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 antonellae Praz & Genoud sp. nov. Figs 11 , 21 , 37 , 47 , 56-61 Type material. Holotype ♀ (Figs 11 , 21 , 56-58 ), Italy • Sardinia, Buggerru, Cala Domestica; 39°22'36"N , 8°22'57"E [ 39.3767°N , 8.3825°E ]; 17.iv.2017; leg. J. Litman, C. Praz; unique identifier: GBIFCH00117710 (PRUN) [DNA extraction number 1209]. Paratypes (Suppl. material 2: Table S2): 15 ♀ 5 ♂. France • ♀; Corse, Agriates ( "Desert des"); [ 42.6615°N , 9.1579°E ]; 24.4.2011; leg. M. Aubert (DGC) [DNA extraction number 1210] • ♀; Corse, Bonifacio; 41.4034°N , 9.1204°E ; 13.4.2017; leg. A. Cornuel; unique identifier: FER17ALCW2660 (MNHN) • ♀; Corse, Bonifacio; 41.3878°N , 9.1578°E ; 6.4.2017; leg. A. Cornuel; unique identifier: FER17ALCW2589 (MNHN) • ♀; Corse, Bonifacio; 41.3878°N , 9.1578°E ; 6.4.2017; leg. A. Cornuel; unique identifier: FER17ALCW2589 (MNHN) • ♀; Corse, Bonifacio; 41.4034°N , 9.1204°E ; 13.4.2017; leg. A. Cornuel; unique identifier: FER17ALCW2660 (MNHN) • ♂; Corse, Bonifacio, Pertusato A; 41.3710°N , 9.1811°E ; 11.4.2017; leg. A. Cornuel; unique identifier: FER17ALCW1806 (MNHN) [DNA extraction number 1301] • ♂; Corse, Bonifacio, Pertusato B; 41.3709°N , 9.1813°E ; 11.4.2017; leg. A. Cornuel; unique identifier: FER17ALCW1811 (MNHN) • ♂; Corse, Bonifacio, Pertusato B; 41.3709°N , 9.1814°E ; 11.4.2017; leg. A. Cornuel; unique identifier: FER17ALCW1811 (MNHN) • ♀; Corse, Bonifacio, Ville; 41.3865°N , 9.1550°E ; 25.4.2017; leg. Perrard & Cornuel; unique identifier: C17 0600 (MNHN) • ♀; Corse, Bonifacio, Ville; 41.3865°N , 9.1550°E ; 25.4.2017; leg. Perrard & Cornuel; unique identifier: C17 0600 (MNHN) • ♀; Corse, Ghisonacia, Pinia; [ 42.0305°N , 9.4796°E ]; 17.5.2018; leg. D. Genoud (DGC) [DNA extraction number 1297] • ♂; Corse, La Trinite ; 41.4035°N , 9.1203°E ; 28.3.2017; leg. A. Perrard; unique identifier: C17 0197 (PRUN) [DNA extraction number 1294] • ♀; Corse, Pertusato; 41.3710°N , 9.1811°E ; 24.4.2017; leg. A. Perrard; unique identifier: C17 0476 (MNHN) • ♀; Corse, Pertusato; 41.3710°N , 9.1811°E ; 24.4.2017; leg. A. Perrard; unique identifier: C17 0476 (MNHN) • ♀; Corse, St-Julien; 41.3902°N , 9.1803°E ; 27.3.2017; leg. A. Perrard; unique identifier: C17 0090 (PRUN) • ♀; Evisa, Col de Vergio; [ 42.2899°N , 8.8786°E ]; 29.6.2019; leg. Bertrand Schatz (DGC) [DNA extraction number 2337] • ♀; Evisa, Col de Vergio; [ 42.2899°N , 8.8786°E ]; 29.6.2019; leg. Bertrand Schatz (DGC) [DNA extraction number 2341] • ♀; Evisa, Col de Vergio; [ 42.2899°N , 8.8786°E ]; 29.6.2019; leg. Bertrand Schatz; unique identifier: GBIFCH00117729 (PRUN) [DNA extraction number 2340]. Italy • ♀; Sardinia, Buggerru, Cala Domestica; 39°22'36"N , 8°22'57"E [ 39.3767°N , 8.3825°E ]; 17.iv.2017; leg. J. Litman; C. Praz (PRUN) • ♂; Sardinia, Monte Crescia; 39.295°N , 9.392°E ; 7.5.2018; leg. D. Benon ; unique identifier: GBIFCH00117711 (PRUN) [DNA extraction number 1437]. Distribution. So far known only from the Islands of Sardinia and Corsica. In Corsica, the species is known from several localities in the south near Bonifacio, one locality along the east coast, as well as one locality in the north of the Island. In Sardinia, the species is known from two localities in the south of the Island. Andrena antonellae sp. nov. is probably widely distributed on both islands. Pollen preferences. Unknown; two females have been captured on an unidentified yellow Fabaceae shrub (CJP, unpublished data). Phenology. Presumably univoltine with one generation from March until the end of June depending on elevation. Diagnosis. Female. Females of Andrena antonellae sp. nov. are characterised by the small body length (8-9 mm), the brown-orange vestiture of the mesosoma and the head (Fig. 56 ; this colour pattern is shared with A. afzeliella on Sardinia and Corsica), the bronze, dark orange colour of the terminal fringe (Fig. 11 ), and the very shallow (nearly indiscernible) punctation of terga 1 and 2 (Fig. 21 ). An unusual feature is the partly orange colour of the hind femora (Fig. 21 ), a character shared by Corso-Sardinian populations of A. afzeliella . Andrena antonellae sp. nov. is morphologically most similar to A. russula (= A. similis ), A. fuliginata and A. croceiventris ; according to current knowledge, these three taxa are absent from Sardinia and Corsica [the map for Andrena russula (= A. similis ) presented in Gusenleitner and Schwarz (2002 : 1175) includes a mention from Corsica; this mention probably refers to A. antonellae sp. nov.]. Compared to A. russula , which also has shallow punctation on T1-2, A. antonellae sp. nov. is smaller, the vestiture on the scutellum is less dense, both scutum and scutellum are usually less shiny (Fig. 57 ) (scutum entirely matt, scutellum weakly shiny, compared to scutum weakly and scutellum strongly shiny in A. russula ), and the tergal hairbands are much shorter (compare Figs 21 and 26 ). Andrena fuliginata is sculpturally similar to A. antonellae sp. nov.; in the latter species, T1 and T2 are shinier (Fig. 21 ), the scutellum less densely punctate (Fig. 57 ) and the clypeus more shiny and more coarsely punctate (Fig. 58 ). In addition, A. fuliginata is characterised by the brown-orange vestiture on the scutum with short dark hairs intermixed with the longer brown-orange hairs, and the terminal fringe is dark brown; in A. antonellae sp. nov., the scutal vestiture is lighter, there are no or only very few short dark hairs intermixed with the longer hairs (Figs 56 , 57 ), and the terminal fringe is brown-orange (Fig. 11 ). Lastly, A. croceiventris is sculpturally close to both A. fuliginata and A. antonellae sp. nov., although the terga are shinier in A. croceiventris (Fig. 22 ), the clypeus weakly punctate with the underlying surface strongly shagreened, unlike in A. antonellae sp. nov. A. croceiventris is characterised by the red integument colour of the terga (Figs 12 , 22 ); the extent of the red colour is variable but at least some parts of the discs of T1-2 are usually red. 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 17, 18. Terminal fringes of females of Taeniandrena 17 Andrena vocifera 18 A. wilkella . The Corso-Sardinian populations of A. wilkella (Kirby, 1802) and A. afzeliella , the only two other species of Taeniandrena known so far from Sardinia and Corsica, also have an orange vestiture on the mesosoma, as A. antonellae sp. nov. Since all three species are found in sympatry, the criteria allowing for the identification of the female are summarised in Table 2 . Male. The males of Andrena antonellae sp. nov. are sculpturally nearly identical to those of A. croceiventris and A. fuliginata . All three species have a comparatively elongate, oval-shaped genitalia with a narrow penis valve (Figs 47 , 48 ), nearly exactly as in A. wilkella (Fig. 54 ). The latter species can be recognised by the comparatively short antennal segment 3 (Fig. 44 ), which is at most 0.7 x as long as A4, and the markedly denser and coarser tergal punctation. In A. croceiventris , A. fuliginata and A. antonellae sp. nov. A3 is 0.8 times as long as A4 (Figs 37 , 38 ), and the tergal discs are strongly shagreened and weakly punctate. In the lone male specimen of A. croceiventris examined, the integument of all tergal margins, of the pregradular area of T2-4, and of the lateral, declivous parts of T1-5 is orange, and the integument of the hind tibiae is dark. In A. antonellae sp. nov. and the lone male specimen of A. fuliginata examined, the integument of the terga is predominantly dark, the tergal margins are slightly lightened apically, and the hind tibiae are orange (partly brown basally and medially in A. fuliginata ). No sculptural difference could be found between the male of A. antonellae sp. nov. and A. fuliginata , although only one specimen of A. fuliginata has been examined. Lastly, the males of these three species are similar to those of A. russula (= A. similis ), but in the latter the broadened, parallel-sided base of the valve is longer, only tapering near the base of the valve opening (Fig. 52 ), and the antennal segment 3 is usually longer (Fig. 42 ). Description. Female . Measurements. Body length 8-9 mm. Head . Head 1.3 times as wide as long. Clypeus dark, flattened over most of its area, densely and uniformly punctate with exception of a narrow central impunctate line, punctures separated by 0.5 puncture diameters, underlying surface weakly shagreened, usually shiny, especially apically (Fig. 58 ). Face, gena, vertex, and scape with light brownish hairs, longest approximately three quarters of scape in length. Antennae dark, A4-12 lightened to light brown below. Foveae broad, occupying almost all area between lateral ocellus and top of compound eye, filled with short, dark brown hairs. Mesosoma . Scutum densely punctate, punctures separated by <0.5 puncture diameters over majority of surface except becoming slightly sparser centrally and posteriorly, underlying surface strongly shagreened (Fig. 57 ). Scutellum with sparser punctures separated by up to 4 puncture diameters medially, shagreenation weaker, surface weakly shiny. Episternum and propodeum with dense raised reticulation, underlying surface dull, propodeal triangle weakly indicated by weak carina, little differentiated from general reticulation. Scutum and scutellum with erect, orange, shortly plumose hairs, episternum with longer light orange hairs. Front legs, mid coxa, trochanter and femur, hind coxa and trochanter dark brown, mid tibia dark basally and orange apically, mid tibia, tarsi and hind femur, tibia and tarsi orange (Figs 21 , 56 ). Leg pubescence light brown basally, becoming orange apically, flocculus, femoral and tibial scopae light brown to golden. Wings hyaline to weakly infuscate, venation brown orange, stigma light brown. Metasoma . Terga dark, strongly shagreened, weakly shiny (Fig. 21 ), margins yellowish brown apically, slightly lighter than tergal discs. T1 nearly impunctate, usually only very few punctures scarcely visible against shagreenation on disc, sometimes weakly punctate with very shallow punctures; vertical anterior area entirely impunctate; margin strongly shagreened, impunctate. T2-3 slightly more densely and visibly punctate, punctures little visible, separated by 0.5-1 puncture diameters (Fig. 21 ). T4 very densely punctate, punctures little visible, separated by less than 0.5 puncture diameters (Fig. 11 ). T1 without hairband, with a few short hairs laterally, hairs not forming dense hairbands. T2-4 with short, narrow lateral spots, hairband covering a third of tergal width on T2 and T4 and half tergal width on T3 (Fig. 21 ). Remaining tergal surface covered with very short, dark hairs visible when viewed obliquely or in profile. Disc of T4 near margin with a few longer, erect dark hairs. Apical fringe of T5 and hairs flanking pygidial plate bronze-orange (Fig. 11 ), pygidial plate rounded, flat, medially slightly raised, without raised margin. Male. Measurements. Body length 8-9 mm. Head . Head 1.3 times as wide as long (Fig. 60 ). Clypeus flattened and densely punctate, punctures separated by <0.5 puncture diameter, underlying surface shiny. Gena, lower part of face, scape and vertex with greyish-white hairs becoming light brown on scape and vertex, longest equalling length of scape. Antennae dark. A3 0.8 x as long as A4 (Fig. 37 ). Mesosoma . Scutum, scutellum, episternum, and propodeum structurally as in female, punctation overall sparser. Scutum and scutellum with fine yellowish grey hairs that equal length of scape. Front legs dark, mid legs dark except tarsi, orange, hind legs dark, tibiae and tarsi orange (Fig. 59 ). Wings hyaline, venation dark orange, stigma dark orange with brown margin. Metasoma . Terga dark, finely shagreened and weakly shiny, apical part of marginal areas lightened, semi-translucent brown (Fig. 61 ). Terga finely but clearly punctate, punctures separated by 2 puncture diameters. T2-5 laterally with weak apical hairbands of whitish hairs, hairbands interrupted on all terga (Fig. 61 ). Sterna apically with loose, long fringe of yellowish hairs. S8 strap-like, slightly broadened apically, uniformly hairy. Genitalia elongated oval-shaped in dorsal view, gonocoxa with inner margins weakly diverging (Fig. 47 ). Penis valve comparatively narrow, basally parallel-sided before tapering apically. Gonostyli comparatively long, apical blades longer than wide, their external margin weakly concave (Fig. 47 ). Variation. Female specimens from one high elevation locality in Corsica (Evisa, Col de Vergio 1477 m; one specimen is included in Fig. 2 with number 2341) have broader and more furnished tergal hair bands. Etymology. This species is named in honour of Antonella Soro for her contributions to the field of conservation genetics of bees. Note. Andrena antonellae sp. nov. is sculpturally highly similar to A. croceiventris and A. fuliginata , both in the female and male sexes. Based on current evidence, these three taxa do not occur in sympatry. The highly similar morphology in these three taxa conflicts with our DNA barcoding results, which suggest that the three taxa are only distantly related. This discrepancy between molecules and morphology is reminiscent of the strong genetic divergences between A. afzeliella and A. ovatula ; these two cases are puzzling and are further discussed below. While A. antonellae sp. nov., A. fuliginata and A. croceiventris are morphologically close, there are still subtle differences among them; these differences include the colour of the terminal fringe, the colour of the integument, the presence or absence of short dark intermixed hairs on the scutum, the sculpture of T1 and T2, and the sculpture of the clypeus. In Taeniandrena , such differences, although subtle, generally correspond to between-species differences. For this reason and based on the strongly divergent DNA barcodes, we treat these three taxa as distinct.