Unexpected levels of cryptic diversity in European bees of the genus Andrena subgenus Taeniandrena (Hymenoptera, Andrenidae): implications for conservation Author Praz, Christophe https://orcid.org/0000-0003-2649-3141 University of Neucha ̂ tel, Institute of Biology, Rue Emile-Argand 11, 2000 Neucha ̂ tel, Switzerland & InfoFauna - Swiss Zoological Records Center, Avenue de Bellevaux 51, 2000 Neuchatel, Switzerland christophe.praz@unine.ch Author Genoud, David Avenue des Roses 2, 87240 Ambazac, France Author Vaucher, Killian University of Neucha ̂ tel, Institute of Biology, Rue Emile-Argand 11, 2000 Neucha ̂ tel, Switzerland & InfoFauna - Swiss Zoological Records Center, Avenue de Bellevaux 51, 2000 Neuchatel, Switzerland Author Benon, Dimitri University of Neucha ̂ tel, Institute of Biology, Rue Emile-Argand 11, 2000 Neucha ̂ tel, Switzerland & InfoFauna - Swiss Zoological Records Center, Avenue de Bellevaux 51, 2000 Neuchatel, Switzerland Author Monks, Joseph Natural History Museum, Cromwell Road, London, UK Author Wood, Thomas J. https://orcid.org/0000-0001-5653-224X Laboratory of Zoology, University of Mons, Avenue du Champs de Mars 6, 7000 Mons, Belgium text Journal of Hymenoptera Research 2022 2022-06-30 91 375 428 http://dx.doi.org/10.3897/jhr.91.82761 journal article http://dx.doi.org/10.3897/jhr.91.82761 1314-2607-91-375 3A5B959985024CB7A83ECAA998B678A9 3F1971FA0B4150AD801E63777D5EB924 Andrena russula Lepeletier, 1841 Figs 5 , 6 , 16 , 26 , 42 , 52 Andrena russula Lepeletier, 1841: 251, ♀, "Oran" [Algeria]. Holotype ♀ (MNHN). Andrena similis Smith, 1849: lx, ♂, "Bristol" [UK], syn. nov. Syntype or holotype ♂ (OUMNH). Andrena ocreata cyprisina Warncke, 1975c: 78, ♀,♂, "Limassol" [Cyprus], syn. nov. Holotype ♀ (OLML). Andrena similis caraimica Osytshnjuk, 1994: 33, ♀,♂ [Crimea], syn. nov. Holotype (SIZK). Material examined. Type material: Holotype of A. russula (MNHN, Figs 5 , 6 ). Holotype of A. o. cyprisina ♀ Cyprus • Limassol (OLML) . Other material (Suppl. material 2: Table S2): Algeria • ♀ ; S. Kabylia , Ait Hassem ; [ 36.583 , 4.197 ]; 16- 17.6.1971 ; leg. A. Hoffer, J . Hordk (OLML) [DNA extraction number 1533] . Crimea • 23♂ 28♀ ; Karadagh [Kara Dag], Vodianja balka; 44°56'22"N , 35°12'44"E ; 21.4.2003 ; leg. Y. Budaschkin (OLML) • ♂ 13♀ ; Kap Kasantyp steppe [Kazantyp]; 45°27'47"N , 35°50'40"E ; 1.5.2003 ; leg. Y. Budaschkin (OLML) Iran • ♀ ; Lorestan province , Dorud Lanjaban env, 960 m ; 33.419°N , 48.986°E ; 10.5.2016 ; leg. M. Kafka (TJWC) [DNA extraction number TJW038] . Italy • ♀ ; Marche , M. Sibillini , M. Vettore ; 42.8011°N , 13.2711°E ; 30.6.2011 ; leg. Trunz , Litman , Praz ; unique identifier: GBIFCH00117708 (PRUN) [DNA extraction number 1425] • ♀ ; Umbria , Castel Viscardo , 10 km NW Orvieto ; 42.7574°N , 12.0023°E ; 28.5.1991 ; leg. J. Gusenleitner (OLML) • ♀ ; Umbria , Panicale , S of L. Trasimeno ; 43.0306°N , 12.0969°E ; 9.5.2011 ; leg. D.W. Baldock (TJWC) • ♂ ; VA, Luino ; 46.0007°N , 8.7463°E ; 2.5.1983 ; leg. H. Teunissen (RMNH) . Libya • ♀ ; Tripolitaine , Djebel Ghariane ; 4.1899; leg. Alluaud (OLML) • ♀ ; Tripolitania , Leptis Magna ; 15.1.1955 ; leg. K. M. Guichard (OLML) • ♀ ; Tripolitania , Sidi Mesri ; 3.1940; leg. G. M. Martelli (OLML) • ♂ ; Tripolitania , Tripoli ; 4.2.1954 ; leg. K. M. Guichard (OLML) • 2♂ ; Tripolitania , Tripoli ; 30.1.1955 ; leg. K. M. Guichard (OLML) . Tunisia • ♀ ; Tunis ; 189? (OLML). Distribution. Widely distributed throughout Europe, including the Iberian Peninsula, France, Central Italy, Bulgaria, Cyprus, Crimea, northern Africa from Morocco to Libya, Turkey, the Caucasus, the Levant (Israel, Syria, Lebanon, Jordan) and Iran. Pollen preferences. Oligolectic on Fabaceae ( Westrich 1989 , as A. similis ). Phenology. Univoltine, in Switzerland from the end of April to early June at low elevations, slightly later at high elevations. Note. This widespread species has so far mostly been referred to as A. similis . A first change to this view was advocated by Warncke (1967 : 174), who stated that A. russula was the "form" with red-coloured vestiture of A. similis in Northern Africa, and that the European form of A. similis should thus be named A. russula ssp. similis . In 1970, he resurrected the name Andrena ocreata (Christ, 1791) for this taxon based on his interpretation of the vague original description of Apis ocreata ( Warncke 1970 ). While he invokes article 23b of the 1958's edition of the International Code of Zoological Nomenclature to uphold numerous junior synonyms in the same article, he does not do so for A. similis , possibly because he considered Hylaeus similis Fabricius, 1793 to be a synonym of Andrena barbilabris (Kirby, 1802), making Andrena similis Smith a junior secondary homonym. The then prevailing version of the code did not allow for a reversal of precedence in the case of secondary homonymy, which possibly explains Warncke's decision to resurrect the name Andrena ocreata for this taxon. The reason why he does not mention A. russula in the 1970 article is unclear. He later designated a neotype for Apis ocreata , selecting a female of " Andrena similis Smith" collected in Erlangen, Germany ( Warncke 1986 ). His interpretation of A. ocreata has not been followed (e.g., Westrich 1989 ; Gusenleitner and Schwarz 2002 ), and the name A. similis has mostly been used until now for this taxon. We consider Hylaeus similis Fabricius and Apis ocreata Christ to be nomina dubia . This species is widely distributed in the Western Palearctic; geographic variation in structural morphology is minimal, and variation that does exist such as in the strength of tergal punctation follows no clear pattern or gradient. In north-western Africa, in populations referred to as " A. ocreata ssp. russula " by Warncke (see Gusenleitner and Schwarz 2002 : 1175), the vestiture of the females is bright red orange, as opposed to brown orange in Europe. This pattern of increased orange intensity in North African populations can clearly be seen in unrelated taxa such as A. lepida Schenck, 1861 and A. numida Lepeletier, 1841 (possibly conspecific with the European taxon A. hypopolia Schmiedeknecht, 1884; Wood, unpublished data). In addition, the integument of the hind legs of the females of north-western African populations of A. russula , in particular of the hind femora, is sometimes entirely orange, although not consistently so. Sculpturally, Andrena " Andrena russula " (or the north-western African populations) and Andrena " Andrena similis " (or the European populations) are morphologically very close and no character allows for an unambiguous separation; in males of " A. russula ", the antennal segment 3 is slightly shorter than A4, while in " A. similis ", A3 is as long as or longer than A4 (Fig. 42 ). The length of A3 and A4 is highly variable throughout the range of " A. similis " and we consider this criterion as variable within this taxon. We were able to obtain a short COI sequence from one specimen of Andrena " Andrena russula " from northern Algeria, likely corresponding to the type locality of Andrena russula (specimen 1533 in Fig. 2 ). This sequence clusters within a large clade that includes numerous specimens of A. similis from Europe, Morocco, Cyprus and Israel. We consequently place A. similis as a synonym of A. russula and consider one largely distributed taxon. As for A. afzeliella and A. albofasciata , a reversal of precedence (article 23.9 of the Code of Zoological Nomenclature) to maintain the prevailing usage of A. similis is not justified since A. russula has been used as a valid name after 1899 (e.g., Warncke 1967 ; Tengoe and Bergstroem 1975 ; Gusenleitner and Schwarz 2002 ), therefore rejecting the first condition of article 23.9 of the code. Diagnosis. See Schmid-Egger and Scheuchl (1997) , Amiet et al. (2010) (both as Andrena similis ) and the identification key below.