Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini) Author Litman, Jessica R. Author Fateryga, Alexander V. Author Griswold, Terry L. Author Aubert, Matthieu Author Proshchalykin, Maxim Yu. Author Divelec, Romain Le Author Burrows, Skyler Author Praz, Christophe J. text Zoological Journal of the Linnean Society 2022 2021-09-29 195 4 1 51 journal article 2789 10.1093/zoolinnean/zlab062 909b187d-8971-417e-a659-46e424c03ffe 0024-4082 5817276 CF1BB523-4E43-486B-9A4F-E510F1854B9B 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 ]. Figure 14. Lectotype Anthidium frontale . A, dorsal view; B, face; C, labels; D, lateral view. ? 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). Figure 15. Neotype Anthidium stigmaticorne . A, dorsal view; B, labels; C, face; D, lateral view. 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. Figure 16. Lectotype Pseudoanthidium karakalense . A, labels; B, dorsal view; C, latero-ventral view S2; D, gonostyli; E, S8; F, S3; G, S4; H, S5 (showing sternal combs), S6. 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 . Figure 17. Dorsal habitus, females. A, vertex Pseudoanthidium stigmaticorne (Arzens, France); B, vertex P. cribratum (Bukhara, Uzbekistan); C, mesonotum P. stigmaticorne (Arzens, France); D, mesonotum P. cribratum (Bukhara, Uzbekistan); E, metasoma P. stigmaticorne (Arzens, France); F, metasoma P. cribratum (Bukhara, Uzbekistan). 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.