Andrena (Warnckandrena) innesi Gribodo 1894

Pisanty, Gideon & Wood, Thomas James, 2024, The early-diverging subgenera of the bee genus Andrena (Hymenoptera: Andrenidae) in the Old World, Zootaxa 5474 (5), pp. 451-488 : 461-462

Andrena (Warnckandrena) innesi Gribodo 1894

( Figs. 10–11 , 30–32, 38, 47–55 )

Andrena innesi Gribodo, 1894: 122 , ♀ [ Algeria: MCSN].

Andrena mastrucata Gribodo, 1894: 123 , ♂ [ Algeria: MCSN], syn. Warncke 1967: 286.

Andrena sefrensis Pérez, 1895: 41 , ♀ [ Algeria: MNHN], syn. Warncke 1967: 307.

Melittoides tunensis Friese, 1921: 179 , ♂ [ Tunisia: ZMB], syn. Warncke 1967: 315 ( Figs. 47–49 ).

Melittoides vulpinus Friese, 1921: 179 , ♂ (nec Apis vulpina Christ, 1791 nec Apis vulpina Panzer, 1798 nec Andrena vulpina Fabricius, 1804 ) [ Egypt: ZMB], syn. Warncke 1967: 317 ( Figs. 50–52 ).

Andrena aemula Alfken, 1926: 105 , ♀ ♂ (nec Andrena aemula Alfken, 1926: 98 ) [ Egypt: SMFD], syn. Gusenleitner & Schwarz 2002: 369 ( Figs. 53–55 ).

Andrena Confalonierii Guiglia, 1929: 410 , ♀ [ Libya: MCSN], syn. Warncke 1967: 262.

Andrena (Melittoides) innesi ssp. undata Warncke 1969: 404 , ♀ ♂ [ Israel: OLML].

Andrena (Melittoides) innesi ssp. tantana Warncke, 1980: 84 , ♀ [ Morocco: NHMUK].

Andrena innesi Gribodo : Warncke 1967: 279.

Andrena (Melittoides) innesi Gribodo : Warncke 1968: 45.

Andrena (Melittoides) innesi Gribodo : Gusenleitner & Schwarz 2002: 369–370.

Distribution: Morocco, Algeria, Tunisia, Libya, Egypt, Israel, West Bank, Jordan, Turkey, Iran ( Gusenleitner & Schwarz 2002; Khodaparast & Monfared 2012).

Flight period: February–May.

Flower records: Asteraceae ; Brassicaceae : Brassica ( Khodaparast & Monfared 2012) ; Moricandia arvensis ( Djouama et al. 2016) ; Sinapis arvensis ( Benarfa et al. 2013; Dermane et al. 2021); Resedaceae : Reseda alba ( Dermane et al. 2021) .

Pollen preferences: Five pollen loads from Morocco, Tunisia, the West Bank and Turkey all consisted of 100% Brassicaceae pollen ( Sinapis - type, Raphanus - type), strongly suggesting that the species is broadly oligolectic on this plant family across its entire distributional range.

Genetic sequences available: BOLD accession nos. ANDGP007-23 ( Tunisia), ANDGP008-23 ( Jordan), WPATW586-22 ( Morocco).

MATERIAL EXAMINED: HOLOTYPE ( Melittoides tunensis Friese ) ( Figs. 47–49 ): TUNISIA • ♂; Tunis, merid.; [O. Schmiedeknecht leg.]; ZMB .— HOLOTYPE ( Melittoides vulpinus Friese ) ( Figs. 50–52 ): EGYPT • ♂; Kingi [Iking Maryut]; 12 Feb. [19]11; ZMB .— HOLOTYPE ( A. innesi undata Warncke ): ISRAEL • ♀; Kinneret; 9 Apr. 1942; Y. Palmoni leg.; OLML .— SYNTYPE ( Andrena aemula Alfken ) ( Figs. 53–55 ): EGYPT • 1♂; Amrich [Amreya, Alexandria]; 16 Feb. [19]08; A. Andres leg.; SMFD .— PARATYPE ( A. innesi tantana Warncke ): MOROCCO • 1♀; km 31 Goulimine [Guelmim]— Tan Tan Rd.; c. 100 m a.s.l.; 17 Mar. 1974; K.M. Guichard & G.R. Else leg.; flowery border of wheat field; OLML .— PARATYPE ( A. innesi undata Warncke ): ISRAEL • 1♂; Tiberias ; 10 Feb. 1944 [?]; H. Bytinski-Salz leg.; OLML. — non-type material: ALGERIA • 1♂; Hammam Bou Hadjar ; 1–30 Apr. 1910; O. Schmiedeknecht leg.; SMFD • 1♂; 6 km E of Saida; 28 Apr. 1983; R. Leys & P. v. d. Hurk leg.; RMNH ZMA.INS.5090150. — ISRAEL • 2♂; Arad ; 7 Feb. 1987; E. Shney-Dor leg.; SMNHTAU • 1♀; Beer Sheva [Beersheba]; 30–31 Mar. 1961; CNC 1287405 • 1♀; Kinneret ; 9 Apr. 1942; Palmoni leg.; OLML • 1♂; Tiberias ; 10 Feb. 1948; Bytinski-Salz leg.; OLML. — JORDAN • 1♂; Al Karak env.; 6 Apr . 2013; Snižek leg.; OLML • 1♂; ibid.; BOLD accession no. ANDGP008-23. — LIBYA • 1♂; Bengasi [Benghazi]; 8 Feb. 1922; Geo C. Krüger leg.; OLML • 1♀; R. U. Agrario [Royal Agricultural Office]; 25 Mar. 1925; Geo C. Krüger leg.; OLML. — MOROCCO • 1♂; Bir Rechid [Berrechid]; 16 Feb. 1934; Rungs leg.; OLML • 1♀, 1♂; 10 km E Guelmin [ Guelmim ]; 15–16 Apr. 1995; M. Halada leg.; OLML • 2♂; 40 km SW Guelmim ; 13 Apr. 1995; M. Halada leg.; OLML • 1♂; Guelmim-Oued Noun, Guelmim, Aferkat , 5 km S; 28.591° N, 10.296° W; 700 m a.s.l.; 18 Mar. 2022; T.J. Wood leg.; BOLD accession no. WPATW586-22; TJWC. — TUNISIA • 1♂; 5 km S Le Kef [El Kef]; 24 Mar. 1976; U. N. Lanham, P. Robinson & B. Solounias leg.; mixed composites; OLML • 1♀; “ Hammam ” [? Hammamet]; OLML • 1♀; 20 km SW Medenine; 13 Apr. 1994; M. Schwarz leg.; OLML • 1♀; Sbeitla ; 12 Apr. 1998; K. Deneš jn. leg.; OLML • 1♀; ibid.; BOLD accession no. ANDGP007-23. — TURKEY • 1♂; Hatay Prov., Topboğazı ; 14 Apr. 1976; K. Warncke leg.; OLML • 1♀; Şanlıurfa Prov., Ceylanpınar ; 25 Apr. 1976; K. Warncke leg.; OLML • 1♀; Şanlıurfa Prov., 10 km N Ceylanpınar ; 25 Apr. 1976; K. Warncke leg.; OLML • 1♂; 20 km W Kilis; 27 Apr. 1976; K. Warncke leg.; OLML • 1♀; 40 km E Urfa; 14 May 1975; K. Warncke leg.; OLML. — WEST BANK • 1♀; Faza’el [Petza'el]; 9 Apr. 1986; I. Susman leg.; SMNHTAU 374708 • 1♂; Jericho ; 11 Apr. 1909; OLML • 1♀; Maale Haadumim [Ma'ale Adumim]; 1 Apr. 1975; H. Bytinski-Salz leg.; SMNHTAU 374709 • 2♂; Masua ; 12 Mar. 1985; A. Hefetz leg.; SMNHTAU 367749 , 367750 .

Remarks. Andrena innesi exhibits considerable variation in specimen colour and size across its range, which has led to the prolific description of multiple synonyms and subspecies by various authors. Our preliminary measurement of several key characters (head, glossa, malar area, flagellomeres, ocelloccipital distance, genal area) exhibited rather consistent allometric proportions across the entire species range. This, as well as the globally conserved genital morphology, argues for maintaining a unified species concept of A. innesi sensu lato. On the other hand, preliminary molecular barcode data suggests a considerable genetic distance (7.6%) between populations in the Levant versus North Africa. Unfortunately, most museum specimens of A. innesi are too old for molecular barcoding, and therefore barcode data are scarce. No nomenclatural action is currently undertaken until further taxonomic research is made on this group.

Among the various synonyms of A. innesi , the case of Andrena aemula requires some discussion for clarity; two of the three specimens mentioned by Alfken (1926: 98) were located by us in the SMFD collection, whereas Warncke (1967: 202) only mentioned a female co-type in the ZMB collection, but we have not been able to locate this specimen. Andrena aemula was proposed by Alfken (1926: 98) as a replacement name for three unavailable names that he considered to belong to the same species: A. bipartita Lepeletier, 1841 nec Brullé, 1839; A. lepeletieri Dalla Torre, 1896 nom. nov. pro A. bipartita nec Lucas, 1849; and Melittoides vulpinus Friese, 1921 nec Andrena vulpina Fabricius, 1804 . There is much to unpack here. The type material of A. bipartita Lepeletier is conspecific with A. nigroaenea (Kirby, 1802) ( Warncke 1967: 173) . Therefore, de facto, the replacement name A. aemula Alfken is a junior synonym of A. nigroaenea , as listed by Gusenleitner & Schwarz (2002: 526). Furthermore, Alfken combined two species concepts together, as the material he selected and named A. aemula contained two species, female A. nigroaenea and male A. innesi . He was therefore correct to associate the female with A. bipartita Lepeletier and the male with M. vulpinus Friese , but the association of the two sexes to one another is incorrect.

Additionally, Alfken did not only propose A. aemula as a replacement name for these combined concepts, but then described A. aemula from male and female specimens ( Alfken 1926: 105). By writing in this order, he unintentionally preoccupied the name A. aemula , making A. aemula Alfken nec Alfken a junior primary homonym. The two specimens marked as type in the SMFD collection are a male from Amrich [Amreya] which is indeed A. innesi ( Figs. 53–55 ), and a female from Kingi [Iking Maryut] which is a specimen of A. nigroaenea ( Figs. 56–58 ). Andrena aemula Alfken nec Alfken could therefore be listed as a synonym of either A. nigroaenea or A. innesi depending on which syntypic specimen is considered to be representative of the species. Given that it was clearly the intention of Alfken to describe Friese’s species under a new name, and Alfken did not explicitly indicate that one specimen was the holotype, we treat the male syntype as representative of the species and treat A. aemula Alfken nec Alfken as a junior synonym of A. innesi . This is consistent with the chresonymy of Gusenleitner & Schwarz (2002), although strictly following Warncke (1967) and the female syntypic specimen reported from the ZMB collection, one would have to list A. aemula Alfken nec Alfken as a junior synonym of A. nigroaenea . A replacement name could be proposed for A. aemula Alfken nec Alfken, but given the many available names for A. innesi , this is considered to be unnecessary.

Figures

FIGURES 10–17. Andrena (Warnckandrena) female habitus (10, 12, 14, 16) and heads (11, 13, 15, 17): 10–11, A. innesi Gribodo; 12–13, A. papio sp. nov.; 14–15, A. ramlehiana Pérez; 16–17, A. otoolei sp. nov.

FIGURES 47–58. Andrena types, habitus (47, 50, 53, 56), heads (48, 51, 54, 57) and labels (49, 52, 55, 58): 47–49, Melittoides tunensis Friese, male holotype; 50–52, M. vulpinus Friese, male holotype; 53–55, Andrena aemula Alfken 1926 (nec Alfken 1926: 98), male syntype; 56–58, A. aemula Alfken 1926 (nec Alfken 1926: 98), female syntype.