Osmia (Allosmia) Tkalců, 1974

Müller, Andreas, 2022, Palaearctic Osmia bees of the subgenera Allosmia and Neosmia (Megachilidae Osmiini): biology, taxonomy and key to species, Zootaxa 5188 (3), pp. 201-232 : 202-216

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 , 1.7.1996, ♀ (leg. M. Halada) ; Ionian Islands: Kefallonia , between Poros and Skala, 28.4.1996, ♀ (leg. C. Schmid-Egger) ; Peloponnese: Mykene , 11.4.2000, 4♀, ♂ (leg. W. Arens) ; Thessaly: 40 km NE Larissa, Mt. Ossa, Kokino Nero , 13.5.2005, ♀ (leg. J. Halada); Western Macedonia: Kastoria , 8.6.1987, ♀ (leg. P. Thomas) . ISRAEL AND PALESTINE: Central District: 0.75 km N Yaqum, 32°15ʹ20ʺN / 34°50ʹ33ʺE, 12 m, 5.3.2010, 2♂ (leg. A. Dorchin); Haifa District : 850 m S Bet Oren , 32°43ʹ18ʺN / 35°0 ʹ20ʺ'E, 256 m, 31.3.2012, 2♀ (leg. A. Dorchin); Jerusalem District : Judean Foothills, Neve Shalom, 6.5.2009, ♀ (leg. G. Pisanty); Northern District : Dovev, 14.4.2016, ♂ (leg. O. Winberger); Southern District : 2 km NW Bet Nir, 31.658°N / 34.855°E, 5.4.2015, ♂ (leg. G. Pisanty) . IRAN: Gilan: Rudsar-Ghazichak , 10.5.2010, 1♀ (leg. A. Nadimi) ; North Khorasan: Ziarat , 36.68°N / 54.563889°E, 5.7.2018, 1♀ (leg. W.- H. Liebig) . JORDAN: Irbid: North Shuna , 22.4.1996, 6♀ (leg. M. Halada) . LEBANON: Boustani et al. (2021) . MOROCCO: Fès-Meknès: Michlifene , 33°24ʹ51ʹʹN / 5°04ʹ35.6ʹʹW, 1900 m, 7.5.2015, 1♂ (leg. V. Soon) . SYRIA: Idlib: Jisr ash Shughur , 26.5.1996, 2♀ (leg. M. Halada) .

Distribution. Osmia rufohirta is a widespread species distributed in northern Morocco (known so far only from a single locality at 1900 m a.s.l. in the Middle Atlas), in southern, central and eastern Europe northwards to about 52 o northern latitude ( Albania, Austria, southern Belarus, southern Belgium, Bulgaria, Croatia, Czech Republic, France, southern and central Germany, Greece including Aegean and Ionian Islands, Hungary, Italy including Sardinia and Sicily, Liechtenstein, Malta, North Macedonia, Portugal, Romania, southernmost Russia, Serbia, Slovakia, Slovenia, Spain, Switzerland, Ukraine including Crimea), in Turkey eastwards over the Caucasus ( Armenia, Azerbaijan, Georgia), and northern Iran to Central Asia (southern Kazakhstan) and China (Xinjiang) as well as in the Levant ( Israel and Palestine, Jordan, Lebanon, Syria).

Pollen hosts. Polylectic (at least 9 plant families): Fabaceae (Fabeae, Galegeae , Genisteae , Hedysareae , Loteae , Trifolieae ; n = 28 pollen loads with Fabaceae pollen), Asteraceae (Asteroideae, Carduoideae, Cichorioideae; n = 19), Brassicaceae (n = 9), Lamiaceae ( Lamioideae , Nepetoideae ; n = 5), Boraginaceae ( Echium ; n = 3); Campanulaceae (n = 2), Convolvulaceae (n = 2), Caryophyllaceae (n = 1) and Cistaceae (n = 1) (based on 31 pollen loads from 23 different localities in France, Greece, Israel and Palestine, Italy, Jordan, Romania, Spain, Syria and Turkey. Flower records: Helianthemum nummularium , Hippocrepis comosa , Lotus corniculatus , Onobrychis viciifolia ( Westrich 1989) ; Hedysarum tauricum ( Fateryga 2017) ; Onobrychis viciifolia ( Özbek 2013) ; Campanula spec. , Centaurea pallescens , Chrysanthemum coronarium , Crepis aspera , Cynoglossum officinale , Echium judaeum , Diplotaxis erucoides , Genista hispanica , Lathyrus aphaca , Scorpiurus muricatus , Spartium junceum , Trifolium resupinatum , T. stellatum , Vicia tenuifolia , V. villosa , Salvia fruticosa , Thymus vulgaris (label records).

Nesting biology. Osmia rufohirta nests in empty snail shells of small to medium size, e.g. of Candidula, Cernuella , Helicella, Pomatias , Rumina , Theba , Xerolenta, Xerophila or Zebrina ( Figs 2, 5 ; Ferton 1894, 1897, 1905; Grandi 1961; Grozdanić 1969; Bonelli 1971, 1972; Bellmann 1981; Westrich 1989; Gogala 1999; G. Le Goff personal communication). After nest site selection, the females cover the surface of the nest shells with numerous small patches of leaf pulp ( Figs 2, 5 ). The nests contain a single brood cell, which lacks a basal wall that seals the larval provisions against the rear end of the shell ( Fig. 5 ). After provisioning, leaf pulp originating from leaves (e.g. Helianthemum ) or occasionally petals (e.g. Lotus ) is amassed in the area of the later cell partition, before an egg is laid and the accumulated leaf pulp is processed to a one-layered partition. Immediately in front of this partition, numerous small stones, earth crumbs, plant fibers or other foreign particles are piled up over a length of about 0.5 cm, before another wall of leaf pulp is constructed, resulting in a three-layered plug ( Fig. 5 ). This plug is usually more or less hidden inside the snail shell ( Fig. 5 ), but may also be built near the shell opening. In rare cases, the nest plug consists of four to five one-layered partitions of leaf pulp enclosing three to four spaces, which are 3-18 mm long and densely filled with foreign particles. After the nest has been sealed, it is rolled over a distance of up to more than 2 m to a previously selected protected place, such as the underside of a stone, below a withered leaf or among the dead blades of a grass tussock. To move the nest shell, the female grasps a plant stem with her mandibles and rolls the shell, which weighs seven to eight times her own weight ( Ferton, 1894), with the help of her legs often through dense vegetation ( Fig. 2 ). Regularly, the nest is also rolled to a more suitable place or turned into a favourable position before or during cell provisioning. In rare cases, the finished nests are buried very shallowly in loose soil. Nesting cycle: Osmia rufohirta overwinters as imago in a self-spun cocoon within the brood cell ( Bellmann, 1981). Brood parasites: Chrysura cuprea (Rossi) , C. dichroa (Dahlbohm) and C. trimaculata (Förster) (Chrysididae) ( Ferton 1905; Berland & Bernard 1938; Bellmann 1981; Kunz 1989).

Male behaviour. The males check snail shells for hatching or nest-seeking females (Müller et al., 1997).

Osmia (Allosmia) soror Pérez, 1896

Osmia cognata Pérez, 1895: 12 . Nomen praeoccupatum (not Osmia cognata Cresson ).

Type material: ♀ (♀), ( Algeria), Muséum National d’Histoire Naturelle Paris .

Osmia soror Pérez, 1896: 1 . Nomen novum with same type specimen for preoccupied Osmia cognata Pérez, 1895 (not Osmia cognata Cresson ).

Literature records. ALGERIA: Tlemcen (Ghazaouet) ( Ferton 1920, as O. rufohirta ). MOROCCO: CasablancaSettat, Fès-Meknès, Tanger-Tetouan-Al Hoceima ( Lhomme et al. 2020, as O. rufohirta ).

New records. ALGERIA: Tissemsilt: Tissemsilt, Cémitiaire , 35°35ʹ39ʺN / 1°48ʹ00ʺE, 12.5.2019, 1♀ (leg. Dermane) . MOROCCO: Fès-Meknès: Mulay Idriss , 26.3.1923, 1♂ (leg. A. von Schulthess) ; Meknès , 33°50.9ʹN / 5°28.9ʹW, 20.3.1992, 2♀, 1♂ (leg. H.- J. Flügel) ; Aghbalou Akourar , 33.8644°N / 4.6714°W, 626 m, 1.4.2019, 1♂ (leg. L. Hamroud) ; Ain Leuh , 33.3224ºN, 5.3359ºW, 1514 m, 12.3.2020, 1♂ (leg. Y. Ben Charki), 3.4.2020, 1♀ (leg. Y. Ben Charki) ; Rabat-Salé-Kénitra: Ouled Ben Hammadi , 34.246605°N / 5.854725°W, 42 m, 24.2.2021, 1♀ (leg. Y. Bencharki) ; Tanger-Tetouan-Al Hoceima: Larache , 70 km S Tanger, 31.3.1996, 1♀ (leg. O. Niehuis) . TUNISIA: Gabès: 10 km SE Matmata , 24.4.2012, 1♀ (leg. C. Praz) ; Kairouan: 48 km S Kairouan , 9.4.1994, 1♀ (leg. M. Schwarz) ; Kasserine: 20 km NW Kasserine , 4.4.2001, 2♀ (leg. J. Halada) ; Nabeul: Grombalia , 18.3.1996, 3♂ (leg. K. Denes) ; Sfax: 30 km SW Sfax , 10.4.1994, 1♀ (leg. J. Gusenleitner) ; Sidi Bouzid: Sidi Bouzid, 12.4.1999, 3♀ (leg. K. Denes) ; Sousse: M'saken, 21.4.1998, 2♀ (leg. K. Denes) ; Zaghouan: Zaghouan, 18.4.1998, 1♀ (leg. K. Denes) .

Distribution. Maghreb ( Morocco, Algeria, Tunisia).

Pollen hosts. Polylectic (at least 5 plant families): Fabaceae ( Hedysareae ; n = 1 pollen load with Fabaceae pollen), Asteraceae (Carduoideae; n = 1), Brassicaceae (n = 1), Caryophyllaceae (n = 1) and Convolvulaceae (n = 1) (based on 1 pollen load from Morocco). Flower record: Thymus serpyllum (label record).

Nesting biology. Osmia soror nests in empty snail shells ( Ferton 1920, as O. rufohirta ; but see species account of O. rutila ).

Note. Osmia soror was treated as the North African subspecies of O. rufohirta Latreille, 1811 by Zanden (1988b), Ungricht et al. (2008) and Scheuchl & Willner (2016). However, as O. soror differs from O. rufohirta by several morphological characters including the male genitalia (see identification key) and as its distribution appears to overlap with that of O. rufohirta in northern Morocco as suggested by a single record of a typical male of O. rufohirta from the Middle Atlas (Michlifene, 1900 m a.s.l.), it is considered here to represent a species of its own.

Osmia sybarita species group

The representatives of the O. sybarita species group differ from those of the O. rufohirta species group in both sexes by the distinctly narrower head and in the male sex by the predominantly flat and bifid tergum 7 as well as the lack of a projection on sternum 2. The two groups probably differ also in several characteristics of their nesting biology (see above).

Osmia (Allosmia) lhotelleriei Pérez, 1887

Osmia Lhotelleriei Pérez, 1887: 178 .

Type material: Lectotype ♀, by designation of Tkalců (1975), “ Égypte ” ( Egypt), Muséum National d’Histoire Naturelle Paris.

Osmia fossoria Pérez, 1890: 201 .

Type material: Lectotype ♀, by designation of Tkalců (1974), “ Alger ” ( Algeria), Muséum National d’Histoire Naturelle Paris. New synonymy based on original description, Tkalců (1974) and a large quantity of O. ( Allosmia ) material from North Africa.

Osmia duckei Friese, 1899: 27 . Type material: Syntypes ♂♂, “ Alger ” ( Algeria), Museum für Naturkunde Berlin. Synonymy with Osmia fossoria Pérez in Friese (1911).

Literature records. ALGERIA: Algiers (Algiers, Zéralda), Tipasa (Tipasa)( Ferton1890; Tkalců1975). MOROCCO: Drâa-Tafilalet, Fès-Meknès ( Lhomme et al. 2020).

New records. ALGERIA: Mila: Redjas , 36º27ʹN / 6º16E, 30.4.2013, 2♀ (leg. Abderrezak) ; Oum El Bouaghi: Ain M'lila, 36°20ʹN / 6°35ʹE, 771 m, 1.5.2011, 1♀ (leg. G. Zineb) . ISRAEL AND PALESTINE: Central District: 1.2 km NW Bene Ziyyon, 32°13ʹ27ʺN / 34°51ʹ22ʺE, 30 m, 16.2.– 14.4.2010, 12♀, 3♂ (leg. A. Dorchin); Haifa District: 1.3 km N Tiv'on , 32°44ʹ14ʺN / 35°07ʹ55ʺE, 163 m, 23.3.2012, 3♂ (leg. A. Dorchin); Jerusalem District : Judean Foothills, Neve Shalom, 12.2.2010, 1♂ (leg. G. Pisanty); Northern District : Ubeidiya, 7.3.2015, 1♀ (leg. T. Jumah); Southern District : Sede Boqer, 16.3.2015, 1♀ (leg. I. Zonstein); Tel Aviv District : Tel Aviv University, Botanical Garden, 32°06ʹ50ʺN / 34°48ʹ31ʺE, 2.– 5.4.2012, 4♀ (leg. J. S. Ascher); West Bank : 3.7 km W Pezael , 32°03ʹ05ʺN / 35°23ʹ52ʺE, - 30 m, 4.4.2017, 1♀ (leg. A. Dorchin) . JORDAN: Amman: Wadi Shu'ayb , 20 km W Amman, 22.4.2007, 1♀ (leg. C. Praz, C. Sedivy, A. Müller); Dscharasch: 10 km N Jerash, 20.4.2002, 12♀ (leg. M. Snizek) ; Karak: Wadi al Hasa S Al-Karak, 20.4.2007, 1♀ (leg. C. Praz, C. Sedivy, A. Müller) ; Ma'an: AlShawbak , 18.4.2007, 9♀ (leg. C. Praz, C. Sedivy, A. Müller) ; Madaba: Wadi Mujib, Kings Highway , 19.4.2007, 5♀ (leg. C. Praz, C. Sedivy, A. Müller) . MOROCCO: Fès-Meknès: Ifrane , 33°31.14ʹN / 05°05.65ʹW, 1680 m, 14.5.2002, 3♀ (leg. H.- J. Flügel) . SYRIA: Al-Quneitra: Golan Heights , Yehudiya Reservoir, 28.6.2009, 1♀ (leg. A. Dorchin) ; Aleppo: Marbij , 9.5.1996, 1♀ (leg. M. Halada) . TUNISIA: Gabès: 30 km N Gabès , 10.4.1994, 1♀ (leg. M. Schwarz) ; Kef: 10 km SW Le Kef , 15.4.2001, 4♀ (leg. J. Halada) ; Medenine: Djerba, SE Houmt Souk , 33.52ºN / 10.55ºE, 19.3.2001, 22♀, 3♂ (leg. C. Saure, C. Schmid-Egger) ; Nabeul: Tazerka , 36°30ʹN / 10°50ʹE, 19.4.2004, 1♀ (leg. F. Amiet) ; Siliana: Makthar , 16.– 17.04.1998, 4♀ (leg. K. Denes) ; Tunis: Carthage , 8.4.2000, 1♀ (leg. P. Hartmann) .

Distribution. From the Maghreb ( Morocco, Algeria, Tunisia) over Libya and Egypt to the Levant ( Israel and Palestine, Jordan, Syria).

Pollen hosts. Polylectic (at least 10 plant families): Fabaceae ( Genisteae , Hedysareae , Loteae , Trifolieae ; n = 38 pollen loads with Fabaceae pollen), Asteraceae (Asteroideae, Carduoideae, Cichorioideae; n = 18), Boraginaceae (e.g. Echium ; n = 11), Brassicaceae (n = 9), Monocots (n = 7), Caryophyllaceae (n = 5), Resedaceae (n = 5); Lamiaceae ( Lamioideae , Nepetoideae ; n = 4), Cistaceae (n = 2) and Apiaceae (n = 1) (based on 44 pollen loads from 20 different localities in Israel and Palestine, Jordan, Morocco and Tunisia). Flower records: Alkanna tinctoria , Echium judaeum , Anthemis melampodina , Chrysanthemum coronarium , Senecio verna , Centaurea cyanoides , Silybum marianum , Crepis aculeata , Leontodon tuberosus , Arenaria leptoclados , Asphodelus ramosus , Campanula spec. , Diplotaxis erucoides , Geranium robertianum , Ononis serrata , Trifolium philistaeum (label records).

Nesting biology. Osmia lhotelleriei nests in empty snail shells of small size, e.g. of Theba pisana ( Ferton 1890; O’Toole & Raw 1991; N. Vereecken personal communication). In contrast to O. bischoffi and O. rufohirta , the females do not cover the shell surface with patches of leaf pulp. The nests contain a single brood cell, which is closed with a one-layered partition of leaf pulp a few millimetres behind the shell opening; this partition forms the base of the 3–4 mm thick nest plug, which is built from broken pieces of gastropod shells embedded into a matrix of pulp consisting of chewed green leaves, occasionally also of chewed petals. After the nest has been sealed, it is rolled to a previously selected sandy place, where it is moved into a slanting burrow of 6–7 cm length and 1–1.5 cm depth excavated in loose sand, before it is completely covered with sand. Brood parasite: Chrysura osiris (Buysson) ( Buysson 1887, 1908).

Note. Tkalců (1974) treated O. fossoria Pérez, 1890 as subspecies of O. sybarita Smith, 1853 , which is erroneous as the latter species does not occur in northern Africa.

Osmia (Allosmia) melanura Morawitz, 1871

Osmia melanura Morawitz, 1871: 203 .

Type material: ♀ (♀), “ Calabria ” ( Italy), type depository unknown.

Osmia decorata Morawitz, 1886: 71 .

Type material: ♂ (♂), “Talysch-Karabach-Jurdi” ( Azerbaijan), Zoological Institute of Russian Academy of Sciences St. Petersburg. New synonymy based on original description and topotypical specimens from Azerbaijan.

Literature records. ITALY: Calabria, Puglia (Brindisi), Sicily (Siracusa) ( Ducke 1900). NORTH MACEDONIA: Southwestern (Ohrid) ( Zanden 1984a). TURKEY: Antalya, Erzurum ( Özbek & Zanden 1992; Özbek 2013). UKRAINE: Odessa: Odessa (Warncke, unpublished distribution maps).

New records. ARMENIA: Ararat: Chor Virap , 1000 m, 12.6.2013, 1♀ (leg. W. Schläfle) ; Sjunik: E Meghri, 12.6.2003, 1♀ (leg. Mücka) . AZERBAIJAN: Nakhichevan: Goynuk , 39°18ʹN / 45°40ʹE, 1680 m, 12.6.2019, 1♀ (leg. M. Proshchalykin) . BULGARIA: Blagoevgrad: SW Kresna, 41°42ʹN / 23°11ʹE, 150 m, 24.6.2008, 1♀ (leg. M. Halada) ; Burgas: Nessebar , 28.6.1982, 1♂ (leg. M. Kocourek) ; Dobrich: Albena , 18.– 29.6.1986, 1♀ (leg. J. Halada) . GREECE: Eastern Macedonia and Thrace: Thasos, Potos , 40.613ºN / 24.6195ºE, 11.4.2012, 1♀ (leg. M. de Courcy) . ITALY: Puglia: Gargano, Varano , 0 m, 8.5.– 16.05.1990, 2♀ (leg. A. Müller) ; Gargano , M. S. Angelo, S. Barnabeo, 800 m, 4.7.1997, 1♀ (leg. A. Müller) ; Sicily: Piazza Armerina , 35km N Gela, 27.– 29.5.2002, 14♀ (leg. J. Halada) . TURKEY: Ankara: Hacettepe University, Beytepe Campus , 17.5.2005, 1♂ (leg. E. Scheuchl) ; Antalya: 10km S Kizilcadag, 26.5.2009, 1♂ (leg. J. Ascher , J. Rozen, H. Özbek) ; Bolu: 17 km S Seben , 17.6.1998, 1♀ (leg. J. Halada) ; Erzurum: Kuslu , 10 km E Ispir, 12.6.2010, 1♀ (leg. J. Halada) ; Konya: Beysehir, around Sarkikaraagaç , 37°45ʹ885"N / 31°40ʹ397"E, 1140 m, 25.5.2005, 1♀ (leg. E. Scheuchl) ; Kütahya: Porsuk Baraji , 30 km N Kutahya, 15.6.1997, 1♀ (leg. M. Halada) ; Van: 10 km N Muradiye , 27.6.1997, 1♀ (leg. M. Halada) ; Sivas: Zara , 40 km E Sivas, 3.6.2002, 5♀ (leg. W.- H. Liebig) ; Yozgat: Bogazliyan , 39°14.67ʹN / 35°12.376ʹE, 7.6.2002, 1♀ (leg. W.- H. Liebig) .

Distribution. From southern and eastern Italy including Sicily over North Macedonia, northern Greece and Bulgaria to southern Ukraine, Turkey and the Caucasus ( Armenia, Azerbaijan). Rasmont et al. (1995) list O. melanura among the bee species occurring in France, which is most probably erroneous.

Pollen hosts. Polylectic (at least 7 plant families): Fabaceae ( Galegeae , Hedysareae , Loteae , Trifolieae ; n = 10 pollen loads with Fabaceae pollen), Boraginaceae (e.g. Echium ; n = 5), Asteraceae (Cichorioideae; n = 1), Cistaceae (n = 1), Lamiaceae ( Lamioideae ; n = 1), Brassicaceae and Caryophyllaceae (based on 11 pollen loads from 9 different localities in Azerbaijan, Bulgaria, Italy and Turkey and on Müller 1992). The importance of Fabaceae as pollen hosts is indicated by the finding that the percentage of Fabaceae pollen in four brood cells ranged from 89.7– 96.4 % ( Müller 1992). Flower records: Calystegia soldanella , Lotus creticus , Medicago marina ( Müller 1992) ; Medicago sativa ( Özbek 2013) .

Nesting biology. Osmia melanura nests in empty snail shells of small size, e.g. of Theba pisana ( Fig. 1 ; Müller 1992). In contrast to O. bischoffi and O. rufohirta , the females do not cover the shell surface with patches of leaf pulp ( Fig. 1 ), but - as in O. rutila - they attach leaf pulp to the inner shell surface few millimeters behind the shell opening immediately after they have selected a suitable nest shell; these leaf pulp markings are later extended to the cell partition. The nests contain a single brood cell, which lacks a basal wall that seals the larval provisions against the rear end of the shell. After provisioning and egg deposition, the cell is closed with a one-layered partition of leaf pulp 2–3 mm behind the shell opening in a considerable distance (0.6–0.8 shell whorls) from the larval provisions ( Fig. 4 ); this partition forms the base of the 3–4.5 mm thick nest plug, which consists of broken pieces of gastropod shells embedded into a matrix of leaf pulp ( Fig. 4 ). After the nest has been sealed, it is rolled over a distance of up to 50 cm to a previously selected sandy place, where it is moved into a 2 cm deep hole excavated among plant roots, before it is completely covered with sand. O. melanura is not strictly confined to sandy areas but occasionally also occurs in stony habitats, where the ground is too hard to dig; here, the females might possibly transport their nests to a protected place under vegetation or stones.

Osmia (Allosmia) rufotibialis Friese, 1920

Osmia rufotibialis Friese, 1920: 50 . Type material: ♀ (♀), “Ain Dilb” ( Israel and Palestine), Museum für Naturkunde Berlin.

Literature records. ISRAEL AND PALESTINE: Jerusalem district (Jerusalem) ( Mavromoustakis 1939b).

New records. ISRAEL AND PALESTINE: Central District: Karmei Yosef, 31.863594ºN / 43.930752ºE, 22.3.2018, 3♀ (leg. T. Roth); Haifa District : 1.3 km N Tiv'on , 32°44ʹ14ʺN / 35°07ʹ55ʺE, 163 m, 30.3.2012, 2♀, 1♂ (leg. A. Dorchin); Jerusalem District : Judean Foothills, Ya'ar Adulam, 25.3.2015, 1♀ (leg. T. Chaprazaro); Northern District : Hermon, 15 km E Qiryat Shemona , 33°15ʹN / 35°44ʹE, 16.5.1996, 1♀ (leg. C. Schmid-Egger); Southern District : Fura NR, 6.3 km E Ruhama , 31°29ʹ47ʺN / 34°46ʹ33ʺE, 196 m, 15.4.2012, 1♀ (leg. A. Dorchin); West Bank : 3.7 km W Pezael , 32°03ʹ05ʺN / 35°23ʹ52ʺE, - 30 m, 4.4.2017, 1♀ (leg. A. Dorchin) . JORDAN: Ajloun: Ajloun, 32°23ʹN / 35°46ʹE, 28.4.2012, 1♀ (leg. M. Kafka); Dscharasch: 10 km N Jerash, 23.4.2007, 3♀ (leg. C. Praz, C. Sedivy, A. Müller); Irbid: Pella, 32°26ʹN / 35°36ʹE, - 80 m, 29.4.2006, 1♀ (leg. K. Denes); Jordan Valley : Mubalath , 27.4.1996, 7♀ (leg. M. Halada); Karak: 30 km S Al Karak, 30°59ʹN / 35°44ʹE, 27.4.2006, 1♀ (leg. Kantner) . SYRIA: Al-Quneitra: Golan Heights, Nahal Mezar , 32.755°N / 35.69°E, 1.5.2015, 1♀ (leg. G. Pisanty) .

Distribution. Israel and Palestine, Jordan and southernmost Syria.

Pollen hosts. Polylectic (at least 5 plant families): Fabaceae ( Genisteae , Hedysareae , Loteae , Psoraleeae , Trifolieae ; n = 14 pollen loads with Fabaceae pollen), Asteraceae (Asteroideae, Carduoideae, Cichorioideae; n = 12), Boraginaceae ( Echium ; n = 7); Brassicaceae (n = 3) and Lamiaceae ( Lamioideae ; n = 3) (based on 19 pollen loads from 14 different localities in Israel and Palestine and Jordan). Flower records: Centaurea , Lepidium ( Mavromoustakis 1939b) ; Trifolium purpureum , Crepis aculea , Picris altissima , Senecio verna , Diplotaxis erucoides , Nepeta curviflora , Stachys neurocalycina , Caylusea hexagyna , Scabiosa prolifera (label records).

Nesting biology. Unknown.

Osmia (Allosmia) rutila Erichson, 1835

Osmia rutila Erichson, 1835: 107 . Type material: Holotype ♀, “ Andalusien ” ( Spain), Museum für Naturkunde Berlin.

Osmia baetica Spinola, 1843: 142 . Type material: Holotype ♀, ( Spain), type depository unknown. Synonymy in Dalla Torre (1896), but see Zanden (1987).

Osmia xanthognatha Pérez, 1895: 12 . Type material: Lectotype ♀, by designation of Zanden (1988a), “ Tanger ” ( Morocco), Muséum National d’Histoire Naturelle Paris. Synonymy in Zanden (1987).

Literature records. MOROCCO: Casablanca-Settat (Ain Diab), Souss-Massa (Agadir, Massa, Tiznit), TangerTetouan-Al Hoceima (Tanger) ( Zanden 1987, 1988a; Haeseler 1997; Lhomme et al., 2020). SPAIN: Barcelona (Pto Real, Canet de Mar), Cádiz (Barbate de Franco, Puerto Real, Puerto St. Maria, Tarifa), Málaga (Costa del Sol, Marbella) ( Mavromoustakis 1947; Zanden 1984b, 1987, 1988a; Haeseler 1997).

New records. MOROCCO: Casablanca-Settat: El Jadida, 5.1964, 1♀ (leg. W. Schläfle); Souss-Massa: Cap Rhir-Tamri, 60 km N Agadir, 25 m, 14.4.2009, 5♀ (leg. A. Müller); Corniche Aglou, 65 km NE Sidi Ifni , 29°48.499ʹN / 09°49.650ʹW, 40 m, 16.4.2019, 1♀ (leg. A. Müller); Tanger-Tetouan-Al Hoceima: Frideq, sand dunes, 35°44.08ʹN / 05°20.49ʹW, 14 m, 3.5.2002, 4♀ (leg. H.- J. Flügel) . SPAIN: Cádiz : Andalucia, Punta Tarifa, 30 km SW Algeciras , 5.5.2003, 32♀, 18♂ (leg. Z. Pedr); Málaga: Estepona, 11.4.1985, 1♀, 1♂ (leg. H. Wolf); Murcia: 25 km SW Cartagena, 12.5.2003, 2♀ (leg. J. Halada) .

Distribution. From southern Morocco along the coast to southern and southeastern Spain; one isolated record is from northeastern Spain near Barcelona. Ferton (1920) mentions several females of O. rufohirta from Nemours (= Ghazaouet) in northwesternmost Algeria, which were characterized by a red metasoma; these records possibly refer to O. rutila and suggest that this species might also occur at the Algerian Mediterranean coast. Osmia rutila seems to exhibit a clear or even exclusive preference for sand dunes or sandy strand walls in coastal areas.

Pollen hosts. Polylectic (at least 5 plant families): Fabaceae ( Loteae , Trifolieae ; n = 18 pollen loads with Fabaceae pollen), Boraginaceae ( Echium ; n = 2), Brassicaceae (n = 2), Asteraceae and Oxalidaceae (based on 20 pollen loads from 10 different localities in Morocco and Spain and on Haeseler 1997). The importance of Fabaceae as pollen hosts is indicated by the finding that brood cells from several nests almost exclusively contained Fabaceae pollen ( Haeseler 1997). Flower records: Lotus creticus , Lotus edulis , Medicago marina , Oxalis pes-caprae ( Haeseler 1997) .

Nesting biology. Osmia rutila nests in empty snail shells of small size, e.g. of Helicella ( Haeseler 1997) . In contrast to O. bischoffi and O. rufohirta , the females do not cover the shell surface with patches of leaf pulp, but - as in O. melanura - they attach leaf pulp to the inner shell surface few millimeters behind the shell opening immediately after they have selected a suitable nest shell; these leaf pulp markings, which are later extended to the cell partition, allow the females to recognize their own nest and signal other females that the shell is already occupied ( Haeseler 1997). The nests contain a single brood cell, which lacks a basal wall that seals the larval provisions against the rear end of the shell. After provisioning, which requires up to 34 foraging bouts, and egg deposition, the cell is closed with a one-layered partition of leaf pulp a few millimetres behind the shell opening in a considerable distance (about 0.6 shell whorls) from the larval provisions; this partition forms the base of the thick nest plug, which consists of up to 28 broken pieces of gastropod and mussel shells, sometimes also small stones, embedded in a matrix of leaf pulp; the construction of the nest plug including the basal partition requires up to 150 flights with leaf pulp or broken shell pieces. After the nest has been sealed, it is rolled over a distance of up to 3 m to a previously selected sandy place in the vicinity of plants, where it is moved into a 3–3.5 cm deep hole and turned in a vertical position with the nest plug directing towards the entrance of the excavated hole, before it is completely covered with sand. Even under good weather conditions, the nesting cycle from nest selection to nest burying lasts two days.

Male behaviour. The males search for females in small areas of 10–15 m 2 around nesting sites and flower patches, where they patrol along flight routes regularly interrupted by short resting periods on the ground or on low vegetation ( Haeseler 1997).

Osmia (Allosmia) sybarita Smith, 1853

Osmia sybarita Smith, 1853: 140 . Type material: Holotype ♀, “ Albania ” ( Albania), Natural History Museum London.

Osmia ruficollis Sichel in Dours, 1873: 297. Type material: Syntypes ♀♀, ♂♂ “Iles de l’Archipel grec” ( Greece), Muséum National d’Histoire Naturelle Paris. Synonymy in Zanden (1988b).

Literature records. CYPRUS: Famagusta, Limassol, Nicosia, Paphos ( Mavromoustakis 1939 a, 1948b). TURKEY: Ankara, Antalya, Elazig, Konya, Mersin ( Zanden 1980; Özbek & Zanden 1996; Özbek 2013; Varnava et al. 2020).

New records. BULGARIA: Burgas: Sozopol , 8.– 22.6.1985, 1♀ (leg. K. Polacek) . CYPRUS: North Cyprus: Besparmak, Gecitköy W Lapta , 300 m, 30.4.2011, 8♀ (leg. C. Sedivy, A. Müller) ; South Cyprus: Agia Napa , 35 km E Larnaka, 34°59ʹ14ʺN / 33°57ʹ45ʺE, 12.4.2012, 1♀ (leg. G. Reder) . GREECE: Aegean Islands: Chios, Emborios , 38.2048ºN / 26.0218ºE, 4.5.2012, 1♀ (leg. M. Taylor) ; Ios, Kambos , 36.7518ºN / 25.2921ºE, 14.4.2013, 1♀ (leg. T. Petanidou) ; Karpathos, Avlona , 35.7689ºN / 27.1849ºE, 28.3.2012, 4♀, 1♂ (leg. T. Petanidou) ; Kos, Psallidi , 36.8785ºN / 27.3352ºE, 7.4.2012, 1♀ (leg. S. Papakonstantinou, A. Kyriazis) ; Limnos , Ag. Sotira, 39.9903ºN 25.3917ºE, 6.4.2012, 1♂, ( J. Devalez) ; Paros, Kamari , 37.0132ºN / 25.1402ºE, 13.5.2013, 1♀ (leg. I. Vavitsas) ; Rhodes, Archangelos , 9.5.2005, 2♀ (leg. A. Müller) ; Thasos, Astris , 40.5871ºN / 24.6528ºE, 11.4.2012, 4♀, 2♂ (leg. M. de Courcy) ; Attica: Aegina, Ag. Asomaton , 37.7502ºN / 23.457ºE, 28.3.2013, 3♀ (leg. S. Margaroni) ; Brauron , 7.4.2000, 3♀ (leg. W. Arens) ; Central Greece: S Delphi , 9.5.2005, 3♀ (leg. J. Halada) ; Central Macedonia: Pieria, Leptokarya , 11.6.1991, 1♀ (leg. Z. Pedr) ; Crete: Pitsidia , 35.01ºN / 24.46ºE, 20 m, 27.5.2009, 1♀ (leg. U. Frommer) . Ionian Islands: Korfu, Linia , 10.5.1984, 2♀ (leg. L. Norén) ; Peloponnese: 50 km SW Patra , 22.4.2005, 30♀, 1♂ (leg. J. Halada) ; Thessalia: Mt. Ossa, Kokino Nero , 40 km NE Larisa, 13.5.2005, 1♀ (leg. M. Kadlecova) ; Western Greece: Kaiaphas lake, Zacharo , 30.4.1995, 1♀ (leg. W. Arens) . ISRAEL AND PALESTINE: Central District : 0.75 km N Yaqum, 32°15ʹ20ʺN / 34°50ʹ33ʺE, 12 m, 11.2.2010, 1♀, 4♂ (leg. A. Dorchin) ; Haifa District: Mt. Carmel, Bet Oren , 13.3.1997, 1♀ (leg. J. Rozen, M. S. Engel, A. Hefetz) ; Jerusalem District: Har Gillo , 12.5.1997, 1♀ (leg. J. Rozen) ; Southern District: Negev, Wadi Loz , 7.5.1997, 1♀ (leg. J. Rozen) . JORDAN: Dscharasch : Jerash, 1.5.1996, 1♀ (leg. M. Halada) ; Irbid: North Shuna , 30.4.1996, 7♀ (leg. M. Halada) . LEBANON: Mount Lebanon: Batloun , 33.7000007629395°N / 35.6500015258789°E, 25.4.1981, 1♂ (leg. Othob) . MONTENEGRO: Ulcinj , 41°52.4ʹN / 19°21.1ʹE, 27.5.2005, 1♀ (leg. Z. Pedr) . SYRIA: Idlib: Jisr ash Shughur , 10.5.1996, 3♀ (leg. M. Halada) ; Latakia: 20 km NE Latakia , 25.5.1996, 2♀ (leg. M. Halada) ; Tartus: Tartus, 25.5.1996, 1♀ (leg. M. Halada) . TURKEY: Ankara: Karakeçili , 39°35.001ʹN / 33°24.672ʹE, 760 m, 6.4.2005, 4♀ (leg. E. Scheuchl) ; Antalya: 60 km SEE Antalya, 10 km NWW Manavgat, 36°49ʹN / 31°21ʹE, 10.4.1998, 1♀ (leg. Geller-Grimm) ; Batman: 10 km N Gercüs , 5.6.1998, 2♀ (leg. M. Halada) ; Elazig: Keban , 38°45ʹ4ʺN / 38°46ʹ52ʺE, 900 m, 16.5.2002, 1♀ (leg. H. Özbek) ; Mersin: Cornelek , 40 km E Mut, 29.5.1996, 3♀ (leg. M. Halada) ; Mugla: Yatagan , 37°12.541ʹN / 28°20.108ʹE, 625 m, 26.5.2005, 1♀ (leg. E. Scheuchl) .

Distribution. From southeastern Europe ( Montenegro, Albania, Bulgaria, Greece including Crete, Ionian and Aegean Islands) over Turkey and Cyprus to the Levant ( Syria, Lebanon, Jordan, Israel).

Pollen hosts. Polylectic (at least 7 plant families): Fabaceae (Fabeae, Genisteae , Hedysareae , Loteae , Psoraleeae , Trifolieae ; n = 24 pollen loads with Fabaceae pollen), Asteraceae (Asteroideae, Carduoideae, Cichorioideae; n = 11), Boraginaceae (e.g. Echium ; n = 8); Brassicaceae (n = 8), Resedaceae (n = 5), Lamiaceae ( Nepetoideae ; n = 1) and Monocots (n = 1) (based on 32 pollen loads from 27 different localities in Cyprus, Greece, Israel and Palestine, Jordan, Syria and Turkey). Flower records: Hymenocarpus , Onobrychis , Vicia ( Mavromoustakis 1939 a, 1948b); Alkanna tinctoria , Hymenocarpus circinnatus , Trifolium palaestinum , Crepis commutata , Echium angustifolium (label records).

Nesting biology. Osmia sybarita nests in empty snail shells of small size, e.g. of Helicella, Pseudoxerophila, Trochoidea or Xeromunda ( Fig. 3 ; Mavromoustakis 1939 a, 1948b; O’Toole & Raw 1991; Haeseler 1997; Vereecken & Le Goff, 2012). In contrast to O. bischoffi and O. rufohirta , the females do not cover the shell surface with patches of leaf pulp ( Fig. 3 ). The nests contain a single brood cell, which lacks a basal wall that seals the larval provisions against the rear end of the shell. After provisioning and egg deposition, the cell is closed with a one-layered partition of leaf pulp a few millimetres behind the shell opening; this partition forms the base of the thick nest plug, which consists of broken pieces of gastropod shells embedded into a matrix of leaf pulp ( Fig. 3 ). After the nest has been sealed, the shell is rolled over a distance of up to 40 cm to a previously selected place, where it is shallowly buried into the sandy ground ( Fig. 3 ), or - if the ground is too hard to dig - hidden under low vegetation (G. Le Goff personal communication). Brood parasites: Chrysura dichroa (Dahlbohm) ( Mavromoustakis 1939a) .

Key to species

The male of Osmia imitatrix is unknown.

Females

1 Head longer, more than 0.9x as long as wide................................................................ 2

- Head shorter, less than 0.9x as long as wide................................................................ 7

2 Terga and sterna with red markings....................................................................... 3

- Terga and sterna without red markings..................................................................... 4

3 Terga 1–3(4) completely or partly orange-red, terga (4)5–6 black. Tibia and tarsus of hind leg usually completely or largely orange-red, tibia and basitarsus of middle and fore leg completely or predominantly black, femora of all legs black. Pilosity of vertex, mesosoma and terga whitish to yellowish. Mandible black. (Body length 7–10 mm; southern and eastern Europe, Turkey, Caucasus)........................................................................ Osmia melanura

- Terga with varying extent of red colour, ranging from completely orange-red to considerably darkened, but at least parts of tergal discs 1–3 reddish. Tibia and tarsus of all legs completely or predominantly orange-red, femora of all legs with orange-red markings of varying extent. Pilosity of vertex, mesosoma and terga orange-red. Mandible more or less reddish except for black teeth. (Body length 8–10 mm; Maghreb, Spain)................................................... Osmia rutila

4 Tibia and basitarsus of hind leg predominantly orange-red. (Body length 7–9 mm; Levant)............. Osmia rufotibialis

- Tibia and basitarsus of hind leg black..................................................................... 5

5 Pilosity of head and mesosoma as well as metasomal scopa whitish to yellowish. Abscissa between first submarginal crossvein and first recurrent vein in second submarginal cell of fore wing about 2x as long as width of recurrent vein. First segment of labial palpus about 0.8x as long as second segment. (Body length 7.5mm; Turkmenistan)................ Osmia imitatrix

- Pilosity of head and mesosoma as well as metasomal scopa orange-red to yellowish-red. Abscissa between first submarginal crossvein and first recurrent vein in second submarginal cell of fore wing at least 3x as long as width of recurrent vein. First segment of labial palpus 0.6–0.7x as long as second segment................................................... 6

6 Lateral hair spots of tergum 1 orange-red and of same colour as mesosomal pilosity. Apical hair bands of terga 2–4 yellowishred, medially interrupted and usually completely rubbed off on terga (2) 3–4 in older specimens. Tergum 5 without whitish apical hair band. Punctation of tergal discs rather fine: punctures on tergal discs 2–4 at most twice to three times as large as punctures on tergal marginal zones. Short pilosity of tergal discs 3–5 often dark brown to black. (Body length 7–9 mm; northern Africa, Levant)......................................................................... Osmia lhotelleriei

- Lateral hair spots of tergum 1 whitish to yellowish, usually differing from orange-red colour of mesosomal pilosity. Apical hair bands of terga 2–5 whitish and not interrupted on terga (3) 4–5 in fresh specimens. Punctation of tergal discs rather coarse: punctures on tergal discs 2–4 usually several times larger than punctures on tergal marginal zones. Short pilosity of tergal discs 3–5 usually whitish to yellowish (Body length 7–9.5 mm; southeastern Europe, Turkey, Cyprus, Levant).... Osmia sybarita

7 Apical margin of clypeus medially emarginate with slightly projecting lateral edges. Pilosity of head, mesosoma and terga whitish. (Body length 7.5–10 mm; southeastern Europe, Turkey).................................... Osmia bischoffi - Apical margin of clypeus medially straight. Pilosity of head, mesosoma and terga predominantly orange-red to yellowish-red.................................................................................................... 8

8 Smaller, body length at most 7 mm. Antennal segment 3 less than 1.25x as long as wide, slightly shorter than total length of segments 4–5. First segment of labial palpus relatively longer, at least 0.6x as long as second segment. (Body length 6–7 mm; southeastern Europe, Turkey)................................................................... Osmia nuda

- Larger, body length usually surpassing 7 mm. Antennal segment 3 more than 1.25x as long as wide, slightly longer than total length of segments 4–5. First segment of labial palpus relatively shorter, about 0.5x as long as second segment........... 9

9 Tergal hair bands longer, hairs at apical margin of terga 2–4 surpassing tergal margin by more than half of their length. (Body length 7–9 mm; Maghreb).................................................................... Osmia soror

- Tergal hair bands shorter, hairs at apical margin of terga 2–4 surpassing tergal margin by less than half of their length..... 10

10 Outside southern Levant. (Body length 7–9.5 mm; Moroccan Middle Atlas, Europe, Turkey, Caucasus, Iran, Levant)............................................................................................. Osmia rufohirta

- Southern Levant..................................................................................... 11

11 Marginal zone of terga 1–5 predominantly black to narrowly dark reddish-brown ( Fig. 6 ). (Body length 7–9 mm; Levant)........................................................................................... Osmia gemina

- Marginal zone of terga 1–4(5) usually broadly yellowish. (Body length 7–9.5 mm; Europe, Turkey, Caucasus, Iran, Levant)........................................................................................ Osmia rufohirta

Males

1 Tergum 7 predominantly flat and apically bifid. Sternum 2 flat and without projection............................... 2

- Tergum 7 laterally distinctly curved downwards and apically rounded to truncate. Sternum 2 with preapical swelling, which is medially extended into a projection of pyramidal or triangular shape............................................. 7

2 Apical margin of clypeus medially emarginate with two projecting flat teeth of triangular shape. Sterna 1–2 with dense and very long yellowish pilosity, which is distinctly longer than tarsal segment 2 of hind leg. Apical margin of sternum 2 laterally slightly concave and medially projecting. Apical margin of sternum 5 with long ciliae, which are almost as long as tarsal segment 4 of hind leg. (Body length 11 mm; southeastern Europe, Turkey)....................................... Osmia bischoffi

- Apical margin of clypeus medially straight to slightly emarginate without tooth-like projections. Sterna 1–2 with sparse and moderately long whitish pilosity, which is barely longer than tarsal segment 2 of hind leg. Apical margin of sternum 2 laterally straight to slightly convex and medially not distinctly protruding. Apical margin of sternum 5 with short ciliae, which are distinctly shorter than tarsal segment 4 of hind leg........................................................... 3

3 Terga and sterna with red markings of varying extent, but at least tergal marginal zones and ventrolateral parts of terga 1–2 always reddish....................................................................................... 4

- Terga and sterna black except for ventrolateral parts of terga 1–2, which are occasionally partly reddish................. 5

4 Emargination at apical margin of sternum 3 at most one fifth as wide as entire sternal width. Tergal discs 1–4(5) partly or completely orange-red, terga 5–6 predominantly black except for reddish marginal zone. Tibia of hind leg black except for orange-red apicalmost part. (Body length 8–9 mm; southern and eastern Europe, Turkey, Caucasus)....... Osmia melanura

- Emargination at apical margin of sternum 3 at least one half as wide as entire sternal width. Terga with varying extent of red colour, ranging from completely orange-red to strongly darkened. Tibia of hind leg at least on inside completely orange-red. (Body length 8–10 mm; Maghreb, Spain)........................................................ Osmia rutila

5 Tarsal segments of hind leg completely orange-red. Emargination at apical margin of sternum 3 shallower, about one forth as deep as wide. (Body length 7.5–9 mm; Levant)................................................ Osmia rufotibialis

- Tarsal segments of hind leg with varying extent of reddish colour, but basitarsus always predominantly black. Emargination at apical margin of sternum 3 deeper, about one third to one half as deep as wide..................................... 6

6 Apical margin of sternum 6 laterally slightly concave and medially protruding ( Fig. 12 ). Gonoforceps in its apical half moderately bent inwards ( Fig. 14 ). Teeth of tergum 7 usually narrower, about 1.5x as long as basally wide or longer ( Figs 12, 14 ). (Body length 7.5–9 mm; northern Africa, Levant).......................................... Osmia lhotelleriei

- Apical margin of sternum 6 laterally straight to convex and evenly rounded ( Fig. 13 ). Gonoforceps in its apical half more strongly bent inwards ( Fig. 15 ). Teeth of tergum 7 usually wider, about 1.25x as long as basally wide or shorter ( Figs 13, 15 ). (Body length 7.5–10 mm; southeastern Europe, Turkey, Cyprus, Levant)............................. Osmia sybarita

7 Sternum 2 with small and acute tubercle of pyramidal shape. Lateral margins of tergum 7 evenly converging towards apex in dorsal view. (Body length 7.5–9 mm; southeastern Europe, Turkey).................................... Osmia nuda

- Sternum 2 with appressed and flat tooth of triangular shape, whose apex reaches sternal margin. Lateral margins of tergum 7 apically almost parallel-sided in dorsal view................................................................ 8

8 Marginal zone of sterna 4–5 covered with non-shaggy yellowish pilosity consisting of rather short and appressed hairs, which form a rather dense band ( Fig. 9 ). Apical half of gonoforceps distinctly broadened and apically strongly bent inwards, its inner margin concave ( Fig. 11 ). Roundish preapical impression of sternum 6 usually more or less dull, more densely punctate and distinctly haired ( Fig. 9 ). (Body length 8.5–12 mm; Moroccan Middle Atlas, Europe, Turkey, Caucasus, Iran, Levant)............................................................................................ Osmia rufohirta

- Marginal zone of sterna 4–5 covered with shaggy yellowish pilosity consisting of rather long and suberect hairs, which do not form a band ( Fig. 8 ). Apical half of gonoforceps weakly broadened and apically slightly bent inwards, its inner margin almost straight ( Fig. 10 ). Roundish preapical impression of sternum 6 more or less polished, more sparsely punctate and almost unhaired ( Fig. 8 )...................................................................................... 9

9 Pilosity along inner margin of apical half of gonoforceps in dorsal view distinctly less than half as long as pilosity along outer margin ( Fig. 10 ). Longest hairs on tergal discs 4–5 distinctly shorter than tarsal segment 2 of hind leg. Marginal zone of terga 1–5 often more or less dark, rarely predominantly reddish. (Body length 8–10 mm; Levant)............... Osmia gemina

- Pilosity along inner margin of apical half of gonoforceps in dorsal view about half as long as pilosity along outer margin. Longest hairs on tergal discs 4–5 almost as long as tarsal segment 2 of hind leg. Marginal zone of terga 1–5 usually predominantly reddish. (Body length 9.5–11 mm; Maghreb)..................................................... Osmia soror

Figures

FIGURES 1–5: Osmia (Allosmia). 1: Female of O. melanura at the entrance of her nest. 2: Female of O. rufohirta rolling her nest towards a protected place. 3: Female of O. sybarita rolling her nest into a self-excavated hole in sandy soil (photo N. Vereecken). 4: Opened nest of O. melanura with one brood cell and a nest plug at the shell opening composed of a one-layered partition of leaf pulp followed by broken pieces of gastropod shells embedded into a matrix of leaf pulp. 5: Opened nest of O. rufohirta with one brood cell and a nest plug at considerable distance from the shell opening consisting of a central layer of earth crumbs sandwiched between two partitions of leaf pulp (photo P. Westrich).

FIGURES 6–15: Osmia (Allosmia). 6: Female of O. gemina. 7: Male of O. gemina. 8: Male sterna 3–6 of O. gemina. 9: Male sterna 3–6 of O. rufohirta. 10: Male genitalia of O. gemina (ventral view). 11: Male genitalia of O. rufohirta (ventral view). 12: Male sterna 3–6 of O. lhotelleriei. 13: Male sterna 3–6 of O. sybarita. 14: Male genitalia of O. lhotelleriei (dorsal view). 15: Male genitalia of O. sybarita (dorsal view).