Subgenus Osmia (Allosmia) Tkalců, 1974 Morphological diagnosis Osmia (Allosmia) species are non-metallic, slender and medium-sized bees (6–12 mm) with short-linear, in O. bischoffi Atanassov punctiform parapsidal lines and distinctly keeled to carinate hind coxae. The females are characterized by their yellowish-red metasomal scopa in combination with the coarsely punctate to rugose clypeus, whose apical zone is impunctate, polished, transversally weakly impressed behind its straight apical margin, and distinctly separated from the clypeal disc, often by a short transverse groove; O. bischoffi differs from all other species in that the apical zone of the clypeus is partly punctate along its base, less distinctly separated from the clypeal disc and apically emarginate. The males are characterized by two-toothed mandibles (indistinctly threetoothed in O. bischoffi) in combination with the shape of tergum 7, which is either predominantly flat and apically bifid or laterally distinctly curved downwards and apically rounded to truncate. Biology Pollen hosts. The species of Osmia (Allosmia) are pollen generalists collecting pollen on up to 13 plant families (see species accounts for details). However, in all species, for which a reasonable number of pollen loads was available for study, flowers of the Fabaceae turned out to be the most important pollen hosts. Depending on the species, pollen of this plant family was recorded in 60.0 % to 90.9 % of the pollen samples analysed. The finding that 80.9 % of the 188 pollen loads analysed contained 2–7 different pollen types often belonging to different plant families suggests low flower constancy of pollen-collecting O. (Allosmia) females. Nesting biology. All species of O. (Allosmia), for which information on the nesting biology is available, utilize empty snail shells as exclusive nesting sites (see species accounts for details), indicating that snail shell nesting is a subgeneric trait (Figs 1–5) In contrast to all other obligate snail shell nesters among the osmiine bees, which build more than one brood cell per shell if the available space allows, species of O. (Allosmia) invariably construct only one cell per shell (Figs 4, 5; Müller et al. 2018). The nesting biology slightly differs among the different O. (Allosmia) species. Osmia rufohirta Latreille (and possibly also other representatives of the O. rufohirta species group) differs from representatives of the O. sybarita species group in that i) the nest plug is usually three-layered consisting of two partitions of leaf pulp enclosing a narrow space filled with small stones, earth crumbs, plant fibers or other foreign particles (Fig. 5), ii) the nest plug is usually more or less hidden inside the shell (Fig. 5), iii) the females cover the shell surface with patches of leaf pulp before and often also during cell provisioning (Figs 2, 5) and iv) the sealed nest is usually concealed under stones, below leaves or in grass tussocks (Fig. 2). In the representatives of the O. sybarita species group, the nest plug consists of a mixture of leaf pulp and mollusc shell fragments (Figs 3, 4), it is built at the shell opening (Figs 3, 4), the shell surface is not covered with leaf pulp patches (Figs 1, 3, 4) and the sealed nest is usually buried in loose soil (Fig. 3). The adaptive value of covering the shell surface with leaf pulp patches is not understood. This behaviour, which is also known from species of O. (Neosmia) and from O. (Helicosmia) aurulenta (Panzer) (Müller et al. 2018), has alternatively been interpreted as an evolutionary relict inherited from an ancestor that did not colonize preexisting cavities but constructed free standing brood cells (Bellmann 1981), as a camouflage strategy to reduce the optical conspicuousness of the white shells (Grozdanić 1969) or as a means to facilitate the movement of the shell (Malyshev 1937). Interestingly, instead of gluing patches of leaf pulp on the outer shell surface, some representatives of the O. sybarita species group, such as O. melanura Morawitz and O. rutila Erichson, attach portions of leaf pulp to the inner shell surface few millimeters behind the shell opening immediately after nest site selection (Müller 1992; Haeseler 1997). Experiments showed that this leaf pulp markings near the nest entrance allow the females to recognize their own nests (Haeseler 1997). Thus, covering the outer shell surface with patches of leaf pulp might possibly serve the same purpose, i.e. to individually mark the shell and to signal other females that the shell is already occupied. Taxonomy Michener (2007) included the representatives of the Osmia sybarita species group (see below) in the subgenus O. (Erythrosmia) Schmiedeknecht and recognized O. rufohirta and its closest relatives as the only members of the subgenus O. (Allosmia). However, this placement is neither supported by molecular phylogenetic analyses, which placed O. sybarita Smith as sister to O. rufohirta (Praz et al. 2008; Rightmyer et al. 2013), nor by morphology and biology, which both are more similar between O. sybarita and O. rufohirta than between O. sybarita and O. andrenoides Spinola, the type species of O. (Erythrosmia). Thus, in the present study, we consider O. sybarita and its relatives to belong to O. (Allosmia), following other European authors (e.g. Tkalců 1974; Zanden 1988b; Ungricht et al. 2008). Species accounts Osmia (Allosmia) bischoffi Atanassov, 1938 Osmia bischoffi Atanassov, 1938: 180. Type material: Holotype ♂, “Im Park des Palais Euxinograd bei Warna am Schwarzen Meer” (Bulgaria), National Museum of Natural History Sofia. Literature records. TURKEY: Bayburt, Erzurum (Özbek & Zanden 1992; Özbek 2013). New records. ALBANIA: Gjirokastra: Tepelena, 40°17ʹ23ʺN / 20°01ʹ35ʺE, 125 m, 24.4.2017, 1♀ (leg. A. Rey). BOSNIA AND HERZEGOVINA: Srpska: Ljubovo, 44.643°N / 15.492°E, 200 m, 2♀ (leg. M. Kafka). BULGARIA: Varna: Zlatni Pjasaci, 26.5.1983, 1♀ (leg. L. Norén). CROATIA: Lika-Senj: Baske Ostarie N Velebit, 26.5.2011, 1♀ (leg. Z. Jozan); Split-Dalmatia: 40 km N Split, 43°47.9ʹN / 16°34.1ʹE, 370 m, 29.5.2005, 3♀ (leg. M. Halada). GREECE: Central Greece: Tymfristos, 1990 m, 1♀ (leg. A. W. Ebmer); Eastern Macedonia and Thrace: Nomos Drama, Falakró, 41°17ʹ40ʺN / 24°02ʹ05ʺE, 1400–1800 m, 15.– 16.6.2012, 2♀ (leg. A. W. Ebmer); Epirus: Igoumenitsa, 200 m, 17.4.1994, 1♀ (leg. S. Becvar); 30 km W Ioannina, 16.5.2005, 2♀ (leg. M. Halada); N Tyria, 400 m, 1♀ (leg. A. W. Ebmer); Peloponnese: ancient Mykene, 27.4.2000, 1♀ (leg. W. Arens); ancient Korinth, 21.4.1995, 2♀ (leg. W. Arens); West Macedonia: Pentalofos pass, 1500–1700 m, 1♀ (leg. A. W. Ebmer). ROMANIA: Mehedinti: SW Orsova, 25.5.2002, 6♀ (leg. M. Snizek); Cozla, 50 km W Turnu Severin, 25.– 26.5.2002, 36♀ (leg. M. Halada, Z. Pedr). TURKEY: Bilecik: Bilecik, 4.4.1987, 1♂ (leg. Menrad); Erzurum: Basakli, 20.7.1979, 1♀ (leg. H. Özbek); Sutkans, Oltu, 2000 m, 10.6.1997, 1♀ (leg. L. Gültekin); 25 km SSW Oltu, 40.29°N / 41.47°E, 18.5.2002, 1♀ (leg. J. Rozen); 2 km NW Gecitköy, 5.7.2007, 1♀ (leg. J. Ascher, J. Rozen, H. Özbek); Ilica, Agziacik Gecidi, 40°16ʹN / 40°59ʹE, 2295 m, 3.7.2008, 1♀ (leg. J. Rozen, H. Özbek); Kütahya: Porsuk Baraji, 30 km N Kütahya, 22.5.1998, 2♀ (leg. M. Halada); Sakarya: Adapazari, 12.5.1964, 1♀ (leg. K. Warncke); Tokat: Yolüstü, 40°28.79ʹN / 37°16.86ʹE, 1250 m, 20.5.2007, 1♀ (leg. W.- H. Liebig). Distribution. From southeastern Europe (Croatia, Bosnia and Herzegovina, Albania, Greece, Romania, Bulgaria) to eastern Turkey. Pollen hosts. Polylectic (at least 13 plant families): Fabaceae (Hedysareae, Loteae, Trifolieae; n = 15 pollen loads with Fabaceae pollen), Asteraceae (Asteroideae, Cichorioideae; n = 9), Brassicaceae (n = 8), Monocots (n = 6), Convolvulaceae (n = 5), Cistaceae (n = 3), Caryophyllaceae (n = 2), Ranunculaceae (n = 2), Boraginaceae (n = 1), Dipsacoideae (n = 1), Lamiaceae (Nepetoideae; n = 1), Plantaginaceae (Plantago; n = 1) and Crassulaceae (n = 1) (based on 25 pollen loads from 14 different localities in Albania, Croatia, Greece, Romania and Turkey). Nesting biology. Osmia bischoffi nests in empty snail shells (Atanassov, 1938; A. W. Ebmer, personal communication). The females cover the surface of the nest shells with patches of leaf pulp. Osmia (Allosmia) imitatrix (Tkalců, 1992) Hoplitis (Allosmia) imitatrix Tkalců, 1992: 219. Type material: Holotype ♀, “Kara-kala” (Turkmenistan), Halada Collection (České Budějovice, Czech Republic). New records. TURKMENISTAN: Ahal: Firjuza-Vanovski, 23.–26.4.?, 1♀ (leg. S. Bečvàř). Distribution. Known so far only from two localities in the Kopet Dag mountain range in southern Turkmenistan. Pollen hosts. Unknown. Nesting biology. Unknown. Note. Male unknown. Osmia rufohirta species group The representatives of the O. rufohirta species group differ from those of the O. sybarita species group in both sexes by the distinctly wider head and in the male sex by the shape of tergum 7, which is laterally distinctly curved downwards and apically rounded to truncate, as well as the presence of a pyramidal or triangular projection on sternum 2. The two groups probably differ also in several characteristics of their nesting biology (see above). Osmia (Allosmia) gemina spec. nov. Holotype. ISRAEL AND PALESTINE: Central District: Tel Yizhaq, 32°14ʹ34ʺN / 34°51ʹ53ʺE, 20 m, 13.2.2010, ♂ (leg. A. Dorchin). Deposited in the Entomological Collection of ETH Zurich. Paratypes. ISRAEL AND PALESTINE: Central District: Bnei Tsiyon, 32°13ʹ16ʺN / 34°51ʹ25ʺE, 35 m, 25.2.2009, 1♀ (leg. A. Dorchin), 16.2.2010, 1♂ (leg. A. Dorchin); Harutsim, 32°13ʹ47ʺN / 34°51ʹ35ʺE, 35 m, 26.2.– 30.3.2009, 6♀, 11♂ (leg. A. Dorchin); Netanya, 11.3.– 12.4.2009, 5♀, 2♂ (leg. A. Dorchin); Tel Yizhaq, 32°14ʹ34ʺN / 34°51ʹ53ʺE, 20 m, 13.2.– 27.2.2010, 2♂ (leg. A. Dorchin); Yakum, 19.3.2009, 1♀, 1♂ (leg. A. Dorchin); Northern District: Gilboa Mt., 1 km N Gan Ner, 32.5478°N / 35.33924°E, 50 m, 24.2.2018, 1♂ (leg. A. Dorchin); Southern District: Fura NR, 6.3 km E Ruhama, 31°29ʹ47ʺN / 34°46ʹ33ʺE, 196 m, 18.3.2010, 1♀ (leg. A. Dorchin), 15.4.2012, 1♀ (leg. A. Dorchin); Lakhish, 31.5562°N / 34.869°E, 18.2.– 9.3.2020, 1♀, 2♂ (leg. K. Levy). JORDAN: Dscharasch: 10 km N Jerash, 20.4.2002, 1♀ (leg. M. Snizek); Alhuna, SW Jerash, 12.4.2009, 1♀ (leg. M. Snizek); Irbid: Tall al Arbatin, 20km S North Shuna, 19.4.1996, 3♀ (leg. M. Halada); North Shuna, 20.– 22.4.1996, 1♀ (leg. M. Halada); S Irbid, 13.4.2009, 1♀ (leg. M. Snizek). Deposited in the entomological collections of ETH Zurich, the Steinhardt Museum of Natural History Tel Aviv, the Hebrew University Jerusalem and the Oberösterreichisches Landesmuseum Linz. Other records. ISRAEL AND PALESTINE: Southern District: Judean Foothills, Lakhish, 6.3.2013, 1♂ (leg. T. Shapira); Lakhish, 31.5562°N / 34.869°E, 4.3.2016, 1♂ (leg. G. Pisanty); Lehavim, 31.370°N / 34.8257°E, 7.3.2015, 3♀ (leg. G. Pisanty); Tel Qeriyyot, 31.342°N / 35.125°E, 27.3.2015, 1♀, 1♂ (leg. G. Pisanty). Diagnosis. Osmia gemina is in both sexes morphologically very close to O. soror and O. rufohirta (Figs 6, 7). The females differ from O. soror by the shorter tergal hair bands, which surpass the apical margin of terga 2–4 by less than half of their length, and from O. rufohirta by the darker colour of the marginal zone of terga 1–5, which is predominantly black to narrowly dark reddish-brown; outside the southern Levant, where O. gemina does not occur, females of O. rufohirta may also have more or less darkened tergal margins. The males differ from O. rufohirta by the shaggy yellowish pilosity at the marginal zone of sterna 4–5 consisting of long and suberect hairs not forming bands (Figs 8, 9), by the polished, sparsely punctate and almost unhaired roundish preapical depression of sternum 6 (Figs 8, 9) and by the apically less broadened and less inwardly bent gonoforceps (Figs 10, 11). They differ from O. soror by the shorter pilosity along the inner margin of the apical half of the gonoforceps, which is distinctly less than half as long as the pilosity along the outer margin (Fig. 10), by the smaller length of the longest hairs on tergal discs 4–5, which are distinctly shorter than tarsal segment 2 of the hind leg, and often also by the dark rather than reddish marginal zone of terga 1–5. Description. Except for the characters given in the diagnosis and the identification key, both sexes of O. gemina are morphologically identical to the widespread and well-known O. rufohirta. Therefore, no detailed description of the new species is given here. Distribution. Central and northern Israel and northern Jordan.At five localities in Israel (Harutsim, Lakhish, Tel Yizhaq, Yakum) and Jordan (North Shuna), O. gemina was found to cooccur with O. rufohirta indicating syntopic occurrence of these two closely related species within the distribution range of O. gemina. Pollen hosts. Polylectic (at least 4 plant families): Fabaceae (Hedysareae, Loteae, Trifolieae; n = 3 pollen loads with Fabaceae pollen), Asteraceae (Asteroideae, Cichorioideae; n = 3), Brassicaceae (n = 1) and Plantaginaceae (Plantago; n = 1) (based on 3 pollen loads from 3 different localities in Israel and Palestine). Nesting biology. Unknown. Etymology. The scientific name refers to the close morphological similarity with the sister species O. rufohirta Latreille, 1811 (lat. “geminus” = twin). Osmia (Allosmia) nuda Friese, 1899 Osmia nuda Friese, 1899: 328. Type material: Lectotype ♀, by designation of G. van der Zanden (unpublished), “ Balkan ” (Balkans), Museum für Naturkunde Berlin. Literature records. TURKEY: Bursa (Bursa) (Friese 1899). New records. BULGARIA: Burgas: 5 km E Zvedec, 42°05ʹ17ʺN / 27°29ʹ01ʺE, 4.5.2006, 1♀ (leg. J. Smit); Varna: Zlatni Pjasaci, 22.– 26.5.1983, 6♀, 1♂ (leg. L. Norén). CROATIA: Zadar: Biograd, 1.7.1966, 1♀ (leg. Hoffer). TURKEY: Adiyaman: Nemrut, 8.6.1992, 1♂ (leg. M. Hradsky); Kütahya: Porsuk Baraji, 30 km N Kütahya, 15.6.1997, 1♂ (leg. M. Halada). Distribution. From southeastern Europe (Croatia, Bulgaria) to central Turkey. Pollen hosts. Polylectic (at least 3 plant families): Fabaceae (Trifolieae; n = 1 pollen load with Fabaceae pollen), Convolvulaceae (n = 1) and Lamiaceae (Nepetoideae; n = 1) (based on 2 pollen loads from 2 different localities in Bulgaria). Nesting biology. Unknown. Note. Warncke (1986) treated O. nuda as junior synonym of Chelostoma ventrale Schletterer, which was correctly rejected by Özbek & Zanden (1992). Osmia (Allosmia) rufohirta Latreille, 1811 Osmia rufo-hirta Latreille, 1811: 580. Type material: Syntypes ♀ ♀, “ France ” (France), “Allemagne” (Germany), type depository unknown. Type species of Allosmia Tkalců. Osmia fulvo-hirta Lepeletier, 1841: 322. Type material: Lectotype ♀, by designation of Tkalců (1974), “Environs de Paris” (France), Muséum National d’Histoire Naturelle Paris. Synonymy in Dalla Torre (1896). Osmia spiniventris Giraud, 1857: 181. Type material: Syntypes ♂♂, “ Autriche ” (Austria), “Carniole” (Slovenia), “ Italie ” (Italy), “ Hongrie ” (Hungary), type depository unknown. Synonymy in Dalla Torre (1896). Literature records. ALBANIA ( Tkalců 1974). AZERBAIJAN: Göygöl (Ducke, 1900). AUSTRIA: Gusenleitner et al. (2012). BELARUS: Prishchepchik (2000). BELGIUM: Pauly (1999), Pauly et al. (2018). CROATIA: Jozan (2009). CHINA: Xinjiang (Hetienkashentaxian) (Wu, 2006). CZECH REPUBLIC: Bogusch et al. (2007). FRANCE including Corsica (Benoist 1931). GEORGIA: Kirkitadze & Japoshvili (2015). GERMANY: Scheuchl & Willner (2016). HUNGARY: Józan (2011). IRAN: Alborz, East Azerbaijan, Mazandaran (Ascher & Pickering, 2020). ITALY including Sardinia and Sicily: Pagliano (1994, 1995). KAZAKHSTAN: Turkistan (Bairkum) (Morawitz, 1875). LIECHTENSTEIN: Bieri (2002). LUXEMBOURG: Rasmont et al. (1995). MALTA: Balzan et al. (2016). NORTH MACEDONIA: Vardar (Stobi) (Zanden (1984a). PORTUGAL: Baldock et al. (2018). ROMANIA: Ban-Calefariu (2009). RUSSIA: Astrachan, North Caucasus (Ducke, 1900; Proshchalykin & Fateryga 2017). SERBIA: Mudri-Stojnić et al. (2021). SLOVAKIA: Bogusch et al. (2007). SLOVENIA: Gogala (1999). SPAIN: Ortiz-Sánchez (2020). SWITZERLAND: Amiet et al. (2004). TURKEY: Adana, Ankara, Antalya, Aydin, Denizli, Diyarbakar, Erzurum, Hatay, Izmir, Karaman (Özbek & Zanden 1992; Özbek 2013). UKRAINE including Crimea (Scheuchl & Willner, 2016; Fateryga et al., 2018). New records. ARMENIA: Kotayk: Gekhard, 17.5.1978, ♀ (leg. M. Kocourek). AZERBAIJAN: Nakhichevan: Julfa, Gazanchi, 39°13ʹN / 45°41ʹE, 1300 m, 15.6.2019, ♀ (leg. M. Proshchalykin). BULGARIA: Burgas: Slancev brjag, 1.6.1972, f (leg. M. Kocourek); Haskovo: 5 km NE Harmanli, 41°57ʹN / 25°57ʹE, 200 m, 14.6.2008, 2♀ (leg. M. Halada); Kardschali: Balabanovo, 41°34ʹN / 25°22ʹE, 300 m, 22.6.2007, 4♀ (leg. M. Halada); Stara Zagora: Galobovo, 1.7.1997, ♀ (leg. A. Zaykov); Vratsa: Voivodovo, 18.5.1996, ♀, ♂ (leg. A. Zaykov). GREECE: Aegean Islands: Lesvos, Eresos, 39.1771°N / 25.946°E, 8.4.2012, ♀ (leg. G. Nakas); Central Greece: 30 km S Lamia Bralos, 10.5.2005, 3♀ (leg. J. Halada); Epirus: 10 km NE Ioannina, Published as part of Müller, Andreas, 2022, Palaearctic Osmia bees of the subgenera Allosmia and Neosmia (Megachilidae Osmiini): biology, taxonomy and key to species, pp. 201-232 in Zootaxa 5188 (3) on pages 202-216, DOI: 10.11646/zootaxa.5188.3.1, http://zenodo.org/record/7091605, {"references":["Tkalcu, B. (1974) Ergebnisse der Albanien-Expedition 1961 des ' Deutschen Entomologischen Institutes'. 89. Hymenoptera: Apoidea V (Megachilidae). Beitrage zur Entomologie, 2 4, 323 - 348.","Muller, A., Praz, C. & Dorchin, A. 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