Osmia (Erythrosmia) andrenoides Spinola, 1808

Müller, Andreas, 2020, Palaearctic Osmia bees of the subgenera Hemiosmia, Tergosmia and Erythrosmia (Megachilidae, Osmiini): biology, taxonomy and key to species, Zootaxa 4778 (2), pp. 201-236 : 227-229

Osmia (Erythrosmia) andrenoides Spinola, 1808

Osmia andrenoides Spinola, 1808: 61 . Type material: Syntypes ♀ ♂, “habitat prope Genuam, in loco dicto Marassi” ( Italy), type depository unknown. Type species of Erythrosmia Schmiedeknecht.

Literature records. ALBANIA: Lezha (Lurja E Kurbneshi), Vlora (Borshi S Vlora) ( Tkalců 1974). AUSTRIA: Burgenland, Carinthia, Lower Austria, Styria, Tyrol, Vienna ( Gusenleitner et al. 2012; Zettel et al. 2015). AZER- BAIJAN: Nakhchivan (Kechili, Ordubad) ( Proshchalykin et al. 2018). BELGIUM: Namur ( Pauly 1999; Pauly et al. 2018). CROATIA: Istria (Cerovica, Kanfanar, Koromačno, Rovinj, Salakovci), Krk (Omišalj), Lika-Senj (Baške Oštarie, Susanj), Međimurje (Sveta Jelena), Primorje-Gorski Kotar (Brseč, Golovik, Hreljin, Lovran, Sveti Vid-Mi- holjice), Split-Dalmatia (Trolokve) ( Józan 2009). CYPRUS: Limassol (Limassol, Polemodia Hills) ( Mavromoustakis 1939 a, 1948 a, 1951). CZECH REPUBLIC: Moravia ( Bogusch et al. 2007; P. Bogusch, personal communication). FRANCE: Allier, Alpes-Maritimes, Bouches-du-Rhône, Corrèze, Essone (Saclas, Bouray), Hautes-Pyrénées, Isère, Val-d’Oise (Forêt de Carnelle), Var ( Benoist 1931). GEORGIA: Adjaria (Kintrishi), Mtskheta-Mtianeti (Kumuros khevi), Samzche-Javakheti (Borjomi reserve), ( Kirkitadze & Japoshvili 2015). GERMANY: Baden-Württemberg, Bavaria, Rhineland-Palatinate, Saxony-Anhalt, Thuringia ( Scheuchl & Willner 2016). GREECE: Aegean Islands (Rhodes), Thessaly (Platania) ( Mavromoustakis 1959, Standfuss et al. 2003). HUNGARY: Great Hungarian Plain, Transdanubian Hills, Transdanubian Mountains, North Hungarian Mountains ( Józan 2011). ISRAEL AND PALESTINE: Jerusalem District (Jerusalem), Gaza (Gaza) ( Mavromoustakis 1939b, 1948b). ITALY: Abruzzi, Basilicata, Emilia-Romagna, Friuli-Venezia Giulia, Lazio, Liguria, Lombardia, Molise, Piemonte, Sardegna, Sicilia, Toscana, Veneto ( Pagliano 1994). POLAND: Western Carpathians (Pieniny Mountains) ( Banaszak 2000; Bogdanowicz et al. 2004). PORTUGAL: Estremadura, Faro (Guia) ( Baldock et al. 2018). ROMANIA: Ban-Cal- efariu (2009). RUSSIA: Krasnodar (Anapa), Rostov ( Popov 1954; Fateryga 2017; Proshchalykin & Fateryga 2017). SERBIA: Ascher & Pickering (2019). SLOVAKIA: Bogusch et al. (2007). SLOVENIA: Obalno-kraška, Goriška (Soča valley) ( Gogala 1999). SPAIN: Menorca ( Baldock et al. 2020). SWITZERLAND: Berne, Geneva, Grisons, Neuchâtel, Soleure, St Gall, Ticino, Valais, Vaud ( Amiet et al. 2004). TURKEY: Antalya (Arapsuyu), Artvin (Altiparmak), Aydin (Milet), Erzurum (Oltu, Uzundere, Yesildere), Osmaniye ( Özbek & Zanden 1992; Özbek 2013). UKRAINE: Crimea, Kirovohrad ( Popov 1954; Proshchalykin & Fateryga 2017; Fateryga et al. 2018).

New records. BULGARIA: Blagoevgrad: Sandanski env., 1♂ (leg. M. Kocourek); Burgas: Lozenec, 42°13ʹN / 27°47ʹE, 15 m, 16.6.2008, 3♀, 1♂ (leg. M. Halada); Dobrich: Albena, 29.6.1986, 3♀ (leg. M. Halada); Kardzhali: Balabanova, 41°34ʹN / 25°22ʹE, 350 m, 22.6.2007, 1♀, 2♂ (leg. M. Halada); Plovdiv: Plovdiv, 1.5.1997, 1♀ (leg. Z. Pedr); Schumen: Schumen, 5.6.1997, 1♀ (leg. Z. Pedr); Sofia: 30 km W Sofia, 12.8.1993, 1♀ (leg. M. Halada); Stara Zagora: Galabovo, 10.6.1997, 1♀ (leg. Z. Pedr); Varna: Zlate Pisky, 18.6.1986, 1♀ (leg. M. Halada). CROA- TIA: Dubrovnik-Neretva: Dubrovnik, 1.4.1933, 3♂ (leg. A. Nadig); Istria: 10km E Rovinj, 16.7.2002, 3♂ (leg. Z. Pedr); Korcula: Korcula, 6.8.1934, 1♂ (leg. A. Nadig); Krk: S Stara Baska, 44.94°N / 14.69°E, 17.7.2014, 1♀ (leg. C. Schmid-Egger); Lika-Senj: Gospic, 44°25.8ʹN / 15°31.5ʹE, 600 m, 24.5.2005, 8♂ (leg. Z. Pedr); Rab: between Misnajak and Barbat, 6.7.2004, 1♀ (leg. E. Scheuchl); Šibenik-Knin: 30 km SE Knin, 43°51.4ʹN / 16°29.0ʹE, 450 m, 25.5.2005, 2♀ (leg. Z. Pedr); Split-Dalmatia: 40 km N Split, 43°47.9ʹN 7 16°34.1ʹE, 370 m, 29.5.2005, 6♀, 2♂ (leg. Z. Pedr). FRANCE: Alpes-de-Haute-Provence: Barcelonette, Forêt Domainiale de Ubaye, 28.8.2003, 1♀ (leg. J. Smit); Alpes Maritimes: Sospel, Col de Brouis, 23.7.2001, 1♂ (leg. J. Smit); Aude: Carcassonne, Couffoulens, 22.7.2002, 1♀ (leg. J. Smit), Bouche du Rhône: Camargue, 25.7.1999, 1♂ (leg. J. Smit); Drôme: Dieulefit, 25 km E Montélimar, 23.5.2000, 1♀ (leg. J. van der Smissen); Gard: St. Chaptes, 16.4.1982, 1♂ (leg. F. Amiet); Hérault: St. Guilhem le Désert, 10.5.1994, 1♂ (leg. W. Arens); Var: Vinon sur Verdon, 7.7.2001, 1♂ (leg. J. Smit); Vaucluse: Le Beaucet-Venasque, 10 km SE Carpentras, 1.6.2000, 1♀ (leg. J. van der Smissen). GEORGIA: Khaketi: Chachuna, 2.5.2018, 1♀ (leg. A. Müller). GREECE: Aegean Islands: Andros, Batsi, 37°50ʹ48.61ʺN / 24°47ʹ28.74ʺE, 13.6.2018, 1♀ (leg. A. Haselböck); Chios, Emborios, 38.204°N / 26.021°E, 30.4.2012, 1♀ (leg. M. Taylor); Ikaria, Airport Ikaria, 37.6836°N / 26.3424°E, 20.4.2012, 1♂ (leg. I. Vavitsas); Ios, Kambos, 14.4.2013, 1♀ (leg. T. Pet- anidou); Kea, Dihala Sklavou, 21.4.2013, 1♀ (leg. T. Petanidou); Kos, 18.4.1974, 1♂ (leg. K. Warncke); Lesvos, Eresos, 39.1458°N / 25.9211°E, 6.4.2012, 1♂ (leg. G. Nakas); Limnos, Karvounolakas, 39.9372°N / 25.0586°E, 7.4.2012, 1♀, 1♂ (leg. J. Devalez); Naxos, Fragma, 37.132°N / 25.4398°E, 11.4.2012, 1♀ (leg. I. Vavitsas); Rhodes, Ialissoss, 36.23.094N / 28.09.710E, 28.4.2003, 1♀ (leg. J. Smit); Samos, Pythagorio, 28.5.2001, 1♂ (leg. A. Spring- er); Sifnos, Sellades, 5.2011, 1♀ (leg. E. Papas); Attica: Sounion, 3.6.1963, 1♂ (leg. W. Schlaefle); Central Greece: Bralos, 30 km S Lamia, 10.5.2005, 1♀ (leg. M. Halada); Central Macedonia: Litochoron, 17.8.1978, 1♀ (leg. K. Warncke); Crete: Palaeochora, 6.4.2002, 1♀ (leg.A. Müller); Eastern Macedonia and Thrace: Pangeo, Kavala, 1700 m, 26.7.1980, 1♀ (leg. K. Warncke); Thasos, Potos, 40.613°N / 24.6195°E, 12.4.2012, 1♀ (leg. M. de Courcy); Epirus: 40 km S Igoumenitsa, 16.5.2005, 1♀ (leg. M. Kadlecova); Ionian Islands: Corfu, Evropouli, 8.5.1984, 2♂ (leg. L. Norén); Peloponnese: Mani, Kap Tenaro, 7.6.1996, 2♂ (leg. W. Arens); Thessaly: Trikala, 21.4.1962, 1♀ (leg. K. Warncke); Western Greece: Andritsena (Vassae), 18.5.1996, 1♂ (leg. W. Arens); Western Macedonia: Proastio, Kozani, 620 m, 7.5.1991, 1♀, 1♂ (leg. Z. Pedr). ISRAEL AND PALESTINE: Central District: Lakhish, 31.578ºN / 34.87ºE, 26.2.2016, 1♂ (leg. G. Pisanty); Haifa District: Lower Galilee, Tiv’on, 32º44ʹ15ʺN / 35º08ʹ00ʺE, 7.3.2011, 1♂ (leg. A. Dorchin); Jerusalem District: Judean Mountains, Ya’ar Kedoshim, 26.3.2014, 1♀ (leg. N. Shamir); Northern District: Mt. Tabor, 580 m, 28.5.1991, 1♂ (leg. K. Warncke); West Bank: Quedumim, Karme, Quedem, 27.2.2015, 1♂ (leg. L. Friedman); Southern District: Central Negev, Giv’at Tamar 2km N Sede Boqer, 2.5.2011, 1♀ (leg. A. Dorchin). ITALY: Abruzzi: Massa d’Alba, 900 m, 1.8.2010, 1♀ (leg. A. Müller); Aosta: St. Pierre, 600 m, 1♂ (leg. C. Schmid-Egger); Calabria: Palmi, 3.8.1999, 1♀ (leg. J. Halada); Lazio: Serrone, Monte Scalambre, 1450 m, 20.8.2000, 1♀ (leg. R. Gigli); Piemonte: San Benedetto Belbo, 21.8.2010, 1♀ (leg. G. Pagliano); Puglia: Monte Gargano, San Giovanni, 650 m, 29.5.1993, 1m (leg. A. Müller); Sardinia: Muravera, 2.7.2000, 1♂ (leg. J. Halada); Sicilia: Morgantia near Piazza Amerina, 6.4.1999, 1♂ (leg. W. Arens); Umbria: Todi, 100 km N Roma, 27.7.1999, 1♂ (leg. J. Halada). JORDAN: Amman: Wadi Shu’ayb, 20 km W Amman, 24.4.2007, 3♀, 4♂ (leg. C. Praz, C. Se- divy, A. Müller); Irbid: North Shuna, 22.4.1996, 2♀ (leg. M. Halada); Jerash: 10 km N Jerash, 20.4.2002, 1♀, 1♂ (leg. M. Snizek); Karak: Wadi al Hasa S Al-Karak, 20.4.2007, 5♀ (leg. C. Praz, C. Sedivy, A. Müller); Madaba: Wadi Mujib, King’s Highway, 19.4.2007, 2♀ (leg. C. Praz, C. Sedivy, A. Müller). NORTH MACEDONIA: Southwest: Ohrid, 17.7.1958, 1♀ (leg. W. Schlaefle); Vardar: Titov Veles, 1.5.1963, 1♀ (leg. W. Schlaefle). PORTUGAL: Faro: 3 km W Salema, 7.4.1997, 1♂ (leg. P. Hartmann). ROMANIA: Mehedinti: Cozla, 50 km W Turnu Severin, 26.5.2002, 1♀, 4♂ (leg. M. Halada). RUSSIA: North Caucasian: Pjatigorsk env., 6.6.2002, 1♀ (leg. O.V. Pak). SPAIN: Alicante: Sierra Bernia, 12.8.1976, 1♀; Almería: Berja, Sierra de Gador , 500 m, 10.5.2006, 1♀ (leg. F.J. Ortiz-Sanchez); Barcelona: Gava National Park, 18.5.1999, 1♀ (leg. J. Smit); Cádiz: Chiclana , 23 km S Cadiz , 36.344°N / 6.162°W, 8.4.2010, 1♀ (leg. C. Schmid-Egger); Gerona: 30 km NW Ripoll, 42°22ʹN / 2°53E, 1750 m, 22.7.2011, 3♀ (leg. J. Halada); Granada: Sierra Alfaguara, Huétor-Santillan, 1400 m, 21.5.2006, 1♀ (leg. F.J. Ortiz-Sanchez); Málaga: Torcal, Antequera, 1220 m, 16.5.2008, 1♂ (leg. F.J. Ortiz-Sanchez); Murcia: Sierra de Maria, 25 km W Lorca, 10.5.2003, 1♀ (leg. M. Halada); Tarragona: Ginestar, 50 km N Tortosa, 4.5.2000, 1♀ (leg. E. Scheuchl); Teruel: Albarracin, 8.6.1989, 1♂ (leg. W. Klein). SYRIA: Al-Quneitra: Golan, 2 km NW Hamat Gader, 32°41ʹ39ʺN / 35°39ʹ08ʺE, 3.5.2010, 1♂ (leg. C. Sedivy, C. Praz); Homs: 50 km S Homs, 24.5.1996, 1♀ (leg. M. Halada). TURKEY: Aksaray: Ihlara valley, 38°15ʹN / 36°18ʹE, 13.6.2008, 2♀ (leg. M. Kafka); Ankara: Ankara, Hacettepe University, Beytepe Campus, 3.6.2005, 1♀ (leg. E. Scheuchl); Antalya: between Kernerköy and Gömbe, 36°25ʹ47ʺN / 29°40ʹ08ʺE, 1540 m, 7.6.2006, 1♂ (leg. E. Scheuchl); Aydin: Kusadasi, 19.4.1981, 1♀ (leg. K. Warncke); Balikesir: Arkent, Aywalik, 28.6. – 10.7.1993, 1♀, 1♂ (leg. P. Tyrner); Bartin: Amasra, 26.9.1986, 1♀ (leg. S. Risch); Bolu: 17 km S Seben, 17.6.1998, 1♀ (leg. J. Halada); Burdur: Tefenni env., 37°16ʹ10ʺN / 29°52ʹ44ʺE, 1150 m, 8.8.2006, 1♀ (leg. E. Scheuchl); Hakkari: 25 km S Hakkari, 1200 m, 13.6.1981, 1♀ (leg. K. Warncke); Konya: Bozkir, 26.5.1998, 1♀ (leg. M. Halada); Malatya: Erkenek, 80 km SW Malatya, 9.7.1997, 1♂ (leg. M. Halada); Manisa: 35 km SEE Salihii, 38°20ʹN / 28°24ʹE, 900 m, 30.6.2006, 1♀ (leg. M. Halada); Mardin: 20 km N Mardin, 21.6.1997, 3♂ (leg. M. Halada); Mersin: Uzuncaburc, 30 km N Silifke, 28.5.1996, 2♀, 1♂ (leg. M. Halada); Nevşehir: Capadocia, Ürgüp, 13.6.1998, 1♂ (leg. M. Halada). UKRAINE: Crimea: Simferopol, 30.6.2002, 1♀ (leg. Gurko).

Distribution. Southern, central and eastern Europe northwards to about 51° northern latitude ( Albania, Austria, southern Belgium, Bulgaria, Croatia, Cyprus, Czech Republic, France, southern and central Germany, Greece including Aegean Islands and Crete, Hungary, Italy including Sardinia and Sicilia, North Macedonia, southernmost Poland, Portugal, Romania, southernmost Russia, Serbia, Slovakia, Slovenia, Spain, Switzerland, southern Ukraine), Turkey, Caucasus ( Azerbaijan, Georgia), Levant ( Israel and Palestine, Jordan, Syria).

Pollen hosts. Polylectic with affinity for nototribic flowers of both Lamiaceae (e.g. Acinos , Ajuga , Ballota , Prasium , Satureja , Stachys , Teucrium ; Fig. 24 ) and Antirrhineae (e.g. Linaria ) as well as for Fabaceae (e.g. Lotus , Onobrychis , Trifolium, Genisteae ) (Tab. 1). Additional pollen sources include Crassulaceae , Cistaceae (e.g. Cistus , Helianthemum ), Brassicaceae, Echium (Boraginaceae) , Campanulaceae , Plantago (Plantaginaceae) and Asteraceae (Cichorioideae) . The females are equipped with a specialized pilosity on clypeus and frons composed of apically twisted bristles ( Fig. 25, 26 ), which are used to comb pollen from the raised anthers of nototribic flowers ( Müller 1996).

Nesting biology. O. andrenoides nests in empty snail shells of small to medium size (e.g. Cepaea, Cernuella, Helix, Pomatias, Theba, Xerolenta or Zebrina ), which are hidden at (semi-)shaded places under prominences of stones and rocks or under large flat stones ( Fig. 27 ), which might explain the species’ distinct preference for rocky and stony habitats ( Ferton 1894, 1908; Stöckhert 1933; Popov 1954; Müller & Richter 2019; G. Le Goff personal communication). The females do not cover the surface of the nest shells with patches of leaf pulp after nest site selection. The nests contain 1 – 3 brood cells, which are separated from each other by one-layered partitions constructed from leaf pulp, e.g. of Helianthemum (Cistaceae) or Potentilla (Rosaceae) ( Ferton 1894; G. Le Goff personal communication). They are not sealed against their rear end with a basal wall. After completion of the outermost brood cell, the shells are barricaded towards the nest opening with densely packed earth crumbs, small stones and other particles followed by two adjacent walls of leaf pulp at the shell opening, which enclose a narrow interspace usually filled with few earth crumbs or small stones ( Fig. 28, 29 ). In sealed nests, the closed shell orifice faces upwards or to the side, suggesting that the nest shells are not moved at the end of the nesting cycle. O. andrenoides overwinters as a fully developed imago in a self-spun cocoon within the brood cell ( Müller & Richter 2019). Phenological data, which cover an extended period from early spring to early fall at some localities (see above and Özbek 2013), suggest that the species has two generations per year at more southern latitudes, such as southern Europe or mediterranean Turkey. Brood parasites: The chrysidid wasps Chrysis analis Spinola , Chrysura cuprea (Rossi) and Chrysura dichroa (Dahlbom) develop as parasitoids in the nests of O. andrenoides ( Berland & Bernard 1938; Rosa et al. 2017).

Figures

FIGURES 24–33: Osmia (Erythrosmia). 24: Female of O. andrenoides on Stachys recta (Lamiaceae; photo A. Krebs). 25: Face of female of O. andrenoides with specialized pollen-collecting bristles on clypeus and frons. 26: Pollen-collecting bristles on female clypeus of O. andrenoides in side view. 27: Nesting site of O. andrenoides in cavity under rock prominence. 28: Nest of O. andrenoides in shell of Zebrina detrita sealed with wall of leaf pulp. 29: X-rayed nest of O. andrenoides with entrance barrier of densely packed small particles. 30–33: Male terga 6–7 of O. andrenoides (30), O. erythrogastra (31), O. relicta (32) and O. sparsipuncta (33).