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PSEUDOANTHIDIUM PALESTINICUM (MAVROMOUSTAKIS, 1938), STAT. NOV. (FIGS 2G, 11E, 12E, 13E, 21B, D, F, 23A) Anthidium cribratum palestinicum Mavromoustakis, 1938: 16–17, ♀ ♂. Type locality: ‘ Migdab’ [possibly ‘Migdal’, Israel but not certain], 19.v.1931 (holotype ♀, allotype ♂, ‘ Tiberias’ [Israel], 3.vii.1931) (BMNH). Material examined: Seven females, 12 males (see Supporting Information, Table S1 for specimen data). Distribution: Israel and Palestine, Jordan, Lebanon, Syria and Turkey (Fig. 22A)., Published as part of Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2022, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), pp. 1287-1337 in Zoological Journal of the Linnean Society 195 on page 1317, DOI: 10.1093/zoolinnean/zlab062, http://zenodo.org/record/5817276, {"references":["Mavromoustakis GA. 1938. II. - On some Anthidiine bees from Palestine. Annals & Magazine of Natural History 11: 15 - 18."]}

PSEUDOANTHIDIUM ROZENI GRISWOLD & BURROWS, SP. NOV. (FIGS 11H, 12H, 13H, 23F, 27B, D, F) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 79963D17-82B6-45EA-BF0E-5B82B9ED3463 Type material: Holotype. PAKISTAN: ♂ Baluchistan, Hanna, 15 km E of Quetta, 23 May 1984, Rozen, Lodhi, Stupakoff (AMNH). Paratypes: PAKISTAN: 3♂♂, 1♀, same data as holotype (AMNH, BBSL); 1♂, Baluchistan, Jalogir, 32 km N of Quetta, 23.v.1984, Rozen, Lodhi, Stupakoff (AMNH). Additional material: AFGHANISTAN: 1♂, Kabul, 1740 m, 1952, J. Klapperich (OLML); TAJIKISTAN: 1♂, Kuljab, 20.viii.1933, V.B. Popov (AMNH); 1♀, Kuljab, 11.viii.1933, V.B. Popov (BBSL). Distribution: Afghanistan, Pakistan and Tajikistan (Fig. 26B). Host plant associations: Unknown. Description female: Measurements BL = 8.3 mm, IW = 2.2 mm. Head: Mandible yellow, except for teeth and anterior margin, which are black. Mouthparts with second segment of labial palpus 1.98 × length of first, 0.95 × length of eye. Clypeus yellow except for small transverse bar of white pre-apically; apical margin of slightly emarginate; basal half shiny with mostly dense, coarse punctation (slightly less coarse than on supraclypeal area) and with distinct flat interspaces between punctures; punctation becomes finer, denser apically where punctures almost indistinguishable and integument matte. Paraocular area yellow to upper margin of antennal socket. Antenna with anterior face of flagellum more or less brownish-ferruginous; scape and pedicel relatively long (3.63 as long as wide and 1.16 as long as wide, respectively) but flagellar segments much shorter than wide (L/W: F1 = 0.88; F2 = 0.49; F3 = 0.49; F4–F5 = 0.49; F6–F7 = 0.68; F8 = 0.89; F9 = 0.96), except the last segment (L/W = 1.63) (measurements made on a single specimen). Frons matte, with honeycomb areolate punctation. Vertex with pair of wedge-shaped lateral yellow marks on posterior margin separated by approximately distance between lateral ocelli (Fig. 27B). Punctation of vertex relatively coarse, mostly contiguous or nearly so with shiny interspaces between punctures (Fig. 27B); punctures as coarser than those on frons, nearly as coarse as those of mesonotum and terga (Fig. 27B, D, F). Gena honeycomb areolate with wide but shallow punctures. Pilosity on head white or whitish, including short, dense pubescence on ventral margin of clypeus, except golden setae on mandible and labrum. Mesosoma: Mesosoma black except as noted (Fig. 27D). Scutum with punctation dense, not honeycombed, surface shiny; long median sulcus reaching more than half scutal length; parapsidal furrow indistinct, approximately three to four puncture diameters in length; anterior margin with pair of lateral, rectangular yellow marks with short pointed extension adjacent to tegula (Fig. 27D). Pronotal lobe yellow (Fig. 27D). Axilla almost entirely yellow (Fig. 27D). Scutellum with yellow band across entire posterior margin slightly narrowed medially; punctation dense, shiny, not honeycombed (Fig. 27D). Tegula with commashaped yellow mark anteriorly, otherwise translucent ferrugineous (Fig. 27D). Mesepisternum with lateral face anteriorly with coarse dense, but not honeycomb areolate punctation, interspaces rounded, becoming shallow and sparse ventrally. Metepisternum with superficial, dense minute punctation. Propodeum laterally finely, densely punctate; posterior face basally with row of coalescent, shallow punctures widely interrupted medially; rest of posterior face smooth, shiny. Tarsi, tibiae, tibial spurs and distal portion of femora yellow; ventral margin of tibia 1 with rake of thick, golden setae; basitarsal brushes golden; tarsal segments with some thick golden setae apically. Wings infuscate, darkest in marginal cell. Metasoma: Integument shiny, terga black with yellow markings (Fig. 27F). T 1 along anterior carina with dense golden fringe mostly two puncture widths long. T 1– T 5 black with yellow bands, widely interrupted on T 1 (Fig. 27F), decreasingly so on T 2– T 3 (Fig. 27F), nearly touching posteriorly on T 4, on T 5 narrowly separated; T 6 with pair of triangular yellow patches almost touching anteriorly, creating apical black triangle. Terga with punctures regular; punctures on discs of T 1– T 3 dense but not contiguous, nearly as large as those of mesonotum, slightly larger than punctation on preapical depressions of T 1– T 3 (Fig. 27F); interspaces (mostly one-third or less puncture diameter apart) flat polished (Fig. 27F); punctures on T 4– T 6 increasingly finer, denser. T 1– T 5 with apical flanges, amber on T 1– T 4 (Fig. 27F), mostly clear on T 5; on T 1– T 4 widely impunctate, on T 1– T 3 with maximal width of about 1.5 times diameter of broadest punctures (Fig. 27F), impunctate margin of T 4 slightly less broad, approximately 1.3 times diameter of broadest punctures, impunctate margin of T 5 narrower still, from approximately 0.8 to less than 0.5 puncture diameters wide medially. T 6 distinctly indented posteromedially. Scopa white. Diagnosis female: The female of P. rozeni may be distinguished from other members of this complex by the following combination of characters: punctation on T 1– T 3 coarse, punctures nearly as large as those on mesonotum and black part of scutellum, with flat smooth interspaces that give integument a shiny appearance (interspaces often reach one-third of a puncture diameter) (Fig. 27D, F); punctation of scutum dense, nearly contiguous and fairly regular, scutum thus appearing shiny (Fig. 27D); punctation on vertex between lateral ocellus, eye and post-occipital margin relatively coarse, mostly contiguous or nearly so, with shiny interspaces between punctures (Fig. 27B); T 1 along anterior carina with dense golden fringe mostly two puncture widths long; T 6 distinctly indented posteromedially. The female of P. rozeni is most similar to P. cribratum and P. tenellum, sharing the relatively coarse punctation on the vertex between the compound eye and lateral ocellus that is no finer than that on T 1 and T 2, the shiny vertex and mesonotum with distinct smooth interspaces between punctures, and punctures of scutellum as dense and as large as largest punctures of T 2 (Figs. 17B, D, F, 21A, C, E, 27B, D, F). It differs from P. tenellum in the punctures of T 2 as large as those on the black part of the scutellum instead of smaller (Figs 21C, E, 27D, F) and the presence of a distinct depression apicomedially on T 6. From P. cribratum it differs by the finer punctation of T 2, with punctures not visibly different from those on the black part of the scutellum, and the mesonotum with the surface between punctures polished, not shagreened (Figs 17D, F, 27D, F). Description male: Measurements BL = 7.2–8.1 mm, IW = 1.8–2.0 mm. Head: Integument black except where noted otherwise. Mandible cream, except for black teeth. Clypeus cream; density of punctation at base of clypeus irregular, with some punctures very close (separated by less than one-half a puncture diameter), others separated by up to several puncture diameters wide; sparsely punctate area extends medially more than halfway down clypeus; apical region with relatively dense punctation, spaces between punctures Mesosoma: Integument mostly black. Scutum and black part of scutellum with punctation dense, coarse, not areolate, with spaces between punctures Metasoma: Terga with integument shiny. T 1– T 5 black with yellow bands, widely interrupted on T 1, decreasingly so on T 2– T 3, on T 4 partially joined, T 5 entire; T 6– T 7 entirely yellow. Punctures on T 1– T 3 broad with shiny spaces between punctures Diagnosis male: Pseudoanthidium rozeni is morphologically most similar to P. tenellum. In males, P. rozeni may be differentiated from P. tenellum by the shape of the mitten-shaped lateral comb on S5, which has a distinct thumb in P. tenellum and just a nub in P. rozeni (Fig. 11D, H); the shape of the gonostylus, which is deeply notched in P. rozeni whereas only slightly notched in P. tenellum (Figs 10C, 23F); S3 with basal patch of golden pubescence rounded apically, versus pointed apically in P. tenellum; and S4 with apical translucent flap a semicircle instead of much broader than long in P. tenellum. Etymology: It is a pleasure to recognize Jerry Rozen, colleague and friend, who has contributed extensively to our knowledge of bee-nesting behaviour and immature stages worldwide and in the process collected many species new to science including P. rozeni. He has contributed greatly to training new generations of bee researchers through his leadership in the Bee Course hosted at the South-West Research Station., Published as part of Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2022, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), pp. 1287-1337 in Zoological Journal of the Linnean Society 195 on pages 1325-1328, DOI: 10.1093/zoolinnean/zlab062, http://zenodo.org/record/5817276

GENUS PSEUDOANTHIDIUM FRIESE, 1898 Pseudoanthidium Friese, 1898: 101. Type species: Anthidium alpinum Morawitz, 1874, by subsequent designation by Sandhouse, 1943: 593., Published as part of Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2022, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), pp. 1287-1337 in Zoological Journal of the Linnean Society 195 on page 1296, DOI: 10.1093/zoolinnean/zlab062, http://zenodo.org/record/5817276, {"references":["Friese H. 1898. Die Bienen Europa's (Apidae europaeae) nach ihren Gattungen, Arten und Varietaten auf vergleichend morphologisch-biologischer Grundlage. Theil 4. Innsbruck & Imst: C. Lampe.","Sandhouse GA. 1943. The type species of the genera and subgenera of bees. Proceedings of the United States National Museum 92: 519 - 619."]}

PSEUDOANTHIDIUM NANUM (MOCSÁRY, 1880) (FIGS 2F, 4, 5, 9A, C, E, 10A, 11A, 12A, 13A) Apis liturata Panzer, 1801: 21, (♀). Type locality: ‘Legi primo vere in sylvaticis 1800’ [collected in early spring in forest (in Germany, based on the title of Panzer’s work, ‘ Faunae Insectorum Germanicae ’)]. Junior primary homonym of Apis liturata Gmelin, 1790: 2789. Anthidium reptans Eversmann, 1852: 85, ♀ ♂, nom. oblit., synon. nov. Type locality: ‘in promontoriis Uralensibus australibus’ [Russia: Urals]. Lectotype, ♂, by present designation: ‘Spask Jun’ [Russia: Orenburg Prov., Saraktash Distr., Spasskoe], ‘ Anthidium reptans ♂. Evers.’, ‘ Lectotypus ♂ Anthidium reptans Eversmann, 1852 design. Fateryga et Proshchalykin 2019’ [red label] (ISZP) (Fig. 4). Paralectotypes: ♀, ‘Spask Jun’, [golden disc], ‘ Paralectotypus ♀ Anthidium reptans Eversmann, 1852 design. Fateryga et Proshchalykin 2019’ [red label]; ♀, ‘Spask Jun’, ‘ Paralectotypus ♀ Anthidium reptans Eversmann, 1852 design. Fateryga et Proshchalykin 2019’ [red label]; ♀, ‘Spask Jun’, ‘ Anthidium reptans ♀. Evers.’, ‘ Paralectotypus ♀ Anthidium reptans Eversmann, 1852 design. Fateryga et Proshchalykin 2019’ [red label]; ♀, ‘Spask Jun’, [golden disc], ‘4’ [purple label], ‘ lituratum Panz. ’, ‘ Anthidium reptans ♀. Evers.’, ‘ Paralectotypus ♀ Anthidium reptans Eversmann, 1852 design. Fateryga et Proshchalykin 2019’ [red label] (ISZP). Anthidium nanum Mocsáry, 1880: 51–53, ♀ ♂, nom. protect. Type locality: in Latin ‘Hungaria centrali, meridionali et orientali‘ [central, southern and eastern Hungary] and in Hungarian ‘Hazánk központi, déli és keleti részében’ [in the central, southern and eastern part of our country]. Lectotype, ♂, by present designation: ‘ nanum Mocs’, ‘ Anthid. lituratum ♂ det. Friese 1897’, ‘ Pécs 6/9 Biró’ [Pécs, Hungary; ‘Biró’ may refer to L. Biró, collections assistant to Sandor Mocsáry at the HMNH (Bohart & French 1986)], ‘Hungarian Natural History Museum Hymenoptera Coll. Budapest’ [blue label], [red label] (blank), ‘ Lectotypus Anthidium nanum Mocsáry 1880 design. J. Litman 2020’ [red label] (HNHM) (Fig. 5). ? Anthidium tenellum var. grandii Alfken, 1936: 111, ♀ ♂. Type locality: ‘ Molina di Quosa, Pisa’ [Italy]. Material examined: 195 females, 117 males (see Supporting Information, Table S1 for specimen data). Distribution: Abkhazia, Albania, Armenia, Austria, Azerbaijan, Belgium, Bulgaria, Crimea, Croatia, Czech Republic, France (including Corsica), Georgia, Germany, Greece, Hungary, Italy (including Sardinia and Sicily), Kazakhstan, Malta, Moldova, the Netherlands, Romania, Russia (European part, Urals), Serbia, Slovakia, Slovenia, Switzerland, Turkey (European part), Turkmenistan and Ukraine (Fig. 6A). The single female specimen mentioned from Portugal in Baldock et al. (2018) was examined and determined to be a male P. stigmaticorne. Introduced in the USA. Host-plant associations: Asteraceae south-western Germany Cirsium vulgare (Savi) Ten., Centaurea paniculata L., C. jacea L., C. scabiosa L., Onopordum acanthium L. (Westrich, 1990); Hungary Centaurea biebersteinii DC. (Mocsáry, 1880); Crimea Arctium sp. (female visit), Centaurea jacea subsp. substituta (Czerep.) Mikheev (female visit), Grindelia squarrosa (Pursh) Dunal (male visit), Inula aspera Poir. (female visit), Helianthus sp. (male visits, personal observations); Fabaceae Crimea Lotus corniculatus L. (female visit) (personal observation, A. V. Fateryga). Remarks: The name of this taxon has been the subject of much discussion. The name Apis liturata Panzer, 1801, long considered valid, was eventually recognized as a junior homonym of Apis liturata Gmelin, 1790 (Warncke, 1980). Nevertheless, Warncke (1980) assigned priority to the name Anthidium lituratum (Panzer, 1801), justifying this decision by referring to Article 59b(ii) from an amendment to the Code published in 1973 (ICZN, 1973), regarding the assignment of priority in cases of secondary homonymy. Yet Apis liturata Panzer, 1801 and Apis liturata Gmelin, 1790, representing identical species names described in the same genus are, in fact, primary homonyms and not secondary homonyms. This was noticed by Schwarz et al. (1996), who subsequently designated Apis liturata Panzer, 1801 as a junior primary homonym of Apis liturata Gmelin, 1790, permanently preventing further use of Panzer’s name. While Articles 23.9.1.1 and 23.9.1.2 of the present version of the code (ICZN, 1999) allow priority to be assigned to a junior primary homonym in ‘prevailing usage’ under certain circumstances, the version of the code available to Schwarz et al. (1996) did not include these articles. The recommendation at the time concerning primary homonyms was simply that junior primary homonyms were to be considered invalid. Therefore, regardless of the fact that certain authors have continued to use the name liturata Panzer, 1801 after the publication of Schwarz et al. (1996), this name was permanently invalidated by Schwarz et al. (1996) in accordance with the Code available at the time. The fourth edition of the code is clear regarding the stability of taxonomic decisions made using previous editions of the code, especially that ‘new names must not upset actions taken by past generations operating under different, and less restrictive, nomenclatural rules or conventions’ (ICZN, 1999). We thus uphold Schwarz’ decision and consider Apis liturata Panzer, 1801 as a junior primary homonym of Apis liturata Gmelin, 1790 and thus invalid. Finally, we consider the decision regarding the validity of a name as a separate issue from the decision regarding the taxon to which that name may be applied. When Schwarz et al. (1996) invalidated the name Apis liturata Panzer, 1801, they assigned priority to what they considered to be the next available name, P. scapulare (as Anthidium scapulare). While we maintain the decision to invalidate the name Apis liturata Panzer, 1801, we do not consider P. scapulare as a synonym of Apis liturata Panzer, 1801, but instead as a distinct taxon (see section below on P. scapulare for discussion). Chronologically speaking, P. reptans (Eversmann, 1852) is the next available name for this taxon and an examination of the type material confirms that the name P. reptans indeed refers to this taxon. Yet to the best of our knowledge, P. reptans has not been used as a valid name since 1899. Anthidium nanum Mocsáry, 1880, on the other hand, has been used as the presumed valid name for this particular taxon in at least 25 works, published by at least ten authors in the immediately preceding 50 years and encompassing a span of not less than ten years (Supporting Information, Appendix S1). Given that the conditions outlined in both Articles 23.9.1.1 and 23.9.1.2 of the Code are fulfilled, and thus in accordance with Article 23.9.1, Pseudoanthidium nanum (Mocsáry, 1880) is the valid name for this taxon and is considered a nomen protectum, while Pseudoanthidium reptans (Eversmann, 1852) is assigned the status of nomen oblitum. No specimens explicitly labelled as belonging to the type series of Anthidium nanum were located, despite extensive inquiries made at different collections, but a male specimen was located at the HNHM bearing labels written in Mocsáry’s handwriting, as well as a simple dark red label often found on type specimens designated by Mocsáry, and collected at a locality compatible with the somewhat vague type locality described in Mocsáry (1880): ‘In Hungaria centrali, meridionali et orientali’ [in central, southern and eastern Hungary] (Fig. 5). We thus consider it likely that this specimen belonged to the type series of P. nanum and designate it as a lectotype. A final note regarding the name Pseudoanthidium nanum: Mocsáry published the name P. nanum (as Anthidium nanum) in Vol. 16, No. 1 of Mathematikai es Termeszettudomdnyi Közlemenyek. A footnote on the first page of this publication suggests a citation date of 1879 but the printing of Volume 16 was not officially closed until 1881 (Baker, 1996). As a result, the name P. nanum appears in most works as either P. nanum (Mocsáry, 1879) or P. nanum (Mocsáry, 1881). However, in a note published in 1996, Baker clearly stated that Vol.16 No. 1 was itself published in April 1880 and proposes this publication year for all taxa described in Vol. 16 No. 1 of Mathematikai es Termeszettudomdnyi Közlemenyek. Few authors, if any, have followed Baker’s recommendation but at the present moment we see no reason not to do so. We thus propose that in the future, this taxon be referred to as P. nanum (Mocsáry, 1880). Anthidium sinuatum Lepeletier de Saint Fargeau, 1841 has also been proposed as a possible synonym for this taxon (Warncke, 1980; Schwarz et al., 1996; Přidal, 2004; Aguib et al., 2010; Kuhlmann et al., 2021). The lectotype and a paralectotype of Anthidium sinuatum, designated as such by Donald B. Baker in 2003 (unpublished), are deposited in the Oxford University Museum of Natural History. Photographs of these specimens were examined and were determined to be the taxon commonly referred to as Anthidium loti Perris, 1852 (Fig. 7). We hereby recognize this unpublished designation as detailed below and remove A. sinuatum from synonymy with P. nanum. We treat A. sinuatum as a nomen oblitum because to the best of our knowledge, it has never been used as a valid name after 1899. We consider A. loti a nomen protectum, since this name has been used as the presumed valid name for this particular taxon in at least 25 works, published by at least ten authors in the immediately preceding 50 years and encompassing a span of not less than ten years (Supporting Information, Appendix S2), thus fulfilling both Articles 23.9.1.1 and 23.9.1.2 of the Code. Anthidium sinuatum Lepeletier de Saint Fargeau, 1841: 374–375, ♀ ♂. Type locality: ‘Espagne. Musées de France et du général Dejean’ [Spain. Museums of France and of the General Dejean]. Lectotype ♀, designated by D. B. Baker 2003, published here: ‘[unreadable]’, ‘sinuatum ♀ ’, ‘ Lectotype Anthidium sinuatum Lep., 1841 D.B. Baker des. 2003’ [red label] (OUMNH) (Fig. 7); Paralectotype: ♀, ‘ Paralectotype Anthidium sinuatum Lep., 1841 D.B. Baker des. 2003’ [yellow label] (OUMNH). Warncke (1980) also proposed Anthidium floripetum Eversmann, 1852 as a synonym for this taxon but an examination of the type material of A. floripetum in the ISZP collection revealed that the latter in fact belongs to the genus Icteranthidium Michener, 1948 and is thus not applicable to this taxon. We hereby transfer the epithet and designate a female lectotype and a male paralectotype for Icteranthidium floripetum. To the best of our knowledge, this species epithet has never been used in combination with the genus Icteranthidium; whether I. floripetum represents a distinct species or a junior or senior synonym of an existing species must still be investigated. We call attention to the fact that while we designate a lectotype for I. floripetum, we are not using this name as the presumed valid name for any particular taxon, but are simply proposing a new combination. The validity of this name must be properly investigated, ideally within the framework of a revision of the genus Icteranthidium. Icteranthidium floripetum (Eversmann, 1852), comb. nov. Anthidium floripetum Eversmann, 1852: 83–84, ♀ ♂. Type locality: ‘in prov. Orenburgensi’ [Russia: Orenburg Prov.]. Lectotype, ♀, by present designation: ‘ Spask Aug’, [golden disc], ‘ floripetum ♀ Evers.’, ‘ Lectotypus ♀ Anthidium floripetum Eversmann, 1852 design. Fateryga et Proshchalykin 2019’ [red label] (ISZP) (Fig. 8). Paralectotype: ♂, ‘ Indersk’ [currently Inder Distr. in Atyrau Prov. of Kazakhstan], ‘ floripetum ♂. Evers.’, ‘ Paralectotypus ♂ Anthidium floripetum Eversmann, 1852 design. Fateryga et Proshchalykin 2019’ [red label] (ISZP). Anthidium peregrinum Costa, 1885, described from Sardinia, has alternately been placed in synonymy with P. nanum (as P. lituratum in Warncke, 1980; Přidal, 2004) and with P. stigmaticorne (as P. leucostoma in Nobile, 1995); other authors have treated this taxon as a subspecies of P. nanum (as P. lituratum Rasmont et al., 1995; Ornosa et al., 2008). Costa’s original description does not provide enough detail to clarify to which taxon this name applies. Only an examination of the type material will clarify the status of P. peregrinum. For the moment we include it as a synonym of P. stigmaticorne (see below). Diagnosis female: The female of P. nanum may be distinguished from other members of this complex by the following combination of characters: overall impression of cuticle shiny; punctation on terga, namely on T1–T2, narrow and dense with shiny interspaces between puncture; posterior margin of T1– T4 shiny (Fig. 9A, C, E). Nevertheless, in their zone of overlap, differentiating females of P. nanum from those of P. scapulare and, in some cases, from those of P. stigmaticorne may be challenging. Diagnosis male: The male of P. nanum may be distinguished from other members of this complex by the following combination of characters: gonostylus nearly parallel-sided, only slightly wider at apex than at base (approximately 1.25 times wider at apex than at base in P. scapulare and considerably more so in other species) (Fig. 10A); notch at apex of gonostylus at least as deep as notch is wide, nearly centred at apex and V-shaped (notch either deeper or less deep in other species or broadly rounded) (Fig. 10A); lateral comb on S5 small, with longest teeth shorter than maximal width of hind basitarsus (either larger in other species, as in P. scapulare, P. tenellum and P. palestinicum, or even smaller in other species, such as P. stigmaticorne and P. cribratum) (Fig. 11A); comb of S5 wider than arm preceding it; posterior, premarginal brush on S3 with hairs hooked at tips (hairs unhooked in P. tenellum and P.cribratum) (Fig. 12A); shiny, hairless zone on S3 between posterior premarginal brush of hairs and anterior zone of dense, velvety pilosity chevronshaped, with the tip of chevron extending anteriorly as a carina along the midline of sternum (Fig. 12A) (no shiny hairless zone in P. stigmaticorne; extending nearly straight across sternum in P. scapulare, with similar medial extension; raised hairless zone matte in P. tenellum, extending across width of sternum but without medial extension; raised hairless zone in P. cribratum trapezoidal, without medial extension); raised zone on S2 shiny, dark and in shape of chevron (Fig. 13A); hairs on ventral surface of trochanter 3 short and of even length but not velvety, tuft of longer hairs present at base of trochanter. The male of P. nanum is morphologically similar to both P. palestinicum and P. kaspareki; for more information concerning the differentiation of these three species, see the section entitled ‘Diagnosis male’ for P. kaspareki. Geographical variation: Males of P. nanum are mostly black, with yellow markings that are restricted to the clypeus (typically entirely pale yellow) and present on the paraocular area laterally to inner margin of the eye and posteriorly as far as the basal margin of the clypeus, as well as a single small yellow marking on the preoccipital margin just posterior to each eye, and paired yellow markings laterally on T1–T4, T1–T5 or T1–T6. In some populations (e.g. in Switzerland, Crimea, Dagestan and Azerbaijan), tarsi, tibiae and apex of femora are dark orange-yellow; in other populations (e.g. those in France), these parts are bright lemon-yellow. While most individuals have lemonyellow markings on the terga, in some populations (e.g. Azerbaijan, Corsica and Germany), these markings are pale yellow. In females, the colour of the clypeus is variable and may range from entirely black, to black with lateral yellow spots, to yellow with black markings, to entirely yellow. It should also be noted that the colour of the mandibles, used as a diagnostic feature in other keys [e.g. that of Aguib et al. (2010)], is variable and can range from nearly black in certain specimens to yellow in others. Paired, lateral yellow markings on the terga are present from T1–T4 or T1–T5., Published as part of Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2022, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), pp. 1287-1337 in Zoological Journal of the Linnean Society 195 on pages 1296-1301, DOI: 10.1093/zoolinnean/zlab062, http://zenodo.org/record/5817276, {"references":["Mocsary S. 1880. Ujabb adatok temesmegye hartyaropu faunajahoz. Mathematikai es Termeszettudomanyi Kozlemenyek 16: 1 - 70.","Panzer GWF. 1801. Favnae insectorvm Germanicae initia, oder, Deutschlands Insecten, Vol. 14, Hf. 80. Nurnberg: Felssecker.","Gmelin JF. 1790. Caroli a Linne. Systema naturae per regna tria naturae: secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tom 1. Pars 5. Leipzig: G. E. Beer, 2225 - 3020.","Eversmann E. 1852. Fauna hymenopterologica volgouralensis (Continuatio). Bulletin de la Societe Imperiale des Naturalistes de Moscou 25: 1 - 137.","Bohart RM, French LD. 1986. Designation of chrysidid lectotypes in the Mocsary collection at the Hungarian National Museum, Budapest (Hymenoptera: Chrysididae). Pan-Pacific Entomologist 62: 340 - 343.","Alfken JD. 1936. Beitrag zur Kenntnis der Bienenfauna von Mittel-Italien. Bollettino dell'Istituto di Entomologia della Universita degli Studi di Bologna 9: 104 - 112.","Baldock D, Wood TJ, Cross I, Smit J. 2018. The bees of Portugal (Hymenoptera: Apoidea: Anthophila). Entomofauna Zeitschrift fur Entomologie, Supplement 22: 1 - 164.","Westrich P. 1990. Die Wildbienen - Baden-Wurttembergs, 2 nd edn. Stuttgart: Eugen Ulmer GmbH & Co.","Warncke K. 1980. Die Bienengattung Anthidium Fabricius, 1804 in der Westpalaarktis und im turkestanischen Becken. Entomofauna 1: 119 - 210.","ICZN. 1973. XVIIth International Congress of Zoology, Monaco, 1972. Minutes of the Meeting of the Section on Nomenclature, 28 September 1972. Bulletin of Zoological Nomenclature 29: 179 - 189.","Schwarz M, Gusenleitner F, Westrich P, Dathe H. 1996. Katalog der Bienen Osterreichs, Deutchlands und der Schweiz. Entomofauna, Supplement 8: 1 - 398.","ICZN. 1999. International Code of Zoological Nomenclature. 4 th edn. London: The International Trust for Zoological Nomenclature.","Baker DB. 1996. Priorities of publication for some nineteenthcentury works describing new taxa of Hymenoptera, principally Apoidea. Archives of Natural History 23: 299 - 302.","Lepeletier de Saint Fargeau ALM. 1841. Histoire naturelle des insectes. Hymenopteres, Vol. 2. Paris: Roret.","Pridal A. 2004. Checklist of the bees in the Czech Republic and Slovakia with comments on their distribution and taxonomy (Insecta: Hymenoptera: Apoidea). Acta Universitatis Agriculturae et Silviculturae Mendelianae Brunensis 52: 29 - 66.","Aguib S, Louadi K, Schwarz M. 2010. Les Anthidiini (Megachilidae, Megachilinae) d'Algerie avec trois especes nouvelles pour ce pays: Anthidium (Anthidium) florentinum (Fabricius, 1775), Anthidium (Proanthidium) amabile Alfken, 1932 et Pseudoanthidium (Exanthidium) enslini (Alfken, 1928). Entomofauna 31: 121 - 152.","Kuhlmann M, Ascher JS, Dathe HH, Ebmer AW, Hartmann P, Michez D, Muller A, Patiny S, Pauly A, Praz C, Rasmont P, Risch S, Scheuchl E, Schwarz M, Terzo M, Williams PH, Amiet F, Baldock D, Berg O, Bogusch P, Calabuig I, Cederberg B, Gogala A, Gusenleitner F, Josan Z, Madsen HB, Nilsson A, Odegaard F, Ortiz-Sanchez J, Paukkunen J, Pawlikowski T, Quaranta M, Roberts SPM, Sa ropataki M, Schwenninger H-R, Smit J, Soderman G, Tomozei B. 2021. Checklist of the Western Palaearctic Bees (Hymenoptera: Apoidea: Anthophila). Available at: http: // westpalbees. myspecies. info (accessed 2020).","Costa A. 1885. Notizie ed osservazioni sulla geo-fauna Sarda, Memoria quarta. Atti della Reale Accademia delle Scienze Fisiche e Matematiche di Napoli 2: 13, 1 - 31.","Nobile V. 1995. Api (Insecta, Hymenoptera) nuove o poco note di Sicilia e di Sardegna. Bollettino delle Sedute della Accademia Gioenia di Scienze Naturali in Catania 28: 147 - 159.","Rasmont P, Ebmer A, Banaszak J, van der Zanden G. 1995. Hymenoptera Apoidea Gallica. Bulletin de la Societe Entomologique de France 100: 1 - 98.","Ornosa C, Ortiz-Sanchez FJ, Torres F. 2008. Catalogo de los Megachilidae del Mediterraneo occidental (Hymenoptera, Apoidea). III. Anthidiini y Dioxyini. Graellsia 64: 61 - 86."]}

Stelis leucostoma Costa described Stelis leucostoma from Sardinia (Costa, 1883). Warncke (1980), Rasmont et al. (1995), Přidal (2004) and Ornosa et al. (2008) each placed S. leucostoma in synonymy with P. nanum (as either Anthidium lituratum or Pseuodanthidium lituratum). However, Nobile (1995) considered this taxon a distinct species. Nobile noted that his decision to assign this name to this particular taxon was based on Costa’s original description of Stelis leucostoma but he gives no indication of having examined the type series. Nobile illustrated the male gonostylus and sterna of a specimen collected close to Nuoro (Sardinia); this specimen, as well as others that we have examined from Sardinia, agree with our concept of P. stigmaticorne. At the present time we place Stelis leucostoma in synonymy with P. stigmaticorne. However, both P. stigmaticorne and P. nanum are known from Sardinia, and Costa’s description of Stelis leucostoma is suggestive of both. Costa’s type series must thus be examined to determine whether the name Stelis leucostoma refers to the taxon that we refer to as P. stigmaticorne or to another., Published as part of Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2022, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), pp. 1287-1337 in Zoological Journal of the Linnean Society 195 on page 1310, DOI: 10.1093/zoolinnean/zlab062, http://zenodo.org/record/5817276, {"references":["Costa A. 1883. Notizie ed osservazioni sulla geo-fauna Sarda, Memoria seconda, Risultamento di ricerche fatte in Sardegna nella primavera del 1882. Atti della Reale Accademia delle Scienze Fisiche e Matematiche di Napoli 2: 2, 1 - 109.","Warncke K. 1980. Die Bienengattung Anthidium Fabricius, 1804 in der Westpalaarktis und im turkestanischen Becken. Entomofauna 1: 119 - 210.","Rasmont P, Ebmer A, Banaszak J, van der Zanden G. 1995. Hymenoptera Apoidea Gallica. Bulletin de la Societe Entomologique de France 100: 1 - 98.","Pridal A. 2004. Checklist of the bees in the Czech Republic and Slovakia with comments on their distribution and taxonomy (Insecta: Hymenoptera: Apoidea). Acta Universitatis Agriculturae et Silviculturae Mendelianae Brunensis 52: 29 - 66.","Ornosa C, Ortiz-Sanchez FJ, Torres F. 2008. Catalogo de los Megachilidae del Mediterraneo occidental (Hymenoptera, Apoidea). III. Anthidiini y Dioxyini. Graellsia 64: 61 - 86.","Nobile V. 1995. Api (Insecta, Hymenoptera) nuove o poco note di Sicilia e di Sardegna. Bollettino delle Sedute della Accademia Gioenia di Scienze Naturali in Catania 28: 147 - 159."]}

PSEUDOANTHIDIUM KASPAREKI LE DIVELEC & LITMAN, SP. NOV. (FIGS 11G, 12G, 13G, 23E, 27A, C, E) Z o o b a n k r e g i s t r a t i o n: u r n: l s i d: z o o b a n k. org:act: 418E5EFF-5484-44C1-B45E-C9FFE45B931E Type material: Holotype. TURKEY: ♂, Side, 70 km E Antalya, 29.vii–07.viii.2001, P. Tymer (CSEC). Paratypes: TURKEY: 1♂, Yalova, Between Izmit & Bursa, 16.vi.1976 (MNHN); 1♂, 1♀, Between Kula & Usak, 2 km before Güre, 19.vi.1976 (MNHN). Additional material: GREECE: 1♂, Southern Aegean, Empona, 18.vi.1958, Mavromoustakis (OLML); TURKEY: 1♂, Alanya, 10 km W Alanya, Konakli, 36.58N, 31.89E, 01.viii.2009, Schmid-Egger (CSEC); 1♂, Mersin, Mut, Sertavul, 1300 m, 07.vi.1968, J. Gusenleitner (OLML); 1♂, Side, 70 km E Antalya, 29.vii–07.viii..2001, P. Tymer (CSEC); 1♂, Antalya, vi.1970, C. Holzschuh (OLML); 3♀♀, Antalya, Avsallar, 15.viii.1989, M. Blösch (ZSM); 1♂, Bursa, Ulu Dagi Mts, Barakli envir., 2000, M. Obořil (OLML); 1♂, Çanakkale, Ezine, 6 km N, 35 m, 27.vi.2006, J. Halada (OLML); 3♂♂, Çanakkale, Ezine, 6 km N, 35 m, 27.vi.2006, M. Halada (OLML); 1♂, Çanakkale, Troya, 1933 (OLML); 1♂, Eskişehir, İnönü, 800 m, 01.viii.1991, K. Warncke (OLML); 1♂, Hakkari, Semdinli, 1350 m, 09.viii.1983, K. Warncke (MSC); 1♀, Hatay, Yayladaği, 11.vi.1998, Ma. Jalada (MSC); 1♂, Kars, Pasli, 50 km S Kars, 01.vii.1997, Ma. Jalada (MSC); 2♂♂, Kütahya, 28 km SSE, 1110 m, 12.vii.2006, J. Halada (MSC); 1♂, 1♀, Mansisa, Salihli, 15 km SEE, 170 m, 02.vii.2006, M. Kadlecová (OLML); 4♂♂, 6♀♀, Mansisa, Salihli, 15 km SEE, 170 m, 02.vii.2006, J. Halada (OLML); 1♂, 1♀, Mansisa, Salihli, 15 km SEE, 170 m, 02.vii.2006, J. Halada (MSC); 2♂♂, Mansisa, Salihli, 15 km SEE, 170 m, 02.vii.2006, M. Halada (OLML); 3♂♂, Mansisa, Salihli, 30 km SEE, 430 m, 29.vi.2006, M. Halada (OLML); 2♂♂, 3♀♀, Mansisa, Salihli, 30 km SEE, 430 m, 29.vi.2006, J. Halada (OLML); 2♂♂, Mansisa, Salihli, 35 km SEE, 900 m, 30.vi.2006, J. Halada (OLML); 2♂♂, Mansisa, Salihli, 35 km SEE, 900 m, 30.vi.2006, M. Halada (OLML); 3♂♂, 3♀♀, Mansisa, Salihli, 40 km SEE, 150 m, 26.vi.2006, J. Halada (OLML); 1♀, Mansisa, Salihli, 40 km SEE, 150 m, 26.vi.2006, M. Halada (OLML); 1♂, 1♀, Mugla, Mugla, Univ. Campus, 700 m, 2015, H. Kavak (OLML); 1♂, Nevşehir, Ürgüp, 5 km E, 1100 m, 24.viii.1991, K. Warncke (OLML); 1♀, Osmaniye, Hasanbeyli, 18–19.vi.1991, Kuorna (OLML); 1♀, Şanlıurfa, Urfa, 29.v.1970, J. Schmidt (SMF); 1♂, Şırnak, Siirt, ‘östl. Şırnak /Siirt’, 04.vii.1977, K. Warncke (OLML); 1♀, Asia Minor, 1890 (ZSM). Distribution: Greece, Turkey (Fig. 26A). Host plant associations: Unknown. Description female: Measurements BL = 6.5 mm, IW = 2.1 mm. Head: Mandible yellow, except for teeth and anterior margin, which are black. Clypeus yellow with diffuse black stains medially, apical margin slightly emarginate, basal half shiny with sparse, coarse punctation (as coarse as in supraclypeal area), with distinct flat interspaces between punctures; punctation becomes finer, denser apically where punctures almost undistinguishable, integument matte. Paraocular area yellow. Antenna with anterior face of antennal flagellum more or less brownish-ferruginous with third and fourth segments entirely light ferruginous; scape and pedicel relatively long (3.66 as long as wide and 1.66 as long as wide, respectively); flagellar segments much shorter than wide (L/W: F1 = 0.8; F2 = 0.72; F3 = 0.57; F4–F5 = 0.53; F6– F7 = 0.66; F8–F9 = 0.73), except last segment (L/W = 1.35) (measurements made on a single specimen). Frons matte, with honeycomb areolate punctation becoming slightly coarser toward clypeus. Vertex with pair of lateral yellow spots on posterior margin, punctation fine, dense with shiny, minute, ridge-like interspaces between punctures (Fig. 27A); punctures as fine as those on frons, clearly finer than those of mesonotum and terga (Fig. 27A, C, E). Gena honeycomb areolate with wide, shallow punctures. Pilosity on head white or whitish, except for short, dense golden setae covering anterior margin of clypeus and on mandible and labrum. Mesosoma: Punctation of scutum and scutellum honeycomb areolate and rather matte, long median sulcus reaching more than half of scutum length, parapsidal furrow distinct, approximately three to four puncture diameters in length (Fig. 27C); integument mostly black, anterior margin of scutum with pair of lateral, rectangular yellow markings (Fig. 27C). Pronotal lobe black with yellow apical spot (Fig. 27C). Axilla black with yellow posterior margin (Fig. 27C). Scutellum black with pair of yellow rectangular markings laterally on posterior margin (Fig. 27C); punctation honeycomb areolate, rather matte (Fig. 27C). Tegula with yellow spot anteriorly, otherwise ferrugineous (Fig. 27C). Mesepisternum and mesosternum honeycomb areolate with dense, coarse punctation anteriorly with sharp carina-shaped interspaces, becoming shallow, sparse posteriorly. Metepisternum with superficial, dense, minute punctation. Propodeum with lateral face finely, densely punctate; posterior face basally with row of coalescent, shallow punctures interrupted medially; rest of posterior face smooth, shiny. Tarsi, tibiae, tibial spurs, apical half of femora ferrugineous; base of tibia, apex of femora with diffuse yellowish stains. Inner surface of foretibia with rake of thick, golden setae. Basitarsal brushes gold. Tarsal segments with some thick gold setae apically. Wings infuscate. Metasoma: Integument shiny, terga black with yellow markings (Fig. 27E). Carina separating dorsal and anterior declivous faces of T 1 with fringe of short, golden hairs mostly two puncture widths long (Fig. 27E); T 1– T 4 with pair of lateral, yellow, ovoid spots increasing in size from T 1 to T 4 (Fig. 27E); T 5 with lateral yellow spots. Terga with coarse, irregular punctation (Fig. 27E); punctures on discs of T 1– T 4 as large or larger than those of mesonotum, much larger than punctation on depressions of T 1– T 4 (Fig. 27E); punctation of T 1 distinctly sparser in basal half, punctures separated by flat smooth interspaces (distances reaching one-third to half the diameter of a puncture), while punctures of apical half separated by narrow ridge-like interspaces; punctation of T 2– T 4 coarse, sparse on disc, punctures separated by flat, smooth interspaces (distances reaching one-third to half of puncture diameter), such that integument appears shiny (Fig. 27E); punctation much denser on apical depression than on discs on T 2– T 4, with punctures separated only by carina-shaped interspaces, nearly coalescent on T 3– T 4 (Fig. 27E). T 5 matte, with dense punctation on disc, punctures separated only by narrow, ridge-like interspaces; on depression punctation coalescent. T 6 coriaceous with deep coalescent punctures on entire surface. T 1– T 4 with broad smooth apical margin, length equals 1.3– 1.4 diameter of largest punctures (Fig. 27E); T 5 with smooth, narrow apical margin, length equals diameter of one large puncture medially. Scopa white. Diagnosis female: The female of P. kaspareki may be distinguished from other members of this complex by the following combination of characters: punctures on terga comparatively broad, wider than those of mesonotum and remarkably sparse on discs of T 2– T 4 and with flat smooth interspaces that give integument a shiny appearance (interspaces often reach onethird to half of a puncture diameter) (Fig. 27C, E); largest punctures on black part of scutellum smaller in diameter and denser than largest punctures on T 2 (Fig. 27C, E); punctation of scutum is contiguous and irregular, scutum thus appearing matte (Fig. 27C); punctation on vertex between lateral ocellus, eye and post-occipital margin small but distinct, with shiny narrow spaces between most punctures (Fig. 27A); clypeus short, less than 0.8 mm from base to apical margin. The females of P. kaspareki and P. cribratum exhibit coarse, sparse punctation on the metasoma (Figs. 17F, 27E). However, the two species may be differentiated based on the punctation of the vertex and mesonotum. In P. kaspareki, the punctation on these zones is finer and denser than that on the metasomal terga and interspaces between punctures are carina-like, so that the integument appears rather matte (Fig. 27A, C, E). In P. cribratum, the punctures of the vertex and mesonotum are almost as large as those of the metasomal terga and punctures are clearly separated by smooth interspaces, so that the integument appears relatively shiny (Fig. 17B, D, F). Due to its coarse metasomal punctation and the relatively fine, dense punctation on the vertex and mesonotum, P. kaspareki is most similar to P.palestinicum. However, it may be distinguished by the presence of sparser punctation on the metasomal terga of the former species. In P. kaspareki, the punctures on discs T 2– T 4 are strongly separated by flat, smooth interspaces, giving the integument a shiny appearance (Fig. 27E). In P. palestinicum, the punctures of discs T 2– T 4 are only separated by narrower, carina-shaped interspaces, making the integument appear matte (Fig. 21F). In P. kaspareki, the punctures on the vertex are separated by shiny, narrow yet distinct interspaces (Fig. 27A); in P. palestinicum, on the other hand, the interspaces between punctures on the vertex are carina-shaped, making the integument appear matte (Fig. 21B). The clypeus is also longer in P. palestinicum (> 0.9 mm measured from base to apical margin). Pseudoanthidium stigmaticorne is a morphologically variable species that is also present in Turkey, where it may be confused with P. kaspareki. However, in P. kaspareki the metasomal punctation is much larger than the punctation on the scutum (Fig. 27C, E), while in P. stigmaticorne, it is as large or only slightly larger (Fig. 17C, E). In P. kaspareki, the punctation at the base of T 1 is distinctly coarser and sparser than it is apically (Fig. 27E), whereas the size and the density of the punctation of T 1 is regular across the whole surface of the tergum in P. stigmaticorne (Fig. 17E). In P. kaspareki, the punctation of T 2– T 4 is distinctly coarser and sparser on the disc than on the apical depression, where punctures are almost coalescent (Fig. 27E). In contrast, in P. stigmaticorne, the punctation is the same size and density across the whole surface of the metasomal terga, where it is usually separated by minute ridge-like interspaces (Fig. 17E). Punctures on discs T 2– T 4 in P. kaspareki are separated by smooth interspaces of irregular width (reaching from one-quarter up to half of a puncture diameter), giving the integument a shiny appearance (Fig. 27E). In P. stigmaticorne, punctures of T 2– T 4 are usually separated by regular, narrow interspaces (reaching at most one-fifth of a puncture diameter, rarely one-quarter or more) across their whole surface, so that the integument appears comparatively matte (Fig. 17E). Description male: Measurements BL = 7.5–8 mm, IW = 2.1–2.2 mm. Head: Integument black except where noted otherwise. Mandible yellow, except for teeth and anterior margin, which are black. Clypeus pale yellow; density of punctation at base of clypeus irregular, with some punctures very close (separated by less than one-half a puncture diameter), others separated by up to several puncture diameters, sparsely punctate area extends medially halfway down clypeus; apical region with relatively dense punctation, spaces between punctures Mesosoma: Integument mostly black. Scutum with punctation dense, coarse, honeycomb areolate, with spaces between punctures Metasoma: Terga with integument shiny. T 1– T 5 black with lateral yellow spots, T 6 in some specimens nearly entirely yellow, except for black band at base and reduced black markings medially between lateral yellow spots, in other specimens, T 6 entirely black; T 7 yellow. Punctures on T 1– T 3 broad with shiny spaces between punctures Diagnosis male: Pseudoanthidium kaspareki is morphologically most similar to P. nanum and P. palestinicum. In males, P. kaspareki may be differentiated from P. palestinicum by the shape of the lateral comb on S5, which is extremely broad and chevron-shaped in P. palestinicum (Fig. 11E) and narrower and more evenly rounded in P. kaspareki (Fig. 11G); and in the shape of the penis valve, which is parallel-sided with a rounded tip in P. kaspareki (Fig. 23E) and which gradually tapers to a pointed end in P. palestinicum. Finally, the gonostylus of P. kaspareki is comparatively slender and only slightly convex laterally (Fig. 23E), whereas the gonostylus of P. palestinicum is broader and more strongly convex laterally (Fig. 23A). In P. kaspareki, the curve of the inner margin of gonostylus is strongest apically, while the curve of the outer margin is strongest about twothirds of way between base and apex (Fig. 23E). In P. palestinicum, the curve of both inner and outer margins is strongest about two-thirds of way between base and apex (Fig. 23A). Pseudoanthidium kaspareki may be differentiated from P. nanum by the size of the lateral comb on S5, which is slightly larger in P. kaspareki than in P. nanum (Fig. 11A, G); in the shape of the gonostylus, which is laterally rounded in P. kaspareki (Fig. 23E) and nearly parallel-sided in P. nanum (Fig. 10A); in the depth of the apical notch of the gonostylus, which is less deep in P. kaspareki (Fig. 23E) than in P. nanum (Fig. 10A); and by the punctures of T 1– T 3, which are considerably broader than those of P. nanum. Etymology: We are pleased to name this species for our colleague Max Kasparek, in recognition of his significant contributions to the systematics of anthidiine bees.

Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler, Praz, Christophe J. (2022): Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini). Zoological Journal of the Linnean Society 195 (4): 1-51, DOI: 10.1093/zoolinnean/zlab062

Pseudoanthidium gregoriense Pseudoanthidium gregoriense was described by Nobile (1990) as an island endemic from Sicily. Originally described as a subspecies of P. alpinum, Nobile later raised P. gregoriense to species status (1995). As mentioned above for Stelis leucostoma, Nobile (1995) draws parallels between P. gregoriense, P. leucostoma and P. lituratum. We examined four specimens of this taxon identified by Nobile (including one of his paratypes) and a photograph of the holotype, plus two other female specimens from Sicily. These specimens agree with our concept of P. stigmaticorne., Published as part of Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2022, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), pp. 1-51 in Zoological Journal of the Linnean Society 195 (4) on page 26, DOI: 10.1093/zoolinnean/zlab062, http://zenodo.org/record/5817276, {"references":["Nobile V. 1990. Il genere Anthidium Fabricius 1804 e affini (Hymenoptera, Apoidea, Megachilidae) in Sicilia, con descrizione di Pseudoanthidium alpinum gregoriense subsp. n. Animalia 16: 131 - 145.","Nobile V. 1995. Api (Insecta, Hymenoptera) nuove o poco note di Sicilia e di Sardegna. Bollettino delle Sedute della Accademia Gioenia di Scienze Naturali in Catania 28: 147 - 159."]}

PSEUDOANTHIDIUM SCAPULARE (LATREILLE, 1809) (FIGS 2E, 9B, D, F, 10B, 11B, 12B, 13B, 14) Anthidium scapulare Latreille, 1809: 46–47, 227, pl. 1, Fig. 8, ♀, Type locality: ‘Patria incognita’ [origin unknown]. Lectotype, ♀, designated by Tkalců, 1966: 63: ‘ scapulare ♀ ’, ‘ Lectotype of Anthidium scapulare Latr. ♀ 1965 Tkalců det.’, ‘Type O.U.M.’ (OUMNH). Anthidium frontale Lepeletier de Saint Fargeau, 1841: 377–378, ♀, synon. nov. Type locality: ‘ Oran; envoyé par mon fils. Ma collection’ [Oran, Algeria; sent by my son. [My collection] Lectotype, ♀, by present designation: ‘anthidium frontale LP. ♀.’, ‘ Pseudoanthidium (Paraanthidiellum) lituratum Panz. J. Pasteels det. 1966’, ‘ Lectotype Anthidium frontale Lep. Le Divelec des. 2020’ [red label] (MNHN), ‘ Pseudoanthidium scapulare (Latreille, 1809) Le Divelec det. 2020’, ‘MNHN, Paris EY24646’ (Fig. 14). Material examined: 89 females, 83 males (see Supporting Information, Table S1 for specimen data). Distribution: Algeria, France, Morocco, Portugal, Spain (including Mallorca) and Tunisia (Fig. 6B). Host-plant associations: A steraceae Algeria Centaurea calcitrapa L., Carduus sp., Onopordum macracanthum Schousb., Atractylis serratuloides (Cass.) DC., Cynara cardunculus L.; Verbenaceae Algeria Verbena officinalis L. (Aguib et al., 2010). Remarks: Tkalců (1966) and Warncke (1980) considered A. scapulare as a junior synonym of P. nanum (= as lituratum Panzer), yet an examination of photographs of the lectotype of P. scapulare (Latreille, 1809) demonstrate that P. scapulare is distinct from P. nanum. Tkalců (1975) also came to this conclusion, but recognized P. scapulare as a subspecies of P. nanum (as lituratum Panzer), rather than a distinct species. Přidal (2004) eventually removed P. scapulare from synonymy with P. nanum (as P. lituratum), considering it a distinct species. The lectotype of P. scapulare is a badly worn female, but still corresponds to the morphological criteria associated with P. scapulare, namely in the relatively fine punctation on the second tergum and the overall aspect of the integument, which is glittery but not shiny. The male of P. scapulare is also clearly distinct from the males of other members of the complex, namely in the shape of the gonostylus, the shape of the sternal comb and the morphology of S2–S4. Ultra-conserved element analyses further demonstrate that P. scapulare is genetically distinct from other taxa in the complex, in particular from P. nanum. We thus agree with Přidal (2004) and Aguib et al. (2010) and consider P. scapulare as a distinct species from P. nanum. A specimen of Anthidium frontale Lepeletier de Saint Fargeau, 1841 was discovered in the collections at the MNHN (Fig. 14). The handwriting on the label bearing the species name is discernible as that of Lepeletier de Saint Fargeau. As is typical of the Lepeletier de Saint Fargeau collection, the specimen bears no locality label; in this particular collection, such information is most often found pinned in the entomological drawer containing the specimens. Unfortunately, the individual in question has been dissociated from both its original drawer and some of its original labels; no information concerning its original collection locality is available. Although the specimen is not explicitly labelled as a type, the unusual yellow markings on the face are detailed in the species description of A. frontale, giving us reason to believe that this individual was used as this basis for the original description. Furthermore, the specimen is prepared on a short, thick pin characteristic of the Lepeletier de Saint Fargeau collection. We hereby designate this specimen as the lectotype for A. frontale. A careful examination indicates that it corresponds to the same taxon that we refer to here as P. scapulare and it bears the extensive yellow markings frequently seen in northern African populations of this species. Anthidium frontale was, until now, considered a synonym of Anthidiellum strigatum contractum Latreille, 1809 (Warncke, 1980). We hereby remove A. frontale from synonymy with Anthidiellum strigatum contractum and place it in synonymy with P. scapulare. Diagnosis female: The female of P. scapulare may be distinguished from other members of this complex by the following combination of characters: overall impression of cuticle glittery but not shiny; punctation on terga, namely on T 1– T 2, narrow and dense with finely shagreened interspaces between punctures; posterior margin of T 1– T 4 glittery but not shiny (Fig. 9B, D, F). In their zone of overlap, differentiating females of P. scapulare from those of P. nanum is challenging (see ‘Diagnosis female’ under P. nanum for comparison). Diagnosis male: The male of P. scapulare may be distinguished from other members of this complex by the following combination of characters: gonostylus approximately 1.25 times wider at apex than at base (nearly parallel sided in P. nanum and considerably more so in other species) (Fig. 10B); notch strongly U-shaped, deeper than notch is wide at opening and slightly offset at apex of gonostylus, so that interior tip of the notch is visibly wider than the exterior (notch either less deep in other species or V-shaped) (Fig. 10B); lateral comb on S 5 in shape of chevron, with longest teeth approximately equal to or greater in length than maximum width of hind basitarsus (either larger in other species, as in P. palestinicum, mitten-shaped, as in P. tenellum or even smaller in other species, such as P. nanum, P. stigmaticorne and P. cribratum) (Fig. 11B); posterior margin of S4 deeply emarginate at centre (not emarginate in other species); posterior, premarginal brush on S3 with hairs hooked at tips (hairs unhooked in P. tenellum and P. cribratum) (Fig. 12B); shiny, hairless zone on S3 between posterior premarginal brush of hairs and anterior zone of dense, velvety pilosity stretching nearly straight across entire width of sternum, with medial extension extending anteriorly along the midline of sternum (no shiny hairless zone in P. stigmaticorne; raised hairless zone matte in P. tenellum, extending across width of sternum but without medial extension; raised hairless zone in P. cribratum trapezoidal, without medial extension) (Fig. 12B); raised zone on S2 shiny, dark and in shape of chevron; posterior margin of S2 usually gently emarginate medially (not so in other species) (Fig. 13B); hairs on ventral surface of trochanter 3 very short, velvety and dense, tuft of longer hairs present at base of trochanter., Published as part of Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2022, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), pp. 1-51 in Zoological Journal of the Linnean Society 195 (4) on pages 16-18, DOI: 10.1093/zoolinnean/zlab062, http://zenodo.org/record/5817276, {"references":["Latreille PA. 1809. Memoire sur le genre d'Anthidie, Anthidium, de Fabricius, classe des insectes, ordre des hymenopteres, famille des Apiaires. Annales du Museum d'Histoire Naturelle de Paris 13: 1 - 507.","Tkalcu B. 1966. Revision of some Latreille's European species of the tribe Anthidi [i] ni with the description of a new species (Hymenoptera; Apoidea, Megachilidae). Acta Entomologica Bohemoslovaca 63: 62 - 66.","Lepeletier de Saint Fargeau ALM. 1841. Histoire naturelle des insectes. Hymenopteres, Vol. 2. Paris: Roret.","Aguib S, Louadi K, Schwarz M. 2010. Les Anthidiini (Megachilidae, Megachilinae) d'Algerie avec trois especes nouvelles pour ce pays: Anthidium (Anthidium) florentinum (Fabricius, 1775), Anthidium (Proanthidium) amabile Alfken, 1932 et Pseudoanthidium (Exanthidium) enslini (Alfken, 1928). Entomofauna 31: 121 - 152.","Warncke K. 1980. Die Bienengattung Anthidium Fabricius, 1804 in der Westpalaarktis und im turkestanischen Becken. Entomofauna 1: 119 - 210.","Tkalcu B. 1975. Sammelergebnisse der von RNDr. A. Hoffer geleiteten Algerien-Expeditionen in den Jahren 1971 und 1972 (Hymenoptera: Apoidea). 1. Teil: Megachilidae. Acta Rerum Naturalium Musei Nationalis Slovaci Bratislava 21: 165 - 190.","Pridal A. 2004. Checklist of the bees in the Czech Republic and Slovakia with comments on their distribution and taxonomy (Insecta: Hymenoptera: Apoidea). Acta Universitatis Agriculturae et Silviculturae Mendelianae Brunensis 52: 29 - 66."]}

PSEUDOANTHIDIUM PALESTINICUM (MAVROMOUSTAKIS, 1938), STAT. NOV. (FIGS 2G, 11E, 12E, 13E, 21B, D, F, 23A) Anthidium cribratum palestinicum Mavromoustakis, 1938: 16–17, ♀ ♂. Type locality: ‘ Migdab’ [possibly ‘Migdal’, Israel but not certain], 19.v.1931 (holotype ♀, allotype ♂, ‘ Tiberias’ [Israel], 3.vii.1931) (BMNH). Material examined: Seven females, 12 males (see Supporting Information, Table S1 for specimen data). Distribution: Israel and Palestine, Jordan, Lebanon, Syria and Turkey (Fig. 22A)., Published as part of Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2022, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), pp. 1-51 in Zoological Journal of the Linnean Society 195 (4) on page 31, DOI: 10.1093/zoolinnean/zlab062, http://zenodo.org/record/5817276, {"references":["Mavromoustakis GA. 1938. II. - On some Anthidiine bees from Palestine. Annals & Magazine of Natural History 11: 15 - 18."]}

Paranthidiellum karakalense Pseudoanthidium karakalense was described from Kara-Kala in Turkmenistan. Popov described P. karakalense as similar to P. astilleroi but with the female differing in certain morphological characters, including the punctation on the frons, the straight edge of the clypeal margin, the length of the first, second and third antennal segments and the paler yellow colour. He further noted that this taxon differed from P. lituratum in its punctation, size, coloration and male genitalia. A careful examination of Popov’s type material confirms the synonymy of this taxon with P. stigmaticorne. Warncke (1980) erroneously placed this taxon in synonymy with P. cribratum (as Anthidium lituratum cribratum). Diagnosis female: The female of P. stigmaticorne may be distinguished from other members of this complex by the following combination of characters: punctation on terga comparatively coarse, as large or larger than punctation on mesonotum and with shiny interspaces between punctures; largest punctures on black part of scutellum smaller in diameter than largest punctures on T2; punctation on vertex between lateral ocellus, eye and postoccipital margin small but distinct, with shiny narrow spaces between most punctures; clypeus either yellow, yellow with black markings or black; small, mostly less than 0.8 mm (Fig. 17A, C, E). In their zone of overlap (e.g. in Crimea), differentiating females of P. stigmaticorne from those of P. tenellum may be challenging in some cases. Diagnosis male: The male of P. stigmaticorne may be distinguished from other members of this complex by the following combination of characters: gonostylus approximately 1.5 times wider at widest point than at base (Fig. 18A–F); exterior margin of gonostylus relatively straight, inner margin more sharply rounded toward apex (Fig. 18A–F); notch at apex of gonostylus either as deep as opening of notch is wide and more or less centred at apex of gonostylus (most European and Central Asian populations) or notch significantly less deep than opening of notch is wide and strongly offset laterally (northern Africa, southern Italy) (notch deeper in other species) (Fig. 18A–F); lateral comb on S5 small, with longest teeth shorter than maximal width of hind basitarsus (comb larger in P. scapulare, P. nanum, P. tenellum and P. palestinicum) (Fig. 11C); comb of S5 no wider than arm preceding it (Fig. 11C); posterior, premarginal brush on S3 with hairs hooked at tips (hairs unhooked in P. tenellum and P. cribratum) and arranged in an open brushy arc (Fig. 12C); no shiny, hairless zone on S3 between posterior premarginal brush of hairs and anterior zone of dense, velvety pilosity (present in other species) (Fig. 12C); darkened raised zone across S2 with only short, sparse hairs, in shape of ‘moustache’ (Fig. 13C); hairs on ventral surface of trochanter 3 shaggy and uneven, not dense, with longer hairs toward base. Geographic variation: A considerable amount of variation with regard to colour pattern is observed in this taxon, even in individuals from the same geographic region. We attempt to describe some of the variation that we have seen and to associate it with particular geographic regions. In populations from Crimea, the clypeus in females is black and the yellow markings on the face are reduced to a rounded spot on either side laterally between the lateral clypeal margin and the eye. The maculations on head, thorax and metasoma are pale yellow. Paired spots behind the eyes are small and those on anterior margin of mesonotum are reduced to a single spot on either side of anterior margin. In some individuals, this spot may be rectangular and relatively long, representing about one-third of the anterior margin of the mesonotum; in others it is much smaller. The scutellum may be entirely black, or mostly black except for a single reduced spot on either side laterally, or with lateral spots larger, giving the scutellum the impression of being yellow with a black triangle in the centre. Yellow maculations are present on T1–T5, although those on T5 are sometimes much reduced. Legs are dark orange yellow and maculations on metasoma sub-rectangular with rounded corners. Maculations on T3–T5 do not reach lateral margin of terga. Some populations in southern mainland Italy are similar, although in these populations the clypeus is mostly yellow with some black medially. Populations in the Middle East are also similar to those in Crimea, except that maculations are brighter yellow, yellow spots on T1–T5 nearly reach the lateral margin of tergum on either side and legs are yellower. Females found in Turkey are similar but with metasoma visibly shinier and with legs and clypeus mostly yellow. Specimens from Spain and France are similar to those from the Middle East. There also appears to be a certain difference in size, with individuals from northern Africa and the Middle East smaller than those from elsewhere. Finally, most individuals are black and yellow but individuals from northern Africa also exhibit shades of red on the metasoma. The colour patterns seen in males are mostly similar to those in females from the same regions except that the clypeus in males is typically pale, creamy yellow. In some populations, such as those in Crimea, pale yellow maculations on metasoma are present on T1– T4, T5–T6 are entirely black and T7 is pale yellow. In the Middle East, paired maculations are present on T1–T6 and T7 is yellow or, alternatively, paired maculations are present on T1–T5, T6 is black and T7 is pale yellow. We have also examined specimens from Dagestan in which lemon-yellow maculations are present on T1–T5, T6 is black and T7 is pale yellow. Specimens from northern Africa are similar to those in Dagestan but the posterior margin of T6 is yellow. In individuals from France and Spain, paired maculations are present on T1–T6 and T7 is yellow, although this varies considerably. The notch at the apex of the gonostylus is approximately as deep as the opening of the notch is wide and more or less centred at apex of gonostylus in most individuals. However, in Middle Eastern populations the gonostylus is more flared at the apex (Fig. 18B) and in northern African populations the gonostylus is strongly flared at the apex and the notch is much less deep than it is wide at the opening (Fig. 18C). Specimens with a similarly shaped gonostylus are also occasionally observed in other regions (Fig. 18D)., Published as part of Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2022, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), pp. 1-51 in Zoological Journal of the Linnean Society 195 (4) on pages 26-27, DOI: 10.1093/zoolinnean/zlab062, http://zenodo.org/record/5817276, {"references":["Warncke K. 1980. Die Bienengattung Anthidium Fabricius, 1804 in der Westpalaarktis und im turkestanischen Becken. Entomofauna 1: 119 - 210."]}

PSEUDOANTHIDIUM TENELLUM (MOCS��RY, 1880) (FIGS 2A, 10C, 11D, 12D, 13D, 19, 20, 21A, C, E) Anthidium tenellum Mocs��ry, 1880: 48���50, ♀ ♂. Type locality: in Latin ��� Hungaria centrali et meridionali��� [central and southern Hungary], in Hungarian ��� Budapest mellett a Gell��rthegyen, Si��fokn��l ��s Greben��cz k��r��l��� [next to Budapest, on Gell��rthegyen, near Si��fok and around Greben��cz, Hungary]. Lectotype, ♀, designated by B. Tkalců 1984, published here, ��� Greben��c 1878��� [now Serbia], ��� Anthidium tenellum Mocs. det. Mocs��ry���, ���497/202.���, ��� Lectotypus Anthidium tenellum Mocs. (Tkalců, 1984) ���, [red label] (blank), ��� Hungarian Natural History Museum Hymenoptera Coll. Budapest��� [blue label] (HNHM) (Fig. 19). Anthidium eversmanni Radoszkowski, 1886: 54���55, ♂, synon. nov. Type locality: ���Orenbourg��� [Russia: Orenburg Prov., Orenburg]. Lectotype, ♂, by present designation: ���Coll. Radosz.���, ��� floripetum ♂. Evers.���, ��� eversmanni ���, ���Zool. Mus. Berlin���, ��� Lectotypus ♂ Anthidium eversmanni Radoszkowski, 1886 design. Fateryga et Proshchalykin 2020��� [red label] (ZMHB) (Fig. 20). Material examined: 15 females, 60 males (see Supporting Information, Table S1 for specimen data). Distribution: Austria, Azerbaijan, Bulgaria, China (Xinjiang), Crimea, Hungary, Kazakhstan, Kyrgyzstan, Romania, Russia (European part, Urals,Western Siberia), Serbia, Slovakia, Tajikistan, Turkey, Turkmenistan, Ukraine and Uzbekistan (Fig. 6D). The reference in Tkalců (1975) to a distribution for this species in southern Europe and northern Africa, and with a collection locality in Ain Zaatout, Algeria, is probably an error. Host-plant associations: Asteraceae Hungary Centaurea scabiosa subsp. sadleriana (Janka) Asch. & Graebn.(Mocs��ry, 1880) Tajikistan Pulicaria salviifolia Bunge (male visits) (Popov, 1967); Plumbaginaceae Crimea Limonium scoparium (Pall. ex Willd.) Stankov (male visit) (personal observation, A. V. Fateryga). Remarks: Pseudoanthidium tenellum was described in the same publication as P. nanum (Mocs��ry, 1880). For the reasons cited in the Remarks section for P. nanum, we propose the use of 1880 as the official publication date of the name P. tenellum, in other words, P. tenellum (Mocs��ry, 1880). A lectotype is designated for P. eversmanni (Radoszkowski, 1886) (ZMHB). This taxon was determined to be a junior synonym of P. tenellum (Fig. 20). Diagnosis female: The female of P. tenellum may be distinguished from other members of this complex by the following combination of characters: punctation on terga comparatively coarse, as large or larger than punctation on mesonotum; largest punctures on black part of scutellum greater in diameter than the largest punctures on T 2; maculations on European specimens creamy white to pale yellow but lemon yellow in Central Asian specimens (Fig. 21A, C, E). The female of P. tenellum is similar to P. cribratum and P. rozeni; for more information concerning the differentiation of these three species, see the section entitled ���Diagnosis female��� for P. rozeni. In their zone of overlap, differentiating females of P. tenellum from those of P. cribratum and, in some cases, from those of P. stigmaticorne may be challenging. Diagnosis male: The male of P. tenellum may be distinguished from other members of this complex by the following combination of characters: gonostylus over 1.5 times wider at widest point than at base (Fig. 10C); notch at apex of gonostylus less deep than opening of notch is wide (Fig. 10C); exterior and interior margin of gonostylus regularly curved (Fig. 10C); notch at apex of gonostylus more or less centred (Fig. 10C); lateral comb on S5 mitten-shaped (Fig. 11D); posterior, premarginal brush on S3 with hairs unhooked at tips (Fig. 12D); hairless zone on S3 between posterior premarginal brush of hairs and anterior zone of dense, velvety pilosity slightly shiny to matte and trapezoidal, without median extension anteriorly along midline (Fig. 12D); posterior margin of S2 strongly depressed, overhung by long fringe across entire width (Fig. 13D); hairs on ventral surface of trochanter 3 shaggy and uneven, not velvety. T6 and T7 predominantly orange or yellow. The male of P. tenellum is most similar to P. rozeni; for more information concerning the differentiation of these two species, see the section entitled ���Diagnosis male��� for P. rozeni. Geographic variation: Populations in Europe, as well as in Siberia, are characterized by pale yellow to cream-coloured markings on the head, thorax and metasoma. Moreover, the posterior half of T5, as well as the entirety of T6 and T7, are a translucent orange in males. Populations from Central Asia, including Turkmenistan, Tajikistan and Xinjiang, on the other hand, have lemon-yellow coloured markings on the head, thorax and metasoma. In these same populations, the posterior half of T5, as well as T6 and T7, are yellow. The population we examined from Kazakhstan is intermediate, with lemon-yellow coloured markings on the head, thorax and metasoma, similar to those of Central Asian specimens, but with the posterior half of T5, as well as T6 and T7 orange, similar to those of European specimens. The lateral comb on S5 is mitten-shaped and the apical notch of the gonostylus is V-shaped and wider than deep in most populations of P. tenellum., Published as part of Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2021, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), pp. 1-51 in Zoological Journal of the Linnean Society CLXVI (CLXVI) on pages 27-30, DOI: 10.1093/zoolinnean/zlab062, http://zenodo.org/record/5817276, {"references":["Mocsary S. 1880. Ujabb adatok temesmegye hartyaropu faunajahoz. Mathematikai es Termeszettudomanyi Kozlemenyek 16: 1 - 70.","Radoszkowski O. 1886. Fauna hymenopterologique transcaspienne. Horae Societatis Entomologicae Rossicae 20: 3 - 56, pl. I - XI.","Tkalcu B. 1975. Sammelergebnisse der von RNDr. A. Hoffer geleiteten Algerien-Expeditionen in den Jahren 1971 und 1972 (Hymenoptera: Apoidea). 1. Teil: Megachilidae. Acta Rerum Naturalium Musei Nationalis Slovaci Bratislava 21: 165 - 190.","Popov VV. 1967. The bees (Hymenoptera, Apoidea) of Middle Asia and their associations with angiosperm plants. Trudy Zoologiceskzo Instituta Akademija Nauk SSSR (Leningrad [St. Petersburg]) 38: 11 - 329 [in Russian]."]}

Pseudoanthidium gregoriense Pseudoanthidium gregoriense was described by Nobile (1990) as an island endemic from Sicily. Originally described as a subspecies of P. alpinum, Nobile later raised P. gregoriense to species status (1995). As mentioned above for Stelis leucostoma, Nobile (1995) draws parallels between P. gregoriense, P. leucostoma and P. lituratum. We examined four specimens of this taxon identified by Nobile (including one of his paratypes) and a photograph of the holotype, plus two other female specimens from Sicily. These specimens agree with our concept of P. stigmaticorne., Published as part of Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2022, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), pp. 1287-1337 in Zoological Journal of the Linnean Society 195 on page 1312, DOI: 10.1093/zoolinnean/zlab062, http://zenodo.org/record/5817276, {"references":["Nobile V. 1990. Il genere Anthidium Fabricius 1804 e affini (Hymenoptera, Apoidea, Megachilidae) in Sicilia, con descrizione di Pseudoanthidium alpinum gregoriense subsp. n. Animalia 16: 131 - 145.","Nobile V. 1995. Api (Insecta, Hymenoptera) nuove o poco note di Sicilia e di Sardegna. Bollettino delle Sedute della Accademia Gioenia di Scienze Naturali in Catania 28: 147 - 159."]}

PSEUDOANTHIDIUM CRIBRATUM (MORAWITZ, 1875) (FIGS 2B, 11F, 12F, 13F, 17B, D, F, 23B, 24) Anthidium cribratum Morawitz, 1875: 130–131, ♀ ♂. Type locality: in Latin ‘Hab. in deserto prope Taschkent et in Kuldscha’ [in the desert near Tashkent and in Kuldscha], in Russian ‘Найдень въ степи между Ташкентомъ и Сыръ- Дарьей 19 мая (700‘-1400‘). Полученъ также изъ Кульджи’ [found in the steppe between Tashkent and the Syr-Darya (Uzbekistan, Kazakhstan) on May 19. Received also from Gulja, probably in China]. Lectotype, ♂, designated byWarncke, 1980: 162:‘Ташкентъ [Tashkent]’, ‘к. Ф Моравица [coll. F. Morawitz]’, ‘ Anthidium cribratum F. Moraw. ♂.’, ‘ Lectotypus Anthidium cribratum Mor. (Warncke 1978) ’ (ZISP). ? Anthidium petechiale Morawitz, 1875: 130, ♀. Type locality: in Latin ‘Hab. in valle Sarafschan; semel captum’ [Captured once in the Zeravshan River Valley, Tajikistan], in Russian ‘Найденъ только разъ въ заравшанской долинѣ между Іори и Дашты- Казы 31 мая (3‘800)’ [Found only once in the Zarafschan River Valley between Yori and Dasthikazy, Tajikistan, May 31, 3‘ 800 m]. Material examined: Seven females, 29 males (see Supporting Information, Table S1 for specimen data). Distribution: Iran, Israel and Palestine, Jordan, Kazakhstan, Kyrgyzstan, Syria, Tajikistan, Turkey, Turkmenistan and Uzbekistan (Fig. 22B). Host-plant associations (all records from Popov, 1967): Asteraceae Tajikistan Centaurea iberica Trevir. ex Spreng. (male visit), Chondrilla juncea L. (female visit), Cirsium turkestanicum (Regel) Petr. (female visit), Cynara scolymus L. (female visit), Erigeron canadensis L. (female and male visits), Inula sp. (male visits), Onopordum acanthium L. (male and female visits), Pulicaria salviifolia Bunge (male and female visits), Rhaponticum repens (L.) Hidalgo (male and female visits), Tripleurospermum disciforme (C.A. Mey.) Sch. Bip. (male visit); Boraginaceae Uzbekistan Echium italicum subsp. biebersteinii (Lacaita) Greuter & Burdet (male visits); Chenopodiaceae Tajikistan Climacoptera transoxana (Iljin) Botsch. (male visit); Dipsacaceae Tajikistan Dipsacus laciniatus L. (female visit); Fabaceae Tajikistan Alhagi kirghisorum Schrenk (female visits), Trifolium repens L. (male visit); Lamiaceae Tajikistan Mentha longifolia (L.) L. (male and female visits), Vitex angus-castus L. (female visit); Onagraceae Tajikistan Epilobium hirsutum L. (male visits); Plumbaginaceae Tajikistan Limonium perfoliatum (Kar. ex Boiss.) Kuntze (male visit); Ranunculaceae Tajikistan Clematis orientalis L. (female and male visits); Tamaricaceae Tajikistan Tamarix sp. (male visits); Verbenaceae Tajikistan Verbena officinalis L. (male and female visits). Diagnosis female: The female of P. cribratum may be distinguished from other members of this complex by the following combination of characters: punctation on terga comparatively coarse, as large or larger than punctation on mesonotum, with shiny interspaces between punctures; largest punctures on black part of scutellum equal in diameter to largest punctures on T 2 (Fig. 17B, D, F). The female of P. cribratum is similar to P. tenellum and P. rozeni; for more information concerning the differentiation of these three species, see the section entitled ‘Diagnosis female’ for P. rozeni. In their zone of overlap (e.g. in Central Asia), differentiating females of P. cribratum from those of P. tenellum may be challenging. Diagnosis male: The male of P. cribratum may be distinguished from other members of this complex by the following combination of characters: apex of coxa 3 with pronounced, flattened, round-tipped tooth, about as long as third tarsal segment is wide at apex (Fig. 24A–B), which is unique within the species complex; gonostylus over 1.5 times wider at widest point than at base (Fig. 23B); notch at apex of gonostylus wide and deeply U-shaped (Fig. 23B); notch is slightly less deep than width of notch at opening; notch slopes laterally, so that interior tip of the notch is visibly wider than the exterior (Fig. 23B); lateral comb on S5 very small, with longest teeth far shorter than maximum width of hind basitarsus (Fig. 11F); posterior, premarginal brush on S3 with hairs unhooked at tips (Fig. 12F); shiny, hairless zone on S3 between posterior premarginal brush of hairs and anterior zone of dense, velvety pilosity trapezoidal, without medial extension extending anteriorly along the midline of sternum (Fig. 12F); posterior margin of S2 strongly depressed (Fig. 13F); hairs on ventral surface of trochanter 3 dense and of even length but not velvety. Geographic variation: In Central Asia, individuals have broad punctures that are dense but not contiguous on the vertex, thorax and terga and the integument is shiny. In specimens from the Middle East, including Israel, Jordan, Syria and eastern Turkey, punctures are smaller and mostly contiguous and the integument is less shiny (lightly reticulate), especially on the scutum and scutellum. In addition, males from Central Asia have a more elongate gonostylus with the outer apical projection cylindrical and the lateral comb of S5 nearly truncate and symmetrical apically, while Middle Eastern males have a shorter gonostylus with the outer projection conical and the lateral comb asymmetrical, not truncate. Specimens from Iran are more similar to specimens from the Middle East than to those from Central Asia, although in a single specimen the punctation on the terga is intermediate in size between the forms seen at the extreme ends of this species’ distribution. PSEUDOANTHIDIUM CANARIENSE (MAVROMOUSTAKIS, 1954) (FIGS 2C, 23C, 25A, C, E) Anthidium canariense Mavromoustakis, 1954: 712– 715, ♀ ♂. Type locality:‘Canary Islands’ (without specific locality) [holotype ♀, allotype ♂, paratype ♀ (NMW); paratypes 2♀ (listed in publication as deposited in the Mavromoustakis personal collection), likely deposited at the MCN]. Material examined: Four females, seven males (see Supporting Information, Table S1 for specimen data). Distribution: Spain (Canary Islands): Santa Cruz de Tenerife, Gran Canaria, La Gomera (Fig. 22C). Host-plant associations: Asteraceae Gran Canaria Argyranthemum cf. frutescens (L.) Sch. Bip. (male visits), Asteriscus graveolens subsp. stenophyllus (Link) Greuter (male and female visits), Carduus tenuiflorus Curtis (female visits); Tenerife and Grand Canaria Carduus sp. (female visits); Tenerife, Grand Canaria, La Gomera Galactites tomentosa Moench (male and female visits); Tenerife Scolymus hispanicus L. (male and female visits); Brassicaceae Gran Canaria Erysimum scoparium (Brouss. ex Willd.) Wettst. (male visits), Hirschfeldia incana (L.) Lagr.-Foss. (female visits); Lamiaceae Tenerife Cedronella canariensis (L.) Webb & Berthel. (male visits) (Hohmann et al., 1993). Diagnosis female: The female of P. canariense may be distinguished from other members of this complex by the following combination of characters: punctation on terga comparatively coarse, as large or larger than punctation on mesonotum, with shiny interspaces between punctures; largest punctures on black part of scutellum approximately equal in diameter to those of largest punctures on T 2; maculations on head, mesosoma and metasoma dark orange; shiny spaces between punctures on T 3 narrow, less than one-quarter of a puncture wide; hairs on inside of third basitarsus dark brown and with individual hairs mostly thicker than hairs on outside surface. Colour of maculations orange-yellow (Fig. 25A). Diagnosis male: The male of P. canariense may be distinguished from other members of this complex by the following combination of characters: gonostylus approximately parallel-sided and unnotched at apex (Fig. 23C); lateral comb on S5 small, with longest teeth shorter than maximal width of hind basitarsus; posterior, premarginal brush on S3 with hairs hooked at tips; shiny, hairless zone on S3 between posterior premarginal brush of hairs and anterior zone of dense, velvety pilosity very short, about one-third of the width of the sternum, dark, chevron-shaped, without medial extension extending anteriorly along the midline of sternum; posterior margin of S2 medially emarginate, S2 otherwise covered in silvery pilosity except for a more or less hairless posterior margin. Colour of maculations orange-yellow (Fig. 25C). Posterior margin of T 7 with deep, nearly semi-circular emargination medially (Fig. 25E). PSEUDOANTHIDIUM TROPICUM (WARNCKE, 1982), STAT. NOV. (FIGS 23D, 25B, D, F) Anthidium lituratum tropicum Warncke, 1982: 172–173, ♀ ♂. Type locality: ‘ Pass E Rudan /V Minab, Bandar Abbas in 570 m, 23-V-1978, an Centaurea spec.’ (holotype ♀) (OLML). Paratypes: ‘ Bandar Abbas: Pass E Rudan /N Minab in 570 m, 23-V-1978, an Centaurea spec’ (7♂, 3♀); ‘ Fars: Daria Namak / 27 km E Shiraz, 7-VII-1965, S’ (1♀) (OLML); ‘ Fars: Persepolis in 1570 m, 16-V-1978, W’ (1♂) (OLML). Material examined: Eight females, four males (see Supporting Information, Table S1 for material examined). Distribution: Iran (Fig. 22D). Host-plant associations: Asteraceae Bandar Abbas Centaurea sp. (Warncke 1982). Remarks: This taxon was originally described as a subspecies of P. nanum (as P. lituratum tropicum). However, an examination of the type series indicates that this taxon is the most morphologically distinct of all the members of the P. scapulare complex, namely in its small size, its light coloration and the rounded apex of the male gonostylus, the last trait shared only with P. canariense in this species complex. Although we were unable to obtain genetic data for this taxon, we consider it morphologically divergent enough to elevate it to species status. Diagnosis female: The female of P. tropicum may be distinguished from other members of this complex by the following combination of characters: punctation on terga comparatively fine, less wide than diameter of punctation on mesonotum; fourth antennal segment less than half as long as fifth (proportionally longer in all other taxa); metasoma brown with pale yellow maculations; comparatively small (~ 5 mm) (Fig. 25B). Diagnosis male: The male of P. tropicum may be distinguished from other members of this complex by the following combination of characters: very small size (≤ 5 mm), gonostylus approximately parallel-sided with apex almost square and unnotched (Fig. 23D); lateral comb on S5 small, with longest teeth shorter than maximal width of hind basitarsus and no wider than basal comb; posterior, premarginal brush on S3 with hairs hooked at tips; posterior margin of S2 gently emarginate medially, S2 otherwise covered in dense, velvety pilosity except for a more or less hairless posterior margin., Published as part of Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2022, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), pp. 1287-1337 in Zoological Journal of the Linnean Society 195 on pages 1317-1321, DOI: 10.1093/zoolinnean/zlab062, http://zenodo.org/record/5817276, {"references":["Morawitz F. 1875. Puteshestviye v Turkestan chlenaosnovatelya obshchestva A. P. Fedtschenko, sovershennoye ot Imperatorskogo obshchestva lyubiteley yestestvoznaniya po porucheniyu turkestanskogo general-gubernatora K. P. fon-Kaufmana. Vypusk 9. Tom 2. Zoogeograficheskiye issledovaniya. Chast 5. Otdel sedmoy. Pchely (Mellifera). Moscow: M. Stasyulevich [in Russian and Latin].","Popov VV. 1967. The bees (Hymenoptera, Apoidea) of Middle Asia and their associations with angiosperm plants. Trudy Zoologiceskzo Instituta Akademija Nauk SSSR (Leningrad [St. Petersburg]) 38: 11 - 329 [in Russian].","Mavromoustakis GA. 1954. LXXXIX. - New and little-known bees of the subfamily Anthidiinae (Apoid [ae]). Part IX. Annals & Magazine of Natural History 12: 711 - 715.","Hohmann H, La Roche F, Ortega G, Barquin J. 1993. Bienen, Vespen und Ameisen des Kanarischen Inseln. Band 1. Bremen: Veroffentlichungen aus dem Ubersee-Museum.","Warncke K. 1982. Beitrag zur Bienenfauna des Iran 15. Die Gattung Anthidium F. Bollettino del Museo Civico di Storia Naturale di Venezia 32: 171 - 196."]}

PSEUDOANTHIDIUM STIGMATICORNE (DOURS, 1873) (FIGS 1B, 2D, 11C, 12C, 13C, 15, 16, 17A, C, E, 18) Anthidium stigmaticorne Dours, 1873: 305–306, ♀. Type locality: ‘Algérie‘ [Algeria]. Neotype, ♂, by present designation: ‘Oran’, ‘MNHN, Paris, EY24647’, ‘ ♂ ’, ‘Museum Paris Collection Ernest André 1914’, ‘ Neotype Anthidium stigmaticorne Dours, 1873 R. Le Divelec des.’ [red label] (MNHN) (Fig. 15). ? Stelis leucostoma Costa, 1883: 96, ♂. Type locality: ‘ Sardegna’ [Sardinia]. ? Anthidium peregrinum Costa, 1885: 21–22, ♀ ♂. Type locality: ‘ Sardegna’ [Sardinia]. ? Anthidium fraternum Pérez, 1895: 22, ♂. Type locality not given. ? Anthidium astilleroi Dusmet y Alsonso, 1915: 301–302, ♀. Type localities: 6♀ ‘ Mogador’ [currently Essaouira, Morocco], 2♀ ‘ Marraquesh’ [Marrakesh, Morocco], 3♀ ‘ Tigui’ [possibly Morocco]. Paranthidiellum karakalense Popov, 1952: 98–101, ♀ ♂, synon. nov. Type locality: ‘Кара- кала [Karakala]’ [currently Magtymguly in Balkan Prov. of Turkmenistan]. Lectotype, ♂, by present designation: ‘Кара- кала, Копетдаг, Туркм, В. Попов [Kara-kala, Kopetdag, Turkm, V. Popov] 7.VIII.934’, ‘ Anthidium karakalense, sp. n. ♂. monotyp. Popov. 1934. det’, ‘ Lectotypus ♂ Paranthidiellum karakalense Popov, 1952 design. Fateryga et Proshchalykin 2020’ [red label] (ZISP) (Fig. 16). Paralectotypes: ♀, ‘Кара- кала, Копетдаг, Туркм, В. Попов [Kara-kala, Kopetdag, Turkm, V. Popov] 9.VIII.934’, ‘ Anthidium karakalense, sp. n. ♀ holotyp. Popov. 1934 det’, ‘ Paralectotypus ♀ Paranthidiellum karakalense Popov, 1952 design. Fateryga et Proshchalykin 2020’ [red label]; ♀, ‘Кара- кала, Копетдаг, Туркм, В. Попов [Kara-kala, Kopetdag, Turkm, V. Popov] 8.VIII.934’, ‘ Anthidium karakalense, sp. n. ♀ paratyp. Popov. 1934 det’, ‘ Paralectotypus ♀ Paranthidiellum karakalense Popov, 1952 design. Fateryga et Proshchalykin 2020’ [red label]; ♀, ‘Кара- кала, Копетдаг, Туркм, В. Попов [Kara-kala, Kopetdag, Turkm, V. Popov] 6.VIII.934’, ‘ Anthidium karakalense, sp. n. ♀ holotyp. Popov. 1934. det’, ‘ Paralectotypus ♀ Paranthidiellum karakalense Popov, 1952 design. Fateryga et Proshchalykin 2020’ [red label] (ZISP). ? Pseudoanthidium alpinum gregoriense Nobile, 1990: 138–140, ♂. Type locality: ‘ Catania, S. Gregorio’ [Sicily], 3.IX.1989 (holotype ♂, paratypes 8 ♂). Material examined: 143 females, 155 males (see Supporting Information, Table S1 for specimen data). Distribution: Algeria, Azerbaijan, Bulgaria, Crimea, Croatia, Cyprus, France (including Corsica), Greece, Iran, Israel and Palestine, Italy (including Sardinia and Sicily), Jordan, Morocco, Portugal, Romania, Russia (European part), Spain, Syria, Tunisia, Turkey and Turkmenistan (Fig. 6C). Host-plant associations: Asteraceae Algeria Silybum marianum (L.) Gaertn., Centaurea algeriensis Durieu & Coss. (Aguib et al., 2010) Crimea Carduus hamulosus Ehrh. (male and female visits), Grindelia squarrosa (Pursh) Dunal (female visit) (personal observation, A. V. Fateryga); Fabaceae Crimea Trigonella procumbens (Besser) Rchb. (male visit) (personal observation, A. V. Fateryga); Lamiaceae Algeria Marrubium vulgare L. (Aguib et al., 2010), Teucrium polium L. (Saunders, 1908) Crimea Teucrium chamaedrys L. (male visit), Thymus tauricus Klokov & Des. -Shost. (male visits) (personal observation, A. V. Fateryga); Plantaginaceae Crimea Linaria genistifolia (L.) Mill. (female visit) (personal observation, A. V. Fateryga); Plumbaginaceae Crimea Limonium sp. (female visit) (information taken from specimen label); Zygophyllaceae Dagestan Zygophyllum fabago L. (male visit) (personal observation, A. V. Fateryga). Remarks: Dours originally described P. stigmaticorne based on an unspecified number of female specimens collected in Algeria, referring to the species as intermediate between P. scapulare (as Anthidium scapulare) and Icteranthidium grohmanni (Spinola, 1838) (as Anthidium rubiginosum). Saunders (1908) used the same name to refer to female specimens collected in Biskra (Algeria) with the ‘4 th and 5 th joints of the antennae testaceous’. Warncke (1980) also referred to P. stigmaticorne but considered it a subspecies of P. lituratum; he mentioned the dark red colour of this taxon in northern Africa and questioned whether the rounded apex of the gonostylus in males from this region was also present elsewhere (‘Ob beim ♂ überall die Gonostylenenden gerundet sind, muss erst noch herausgefunden werden’). Aguib et al. (2010) applied the name P. stigmaticorne to a member of the P. scapulare complex ‘endemic’ to northern Africa in which males exhibit an apically widened gonostylus bearing a rounded notch and females have evenly spaced punctures on T 1– T 2, separated by shiny interspaces (see below for complete diagnosis). The description of the antennal segments given by Saunders (1908), as well as of the rounded gonostylus mentioned by Warncke (1980), both characters typical of P. stigmaticorne in northern Africa and elsewhere, suggest that both authors may have been referring to the same taxon clearly described and illustrated in Aguib et al. (2010). Yet none of these works make any mention of having seen Dours’ original type material. Most of the Dours collection is said to have been destroyed in a fire in the United States (Horn & Kahle, 1935). While a small amount of material from Dours is present in the collections at the MNHN, no specimens bearing any evidence of belonging to the type series of P. stigmaticorne were located, despite extensive searches of the collections. At this point we consider this material lost. We draw attention to the fact that some of the criteria mentioned by Dours in his original description of P. stigmaticorne do not correspond with the specimens that we have examined from northern Africa. For example, Dours describes the abdominal scopa of P. stigmaticorne as red (‘palette ventrale rousse’) but in the specimens that we have examined from northern Africa (Algeria, Morocco and Tunisia), the scopa is nearly white. In the absence of the type series, it is impossible to know whether Dours’ description was made in reference to another taxon (although no taxon fitting this description is thus far known to us) or whether his description was partially erroneous. The name P. stigmaticorne has been applied by other authors (Warncke, 1980; Aguib et al., 2010) in a clear and descriptive manner, leaving no doubt to which taxon this name has been applied. We thus find it appropriate to designate a neotype in order to clarify the taxonomic status of P. stigmaticorne and to do so in keeping with the nomenclature in common usage. The characters that may be used to differentiate this taxon from other, closely related taxa are detailed in the species diagnoses found below; morphological variation is further discussed in the section below entitled ‘Geographic variation’. The neotype was collected in Algeria, in keeping with the original type locality (only given as ‘Algeria’ in Dours, 1873). Although the original description was based on female specimens, we have chosen a male specimen as a neotype (Fig. 15). The identification of males is unambiguous and the selection of a male neotype promotes a greater degree of taxonomic stability than would a female. The neotype is deposited in the MNHN. Both Warncke (1980) and Aguib et al. (2010) describe P. stigmaticorne as restricted to northern Africa, yet morphologically similar specimens exist throughout the western Palaearctic. Both the DNA barcode and the UCE results indicate that the ensemble of these individuals forms a monophyletic clade exhibiting little genetic differentiation and at the present time we refer to them collectively as P. stigmaticorne. Our results also highlight the presence of P. stigmaticorne throughout southern mainland Europe, thus revealing a previously overlooked taxon in this region. Numerous taxa corresponding to regional variants of P. stigmaticorne have been described from different regions of the Palaearctic, including several island endemics. We place a certain number of these taxa in synonymy. Some were placed in synonymy by other authors (Warncke, 1980; Nobile, 1995; Rasmont et al., 1995; Ornosa et al., 2008); one synonym is published for the first time here. All are discussed below., Published as part of Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2022, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), pp. 1287-1337 in Zoological Journal of the Linnean Society 195 on pages 1306-1310, DOI: 10.1093/zoolinnean/zlab062, http://zenodo.org/record/5817276, {"references":["Dours JA. 1873. Hymenopteres du bassin mediterraneen. Andrena (suite). Revue et Magasin de Zoologie Pure et Applique 3: 274 - 325.","Costa A. 1883. Notizie ed osservazioni sulla geo-fauna Sarda, Memoria seconda, Risultamento di ricerche fatte in Sardegna nella primavera del 1882. Atti della Reale Accademia delle Scienze Fisiche e Matematiche di Napoli 2: 2, 1 - 109.","Costa A. 1885. Notizie ed osservazioni sulla geo-fauna Sarda, Memoria quarta. Atti della Reale Accademia delle Scienze Fisiche e Matematiche di Napoli 2: 13, 1 - 31.","Perez J. 1895. Especes nouvelles de melliferes de Barbarie (diagnoses preliminaires). Bordeaux: Gounouihou.","Popov VV. 1952. Bee fauna (Hymenoptera, Apoidea) of southwestern Turkmenistan and its landscape distribution. Trudy Zoologicheskogo Instituta Akademii Nauk SSSR (Leningrad [St. Petersburg]) 10: 61 - 117 [in Russian].","Nobile V. 1990. Il genere Anthidium Fabricius 1804 e affini (Hymenoptera, Apoidea, Megachilidae) in Sicilia, con descrizione di Pseudoanthidium alpinum gregoriense subsp. n. Animalia 16: 131 - 145.","Aguib S, Louadi K, Schwarz M. 2010. Les Anthidiini (Megachilidae, Megachilinae) d'Algerie avec trois especes nouvelles pour ce pays: Anthidium (Anthidium) florentinum (Fabricius, 1775), Anthidium (Proanthidium) amabile Alfken, 1932 et Pseudoanthidium (Exanthidium) enslini (Alfken, 1928). Entomofauna 31: 121 - 152.","Saunders E. 1908. Hymenoptera aculeata collected in Algeria by the Rev. A. E. Eaton M. A., F. E. S. and the Rev. Francis David Morice M. A., F. E. S. Part III. Anthophila. Transactions of the Entomological Society of London 1908: 177 - 274.","Warncke K. 1980. Die Bienengattung Anthidium Fabricius, 1804 in der Westpalaarktis und im turkestanischen Becken. Entomofauna 1: 119 - 210.","Horn W, Kahle I. 1935. I. Kapitel: Sammlungen, welche ihren Eigentumer gewechselt haben. Entomologische Beihefte Berlin-Dahlem 2: 1 - 162 pl. I - XV.","Nobile V. 1995. Api (Insecta, Hymenoptera) nuove o poco note di Sicilia e di Sardegna. Bollettino delle Sedute della Accademia Gioenia di Scienze Naturali in Catania 28: 147 - 159.","Rasmont P, Ebmer A, Banaszak J, van der Zanden G. 1995. Hymenoptera Apoidea Gallica. Bulletin de la Societe Entomologique de France 100: 1 - 98.","Ornosa C, Ortiz-Sanchez FJ, Torres F. 2008. Catalogo de los Megachilidae del Mediterraneo occidental (Hymenoptera, Apoidea). III. Anthidiini y Dioxyini. Graellsia 64: 61 - 86."]}

Anthidium astilleroi The type series of A. astilleroi includes 11 females described from Morocco. No males were described. The authors describe A. astilleroi as similar to both ‘ lituratum ’ (= P.nanum) and ‘ ferrugineum ’ [= Icteranthidium ferrugineum (Fabricius, 1787)] based on the colour of the metasoma yet different from ‘ ferrugineum ’ in the shape of the scutellum. They conclude that A. astilleroi is more similar to A. lituratum but also mention that there are no two identical specimens in the series but rather a series of ‘modifications’. The type material should thus be carefully examined to determine to which taxon the name should be applied, as well as to verify that the type series does not include multiple taxa. We include this taxon here in the discussion because it was synonymized with P. stigmaticorne by Warncke (1980) but we have not examined the type series., Published as part of Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2022, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), pp. 1287-1337 in Zoological Journal of the Linnean Society 195 on page 1311, DOI: 10.1093/zoolinnean/zlab062, http://zenodo.org/record/5817276, {"references":["Warncke K. 1980. Die Bienengattung Anthidium Fabricius, 1804 in der Westpalaarktis und im turkestanischen Becken. Entomofauna 1: 119 - 210."]}