The genus Andrena Fabricius, 1775 in the Iberian Peninsula (Hymenoptera, Andrenidae) Author Wood, Thomas J. https://orcid.org/0000-0001-5653-224X University of Mons, Research Institute for Biosciences, Laboratory of Zoology, Place du Parc 20, 7000, Mons, Belgium thomasjames.wood@umons.ac.be text Journal of Hymenoptera Research 2023 2023-05-22 96 241 484 http://dx.doi.org/10.3897/jhr.96.101873 journal article http://dx.doi.org/10.3897/jhr.96.101873 1314-2607-96-241 15A2B06B92F34E70AC8F6FEABF365E71 A5722C06212C5BA7A2099B77C6A8DF54 Andrena (Truncandrena) ghisbaini Wood, sp. nov. Type material. Holotype . Spain : Malaga , PN Sierra de las Nieves, mountain peak S of Pinsapo Escalereta, 36.6621°N , - 5.0362°W , 1600 m, 30.v.2021, 1♀, leg. T.J. Wood, OOELM [BOLD accession number WPATW239-21]. Paratypes . Spain : Malaga , PN Sierra de las Nieves, mountain peak S of Pinsapo Escalereta, 1600 m, 30.v.2021, 1♀, leg. G. Ghisbain, TJWC; Malaga - Elvira, 11.ii.1981, 4♂, leg. H. Teunissen, RMNH. Description. Female. Body length: 15-16 mm (Fig. 51A ). Head : Dark, 1.2 times wider than long (Fig. 51B ). Clypeus weakly domed, clearly punctate, punctures separated by 0.5-1 puncture diameters with exception of median longitudinal impunctate line, narrow basally, broadening apically, thus elongate triangular; underlying surface finely shagreened, weakly shiny. Process of labrum broadly trapezoidal, twice as broad as long, apical margin clearly emarginate. Gena broad, almost two times width of compound eye; ocelloccipital distance 1.5 times diameter of lateral ocellus. Fovea dorsally occupying slightly more than 1/2 space between lateral ocellus and compound eye, extending ventrally to lower margins of antennal insertions, filled with black hairs. Gena ventrally and laterally with long light brown hairs, longest equalling length of scape, hairs becoming black on vertex, scape, and majority of face, with shorter light brown hairs around antennae insertions. Antennae basally dark, A4 apically, A5-12 ventrally extensively lightened orange; A3 exceeding A4+5, shorter than A4+5+6. Mesosoma : Scutum and scutellum with extremely shallow and obscure punctures, punctures separated by 0.5-1 puncture diameters, punctures disappearing into underlying fine granular shagreen, surface dull to very weakly shiny. Pronotum rounded. Mesepisternum and dorsolateral parts of propodeum with fine granular shagreen, weakly shiny, with fine and scattered raised hair-bearing punctures, punctures separated by 2-3 puncture diameters; propodeal triangle broad, with extremely fine granular shagreen, without hair-bearing punctures, propodeal triangle thus defined by change in surface sculpture from dorsolateral parts of propodeum. Mesepisternum and propodeum with long finely plumose light brown hairs, clearly exceeding length of scape; scutum and scutellum medially with long black hairs and occasional isolated pale hairs covering majority of disc, laterally becoming intermixed with light brown hairs. Propodeal corbicula incomplete, very weakly defined, dorsal fringe not differentiated from hairs of internal surface, both parts composed of long finely plumose long light brown hairs. Legs dark, apical tarsal segments lightened dark reddish, pubescence dark brown. Flocculus complete, composed of long weakly plumose and upturned light brown hairs; femoral scopae composed of light brown simple hairs; tibial scopa long, hairs exceeding apical width of hind tibia, hairs dorsally dark brown, ventrally golden orange (Fig. 51D ). Hind tarsal claws with inner tooth. Wings hyaline, stigma dark brown, venation dark brown to orange, nervulus interstitial. Metasoma : Terga dark, apical rim of marginal areas narrowly lightened hyaline-yellow; discs with extremely fine granular shagreen, weakly shiny, with fine and scattered hair-bearing punctures, punctures separated by 3-4 puncture diameters (Fig. 51C ). Disc of T1 with long light brown hairs, exceeding length of scape, decreasing in length over T2-3, disc of T3 with intermixed short light brown and black hairs, becoming predominantly black on discs of T4-5. T2-4 with weak apical fringes of short light brown hairs, not obscuring underlying surface. Apical fringe of T5 and hairs flanking pygidial plate dark brown, pygidial plate rounded triangular, with weakly raised medial area, otherwise featureless. Figure 51. Andrena (Truncandrena) ghisbaini sp. nov. female A profile B face, frontal view C terga, dorsal view D tibial scopa, profile view. Male. Body length: 13-14 mm (Fig. 52A ). Head : Dark, 1.3 times wider than long (Fig. 52B ). Clypeus weakly domed, entirely yellow-marked with exception of two dark rounded spots medio-laterally. Clypeus punctured, punctures separated by 0.5-1 puncture diameters with exception of median longitudinal impunctate line, essentially non-existent basally, broadening apically, thus elongate triangular; underlying surface finely shagreened, weakly shiny. Process of labrum broadly rectangular, 2.5 times wider than long, apical margin weakly emarginate, surface smooth and shiny. Gena broad, 2 times width of compound eye; ocelloccipital distance 1.5 times diameter of lateral ocellus. Gena ventrally and laterally with long light brown hairs clearly exceeding length of scape, becoming intermixed with black hairs on vertex. Face medially with extensive whitish to light-brownish hairs on clypeus, antennae insertions, and scape intermixed with black hairs along inner margins of compound eyes and scape. Antennae basally dark, A4-13 ventrally lightened orange; A3 longer than A4, shorter than A4+5; A4 rectangular, longer than broad, slightly shorter than A5. Mesosoma : Mesosoma structurally as in female; pubescence as in female. Legs basally dark, apical tarsal segments and hind tibiae lightened dark reddish-brown, pubescence dark brown to orange brown. Hind tarsal claws with inner tooth. Wings hyaline, stigma orange, venation dark brown to orange, nervulus interstitial. Metasoma : Terga structurally as in female. Discs of T1-4 with long light brown hair, on T1 clearly exceeding length of scape, becoming progressively shorter to T4; T5-6 with short black hairs on disc (Fig. 52C ). T2-4 with weak apical hair fringes apically, not obscuring underlying surface. S8 columnar, apex rounded, ventral surface covered with short brown hairs. Genital capsule slightly elongate, gonocoxae produced into strong apical teeth, teeth pointed with apical margins diverging (Fig. 52D, E ). Gonostyli basally narrow, apically produced and flattened into rounded triangular plates, internal margin strongly raised and reflexed. Penis valves basally broad, occupying more than 1/2 space between gonostyli, with narrow laterally produced hyaline extensions; penis valves strongly narrowing apically. Figure 52. Andrena (Truncandrena) ghisbaini sp. nov. male A profile B face, frontal view C terga, dorsal view D genital capsule, dorsolateral view E genital capsule, dorsal view; Andrena (Truncandrena) villipes Perez , 1895 male F genital capsule, dorsal view. Diagnosis. Andrena ghisbaini can be recognised within Truncandrena due to its characteristically smooth and finely granulate propodeal triangle which contrasts with the similarly granulate dorsolateral parts of the propodeum which bear fine and scattered raised hair-bearing punctures, the rounded pronotum, the linear malar space, the large body size (>13 mm), yellow male clypeus, and typical genital capsule with the inner margins of the flattened apical parts of the gonostyli strongly raised. It can be placed closest to A. villipes Perez , 1895 (Fig. 30 ) due to the antennae that are ventrally extensively lightened orange, the intermixed light and dark pubescence of the face, with pale hairs medially and dark hairs laterally, the intermixed light and dark pubescence of the scutum, with dark hairs medially and lighter hairs laterally, by the entirely yellow-marked male clypeus, and by the penis valves basally broad with lateral hyaline extensions, valves strongly narrowing apically. The immediate difference between the two taxa is size, with A. villipes averaging 12-13 mm in length in females and 11-12 mm in males, compared to 15-16 mm and 13-14 mm respectively in A. ghisbaini . Structurally, A. ghisbaini females can be separated by the bicoloured scopa, black dorsally and orange ventrally (Fig. 51D ; A. villipes with scopa unicolourous orange, Fig. 30B ), the longer face, clypeus clearly projecting ventrally well below a line drawn between the lower margins of the compound eyes ( A. villipes with face shorter, clypeus only slightly projecting below this line in direct comparison), the clypeus clearly punctured with a longitudinal impunctate midline that broadens apically ( A. villipes with clypeus obscurely and shallowly punctate, without obvious impunctate midline), and the reduced pale pubescence of the face, with light hairs restricted to the area around the antennal insertions ( A. villipes with extensive pale hairs covering majority of face and clypeus, with black hairs predominantly along the inner margins of the compound eyes). In the male sex, A. ghisbaini can be separated by the same clypeal punctation character (stronger in A. ghisbaini with clearer impunctate midline), but this is slightly more subtle than in the female sex. Direct comparison of the genital capsule shows that flattened apical part of the gonostyli are more strongly elongate and longer than broad, thus appearing triangular (Fig. 52D, E ; in A. villipes with the flattened apical part of the gonostyli more rounded, about as long as broad, Fig. 52F ), the inner margins of these parts more strongly and acutely raised, slightly reflexed (in A. villipes with the inner margin less strongly raised and not reflexed). Remarks. The two females from the Sierra de las Nieves (Fig. 50B ) were collected from Cistus albidus Linnaeus ( Cistaceae ). Like A. villipes (Table 1 ), this species is likely to be oligolectic on Cistaceae . Additional surveys are needed to clarify the limits of its range. Male specimens from Elvira in the Naturalis collection were incorrectly determined by Teunissen as A. maroccana Benoist, 1950 which is a synonym of A. leptopyga Perez , 1895. Table 1. Host plant use and dietary classification for selected Iberian Andrena species. n , total number of pollen loads; N , number of pollen loads from different localities. Plant taxa: ADO, Adoxaceae ; AMA, Amaryllidaceae ; API, Apiaceae ; ASP, Asparagaceae ; AST, Asteraceae ; BOR, Boraginaceae ; BRA, Brassicaceae ; CAM, Campanulaceae ; CAP, Caprifoliaceae ; CAR, Caryophyllaceae ; CIS, Cistaceae ; CRA, Crassulaceae ; EUP, Euphorbiaceae ; FAB, Fabaceae ; FAG, Fagaceae ; FRA, Frankeniaceae ; GER, Geraniaceae ; HYP, Hypericaceae ; PAP, Papaveraceae ; PLA, Plantaginaceae ; PLU, Plumbaginaceae ; RES, Resedaceae ; RHA, Rhamnaceae ; ROS, Rosaceae ; SAL, Salicaceae ; SAP, Sapindaceae ; SCR, Scrophulariaceae . Countries: BE, Belgium; BG, Bulgaria; DZ, Algeria; ESP, Spain; FRA, France; IL, Israel; IR, Iran; MA, Morocco; PT, Portugal; SY, Syria; TJ, Tajikistan; TN, Tunisia.

Species	n	N	Origin (and number) of pollen loads	Result of microscopic analysis of pollen grains (% of pollen grains)	Percentage of pure loads of preferred host	Percentage of loads with preferred host	Host range
Aciandrena Warncke
A. fulica Warncke	12	7	ESP (10), PT (2)	BRA 99.6, CIS 0.4	91.7	100.0	Broadly oligolectic ( Brassicaceae )
A. vacella Warncke	2	2	ESP (2)	BRA 100.0	100.0	100.0	Broadly oligolectic ( Brassicaceae )
Andrena aegyptiaca -group
A. alluaudi Benoist	4	3	MA (2), PT (2)	AST 100.0	100.0	100.0	Broadly oligolectic ( Asteraceae ; Cichorioideae )
Aenandrena Warncke
A. aeneiventris Morawitz	15	7	ESP (15)	API 100.0	100.0	100.0	Possibly broadly oligolectic ( Apiaceae )
A. hedikae Jaeger	22	10	ESP (11), MA (9), PT (2)	API 100.0	100.0	100.0	Possibly broadly oligolectic ( Apiaceae )
A. hystrix Schmiedeknecht	9	5	ESP (8), PT (1)	BRA 100.0	100.0	100.0	Broadly oligolectic ( Brassicaceae )
Avandrena Warncke
A. avara Warncke	2	2	ESP (2)	GER 100.0	100.0	100.0	Broadly oligolectic ( Geraniaceae )
A. melacana Warncke	6	2	ESP (6)	GER 100.0	100.0	100.0	Broadly oligolectic ( Geraniaceae )
A. panurgina De Steffani	11	5	ESP (4), FRA (4), PT (3)	GER 93.6, AST 5.0, BRA 1.4	81.8	100.0	Broadly oligolectic ( Geraniaceae )
Andrena Blandandrena subgen. nov.
A. blanda Perez	27	9	ESP (8), MA (19)	RES 100.0	100.0	100.0	Narrowly oligolectic ( Reseda , Resedaceae )
Brachyandrena Pittioni
A. colletiformis Morawitz	6	4	ESP (5), PT (1)	API 100.0	100.0	100.0	Possibly broadly oligolectic ( Apiaceae )
A. miegiella Dours	5	4	ESP (2), MA (2), TN (1)	API 99.8, AST 0.2	80.0	100.0	Possibly broadly oligolectic ( Apiaceae )
Chlorandrena Perez
A. abrupta Warncke	1	1	PT (1)	AST 100.0	100.0	100.0	Probably broadly oligolectic ( Asteraceae ; Asteroideae )
A. cinerea Brulle	22	15	ESP (6), FRA (3), PT (12), TN (1)	AST 100.0	100.0	100.0	Broadly oligolectic ( Asteraceae ; Cichorioideae )
A. curtivalvis Morice	1	1	ESP (1)	AST 100.0	100.0	100.0	Broadly oligolectic ( Asteraceae ; Cichorioideae )
A. elata Warncke	13	5	ESP (13)	AST 100.0	100.0	100.0	Broadly oligolectic ( Asteraceae ; Asteroideae )
A. leucolippa Perez	22	10	ESP (12), FRA (10)	AST 100.0	100.0	100.0	Broadly oligolectic ( Asteraceae ; Asteroideae )
A. rhenana Stoeckhert	8	4	ESP (2), PT (6)	AST 100.0	100.0	100.0	Broadly oligolectic ( Asteraceae ; Cichorioideae )
A. senecionis Perez	21	15	ESP (11), FRA (3), MA (3), PT (4)	AST 100.0	100.0	100.0	Broadly oligolectic ( Asteraceae ; Cichorioideae )
Chrysandrena Hedicke
A. fertoni Perez	4	3	ESP (4)	AST 100.0	100.0	100.0	Broadly oligolectic ( Asteraceae ; Cichorioideae )
Cordandrena Warncke
A. vaulogeri Perez	10	5	ESP (3), MA (7)	BRA 61.8, ROS 22.6, AST 9.6, FAB 6.0	30.0	80.0	Polylectic s. str.
Cryptandrena Pittioni
A. ventricosa Dours	39	10	ESP (29), FRA (10)	FAB 92.8, API 6.1, others 1.1	76.9	97.4	Polylectic with a strong preference ( Fabaceae )
Didonia Gribodo
A. mucida Kriechbaumer (1st generation)	3	3	ESP (1), PT (2)	ASP 100.0	100.0	100.0	Possibly narrowly oligolectic ( Muscari ; Asparagaceae )
A. mucida Kriechbaumer (2nd generation)	12	8	BG (2), ESP (9), MA (1)	CAP 100.0	100.0	100.0	Broadly oligolectic ( Caprifoliaceae )
Euandrena Perez
A. lavandulae Perez	5	5	ESP (3), FRA (1), PT (1)	FAB 27.6, CIS 24.3, SCR 20.7, PLA 10.3, AST 8.3, CAM 4.7, GER 2.9, CAR 1.2	20.0	20.0	Polylectic s. str.
Graecandrena Warncke
A. nebularia Warncke	5	3	ESP (1), MA (4)	BRA 100.0	100.0	100.0	Probably broadly oligolectic ( Brassicaceae )
A. verticalis Perez	30	21	ESP (20), MA (8), PT (2)	BRA 56.3, API 43.8	53.3	53.3	Mesolectic ( Apiaceae & Brassicaceae )
Andrena incisa -group
A. lateralis Morawitz	7	3	ESP (3), IR (1), TJ (3)	API 100.0	100.0	100.0	Broadly oligolectic ( Apiaceae )
Leucandrena Hedicke
A. leptopyga Perez	19	12	DZ (1), ESP (1), MA (12), PT (5)	RES 90.8, BRA 6.6, BOR 1.8, SCR 0.7	78.9	94.7	Polylectic with a strong preference ( Reseda , Resedaceae )
A. tunetana Schmiedeknecht	4	4	DZ (1), ESP (2), MA (1)	BRA 100.0	100.0	100.0	Probably broadly oligolectic ( Brassicaceae )
Melanapis Cameron
A. fuscosa Erichson	18	12	ESP (15), FRA (1), IL (1), PT (1)	BRA 52.4, API 15.5, PAP 12.7, AST 9.5, ROS 6.1, EUP 3.1, others 0.7	50.0	72.2	Polylectic s. str.
Melandrena Perez
A. albopunctata (Rossi)	15	7	ESP (14), MA (1)	AST 47.4, API 25.3, CAP 11.2, BRA 6.4, FAB 3.9, PAP 3.1, others 2.7	20.0	86.7	Polylectic s. str.
A. assimilis Radoszkowski	15	6	ESP (4), FRA (11)	AST 34.8, ROS 21.1, API 21.0, PLU 7.9, AMA 6.6, SAL 2.7, others 6.1	6.7	73.3	Polylectic s. str.
A. bicolorata (Rossi)	8	5	ESP (2), PT (6)	BRA 100.0	100.0	100.0	Broadly oligolectic ( Brassicaceae )
A. florentina Magretti	9	5	MA (1), PT (8)	BRA 100.0	100.0	100.0	Broadly oligolectic ( Brassicaceae )
A. morio Brulle (including A. hispania Warncke)	9	8	ESP (3), PT (6)	CIS 69.6, API 17.2, AST 10.0, others 3.2	22.2	77.8	Polylectic s. str.
Micrandrena Ashmead
A. ampla Warncke	21	10	ESP (10), FRA (10), PT (1)	API 100.0	100.0	100.0	Broadly oligolectic ( Apiaceae )
A. bayona Warncke	2	2	ESP (2)	API 50.0, BRA 50.0	50.0	50.0	Probably polylectic
A. djelfensis Perez	20	13	ESP (3), MA (7), PT (10)	CIS 99.9, FAB 0.1	95.0	100.0	Broadly oligolectic ( Cistaceae )
A. fabrella Perez	22	13	ESP (8), FRA (1), MA (5), PT (8)	CIST 99.9, AST 0.1	90.0	100.0	Broadly oligolectic ( Cistaceae )
A. icterina Warncke	9	6	ESP (9)	BRA 69.1, CIS 10.6, SAL 10.3, EUP 5.8, others 4.2	22.2	88.9	Polylectic s. str.
A. longibarbis Perez	13	8	ESP (2), MA (8), PT (3)	BRA 99.6, AST 0.4	92.3	100.0	Broadly oligolectic ( Brassicaceae )
A. nana (Kirby)	51	28	ESP (33), FRA (1), MA (8), PT (9)	API 71.2, BRA 28.6, EUP 0.2	68.6	74.5	Polylectic with a strong preference ( Apiaceae )
A. nitidula Perez	39	18	ESP (16), MA (21), PT (2)	BRA 100.0	100.0	100.0	Broadly oligolectic ( Brassicaceae )
A. omnilaevis Wood	6	5	ESP (2), PT (4)	CRA 100.0	100.0	100.0	Probably narrowly oligolectic ( Sedum , Crassulaceae )
A. orana Warncke	17	3	DZ (9), MA (5), PT (3)	BRA 100.0	100.0	100.0	Broadly oligolectic ( Brassicaceae )
A. pauxilla Stoeckhert	11	5	ESP (11)	CRA 82.1, BRA 17.9	81.8	81.8	Possibly polylectic with a strong preference ( Sedum , Crassulaceae )
A. spreta Perez	19	11	ESP (15), MA (3), PT (1)	BRA 93.1, AST 3.0, EUP 2.8, FAB 1.2	78.9	100.0	Polylectic with a strong preference ( Brassicaceae )
A. tenuistriata Perez	39	27	ESP (17), FRA (3), MA (6), PT (13)	BRA 99.8, others 0.2	94.9	100.0	Broadly oligolectic ( Brassicaceae )
Nobandrena Warncke
A. funerea Warncke	12	6	ESP (12)	BRA 100.0	100.0	100.0	Broadly oligolectic ( Brassicaceae )
Notandrena Perez
A. aerinifrons Dours	25	8	DZ (3), ESP (3), MA (11), PT (8)	BRA 100.0	100.0	100.0	Broadly oligolectic ( Brassicaceae )
A. bellidis Perez	3	3	ESP (2), PT (1)	AST 38.9, RES 36.4, RAN 18.2, BOR 6.5	0.0	66.7	Polylectic s. str.
A. juliana Wood	35	2	ESP (35)	API 82.6, FRA 13.1, CIS 2.8, others 1.4	74.3	82.9	Polylectic with a strong preference ( Apiaceae )
A. leucophaea Lepeletier	2	2	ESP (2)	AST 100.0	100.0	100.0	Possibly oligolectic ( Asteraceae ; Asteroideae )
A. nigroviridula Dours	9	8	ESP (4), MA (4), PT (1)	BRA 100.0	100.0	100.0	Broadly oligolectic ( Brassicaceae )
A. varuga Warncke	3	2	ESP (3)	BRA 100.0	100.0	100.0	Probably broadly oligolectic ( Brassicaceae )
Andrena numida -group
A. hypopolia Schmiedeknecht	8	7	ESP (4), FRA (1), PT (3)	BRA 64.8, API 34.6, AST 0.6	50.0	62.5	Mesolectic ( Apiaceae & Brassicaceae )
A. ranunculorum Morawitz	17	17	FRA (17)	BRA 75.8, API 10.2, ROS 5.7, FAG 5.6, AST 2.1, ADO 0.5	64.7	100.0	Polylectic with a strong preference ( Brassicaceae )
Orandrena Warncke
A. monilia Warncke	2	2	ESP (1), MA (1)	BRA 100.0	100.0	100.0	Probably broadly oligolectic ( Brassicaceae )
Andrena Ovandrena subgen. nov.
A. farinosa Perez	9	5	ESP (9)	FAB 100.0	100.0	100.0	Broadly oligolectic ( Fabaceae )
A. oviventris Perez	28	11	ESP (9), FRA (4), MA (12), PT (3)	RES 98.3, others 1.7	82.1	100.0	Narrowly oligolectic ( Reseda , Resedaceae )
Plastandrena Hedicke
A. asperrima Perez	56	29	ESP (8), FRA (5), MA (43)	BRA 77.6, RES 18.6, ROS 2.5, AST 1.3	69.6	87.5	Polylectic with a strong preference ( Brassicaceae )
A. pilipes Fabricius s. str.	28	21	ESP (15), FRA (8), PT (5)	BRA 54.4, ROS 18.9, AST 13.8, CIS 6.6, API 5.7, others 0.7	35.7	57.1	Polylectic s. str.
Andrena relata -group
A. corax Warncke	10	4	ESP (8), PT (2)	RES 99.7, AST 0.3	90.0	100.0	Narrowly oligolectic ( Reseda , Resedaceae )
A. laurivora Warncke	3	1	MA (1)	RES 100.0	100.0	100.0	Probably narrowly oligolectic ( Reseda , Resedaceae )
A. relata Warncke	2	2	ESP (2)	RES 100.0	100.0	100.0	Probably narrowly oligolectic ( Reseda , Resedaceae )
Rufandrena Warncke
A. orbitalis Morawitz	9	6	ESP (2), FRA (4), PT (3)	PLA 100.0	100.0	100.0	Narrowly oligolectic ( Plantago , Plantaginaceae )
A. rufiventris Lepeletier	3	1	MA (3)	PLA 100.0	100.0	100.0	Narrowly oligolectic ( Plantago , Plantaginaceae )
Simandrena Perez
A. antigana Perez	25	14	ESP (6), MA (7), PT (12)	BRA 99.8, others 0.2	96.0	100.0	Broadly oligolectic ( Brassicaceae )
A. cilissaeformis Perez	5	5	ESP (2), MA (3)	BRA 83.6, EUP 8.8, RHA 7.6	60.0	80.0	Probably polylectic with a strong preference ( Brassicaceae )
A. propinqua Schenck	43	31	BE (4), ESP (21), FRA (5), MA (3), PT (10)	BRA 46.1, ROS 25.8, FAB 10.2, CIS 4.4, CRA 3.2, BOR 2.5, others 7.7	30.2	58.1	Polylectic s. str.
A. rhypara Perez	4	3	MA (4)	RES 100.0	100.0	100.0	Possibly narrowly oligolectic ( Reseda ; Resedaceae )
A. vetula Lepeletier	31	16	ESP (20), FRA (2), MA (7), SY (1), TN (1)	BRA 99.8, others 0.2	93.5	100.0	Broadly oligolectic ( Brassicaceae )
Truncandrena Warncke
A. doursana Dufour	8	3	MA (7), PT (1)	BRA 100.0	100.0	100.0	Broadly oligolectic ( Brassicaceae )
A. ferrugineicrus Dours	28	18	DZ (1), ESP (16), MA (2), PT (9)	BRA 100.0	100.0	100.0	Broadly oligolectic ( Brassicaceae )
A. nigropilosa Warncke	23	8	ESP (16), FRA (5), MA (2)	BRA 100.0	100.0	100.0	Broadly oligolectic ( Brassicaceae )
A. villipes Perez	6	2	FRA (1), PT (5)	CIS 100.0	100.0	100.0	Probably broadly oligolectic ( Cistaceae )

Etymology. Dedicated to my friend and colleague Guillaume Ghisbain (Mons, Belgium) who accompanied me during fieldwork in Malaga province, and who is an accomplished hymenopterist in his own right. Distribution. Spain ( Malaga province). Description of missing sexes Andrena (Micrandrena) alma Warncke, 1975 Description. Male. Body length 6.5-7 mm (Fig. 53A ). Head : Dark, 1.2 times wider than long. Clypeus flattened, unevenly punctate with large punctures, punctures separated by 0.5-2 puncture diameters, underlying surface shagreened in basal half, polished and shiny in apical half. Process of labrum trapezoidal, slightly wider than long, apical margin slightly thickened. Gena 1.3 times width of compound eye (Fig. 53B, C ); ocelloccipital distance 0.5 times diameter of lateral ocellus. Face medially with whitish hairs, scape with mixture of black and white hairs, inner margin of compound eyes with black hairs, gena ventrally with white hairs, becoming black dorsolaterally, vertex and frons with mixture of black and pale hairs, none exceeding length of scape. Antennae dark, A3 exceeding length of A4, shorter than A4+5. Mesosoma : Scutum and scutellum finely granularly shagreened and weakly shiny, shallowly and obscurely punctate, punctures separated by 2-3 puncture diameters. Pronotum with clear humeral angle. Mesepisternum and dorsolateral surfaces of propodeum with fine granular microreticulation, with regular slightly raised hair bearing punctures. Propodeal triangle with regular granular shagreen, basally and medially with obscure and finely raised rugosity, propodeal triangle thus defined by change in surface sculpture compared to dorsolateral parts of propodeum (Fig. 53D ). Mesosoma laterally with long white pubescence, exceeding length of scape, pubescence becoming brownish and shorter dorsally, not exceeding length of scape. Legs dark, pubescence whitish. Hind tarsal claws with inner tooth. Wings hyaline, venation and stigma dark brown, nervulus interstitial. Metasoma : Terga dark, marginal areas obscurely lightened dark hyaline brown apically (Fig. 53E ). Tergal discs with obscure and weak punctures that disappear into even underlying microreticulation, surface weakly shiny. Tergal discs with sparse long light brown hairs, T2-4 laterally with weak, widely interrupted apical hair fringes. S8 columnar, slightly broadened apically, ventrally covered with short yellowish hairs. Genital capsule compact, gonocoxae apically weakly produced into short rounded teeth; gonostyli parallel-sided, spatulate; penis valves slightly broadened basally (Fig. 53F ). Figure 53. Andrena (Micrandrena) alma Warncke, 1975 male A profile B head, dorsal view C head, profile view D propodeal triangle, dorsal view E terga, dorsal view F genital capsule, dorsal view. Diagnosis. Andrena alma can be recognised due to its combination of small body size, dark integument, pronotum with humeral angle, evenly shagreened and weakly shiny terga, gena exceeding the width of the compound eye (Figs 53B, C ), flattened and striation-free clypeus, simple genital capsule (Fig. 53F ), and smooth and granulate propodeal triangle with obscure and finely raised rugosity basally (Fig. 53D ). This smooth propodeal triangle places it close to former Distandrena species, but the flattened and striation-free clypeus excludes A. longibarbis Perez , 1895 (clypeus domed, obscurely striate) and A. orana Warncke, 1974 (clypeus domed and striate). The evenly shagreened terga and obscure basal rugosity on the largely granularly shagreened propodeal triangle place it superficially close to A. djelfensis , but this species can easily be separated by the distinctive genital capsule with elongate and strongly medially bent gonostyli, whereas the genital capsule is simple and lacking distinctive features in A. alma . Finally, A. alma has a distinctly broadened gena that slightly but distinctly wider than the width of the compound eye, an unusual character in Micrandrena . This allows separation from A. abjecta which has the gena equalling the width of the compound eye. Collectively, these characters make recognition of A. alma straightforward, though as the males of A. tenostra and A. aff mica are unknown, diagnosis may become more challenging in the future. A level of caution should therefore be applied when identifying material from the extreme south and south-east of Spain. Distribution. Central and southern Portugal and Spain. Material examined. Portugal : Algarve, Monte Gordo, Retur, Praia do Cabeco , 29.iii.2022, 1♂, leg. T.J. Wood, TJWC; Algarve, Tavira, Cacela Velha, 28.iii.2022, 1♂, leg. T.J. Wood, TJWC; Spain : Almodovar del Campo (Ciudad Real), 700 m, 24.iii.2005, 1♂, leg. F.J. Ortiz-Sanchez , FJOS; Santa Ana la Real, Sierra Aracena (Huelva), 630 m, 13.iv.2006, 2♂, leg. F.J. Ortiz-Sanchez , FJOS; El Hongo (P.N. Donana ), 30.iii.2018, 1♂, leg. F. Molina, EBDC. Andrena (? Euandrena ) Andrena ramosa Wood, 2022 Description. Male. Body length 8-10 mm (Fig. 54A ). Head : Dark, 1.05 times wider than long (Fig. 54C ). Clypeus long, weakly domed, unevenly punctate, punctures separated by 0.5-2 puncture diameters, underlying surface strongly shagreened to microreticulate in basal half, becoming smooth and shiny in apical half. Process of labrum trapezoidal, 3 times wider than long, ventral surface smooth and polished. Gena equalling width of compound eye; ocelloccipital distance 1.5 times diameter of lateral ocellus. Face medially and gena ventrally with long yellowish hairs, face laterally, frons, and scape with long black hairs, mixing medially on face with yellowish hairs, longest exceeding length of scape. Antennae dark, A3 exceeding length of A4, shorter than A4+5, A4 slightly longer than wide, A5-13 elongate, clearly longer than wide. Mesosoma : Scutum and scutellum obscurely punctate, punctures separated by 1-2 puncture diameters, disappearing into extremely strong underlying granular microreticulation, surface dull (Fig. 54D ). Pronotum rounded. Mesepisternum and dorsolateral surface of propodeum with fine granular microreticulation, with finely raised network of reticulation that gives impression of large shallow punctures. Propodeal triangle narrow, surface with fine granular shagreen, basally and medially with finely raised rugosity, propodeal triangle thus defined by change in surface sculpture compared to dorsolateral parts of propodeum. Mesosoma with long, densely branches and plumose yellowish hairs clearly exceeding length of scape, black plumose hairs intermixed around wing bases, on scutum, and propodeum. Legs dark, pubescence brownish to black. Hind tarsal claws with inner tooth. Wings hyaline, stigma dark brown, venation dark to light brown, nervulus weakly antefurcal. Metasoma : Terga dark, apical rim of marginal areas very narrowly lightened hyaline brown (Fig. 54E ). Tergal discs with obscure hair-bearing punctures, disappearing into background sculpture, becoming more strongly defined laterally, underlying surface shagreened and weakly shiny. T1-3 with long but loose plumose yellowish-brown hairs, these becoming black on T4-5. S8 relatively short, rectangular, apically truncate, ventrally covered with dark brown hairs. Genital capsule moderately elongate, gonocoxae apically weakly produced into short rounded projections, gonostyli more or less parallel-sided, spatulate (Fig. 54F ). Penis valves occupying 1/2 space between gonostyli, slightly narrowing towards their apexes. Figure 54. Andrena (? Euandrena ) Andrena ramosa Wood, 2022 male A profile B head, profile view C head, frontal view D scutum, dorsal view E terga, dorsal view F genital capsule. Diagnosis. The male of A. ramosa is morphologically most similar to A. (Euandrena) solenopalpa due to the long head (only marginally wider than long) and clypeus that is shiny at least in its apical half. The two species are easily separated by the mouthparts, as in A. ramosa the mouthparts that protrude in front of the head are at most as long as the head (viewed frontally or laterally, Fig. 54B ), whereas in A. solenopalpa the mouthparts are extremely long, the labial palpi alone exceed the length of their head, the part of the mouthparts protruding in front of the head therefore collectively greatly exceed the length of the head. The clypeus of A. solenopalpa is also more extensively shiny, the gonostyli have their outer margin emarginate and are apically produced into narrow points, and A3 slightly exceeds the length of A4+5, whereas in A. ramosa the clypeus is shiny only in its apical half, the gonostyli are spatulate and apically truncate, and A3 only slightly exceeds A4, and is shorter than A4+5. The two species do not occur in sympatry, with A. ramosa restricted to south-western Spain, whereas A. solenopalpa is found in central and eastern Spain to southern France. Distribution. South-western Spain ( Cadiz , Sevilla). Remarks. The phylogenetic placement of A. ramosa remains somewhat obscure even following the discovery of the male sex and generation of a barcode sequence. A 658-bp fragment was generated from the female type specimen [BOLD accession number: IBIHM524-21], but this did not fall unambiguously close to any species or subgenus. The most similar sequences belonged to the subgenus Andrena Euandrena , specifically to A. symphyti (90.26%), A. montana Warncke, 1973 (90.31%), A. fulvida Schenck, 1853 (89.98%), and A. rufula Schmiedeknecht, 1883 (89.84%). Morphologically, A. ramosa does not fall nicely into Euandrena , as the female sex has foveae which are narrow but which do not narrow ventrally. However, Euandrena are part of the most highly derived clade of Andrena ( Pisanty et al. 2022b ), and subgeneric classification in this clade has been highly problematic due to the lack of clear delineating characters. Andrena ramosa does not belong to Margandrena Warncke, 1968 due to the lack of a strong humeral angle on the pronotum. It does not belong to the crocusella -group due to the lack of a humeral angle and the lack of lateral projections on the male penis valves (in addition to the lack of barcode similarity), and whilst it has strongly plumose pollen collecting hairs comprising the propodeal corbiculae and femoral scopae, those of the tibial scopae are simple, and the foveae do not narrow below which makes placement in Chrysandrena Hedicke, 1933 difficult. Andrena ramosa is therefore best considered to be affiliated with the subgenus Andrena Euandrena , pending investigation with more powerful genetic techniques. It clearly possesses a unique and unusual morphology within the West Palaearctic Andrena fauna. Examination of additional material from the province of Cadiz has shown that A. ramosa is commonly encountered in the Parque Natural Los Alcornocales area. Here it can be encountered between January and March, and is most frequently observed on Erica ( Ericaceae ; Perez Gomez in litt.). However, the pollen host is still obscure, since none of these bees have been observed collecting pollen. Moreover, Ericaceae pollen is small, with the grains typically having a diameter of 25 μm . The widely spaced and strongly branched and plumose pollen collecting hairs of A. ramosa (described and illustrated by Wood et al. 2022a ) would not seem to be an adaptation to the collection of small Ericaceae pollen grains, and indeed this adaptation is absent in the Ericaceae specialist A. (Cnemidandrena) fuscipes (Kirby, 1802) which has simple pollen collecting hairs. Further study is required; the assumption remains that A. ramosa collects pollen from a plant species with large pollen grains, thus necessitating these branched and plumose hairs. Material examined. Spain : Carretera Marrufo, Herriza ( Cadiz ; 3 km E Puerto de Galiz ), 11.xi.2020, 1♀, leg. A . Perez Gomez , APGC; Sevilla, Los Pinares de Aznalcazar [ 37.2782°N , - 6.2356°E ], 10.iii.2020, 1♀, leg. F. Molina, OOELM (holotype); Cadiz , Sierra de Montecoche, 31.i.2022, 4♂, 1♀, leg. A . Perez Gomez , APGC/TJWC; 18.i.2021, 1♂, leg. A . Perez Gomez , APGC; Cadiz , Pico del Montero, 2.ii.2022, 3♂, 1♀, leg. A . Perez Gomez , APGC/TJWC; Cadiz , Sierra de Fates, 21.iii.2022, 1♀, leg. A . Perez Gomez , APGC; Cadiz , Pico del Montero, Alcala de los Gazules, 26.iii.2022, 1♂, leg. A . Perez Gomez , APGC. Additional designation of lectotypes Andrena (Aenandrena) hystrix Schmiedeknecht, 1883 Andrena (Aenandrena) hystrix Schmiedeknecht, 1883: 618, ♀ [France, lectotype by present designation: RMNH]. Remarks. Schmiedeknecht (1883) described several Andrena species from material that had been sent to him by Perez . In each case, he indicated this in his title, e.g. ' Andrena hystrix Perez in litt' [sic]. For several of these species, the location of type material has been unclear, as they mostly did not seem to be in the MNHN in the Perez collection (see Le Divelec 2021 ), or elsewhere (see Gusenleitner and Schwarz 2002 ). Searches in the RMNH unexpectedly uncovered specimens of four species described by Schmiedeknecht with labels written in Perez's distinctive handwriting. The exact providence of these specimens is unclear, but the RMNH collection is known to contain material from Schmiedeknecht's collection (F. Bakker, pers. comm.). A lectotype was designated for one of these species ( A. (Andrena) mitis Schmiedeknecht, 1883) in a previous publication ( Wood 2023a ); the others are designated here. Schmiedeknecht (1883) described A. hystrix from female specimens from Hungary ( Hungaria ) and southern France ( Gallia meridionali ). He did not specify a type, though Gusenleitner and Schwarz (2002) list south France as the locus typicus . A specimen of A. hystrix from Marseille is labelled with Perez's handwriting, and this is considered to be part of the original syntypic series; it is here designated as a lectotype (Fig. 55A, B ). No specimens from Hungary s.l. could be found. Figure 55. Andrena (Aenandrena) hystrix Schmiedeknecht, 1883, female lectotype A label details B profile; Andrena (Notandrena) ranunculi Schmiedeknecht, 1883, female lectotype C label details D profile; Andrena (Euandrena) symphyti Schmiedeknecht, 1883, female lectotype E label details F profile. Material examined. France : Marseille [ 43.3612°N , 5.3942°E ], 1♀, RMNH (lectotype by present designation; Fig. 55A, B ). Andrena (Notandrena) ranunculi Schmiedeknecht, 1883 Andrena (Notandrena) ranunculi Schmiedeknecht, 1883: 617, ♀♂ [France, lectotype by present designation: RMNH]. Remarks. Schmiedeknecht (1883) described A. ranunculi , comparing it to A. ranunculorum . He gave the habitat as Russia (referring to A. ranunculorum ) and southern France (referring to the specimens received from Perez ). A series of males and females labelled by Perez as being from Bordeau [sic, = Bordeaux] were found in the RMNH. These conform to Schmiedeknecht's description, and the concept used by subsequent authors. A female is here designated as a lectotype (Fig. 55C, D ). Material examined. France : Bordeaux [ 44.8352°N , - 0.5888°E ], 1♀, RMNH (lectotype by present designation; Fig. 55C, D ); Bordeaux, 8♂, 5♀, RMNH. Andrena (Euandrena) symphyti Schmiedeknecht, 1883 Andrena (Euandrena) symphyti Schmiedeknecht, 1883: 583, ♀♂ [France, lectotype by present designation: RMNH]. Remarks. As for the previous two species, material labelled by Perez was found in the RMNH collection. Two females and one male labelled as being from Bordeau [sic, = Bordeaux]. Schmiedeknecht (1883) specifically states that the type material comes from Bordeaux, writing " In Gallia prope Bordeaux a Dom. Illustrissimo Perez detecta ". A female is here designated as a lectotype (Fig. 55E, F ). Material examined. France : Bordeaux [ 44.8352°N , - 0.5888°E ], 1♀, RMNH (lectotype by present designation; Fig. 55E, F ); Bordeaux, 1♂, 1♀, RMNH. Designation of neotypes Andrena (Chlorandrena) boyerella Dours, 1872 Andrena (Chlorandrena) distincta Lucas, 1849 nec. Smith, 1847 [Algeria: MNHN, not examined]. Andrena (Chlorandrena) boyerella Dours, 1872: 429, ♀♂ [Morocco: OOELM ]. Neotype. Morocco : Fes-Meknes , Azrou, 4 km SWW of Bakrit, Cascades Bakrit, 33.0466°N , - 5.2681°E , 1650 m, 17.v.2022, 1♂, leg. T.J. Wood, OOELM [BOLD accession number WPATW495-22] (Fig. 56 ). Figure 56. Andrena (Chlorandrena) boyerella Dours, 1872, male neotype A profile B face, frontal view C terga, dorsal view D genital capsule dorsal view. Remarks. As discussed above, Dours (1872) described A. boyerella from southern France and Algeria. No material is available for study, as all of Dours' types were destroyed in a fire. Given that two taxa are present in these regions, in order to fix the name A. boyerella on the North African population, a barcoded neotype is designated from Moroccan material (Fig. 56 ). Distribution. Morocco, Algeria, Tunisia, Italy (Sicily). Andrena (Notandrena) griseobalteata Dours, 1872 Andrena (Notandrena) erythrocnemis auctorum. nec. Morawitz, 1871. Andrena (Notandrena) griseobalteata Dours, 1872: 427, ♀ [France: RMNH]. Neotype. France : Pyrenees-Atlantiques , Berenx [ 43.4994°N , - 0.8575°W ], 6.vi.1987, 1♀, leg. E. A. M. Speijer, RMNH (Fig. 57 ). Figure 57. Andrena (Notandrena) griseobalteata Dours, 1872, female neotype A profile B face, frontal view C scutum, dorsal view D terga, dorsal view. Remarks. The correct name to apply to this distinctive taxon has been confused for many years. Through the combination of its large size (for a Notandrena ) and densely punctate scutum it is comparable only to A. ungeri Mavromoustakis, 1952. The name A. erythrocnemis Morawitz, 1871 was used by many authors to refer to this taxon (e.g. Warncke 1967 ), but the lectotype of A. erythrocnemis is actually A. chrysosceles (see Proshchalykin et al. 2017 ; Astafurova et al. 2021 ). Gusenleitner and Schwarz (2002) resolved this issue by resurrecting A. griseobalteata to species status. Finally, Wood and Monfared (2022) removed A. emesiana Perez , 1911 (southern Turkey, Syria, Iran) from synonymy with this taxon. Although Gusenleitner and Schwarz (2002) would seem to have resolved the issue, there is no type specimen for A. griseobalteata due to the loss of Dours' collection. In the original description, Dours (1872 : 428) listed the species from Saint-Sever in the department of Landes in south-western France, but also from Algeria. This is peculiar, because A. griseobalteata is not known from North Africa. The original description also does not allow for completely unambiguous recognition of the species, though the dense punctation of the scutum is mentioned. In the interests of nomenclatural stability, a neotype from Berenx in south-western France (36 km from Saint-Sever) is designated in order to fix the species concept for the future (Fig. 57 ). Finally, though listed from Spain by Ortiz-Sanchez (2011 , as A. griseobalteata ; 2020, as A. erythrocnemis ), the presence of this taxon in Spain is somewhat doubtful. The distribution maps of Warncke ( Gusenleitner and Schwarz 2002 ) show dots around south-western France into the Pyrenees, but there are no unambiguous dots for Spain. However, I have been able to examine one specimen of A. griseobalteata from Spain, from the Sistema Central. The species is also expected to occur in the western Pyrenees; more recent material should be found to establish whether this taxon remains present in Iberia. Material examined. Spain : Sierra de Gredos, 12 km SSW Hoyos del Espino, 1950-2100 m, 4.vii.1972, 1♀, leg. J.A.W. Lucas, RMNH. Distribution. Spain, France, Italy, Croatia, Hungary, Albania, Romania, North Macedonia, Bulgaria, Greece, Turkey (western and northern Turkey; Gusenleitner and Schwarz 2002 ). The species is not considered to be present in North Africa, or in the Levant, as it is not found in very dry environments. Andrena (Taeniandrena) poupillieri Dours, 1872 Andrena (Taeniandrena) poupillieri Dours, 1872: 430, ♀ [Algeria: OOELM ]. Andrena (Taeniandrena) poupillieri incana Warncke, 1975a: 310, ♀♂ [Spain, Mallorca: OOELM , examined]. Neotype. Algeria : Tizi-Ouzou, Tigzirt, 36.8877°N , 4.1140°E , 6 m, 31.iii.2017, 1♀, leg. H. Ikhlef, OOELM [BOLD accession number HYMAA322-22] (Fig. 58 ). Figure 58. Andrena (Taeniandrena) poupillieri Dours, 1872, female neotype A profile B face, frontal view C dorsal view D terga, dorsal view. Remarks. This is the taxon referred to as ' A. poupillieri 2' by Praz et al. (2022) . The taxon A. poupillieri incana Warncke, 1975 which was described from and restricted to the Balearic Islands is considered a simple synonym of A. poupillieri due to the lack of genetic differentiation observed (see above), even though the tergal punctation is slightly reduced compared to the nominate taxon. The specimen used in the analysis of Praz et al. (2022) is designated as a lectotype (Fig. 58 ). Distribution. Morocco, Algeria, Tunisia, Spain (mainland and Balearic Islands). Records ( Gusenleitner and Schwarz 2002 ) from Libya must be confirmed, though they probably do refer to true A. poupillieri . Records from Crete probably refer to unrecognised A. ovata specimens, and so A. poupillieri is not considered to be present there until definitive material is located. Andrena (Pruinosandrena) succinea Dours, 1872 Andrena succinea Dours, 1872: 424, ♀ [Morocco: OOELM ]. Neotype. Morocco : Oriental, Guercif, P5427, 2 km SW of Bou Rached, 33.8844°N , - 3.6154°W , 950 m, 13.v.2022, 1♀, leg. T.J. Wood, OOELM [BOLD accession number WPATW389-22] (Fig. 59 ). Remarks. As discussed above, it is preferable to designate a neotype for A. succinea in order to maintain nomenclatural stability. The barcoded specimen pictured in Fig. 34B is selected as a neotype (Fig. 59 ) in order to fix the species concept for the future. Figure 59. Andrena (Pruinosandrena) succinea Dours, 1872, female neotype A profile B face, frontal view C mesosoma, dorsolateral view D terga, dorsal view. Distribution. Morocco, Algeria, Tunisia, Libya, Egypt, Israel and the West Bank, Jordan, Syria, Saudi Arabia, Iran ( Wood and Monfared 2022 ). Andrena (incertae sedis) Andrena numida Lepeletier, 1841 Andrena numida Lepeletier, 1841: 252, ♀ [Morocco: OOELM ]. Neotype. Morocco : Fes-Meknes , Azrou, P7311, 10 km S of Ain Leuh, 1750 m, 33.2220°N , - 5.3411°W , 18.v.2022, 1♀, leg. T.J. Wood, OOELM [BOLD accession number WPATW484-22] (Fig. 60 ). Figure 60. Andrena (incertae sedis) Andrena numida Lepeletier, 1841, female neotype A profile B face, frontal view C dorsal view D T2-5, dorsal view detail. Remarks. As discussed above, it is beneficial to designate a neotype for A. numida since the original type series cannot be located in the MNHN, and so that the name and genetic identity of North African populations can be fixed. Distribution. Morocco, Algeria, Tunisia, Libya, Italy (Sicily, Calabria, Campania).