JHR 96: 241-484 (2023) igo.) JOURNAL OF. Ah entensescieat doi: 10.3897/jhr.96. 101873 MONOGRAPH () Hymenopter a https://jhr.pensoft.net Thelnternaonl Scie of Hymenoptxriss, RESEARCH The genus Andrena Fabricius, |775 in the Iberian Peninsula (Hymenoptera, Andrenidae) Thomas J. Wood! | University of Mons, Research Institute for Biosciences, Laboratory of Zoology, Place du Parc 20, 7000, Mons, Belgium Corresponding author: Thomas J. Wood (thomasjames.wood@umons.ac.be) Academic editor: Jack Neff | Received 13 February 2023 | Accepted 3 May 2023 | Published 22 May 2023 https.//zoobank.org/1 5A2B06B-92F3-4E70-AC8F-6FEABF365E71 Citation: Wood TJ (2023) The genus Andrena Fabricius, 1775 in the Iberian Peninsula (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 96: 241-484. https://doi.org/10.3897/jhr.96.101873 Abstract The Iberian Peninsula is a global hotspot for bee diversity due to its large number of different habitats, par- ticularly Mediterranean scrubland, mountains, and hot and cold steppe. In line with its status as a hotspot of bee diversity, the peninsula hosts a very large Andrena fauna, which despite progress in recent years re- mains incompletely studied, particularly with reference to genetic investigation. Here the Iberian Andrena fauna is comprehensively revised, resulting in a total of 228 recorded species. Numerous taxonomic changes are necessary following inspection of museum specimens, type material, and genetic investiga- tion. The following subgenera are described: Pruinosandrena subgen. nov., containing six taxa previously placed in the subgenus Campylogaster Dours, 1873, and Blandandrena subgen. nov., Bryandrena subgen. nov., Limbandrena subgen. nov., and Ovandrena subgen. nov., containing one, one, one, and four taxa previously placed in the subgenus Poliandrena Warncke, 1968. Andrena (Limbandrena) toelgiana Friese, 1921 syn. nov. is synonymised with A. (Limbandrena) limbata Eversmann, 1852. The current lectotype of A. (Micrandrena) obsoleta Pérez, 1895 was incorrectly designated by Warncke; the taxon differs from A. obsoleta sensu Warncke, belonging instead to a taxon within the A. mariana Warncke, 1968 complex. A new lectotype is designated for A. obsoleta sp. resurr. from Algeria, and A. mariana solda Warncke, 1974 syn. nov. is synonymised with it; A. (Micrandrena) alma Warncke, 1975 stat. nov., A. (Micrandrena) mica Warncke, 1974 stat. nov., and A. (Micrandrena) tenostra Warncke, 1975 stat. nov. are raised to species status. Andrena (Truncandrena) abunda Warncke, 1974 stat. nov., A. (Micrandrena) lecana Warncke, 1975 stat. nov., A. (Pruinosandrena) parata Warncke, 1967 stat. nov., A. (Micrandrena) pauxilla Stockhert, 1935 sp. resurr., A. (Pruinosandrena) succinea Dours, 1872 sp. resurr., and A. (Notandrena) varuga Warncke, 1975 stat. nov. are also returned or elevated to species status. A lectotype is designated for A. (Euandrena) lavandulae Pérez, 1902 sp. resurr. which is returned to species status, and A. (Euandrena) impressa Copyright Thomas J.Wood. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 242 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Warncke, 1967 syn. nov. is synonymised with it. Andrena (Truncandrena) nigropilosa Warncke, 1967 stat. nov. is elevated to species status, and A. (Truncandrena) truncatilabris espanola Warncke, 1967 syn. nov. is synonymised with it as a junior subjective synonym. A lectotype is designated for A. (Melandrena) vachali Pérez, 1895; A. (Melandrena) creberrima Pérez, 1895 syn. nov. and A. (Melandrena) vachali syn. nov. are synonymised with A. (Melandrena) discors Erichson, 1841, and Andrena (Melandrena) hispania Warncke, 1967 syn. nov. is synonymised with A. (Melandrena) morio Brullé, 1832. Andrena (Pruinosandrena) mayeti Pérez, 1895 syn. nov. is newly synonymised with A. (Pruinosandrena) caroli Pérez, 1895 and A. (incertae sedis) setosa Pérez, 1903 syn. nov. is newly synonymised with A. (incertae sedis) nranunculorum Morawitz, 1877. Andrena (Simandrena) cilissaeformis Pérez, 1895 sp. resurr. is returned to species status, and is the correct name for A. (Simandrena) breviscopa auctorum. Andrena (incertae sedis) breviscopa Pérez, 1895 is returned to synonymy with A. (incertae sedis) numida Lepeletier, 1841, and A. (incertae sedis) inconspicua Morawitz, 1871 is newly synonymised syn. nov. with A. numida. Andrena (Euandrena) isolata sp. nov. and A. (Micrandrena) ortizi sp. nov. are described from the Sierra Nevada (Granada), A. (Zruncandrena) ghisbaini sp. nov. is described from Malaga province, and A. (Avandrena) juliae sp. nov. is described from Cadiz province. The males of A. (Micrandrena) alma and A. (?Euandrena) ramosa Wood, 2022 are described. Additional lectotypes are designated for A. (Plastandrena) asperrima Pérez, 1895, A. (Plastandrena) atrica- pilla Pérez, 1895, A. (Aenandrena) hystrix Schmiedeknecht, 1883, A. (Pruinosandrena) lanuginosa Spinola, 1843, A. (Notandrena) ranunculi Schmiedeknecht, 1883, and A. (Euandrena) symphyti Schmiedeknecht, 1883. Neotypes are designated for A. (Chlorandrena) boyerella Dours, 1872, A. (Notandrena) griseobalteata Dours, 1872, A. (Taeniandrena) poupillieri Dours, 1872, A. (Pruinosandrena) succinea Doutrs, 1872, and A. (incertae sedis) numida Lepeletier, 1841. Type photographs and diagnostic characters are presented in each case, as well as new dietary information for understudied species. Finally, an identification key is presented in order to facilitate future research on this hyper-diverse genus in one of their global diversity hotspots, and current and future research perspectives for Iberian Andrena are discussed. Keywords Cryptic species, DNA barcoding, Iberian endemic species, solitary bees, taxonomy Table of contents Pratroe ere iia sf) 28 Ache t LEE he Mee PORTE sl ela te tet Rael Mee! Aleks Me BAS oe Mads 244 MCE OCOLO OY tachabenoerenttirabesin: Nairmnd it dibenmmiet bnicminremdnbecme sy daliceecmrednles 245 SPECIES CONCEP ES a A ic Fie Sh Ae A la eR AS eer he Rela. nce 245 Geneticsanpliie atid Atialyeisa. A550 0eh Adder Bs. ted ese nbaas top haa nese dodd 246 iChiceklistiamdvid entileatlon, 7.2.0 ordet. eed apace alt tea baet eds deere el 247 Identification-key- and! seogtapliic SCOpe:swiesecvectecsteduewewseoneeiteancvcrede dete neuncaniets 247 TPIS CTO TAA SoM Fs cloaks tau cant eect eMh tod Meroe ial lubes Moet nosis nat uA le titak ebbede 248 Dieraryeniches Otel Dera Andienaspeelese t.5.. Ri acents A ee Scart ad 249 Morpholodical term tology ..s0te.8. 5c oot cs Ssoe. ki et areecee lp agence deans 249 OLECHO MSS tM eG teeta ad inte ME vacua eee eh Ae Benard Nae MI boars ae 249 RUA Rast «iets aAhanlOh ale it Rt Veh KAR O25 oe, ns A nc ee in ae A 250 Cleiietio Pestilts-an d CaxOnOmiGre Aan Ges neasetiesuiseoa pebeens teu pernteeentan dein taesneorueceneee 250 Subgenus Acizndneng Warnecke 1968. css. trtstecsstst co tiapedeanctentsdeoeata te aneseasee 250 The genus Andrena in the Iberian Peninsula 243 SubpeniswA ciara rea Warnecke; VIGS A irate. acetates: sed hevens cons aecndvtaoncaetaehes 250 SUDPEMUS@ lorena icra PETEZ SIO): Nawecss es secemea nuns auiiealg abut anedieaneenien cst D5? Sub eens: dont a GribOd Oy 1894. x cscienssncestne vss recqusteneecesactivcngietelucser boca. 2D) SubeenusEuai7a rend Tledicke, 193 3% deren ticsagtaantieretlseleoscadacstupboneds pladben rsoets 257 Subgenus:Graecundrenad Warncke; WI GSS. i. sasraacvivesansivs codon uavtstinpesbGaeeonghheee 261 Subgents Melanapes Gamerons FOOD Aiieasn coat canes tinoriads esl sadnnsacunnctabsosedisbaal 262 Suber VMelanavendPEres,: LODO tap srncr ste we nib nd one oth lalclOa de dS Ponce Soa 263 Subgenus Micranarena ASWMiead, VSI a. avd cs ncrventeses one ecssssedpisonincnessrsttseettns RD SUDRENs IND NareNA VErer, GE SIO: sccravconctovesan sop see saulecronvedeadasec is mabocess escent 290 Subgenus Pasadena Vedicke, 1933+ sz.victsuoctene tacts tvaseanaecsiguectigrteugine detec, 292 Subgenus. oecianarena ledicke, VIGS i... 2. bitstachse West. wdkeo seine yuneabedaeuie oak 296 SUDPEMUS SA mAanAreNa Ere, NS IO etic remiteanh decartieeeniinhmasp MELE Cio Esk DOF Subgenus dzeniindrena Fledicke, 933+... cies. acatedetae esc dedvhaws duipdudesonnsbsties 299 Subgenuispellier 2. — Tebessa, avril, 2. - Biskra, mars, 39, rares. — Bone, 2..— Marseille, &. - Bone, 27liv BY. in cop. Souci et autres [illegible — presumably a type of flower]”. Babor is not mentioned, and so Warncke’s lectotype designation is rejected. Examination of specimens labelled as obsoleta by Pé- rez from the localities indicated shows that they belong to a different taxon to that of Warncke’s false lectotype. Warncke’s false lectotype is a female Distandrena (=Micran- drena) with a clearly striate clypeus, and T1 is polished. This conforms to the concept of A. (Micrandrena) nitidula Pérez, 1903, and indeed Warncke (1967) synonymised A. nitidula with A. obsoleta. Only female specimens of A. obsoleta from Béne [=Annaba] in northern Algeria could be found in the Pérez collection, and almost all were in very poor condition, missing their metasomas. A single female from Béne was however in good condition, and it is here designated as a new lectotype (Fig. 17). This new lectotype specimen (i.e. the true A. odsoleta) morphologically falls within the A. (Micrandrena) mariana Warncke, 1968 complex within the former Distandrena, as it lacks striations on the clypeus. The larger A. (Micrandrena) merimna Saunders, 1908 can be excluded as this species has a very long A3 that clearly exceeds the length of A4+5, whereas members of the A. mariana complex have A3 at most slightly exceeding A4+5, A4 and A5 are sub-quadratic and slightly shorter than wide, AG—12 are as long as wide. 274 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) | MUSEUM FAHD Andrena 21 . & COsSO!ETa Ag et. Dr. Wa rick “ ™ I P “> OLL. J, PEREZ 1015 MUSEUM Paris EY0000002005 ra obsolels olesignahion ypeorrecte, °*, ° Schenefe = al's tin guendd Fy é oh Figure 16. Andrena (Micrandrena) obsoleta Pérez, 1895, false female lectotype, =Andrena (Micrandrena) nitidula Pérez, 1903 A label details B profile C face, frontal view D dorsal view. Warncke described A. mariana s. str. from the island of Fuerteventura in the Canary Islands, stating that the species could potentially be found in Morocco (Warncke 1968b). He then described five subspecies in subsequent publications (Warncke 1974, 1975a): A. m. mica Warncke, 1974, A. m. leptura Warncke, 1974, A. m. solda Warncke, 1974, A. m. alma Warncke, 1975, and A. m. tenostra Warncke, 1975. For North African taxa, Warncke (1974) described A. mariana solda from Morocco (Tangier) and Algeria (Algiers, including Husseyn-Dey [Hussein Dey] the locus typicus, Constantine, Bone, Blida) and Tunisia (Carthago), and noted that this species corresponds to the Andrena forms from Bone identified as iota, kappa, and lambda by Saunders (1908), though this is clearly wrong for lambda as the genital capsule illustrated by Saunders shows A. (Micrandrena) djelfensis Pérez, 1895. The A. mariana solda taxon differs from A. mariana s. str. in the female sex by the darker tergal margins, the darker antennae, and the more extensively shagreened clypeus. Warncke (1974) described A. mariana mica from southern Algeria (locus typicus Ghardaia) and Morocco, giving it a more arid distribution. It also differs from A. mariana s. str. in the female sex by the darker tergal margins and darker antennae, but the clypeus is more extensively shiny than in A. mariana solda, and the nervulus is said to be more antefurcal. Andrena m. leptura is a more eastern taxon, being described from Libya and Egypt, and having a broader process of the labrum than any of these species. The genus Andrena in the Iberian Peninsula 275 Figure 17. Andrena (Micrandrena) obsoleta Pérez, 1895, true female lectotype A label details B profile C face, frontal view D scutum, dorsal view E propodeal triangle, dorsal view F terga, dorsal view. Examination of A. mariana solda material reveals that it is conspecific with the newly designated lectotype of A. obsoleta and is synonymised with it syn. nov. As iden- tified by Warncke, the taxon has a wide distribution across Mediterranean parts of Morocco, Algeria, and Tunisia (Warncke 1974, as A. m. solda). Pérez’s reference to specimens from France indicates that his original syntypic series would have been poly- typic, or he may have changed his mind between writing this catalogue entry and his 1895 publication, or even after this point. Warncke (1974: 40) indicated that A. m. solda is present in Sicily which is supported by a newly examined specimen (see below), 276 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) though he did not designate any Italian specimens as paratypes. As no specimens from France labelled as obsoleta could be found in Pérez collection, the conservative position is taken here that this species does not occur in France. The implications for Warncke’s misinterpretation of A. obsoleta are discussed below. As A. mariana solda is actually a synonym of a valid species that was described prior to A. (Micrandrena) mariana s. str., this has implications for the species-status of the other taxa lumped under A. mariana by Warncke. Genetic sequences were available for A. mariana mica Warncke, 1974 from southern Morocco, A. mariana s. str. from south- western Morocco, and A. mariana alma Warncke, 1975 from Portugal (locus typicus near Cérdoba, southern Spain; Fig. 18). Contrary to the position of Warncke (1968b), examination of material from Morocco, Algeria, and Tunisia shows that A. mariana is actually widely distributed across this region, predominantly in southern, more arid ar- eas. Genetically, A. mariana s. str. was more closely related to A. (Micrandrena) abjecta Pérez, 1895, separated by an average genetic distance of 5.55% (range 5.49-5.61%), and was strongly separated from A. m. mica by 12.73% (range 12.50—12.95%) and A, m. alma by 10.94% (range 10.65—11.39%). Andrena abjecta + A. mariana s. str. therefore formed a clade with bootstrap support of 99. Andrena m. mica and A. m. alma formed a clade, but were strongly separated from each other by 11.26% (range 10.97-11.41%). The conclusion therefore is that each of these taxa is distinct, and so A, mica stat. nov. and A. alma stat. nov. are raised to species status. Morphologically, all three taxa can clearly be separated by the structure of the female clypeus, the colour of the antennae, the extent of lightening on the tergal margins, the strength of the scutal punctures, and the shape of the process of the labrum. Warncke described two further subspecies of A. mariana: A. mariana leptura from Egypt and A. mariana tenostra from south-eastern Spain, the latter specifically from a single female specimen from Villajoyosa in the province of Alicante, then listing addi- tional specimens from Almeria and Murcia (Warncke 1976). The status of A. mariana leptura will be dealt with in a future publication, as its exact species concept and its eastern and western limits are unclear, though it will be a valid species as it clearly differs from A. mariana s. str.; its relationship with A. mariana s.|. specimens from southern Israel must also be clarified (Pisanty, in litt.). In Iberia, A. m. tenostra is very poorly known. The distinguishing features given by Warncke (1975a) are that the process of the labrum is narrowly trapezoidal and that the clypeus is arched and somewhat flattened in the middle, in contrast to A. alma where the clypeus is largely flattened. Additional female specimens were found in Granada which highlight a difference not noted by Warncke, which is that the scutellum is polished and shiny, whereas it is dull in A. a/ma, and more broadly the scutal punctures are less strongly pronounced. Given that subtle differences within this group represent large genetic differentiation, A. tenostra stat. nov. is raised to species status. Finally, a number of specimens were found in the very south of Spain (Malaga, Sevilla) which show a morphology very close to that of A. mica. However, ecologically this does not make sense as A. mica is not known from the more humid and Mediter- ranean areas north of the Atlas Mountains. Examination of these specimens shows that A3 is much longer than A4+5, whereas it is as long as A4+5 in A. mica. These specimens probably represent an additional undescribed species in the A. mariana complex, but The genus Andrena in the Iberian Peninsula 277 WPATW124-21_Andrena_murana_Spain_Lupiana WPATW119-21_Andrena_nitidula_Spain_Alcolea 98 WPATW634-22_Andrena_nitidula_TJW_822_Morocco_Oriental_Region WPATW635-22_Andrena_nitidula_TJW_823_Morocco_Fes-Meknes 98 KJ837117_Andrena_distinguenda_Germany_Thuringia KJ839472_Andrena_distinguenda_Germany_Thuringia IBIHM1043-22_Andrena_nitidula_Spain_Malaga distinguenda 84 WPATW187-21_Andre! itidula_Spain_Ucles pie KJ839833_Andrena_nitidula_Spain_Murcia / nitidula IBIHM1215-22_Andrena_nitidula_Portugal_Santo_Estevao KJ836688_Andrena_nitidula_Spain_Murcia WPATW217-21_Andrena. lula_Spain_Cortes_Frontera GU705971_Andrena_nitidula_France_Alsace GU705964_Andrena_nitidula_France_Alsace 100 WPATW4B88-22_Andrena_fumida_TJW_676_Morocco_Fes-Meknes | fumida WPATW489-22_Andrena_fumida_TJW_677_Morocco_Fes-Meknes 100 IBIHM513-21_Andrena_orana_Portugal_Pera_Praia_Grande | orana IBIHM512-21_Andrena_orana_Portugal_Pera_Praia_Grande WPATW738-22_Andrena_abjecta_TJW_926_Morocco_Guelmim-Oued_Noun_Region WPATW740-22_Andrena_abjecta_TJW_928_Morocco_Guelmim-Oued_Noun_Region 76 97; WPATW739-22_Andrena_abjecta_TJW_927_Morocco_Guelmim-Oued_Noun_Region a b, rj ecta 99 WPATW476-22_Andrena_abjecta_TJW_664_Morocco_Fes-Meknes WPATW737-22_Andrena_abjecta_TJW_925_Morocco_Guelmim-Oued_Noun_Region WPATW687-22_Andrena_mariana_s_str_TJW_875_Morocco_Guelmim-Qued_Noun_Region "9 88 WPATW688-22_Andrena_mariana_s_str_TJW_876_Morocco_Guelmim-Oued_Noun_Region | mariana s. str. WPATW620-22_Andrena_mariana_mica_TJW_808_Morocco_Draa WPATW621-22_Andrena_mariana_mica_TJW_809_Morocco_Draa IBIHM947-21_Andrena_mariana_alma_Portugal_Castelo_Branco 93 IBIHM1229-22_Andrena_mariana_alma_Portugal_Algarve i alma stat. nov. IBIHM1230-22_Andrena_mariana_alma_Portugal_Algarve 100 Bf mica stat. nov. IBIHM1219-22_Andrena_djelfensis_Portugal_Algarve ir A 100 WPATWS557-22_Andrena_djelfensis_TJW_745_Morocco_Fes-Meknes | djelfensis IBIHM482-21_Andrena_djelfensis_Portugal_Praia_Zavial IBIHIM1040-22_Andrena_fabrella_Spain_Guadalajara WPATW414-22_Andrena_fabrella_TJW_602_Morocco_Fes-Meknes 100 WPATW167-21_Andrena_fabrella_Spain_Madrona fabrella IBIHM1046-22_Andrena_fabrella_Spain_Malaga WPATW233-21_Andrena_fabrella_Spain_Sierra_Nieves KJ838631_Andrena_nana_Germany_Badden_Wurttemberg 'WPATW235-21_Andrena_nana_S| |_Sierra_Nieves. 100 'WPATW161-21_Andrena_nana_Spain_Madrona IBIHM509-21_Andrena_nana_Portugal_Gondesende 'WPATW473-22_Andrena_nana_TJW_661_Morocco_Fes-Meknes 'WPATW325-21_Andrena_nana_Spain_Veguillas 'WPATW282-21_Andrena_nana_Spain_Tocon_Quentar 99 HYMAA496-22_Andrena_rugulosa_Switzerland WPATW973-22_Andrena_rugulosa_1579_Lebanon_Chouf 91 WPATW413-22_Andrena_pauxilla_TJW_601_Morocco_Fes-Meknes WPATW285-21_Andrena_pauxilla_Spain_Sierra_Nevada WPATW363-21_Andrena_pauxilla_Spain_Atienza 99. WPATW366-21_Andrena_pauxilla_Spain_Villar_del_Cobo WPATW242-21_Andrena_pauxilla_Spain_Sierra_Nieves 87 WPATW314-21_Andrena_pauxilla_Spain_Guadalaviar WPATW302-21_Andrena_pauxilla_Spain_Valdecabras WPATW364-21_Andrena_pauxilla_Spain_Sierra_Baza KJ837178_Andrena_curtula_France_Province_Alpes_Cote_dAzur 91 100 IBIHM1036-22_Andrena_ampla_Spain_Avila WPATW674-22_Andrena_ampla_TJW_862_Morocco_Fes-Meknes WPATW673-22_Andrena_ampla_TJW_861_Morocco_Fes-Meknes: WPATW271-21_Andrena_ampla_Spain_Sierra_Nevada 79 97 WPATW100-21_Andrena_pusilla_Belgium_Damme POLLE2039-19 Andrena_pusilla_France_Tours. POLLE2802-19_Andrena_pusilla_France_Loire KJ839537_Andrena_pusilla_Germany_Rhineland_Palatinate 99 KJ836546_Andrena_pusilla_Germany_Rhineland_Palatinate WPATW186-21_Andrena_spreta_Spain_Ucles IBIHM532-21_Andrena_spreta_Portugal_Altura 85 | IBIHMS533-21_Andrena_spreta_Portugal_Almograve IBIHM531-21_Andrena_spreta_Portugal_Assaflora IBIHM536-21_Andrena_spreta_Portugal_Algoceira WPATW343-21_Andrena_spreta_Spain_Pinto WPATW133-21_Andrena_spreta_Spain_Toledo WPATW352-21_Andrena_spreta_Spain_Ucles ANDIL176-22_Andrena_spreta_Sa_ad ANDIL281-22_Andrena_spreta_Israel_Montfort ANDIL189-22_Andrena_spreta_Israel_Tel_Aviv ANDIL219-22 Andrena_spreta_Israel_Lakhish ANDIL207-22_Andrena_spreta_Israel_Yagur WPATW404-22_Andrena_spreta_TJW_592_Morocco_Guelmim-Oued_Noun_Region WPATW405-22_Andrena_spreta_TJW_593_Morocco_Guelmim-Oued_Noun_Region WPATW407-22_Andrena_spreta_TJW_595_Morocco_Draa WPATW411-22_Andrena_spreta_TJW_599_Morocco_Oriental_Region WPATW410-22_Andrena_spreta_TJW_598_Morocco_Draa WPATW409-22_Andrena_spreta_TJW_597_Morocco_Draa WPATW408-22_Andrena_spreta_TJW_596_Morocco_Draa 100 WPATW157-21_Andrena_simontornyella_Spain_Madrona WPATW108-21_Andrena_simontornyella_Spain_Balaguer MT871628_Andrena_simontomyella_France_Loire 76 100 KJ839781_Andrena_alfkenella_Germany_Brandenburg IBIHM474-21_Andrena_alfkenella_Portugal_Braganca ‘|, WPATW357-21_Andrena_alfkenella_Spain_Sierra_Baza WPATW029-21_Andrena_alfkenella_Spain_Ourense 98 WPATW338-21_Andrena_bayona_Spain_Valdeprados WPATW373-21_Andrena_bayona_Spain_Sierra_Nevada KJ839000_Andrena_minutuloides_Germany_Brandenburg 99 WPATW458-22_Andrena_minutuloides_TJW_646_Morocco_Fes-Meknes WPATW457-22_Andrena_minutuloides_TJW_645_Morocco_Fes-Meknes 98 WPATW456-22_Andrena_minutuloides_TuW_644_Morocco_Fes-Meknes WPATW016-21_Andrena_minutuloides_Belgium_Mons. WPATW342-21_Andrena_minutuloides_Spain_Bernuy_Porreros IBIHMS08-21_Andrena_minutuloides_Portugal_Vilar_do_Conde WPATWO93-21_Andrena_minutuloides_Belgium_Liege WPATW096-21_Andrena_minutuloides_Belgium_Chokier WPATWO095-21_Andrena_minutuloides_Belgium_Liege IBIHM541-21_Andrena_tenuistriata_Portugal_Curalha WPATW653-22_Andrena_tenuistriata_TJW_841_Morocco_Marrakech-Safi_Region WPATW654-22_Andrena_tenuistriata_TJW_842_Morocco_Fes-Meknes 95 WPATW655-22 Andrena_tenuistriata_TJW_&43 Morocco _Souss-Massa_Region WPATW656-22_Andrena_tenuistriata_TJW_844_Morocco_Souss-Massa_Region 100 WPATW007-21_Andrena_subopaca_Belgium_Han_sur_Lesse 84 WPATWO86-21_Andrena_subopaca_Luxembourg_Dreps WPATW597-22_Andrena_minutula_TJW_785_Morocco_Ouarzazarte L WPATW512-22_Andrena_minutula_TJW_700_Moracco_Souss-Massa_Region WPATW596-22_Andrena_minutula_TJW_784_Morocco_Ouarzazarte WPATWS595-22_Andrena_minutula_TJW_783_Morocco_Fes-Meknes 98 (BIHMS506-21_Andrena_minutula_Portugal_Odeceixe WPATW232-21_Andrena_minutula_Spain_Sierra_Nieves WPATW353-21_Andrena_minutula_Spain_Grazalema 89 'WPATW238-21_Andrena_minutula_Spain_Sierra_Nieves IBIHM535-21_Andrena_minutula_Portugal_Lindoso KJ838202_Andrena_minutula_ltaly_ Lombardy IBIHM507-21_Andrena_minutula_Portugal_Cabril 99 KJ836685_Andrena_minutula_Germany_ Rhineland WPATW328-21_Andrena_minutula_Spain_Atienza 99 WPATW307-21_Andrena_saxonica_Spain_Griegos. 7 ‘WPATW162-21_Andrena_saxonica_Spain_Madrona saxonica IBIHM1037-22_Andrena_saxonica_Spain_Avila 76 os WPATWS551-22_Andrena_atlantea_TJW_739_Morocco_Fes-Meknes ——— atlantea WPATW685-22_Andrena_sp._TJW_873_Morocco_Fes-Meknes = 96 WPATW686-22_Andrena_sp._ TJW_874_Morocco_Fes-Meknes. + undescribed WPATW$971-22_Andrena_Micrandrena_nov_TJW571_Spain_Sierra_Nevada a 98 WPATW972-22_Andrena_Micrandrena_nov_TJWS72_Spain_Sierra_Nevada | ortizi sp. nov. j WPATW615-22_Andrena_purpurascens_TJW_803_Morocco_Fes-Meknes ——_—_—_—_—_—_—_—_— purpurascens 98 7 WPATW684-22_Andrena_longibarbis_TJW_872_Morocco_Souss-Massa_Region 100 WPATW147-21_Andrena_longibarbis_Spain_Sesena_Nuevo IBIHM504-21_Andrena_longibarbis_Portugal_Nazare longibarbis IBIHM505-21_Andrena_longibarbis_Portugal_Fornelos WPATW188-21_Andrena_longibarbis_Spain_Ucles nana rugulosa pauxilla sp. resurr. ampla pusilla spreta simontornyella alfkenella bayona minutuloides 100 tenuistriata subopaca 96 minutula cy Figure 18. Phylogenetic tree (maximum likelihood) of Andrena subgenus Micrandrena Ashmead, 1899 based on the mitochondrial COI gene. Andrena (incertae sedis) murana Warncke, 1975 is used as an out- group. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted). 278 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) they are not described at this time, as it would be beneficial to have barcoded speci- mens to confirm this differentiation. They are therefore referred to as aff mica in the identification key. It is important to note that this material does not belong to A. abjecta, the status of which in Iberia is unclear. Warncke (1976) did not record this species from Iberia, and only his distribution maps (Gusenleitner and Schwarz 2002) indicate the pres- ence of this species in northern and eastern Spain. This distribution does not make sense given that all other known records of A. abjecta come from north-western Africa (Morocco, Algeria, Tunisia), so the species would naturally be expected to occur in southern Iberia. Dardén (2014) listed two female specimens from Huelva (22.iv.1987) and Zaragoza (9.iii.1991). The specimen from Huelva may represent the undescribed aff mica taxon, but it is more difficult to conclude on the specimen from Zaragoza. For now, A. abjecta is retained on the Iberian list, but I have not examined any specimens and I consider its presence in Iberia doubtful. Material examined. Andrena alma: Porrucat: Albandeira, near Lagoa, 20.iv.2005, 19, leg. D.W. Baldock, TJWC; Algarve, Casaqueimada (7 km N of Silves), 20.ii1.1995, 19, leg. T. & M. Simon Thomas, RMNH; Algarve, Monte Gordo, Re- tur, Praia do Cabeco, 29.iii.2022, 13, leg. TJ. Wood, TJWC; Algarve, Tavira, Cacela Velha, 28.iii.2022, 1, leg. TJ. Wood, TJWC; Castelo Branco, Fundao Vale de Praz- eres, 17.iv.2021, 19, leg. C. Siopa, FLOW; Spain: El Soldado, Cérdoba [38.3747°N, -5.0001°E], 19, leg. Seyrig, OOLM (holotype); Almodévar del Campo (Ciudad Real), 700 m, 24.iii.2005, 13, leg. EJ. Ortiz-SAnchez, FJOS; Santa Ana la Real, Sierra Aracena (Huelva), 630 m, 13.iv.2006, 24, leg. EJ. Ortiz-Sanchez, FJOS; El Hongo (PN. Donana), 30.iii.2018, 13, leg. F Molina, EBDC. Andrena mariana s. str.: ALGERIA: Saida, 15 km S of Sfissifa, Ben Ikhou, st. 6, 6.iv.1983, 69, leg. R. Leys & P. v. d. Hurk, RMNH; 23 km NE of Ras El Ma, st. 38, 26.iv.1983, 19, leg. R. Leys & Pv. d. Hurk, RMNH; Dayet el Kerch [Daiet el Kerch], st. 5, 5.iv.1983, 29, leg. R. Leys & P. v. d. Hurk, RMNH; Morocco: Guelmim- Oued Noun, Guelmim, Asrir, 1 km W, 20.iii.2022, 19, leg. TJ. Wood, TJWC; Spain: Fuerteventura, Valley Granadillos [28.3864°N, -14.0865°W], 17.iv.1934, 19, OOLM (holotype); Tunisia: Ksar Hadada, 4—5.iv.1996, 469, leg. K. Dene§, OOLM; M’saken, 20—21.iv.1998, 29, leg. K. Denes, OOLM; Wadi Ram, 45 km E Douz, 4.iv.2006, 19, leg. J. Straka, OOLM. Andrena mica: Auceria: Ghardaia [32.5047°N, 3.6419°E], 19, OOLM (holo- type); Morocco: 30 km E Midelt, 13.v.1995, 19, leg. Mi. Halada, OOLM; Dria- Tafilalet, Ouarzazate, P1506, Telouet, Adaha, 1700 m, 18.iv.2022, 69, leg. TJ. Wood, TJWC; Drda-Tafilalet, Ouarzazate, 2 km W Agouim, 1800 m, 13.iv.2022, 39, leg. TJ. Wood, TJWC; Ifkern, 25 km E Boulemane, 24.v.1995, 19, leg. Mi. Halada, OOLM; M’rirt (30 km N), 11.iii.1989, 1Q, leg. H. Teunissen, RMNH; Tunisia: Hammamet eny, 15.iii.1996, 19, leg. K. Denes, OOLM; Kasserine, 13.iv.1998, 19, leg. K. Denes, OOLM; Nefta [Naftah], 20.v.1993, 19, leg. J. Batelka, OOLM. Andrena aff mica: Spain: 40 km W Malaga, Yunquera, 800 m, 29.iv.2003, 52, leg. J. Halada, OOLM/TJWC; La Corchuela (Dos Hermanas, Sevilla), 35 m, The genus Andrena in the Iberian Peninsula 279 TAT 2009 26.22, leg. EJ. Ortiz-Sanchez, FJOS; Rio Blanco, Aguadulce (Sevilla), 300 m, 17.v.2008, 19, leg. FJ. Ortiz-Sanchez, FJOS. Andrena nitidula: Acer: Babor, 19, MNHN (false lectotype of A. obsoleta). Andrena obsoleta: ALGERIA: Bone [=Annaba, 36.9092°N, 7.7264°E], 12, MNHN (lectotype of A. obsoleta, by present designation); 23 km NE of Ras El Ma, st. 38, 26.iv.1983, 19, leg. R. Leys & P. v.d. Hurk, TJWC; 5 km N of Mecheria, Djebel Antar, st. 10, 7.iv.1983, 19, leg. R. Leys & P. v.d. Hurk, RMNH; Ivar: Sicilia orient., Taormina, 200 m, Ghardino [Giardini Naxos], 25.iii.1950, 14, Hartig & Grisham, MZUR; Tunisia: Tunis, 1911, 39, leg. O. Schmiedeknecht, SMFD. Andrena tenostra: Spain: Villajoyosa [38.5097°N, -0.2299°E], 11.v.1936, 19, leg. Andréu, OOLM (holotype); Salobrefia, Granada, 8.v.1983, 39, leg. W. Perrandin, OOLM/TJWC. Andrena (Micrandrena) distinguenda Schenck, 1871 and Andrena (Micrandrena) nitidula Pérez, 1903 The status of these two species has been extensively argued over, and despite much attention the position remains somewhat unclear. Warncke (1967) argued that A, distinguenda Schenck, 1871 was an unavailable name because it was preoccupied by A. distinguenda Schenck, 1853. He then authored the replacement name A. obsoleta spongiosa Warncke, 1967, using the incorrect concept that A. obsoleta was the oldest available name for this group of species, as discussed above. Schénitzer et al. (1992) clarified the situation, and argued that A. distinguenda Schenck, 1853 was a nomen nudum because it appears without a description in a discussion concerning species related to A. bicolor. As a nomen nudum, the name is available and cannot preoccupy its later usage. Therefore, the name A. distinguenda Schenck, 1871 is both available and valid as the senior synonym; A. obsoleta spongiosa is consequently an unnecessary replacement name. Andrena distinguenda was described from western Germany, and A. nitidula was described from south-western France (lectotype from Bordeaux [though this is not indicated on the specimen, it bears the number ‘675’ which refers to the entry for A. nitidula in the catalogue of Pérez], designated by Warncke 1967, Fig. 19). As dis- cussed above, Warncke (1967) incorrectly recognised a specimen that morphologi- cally corresponds to A. nitidula as the lectotype of A. obsoleta, and then synonymised A. nitidula with A. obsoleta. He then treated A. distinguenda Schenck, 1871 as a sub- species of A. obsoleta in the form A. obsoleta spongiosa. He then later adopted a three subspecies model, A. 0. obsoleta (North Africa), A. 0. nitidula (Western Europe), and A. 0. spongiosa (Central Europe) (see Schénitzer et al. 1992), finally introducing a fourth subspecies A. 0. puella Alfken, 1938 in Sardinia and Sicily (see Gusenleitner and Schwarz 2002). Schénitzer et al. (1992) argue that both A. distinguenda and A. nitidula can be morphologically separated in north-western Italy (Bordighera). They argue that A, nitidula occurs in North Africa, citing Pérez (1903) and Alfken (1914). They also argue that specimens from southern Spain (Tarifa) appear to be somewhat intermedi- 280 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) MUSEUM PARIS] — EY0000002004 | Andrena #¥t ' Figure 19. Andrena (Micrandrena) nitidula Pérez, 1903, female lectotype A label details B profile C face, frontal view D terga, dorsal view. ate between A. distinguenda and A. nitidula, concluding that the question as to whether there are two species or simply a very variable single species remains open. Burger and Herrmann (2003) revised European material from this species pair using the names A. distinguenda and A. nitidula, after having examined syntypes of A, nitidula in the ZMHB collection. Using morphological characters, they conclude that (in a European context) A. distinguenda has a predominantly eastern distribution, from Central Europe to Greece, with populations also in the south of France and a few records from north-eastern Spain. In contrast, A. nitidula is a western species, from Iberia to the south and east of France and into south-western Germany and the north-west of Italy. They reach the same conclusion as Sch6nitzer et al. (1992), that where these two forms occur in sympatry (Germany, France, Italy) they can be mor- phologically separated. They do not conclude on the status of A. obsoleta, but note that A. nitidula occurs in North Africa, and that Warncke’s treatment is unclear. This prob- lem arises because it does not appear that any of the authors of Sch6nitzer et al. (1992) or Burger and Herrmann (2003) inspected the lectotype of A. obsoleta designated by Warncke in the MNHN. This lectotype bears a handwritten label that the lectotype designation is incorrect, and that the specimen corresponds to A. distinguenda Schenck (Fig. 16A). However, it is unclear who wrote this, as neither Schénitzer et al. (1992) The genus Andrena in the Iberian Peninsula 281 nor Burger and Herrmann (2003) mention this explicitly, and I do not recognise the handwriting on the label. Barcode analysis complicates this matter further (Fig. 18). Analysis of sequences from France, Germany, Morocco, Portugal, and Spain shows the presence of two clear clades. One clade is composed of specimens identified as A. nitidula from France, Portugal, and Spain, including specimens from the south of Iberia (Algarve, Malaga, Murcia). This clade would appear to be the true A. nitidula. However, the second clade contained sequences from specimens identified as A. distinguenda from central Germany, but also from a specimen identified as A. nitidula from central Spain (Guadalajara) and two specimens identified as A. nitidula from northern Morocco (Fés-Meknes, Oriental). These two clades were consistently separated genetically, by an average genetic distance of 7.08% (range 5.58-8.77%). The nominally true A. nitidula clade showed low in- traspecific variation of 0.26% (range 0.00—0.63%). However, the situation in the clade containing A. distinguenda sequences from Germany showed intraspecific variation of 2.84% (range 0.00—4.10%), with the 4.10% genetic distance recorded between speci- mens from Germany and the Middle Atlas (Bakrit, near Azrou) in Morocco. Morphologically, both Spanish and Moroccan specimens falling into the A. dis- tinguenda clade conform to the concept of A. nitidula using the criteria specified by Burger and Herrmann (2003). Indeed, all material I have identified from north- western Africa morphologically conforms to the concept of A. nitidula sensu Burger and Herrmann. Although the genetic data strongly supports the existence of two spe- cies, the conflict between the genetic and morphological results suggests that, at the moment, they cannot be consistently separated morphologically across their range. No taxonomic action is taken here, and the morphological criteria of Burger and Herrmann (2003) are followed for the purpose of the identification key. In Iberia, A, nitidula is considered to be present throughout, but A. distinguenda is limited to the extreme north-east of Spain. However, it is clear that there is a major disagree- ment between the morphological and genetic methods, and these two species may ultimately not be consistently separable. Further analysis using more powerful genetic techniques is necessary. Relictual species morphologically related to Andrena (Micrandrena) rugulosa Stéckhert, 1935 Inspection of Micrandrena specimens from high altitude in the Sierra Nevada revealed the presence of a species that morphologically resembles A. (Micrandrena) rugulosa Stéckhert, 1935 due to its head that is only slightly shorter than wide rather than clear- ly shorter than wide, an unusual character in Micrandrena. This finding is remarkable, because although widely distributed in Central and Eastern Europe, A. rugulosa has a western limit in the Swiss Alps, and has not been previously recorded from France, Spain, or Portugal (Gusenleitner and Schwarz 2002). Samples collected from the Sierra Nevada at high altitude (2000-2100 m) during 2021 are strongly differentiated from A, rugulosa, separated by an average genetic distance of 16.01% (range 15.81-16.41%; 282 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Fig. 18). Instead, they form a clade with bootstrap support of 96 with sequences of A, (Micrandrena) atlantea Wood, 2021 (High and Middle Atlas in Morocco) and a morphologically similar undescribed Micrandrena from high altitude (1800-1900 m) in the Moroccan Middle Atlas. The Sierra Nevada sequences were separated from A. atlantea by an average genetic distance of 6.04% (range 5.71—6.38%) and from the undescribed Micrandrena by an average genetic distance of 6.34% (range 6.31-6.38%). These three species would therefore seem to represent an isolated Micrandrena lineage that is restricted to the Sierra Nevada and the High and Middle Atlas Mountains in Morocco, with conse- quent genetic and morphological divergence. This is the same pattern as observed in the subgenus Euandrena, suggesting that the Sierra Nevada hosts the remnants of a fauna that was presumably once more widespread across Iberia and North Africa. Additional sampling and genetic analysis is required to determine if this pattern holds true for other bee groups. The new Micrandrena species from the Sierra Nevada is described below, and the undescribed Micrandrena species from Morocco will be de- scribed in an upcoming publication. Andrena (Micrandrena) niveata Friese, 1887 sensu lato Andrena niveata was described from Germany and Hungary, without further informa- tion (Friese 1887). The exact type locality is unclear, as well as the type depository, with no clearly identifiable syntypes located during searches in the SMFD and ZMHB collec- tions, the suggested possible depositories (Gusenleitner and Schwarz 2002). Nevertheless, the concept of A. niveata in Central Europe is clear, and A. niveata is a well-defined spe- cies in this region (Schmid-Egger and Scheuchl 1997; Gusenleitner and Schwarz 2002; Amiet et al. 2010). Warncke described three subspecies; A. 1. lecana Warncke, 1975 (Spain; ocus typicus: Rivas-Vaciamadrid), A. 2. haloga Warncke, 1980 (Italy; locus typicus: Lecce), and A. n. bubulca Warncke, 1975 (Turkey; locus typicus: Erzurum); the status of the Italian and Turkish subspecies is not addressed here. Unfortunately, all nine freshly collected A. 7. lecana specimens from central and southern Spain (including from the locus typicus) sent for genetic analysis failed to produce sequences. More specific primers will be required to address this question using genetic evidence. Warncke (1975a) described A. n. lecana extremely briefly, noting that in both sexes the first tergum was finely but clearly shagreened, and finely but more strongly punc- tured than in A. niveata s. str., with the following terga also more clearly and strongly punctured. He then gave a distribution of A. 2. lecana across much of central Spain south to hilly parts of the provinces of Granada (Huéscar) and Almeria (Tijola). In contrast he gives a distribution for A. niveata s. str. of the whole of Iberia, though his distribution maps (Gusenleitner and Schwarz 2002) show a distribution covering eastern, central, and south-eastern Spain. ‘There is therefore the possibility that the two subspecies exist in sympatry, suggesting that they may be distinct. Examination of material from Spain shows that the two taxa are distinct, but morphological separation is challenging, may not be possible in all cases when old, The genus Andrena in the Iberian Peninsula 283 abraded, or dirty specimens are available, and is best made with reference to the male genital capsule. Taking male specimens, the shape of the gonostyli are distinctive. In A, niveata s. str., the gonostyli are elongate, with the inner margins of the gonostyli only weakly produced towards the penis valves (Fig. 20A). In contrast, in A. n. lecana, the inner margins of the gonostyli are strongly and clearly produced towards the pe- nis valves (Fig. 20B); the apical spatulate part of the gonostyli are also comparatively shorter, thus the gonostyli appear shorter and more compact in A. n. lecana and longer and more elongate in A. niveata s. str. The structure of the genital capsule of A. niveata s. str. is consistent across Spain to the province of Malaga (the most southerly A. ni- veata s. str. male examined). Furthermore, the male scutum is shagreened in A. niveata s. str., whereas it is extensively shiny in A. 7. lecana, and the terga are also more strongly shagreened in A. niveata s. str. compared to A. n. lecana (Fig. 20C, D). Separation of females (recognised by their wide and poorly defined propodeal triangle, Fig. 20E, F in combination with their depressed tergal margins) is more challenging and may not be possible in all situations. Typically, A. niveata s. str. has the terga shagreened, with punctures on the tergal discs partially disappearing into this shagreenation (Fig. 20G). In contrast, A. 7. lecana has the terga shiny, almost without shagreen, with the dense punctures on the tergal discs strong and clearly visible, not disappearing due to the absence of background shagreenation (Fig. 20H). The scutum and scutellum are also more extensively shiny, whereas in A. niveata s. str. these areas usually show at least some shagreenation and are never polished and smooth. The problem comes that some individuals nominally assigned to A. niveata s. str. have partially shing terga, and with- out genetic sequences it is not possible to say with complete confidence if they belong to A. niveata s. str. or A. n. lecana. Moreover, all Iberian A. niveata s. str. have tergal sha- greenation that is weaker than that of A. niveata s. str. populations in Central Europe. The position is taken here that the strength of tergal shagreenation in A. niveata s. str. females is variable across Europe, but that A. /ecana stat. nov. is a valid species based on the overlapping range in combination with the consistent difference in the shape of the male genital capsule. It has a distribution across steppic parts of central Spain, extending into mountainous areas in south-eastern Spain. Females displaying any level of shagreenation on the tergal discs are considered to represent Iberian popu- lations of A. niveata s. str., and females with completely polished tergal discs without a trace of shagreenation represent A. /ecana (see identification key). Future genetic inves- tigation using more targeted primers will be necessary to confirm this position. Finally, specimens of A. /ecana from high altitude in the Sierra Nevada show slightly different antennal ratios in the male sex, though the male genital capsule is otherwise identical; this requires further investigation. Material examined. Andrena lecana: SPAIN: Ribas [Rivas-Vaciamadrid, 40.3503°N, -3.5390°E], 6.v.1908, 19, leg. Dusmet, OOLM (holotype); Carbon- eras de Guadazaén (Cuenca), 1030 m, 16.v.2009, 19, leg. FJ. Ortiz-Sanchez, FJOS; Guadalajara, Alcolea del Pinar, 12.v.2021, 19, leg. TJ. Wood, TJWC; Guadalajara, Lupiana, 12.v.2021, 19, leg. TJ. Wood, TJ WC; Huéscar (Granada), 1900, 14, leg. Escalera, OOLM (paratype); La Cabrilla, Sierra Cazorla (Jaén), 1600 m, 1—3.vi.2022, 284 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Figure 20. Andrena (Micrandrena) niveata Friese, 1887 A male genital capsule C male terga, dorsal view E female propodeal triangle G female terga, dorsal view; Andrena (Micrandrena) lecana Warncke, 1975 B male genital capsule D male terga, dorsal view F female propodeal triangle H female terga, dorsal view. The genus Andrena in the Iberian Peninsula 285 1Q, leg. C.M. Hererra, CMHC; Madrid, Rivas-Vaciamadrid, Canal de Manzanares to Camino de Uclés, 19.v.2021, 29, leg. TJ. Wood, TJ WC; Orgiva, N, 1300 m, Sierra Nevada, 26.vi.1988, 19, leg. M. Schwarz, OOLM;; Pozuelo, La Fuente, 14, OOLM (paratype); Segovia, Madrona, 500 m NE, Arroyo del Hocino, 15.v.2021, 29, leg. TJ. Wood, TJWC; Sierra Nevada, Trevélez, Refugio La Campifuela, 2400 m, 14.vi.2021, Tied Oe leg. TJ. Wood, TJ WC. Andrena niveata s. str.: SPAIN: Caceres, Cuacos de Yuste, 500 m, 11.v.1999, 19, leg. H. & J.E. Wiering, RMNH; Cadiz, Grazalema, Rio Guadalete, 28.v.2021, 19, leg. TJ. Wood, TJ WC; Cadiz, Embalse de Barbate Sw, 14, 39, 6.v.2017, leg. Bartak & Kubik, OOLM/TJWC; Granada, Sierra de Baza, Prados del Rey, 2000 m, 19.vi.2021, 119, leg. TJ. Wood, TJWC; Malaga, 5 km W of Alhaurin de la Torre, 4.vi.1962, 19, leg. Jeekel & Wiering, RMNH; Malaga, Cortes de la Frontera, path to Llano de las Labores, 26.v.2021, 39, leg. T.J. Wood, TJWC; Malaga, Estepona, 21.iv.1983, 19, leg. H. Teunissen, RMNH; Navarra, Tudela, 16.iv.1978, 19, leg. C. Gielis, RMNH; Gerona, Figueras, 15.iv.1971, 19, leg. J. Leclercq, UMONS. Andrena (Micrandrena) spreta Pérez, 1895, Andrena (Micrandrena) curtula Pérez, 1903, Andrena (Micrandrena) pusilla Pérez, 1903, and Andrena (Micrandrena) pauxilla Stéckhert, 1935 Members of this species group are challenging to identify and have been inconsistently treated in the literature, with variable species concepts. Andrena spreta was described from Algeria, with Warncke (1967) designating a lectotype from Biskra (Fig. 21A; Pérez also mentions a female from Constantine in his catalogue). Andrena curtula was described from north-eastern Spain, with Warncke (1967) designating a lectotype from Barcelona (Fig. 21B). Andrena pusilla was described from the south of France (Bordeaux, Nantes) and Spain, with Warncke (1967) designating a lectotype from Nantes (Fig. 21C). Finally, A. pauxilla was described from the Upper Rhine valley in south-western Germany, with Stéckhert (1935) designating a female from Karlsruhe as the holotype. Warncke (1967) recognised three species, A. spreta, A. curtula, and A. pauxilla, treat- ing A. pusilla as a subspecies of A. spreta. Warncke (1974) recognised only one taxon in North Africa (A. spreta spreta), but Warncke (1976) surprisingly recognised only A. spre- ta pauxilla in Iberia, not mentioning A. curtula as a species despite its description from Iberia, only referring to its listing in Ceballos (1956) but indicating that this referred to A. (Micrandrena) tenuistriata Pérez, 1895. This treatment is difficult to understand. Warncke later adopted an extremely broad position (see distribution maps in Gusenleit- ner and Schwarz 2002), lumping all members of the group into a broad A. spreta. In this model, A. spreta s. str. was restricted to North Africa, A. spreta curtula was distributed across Iberia to southern France, A. spreta pusilla was found in northern and Central Europe, with two more subspecies in i) Italy and ii) south-eastern Europe to the Levant. This position was not adopted by subsequent authors. Schwarz et al. (1996) ac- cepted A. pusilla as a valid species, as well as A. curtula. However, they synonymised A, pauxilla with A. curtula. Schmid-Egger and Scheuchl (1997) did not follow this inter- 286 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) it USEUM PARIS P11 COLL. J. VACHAL | tein ahdiaimahal Figure 21. Andrena (Micrandrena) spreta Pérez, 1895, female lectotype A label details B profile; Andrena (Micrandrena) curtula Pérez, 1903, female lectotype C label details D profile; Andrena (Micrandrena) pusilla Pérez, 1903, female lectotype E label details F profile. pretation, treating A. pauxilla as a distinct species, though Amiet et al. (2010) followed the position of Schwarz et al. (1996). Dardén (2010) and Dardon et al. (2014) accepted only a broad _ A. spreta taxon, including A. curtula, A. pusilla, and A. pauxilla, recognis- ing A. spreta s. str. and_A. spreta pusilla in an Iberian context. The situation is therefore unclear, both for the number of species present in this complex, and their distributions. Analysis of barcodes provides unambiguous support for the existence of three dis- tinct species (Fig. 18), A. spreta including A. curtula, A. pusilla, and A. pauxilla sp. The genus Andrena in the Iberian Peninsula 287 resurr. Specimens of A. spreta from Israel, Morocco, Portugal, Spain formed a clear clade with bootstrap support of 85. These sequences were separated by an average genetic distance of 0.45% (range 0.00—1.07%). They were clearly separated from a sister clade of A. pusilla sequences from Belgium, France, and Germany by an average genetic distance of 4.33% (range 3.88—4.84%). The A. pusilla clade had low intraspe- cific genetic distance of 0.29% (range 0.00—-0.71%). Andrena pauxilla was strongly differentiated and was found as sister to A. (Micrandrena) rugulosa, showing average genetic differentiation of 10.96% (range 10.38-11.85%) from A. spreta and 11.69% (range 10.75—12.59%) from A. pusilla. This A. pauxilla clade includes a specimen from the south of France (KJ837178, Lac St. Croix) that was identified as A. curtula, the ge- netic results suggest that it is misidentified. Morphologically, A. pauxilla is clearly dif- ferentiated from A. spreta and A. pusilla due to the absence of a gradulus at the base of the terga. Confusion is impossible in the female sex after recognition of this character; the foveae are also longer and narrower and filled with white rather than light brown hairs, the scutum is more densely and uniformly punctate when compared to A. spreta. Examination of the lectotypes of A. spreta and A. curtula shows no clear structural difference. The degree of shagreenation varies, but this is typical for A. spreta across its range; both lectotype specimens show the presence of a gradulus at the base of the terga, excluding their conspecificity with A. pauxilla. Due to these genetic results com- bined with the morphology of the lectotype specimens, the synonymy of A. curtula with A. spreta as proposed by Dardén (2010) and Dardon et al. (2014) is therefore fol- lowed, though their synonymies of A. pusilla and A. pauxilla with A. spreta are rejected. A single barcode was available from Moroccan specimens from the Middle Atlas tentatively identified as A. pauxilla which showed an average genetic distance to Eu- ropean A. pauxilla specimens of 2.00% (range 1.48%—2.29%). This is considered to represent only separation by distance, and thus A. pauxilla is recorded for the first time in North Africa and unambiguously recorded in Spain. Within Spain, A. pauxilla ap- pears to be principally recorded from mountain ranges such as the Sierra de las Nieves, the Sierra Nevada (Fig. 22A, B), Sierra Cazorla, Sistema Central, and Sistema Ibérico (Fig. 22C, D). Andrena pauxilla would therefore have a currently known distribution of Morocco, Spain, France, and Germany. On the basis of these results, A. spreta would appear to be a pan-Mediterranean species, and A. pusilla its predominantly northern counterpart (though the species descends south into Italy). The two species can be found in sympatry in the south of France (Praz, in litt.), but to date I have examined no A, pusilla specimens from Iberia. Due to the confusion in this group, it must be consid- ered absent until it can be positively demonstrated to be present south of the Pyrenees. Material examined. Andrena curtula: Spain: Barcelona [41.4028°N, 2.1332°E], 12, MNHN (lectotype; Fig. 21B). Andrena pauxilla: France: Bischenberg, 28.vi.1936, 19, leg. M. Klein, det. E. Stockhert, OOLM; Hausbergen, 29.vi.1930, 19, leg. M. Klein, det. H.R. Schwen- ninger, OOLM: Morocco: Fés-Meknés, Boulemane, R503, 7 km SE of Boule- mane, 1900 m, 22.v.2022, 14, 19, leg. TJ. Wood, TJWC; Spain: Canet de Mar, 26.iii.1963, 19, leg. F. Vergés, det. H.R. Schwenninger, OOLM; Cuenca, Huerta del 288 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Figure 22. Andrena (Micrandrena) pauxilla Stéckhert, 1935 A habitat, Granada, Sierra Nevada, Mirador Monte Ahi de Cara, 2100 m, 12.vi.2021 B female collecting pollen from Vella spinosa (Brassicaceae) C habitat, Cuenca, Mirador Valle de Valdecabras, 21.vi.2021 D female collecting pollen from Sedum spp. (Crassulaceae). Marquesado, environs north of town, 26.vi.2021, 39, leg. TJ. Wood, TJ WC; Gra- nada, Sierra Nevada, Jardin Botanico Hoya de Pedraza environs, 1900 m, 9.vi.2021, 1Q, leg. TJ. Wood, TJWC; Granada, Sierra Nevada, Mirador Monte Ahi de Cara, The genus Andrena in the Iberian Peninsula 289 2100 m, 12.vi.2021, 19, leg. TJ. Wood, TJ WC; Guadalajara, Aldeanueva de Atienza, 9.vii.2021, 19, leg. TJ. Wood, TJWC; Malaga, PN Sierra de las Nieves, mountain peak S of Pinsapo Escalereta, 30.v.2021, 19, leg. T.J. Wood, TJ WC; La Cabrilla, Sierra Cazorla (Jaén), 1600 m, 3.vi.2022, leg. C.M. Herrera, CMHC; Sierra Cazorla, Puerto Llano, 1800 m, 11.vi.2022, 13, 39, leg. J. Valverde, CMHC; Teruel, Guadalaviar, Rambla de los Ojos, 27.vi.2021, 19, leg, TJ. Wood, TJWC; Teruel, Villar del Cobo, Barranco de los Oncenachos, 27.vi.2021, 69, leg, TJ. Wood, TJWC; Cuenca, Mira- dor Valle de Valdecabras, 21.vi.2021, 29, leg, T.J. Wood, TJWC. Andrena pusilla: France: Nantes [47.2233°N, -1.5542°W], 12, MNHN (lecto- type; Fig. 21C). Andrena spreta: ALGERIA: Biskra [34.8600°N, 5.6995°E], 12, MNHN (lecto- type; Fig. 21A). Andrena (Micrandrena) strohmella Stéckhert, 1928 and Andrena (Micrandrena) icterina Warncke, 1974 Andrena strohmella was described from southern Germany and is a typically early spring species in the Central European Andrena fauna, with records extending south to the High and Maritime Alps in France, and west to the Bordeaux region; it has not previously been reported from the Pyrenees (Gusenleitner and Schwarz 2002). Warncke (1974) later described A. icterina from northern Algeria. He later indicated the presence of this species in south-eastern Spain in his distribution maps (Gusenleit- ner and Schwarz 2002), though he did not list its present in Iberia (Warncke 1976). Wood et al. (2020b) later reported the presence of this species in northern Morocco. In his description of A. icterina, Warncke draws comparison with A. strohmella, noting that the process of the labrum is half as broad, the clypeus is more sparsely punctate and lacking an unpunctured midline, that the terga are more strongly shagreened, and that the carinae on the dorsolateral corners of the first tergum are more weakly pro- nounced. Dardén (2010) and Dardén et al. (2014) did not actually examine any mate- rial of Iberian A. icterina, but inspection of new Iberian material has demonstrated that A, icterina is more widespread in Iberia than previously thought, being found from the Los Alcornocales to the Sierra de las Nieves, the Sierra Nevada, Sierra de Baza, Sierra Cazorla, and Sistema Central, and is also newly reported from northern Portugal (see below). In all of these localities it is found in mountainous areas, generally at high altitude in excess of 1000 m. Dardén (2010) and Dardén et al. (2014) reported A. strohmella from north-east- ern Spain, from Tarragona: Alcanar, v.2002, leg. Kadlec, OOLM. I have not been able to locate and examine this specimen for myself in the OOLM collection, but it is slightly problematic for ecological regions. Andrena strohmella is clearly a species of deciduous temperate woodland; this is reflected in its distribution across Central Eu- rope, and in its association with flowering trees such as Crataegus (Rosaceae), Prunus (Rosaceae), and Salix (Salicaceae). Andrena strohmella has therefore not previously been reported from Mediterranean regions. The record from Alcanar does not therefore match the ecology of this species, Alcanar being a low-elevation (c. 70 m) village close 290 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) to the Mediterranean coast. However, A. strohmella is present in north-eastern Catalo- nia based on recently collected specimens (Alvarez Fidalgo, in litt.), and so the species is retained on the Spanish and Iberian lists. More broadly, morphological differences between A. strohmella and A. icterina are slight, though they do not appear to introgress based on examined specimens. Some characters such as the strength of the carinae on the dorsolateral corners of the first tergum are not completely consistent, because some specimens in Central Europe can be found in which these are very weakly produced. Genetic data will hopefully clarify the status of A. icterina, but unfortunately, like A. /ecana, this taxon appears to be chal- lenging to barcode, as all seven Iberian specimens sent for genetic analysis failed or returned corrupted sequences. Material examined. Andrena icterina: PortuGaL: Braganga, Serapicos, 16.v.2021, 1Q, leg. A. Soares, A. Soares Coll.; Spain: Avila, Hoyocasero, El Pinar de Hoyocasero, 16.v.2021, 19, leg. TJ. Wood, TJ WC; Avila, Navalsauz, 1 km E, Alberche stream, 16.v.2021, 19, leg. TJ. Wood, TJWC; Cadiz prov., Vent. L. Canillas Hozgarganta- Tal b. Jimena 250 m, 14.iv.1985, 39, leg. W. Schacht, OOLM; Campamento Alf- aguara (Alfacar, Granada), 1420 m, 13.v.2007, ieee, leg. EJ. Ortiz-Sanchez, FJOS; Cortijo Tortas, Paterna del Madera (Albacete), 1310 m, 30.iv.2022, 24, leg. EJ. Ortiz- Sanchez, FJOS; Granada, Sierra de Baza, Prados del Rey, 2000 m, 19.vi.2021, 19, leg. T.J. Wood, TJWC; Granada, Sierra Nevada, Capileira to La Cebadilla, 1500 m, 8.vi.2021, 19, leg. TJ. Wood, TJ WC; Granada, Sierra Nevada, Jardin Botanico Hoya de Pedraza environs, 1900 m, 29, leg. T.J. Wood, TJ WC; Granada, Venta de los Ala- zores, 25.v.1982, 19, leg. R. Leys, RMNH; La Cabrilla, Sierra Cazorla (Jaén), 1600 m, 3.vi.2022, leg. C.M. Hererra, CMHC; Malaga, PN Sierra de las Nieves, mountain peak S of Pinsapo Escalereta, 1600 m, 30.v.2021, 39, leg. TJ. Wood, TJWC; Sierra Cazorla, Puerto Llano, 1800 m, 11.vi.2022, 19, leg. J. Valverde, CMHC; Puerto Cru- cetillas, Riépar (Albacete), 1340 m, 30.iv.2022, 12, leg. FJ. Ortiz-Sanchez, FJOS; Sierra de Cazorla, Nava de las Correhuelas, 1.v.2021, 1, leg. C.M. Hererra, CMHC. Subgenus Notandrena Pérez, 1890 The definition of this subgenus was expanded by Pisanty et al. (2022b) to include Carandrena Warncke, 1968 (type species Andrena aerinifrons Dours, 1873). It is im- portant to note that several species formerly placed within the Cavandrena fall else- where into an undescribed subgenus, but these are desert-living species, none of which are present in Iberia. In the context of this revision, Carandrena is treated as a strict synonym of Notandrena. Andrena (incertae sedis) urdula Warncke, 1965 was described from Greece and is a rare and poorly understood taxon known only from a small number of specimens. It is reliably known only from Greece (type series), Spain (central Spain), and Morocco, as the distribution map presented by Gusenleitner and Schwarz (2002: 1201) actu- ally refers to A. (Notandrena) ungeri Mavromoustakis 1952; the distribution map for A, ungeri (p. 1200) is actually the distribution map for A. urdula. A barcoded specimen The genus Andrena in the Iberian Peninsula 291 from south of Madrid [WPATW351-21] does not clearly fall into the Notandrena group (Fig. 23); its true placement is unclear because it does not have any strong char- acters that lead to obvious affinities with specific groups. It is best treated as incertae sedis until more genetic data are available. Warncke described two similar taxa in the group of small metallic green Notan- drena, A. (Notandrena) reperta Warncke, 1974 and A. (Notandrena) reperta varuga Warncke, 1975. Warncke actually described A. reperta as a subspecies of A. varuga, but due to the order of publication, A. reperta has priority. Both species can be recognised because the hind tibial spur is apically curved, though this character is more pro- nounced in Iberian specimens. Genetically, specimens of A. reperta from Morocco and A, reperta varuga from central Spain formed a clade with a specimen of A. (Notandrena) nigroviridula Dours, 1873 from Morocco. All three taxa were well separated; A. reperta and A. reperta varuga by 12.84%, A. reperta and A. nigroviridula by 11.67%, and WPATW348-21_Andrena_relata_Spain_Madrona 96 WPATW351-21_Andrena_urdula_Spain_Ucles urdula WPATW786-22_Andrena_urdula_Morocco_Laanoucer 100 ANDIL253-22_Andrena_ungeri_Israel_Mishmar_Ayalon B ungeri ANDIL252-22_Andrena_ungeri_Israel_Mishmar_Ayalon IBIHM496-21_Andrena_juliana_Spain_Laguna_Dulce juliana WPATWO055-21_Andrena_foeniculae_Portugal_Santana foen iculae 100 HM401021_Andrena_pallitarsis_ltaly_Lombardy < ; pallitarsis HM401022_Andrena_pallitarsis_ltaly_Lombardy WPATW047-21_Andrena_griseobalteata_Bulgaria_Golica griseobalteata 79 90 WPATW451-22_Andrena_reperta_TJW_639_Morocco_Fes-Meknes Bi repertas. str. WPATW619-22_Andrena_nigroviridula_TJW_807_Morocco_Fes-Meknes i nigroviridula 90 WPATW344-21_Andrena_reperta_varuga_Spain_Chinchon | | varuga stat. nov. 99 IBIHM1034-22 Andrena_aerinifrons_Spain_Toledo IBIHM1228-22_Andrena_aerinifrons_Portugal_Algarve 95 85 WPATW659-22_Andrena_aerinifrons_TJW_847_Morocco_Guelmim-Oued_Noun_Region [mm aerinifrons S.|. ANDPHO13-21_Andrena_aerinifrons_levantina_Israel_Lakhish 99 ANDIL214-22_Andrena_aerinifrons_levantina_Israel_Lakhish 98 100 IBIHM1226-22_Andrena_bellidis_Portugal_Baixo_Alentejo b llidi ~| eras IBIHM1225-22_Andrena_bellidis_Portugal_Baixo_Alentejo 85 WPATW251-21_Andrena_ranunculi_Spain_Benacoaz . ranunculi 100 WPATW313-21_Andrena_ranunculi_Spain_Zorra 100 HQ954749_Andrena_chrysosceles_Germany_Bavaria chrysosceles JN262171_Andrena_chrysosceles_Germany_Bavaria 96 | WPATW200-21_Andrena_fulvicomis_Spain_Aznalcazar ¥ e - fulvicornis #1 WPATW259-21_Andrena_fulvicornis_Spain_Laguna_Dulce 92 ANDPHO20-21_Andrena_nitidiuscula_Czech_Republic_Hradec_Kralove KJ837911_Andrena_nitidiuscula_Germany_Thuringia . ‘di | nitidiuscula 98 90) KJ838927_Andrena_nitidiuscula_Germany_Thuringia HQ954765_Andrena_nitidiuscula_Germany_Brandenburg ANDILO19-22_Andrena_fulvicornis_lIsrael_Nashut | 97 |; FBAPD917-11_Andrena_fulvicornis_Germany_Baden_Wuerttemberg KJ837677_Andrena_fulvicornis_Turkey_Burdur KJ839658_Andrena_fulvicornis_Germany_Brandenburg a Ee fulvicornis #2 KJ837634_Andrena_fulvicomis_Germany_Brandenburg KJ837732_Andrena_fulvicornis_Turkey_Mersin KJ839481_Andrena_fulvicoris_Germany_Brandenburg HM376237_Andrena_fulvicornis_Germany_Brandenburg 0.03 Figure 23. Phylogenetic tree (maximum likelihood) of Andrena subgenus Notandrena Pérez, 1890 based on the mitochondrial COI gene. Andrena (incertae sedis) relata Warncke, 1975 is used as an outgroup. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted). 292 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) A, reperta varuga and A. nigroviridula by 10.12% (Fig. 23). Moreover, A. reperta and A, reperta varuga do not form a monophyletic clade, being rendered paraphyletic by A. nigroviridula. On this basis, A. varuga stat. nov. is raised to species status. Andrena varuga is endemic to Spain, and A. reperta is found in Morocco, Algeria, and Tunisia. Dours (1873) described A. aerinifrons from Algeria, and A. aerinifrons levantina Hedicke, 1938 was later described from the Levant. Barcode analysis shows that speci- mens from Israel, Morocco, Portugal, and Spain show high average intraspecific varia- tion of 8.64% (range 0.00—11.67%). The Moroccan specimen is intermediate between the Iberian and Levantine sequences, being separated from the Iberian sequences by an average genetic distance of 9.92% (range 9.73—10.12%) and from the Levantine se- quences by 10.12% (range 10.12—10.12%). Given that these sequences form a mono- phyletic clade, a broad interpretation of A. aerinifrons is taken here; a neotype can be designated for A. aerinifrons at a later date when the situation becomes clearer, as Dours’ collection was destroyed in a fire and is not available for study (see below). Andrena (Notandrena) fulvicornis Schenck, 1861 has been recognised as distinct from A. (Notandrena) nitidiuscula Schenck, 1853 (Schmid-Egger and Doczkal 1995; Schmid-Egger and Scheuchl 1997; Schwenninger 2013; Bénon and Praz 2016). Barcode analysis shows that specimens from Germany, Israel, and Turkey are indeed clearly distinct from A. nitidiuscula, not forming a sister clade and showing average separation of 3.79% (range 3.50—3.89%), with bootstrap support of 96. However, sequences of putative A. fulvicornis from Spain fall into a clade of A. nitidiuscula and A, (Notandrena) chrysosceles (Kirby, 1802). Spanish specimens are separated from A. nitidiuscula by 1.95% (range 1.95-1.95%), from A. chrysosceles by 4.67% (range 4,28—5.06%), and from the other clade of A. fulvicornis by 4.18% (range 3.89-4.28%). This Iberian ‘fulvicornis’ is also well-supported, with bootstrap support of 96. At the present time, no taxonomic action can be taken until more genetic data are avail- able. Morphologically, Iberian ‘fulvicornis do not appear to be different from Central European/Turkish/Levantine specimens. Additional sequences from Iberia and France are needed; this barcode difference may be the result of an isolated population in the Iberian glacial refugium. Subgenus Plastandrena Hedicke, 1933 Pisanty et al. (2022b) synonymised the subgenus Agandrena with Plastandrena, and this broader definition is used here. Some taxonomic problems within this subge- nus were discussed by Wood (2023a), who found support for the species-status of A, (Plastandrena) nigrospina Thomson, 1872. Standard barcodes have limited utility for this group for unclear reasons, so care must be taken when interpreting results, as morphologically distinct taxa can form unclear clades, for example A. (Plastandrena) tibialis (Kirby, 1802). This result was reproduced here (Fig. 9), and no further discus- sion or action is taken; in an Iberian context, members of this subgenus appear to be identifiable using barcodes, but work is needed to revise this group at a Palaearctic scale using more powerful genetic techniques. The genus Andrena in the Iberian Peninsula 293 Andrena (Plastandrena) pilipes Fabricius, 1781 and Andrena (Plastandrena) nigrospina Thomson, 1872 Ortiz-Sanchez et al. (2022) reported A. nigrospina as new for Spain from the Sierra de Cazorla based on male specimens with their distinctive genital capsule. Wood (2023a) further reported two barcoded specimens from Spain (Sistema Central and Sistema Ibérico) belonging to A. nigrospina. This genetic result was further supported with the addition of more sequences from Morocco (Fig. 9), revealing that A. nigrospina is also present in the Middle Atlas. The A. nigrospina clade maintained a low intraspe- cific genetic distance of 0.41% (range 0.00—1.06%) from Morocco to Kyrgyzstan, and average separation from A. pilipes was 1.63% (range 1.06—2.35%). As this difference is still small in absolute terms, bootstrap support was moderate, with 77 for A. pilipes and 84 for A. nigrospina, but both are considered to be distinct species. Females can- not currently be consistently separated morphologically, so no characters are given in the identification key. In Iberia, A. nigrospina is newly reported for northern Portugal from close to the Peneda-Gerés National Park. The species is likely to be restricted to cooler parts of northern, central, and eastern Iberia where it will probably be found only in mountainous regions, or at least at elevation. More sampling and genetic study is required to define its exact range limits. Material examined. Andrena nigrospina: PortucaL: Minho, Ruivaes, N103, 12522019; 1s ADs leg. Wood, TJWC; Spain: Cuenca, Pajaroncillo, 3 km SW, Arroyo de Pefia Quebrada, 26.vi.2021, 49, leg. TJ. Wood, TJWC (barcoded); Guadalajara, Bustares, 2 km N, Alto Rey, 1780 m, 19, 9.vii.2021, leg. TJ. Wood, TJWC (barcoded). Andrena (Plastandrena) agilissima Scopoli, 1770 and Andrena (Plastandrena) asperrima Pérez, 1895 Andrena agilissima is a widespread West Palaearctic species that is well-known in Cen- tral and Southern Europe to north-western Africa. In contrast, A. asperrima is much less well known, having a more Mediterranean distribution in France, Spain, Morocco, Algeria, and Tunisia. Unlike A. agilissima, A. asperrima is bivoltine and is exception- ally variable in the density, size, and strength of the integumental punctation. In the female sex, the typical form has strong and dense punctures on the terga, allowing easy separation from A. agilissima in which the terga have small and subtle punctures. However, many specimens of A. asperrima can be found which have greatly reduced tergal punctation and which are therefore extremely similar to A. agilissima; they can be separated by the smaller body size and the sparser punctation of the scutum. ‘This sparsely punctate form is more common in the south-west and was described from Morocco by Warncke as A. asperrima alascana Warncke, 1974. Overall, the two species are clearly separable by their genital capsules in the male sex. Because of this variation, it is important to ensure that Iberian material is conspe- cific with North African material, since the oldest names of Pérez (1895) were described based on North African material. Warncke (1967) designated a male lectotype for 294 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) A. asperrima using a specimen from Biskra from the collection of Pic (Fig. 24A, B). This is unjustified, as it is not part of the original syntypic series, and Warncke’s desig- nation is here rejected. In Pérez’s catalogue, under entry “1030 Andrena trachodes J.P’ [an unpublished name; Pérez sometimes changed his mind and therefore a different name to the published name can be present in the catalogue] Pérez writes that the spe- cies comes from Constantine [in Algeria], drawing comparison with A. flessae Panzer, 1805 (= A. agilissima (Scopoli, 1770)) and arguing it differs by the stronger punctation of the metasoma. Inspection of the Pérez collection shows the presence of a female Figure 24. Andrena (Plastandrena) asperrima Pérez, 1895, false male lectotype A label details B profile; true female lectotype C label details D profile E scutum, dorsal view F terga, dorsal view. The genus Andrena in the Iberian Peninsula 295 specimen from Constantine which bears the label ‘1030’, the code used by Pérez for this species. This specimen is designated as a new lectotype (Fig. 24C—F); it conforms to the concept of A. asperrima used by subsequent authors. Andrena atricapilla Pérez, 1895 was also described from Algeria, but only in the male sex. Warncke (1967) listed this taxon as a synonym of A. asperrima, though he did not mention a lectotype. A male specimen labelled as atricapilla J.P. in the handwriting of Pérez is separated in the MNHN collection and labelled as a lectotype, probably by Teunissen as one of his determination labels is present. This lectotype designation was never published, and so it is here designated as a lectotype (Fig. 25). The synonymy with A. asperrima is maintained, as the genital capsule is typical for the species concept. Genetic analysis of specimens of A. agilissima and A. asperrima from Croatia, Morocco, Portugal, and Spain showed two clear clades (Fig. 9). One clade contained only specimens determined as A. agilissima from Croatia, Portugal, and Spain. These sequences showed average intraspecific variation of 0.07% (range 0.00-—0.25%), and were strongly separated from the second clade by an average genetic distance of 9.33% (range 8.74—10.20%). This second clade contained specimens identified as A. asperrima from Spain and Morocco, but also three specimens identified as A. agilissima from the Middle Atlas. There can be no doubt as to the identity of these specimens, as two are males, and their genital capsule is that of A. agilissima and not A. asperrima. However, the genetic differentiation within this clade is very low at an average of 0.42% (range 0.00-0.91%). As for A. bimaculata and A. tibialis, there is clearly genetic complexity here, as two species with distinct genital capsules have barcodes that form intermixed clades. No taxonomic action is taken on the basis of these barcoding results. However, what can be concluded is that Iberian material of A. asperrima is conspe- cific with North African material, including the weakly punctate form that dominates in Morocco. In combination with the new lectotype designations, the invariant genital capsule, and these genetic results, the concept of Warncke (1967; 1974) and Gusen- leitner and Schwarz (2002) regarding A. asperrima is maintained. ‘The full synonymy is given below. Andrena (Plastandrena) asperrima Pérez, 1895 Andrena (Plastandrena) asperrima Pérez, 1895: 33, ed [Algeria, lectotype by present designation: MNHN]. Andrena (Plastandrena) atricapilla Pérez, 1895: 33, J [Algeria, lectotype by present designation: MNHN]. Andrena (Plastandrena) flessae var. elcheensis Friese, 1922: 211, 9 [Spain: ZMHB, not examined]. Andrena (Plastandrena) hemicyanea Cockerell, 1930: 112, 9 [Tunisia: type lost?] Andrena (Plastandrena) asperrima alascana Warncke, 1974: 36, 93 [Morocco: OOLM, examined]. Distribution. Spain, France, Morocco, Algeria, Tunisia. 296 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Figure 25. Andrena (Plastandrena) atricapilla Pérez, 1895, male lectotype A label details B profile C terga, dorsal view D genital capsule. Material examined. ALGERIA: Constantine [36.3645°N, 6.6409°E], 19, MNHN (lectotype of A. asperrima, by present designation; Fig. 24C—F); Biskra, v.1885, eae leg. Blause, MNHN (false lectotype of A. asperrima; Fig. 24A, B); Biskra [34.8600°N, 5.6995°E], 14, MNHN (lectotype of A. atricapilla, by present designation; Fig. 25); Morocco: Tizi-n-Talrhemt nr. Midelt [32.6821°N, -2.9344°E], 1900 m, 1.vi.1968, leg. M.A. Lieftinck, OOLM (holotype of A. asperrima alascana). Subgenus Poecilandrena Hedicke, 1933 This subgenus is strongly polyphyletic (Pisanty et al. (2022b), but these classification issues relate to eastern taxa; in Iberia, the species belong to Poecilandrena s. str. Warncke (1976) listed only one Poecilandrena species in Iberia, A. (Poecilandrena) labiata Fabricius, 1781. Ceballos (1956) and Pérez-Inigo (1984) also listed A. (Poecilandrena) potentillae Panzer, 1809, though this species was not listed as present by Warncke and his distribution maps (Gusenleitner and Schwarz 2002) indicate that this species has a western range limit in central France. Collection of material from the Sierra Nevada produced red-marked Poecilandrena females [WPATW281-21] that morphologically resemble A. potentillae in the reduced The genus Andrena in the Iberian Peninsula 297 punctation density at the edge of the clypeus. No ‘potentillae’ males with their distinc- tive genital capsule could be found. A female sequence clearly fell into a clade with an A. labiata sequence from Belgium, the two specimens separated by 2.87% (Fig. 1). The four A. /abiata sequences from Belgium, Portugal, and Spain showed average intraspecific variation of 2.52% (range 0.78—-3.92%), much lower than the average separation from A, potentillae sequences from Belgium, France, Germany, and Hungary of 11.81% (range 11.23—12.27%). The position is taken here that true A. potentillae is not present in Iberia, even though some females of A. /abiata can resemble it morphologically. These speci- mens conform to the subspecific concept of A. labiata bellina Warncke, 1967 that was described from Madrid. Warncke (1967) noted that this form had weaker shagreenation and finer punctation than the nominate form, and I believe that this is the source of the confusion and the erroneous reports of A. potentillae in Iberia. Andrena potentillae is there- fore not included in the identification key as typical female characters that can be used to separate the two species in Central Europe do not appear to work universally in Iberia. Finally, Ortiz-Sanchez (2011, 2020) lists A. (Poecilandrena) viridescens Viereck, 1916 as part of the Spanish fauna. The distribution maps of Warncke (Gusenleitner and Schwarz 2002) indicate the possible presence of this species on the Spanish side of the Pyrenees. I have not examined any specimens of A. viridescens from Iberia, but its presence in the Pyrenees is plausible and so it is retained on the list. Several Andrena taxa with apparent range limits in the Pyrenees have recently been confirmed to occur in northern Spain (e.g. A. (Ulandrena) polita Smith, 1847 and A. (Micrandrena) nanula Nylander, 1848, see Alvarez Fidalgo et al. 2022), and so additional searches may well confirm the presence of A. viridescens as well. Subgenus Simandrena Pérez, 1890 Barcode analysis returned Simandrena as paraphyletic (Fig. 4), but this means very lit- tle, as the subgenus is very well characterised morphologically and genetically based on UCE analysis (Pisanty et al. 2022b). No major changes in Simandrena taxonomy are made here, but there are a number of issues to discuss. Warncke (1967) described A. (Simandrena) combinata crudelis Warncke, 1967 from Spain. Comparison of sequences shows that Iberian material is only weakly dif- ferentiated, being separated from A. (Simandrena) combinata (Christ, 1791) sequences from Germany by an average genetic distance of 1.12% (range 1.06—1.29%). Based on this evidence, a subspecific status is not justified. Andrena (Simandrena) vetula Lepeletier, 1841 was recently placed in the Siman- drena, as its unusual male morphology had led to confused previous placement (Pisanty et al. 2022b). Genetically there was a large difference between Iberian and North African sequences, separated by an average genetic distance of 6.33% (range 6.08-6.57%). Andrena vetula has an enormous range, from Morocco and Iberia to Central Asia (Wood and Monfared 2022). Sequences are required over this range be- fore any taxonomic changes can be made, as morphologically there are no obvious differences between Iberian and North African specimens. 298 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) The species pair of A. (Simandrena) confinis Stéckhert, 1930 and A. (Simandrena) congruens Schmiedeknecht, 1884 continues to pose problems. Warncke (1967) treated A. confinis as a synonym of A. congruens, but others have not followed this interpreta- tion. The two taxa can be separated morphologically in Central Europe (Schmid-Egger and Scheuchl 1997), and there are also ecological differences with A. confinis prefer- ring cooler northern climates and A. congruens preferring warmer and drier southern climates, with overlap in Central Europe. However, in southern Europe the situation is complex and confused. Wood et al. (2021) added A. confinis to the Iberian list on the basis of a specimen from the Picos de Europa Mountains. Genetically, this specimen clusters with A. confinis sequences from Germany. However, sequences from specimens provisionally identified as A. congruens from northern Portugal as well as a specimen of A. congruens from Bulgaria render A. congruens paraphyletic. The genetic distance between all sequences is low. The Portuguese specimens are strongly divergent mor- phologically from the Spanish A. confinis specimen, with dense and obvious tergal punctation compared to terga which are almost impunctate, but the average genetic differentiation is low at 0.88% (range 0.60—1.15%). No major taxonomic decision is made here; much more genetic data are required for a dedicated study focused on this group. Both nominal taxa are included in the identification key. Andrena (Simandrena) cilissaeformis Pérez, 1895, sp. resurr. Andrena (Simandrena) cilissaeformis Pérez, 1895: 42, 2 [Spain, lectotype by present designation: MNHN] Andrena breviscopa auctorum. Remarks. Andrena breviscopa Pérez, 1895 was described in the female and male sexes from North Africa. Warncke’s treatment of A. breviscopa is curious, because he desig- nated a lectotype (Fig. 26) and listed the taxon as a synonym of A. numida Lepeletier, 1841 (Warncke 1967). However, just a few years later he listed A. breviscopa as a valid taxon in the subgenus Simandrena (Warncke 1974). Examination of the female lecto- type designated by Warncke shows that his original synonymy was correct; the taxon is clearly not a Simandrena, and is indeed a synonym of A. numida. The use of the name A. breviscopa to apply to the taxon present in Spain, Morocco, and Algeria is therefore incorrect. The correct name is A. cilissaeformis Pérez, 1895 sp. resurr. Andrena cilissaeformis was described from Spain, not Algeria as stated in Warncke (1967) and Gusenleitner and Schwarz (2002). This is because Pérez (1895) does not directly state the type locality (or indeed, any information about the collecting locality of any of the species described in this work), but this information is included in his personal catalogue. Warncke (1967) did not examine material of A. cilissaeformis, stating that whilst the label was present, material was missing. Examination of material in the MNHN has located a specimen labelled with ‘Esp’ [Espagne = Spain] in Pérez’s handwriting (Fig. 27). This specimen was separated by Teunissen, but he never pub- lished this information. This specimen is badly damaged, but it is a Simandrena and The genus Andrena in the Iberian Peninsula 299 2] niet chp det .H.Teunissen Figure 26. Andrena (incertae sedis) breviscopa Pérez, 1895, female lectotype A label details B profile C face, frontal view D terga, dorsal view. conforms to A. breviscopa auctorum sensu Warncke (1974) and subsequent publica- tions. It is designated as a lectotype to fix the name on the Iberian population. Andrena breviscopa is returned to its original synonymy with A. numida. Distribution. Spain, Morocco, Algeria. Material examined. Andrena breviscopa: AuGeria: Ghardaia [32.5047°N, 3.6419°E], 12, MNHN (lectotype; Fig. 26); (Andrena cilissaeformis): SPAIN: no collec- tion information, 12, MNHN (lectotype by present designation; Fig. 27). Subgenus Taeniandrena Hedicke, 1933 Large parts of this subgenus have been revised recently by Wood et al. (2021), Praz et al. (2022), Wood (2022), and Wood and Ortiz-Sanchez (2022). There is relatively little new information to present here, other than to revise the status of A. (Zaeniandrena) poupillieri Dours, 1872, and so a reduced phylogenetic tree is presented given the results presented in these previous publications (Fig. 28). However, it is clear that there is still unfinished work to be completed in this subgenus in an Iberian context. Specifically, Praz et al. (2022) identified an unclear lineage “sp. nov. 2” from northern Portugal. This specimen is a male that has a genital capsule that diverges from any known Iberian 300 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Figure 27. Andrena (Simandrena) cilissaeformis Pérez, 1895; female lectotype A label details B profile C face, frontal view D terga, dorsal view. species. Furthermore, I have an unbarcoded male from Guadalajara province that also possesses a genital capsule that does not match any known species. These may represent additional undescribed Taeniandrena species endemic to the peninsula. No further ac- tion is taken until more genetic samples are available; it is extremely likely that addi- tional barcoding will discover more diversity in this challenging and speciose subgenus. Andrena (Taeniandrena) poupillieri Dours, 1872 Two further issues require discussion. ‘The first is the identity of A. poupillieri. Praz et al. (2022) identified two potential mitochondrial lineages that could correspond to this species. Additional sampling in Morocco has clarified the situation; poupillieri 1 sensu Praz et al. (2022) corresponds to A. (Taeniandrena) gregaria Warncke, 1974 and poupillieri 2 sensu Praz et al. (2022) corresponds to the concept of A. poupillieri used by Warncke. As the type of A. poupillieri is lost, it is beneficial to designate a neotype in order to fix the concept of this species in line with the existing literature; a specimen from Algeria is chosen, as this is the original Jocus typicus. As A. gregaria does not oc- cur in Iberia, it is not included in the phylogenetic tree presented here; it will be dealt with in the upcoming North African Andrena revision. The species A. poupillieri is 0.02 The genus Andrena in the Iberian Peninsula 301 KJ839310_Andrena_lathyri_Thuringia HYMAA325-22_Andrena_ovata_Greece_Arkadia 96 HYMAA324-22 Andrena_ovata_Switzerland_Mendrisio WPATW160-21_Andrena_ovata_Spain_Madrona ovata HYMAA326-22_Andrena_ovata_France_Marseille HYMAA328-22_Andrena_ovata_France_Villefranque MIRADA na 2 1 Apearie: Reagee Spain Haynceserc fi b eno i st i 99 IBIHM1038-22_Andrena_benoisti_Spain_Hoyocasero ANDIL298-22_Andrena_russula_Israel_Hermon HYMAA236-21_Andrena_russula_Cyprus 96 ANDIL289-22_Andrena_russula_lsrael_Karmei_Yosef ANDIL369-22_Andrena_russula_Israel_Hermon HYMAA237-21_Andrena_russula_Morocco_Oukaimeden BEEEE020-15_Andrena_russula_United_Kingdom_Shropshire MZ623867_Andrena_russula_Finland_Nylandia HYMAA336-22_Andrena_russula_France_Porte_Puymorens KJ839751_Andrena_russula_ltaly_Aosta WPATW125-21_Andrena_russula_Spain_Lupiana HYMAA335-22_Andrena_russula_Italy_Marche IBIHM945-21_Andrena_russula_Portugal_Castelo_Branco IBIHM529-21_Andrena_russula_Portugal_Santarem HYMAA337-22_Andrena_russula_Algeria_Ait_Hassem HYMAA233-21_Andrena_russula_Portugal_Serra_do_Geres IBIHM978-21_Andrena_russula_Portugal_Castelo_Branco HYMAA267-22_Andrena_afzeliella_United_Kingdom_Cornwall 96 HYMAA269-22_Andrena_afzeliella_Switzerland_Leuk HYMAA272-22_Andrena_afzeliella_France_Evisa HYMAA270-22_Andrena_afzeliella_Italy_Abruzzo 100 HYMAA266-22_Andrena_afzeliella_Switzerland_Cormondreche KJ837459_Andrena_afzeliella_Germany_Thuringia ANDIL309-22_Andrena_afzeliella_lsrael_Ein_Tina 78 WPATW9336-21_Andrena_afzeliella_Spain_Brieva IBIHM530-21_Andrena_russula_cryptic_Portugal_Cerro_Miguel IBIHM528-21_Andrena_russula_cryptic_Portugal_Benagil IBIHM527-21_Andrena_russula_cryptic_Portugal_Vale_da_Telha IBIHM526-21_Andrena_russula_cryptic_Portugal_Cercal WPATW641-22_Andrena_russula_TJW_829_Morocco_Fes-Meknes 93 WPATW642-22_Andrena_russula_TJW_830_Morocco_Fes-Meknes WPATW6813-22_Andrena_poupillieri_Morocco_Oukaimeden WPATW@805-22_Andrena_poupillieri_Morocco_Aroumd 100 WPATW810-22_ Andrena_poupillieri_incana_Spain_Mallorca WPATW524-22_Andrena_poupillieri_TJW_712_Morocco_Fes-Meknes WPATW811-22_Andrena_poupillieri_Oukaimeden HYMAA322-22_Andrena_poupillieri_2_ Algeria WPATW806-22_Andrena_poupillieri_Morocco_Agelmouss WPATW528-22_Andrena_poupillieri_TJW_716_Morocco_Fes-Meknes WPATW812-22_Andrena_poupillieri_Morocco_Tizi_n_Tichka WPATW527-22_Andrena_poupillieri_TJW_715_Morocco_Beni WPATW526-22_Andrena_poupillieri_TJW_714_Morocco_Fes-Meknes | WPATW525-22_Andrena_poupillieri_TJW_713_Morocco_Fes-Meknes WPATW519-22 Andrena_poupillieri_TJW_707_Morocco_Ouarzazarte WPATW520-22_Andrena_poupillieri_TJW_708_Morocco_Fes-Meknes FBAPC317-10_Andrena_wilkella_Germany_Bavaria HYMAA351-22_Andrena_wilkella_France_Corsica russula #1 afzeliella russula #2 russula #3 poupillieri 95 IBIHM547-21_Andrena_wilkella_Portugal_Avecao_do_Cabo Wi lke ! ! a BEEEE173-15_Andrena_wilkella_United_Kingdom_London WPATW299-21_Andrena_aff_intermedia_Spain_Sierra_Baza 91 WPATW284-21_Andrena_aff_intermedia_Spain_Tocon_de_Quentar g elr iae-g rou p 79 WPATW292-21_Andrena_contracta_Spain_Sierra_Nevada IBIHM1045-22 Andrena_ovatula_Spain_Yunquera 92 IBIHM996-21_Andrena_ovatula_Portugal_Castelo_Branco B ovetue HYMAA254-21_Andrena_ovatula_Portugal_Serra_do_Geres Figure 28. Phylogenetic tree (maximum likelihood) of Andrena subgenus Taeniandrena Hedicke, 1933 based on the mitochondrial COI gene. Andrena (Taeniandrena) lathyri Alfken, 1900 is used as an out- group. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted). commonly encountered in Morocco, and is clearly identifiable from barcodes, forming a clade with bootstrap support of 100. Separation of females from A. (Zaeniandrena) ovatula (Kirby, 1802) can be made by the tergal punctation, this being much stronger in A. ovatula. Generally, the two taxa are well-separated by an average genetic distance of 6.12% (range 5.86—7.79%). Males of A. poupillieri can be recognised based on the genital capsule, in which the gonostyli are apically produced into acute points. 302 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Warncke changed his mind about the status of A. poupillieri — in Warncke (1967), he lists the taxon as A. ovatula poupillieri, but then in Warncke (1975a) he described A, poupillieri incana from the Balearic islands, listing A. poupillieri s. str. from southern Iberia (Warncke 1976). In his distribution maps (Gusenleitner and Schwarz 2002), he gives a distribution of southern Iberia, the Balearic islands, north-western Africa, Lib- ya, and Crete. Records from Crete are likely to refer to unrecognised A. (Taeniandrena) ovata Schenck, 1853 (Wood, unpublished data). Examination of male specimens from the extreme south of Spain shows that A. poupillieri is present based on the distinctive genital capsule, though as females cannot be separated morphologically from_.A. ovatula the exact range of the two species is unclear. Andrena ovatula reaches at least as far south at the Sierra de las Nieves in Malaga province [I[BIHM1045-22], though this far south it may be restricted to mountainous areas whereas the two examined male A. poupillieri specimens come from the coast. More collection is required. For now, A. poupillieri is considered to be a rare and little-collected taxon in Iberia, probably restricted to the coast in the south and south-east of the peninsula. A single sequence for A. p. incana was available from Mallorca that unambiguously nests within the North African A, poupillieri sequences; it is identical to 12 of the North African sequences, and differs from the thirteenth by just 0.15%. As such, A. poupillieri including A. p. incana shows extremely low intraspecific variation of 0.03% (range 0.00—0.15%), and subspecific status is unnecessary for the population on the Balearic islands. Material examined. Spain: Malaga, Estepona, 24.iii.1986, 1, leg. J. van Oosterhout, RMNH,;; Malaga, San Pedro de Alcantara, 15.iii.1986, 14, leg. C. v. Achterberg, RMNH. Andrena (Taeniandrena) russula Lepeletier, 1841 Praz et al. (2022) synonymised A. (Tazeniandrena) similis Smith, 1849 with A. russula, taking a broad, pan-Mediterranean approach. In an Iberian context, sequences from central and northern Iberia clearly fall into a broad A. russula clade along with se- quences from Morocco to Cyprus and Israel and north to the United Kingdom and Finland (Fig. 28). However, in the extreme south-west of Iberia in southern Portugal, sequences from specimens that are morphologically indistinguishable from A. russula from the rest of Iberia form a distinct clade that falls closest to A. (Jaeniandrena) afzeliella (Kirby, 1802). These specimens come from the south of Baixo Alentejo (Cer- cal) and the Algarve (Aljezur, Benagil, Moncarapacho). These sequences are consist- ently separated from the broad A. russula clade by 5.50% (range 3.68-11.11%). Two specimens from Morocco that correspond to A. russula form a third clade of A. russula s.l. At the present time, no taxonomic action is taken; these three lineages may repre- sent different isolated populations. More powerful genetic techniques are required to resolve this problem, as for Iberian members of the ge/riae-group (see Praz et al. 2022). Andrena (Taeniandrena) gredana Warncke, 1975 Wood et al. (2021) elevated A. gredana to species status, and gave a distribution across the Sistema Central, central and northern Portugal, and northern Spain across to the The genus Andrena in the Iberian Peninsula 303 Pyrenees in the province of Huesca. Examination of material from the Hautes-Pyrénées in France approximately 80 km north-east of the Huesca site revealed the presence of A. gredana in France. ‘The species is therefore not endemic to the Iberian Peninsula, though it is likely to have a French range restricted to high altitude sites in the Pyrenees. Material examined. France: Hautes-Pyrénées, Eget Cité, 4.v.2017, 16, leg. R. Rudelle, R. Rudelle Colln.; Spamn: Huesca, San Juan de la Pefa, 14.v.1995, 12, leg. H. & J.E. Wiering, RMNH (see also records in Wood et al. 2021). Subgenus Truncandrena Warncke, 1968 This subgenus contains species that often vary extensively in the colouration of their pubescence, sometimes display minimal variation in structural characteristics in the female sex, and can sometimes only be reliably identified in the male sex. These iden- tification difficulties have led to a large number of subspecific concepts in the litera- ture, the integrity of which must be examined using molecular data. There are a num- ber of taxonomic changes to make which affect the Iberian and more broadly West Mediterranean fauna. Andrena (Truncandrena) doursana Dufour, 1853 sensu lato This nominal species is highly variable across its range which was previously consid- ered to be from Morocco and Iberia to Turkey and the Levant. Andrena doursana was originally described from Algeria, and Warncke (1975a, 1975b, 1980) described the subspecies A. d. citreola Warncke, 1975 from Spain, A. d. agadira Warncke, 1980 from southern Morocco, A. d. bengasia Warncke, 1980 from Libya, and A. d. mizorhina Warncke, 1975 from Turkey. Pisanty et al. (2022b) elevated the eastern subspecies A. (Truncandrena) doursana mizorhina to species status, leaving populations in Iberia and north-western Africa to Libya. The differentiation between these remaining subspecies relies on the colour of the female pubescence, as there are no structural differences in the males; indeed, the sub- species A. d. agadira and A. d. bengasia were described only from the female sex. Andrena d. citreola is bright, and has predominantly white hairs on the face and a light brown terminal fringe with scattered white hairs laterally. Andrena d. agadira is much darker, with dark facial hairs and a uniformly dark terminal fringe. Wood et al. (2020b) found the undescribed female of a similar species endemic to Morocco, A. (Truncandrena) alchata Warncke, 1974, which is structurally very similar to the female of A. d. agadira, but these authors were unable to conclude if the two taxa were synonymous. Analysis of barcodes from southern and northern Morocco and Iberia shows that female specimens identified as A. doursana s. str., A. d. agadira, and A. d. citreola did not form differentiated clades (Fig. 29). Light forms from southern Portugal [IBIHM 1223-22] and the Moroccan Anti-Atlas near Tazenakht [WPATW505-22] were separated, and dark forms from south-western Morocco near Guelmim [WPATW506-22] and Tiznit [WPATW503-22] were clustered closer to the light specimen from Tazenakht. The light specimen from Portugal was clustered close to 304 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) WPATW156-21_Andrena_limbata_Spain_La Losa 98 — WPATW239-21_Andrena_spec_nov_Spain_Sierra_Nieves fl ghisbaini sp. Nov. IBIHM545-21_Andrena_villipes_Portugal_Carrapateira giida = alchata 99 MT466582_Andrena_alchata_Morocco_Laanoucer WPATW503-22_Andrena_doursana_agadira_TJW_691_Morocco_Souss-Massa_Region 78 92 88 WPATW5S04-22_Andrena_doursana_TJW_692_Morocco_Souss-Massa_Region WPATW505-22_Andrena_doursana_citreola_TJW_693_Morocco_Tazenakht WPATW506-22_ Andrena_doursana_TJW_694_Morocco_Guelmim-Oued_Noun_Region 78 WPATW507-22_Andrena_doursana_agadira_TJW_695_Morocco_Guelmim-Qued_Noun_Region do ursana WPATW502-22_Andrena_doursana_TJW_690_Morocco_Souss-Massa_Region WPATW508-22_Andrena_doursana_TJW_696_Morocco_Ouarzazarte IBIHM1223-22 Andrena_doursana_citreola_Baixo_Alentejo 97 ] WPATW509-22 Andrena_doursana_s str TJW_697_Morocco_Fes-Meknes WPATW683-22_Andrena_varia_TJW_871_Morocco_Fes-Meknes Var i a 99 TAU236062_1_Andrena_truncatilabris_Israel_Lakhish L a TAU240823_Andrena_truncatilabris_Israel_Ziv_on truncatila briss. str. IBIHM483-21_Andrena_ferrugineicrus_Portugal_Figueria_Portimao WPATW543-22_Andrena_ferrugineicrus_TJW_731_Morocco_Fes-Meknes. 99 WPATW120-21_Andrena_ferrugineicrus_Spain_Alcolea_Pinar WPATW121-21_Andrena_ferrugineicrus_Spain_Alcolea_Pinar ‘ * 80 WPATW541-22_Andrena_ferrugineicrus_TJW_729_Morocco_Souss-Massa_Region ferrug Inelcrus WPATW542-22_Andrena_ferrugineicrus_TJW_730_Morocco_Draa WPATW647-22_Andrena_ferrugineicrus_TJW_835_Morocco_Tazenakht 72 WPATW629-22_Andrena_ferrugineicrus_TJW_817_Morocco_Fes-Meknes IBIHM1042-22 Andrena_truncatilabris_espanola_Spain_Malaga ———— WPATW216-21_Andrena_truncatilabris_Spain_Cortes_Frontera 100 WPATW212-21_Andrena_truncatilabris_Spain_Ronda WPATW222-21_Andrena_truncatilabris_Spain_Grazalema WPATW206-21_Andrena_aff_truncatilabris_Spain_Castellar_Frontera é / WPATW604-22_Andrena_truncatilabris_nigropilosa_TJW_792_Morocco_Fes-Meknes n Ig r op I ! osa WPATW600-22_Andrena_truncatilabris_nigropilosa_TJW_788_Morocco_Fes-Meknes stat. nov. WPATW605-22_Andrena_truncatilabris_nigropilosa_TJW_793_Morocco_Fes-Meknes WPATW601-22_Andrena_truncatilabris_nigropilosa_TJW_789_Morocco_Fes-Meknes WPATW599-22_Andrena_truncatilabris_nigropilosa_TJW_787_Morocco_Fes-Meknes " WPATW603-22_Andrena_truncatilabris_nigropilosa_TJW_791_Morocco_Fes-Meknes TAU290964_Andrena_urfanella_Israel_Revadim urfanella 96 100 WPATW680-22_Andrena_medeninensis_abunda_TJW_868_Morocco_Fes-Meknes WPATW681-22_Andrena_medeninensis_abunda_TJW_869_Morocco_Fes-Meknes abunda stat. nov. 91 WPATW646-22_Andrena_medeninensis_s_str_TJW_834_Morocco_Tazenakht 86 WPATW432-22_Andrena_medeninensis_tiznita_TJW_620_Morocco_Guelmim-Qued_Noun_Region : ‘i 94 WPATW430-22_Andrena_medeninensis_tiznita_TJW_618_Morocco_Guelmim-Oued_Noun_Region medeninensis Sl. 75 WPATW431-22_Andrena_medeninensis_tiznita_TJW_619 Morocco_Guelmim-Oued_Noun_Region 100 TAU123531_Andrena_tscheki_tritica_Israel_Har_Meron | TAU268801_Andrena_tscheki_tritica_Israel_Har_Haruah 98 | KJ839805_Andrena_tscheki_Germany_Thuringia tschekis.l. KJ836443_Andrena_tscheki_Germany_Thuringia KJ838058_Andrena_tscheki_Germany_Thuringia 0.04 Figure 29. Phylogenetic tree (maximum likelihood) of Andrena subgenus Truncandrena Warncke, 1968 based on the mitochondrial COI gene. Andrena (Limbandrena) limbata Eversmann, 1852 is used as an outgroup. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted). a specimen of A. doursana s. str. from the Middle Atlas near Taza [WPATW509-22]. This pattern strongly suggests simple separation by distance, with geographically closer specimens displaying more similar barcode sequences. Accepting a broad A. doursana species concept, these sequences showed an average intraspecific distance of 2.79% (range 0.29-5.93%). More broadly, this A. doursana clade had bootstrap support of 78, and was sis- ter to the A. alchata sequences generated by Wood et al. (2020b), a species that has a clearly different male morphology. Andrena alchata has bootstrap support of 91, and was The genus Andrena in the Iberian Peninsula 305 separated from A. doursana by an average of 6.29% (range 5.62—8.03%). Finally, the single sequence of A. mizorhina was strongly separated from A. doursana by an average of 10.45% (range 10.18—10.94%). These genetic results justify the decision of Pisanty et al. (2022b) to elevate A. mizorhina to species level, the findings of Wood et al. (2020b) who identifed the female of A. alchata, and the original description of A. d. agadira by Warncke as a subspecies, correctly associating dark specimens from south-western Mo- rocco with A. doursana. Given the variation in colour forms observed here, no subspecies framework is employed, and so the Iberian taxon is referred to simply as A. doursana. Andrena (Truncandrena) medeninensis Pérez, 1895 sensu lato Andrena medeninensis was described from Tunisia, and like A. doursana, it nominal- ly displays great variation across its range from Morocco and Iberia to Turkey and the Levant. Warncke (1967, 1974, 1980) described several subspecies, A. m. donata Warncke, 1967 from Spain, A. m. abunda Warncke, 1974 from Morocco, A. m. tiznita Warncke, 1980 from south-western Morocco, and A. m. usura Warncke, 1967 from Turkey. Wood (2023b) synonymised A. m. usura with A. pareklisiae Mavromoustakis, 1957, leaving the North African and Iberian populations. Sequences of A. medeninensis s. str. and A. m. tignita formed a clade with bootstrap support of 91 (Fig. 29). However, the single available sequence of A. medeninensis s. str. differed from A. m. tignita sequences by an average of 4.80% (range 4.70—4.85%), A, m. tignita forming a subclade with bootstrap support of 94. However, A. m. abunda was strongly separated from this clade of A. medeninensis s.l. by an average genetic dis- tance of 10.19% (range 9.97-10.57%). This A. m. abunda clade had bootstrap support of 100, and was sister to an A. medeninensis s.l. + A. tscheki Morawitz, 1872 s.l. clade. Andrena abunda stat. nov. is therefore raised to species status; it restricted to Algeria and Morocco. Morphologically, it is extremely similar to A. medeninensis s.|., but has much darker pubescence. In the female sex (the male is unknown), the only clear struc- tural difference is that A3 exceeds the length of A4+5+6, whereas in A. medeninensis s.1. A3 is shorter than A4+5+6. As it was unfortunately not possible to sample the Iberian subspecies A. m. donata, and no genetic sequences are available from Tunisia, the Jocus typicus for A. medeninensis s. str., no further taxonomic action is taken here. Given the large genetic difference dis- played by A. abunda despite almost no morphological differentiation (at least in the female sex), it is difficult to comment on the Iberian subspecies which simply appears to be a colour variant of this nominally widespread species. Andrena (Truncandrena) truncatilabris Morawitz, 1877, Andrena (Truncandrena) truncatilabris espanola Warncke, 1967, and Andrena_ (Truncandrena) truncatilabris nigropilosa Warncke, 1967 Andrena truncatilabris is a widespread species that was originally described from the Caucasus from what is today Armenia (Astafurova et al. 2021). The species is nominally 306 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) distributed across the Mediterranean basin, from Morocco and Iberia to the Urals and Iran (Gusenleitner and Schwarz 2002). However, in the east it descends only to the Levant and does not enter the eastern part of North Africa. In the west, Warncke (1967) described two subspecies: A. ¢. espanola from Spain and A. ¢. nigropilosa from Algeria that differed from A. truncatilabris s. str. in the structure of their clypeus and their male genital cap- sule, though Warncke noted that the difference between the two subspecies was minimal, pointing to the darker pubescence of North African specimens as a point of difference. Sequences from specimens from Spain and Morocco showed almost no genetic differentiation, with an average intraspecific distance of 0.99% (range 0.00—1.85%; Fig. 29). They were strongly separated from A. truncatilabris s. str. sequences from northern Israel by an average genetic distance of 9.66% (range 9.12—10.12%). Andrena truncatilabris s. str. formed a clade with bootstrap support of 99, and was sister to A, (Truncandrena) ferrugineicrus Dours, 1872, whereas A. t. nigropilosa + A. t. espanola formed a clade with bootstrap support of 100 that was sister to the A. truncatilabris s. str. + A. ferrugineicrus clade. Given this genetic difference, it is clear that specimens from Iberia and north- western Africa are both conspecific and distinct from A. truncatilabris s. str. Given this distribution, the use of the name A. ¢. espanola is undesirable, and so A. nigropilosa stat. nov. is elevated to species status and A. ¢. espanola syn. nov. is synonymised with it as a subjective junior synonym, as the two names were described in the same publication. The updated synonymy is therefore as follows: Andrena (Truncandrena) nigropilosa Warncke, 1967, stat. nov. Andrena (Truncandrena) truncatilabris nigropilosa Warncke, 1967: 225, 2 [Algeria: OOLM, examined]. Andrena (Truncandrena) truncatilabris espanola Warncke, 1967: 224, 93 [Spain: OOLM, examined] syn. nov. Distribution. Portugal, Spain, France, Morocco, Algeria, Tunisia (newly recorded). Material from south-eastern France and northern Italy must be carefully revised, but the position is taken here that the Maritime Alps represent a barrier between A. nigro- pilosa and A. truncatilabris s. str. This should be confirmed with genetic evidence. Material examined. Atcerta: S. Algeria, Laghouat [33.8082°N, 2.8316°E], iii—iv.1929, 19, leg. Meyer, OOLM (holotype of A. ¢. nigropilosa); Tlemcen, 20.iv.1910, 13, leg. de Bergeoin, OOLM (paratype of A. ¢ nigropilosa); SPAIN: Sierra Nevada [37.0732°N, -3.3948°E], vi.1891, 19, leg. Handl., OOLM (holotype of A. t. espanola); Montarco, 28.iv.1924, 1, leg. J.M. Dusmet y Alonso, OOLM (paratype of A. £. espanola); Tunisia: Kef, 5 km SW Touiref, 28.iv.2012, 41, leg. C. Sevidy & A. Miller, AMC/TJWC. Andrena (Truncandrena) villipes Pérez, 1895 Pérez (1895) described A. villipes from north-eastern Spain (Fig. 30), later describ- ing the synonymous A. (Truncandrena) squalida Pérez, 1903 from south-western The genus Andrena in the Iberian Peninsula 307 Figure 30. Andrena ( Truncandrena) villipes Pérez, 1895; female lectotype A label details B profile C face, frontal view D dorsal view. France. It has a restricted distribution, known from Cistaceae-rich habitats from northern Morocco, Portugal, Spain, southern France, and north-western Italy (Gusenleitner and Schwarz 2002; Lhomme et al. 2020). Searches in the Sierra de las Nieves in southern Spain produced two particularly large and dark specimens provisionally identified as A. villipes. A sequence from one of these specimens is separated from A. villipes sequences from southern Portugal by an average genetic distance of 11.85% (range 11.85—11.85%). This new species is described and di- agnosed below. Material examined. Spain: Barcelona [41.4028°N, 2.1332°E], 12, MNHN (lectotype of A. villipes; Fig. 30). Undescribed subgenera As a result of the ground-breaking analysis of Pisanty et al. (2022b), we now have unprecedented phylogenetic resolution for the genus Andrena, and are now able to seriously deal with the problems inherent to the subgeneric classification system largely solidified by Warncke (1968a). Building on the work of Pisanty et al. (2022b), the Ibe- rian fauna contains representatives of 44 described subgenera, but also representatives for nine additional clades that are currently undescribed. Some of these are dealt with 308 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) here, though the aegyptiaca-group (Iberian representative: A. alluaudi Benoist, 1961) is not treated here; this lineage will be described in a planned future comprehensive revision of the subgenus. The former Poliandrena Warncke, 1968 Pisanty et al. (2022b) demonstrated that Andrena polita Smith, 1847 falls into an expanded Ulandrena, and that Poliandrena is a strict synonym of this subgenus, and additionally that Poliandrena sensu Warncke contains at least five different polyphyl- etic clades, with the possibility of additional distinct clades when taxonomic sampling and phylogenetic analysis is more complete. Warncke, who described the subgenus Poliandrena, understandably used it as a ‘waste-basket’ for unclear taxa that displayed no clear defining characters. It is therefore necessary to split up this old grouping and to describe new subgenera. In Iberia, representatives of all five subgenera can be found. ‘These can be broadly summarised as the blanda-group, the florea-group, the limbata-group, the oviventris- group, and the re/ata-group. Four of these five lineages are represented in the analysis of Pisanty et al. (2022b) which is based on UCE analysis. UCE analyses are based on thousands of loci, and hence offer a high degree of confidence that groups are or are not related. Whilst single locus COI analyses cannot compare to those based on UCEs, they can demonstrate if individual species are closely related, and hence complement a UCE-informed phylogeny. No members of the oviventris-group were included in the analysis of Pisanty et al. (2022b), but analysis of barcodes (Fig. 31) places A. farinosa Pérez, 1895, A. farinosoides Wood, 2020, and A. oviventris Pérez, 1895 ina single clade with bootstrap support of 99, well-separated from A. corax Warncke, 1975, A. murana Warncke, 1975, and A. relata Warncke, 1975 which belong to the re/ata-group and which are morphologically the most similar to members of the oviventris-group. The two clades are separated by A. imbata Eversmann, 1852. On the basis of the analysis of Pisanty et al. (2022b) combined with these barcodes, subgenera are described for the blanda-group, florea-group, limbata-group, and oviventris-group. No action is currently taken for members of the re/ata-group, as the status of mor- phologically similar species in the Eastern Mediterranean to Central Asia is unclear, and it must be genetically demonstrated if they belong to the re/ata-group or not. The members of the newly described subgenera are detailed below; in an Iberian con- text, the following species can be considered to be part of the relata-group: A. corax, A, laurivora Warncke, 1974, A. macroptera Warncke, 1974, A. murana, and A. relata. The caroli-group Members of this group of species have been placed in the subgenus Campylogaster Dours, 1873 (Warncke 1968a) that has the type species A. erberi Morawitz, 1871. Pisanty et al. (2022b) identified that Campylogaster is polyphyletic, with A. caroli Pérez, 1895 falling far away from A. erberi. Campylogaster therefore can probably only be The genus Andrena in the Iberian Peninsula 309 WPATW586-22_Andrena_innesi_TJW_774__Morocco_Guelmim-Oued_Noun_Region IBIHM476-21_Andrena_alluaudi_Portugal_Vale_do_Telha | . aegyptiaca-group IBIHM475-21_Andrena_alluaudi_Portugal_Praia_Cabeco WPATW445-22_Andrena_blanda_TJW_633__Morocco_Draa WPATW444-22_Andrena_blanda_TJW_632__Morocco_Draa | B la n dan d rena su bg en. nov. WPATW446-22_Andrena_blanda_TJW_634_ Morocco_Fes-Meknes IBIHM485-21_Andrena_florea_Portugal_Odeceixe Bryan drena WPATW591-22_Andrena_florea_TJW_779__Morocco_Fes-Meknes 98 subgen. nov. WPATW590-22_Andrena_florea_TJW_778__Morocco_Souss-Massa_Region WPATW166-21_Andrena_murana_Spain_Madrona WPATW165-21_Andrena_murana_Spain_Madrona 89 WPATW124-21_Andrena_murana_Spain_Lupiana WPATW348-21_Andrena_relata_Spain_Madrona relata-group WPATW349-21_Andrena_relata_Spain_Madrona WPATW138-21_Andrena_corax_Spain_Sesna_Nuevo 43 WPATW148-21_Andrena_corax_Spain_Chinchon WPATW156-21_Andrena_limbata_Spain_La_Losa # Limbandrena subgen. nov. WPATW164-21_Andrena_limbata_Spain_Madrona WPATW554-22_Andrena_farinosoides_TJW_742_Morocco_Midelt WPATW113-21_Andrena_farinosa_Spain_Balaguer IBIHM515-21_Andrena_oviventris_Portugal_Serra_do_Geres Ovandrena subgen. nov. 60 WPATW346-21_Andrena_oviventris_Spain_Sesena_Nuevo. WPATW149-21_Andrena_oviventris_Spain_Chinchon WPATW622-22_Andrena_oviventris_TJW_810_ Morocco_Oriental_Region 74 WPATW623-22_Andrena_oviventris_TJW_811__Moracco_Fes-Meknes 0.04 Figure 31. Phylogenetic tree (maximum likelihood) of Andrena from currently undescribed subgenera based on the mitochondrial COI gene. Andrena (incertae sedis) innesi Gribodo, 1894 is used as an out- group. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted). applied to the species around A. erberi that have strongly depressed tergal margins, e.g. A. iranella Popov, 1940 (Iran, Turkemenistan, ?Arabian Peninsula), A. nanshanica Popov, 1940 (China, Mongolia), A. chengtehensis Yasumatsu, 1935 (China, South Ko- rea). This subgenus therefore appears to be eastern and predominantly Asian, with a western limit of A. erberi in the southern Balkans. In contrast, A. caroli has the ter- gal margins flat and not noticeably depressed. Based on the analysis of Pisanty et al. (2022b) combined with this morphological difference, a new subgenus Pruinosandrena subgen. nov. is described below for the species around A. caroli. Importantly, A. lateralis Morawitz, 1876 and A. incisa Eversmann, 1852 were also placed into Campylogaster by Warncke (1968a). Although not sampled by Pisanty et al. (2022b), A. lateralis falls far away from the Pruinosandrena based on COI analysis (Fig. 32). Morphologically, A, lateralis and A. incisa form a species pair (that can be referred to at the incisa-group) that lacks many of the characters shared by members of the Pruinosandrena (see be- low), and they probably represent an additional undescribed subgenus. Without ad- ditional genetic data, no further steps are taken for this species pair at the present time. Finally, clarity is required for the status of taxa lumped under a broad concept of A, pruinosa Erichson, 1835, specifically A. pruinosa succinea Dours, 1872 and_A. pruinosa parata Warncke, 1967. Erichson (1835) described A. pruinosa from southern Spain 310 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) WPATW349-21_Andrena_relata_Spain_Madrona WPATW330-21_Andrena_colletiformis_Spain_Villar_del_Olmo WPATW151-21_Andrena_lateralis_Spain_Chinchon lateralis; subgenus undescribed WPATW193-21_Andrena_lateralis_Spain_Ucles WPATW648-22_Andrena_sardoa_TJW_836_Morocco_Fes-Meknes WPATW649-22_Andrena_sardoa_TJW_837_Morocco_Fes-Meknes WPATW192-21_Andrena_pruinosa_parata_Spain_Madrid_Ucles i parata stat. nov. WPATW136-21_Andrena_pruinosa_Spain_Madrid_Pinto WPATW137-21_Andrena_pruinosa_Spain_Madrid_Pinto pru inosa s. str. WPATW152-21_Andrena_pruinosa_Spain_Madrid_Chinchon 99 Pruinosandrena WPATW423-22_Andrena_caroli_TJW_611_Morocco_Guelmim-Qued_Noun_Region caroli subgen. nov. ANDPHO35-21_Andrena_caroli_Israel_Nahal_Zin WPATW388-22_Andrena_pruinosa_succinea_TJW_576_Morocco_Souss-Massa_Region WPATW389-22_Andrena_pruinosa_succinea_TJW_577_Morocco_Oriental_Region WPATW390-22_Andrena_pruinosa_succinea_TJW_578_Morocco_Fes-Meknes succinea § p. resurr, WPATW391-22 Andrena_pruinosa_succinea_TJW_579_Morocco_Fes-Meknes ANDIL162-22_Andrena_succinea_lIsrael_HaMakhtesh 0.03 Figure 32. Phylogenetic tree (maximum likelihood) of Andrena from the subgenera Brachyandrena Pittioni, 1948, Lepidandrena Hedicke, 1933, and currently undescribed subgenera based on the mito- chondrial COI gene. Andrena (incertae sedis) relata Warncke, 1975 is used as an outgroup. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted). (Andalusia); a single female labelled as ‘type’ is conserved in the ZMHB which may au- tomatically be the holotype (Fig. 33), but this is ambiguous as it is unclear if Erichson described the species from multiple specimens or not. Dours (1872) described A. suc- cinea from Algeria, noting the clear red colouration of the metasoma (hence the species name, succin = amber). Warncke (1967) used A. succinea as a subspecies in combina- tion with A. pruinosa, arguing that males from North Africa could not be clearly sepa- rated from Spanish males. He then described A. pruinosa parata from south-eastern Spain, giving characters related to colouration and antennal ratios. Genetic analysis of members of the Pruinosandrena demonstrates that the broad concept of A. pruinosa used by Warncke was overly conservative (Fig. 32). A single red-marked female specimen initially identified as A. pruinosa was separated by 7.45% from three additional A. pruinosa s. str. specimens. Inspection of Warncke’s original description of A. pruinosa parata and comparison of the antennal ratios demonstrates that this divergent specimen has A3 clearly longer than A4+5, whereas A3=A4+5 in A, pruinosa s. str. The other characters mentioned by Warncke relating to colour do not The genus Andrena in the Iberian Peninsula eilal Figure 33. Andrena (Pruinosandrena) pruinosa Erichson, 1835, female holotype/syntype A label details B head, dorsal view € mesosoma, dorsolateral view D dorsal view. work consistently, as one of the barcoded A. pruinosa s. str. has the terga partially red- marked and the hind tibiae lightened orange. Though small, this morphological differ- ence is meaningful, as the two taxa are found essentially in direct sympatry; the distance between the sampling location of barcoded specimens of A. pruinosa s. str. from Pinto was approximately 11 km from the Camino de Uclés sampling site for A. pruinosa parata. \n this context, a genetic separation of 7.45% combined with the morphologi- cal difference is highly significant, and A. parata stat. nov. is treated as a valid species. Andrena pruinosa succinea was strongly separated from A. pruinosa s. str. by an aver- age genetic distance of 9.45% (range 8.97—10.33%). Sequences of A. pruinosa s. str. were identical, which is not surprising as they all came from a small part of the prov- ince of Madrid. Andrena pruinosa succinea samples came from a large geographic area from south-western Morocco to Israel, but still showed low average intraspecific varia- tion of 2.06% (range 0.14—3.80%). The two clades were not sister, being separated by A, parata and A. caroli, and were supported by bootstrap support of 99 and 95, respec- tively. Andrena succinea sp. resurr. is therefore considered to be a valid species, distinct from A. pruinosa s. str. Morphologically, separation of males is straightforward, and it is unclear why Warncke considered the difference unclear. Andrena succinea males have a yellow marked clypeus (see illustrations in Wood et al. 2020b), with the yellow 312 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) markings sometimes extending onto the lower part of the paraocular areas (uniformly black in A. pruinosa s. str. and A. parata) and, viewed ventrally, A4 is short, as long as broad (A4 is elongate and clearly longer than broad in A. pruinosa s. str.). Andrena parata males can easily be separated as A3 is longer than A4+5, whereas A3 is shorter than A4+5 in both A. pruinosa s. str. and A. succinea. There are also ecological differences. Andrena succinea can be found in dry desert- edge steppe habitats, as opposed to A. pruinosa which in Iberia is found in grasslands and cold steppe that are lightly more lush, at least during the spring. For example, in Morocco, A. succinea can be found in stipa steppe habitat around Bou Rached (Orien- tal region, south of Guercif) on the eastern edge of the Middle Atlas as it transitions into the desert (Fig. 34A, B), whereas it has never been recorded from the more humid parts of the Middle Atlas that have grasslands resembling those that can be found in central Iberia (e.g. Madrid, north of Chinchon, Fig. 34C, D). Although the type of A. succinea is lost, and the type for a more recently described taxon is preserved in the MNHN collection (A. sitifensis Pérez, 1895; Fig. 35), A. suc- cinea is the name that is established in the literature, either as a species itself or in com- bination with A. pruinosa (Benoist 1961; Warncke 1967; Warncke 1974; Gusenleitner and Schwarz 2002; Wood et al. 2020b; Dermane et al. 2021). In order to conserve this use, a neotype is designated below for A. succinea. As a result of these numerous changes, the updated statuses and synonymies are given here: Andrena (Pruinosandrena) parata Warncke, 1967, stat. nov. Andrena pruinosa parata Warncke, 1967: 233, 23 [Spain: OOLM, examined]. Remarks. Though described from south-eastern Spain, Warncke (1976) noted that he had examined a single specimen of A. parata (as A. pruinosa parata) from Madrid: Ribas de Jarama, but he expressed doubts as to whether or not it had been correctly labelled given the collecting localities of all other known specimens. The contemporary Camino de Uclés site is approximately 7 km from Ribas de Jarama, confirming the presence of this taxon in central Spain. Distribution. Spain. Material examined. Spain: Alicante [38.3628°N, -0.5093°W], 16, leg. G. Mercet, OOLM (holotype); Benidorm, 2.vi.1952, 19, leg. J. de Beaumont, OOLM (para- type); Fortuna [Murcia], v.1928, 13, leg. J. M. Dusmet y Alonso, OOLM (paratype); Madrid, Rivas-Vaciamadrid, Canal de Manzanares to Camino de Uclés, 19.v.2021, 19, leg. TJ. Wood, TJWC [BOLD accession number WPATW192-21]. Andrena (Pruinosandrena) pruinosa Erichson, 1835 Andrena pruinosa Erichson, 1835: 104, 2 [Spain: ZMHB, examined]. Andrena lanuginosa Spinola, 1843: 137, 2 [Spain, lectotype by present designation: MRSN]. The genus Andrena in the Iberian Peninsula bul) Figure 34. Andrena (Pruinosandrena) succinea Dours, 1872 A habitat, Oriental, Guercif, P5427, 2 km SW of Bou Rached, 950 m, 13.v.2022 B female collecting pollen from Brassicaceae spp.; Andrena (Prui- nosandrena) pruinosa Erichson, 1835 C habitat, Madrid, Chinchén, 6 km N, M-311, 14.v.2021 D male, in hand. 314 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Remarks, Examination of the type material of both.A. pruinosa and A. lanuginosa (Fig. 36) showed that both were female specimens with dark terga, conforming to the classical con- cept of this species. Neither represent A. parata, and hence the synonymy of A. lanuginosa with A. pruinosa is maintained. Spinola (1843) did not specify how many specimens he de- scribed the species from. The specimen examined here may be automatically the holotype, but as this is not clear from the original description, it is here designated as the lectotype. Distribution. Spain. Material examined. Sparn: Andalusia, 19, leg. Waltl, ZMHB (holotype/syntype; Fig. 33); Andalusia, 19, leg. Ghilinni, MRSN (lectotype of A. lanuginosa, by pre- sent designation; Fig. 36); Madrid, Madrid, Chinchén, 6 km N, M-311, 14.v.2021, 2 Gals leg. TJ. Wood, TJ WC; Madrid, Madrid, Pinto, 1 km E, Carr. la Marafosa, 132021 alidn UO% leg. T.J. Wood, TJWC. Andrena (Pruinosandrena) succinea Dours, 1872, stat. nov. Andrena succinea Dours, 1872: 424, 9 [Algeria: type lost, neotype designated below, OOLM]. Andrena chrysopyga Dours, 1872: 423, 2 (nec. Andrena chrysopyga Schenck, 1853) [Algeria: type lost]. ] Siti fensis F e val ee det. Dr. Warnecke At ; Figure 35. Andrena (Pruinosandrena) sitifensis Pérez, 1895, female lectotype A label details B profile C scutum, dorsal view D terga, dorsal view. The genus Andrena in the Iberian Peninsula 315 Andrena commixta Dalla Torre & Friese, 1895: 43. nom. nov. for Andrena chrysopyga Dours, 1872 Andrena sitifensis Pérez, 1895: 46, 9 [Algeria: MNHN, examined] Andrena fulvisquama Popov, 1940: 260, 9 [Algeria: ZISP, not examined] Remarks. The synonymy of A. mayeti Pérez, 1895 (described from Tunisia) with A. succinea reported by Warncke (1967, see also Benoist 1961) is incorrect. Examina- tion of the female lectotype (Fig. 37) shows that A. mayeti syn. nov. is a synonym of A, caroli Pérez, 1895 (described from Algeria; Fig. 38) because the foveae are wide and there are well-developed hair bands on the tergal margins (see identification key for Pruinosandrena below). Distribution. Morocco, Algeria, Tunisia, Libya, Egypt, Israel and the West Bank, Jordan, Syria, Saudi Arabia, Iran (Wood and Monfared 2022). Material examined. Atcerta: Setif [36.2059°N, 5.3965°E], 12, MNHN (lectotype of A. sitifensis; Fig. 35); Morocco: Oriental, Guercif, P5427, 2 km SW of Bou Rached, 33.8844°N, -3.6154°W, 950 m, 13.v.2022, 19, leg. T-J. Wood, OOLM [BOLD accession number WPATW389-22] (neotype of A. succinea, see below). Ved: Wess pit Sa PY | MedgceManues, — Andyens scree vm. gy ace bial, Joe. Ets... De +: i-. 2. fee. FF, fed: Hl -(137. V.Ghitiani., Anda fousic. Figure 36. Andrena (Pruinosandrena) lanuginosa Spinola, 1843, female lectotype A label details B profile C head, dorso-frontal view D terga, dorsal view. 316 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) 4 Pér © det, msi Ware pe Figure 37. Andrena (Pruinosandrena) mayeti Pérez, 1895, female lectotype A label details B profile C face, frontal view D terga, dorsal view. Andrena (Pruinosandrena) caroli Pérez, 1895 Andrena (Pruinosandrena) caroli Pérez, 1895: 47, 2 [Algeria: MNHN, examined] Andrena (Pruinosandrena) mayeti Pérez, 1895: 47, 2 [Tunisia: MNHN, examined] syn. nov. Distribution. Morocco, Algeria, Tunisia, Egypt, Israel. Material examined. Atcerta: Biskra [34.8600°N, 5.6995°E], 12, MNHN (lec- totype of A. caroli; Fig. 38); Tunisia: Chott el Djerid [33.8806°N, 8.1435°E], 19, MNHN (lectotype of A. mayeti; Fig. 37). The numida-group This group of Palaearctic species was previously placed in the subgenus 7hysandrena Lanham, 1949 by Warncke (1968a). However, true 7hysandrena occur only in North America, and Palaearctic species fall elsewhere (Pisanty et al. 2022b). Four species are recognised in the Palaearctic; A. hypopolia Schmiedeknecht, 1884, A. numida, A, ranunculorum Morawitz, 1877, and A. lunata Warncke, 1975, though additional The genus Andrena in the Iberian Peninsula 317 . Hi Figure 38. Andrena (Pruinosandrena) caroli Pérez, 1895, female lectotype A label details B profile C face, frontal view D terga, dorsal view. genetic work is required to ensure that they all belong together; no subgenus is there- fore described in the current work. The status of A. hypopolia (described from southern France) has been somewhat un- clear, as no major morphological differences from A. numida (described from Algeria) are apparent. Warncke used A. /ypopolia in combination with A. numida as the subspe- cies for south-western Europe, using several other taxa as subspecies for populations in Central and Eastern Europe (ssp. ?holosericea Bramson, 1879, considered a nomen du- bium by Gusenleitner and Schwarz 2002), southern Italy (ssp. syracusae Strand, 1921), and Turkey (ssp. albiscopa Warncke, 1967, see Gusenleitner and Schwarz 2002). No members of this group are known from the Levant. The situation is further complicated because the type of A. Aypopolia is lost (and could not be found amongst undesignated Schmiedeknecht type material located in the RMNH collection, see below), and the type of A. numida can also not be lo- cated at the MNHN. Furthermore, A. inconspicua Morawitz, 1871 was described from Calabria in southern Italy. Warncke (1967) treated this as A. (Taeniandrena) russula Lepeletier, 1841 based on the description; Gusenleitner and Schwarz (2001) then listed it as a synonym of A. numida, before finally Gusenleitner and Schwarz (2002) listed it as a synonym of A. /ypopolia despite the fact that it is an older name. ‘This is because, 318 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) under the broad treatment of Warncke, A. inconspicua would be junior to A. numida. However, Gusenleitner and Schwarz (2002) treated A. hypopolia as a distinct species, and did not resolve the status of A. inconspicua, even though it is an older name than A. hypopolia. The lectotype of A. inconspicua was recently illustrated by Astafurova et al. (2021), confirming its afhnity with the numida-group, and not the subgenus Taenian- drena. The use of the name A. inconspicua therefore depends on two things; 1) whether European and North African material are distinct, and 2) whether material from south- ern Italy shows a stronger affinity with the European or North African taxon. Genetically, barcoded specimens from Iberia showed almost no differentiation from specimens from Germany (average genetic distance 0.26%; Fig. 1), thus demonstrat- ing that the name A. /ypopolia can be applied to both the south-western and Central European populations. However, there was a consistent separation between Moroc- can and German/Iberian sequences, these separated by an average genetic distance of 3.05% (range 2.87—3.13%). A single sequence was available from Sicily. This sequence differed from German/Iberian sequences by an average genetic distance of 6.11% (range 3.87—6.82%) and from Moroccan sequences by an average of 2.39% (range 2.17-2.64%). It clustered with Moroccan sequences, with the clades Morocco+Sicily showing bootstrap support of 92 and Germany+lIberia showing bootstrap support of 86. Examination of these barcoded specimens shows that there is a subtle but consist- ent morphological difference, which is the density of punctures on T3 of females. In Iberian specimens, the disc of T3 is densely punctate, with punctures separated by <1 puncture diameter. In contrast, in Moroccan and Sicilian specimens, the disc of T3 is shallowly and obscurely punctate, with punctures separated by >1 puncture diameter. Examination of the female lectotype of A. inconspicua (Astafurova et al. 2021: fig. 24) shows that this specimen morphologically conforms to A. numida, with weak and obscure punctures on the disc of T3. On this basis, A. inconspicua is synonymised syn. nov. with A. numida, and A. hypopolia is maintained as the senior name for populations in Iberia and Central Europe. A barcoded neotype of A. numida is designated from Moroccan material (see below). This action largely maintains the status quo of Gusenleitner and Schwarz (2002), and A. numida is considered to have a distribution of Morocco, Algeria, Tunisia, Libya, and Sicily and the extreme southern parts of Italy (Calabria, Campania). The inclusion of Campania derives from treatment of the taxon A. syracusae. This taxon was described by Strand in the male sex only as a variety of A. (Simandrena) propinqua Schenck, 1853. Warncke (1967) treated this as a subspecies of A. numida with dark facial hair. The barcoded specimen from Sicily is a female, and whilst it should represent A. syra- cusae since it was collected only 150 km west of the /ocus typicus of Syracuse, since the female of A. syracusae is unknown and undescribed this cannot be said for certain. | have not seen male A. numida material from Sicily, or the type material of A. syracusae itself. However, I have seen males from the island of Ischia by Naples. These conform to the concept of A. syracusae, with extensive dark facial pubescence, and show ex- tremely weak and sparse punctation on T3. The island of Ischia and the nearby Sor- rento Peninsula on the Italian mainland are unusual and host an Andrena fauna with The genus Andrena in the Iberian Peninsula S19 several species that are otherwise found in Italy only in Sicily, such as A. (Micrandrena) fumida Pérez, 1895. On the basis of its morphology, A. syracusae is considered to rep- resent only a form of A. numida. This should ultimately be confirmed through the direct barcoding of males with dark facial pubescence; these are expected to fall into the A. numida clade. The status of A. numida albiscopa is unclear, but based on its distribution and morphology (T3 is clearly punctured), it is transferred to A. hypopolia albiscopa comb. nov. [he punctures of T3 are sparser than in A. Aypopolia s. str., and the interspaces are shinier. This is true also of A. Aypopolia material from Central Asia (Kyrgyzstan). For now, a conservative position is taken that A. hypopolia ranges from Iberia to Central Asia and western Siberia, though the eastern limit and the status of material from Tur- key requires validation through genetic analysis across this range. Finally, examination of the lectotype of A. setosa Pérez, 1903 (Fig. 39; des. Warncke 1967) reveals that it is not a synonym of A. numida hypopolia as given by Warncke (1967), but a synonym (syn. nov.) of Andrena ranunculorum. The specimen comes from Arreau in France which is in the Hautes-Pyrénées department, and is situated immediately adjacent to mountains that ascend to 3,000 m. Andrena ranunculorum is known from alpine habitats in the central and eastern Pyrenees on both the French and Spanish sides, so this synonymy makes both morphological and ecological sense. ; Andrena \ | Andrena A 3 f 2 det by Wwiecdke det. T.Wood 2021 — Figure 39. Andrena (incertae sedis) setosa Pérez, 1903, female lectotype A label details B profile C face, frontal view D terga, dorsal view. 320 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Material examined. Andrena ranunculorum: FRANCE: Arreau [42.9064°N, 0.3557°E], 12, MNHN (lectotype; Fig. 39). Andrena numida f. syracusae: \raty: Campania, Is. Iscia, Panza, 9.iv.1991, 64, leg. J. Gusenleitner, OOLM/TJWC. Andrena fumida: Yraty: Kampanien, Salerno, Monti Alburini (NP), SE Petina, 1100 m, 8.vi.2003, 19, leg. H. & R. Rausch, OOLM; Mondello [Palermo, Sicily], 10.iv.1979, 16, leg. J.A.W. Lucas, OOLM: Monte Faito (Campania), 13.v.1976, 14, leg. Pagliano, MRSN; Sorrento [Naples], 3.v.1970, 14, leg. J.P. van Lith, RMNH. Description of new subgenera Subgenus Blandandrena subgen. nov. https://zoobank.org/E78F170A-60CC-4D38-9D56-9727B 1347 DDC Type species. Andrena blanda Pérez, 1895. Diagnosis. Blandandrena is monotypic, and hence diagnosis of A. blanda (Fig. 40) is de facto diagnosis of the subgenus. Through the combination of slightly upturned fore margin of the clypeus, fovea broad and occupying over ¥2 the space between the lateral ocellus and the inner margin of the compound eye, weak but distinct humeral angle, unmodified posterior face of the hind femur (without teeth, carinae, or spines), lack of squamous hairs, simple hind tibial spur (not broadened basally or medially), dark integument, black male clypeus, and essential absence of defining features it falls very close to members of the relata-group and to Ovandrena subgen. nov. that were formerly lumped together under the subgenus Poliandrena (see above). Andrena blanda females can provisionally be separated from the relata-group by the weakly punctate terga, punctures shallow and somewhat obscure, separated by 1—2 puncture diameters (Fig. 40D) whereas in the re/ata-group, tergal punctures are typically much stronger, clear and dense, separated by 1—2 puncture diameters but often by only 1 puncture di- ameter. [his character works for West Mediterranean members of the re/ata-group, but additional work is needed to define this subgenus in the east, and so it is not defined and described here. Andrena blanda females can be separated from the Ovandrena by their simple scutal hairs (Fig. 40A—C; hairs semi-squamous in Ovandrena) and poorly deline- ated propodeal triangle that lacks lateral carinae (Fig. 40C; propodeal triangle clearly defined by presence of lateral carinae in Ovandrena; see below for additional detail). Andrena blanda males can be separated by their black clypeus (Fig. 40F) in combina- tion with their distinctive genital capsule (Fig. 40G). Most members of the relata-group have a yellow-marked clypeus, but for those with a black clypeus (e.g. A. corax), the geni- tal capsule allows recognition. In A. blanda, the genital capsule is short and compact, more or less round, with gonocoxae with apically diverging inner margins and which are produced into short pointed teeth, and the gonostyli are apically strongly broadened and flattened, the disc being slightly broader than long. In the re/ata-group, the genital capsule is typically simple, without strongly apically broadened gonostyli, or when this is the case (e.g. A. murana Warncke, 1975a) then the clypeus is yellow-marked and the The genus Andrena in the Iberian Peninsula oon z Figure 40. Andrena (Blandandrena) blanda Pérez, 1895, female A profile B face, frontal view C propodeum, dorsal view D terga, dorsal view; male E profile F face, frontal view G terga, dorsal view H genital capsule. 322 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) gonocoxae are not produced into pointed teeth. Andrena blanda can be separated from the Ovandrena by the genital capsule, as in Ovandrena the gonostyli are apically flattened and spatulate, but the disc is always longer than broad. The gonocoxae are also produced into apically projecting teeth, but the inner margins of the gonocoxae are parallel and do not diverge. The propodeal triangle of Ovandrena is also triangular and strongly defined by lateral carinae, whereas in A. blanda it is poorly defined and lacks lateral carinae. Description. Medium-sized bees (9-10 mm) with dark integument. Head broad, 1.4 times broader than long. Gena slightly exceeding width of compound eye; ocel- loccipital distance 1.5—2 times diameter of lateral ocellus, slightly broader in male sex. Facial fovea broad, occupying almost entire distance between lateral ocellus and inner margin of compound eye. Mesosoma dorsally with moderately long light brown hairs, laterally with white hairs. Pronotum laterally with humeral angle. Dorsolateral surface of propodeum with obscure and finely raised rugosity; propodeal triangle broad, poor- ly delineated laterally, surface with fine granular reticulation, basally with weakly raised rugosity, propodeal triangle thus defined by change in surface sculpture. Forewing with nervulus antefurcal. Hind tibial spurs simple, not broadened basally or medially. Terga weakly and obscurely punctate, punctures separated by 1—2 puncture diameters. Male genital capsule rounded, more or less circular in outline, gonocoxae with inner margins apically diverging, produced into apically projecting short pointed teeth. Gonostyli apically broadened and flattened, apical disc slightly broader than long. Penis valves relatively narrow, occupying less than half space between gonostyli. Etymology. The name is taken from the name of the type species A. blanda, with blan- da being the feminine singular of the adjective blandus which can mean pleasant, agree- able, smooth. It can be used to refer to the generally unremarkable nature of the species which has made it hard to assign to a particular group of species. The gender is feminine. Included species. Andrena blanda (Spain, including mainland Spain and newly re- corded for Fuerteventura), Morocco, Algeria, Tunisia; Gusenleitner and Schwarz 2002). Material examined (illustrative). ALGERIA: Biskra [34.8600°N, 5.6995°E], 19, MNHN (lectotype des. Warncke 1967); 5 km N of Mecheria, st. 9, 7.iv.1983, 19, leg. R. Leys & P. v. d. Hurk, RMNH; 5 km SE of Sfissifa, st. 8, 6.iv.1983, 12, leg. R. Leys & Pv. d. Hurk, RMNH; Morocco: Draa-Tafilalet, Tazenakht, 1 km W Anezal, c. 1600 m, 15.iv.2022, 19, leg. TJ. Wood, TJ WC; Foum Zguid, 50 km N, 30.iii.1986, 13, 169, leg. M. Schwarz, MSC; Ifkern, 25 km E Boulemane, 25.v.1995, 109, leg. Mi. Halada, OOLM; Spain: Fuerteventura, Costa Calma, ESE Montajfia Pelada, 29.iii.2015, 3 Q, leg. A. Miller, AMC/TJWC; Fuerteventura, S Costa Calma, Montafeta de los Verdes, 1.iv.2015, 52, leg. A. Miiller, AMC; Sierra de Marta, 25 km W Lorca, 10.v.2003, 19, leg. J. Halada, OOLM; Granada, Pantano de los Bermejales, 26.v.1982, 19, leg. R. Leys, RMNH. Subgenus Bryandrena subgen. nov. https://zoobank.org/54B1C9C7-8D5A-441B-B49B-DDBA8FB5FE39 Type species. Andrena florea Fabricius, 1793. The genus Andrena in the Iberian Peninsula 323 Diagnosis. Bryandrena is monotypic, and hence diagnosis of A. florea is de facto diagnosis of the subgenus. The combination of broad head (Fig. 41B, F; 1.3—-1.4 times broader than long), the inner margins of the compound eyes diverging ventrally, hu- meral angle (stronger in male), punctate clypeus with underlying surface shiny over the majority of its area, propodeal triangle not defined by lateral carinae and comparatively smooth relative to the microreticulate dorsolateral parts of the propodeum (Fig. 41C), terga that are always at least partially red-marked (Fig. 41D, G), unmodified posterior face of the hind femur (without teeth, carinae, or spines), lack of squamous hairs, simple hind tibial spur (not broadened basally or medially), black male clypeus, and unique genital capsule (Fig. 41H; see description below) allows separation from any other Andrena species. Description. Medium-sized bees (11-13 mm), integument predominantly dark, with red markings on at least one tergum, sometimes all terga extensively red marked. Head broad, 1.3—1.4 times wider than long, inner margins of compound eyes diverg- ing ventrally. Gena slightly exceeding width of compound eye in females, clearly ex- ceeding width of compound eye in males; ocelloccipital distance 1.5—2 times diameter of lateral ocellus. Facial fovea broad, occupying %4 of distance between lateral ocellus and inner margin of compound eye. Pronotum laterally with humeral angle, more pro- nounced in male sex. Dorsolateral surface of propodeum microreticulate, with weakly raised reticulation; propodeal triangle poorly defined laterally, comparatively smooth and lacking microreticulation, basally with raised rugosity covering variable extent, never entire propodeal triangle. Forewing with nervulus interstitial. Terga regularly and densely punctate, punctures separated by 1 puncture diameter. Male genital capsule strongly elongate, gonocoxae essentially truncate with inner margin rounded, gono- styli apically produced, elongate, strongly flattened and spatulate, 3 times longer than broad; penis valves basally broad, strongly narrowing medially to become elongate and acutely pointed apically. Etymology. The name is taken from the pollen host plant Bryonia (Cucurbitaceae) which ultimately derives from the Greek Bevwvia [brudnia]. Andrena florea can be found frequently almost wherever Bryonia species are in flower. The gender is feminine. Included species. Andrena florea (West Palaearctic, from Morocco and Iberia to Iran and the Ural Mountains; Gusenleitner and Schwarz 2002). Subgenus Limbandrena subgen. nov. https://zoobank.org/E0948E05-DB8C-4AD2-B260-01D335F6EA96 Type species. Andrena limbata Eversmann, 1852 (illustrated by Astafurova et al. 2022). Remarks. Historically, A. toelgiana Friese, 1921 has been considered the sister spe- cies to A. limbata, differing by the yellow clypeus in the female sex (see Gusenleitner and Schwarz 2002). However, the male is unknown. Structurally, there are no differ- ences. Examination of specimens from Bulgaria and Turkey show a gradient of yellow colouration on the clypeus, from entirely black, with a narrow longitudinal yellow 324 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Figure 41. Andrena (Bryandrena) florea Fabricius, 1793, female A profile B head, frontal view C propo- deum, dorsal view D dorsal view; male E profile F face, frontal view G terga, dorsal view H genital capsule. The genus Andrena in the Iberian Peninsula 325 strip, predominantly yellow-marked, and entirely yellow-marked (Fig. 42). Across this gradient there is also variation in the strength of the tergal hair bands, with material nominally conforming to A. toelgiana falling closer to A. limbata s. str. in displaying clear hair bands, whereas the subspecies A. limbata dusmeti Warncke, 1975 (Portugal, Spain, southern France, north-western Italy) has hair bands that are almost absent in fresh specimens (Figs 43C—H, 44A, B, E, F). On the basis of this colour gradient, ab- sence of structural characters, and lack of a described male that could offer a distinctive difference, A. toelgiana syn. nov. is synonymised with A. limbata. Figure 42. Andrena (Limbandrena) limbata Eversmann, 1852 female head in frontal view on west-east gra- dient A Spain B Bulgaria C Turkey (Giiriin) D Turkey (Bing6l) E Turkey (Bingél) F Turkey (Diyarbakur). 326 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Figure 43. Andrena (Limbandrena) limbata Eversmann, 1852, female A scutum, dorsal view B head, dorso-frontal view; Andrena limbata dusmeti Warncke, 1967 female C profile D terga, dorsal view; A. limbata s. str. female, Bulgarian specimen E profile F terga, dorsal view; A. limbata s. str. female, Turk- ish specimen G profile H terga, dorsal view. The genus Andrena in the Iberian Peninsula 327 Diagnosis. Limbandrena (and, de facto, A. limbata) can be recognised in the female sex due to the combination of squamous brown hairs on the scutum, scutellum, and metanotum (Fig. 43A), the long ocelloccipital distance equal to three times the diam- eter of a lateral ocellus (Fig. 43B), the posterior face of the hind femur which lacks a transverse carina and transverse row of raised teeth or spines, the weakly and shallowly punctate dorsolateral surfaces of the propodeum, the laterally clearly delineated and in- ternally finely rugose propodeal triangle (not rugosely areolate), the pronotum with at most a weak lateral angle, the more or less squarish head which is only 1.1 times wider than long, and the simple hind tibial spur that is not broadened basally or medially. Males can be recognised by most of the same characters: the yellow clypeus (Fig. 44C, D), the relatively rounded head which is only 1.1—1.2 times broader than long (Fig. 44C, D), the long ocelloccipital distance equal to three times the diameter of a lateral ocellus, the pronotum with at most a weak lateral angle, the weakly and shallowly punctate dorsolateral surfaces of the propodeum, the laterally clearly deline- ated and internally finely rugose propodeal triangle (not rugosely areolate), and the genital capsule with weakly produced rounded gonocoxal teeth, gonostyli with weakly raised and rounded projection on inner margin, and penis valves with rounded lateral hyaline extensions (Fig. 44G, H). No other Andrena species shows this combination of characters. Description. Medium-sized bees (11-14 mm) with dark integument with excep- tion of yellow maculations on female (sometimes) and male clypeus (always). Head 1.1—1.2 times broader than long, compound eyes with inner margins weakly converg- ing apically. Gena slightly exceeding width of compound eye; ocelloccipital distance long, 3 times diameter of lateral ocellus. Facial fovea moderate, occupying 2 space between lateral ocellus and compound eye. Female scutum, scutellum, and metano- tum covered with short brown squamous hairs. Pronotum laterally with weak humeral angle. Dorsolateral surface of propodeum weakly and shallowly but regularly punctate, punctures separated by 0.5—1 puncture diameter. Propodeal triangle clearly delineated laterally by raised carinae, internal surface with clear pattern of fine rugosity medially, not extending over entire area. Forewing with nervulus strongly postfurcal. Hind tibial spurs simple, not broadened basally or medially, apically weakly bent. Terga densely and finely punctate, punctures separated by 0.5 puncture diameters. Male genital cap- sule slightly elongate, with produced and weakly rounded gonocoxal teeth. Gonostyli with weakly raised and rounded projection on inner margin. Penis valves produced into rounded hyaline extensions laterally, occupying majority of space between gonostyli. Etymology. The name is taken from the name of the type species A. limbata, with limbata being the feminine singular of the adjective /imbatus which means edged or fringed, probably in reference to the distinct squamous hairs on the female scutum and scutellum. The gender is feminine. Included species. Andrena limbata (Europe from Portugal and Spain to Turkey, Israel, northern Iran, and the Ural Mountains; Gusenleitner and Schwarz 2002). Material examined (illustrative). ALBANIA: Lopan [Lapanj], 14.vi.2018, 29, leg. Kobe Janssen collection (Belgium); Butcarta: Lozenec [Lozenets, Aosenen]/Micurin, 24.vi.1988, 59, leg. B. & O. Tkalct, OOLM: Croat: Istrien, Rovinjsko Selo, 8-9. 328 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Figure 44. Andrena (Limbandrena) limbata Eversmann, 1852, Turkish male A profile C face, fron- tal view E terga, dorsal view G genital capsule; Andrena limbata dusmeti Warncke, 1967, Spanish male B profile D face, frontal view F terga, dorsal view H genital capsule. The genus Andrena in the Iberian Peninsula 20) vi.2012, 19, leg. Holzmann, OOLM; France: B. d. R., Fontvieille, 28.v.1993, 49, leg. H. & J.E. Wiering, RMNH; Georcua: E Tbilisi, lori river S Sagarejo, 19.vi.2015, 39, leg. M. Snizek, OOLM: Greece: Kos I. Kefalos, 4.v.1989, 29, leg. K.M. Guich- ard, NHMUK; Peloponnese, 12 km NWW Sparti, 26.v.2005, 19, leg. M. Kadlecova, OOLM; Ivaty: Arezzo, Oliveto, 28.v.1989, 3Q, leg. H. & J.E. Wiering, RMNH; Pie- monte, S. Benefetto Belbo, 10.vi.1979, 19, leg. Pagliano, MRSN; PorrucaL: Man- teigas, Serra da Estrela [40.4009°N, -7.5396°E], 28.iv.—9.vii.1929, 19, leg. Krichel- dorf, OOLM (holotype of A. limbata dusmeti); Spain: Segovia, Madrona, 500 m NE, Arroyo del Hocino, 15.v.2021, 14, 29, leg. T.J. Wood, TJWC; Avila, Hoyocasero, 1350 m, 20.v.1995, 14, 29, leg. H. & J.E. Wiering, RMNH; Turkey: Tanin-Tanin- Pass, 1700 m, 12.vi.1984, 19, OOLM; Karatepe/Adana, 17.iv.1984, SG5 leg. K. Warncke, OOLM; Akyaka, Yesilova, 20.vi.2016, 19, leg. M. Kasparek, OOLM. Subgenus Ovandrena subgen. nov. https://zoobank.org/A122C278-C33D-4790-A34A-FC270AE6A962 Type species. Andrena oviventris Pérez, 1895. Diagnosis. Through the combination of slightly upturned fore margin of the cl- ypeus, broad fovea occupying at least ¥2 the space between the lateral ocellus and the inner margin of the compound eye, weak but distinct humeral angle, unmodified posterior face of the hind femur (without teeth, carinae, or spines), simple hind tibial spur (not broadened basally or medially), dark integument, black male clypeus, and essential absence of defining features it falls very close to members of the relata-group and to Blandandrena that were formerly lumped together under the subgenus Polian- drena (see above). In the female sex, Ovandrena species differ from these groups by only a single major character: the structure of the propodeal triangle (Fig. 45A, B). Ovandrena spe- cies have scutal hairs that are not as short and clearly squamous as in members of the Aenandrena, Limbandrena, or Lepidandrena Hedicke, 1933, but they are partially squa- mous in some species (Fig. 45C, D), whereas the scutal hairs in the re/ata-group and Blandandrena are typically simple, though in some species such as A. corax the hairs can be semi-squamous. The structure of the propodeal triangle is therefore the only character that allows consistent separation. In Ovandrena, the propodeal triangle is a broad and well-defined triangle (lateral margins extending almost to the lateral edges of the metanotum) that is defined by raised carinae; the internal surface is covered in raised irregular carinae of a similar width, therefore appearing homogenous and consistent (Fig. 45A, B). In the re/ata-group and Blandandrena, the propodeal triangle varies from smooth and undefined to weakly defined with at most irregular rugosity on its internal surface, but never approaches the condition in Ovandrena. The com- bination of these two characters therefore always allows recognition. ‘They can also be partially recognised by the extremely dense and fine punctation of the terga, punctures separated by <0.5 puncture diameters; in comparison groups, the punctures are never this fine or this dense. 330 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Figure 45. Andrena (Ovandrena) subgen. nov. characters. Andrena (Ovandrena) marsae Schmiedeknecht, 1900, female A propodeal triangle; Andrena (Ovandrena) oviventris Pérez, 1895 B female propodeal tri- angle C female scutal hairs, profile view F male genital capsule; Andrena (Ovandrena) farinosa Pérez, 1895 D female scutal hairs, profile view E male genital capsule. Males can be recognised through their combination of dark clypeus with upturned fore margin, distinctive propodeal triangle as in the female sex, pronotum with weak or strong humeral angle, A3 exceeding A4 but shorter than A4+5, and genital cap- sule which is compact with pronounced and rounded gonocoxal teeth (Fig. 45E, F). Members of the re/ata-group often have a yellow-marked clypeus, but for species show- ing a dark clypeus (e.g. A. corax), the clearly defined triangular propodeal triangle allows separation (smooth to at most weakly defined in the re/ata-group). Separation The genus Andrena in the Iberian Peninsula 331 can be made from Blandandrena by the genital capsule and the propodeal triangle, as detailed in the diagnosis section for that subgenus. Description. Small to moderately sized bees (7-11 mm) typically with dark in- tegument, one species with red tergal markings; male clypeus always dark. Head broad, 1.3—1.4 times broader than long, compound eyes with inner margins weakly converg- ing apically. Gena slightly exceeding width of compound eye; ocelloccipital distance short, slightly less than to slightly more than diameter of lateral ocellus. Facial fo- vea moderately broad, occupying 2 distance between lateral ocellus and compound eye. Female scutum, scutellum, and metanotum covered with shortish light brown to whitish semi-squamous hairs (Fig. 45C, D). Pronotum laterally with weak to strong humeral angle. Dorsolateral surface of propodeum with regular and dense network of raised reticulation; propodeal triangle wide, lateral margins extending almost to lateral edges of metanotum, laterally defined by straight raised carinae, thus strongly triangu- lar (Fig. 45A, B). Internal surface of propodeum covered by raised irregular carinae of a similar width, therefore homogenous and consistent. Forewing with nervulus intersti- tial to weakly antefurcal. Hind tibial spurs simple, not broadened basally or medially. Terga densely punctate, in females punctures extremely dense and fine, separated by <0.5 puncture diameters, in males punctures slightly sparser and coarser, separated by 0.5—1 puncture diameters. Male genital capsule compact, consistent across species, with gonocoxae strongly produced into apical rounded teeth (Fig. 45E, F). Gonostyli apically spatulate, with a more or less strongly raised inner margin. Penis valves basally narrow or moderately broad, in both cases becoming narrow medially. Etymology. The name is taken from the type species for the genus, A. oviventris. It derives from the Latin ovum meaning egg, in reference to the egg-shaped metasoma. The gender is feminine. Included species. Andrena farinosa Pérez, 1895 (Spain and France), Andrena fa- rinosoides Wood, 2020 (Morocco), Andrena marsae Schmiedeknecht, 1900 (Morocco, Algeria, Tunisia), and Andrena oviventris (Morocco, Algeria, Portugal, Spain, France). The subgenus is therefore currently restricted to the Western Mediterranean, and the centre of diversity is Morocco. The status of A. (incertae sedis) inusitata Pisanty, 2022 must be resolved through genetic analysis (see Pisanty et al. 2022a). Material examined. Andrena oviventris: ALGERIA: Teniet el Had [35.8727°N, 2.0007°E], 12, MNHN (lectotype; Fig. 46). Andrena farinosa: Spain: Murcie [Murcia], 12, MNHN (lectotype). Identification key to subgenus Ovandrena subgen. nov. 1 Note, males of these four species are extremely similar, and great care must be taken when separating them. Association with females should be made, as well as reference to their known biogeographical distributions and to confidently determined reference material. 332 Subgenus Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Terga almost entirely red-marked, with at most slight black marks basally on T1 and two black spots laterally on T2 (north-western Africa)...........eeeee eae ae eee ee Saar tical nee a raetg Scene See aetrasina cae: marsae Schmiedeknecht ered ark wit hOtei i edsMMACKUAGS sharp tis eldest tue ohueesesbeins slant cemasecuehie 3 Scutum less densely punctate, punctures separated by at least 1 puncture diameter, surface clearly smooth and shiny between punctures (Morocco).. Scutum densely punctate, punctures separated by 0.5 puncture diameters to con nent gratrow. Unters PACES: SIMI. .cnhetecsystos's Pate oet revnenutheseceenveuce tices 4, Tergal discs glabrous, with hairs restricted to marginal areas. Larger, 10-11 mm (north-western Africa and south-western Europe)........::+ oviventris Pérez Tergal discs extensively covered with extremely short hairs, forming a vel- vety pubescence in addition to denser and longer hairs on tergal margins. Smaller, 8-9 mm (Spain and France only) oe eee farinosa Pérez Larger, 9-10 mm. Tongue with outer surface of galea clearly punctate, punctures separated by 1—2 puncture diameters. Sterna with weak and sparse fringes on apical margins. Tergal punctation comparatively larger and coarser (north-western Africa and south-western Europe)..........:.006 wish haats culate et nchilchle tag tsareg Satan Setog ets oe clbatee Seaton Detect oviventris Pérez Smaller, 7-8 mm. Tongue with outer surface of galea more or less smooth and shiny, without obvious punctures. Sterna with strong and dense fringes on apical margins. Tergal punctation comparatively fine... eects 6 Clypeus comparatively less densely punctate, punctures separated by 0.5—1 puncture diameters, with shiny interspaces, thus appearing shiny. Pronotum with humeral angle comparatively strong (Spain and France only).............. sa Seats baat ha conta gh tu ceten op Seth sev aden aah outer See ohoa muah odetiaa asd gst farinosa Pérez Clypeus comparatively more densely punctate, punctures separated by 0.5 puncture diameters, interspaces dull, thus appearing dull (north-western Af- FLOeL ire a cee ad Mee PRs Ree aes main nn Rae elena Otats aus na AY eee a oe ante CRRA al 7. Viewed laterally and ventrally, tergal margins, ventrolateral parts of terga, and sternal margins usually lightened reddish orange-brown (north-western trical = omen: OR cip Nee eA, ie Min @: bec ieee on a marsae Tergal and sternal margins dark to hyaline, never extensively lightened red- dish orange-brown (Morocco only)? oo... .ecesesesceseeseeeeeeeseeees farinosoides Pruinosandrena subgen. nov. https://zoobank.org/62A1E8AF-01AD-4D69-8CD5-E20843A286D5 Type species. Andrena pruinosa Erichson, 1835. 2 ‘These two species are almost identical in the male sex, and no structural characters are currently apparent. The genus Andrena in the Iberian Peninsula 333 Figure 46. Andrena (Ovandrena) oviventris Pérez, 1895, female lectotype A label details B profile C face, frontal view D terga, dorsal view. Diagnosis. This group of species was formerly placed in the subgenus Campylogaster due to the unusual character of the strongly and densely punctate mesepisternum and dorsolateral parts of the propodeum (punctures separated by <0.5 puncture diameters). However, as discussed above, Campylogaster sensu Warncke is polyphyletic and can be broken into three distinct clades (Campylogaster s. stt., Pruinosandrena, and the incisa-group). All three share the distinctly punctate mesepisternum and also an extremely long ocelloccipital distance exceeding three times the diameter of the lateral ocellus. However, separation is straightforward. The true Campylogaster have the marginal area of the terga clearly and distinctly im- pressed with the apical margin reflexed; the impression therefore forms a latitudinal depressed furrow. In Pruinosandrena, the marginal areas of the terga are flat, without any kind of depression. In the incisa-group, the tergal margins are flat and the mese- pisternum is densely punctate, but the dorsolateral parts of the propodeum have only raised reticulation, without punctures. The propodeal triangle is also clearly delineated by raised lateral carinae, whereas these are absent in Pruinosandrena. The combination of extremely long ocelloccipital distance, dense punctures on the mese- pisternum and dorsolateral parts of the propodeum, and flat tergal marginal areas is therefore unique and characterises Pruinosandrena. 334 Thomas J. Wood / Journal of Hymenoptera Research 96: 241-484 (2023) Description. Medium-sized bees (10-14 mm). Integument variable, from dark with at most tergal margins lightened hyaline-yellow to entirety of metasoma and legs red-marked; male clypeus yellow-marked in one species. Head moderately broad, 1.2 times broader than long. Gena broad, exceeding width of compound eye; ocelloccipital distance extremely long, at least 3 times diameter of lateral ocel- lus. Facial fovea variable, from narrow to occupying entirety of distance between lateral ocellus and inner margin of compound eye. Female scutum, scutellum, and metanotum with pubescence variable, in some species with extremely short squa- mous hairs, hairs longer and non-squamous in other species. Pronotum laterally with humeral angle. Mesepisternum and dorsolateral parts of propodeum densely and clearly punctate, punctures confluent to separated by <0.5 puncture diameters. Propodeal triangle without lateral carinae, internal surface with dense network of irregularly raised rugosity, thus contrasting punctate dorsolateral surface. Forewing with nervulus interstitial. Hind tibial spurs simple, not broadened basally or medi- ally. Terga typically densely and finely punctate, punctures separated by 1 puncture diameter, at least on T2—5. Male genital capsule simple, compact, with gonocoxae apically truncate to produced into weak rounded teeth. Gonostyli apically spatu- late, penis valves more or less narrow, parallel-sided, occupying less than 2 space between gonostyli. Etymology. The name is taken from the type species for the subgenus, A. pruinosa. The Latin word pruinosa is the feminine singular of pruinosus which means ‘frosty’, in reference to the squamous hairs of the mesosoma. The gender is feminine. Included species. Andrena caroli (Morocco to Israel); Andrena nilotica Warncke, 1967 (Spain); Andrena parata (Spain); Andrena pruinosa (Spain); Andrena sparsipunc- tata Wood, 2020 (Morocco); Andrena succinea (Morocco to Iran and Saudi Arabia). The centre of diversity is therefore Spain + Morocco, with all six species occurring here. Identification key to subgenus Pruinosandrena subgen. nov. Note, the male of A. sparsipunctata is unknown. The males described by Wood et al. (2020b) are actually misidentified A. mediovittata Pérez, 1895; A. sparsipunctata is expected to be restricted to south-western Morocco. See Wood et al. (2020b) for ad- ditional useful images. 1 Berane si & ere, Re bi reat Betas hiss tents cnsatr Roa tenia Rell aoc Baanas cians 2 - A alletai ts, 2e 3.68, O85. O hts chet thet bee SM ds F Sa SUA ta Bh OE elie okie eas eae 7 2 In dorsal view, fovea broad, clearly occupying more than half of the distance between the lateral ocellus and the compound eye (Fig. 37C). Terga with strong apical pale hair bands that uniformly cover the entirety of the tergal depressions and obscure the underlying surface (Figs 37D, 38D; North Af PICA COSTAL)... ce ceveeveseveestcaveecevsaveceveronscvianucencevecsssesvegesareecssaneee caroli Pérez = Fovea narrow, occupying at most half the distance between the lateral ocel- lus and the compound eye (Figs 33B, 36C). Terga with or without clear hair co | \9 10 The genus Andrena in the Iberian Peninsula 335 bands; if with hair bands, then hair bands typically do not uniformly cover the tergal depressions, either weaker basally or stronger apically «0.0.0.0... 3 T1 with extremely sparse punctures, punctures separated by 2—4 puncture diameters. Facial fovea along its entire length separated from the inner mar- gin of the compound eye by a distance equal to its own diameter (south- western Woroceo, orily: tee cot noc eaece eater eae sparsipunctata Wood T1 with dense punctures, punctures separated at most by 2 puncture diam- eters, usually by 1 puncture diameter (Fig. 59D). Fovea not strongly sepa- rated from the inner-margin.of the-compound eyes... c.