interspaces (4–8 PD); mesoscutum and mesoscutellum with similar shallow and small punctures; with polished, wide interspaces (4–6 PD) (Figure 3C); notauli deeply impressed; parapsidal lines barely impressed; with rugose axillary trough, not irregularly wrinkled; upper mesopleuron impunctate, medially and anteriorly with sparse punctures not aligned; deep, short scrobal pit (Figure 3E); metascutellum shallowly punctate; dorsal surface of propodeum short, slightly longer than mesoscutellum length, irregularly and shallowly reticulate, with ridges barely visible (Figure 3C). Posterior propodeal projections elongate, subparallel and with right angle.

Metasoma. T1 impuctate; T2 medially micropunctate; T3 with larger, shallow punctures and without tiny dots on interspaces; punctation over metasoma is progressively deeper from T2-T4; all terga with impunctate stripe along posterior margins (Figure 4C).

Colouration. Face and most of frons dull black; rest of the dorsal head metallic blue to violet; mesosoma blue to violet, blackish on mesoscutum and propodeum; metasoma orange on T1, T2 and T3; black without metallic reflection on apical half of T3 and T4; antenna light brown on P and F1, rest of antennal articles brown; tegula non-metallic brown; legs dark brown to black without metallic reflections on femora; foretibia brownish, lighter; tarsi testaceous.

Vestiture. Vertex with long (up to 2.0 × MOD), sparse, erect setae; black setae on face, vertex, and temples, greyish setae on occiput and gena; pronotum and mesonotum with long (up to 3.0 × MOD), black setae; greyish, shorter setae on rest of lateral and ventral mesosoma, and propodeum; latero-posterior border of T3, with thick, long (up to 3.0 × MOD), erected, greyish setae, darker on T3 and T4; meso- and metaleg with sparse, thick, dark, erect, and long setae on outer side, with short, appressed whitish setae on inner side.

Male. Unknown.

Distribution. Cyprus.

Etymology. The specific name christi (masculine noun in genitive) is dedicated to Christos Makris, the son of one of the authors, C. Makris.

Habitat. The two specimens of Cleptes christi Rosa & Makris, sp. nov. were collected in May on leaves of Quercus coccifera calliprinos, near the village Eledio in Pafos District. The habitat is a sparse shrubland on gypsum formation dominated by Quercus coccifera calliprinos, wild olive trees (Olea europaea), Pistacia terebinthus and Crataegus azarolus (Figure 8).

Comparative diagnosis. The female of Cleptes christi sp. nov. is characterized by androchrome colouration, shallow and sparse punctation and small size. These features distinguish C. christi sp. nov. from all the other species of Leiocleptes. The female is somehow similar to C. sfenthourakisi Rosa & Makris, sp. nov. and C. adonis Rosa & Boustani, sp. nov. for the colour pattern. It can be separated from the latter two species by: smaller dimensions (4.5 mm vs. 7.0–7.5 mm) (Figure 7A); shallower and sparser punctation on mesosoma (Figure 2C, Figure 3C and Figure 4C) (vs. mesosoma punctation deeper and denser, particularly on pronotum); T1 polished (vs. medially punctate); T3 with sparse, shallow punctures (vs. double punctation, with tiny dots on interspaces); mesopleuron with sparse punctures (vs. medially larger and transversally aligned); posterior propodeal projections narrow, subparallel (vs. sinuous at sides, with divergent pointed angles in C. adonis Rosa & Boustani sp. nov., and more expanded at base in C. sfenthourakisi Rosa & Makris, sp. nov.); lateral pits of posterior ocelli small and rounded (vs. elongate); face with a trace of medial sulcus, on frons and before toruli (Figure 2C) (vs. deep medial sulcus extended from anterior ocellus to clypeus).

3.2.3. Cleptes (Leiocleptes) adonis Rosa & Boustani, sp. nov.

LSID: urn:lsid:zoobank.org:act:6FAFE17D-8559-4E19-A410-6DECB074252F

Figure 9

Type Material

Holotype: female, Lebanon: Horch Ehden, Al Bayada, WGS84, 34°17′57′’ N 35°58′52′’ E, 1686 m, 12.vi.2019, leg. X. Van achter, yellow pan trap, XVA19–0837 (NMLU).

Diagnosis. Cleptes adonis sp. nov. female is recognised by fully black antennae; androchrome colouration, metasoma without metallic reflections; metascutellum subrectangular; posterior propodeal angles sinuous laterally, with apically pointed angles. The male is unknown.

Description. Female. Body length 7.2 mm (Figure 9A).

Head. Vertex with small, deep, dense punctures and narrow polished interspaces (1 PD); face medially with sparser punctures (2–3 PD); clypeus apically emarginate, with straight, protruding medial lobe, with right angle and without teeth at corners; frontal sulcus broad and deep, complete from anterior ocellus to clypeal margin, deeper and broadened below anterior ocellus and before toruli; ocellar triangle isosceles, with post-ocellar line as weak depression; lateral pits of posterior ocelli deep, elongated, as large as 0.5 × MOD; MS = 1.7 × MOD; POL = 1.7 × MOD; OOL = 2.4 × MOD. P:F1:F2:F3 = 1.0:1.4:0.8:0.6.

Mesosoma. Pronotum unmodified; with deep punctures slightly larger than those on vertex and polished interspaces (1–2 PD), punctures denser laterally and smaller along posterior margin; mesoscutum and mesoscutellum with similar punctures, with polished, large interspaces (1–3 PD); notauli and parapsidal lines deep and complete; upper mesopleuron with wide polished interspaces (2–3 PD), punctures small and shallow; deep, short scrobal pit (Figure 9D); metanotum with large axillary trough irregularly and longitudinally wrinkled; metascutellum subrectangular (l/w = 1.4), metapleuron transversely, deeply striate, with small punctures amongst striae; dorsal surface of propodeum short, as long as mesoscutellum length, irregularly reticulate (Figure 9B); posterior propodeal projections convex laterally with pointed and divergent angles (Figure 9B).

Metasoma. T1 with tiny, shallow and sparse punctures; T2 densely micropunctate; T3 with double punctation with small dots among punctures in the first 2/3rds, with sparse punctures in the apical third; all terga with impunctate stripe along posterior margin (Figure 9C).

Colouration. Face and most of frons dull black; rest of the head metallic blue; mesosoma blue with darker median area of mesoscutum and black propodeal declivity; T1, T2 and basal half of T3 orange; apical half of T3 and T4 dark brown without metallic reflection. Antennal articles fully black. Tegula non-metallic brown; legs dark brown to black without metallic reflections on femora; fore tibia brownish, light; fore tarsi yellowish, mid- and hind tarsi dark brown.

Vestiture. Long (1.5 × MOD) and thick black setae on head, becoming clearer to yellowish on mandibles; mesosoma with long (up to 2.0 × MOD), dark grey setae on pronotum and mesonotum laterally; dense, short and whitish setae on mesopleuron and propodeum laterally; elongate, erect, greyish setae on femora; metasoma laterally and posteriorly with long (up to 3 × MOD) sparse, erect, greyish setae; T3 with pubescence of variable size, short setae among long ones, whitish and appressed.

Male. Unknown.

Distribution. Lebanon.

Etymology. The specific epithet is named after Adonis, the god of beauty and desire in Greek mythology. Originally, he was a god worshiped in the area of Phoenicia (modern-day Lebanon), and later adopted by the Greeks. The species name was chosen for the beauty of this Cleptes.

Habitat. The locality is in the buffer zone of Horch Ehden Nature Reserve, in the Montane Mediterranean (Sub-alpine) vegetation level, with sparsely wooded areas and with mainly Quercus sp. and Prunus ursina, transitioning towards the Oro-Mediterranean (Alpine) with bare soil in the background (Figure 10). The herbaceous cover of the site is dominated by Vicia tenuifolia earlier in spring and by Medicago sp. at the time of the specimen collection in July.

Comparative diagnosis. Cleptes adonis sp. nov. is similar to C. sfenthourakisi Rosa & Makris, sp. nov. It can be separated by antennae fully black (vs. antennae with pedicel and flagellomere I light brown, and the rest of flagellomeres brownish); metascutellum subrectangular, l/w = 1.4 (vs. metascutellum subsquare, l/w = 1.2); posterior propodeal angles sinuous at sides, with angles pointed, divergent (vs. posterior propodeal angles with straight sides, subparallel, apically right angled). It can be separated from the other known Cleptes species from Lebanon, C. libanoticus Linsenmaier, 1959, by subgeneric features, such as the pronotum with a basal transverse groove, and the colouration with entirely metallic green head, and T3, T4 with blue highlights (see pictures in Rosa et al. [25]).

3.3. Changes on the Species Groups

Móczár [2] provided a revision for the Cleptes nitidulus group, including a key to 25 species. He described two new species, proposed three synonymies, reinstated two species previously synonymized, and designated some lectotypes. A few months later, Móczár [3] reintroduced the subgeneric system, included the C. nitidulus and the C. alienus groups (with three Holarctic species) in the subgenus Leiocleptes Móczár, 1962 and established other two new groups: C. morawitzi and C. rugulosus groups, based on specimens previously included in the C. nitidulus group. He also revalidated C. splendidus (Fabricius, 1794), synonymized C. consimilis du Buysson, 1887 and provided a key to the Leiocleptes groups. Finally, Móczár [6] completed the revision of the subgenus Leiocleptes with an additional supplement, including new synonyms and corrections for taxa of the nitidulus group.

After Móczár’s revision [2,3,6], six other species were described in the C. nitidulus group: C. laevifacies Linsenmaier, 1999, from Morocco; C. mareki Rosa, 2003, from China; C. hungaricus Móczár, 2009, from Hungary; C. helanshanus Wei, Rosa & Xu, 2013 and C. sinensis Wei, Rosa & Xu, 2013, from China; C. transoxianus Rosa, 2018, from Kazakhstan. Lastly, Rosa [26] described the male and illustrated specimens of C. dauriensis Móczár, 1997a from Mongolia, and Rosa et al. [25] illustrated all Cleptes types described by Linsenmaier, including members of the C. nitidulus group.

We transfer the two members of the C. morawitzi group and the single member of the C. rugulosus group to the C. nitidulus group. The Cleptes semenovi group is transferred from the subgenus Chrysocleptes to the subgenus Leiocleptes, C. nitidulus group; C. viridis, one of the two members of the C. semenovi group, is considered as a nomen dubium (see below).

3.3.1. The Case of Cleptes morawitzi and C. rugulosus Groups

Móczár [3] subdivided the subgenus Leiocleptes into four species groups: Cleptes nitidulus, C. alienus, C. morawitzi, and C. rugulosus groups. Nearctic species in the C. alienus group are not discussed here.

Móczár [3] included two species in the newly established C. morawitzi group: C. canadensis Kimsey, 1987 (Figure 11) and Cleptes morawitzi (Figure 12). He characterized this group based on inconsistent characters, such as the metasoma colouration without metallic highlights (as in most of the species in the C. nitidulus group); head and mesosoma unicolour; posterior propodeal angle stumpy and third metasomal tergum without double punctures. The metasomal colour in different in the two species, being fully black in C. morawitzi and largely yellowish in C. canadensis; head and mesosoma are shiny, metallic red in C. morawitzi but coppery, with greenish reflections, in C. canadensis, with contrasting bluish propodeum; the posterior propodeal angles are variable, as observed in Central Asian specimens belonging to undescribed species of the C. morawitzi group with pointed propodeal angle (unpubl. data).

Examination of the C. morawitzi type series (Figure 12; Rosa et al. [23]) and other undescribed Central Asian species of this group, found in Berlin and St. Petersburg, shows a broad variability for all the diagnostic characters given in the diagnosis of the group. At the same time, C. canadensis (Figure 11) seems to be similar to other species included in the nitidulus group, i.e., C. dauriensis (compare Figure 1 in Rosa [26]). Therefore, we include these two species in the C. nitidulus group.

The Cleptes rugulosus group was proposed based on a single species with coarse punctation on mesosoma, without any other morphological diagnostic character, and with colouration similar to other common species in the nitidulus group, i.e., C. ignitus (Fabricius, 1787). We consider the coarse punctation of the mesosoma a diagnostic character valid for species identification only and consider C. rugulosus as another member of the C. nitidulus group.

3.3.2. The Case of Cleptes semenovi Group

Móczár [4] revalidated the subgenus Chrysocleptes Móczár, 1962 previously synonymized with Cleptes Latreille, 1802 by Kimsey and Bohart [1] based on two features: the unmodified pronotum and the metallic metasoma. He subdivided the subgenus into two species group: the C. putoni group, based on a single species with entirely metallic metasoma, and the C. semenovi group, including two species, Cleptes semenovi Kuznetzov-Ugamszkii, 1927 and Cleptes viridis Gravenhorst, 1807, based on the laterally extensively metallic metasoma.

Móczár [4] was unable to examine any type or specimen of the C. semenovi group, based the diagnosis of the newly established group on the short original descriptions only. According to him: “This group includes two species at present: Cleptes semenovi Kuznetzov-Ugamszkii, 1927 and Cleptes viridis Gravenhorst, 1807. The type material is unknown for both species. Unfortunately, the diagnosis of viridis is very short. However, semenovi was compared by the author with C. dahlbomi Semenov; otherwise it would belong to Leiocleptes subgenus because of the simple pronotum. The body of both species is nearly entirely concolourous and with metallic highlights. The simplest solution to separate these is to form an isolated group, being in equal distance from the other groups and subgenera”.

We identified a specimen collected in Kazakhstan from IBPB as Cleptes semenovi (Figure 13). Members of the C. semenovi group show the same diagnostic characters of members in the C. nitidulus group, with metallic colouration at sides of metasoma, as in the case of C. transoxianus described from Central Asia, with red and purplish colour instead of green. We consider the colouration of C. semenovi as a diagnostic character for species identification in the C. nitidulus group. Any distinct morphological character is observed to consider C. semenovi a member of a different group or subgenus.

We consider the second member of this group, Cleptes viridis Gravenhorst, 1807 nomen dubium because the type depository is unknown [1,4], the description is too short to recognise this species, and the type locality is not given. Therefore, it is not possible to have reliable identification and designation of a neotype.

Based on the above observations on Cleptes semenovi, we here synonymise the C. semenovi group with the C. nitidulus group.

3.3.3. The Cleptes putoni Species Group

This group was established by Móczár [4] based on the simple pronotum and the fully metallic colouration of the metasoma. Cleptes putoni du Buysson, 1886 is the only member of the species group and the type species of the subgenus Chrysocleptes Móczár, 1962. The metallic pattern could be related to the ecology of this species. As explained by Rosa [27], Cleptes putoni is the most eurizonal Cleptes, and the only one repeatedly observed in dry, xerothermic habitats, entering in nests in clay walls. The metallic and heavily sclerotised metasomal cuticle is likely a response to the incremental risk of body dehydration in dry environments, in comparison to other Cleptes species which live in wet environments, in forests or in the vegetation.

We do not consider the metallic metasomal colouration as a diagnostic character for species groups; however, other diagnostic characters should be taken into consideration for its taxonomic placement. The pronotum is described as simple or at most hardly depressed posteriorly [4]. However, both sexes show an additional postero-medial keel (Figure 14A,B), more evident in the male, which was not cited by Móczár [4]. This characteristic, well visible in the neotype, combined with the extremely shortened pronotum of both sexes, place C. putoni out from the C. nitidulus group and the Leiocleptes subgenus. Pauli et al. [28] confirmed by means of molecular analyses that C. putoni belongs to a separate clade from the C. nitidulus group, and we still consider this species group and its inclusion in a separated subgenus to be valid. A molecular comparison with specimens in the Cleptes (Cleptes) satoi group is needed for a better understanding of the real placement of this species and species group, because they share short pronotum and postero-median keel, which are diagnostic characters for the C. satoi group.

3.4. Proposed Synonyms in the Cleptes nitidulus Group

During our research on Cleptes, we collected some new taxonomic data related to members of the C. nitidulus species, some of which were not mentioned in the catalogues of Kimsey and Bohart [1] and/or Móczár [2,3,6]. The remarks are listed under the original combination.

3.4.1. Cleptes afra var. medina du Buysson in André, 1891

Cleptes afra var. medina du Buysson in André, 1891: 79. Holotype female; Spain: Sierra Morena, leg. Medina (depository unknown).

Remarks. Taxon not mentioned in Móczár’s revisions [2,3,6]. Gogorza [29] described Cleptes perezii, another Spanish taxon closely related to the northern African C. afer Lucas, 1849. Du Buysson [30] synonymised C. perezii with C. afer and described a female as C. afra var. medina. C. perezii and C. medina most likely belong to the same taxon, and we here consider C. afra var. medina du Buysson in André, 1891 as a possible synonym of C. perezii Gogorza, 1887, waiting for the examination of the type materials. The current subspecific status of this taxon has still to be evaluated.

3.4.2. Cleptes chevrieri Frey-Gessner, 1887

Cleptes ignitus var. chevrieri Frey-Gessner, 1887: 30. Syntypes females; Switzerland: Champel (Geneva).

Remarks. Taxon not mentioned in Móczár’s revisions [2,3,6], yet Móczár [24] synonymized Cleptes chevrieri Frey-Gessner, 1887 with C. consimilis du Buysson, 1887. Linsenmaier [31,32] revalidated C. chevrieri and proposed C. consimilis du Buysson, 1887 as its synonym. In Geneva, we examined the two females syntypes cited by Frey-Gessner [33]. They are conspecific with C. splendidus (Fabricius, 1794). Since we follow Móczár’s species interpretation, we propose C. ignitus var. chevrieri Frey-Gessner, 1887 syn. nov. of C. splendidus (Fabricius, 1794). In fact, Móczár [2,3,6] already synonymized the following species with C. splendidus: C. caucasicus Semenov-Tian-Shanskij, 1920, C. chyzeri Mocsáry, 1889, C. consimilis Buysson, 1887, C. hyrcanus Semenov-Tian-Shanskij, 1920.

3.4.3. Cleptes nitidula var. nigricans du Buysson, 1901

Cleptes nitidula var. nigricans du Buysson, 1901: 97. Holotype male, Greece (Vienna).

Remarks. Not mentioned in Kimsey and Bohart [1] neither Móczár’s papers [2,3,6]. As mentioned by Rosa et al. [34], this taxon was correctly described, and the name is available. The original description is short: “Diffère du type par l’abdomen entièrement noir, sauf sur la base du premier segment.—Graecia, Oertzen”. This taxon could be a valid species.

3.4.4. Cleptes perezii Gogorza, 1887

Cleptes perezii Gogorza, 1887: 30. Syntypes males; Spain: Real Casa de Campo (Madrid).

Cleptes perezi: du Buysson in André 1891. Incorrect subsequent spelling.

Remarks. Gogorza [29] described Cleptes perezii based on two male specimens: “Solo poseo en mi colección dos ♂ recogidos en el mes de Junio, sobre unas umbelas de Thapsia, en la Real Casa de Campo”. Kimsey and Bohart [1] listed the holotype in Madrid, yet this specimen cannot be considered as either the holotype or the lectotype by inference of a “holotype” according to the Code (ICZN [35]: Article 74.5).

3.5. Updated Checklist of the Cleptes nitidulus Species Group

1. Cleptes adonis Rosa & Boustani, sp. nov.—Lebanon

2. Cleptes afer Lucas, 1849—North Africa, Turkey

3. Cleptes afer perezii Gogorza, 1887—Spain

     = ? Cleptes afra var. medina du Buysson in André, 1891

4. Cleptes anceyi du Buysson, 1891—Algeria, Tunisia, Morocco

5. Cleptes blaisdelli Bridwell, 1919—west USA, north-west Mexico

6. Cleptes canadensis Kimsey, 1987—Canada

7. Cleptes christi Rosa & Makris, sp. nov.—Cyprus

8. Cleptes dauriensis Móczár, 1997a—Mongolia, East Russia

9. Cleptes doii Tsuneki, 1959—Korea

10. Cleptes femoralis Mocsáry, 1890—Greece, Caucasus, Russia, Turkey

     = Cleptes antakyensis Linsenmaier, 1968

11. Cleptes helanshanus Wei, Rosa & Xu, 2013—China

12. Cleptes hungaricus Móczár, 2009—Hungary

13. Cleptes ignitus (Fabricius, 1787)—south-east Europe, North Africa, Russia, West Asia,

     Turkey, Palestine

14. Cleptes insidiosus du Buysson, 1891—Caucasus

15. Cleptes kusdasicus Móczár, 1968—Turkey

16. Cleptes laevifacies Linsenmaier, 1999—Morocco

17. Cleptes maroccanus Linsenmaier, 1987—Morocco

18. Cleptes mareki Rosa, 2003—China

19. Cleptes mayeti du Buysson, 1891—Algeria (possible synonym of C. afer Lucas, 1849)

20. Cleptes morawitzi Radoszkowski, 1877—Central Asia

21. Cleptes muti Móczár, 1968—Turkey

22. Cleptes nitidulus (Fabricius, 1793)—Europe, Russia, Turkey

     = Cleptes thoracicus Laporte in Guérin-Méneville, 1835

     = Cleptes fallax Mocsáry, 1889

23. Cleptes nyonensis Móczár, 1997a—Switzerland [not France]

24. Cleptes obsoletus Semenow, 1891—Russia (Sarepta), South-east Asia

25. Cleptes putoni du Buysson, 1886—south and central Europe to Turkey, Jordan

26. Cleptes rugulosus Linsenmaier, 1968—Palestine, Syria

27. Cleptes scutellaris Mocsáry, 1889—Hungary, Austria, south Europe, Turkey, Israel

28. Cleptes scutellaris gurunensis Linsenmaier, 1987 (possible synonym of C. anceyi du Buysson, 1891)

29. Cleptes semiatrus Linsenmaier, 1959—Palestine

30. Cleptes semicyaneus Tournier, 1879—Europe, Russia, Ukraine

     = Cleptes elegans Mocsáry, 1901

     = Cleptes nitidulus var. erdosi Móczár, 1951

31. Cleptes semenovi Kuznetzov-Ugamszkii, 1927—Central Asia: Kazakhstan (new record), Uzbekistan

32. Cleptes sfenthourakisi Rosa & Makris, sp. nov.—Cyprus

33. Cleptes sinensis Wei, Rosa & Xu, 2013—China

34. Cleptes splendidus (Fabricius, 1794)—Europe, North Africa, Georgia, Russia, Turkey, Israel, Iran

     = Cleptes caucasicus Semenov-Tian-Shanskij, 1920

     = Cleptes chyzeri Mocsáry, 1889

     = Cleptes consimilis du Buysson, 1887

     = Cleptes hyrcanus Semenov-Tian-Shanskij, 1920

     = Cleptes ignitus var. chevrieri Frey-Gessner, 1887 syn. nov.

35. Cleptes syriacus du Buysson, 1887—Israel, Turkey

36. Cleptes transoxianus Rosa, 2018—Kazakhstan

4. Discussion

The three newly described species, Cleptes adonis Rosa & Boustani, sp. nov., C. christi Rosa & Makris, sp. nov. and C. sfenthourakisi Rosa & Makris, sp. nov. show an unusual androchrome colouration of the female. Androchromy is when the female has a similar colour pattern to the male (from ancient Greek “andreîos/ἀνδρεῖος”, meaning masculine, and “chroma/χρῶμα, meaning colour). They represent the first case of this colour pattern observed in the West Palaearctic species of the C. nitidulus group. This chromatism is extremely rare in West Palaearctic species and it was previously recorded only for Cleptes (Maculosicleptes) schmidti Linsenmaier, 1968, a member of the C. dahlbomi group and C. (Holcocleptes) libanoticus Linsenmaier, 1959 a member of the C. aerosus group (see pictures of the type in Rosa et al. [25]).

In the C. nitidulus group, the female of C. mareki Rosa, 2003 (from China) is mainly black, with bluish reflections on pronotum, metanotum and mesopleuron, similarly coloured as the male, which is more blue on the head and mesosoma. Some members of the C. nitidulus group have both sexes with a red to bronze head and mesosoma: C. morawitzi, C. canadensis and C. dauriensis. In other species of the C. nitidulus group, the male is currently unknown (e.g., C. semenovi and C. transoxianus) and may have the same female colouration.

In other species groups, both sexes share the same red forebody colouration, namely in the C. (Holcocleptes) aerosus group (C. aerosus Förster, 1853; C. flammifer Semenow, 1892; C. radoszkowskii Mocsáry in Radoszkowski, 1889), and in the C. juengeri group (C. juengeri Linsenmaier, 1994). A different situation is found in Oriental groups, such as in the C. (Cleptes) asianus group and the C. (Holcocleptes) fudzi group, in which both sexes are fully metallic blue, including the metasoma.

5. Conclusions

The recent findings of two outstanding species in one Mediterranean island, where both professional entomologists, such as G.A. Mavromoustakis and amateur collectors that were active over the past seventy years, show how low the taxonomic knowledge of this genus is.