taxonID	type	description	language	source
AE06D043FFF1FF93FF679477FD8CF93F.taxon	description	(FIGS 2 F, 4, 5, 9 A, C, E, 10 A, 11 A, 12 A, 13 A)	en	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
AE06D043FFF1FF93FF679477FD8CF93F.taxon	materials_examined	Material examined: 195 females, 117 males (see Supporting Information, Table S 1 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. 6 A). 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 59 b (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 S 1). 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).	en	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
AE06D043FFF1FF93FF679477FD8CF93F.taxon	diagnosis	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 T 1 – T 2, narrow and dense with shiny interspaces between puncture; posterior margin of T 1 – T 4 shiny (Fig. 9 A, 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. 10 A); 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. 10 A); lateral comb on S 5 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. 11 A); comb of S 5 wider than arm preceding it; posterior, premarginal brush on S 3 with hairs hooked at tips (hairs unhooked in P. tenellum and P. cribratum) (Fig. 12 A); shiny, hairless zone on S 3 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. 12 A) (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 S 2 shiny, dark and in shape of chevron (Fig. 13 A); 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 T 1 – T 4, T 1 – T 5 or T 1 – T 6. 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 T 1 – T 4 or T 1 – T 5.	en	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
AE06D043FFF7FF8FFF0591B7FE48F902.taxon	description	(FIGS 2 E, 9 B, D, F, 10 B, 11 B, 12 B, 13 B, 14)	en	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
AE06D043FFF7FF8FFF0591B7FE48F902.taxon	materials_examined	Material examined: 89 females, 83 males (see Supporting Information, Table S 1 for specimen data). Distribution: Algeria, France, Morocco, Portugal, Spain (including Mallorca) and Tunisia (Fig. 6 B). 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 S 2 – S 4. 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. 9 B, 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. 10 B); 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. 10 B); 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. 11 B); posterior margin of S 4 deeply emarginate at centre (not emarginate in other species); posterior, premarginal brush on S 3 with hairs hooked at tips (hairs unhooked in P. tenellum and P. cribratum) (Fig. 12 B); shiny, hairless zone on S 3 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. 12 B); raised zone on S 2 shiny, dark and in shape of chevron; posterior margin of S 2 usually gently emarginate medially (not so in other species) (Fig. 13 B); hairs on ventral surface of trochanter 3 very short, velvety and dense, tuft of longer hairs present at base of trochanter.	en	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
AE06D043FFEBFF8BFF0191B2FAE4FA29.taxon	description	(FIGS 1 B, 2 D, 11 C, 12 C, 13 C, 15, 16, 17 A, C, E, 18)	en	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
AE06D043FFEBFF8BFF0191B2FAE4FA29.taxon	materials_examined	Material examined: 143 females, 155 males (see Supporting Information, Table S 1 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. 6 C). 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.	en	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
AE06D043FFE0FF84FF229193FAA6F8F7.taxon	materials_examined	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.	en	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
AE06D043FFE1FF86FF109503FB90FB25.taxon	description	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).	en	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
AE06D043FFE1FF86FF109503FB90FB25.taxon	diagnosis	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 T 2; 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. 17 A, 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. 18 A – F); exterior margin of gonostylus relatively straight, inner margin more sharply rounded toward apex (Fig. 18 A – 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. 18 A – F); lateral comb on S 5 small, with longest teeth shorter than maximal width of hind basitarsus (comb larger in P. scapulare, P. nanum, P. tenellum and P. palestinicum) (Fig. 11 C); comb of S 5 no wider than arm preceding it (Fig. 11 C); posterior, premarginal brush on S 3 with hairs hooked at tips (hairs unhooked in P. tenellum and P. cribratum) and arranged in an open brushy arc (Fig. 12 C); no shiny, hairless zone on S 3 between posterior premarginal brush of hairs and anterior zone of dense, velvety pilosity (present in other species) (Fig. 12 C); darkened raised zone across S 2 with only short, sparse hairs, in shape of ‘ moustache’ (Fig. 13 C); hairs on ventral surface of trochanter 3 shaggy and uneven, not dense, with longer hairs toward base.	en	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
AE06D043FFE1FF86FF109503FB90FB25.taxon	description	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 T 1 – T 5, although those on T 5 are sometimes much reduced. Legs are dark orange yellow and maculations on metasoma sub-rectangular with rounded corners. Maculations on T 3 – T 5 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 T 1 – T 5 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 T 1 – T 4, T 5 – T 6 are entirely black and T 7 is pale yellow. In the Middle East, paired maculations are present on T 1 – T 6 and T 7 is yellow or, alternatively, paired maculations are present on T 1 – T 5, T 6 is black and T 7 is pale yellow. We have also examined specimens from Dagestan in which lemon-yellow maculations are present on T 1 – T 5, T 6 is black and T 7 is pale yellow. Specimens from northern Africa are similar to those in Dagestan but the posterior margin of T 6 is yellow. In individuals from France and Spain, paired maculations are present on T 1 – T 6 and T 7 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. 18 B) 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. 18 C). Specimens with a similarly shaped gonostylus are also occasionally observed in other regions (Fig. 18 D).	en	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
AE06D043FFE2FF82FCF493D0FE9EFF51.taxon	description	(FIGS 2 A, 10 C, 11 D, 12 D, 13 D, 19, 20, 21 A, C, E)	en	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
AE06D043FFE2FF82FCF493D0FE9EFF51.taxon	description	Fateryga et Proshchalykin 2020 ’ [red label] (ZMHB) (Fig. 20). Material examined: 15 females, 60 males (see Supporting Information, Table S 1 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. 6 D). 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. (Mocsáry, 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 (Mocsáry, 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 (Mocsáry, 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. 21 A, 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. 10 C); notch at apex of gonostylus less deep than opening of notch is wide (Fig. 10 C); exterior and interior margin of gonostylus regularly curved (Fig. 10 C); notch at apex of gonostylus more or less centred (Fig. 10 C); lateral comb on S 5 mitten-shaped (Fig. 11 D); posterior, premarginal brush on S 3 with hairs unhooked at tips (Fig. 12 D); hairless zone on S 3 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. 12 D); posterior margin of S 2 strongly depressed, overhung by long fringe across entire width (Fig. 13 D); hairs on ventral surface of trochanter 3 shaggy and uneven, not velvety. T 6 and T 7 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 T 5, as well as the entirety of T 6 and T 7, 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 T 5, as well as T 6 and T 7, 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 T 5, as well as T 6 and T 7 orange, similar to those of European specimens. The lateral comb on S 5 is mitten-shaped and the apical notch of the gonostylus is V-shaped and wider than deep in most populations of P. tenellum.	en	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
AE06D043FFE6FF82FF4C965AFE78FD13.taxon	description	(FIGS 2 G, 11 E, 12 E, 13 E, 21 B, D, F, 23 A)	en	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
AE06D043FFE6FF82FF4C965AFE78FD13.taxon	materials_examined	Material examined: Seven females, 12 males (see Supporting Information, Table S 1 for specimen data). Distribution: Israel and Palestine, Jordan, Lebanon, Syria and Turkey (Fig. 22 A).	en	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
AE06D043FFE6FFBCFC8E94E5FAB2FC51.taxon	description	(FIGS 2 B, 11 F, 12 F, 13 F, 17 B, D, F, 23 B, 24)	en	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
AE06D043FFE6FFBCFC8E94E5FAB2FC51.taxon	materials_examined	Material examined: Seven females, 29 males (see Supporting Information, Table S 1 for specimen data). Distribution: Iran, Israel and Palestine, Jordan, Kazakhstan, Kyrgyzstan, Syria, Tajikistan, Turkey, Turkmenistan and Uzbekistan (Fig. 22 B). 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. 17 B, 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. 24 A – B), which is unique within the species complex; gonostylus over 1.5 times wider at widest point than at base (Fig. 23 B); notch at apex of gonostylus wide and deeply U-shaped (Fig. 23 B); 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. 23 B); lateral comb on S 5 very small, with longest teeth far shorter than maximum width of hind basitarsus (Fig. 11 F); posterior, premarginal brush on S 3 with hairs unhooked at tips (Fig. 12 F); shiny, hairless zone on S 3 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. 12 F); posterior margin of S 2 strongly depressed (Fig. 13 F); 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 S 5 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.	en	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
AE06D043FFD8FFBEFC9C934AFDAEFDA5.taxon	description	(FIGS 2 C, 23 C, 25 A, C, E)	en	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
AE06D043FFD8FFBEFC9C934AFDAEFDA5.taxon	materials_examined	Material examined: Four females, seven males (see Supporting Information, Table S 1 for specimen data).	en	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
AE06D043FFD8FFBEFC9C934AFDAEFDA5.taxon	distribution	Distribution: Spain (Canary Islands): Santa Cruz de Tenerife, Gran Canaria, La Gomera (Fig. 22 C).	en	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
AE06D043FFD8FFBEFC9C934AFDAEFDA5.taxon	biology_ecology	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).	en	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
AE06D043FFD8FFBEFC9C934AFDAEFDA5.taxon	diagnosis	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. 25 A). 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. 23 C); lateral comb on S 5 small, with longest teeth shorter than maximal width of hind basitarsus; posterior, premarginal brush on S 3 with hairs hooked at tips; shiny, hairless zone on S 3 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 S 2 medially emarginate, S 2 otherwise covered in silvery pilosity except for a more or less hairless posterior margin. Colour of maculations orange-yellow (Fig. 25 C). Posterior margin of T 7 with deep, nearly semi-circular emargination medially (Fig. 25 E).	en	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
AE06D043FFDAFFBEFF1F952DFAFCFB90.taxon	description	(FIGS 23 D, 25 B, D, F)	en	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
AE06D043FFDAFFBEFF1F952DFAFCFB90.taxon	materials_examined	Material examined: Eight females, four males (see Supporting Information, Table S 1 for material examined).	en	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
AE06D043FFDAFFBEFF1F952DFAFCFB90.taxon	distribution	Distribution: Iran (Fig. 22 D).	en	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
AE06D043FFDAFFBEFF1F952DFAFCFB90.taxon	biology_ecology	Host-plant associations: A steraceae Bandar Abbas Centaurea sp. (Warncke 1982).	en	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
AE06D043FFDAFFBEFF1F952DFAFCFB90.taxon	discussion	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.	en	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
AE06D043FFDAFFBEFF1F952DFAFCFB90.taxon	diagnosis	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. 25 B). 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. 23 D); lateral comb on S 5 small, with longest teeth shorter than maximal width of hind basitarsus and no wider than basal comb; posterior, premarginal brush on S 3 with hairs hooked at tips; posterior margin of S 2 gently emarginate medially, S 2 otherwise covered in dense, velvety pilosity except for a more or less hairless posterior margin.	en	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
