Thyreus hellenicus Lieftinck, 1968

Wood, Thomas, Leclercq, Vincent, Schmid-Egger, Christian & Praz, Christophe, 2025, A contribution to the knowledge of the genus Thyreus Panzer in the West and Central Palaearctic (Hymenoptera, Apidae), with two new species, taxonomic updates, host relationships, and a key to European species, Deutsche Entomologische Zeitschrift 72 (2), pp. 259-302 : 259-302

Thyreus hellenicus Lieftinck, 1968

Thyreus hellenicus Lieftinck, 1968: 71, ♀ ♂ [ Greece, RMNH, examined] (Fig. 15 A – D).

Material examined.

Greece • 1 ♂; Parnass [us]; 19 Jul. 1956; Bytinski-Salz leg.; M. A. Lieftinck det.; RMNH; RMNH.INS.1714335 ( holotype) • 1 ♂; Parnass [us]; 19 Jul. 1956; Bytinski-Salz leg.; M. A. Lieftinck det.; RMNH; RMNH.INS.1662710 ( paratype) • 1 ♀; Graecia, Peloponnesus, Alt-Korinth ; 3 Jun. 1964; M. Schwarz leg.; M. A. Lieftinck det.; RMNH; RMNH.INS.1662711 ( paratype) • 1 ♀; I. Kalymnos, Pothia ; 2–4 Jun. 1935; O. Wettstein; NHMW • 1 ♂; Legrena ; 1 Jun. 1965; K. V. Krombein leg.; M. A. Lieftinck det.; RMNH; RMNH.INS.1662712 ( paratype) • 3 ♀; Amorgos, Agioi Saranta ; 1 m a. s. l.; 19 Jun. 2023; V. Leclercq leg.; V. Leclercq det.; VLC / EDC • 1 ♂; Gythion [Gytheio] ; 3 Jun. 1937; F. Werner leg.; NHMW ( paratype) ; Kazakhstan • 2 ♂; Uralsk ; M. Bartel leg.; NHMW ( paratypes) ; Spain • 1 ♂; Madrid [no further information]; G. Mercet collection; MNCN; MNCN_Ent 436609 ; ( paratype) ; Syria • 1 ♂; S. Syria: Mount Hermon ; 30 Jul. 1945; G. H. Q. & M. E. F. leg.; M. A. Lieftinck det.; NHMUK ( paratype) ; Turkey • 1 ♂; Cankiri, 15 kms Ilgaz-Cankiri Rd. ; 1400 m a. s. l.; 23 Jul. 1962; Guichard & Harvey leg.; M. A. Lieftinck det.; NHMUK ( paratype) • 1 ♂; Cankiri, 15 kms Ilgaz-Cankiri Rd. ; 1400 m a. s. l.; 23 Jul. 1962; Guichard & Harvey leg.; M. A. Lieftinck det.; RMNH; RMNH.INS.1714265 ( paratype) • 1 ♂, 1 ♀; Konya, Sultan Dağları , 10 km S of Çay; 1300 m a. s. l.; 18–25 Jul. 1980; H. v. Oorschot leg.; RMNH; ZMA.INS.5144951 .

Notes.

Thyreus hellenicus is a poorly known species most commonly encountered in the East Mediterranean. Lieftinck (1968: 76) imprecisely stated “ the Whole Mediterranean region ” when discussing the distribution, despite presenting specimens from only Spain, Greece, Turkey, Syria, and north-western Kazakhstan (the southern tip of the Ural Mountains). Discussion of the range of this species combined with its host is necessary.

Thyreus hellenicus was described from Mount Parnassus in southern Greece, and most specimen records presented by Lieftinck were from this country. Specimens from Greece, Turkey, and Syria (Mount Hermon) are morphologically consistent and recognisable (see characters in the identification key below). It was possible to examine the paratype from Spain ( MNCN) from the collection of Ricardo García Mercet (Fig. 16 ). It has been dissected and is indeed T. hellenicus based on the size and genital capsule (Fig. 16 D ), as indicated by Lieftinck. The presence of this species in Spain remains perplexing, but we suggest that it may be plausible with some caveats (see below concerning host use). The two specimens from Uralsk ( NHMW) are morphologically consistent with T. hellenicus ; they were not actually labelled as T. hellenicus by Lieftinck and bore labels of “ Thyreus praevalens det. M. A. Lieftinck ” with no date, but they are clearly the specimens to which he was referring in his 1968 publication. Paratype labels have been added. The presence of T. hellenicus in western Kazakhstan is possible if broad host use of Anthophora bees of the subgenus Paramegilla Friese, 1897, is the life-history choice of this parasite (see below). Finally, we believe that Lieftinck’s comments that T. hellenicus could be present in Romania (based on the genital illustrations of T. histrionicus presented by Iuga 1958: 214) are unjustified. The genital capsule illustrated by Iuga has the gonostylus small, slender, and sparsely haired, suggesting a member of the scutellaris - group. We therefore consider the presence of T. hellenicus in Romania to be implausible based on the currently available evidence, including revision of Thyreus material from northern Greece and Bulgaria.

In addition, based on new observations on the island of Amorgos, we can demonstrate that at least one of the hosts of T. hellenicus is Anthophora ( Paramegilla) superans Walker, 1871 . On 19 June 2023, in a bare, sandy area of the preserved beach of Agioi Saranta on the island of Amorgos, Greece, one of us (VL) observed three females of T. hellenicus flying in and out of nests located in the ground, with simultaneous visitation of the same nests by Anthophora females, specifically Anthophora ( Paramegilla) superans . Moreover, An. superans was the only large anthophorine bee of the same size as T. hellenicus present at the collecting event (and possibly even on the island of Amorgos) at this time. Further support for this association is the paratype of T. hellenicus collected by Max Schwarz at Corinth on 3 June 1964; he also collected a long series of An. superans at the same moment (specimens OÖLM / TJWC; see Suppl. material 1). The same can be said for the paratype from Gytheio on 3 June 1937 ( NHMW), which was also collected with several An. superans (specimens NHMW).

The name of the Anthophora host requires explanation, as this name has not previously been used for the European fauna. The presence of a large Paramegilla species in southern and Aegean Greece has been known for a long time; Dours (1869: 120) used the name Anthophora dubia Eversmann, 1852 , giving a distribution of “ Îles de l’Archipel grec, montagnes de l’Oural, province d’Orenbourg, d’après Eversman ”. The identity of An. dubia is currently being dealt with elsewhere (Rasmont et al., in review), but in short, An. dubia was described from Orenburg in southern European Russia and comprises a mixed type series (males and females incorrectly associated). Neither taxon within this type series is conspecific with the Greek bee, which is an East Mediterranean species. Searches were therefore made for the appropriate name to apply to this large and conspicuous Anthophora species.

Examination of type material has concluded that the oldest names that can be applied are either Anthophora ( Paramegilla) inclyta Walker, 1871 , or Anthophora ( Paramegilla) superans Walker, 1871 . These two names come from Walker’s (1871) work on Hymenoptera collected around the Red Sea in Egypt, Arabia, and Eritrea. Francis Walker (1809–1874) was a prolific worker who has been much criticised for his abundant and superficial descriptions; the lack of respect for the work of Walker by his contemporaries was clear at the time (see the 1874 obituary cited in Evenhuis 2008), a sentiment echoed by Baker (1993: 300), who commented on the case of An. inclyta and An. superans . Much of the material described in Walker (1871) has been lost (e. g. see Monks et al. 2024), but some specimens of An. inclyta and An. superans were preserved in the collection of Frederick Smith and are now held in the NHMUK ( Baker 1993; see Other material examined, below).

Walker (1871: 58, no. 287) described An. inclyta in the female sex from “ Rafla, Wady Ferran, Mount Sinai ”. This corresponds to Arafali / Irafayle in Eritrea, Wadi Feiran in Sinai, and Mount Sinai in the southern part of the Sinai Peninsula. He then ( Walker 1871: 58, no. 288) described An. superans in the “ female ” sex from Mount Sinai only. In the NHMUK collection, there are 3 ♀ of An. inclyta and 4 ♂ of An. superans . Although Walker wrote “ female ” for An. superans , he described the head as “ Head, excepting the vertex white and without hairs ”, suggesting a male, since the males have the clypeus, supraclypeal area, and lower paraocular areas with the integument yellow-white (Fig. 17 C ). It is therefore considered that An. superans was described from the male sex. A dissected male was labelled as “ lectotype ” by Baker in 1979 and by Brooks in 1983 (Fig. 17 A ), but neither of these designations was published. Although Baker (1993) wrote that it was designated as such, this was published in his PhD thesis, which does not meet the ICZN (1999) criteria for a valid publication. Under Articles 8 and 9, as the thesis was not a formally published work and only seven hard copies were produced, and these limited numbers were not obtainable either for free or for a charge, it fails Articles 8.1. 2 and 8.1. 3. This specimen from the Convent Garden on Mount Sinai (Saint Catherine’s Monastery; 28.55°N, 33.98°E) is therefore finally published here as the lectotype of An. superans , by present designation. It is conspecific with the material from Greece based on the hind basitarsi, which have a rounded bump apically (Fig. 17 D ; no tooth or teeth), and based on the ventro-lateral projection of the gonocoxa, which is reduced (Fig. 17 F ; lectotype specimen with the genital capsule fixed in a capsule with a glycerine-like substance). This reduction in the lateral projection of the gonostylus allows separation from the West Mediterranean Anthophora ( Paramegilla) ferruginea Lepeletier, 1841 (compare Fig. 18 E, F ). Indeed, the genital capsule is so similar (Fig. 18 C, D ) that Lieftinck treated material ( RMNH) from the East Mediterranean as An. ferruginea ssp. dubia ; these specimens have been re-determined (see Suppl. material 1). The two species can also be separated by their tergal pubescence (Fig. 18 A, B ).

For the specimens of An. inclyta , the situation is more complex. The three available female specimens are from the Convent Garden on Mount Sinai (Fig. 19 A ) and Rafla in Eritrea (Fig. 20 A ). Whilst morphologically similar, it is not clear if they are actually conspecific. One character often seen in the East Mediterranean taxon is that the female has a slightly raised longitudinal carina medially, which is sometimes coloured yellow, and the anterior margin of the clypeus slopes towards the apical margin with weak sculpturing, becoming almost shining. This is present in the specimens from Sinai but not in the specimen from Eritrea. Although the specimen from Eritrea was designated as the lectotype by Baker in 1979, this was never validly published, as with An. superans . Baker (1993: 310) selected the Eritrean specimen as the lectotype, as he considered the females from Sinai to be conspecific with An. superans .

Given the lack of a validly published lectotype, the existence of syntypic specimens from Sinai, and the fact that no one has previously acted as a first reviser in a way compliant with ICZN rules ( Brooks 1988 considered both taxa as distinct), under Article 24.2, as First Reviser, we select a syntypic female of An. inclyta from the Convent Garden (Mount Sinai) as lectotype by present designation, and given the simultaneous publication of An. superans and An. inclyta by Walker (1871) with no objective way to determine precedence, we select An. superans to have precedence over An. inclyta syn. nov. Based on this synonymy and establishment of An. superans as the priority name for An. dubia sensu auctorum in the East Mediterranean, a total of 48 ♀ and 34 ♂ could be examined (including type material; see Suppl. material 1), producing a revised distribution of An. superans covering Italy (first record), Greece, Turkey, Armenia, Israel and the West Bank, and Egypt (Fig. 21 ), and (nominally based on its status as a syntype, though the specimen is probably not conspecific due to its differently sculptured clypeus; revision is required) Eritrea. The species is likely to be present in Jordan, Lebanon, Syria, and Iran once additional material has been studied. Baker (1993: 310) gave the range as “ from the Balkans eastwards through the Levant to the Urals and Caucasus ”. We partially agree, citing a need to examine additional material from the Caucasus and Iran, though we consider the Ural Mountains implausible for ecological and biogeographical reasons given the apparent thermophilic tendencies of this East Mediterranean species.

Given the morphology of An. superans , it is relevant to consider the West Mediterranean An. ferruginea . This species is found in southern Spain, Morocco, Algeria, and Tunisia (Fig. 21 ; described from Oran in Algeria); it is particularly common in south-eastern Spain. Given the very scant label details of the MNCN paratype of T. hellenicus , it is not impossible that the specimen (the only known specimen of T. hellenicus collected in Spain) was actually from south-eastern Spain, rather than “ Madrid ”. Indeed, we have never seen any An. ferruginea from the Madrid region, with other Anthophora ( Paramegilla) species occurring there, such as An. balneorum and An. femorata . Given these uncertainties, we cautiously suggest An. ferruginea as a possible host of T. hellenicus in Spain, pending further study.

In conclusion, given the association between T. hellenicus and an Anthophora ( Paramegilla) species, a subgenus strongly diversified in Turkey, Iran, the Caucasus, southern Russia, and Central Asia, it is difficult to comment on either (i) the overall eastern distribution of T. hellenicus or (ii) the number of closely related and insufficiently characterised Thyreus species that may be present in this area.

Distribution.

Spain (precise location uncertain), Greece, Turkey, Syria, Iran, Kazakhstan ( Lieftinck 1968; Ascher and Pickering 2025). The listing of Iran comes from material determined by Baker hosted in his collection at the Kansas museum. Given Baker’s knowledge of this genus, we accept the record as plausible. Listings from Morocco, Liechtenstein, and Ukraine ( Ascher and Pickering 2025) are from the American Museum of Natural History species database and are excluded at the present time due to a lack of precise supporting data.

Other material examined.

Anthophora superans Egypt • 1 ♂; Convent Garden, Mount Sinai; NHMUK ( lectotype by present designation) • 2 ♂; Gebel Musa ( Mt. Sinai) [ Jabal Mousa]; NHMUK ( paralectotypes) • 1 ♂; Sandy plains ( Mt. Sinai); NHMUK ( paralectotype) ; Italy • 1 ♀; Sicilia; 1858; Mann leg.; NHMW .

Anthophora inclyta Egypt • 2 ♀; Convent Garden, Mount Sinai; NHMUK ( lectotype by present designation and paralectotype) ; Eritrea • 1 ♀; Rafla ( Annesley Bay ) [Arafali / Irafayle]; NHMUK ( paralectotype) .

Additional examined non-type specimens (all under the name An. superans ) are presented in Suppl. material 1.

Figures

Figure 15. Thyreus hellenicus Lieftinck, 1968 holotype male (RMNH). A. Label details; B. Habitus, lateral view; C. Face, frontal view; D. Metasoma, dorsal view; Paratype male (RMNH); E. Genital capsule, dorsal view; F. Genital capsule, lateral view.

Figure 16. Thyreus hellenicus Lieftinck, 1968 paratype male (MNCN). A. Label details; B. Profile; C. Metasoma, dorsal view; D. Genital capsule, dorsal view.

Figure 17. Anthophora (Paramegilla) superans Walker, 1871 lectotype male (NHMUK). A. Label details; B. Habitus, lateral view; C. Head, frontal view; D. Hind leg, lateral view; E. Metasoma, dorsal view; F. Genital capsule, inverted along the transverse plane, lateral view.

Figure 18. Anthophora (Paramegilla) superans Walker, 1871 male. A. Metasoma, dorsal view. C. Genital capsule, dorsal view. E. Genital capsule, dorso-lateral view. Anthophora (Paramegilla) ferruginea Lepeletier, 1841 male; B. Metasoma, dorsal view. D. Genital capsule, dorsal view. F. Genital capsule, dorso-lateral view.

Figure 19. Anthophora (Paramegilla) inclyta Walker, 1871 female lectotype (NHMUK). A. Label details; B. Habitus, lateral view; C. Head, frontal view; D. Metasoma, dorsal view.

Figure 20. Anthophora (Paramegilla) inclyta Walker, 1871 female paralectotype (NHMUK). A. Label details; B. Habitus, lateral view; C. Head, frontal view; D. Metasoma, dorsal view.

Figure 21. Distribution of Thyreus hellenicus Lieftinck, 1968 (red circles) along with its host Anthophora (Paramegilla) superans Walker, 1871 (open blue triangles) and possible host Anthophora (Paramegilla) ferruginea Lepeletier, 1841 (filled yellow squares). Distributional data are not comprehensive.