Pseudoanthidium nanum, (MOCSARY, 1880)

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), pp. 1-51 : 10-15

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.

Figures

Figure 2. Best-scoring maximum likelihood tree based on analysis of COI data. Numbers shown at nodes are maximum likelihood bootstrap values based on 1000 bootstrap replicates in RAxML. Only bootstrap values greater than 50% are shown. Terminals are labelled with a DNA extraction code, species name, collection locality and either as male (m) or female (f). Barcodes were obtained using Sanger sequencing technology, except for those corresponding to specimens 1802 and 1805, which were obtained from non-UCE assemblies generated during UCE sequencing. A, Pseudoanthidium tenellum, Burgenland, Austria (m), photo Bernhard Jacobi; B, P. cribratum, Bukhara, Uzbekistan (f), photo Jessica Litman; C, P. canariense, Santa Cruz de Tenerife, Canary Islands, Spain (m), photo Jessica Litman; D, P. stigmaticorne, Crimea, Russia, photo Alexander V. Fateryga; E, P. scapulare, Portugal, photo Ian Cross; F, P. nanum, photo Entomologie/Botanik, ETH Zürich / Albert Krebs; G, P. palestinicum, photo Jessica Litman.

Figure 4. Lectotype, Pseudoanthidium reptans. A, face; B, dorsal view; C, labels; D, ventral view; E, apex of metasoma, showing tips of gonostyli.

Figure 5. Lectotype, Pseudoanthidium nanum. A, lateral view; B, dorsal view; C, labels.

Figure 6. Palaearctic distribution of A, Pseudoanthidium nanum; B, P. scapulare; C, P. stigmaticorne; D, P. tenellum.

Figure 7. Lectotype, Anthidium sinuatum. A, lateral view; B, labels; C, metasoma; D, T5–T6; E, mesonotum.

Figure 8. Lectotype Icteranthidium floripetum A, face; B, dorsal view; C, labels; D, vertex; E, T1–T3.

Figure 9. Dorsal habitus, females. A, vertex Pseudoanthidium nanum (Kalpetran, Switzerland); B, vertex P. scapulare (Rich, Morocco); C, mesonotum P.nanum (Kalpetran, Switzerland); D, mesonotum P.scapulare (Rich, Morocco); E, metasoma P. nanum (Kalpetran, Switzerland); F, metasoma P. scapulare (Rich, Morocco).

Figure 10. Gonostyli. A, Pseudoanthidium nanum (Thüringen, Germany); B, P. scapulare (Villeneuve-lès- Maguelone, France); C, P. tenellum (Krasnoperekopsk, Crimea).

Figure 11. Sternal combs. A, Pseudoanthidium nanum (Dordogne, France); B, P. scapulare (Argèles-sur-Mer, France); C, P. stigmaticorne (Cabanes de Fleury, France); D, P. tenellum (Krasnoperekopsk, Crimea); E, P. palestinicum (Jerusalem, Israel); F, P. cribratum (Ayelet Hashar, Israel); G, P. kaspareki (Side, Turkey); H, P. rozeni (Hanna, Pakistan).

Figure 12. S3, males. A, Pseudoanthidium nanum (Dordogne, France); B, P. scapulare (Argèles-sur-Mer, France); C, P. stigmaticorne (Cabanes de Fleury, France); D, P. tenellum (Krasnoperekopsk, Crimea); E, P. palestinicum (Jerusalem, Israel); F, P. cribratum (Ayelet Hashar, Israel); G, P. kaspareki (Side, Turkey); H, P. rozeni (Hanna, Pakistan).

Figure 13. S2, males. A, Pseudoanthidium nanum (Dordogne, France); B, P. scapulare (Argèles-sur-Mer, France); C, P. stigmaticorne (Cabanes de Fleury, France); D, P. tenellum (Krasnoperekopsk, Crimea); E, P. palestinicum (Jerusalem, Israel); F, P. cribratum (Ayelet Hashar, Israel); G, P. kaspareki (Side, Turkey); H, P. rozeni (Hanna, Pakistan).