Andrena russula Lepeletier, 1841

Praz, Christophe, Genoud, David, Vaucher, Killian, Benon, Dimitri, Monks, Joseph & Wood, Thomas J., 2022, Unexpected levels of cryptic diversity in European bees of the genus Andrena subgenus Taeniandrena (Hymenoptera, Andrenidae): implications for conservation, Journal of Hymenoptera Research 91, pp. 375-428 : 375

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; Tengö and Bergström 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.

Figures

Figure 2. Maximum clade credibility tree found in Bayesian analyses of sequence data of the mitochondrial gene COI, showing the phylogenetic relationships among the different species of Andrena subgenus Taeniandrena investigated in the present study. Numbers above branches represent posterior probabilities (values <0.5 are omitted). Taxa mentioned in the text are indicated by arrows or by vertical bars.

Figures 3 - 6. 3, 4 Lectotype female of Andrena afzeliella (Kirby, 1802) 3 profile 4 terga and terminal fringe. Copyright for these two images belongs to the trustees of the Natural History Museum, London 5, 6 lectotype female of A. russula Lepeletier, 1841 5 profile 6 terga.

Figures 9 - 16. Terminal fringes of females of Taeniandrena 9 Andrena afzeliella 10 A. ovatula 11 A. antonellae sp. nov. 12 A. croceiventris 13 A. gelriae 14 A. intermedia 15 A. ovata 16 A. russula.

Figures 19 - 26. Metasomal terga 1 - 3 of females of Taeniandrena 19 Andrena afzeliella 20 A. ovatula 21 A. antonellae sp. nov. 22 A. croceiventris 23 A. gelriae 24 A. intermedia 25 A. ovata 26 A. russula.

Figures 35 - 44. Male antennae 35 Andrena afzeliella 36 A. ovatula 37 A. antonellae sp. nov. 38 A. croceiventris 39 A. gelriae 40 A. intermedia 41 A. ovata 42 A. russula 43 A. vocifera 44 A. wilkella.

Figures 45 - 54. Male genitalia 45 Andrena afzeliella 46 A. ovatula 47 A. antonellae sp. nov. 48 A. croceiventris 49 A. gelriae 50 A. intermedia 51 A. ovata 52 A. russula 53 A. vocifera 54 A. wilkella.