Pseudoanthidium kaspareki, Litman & Fateryga & Griswold & Aubert & Proshchalykin & Divelec & Burrows & Praz, 2022

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, pp. 1287-1337 : 1321-1325

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 T1 with fringe of short, golden hairs mostly two puncture widths long ( Fig. 27E ); T1– T4 with pair of lateral, yellow, ovoid spots increasing in size from T1 to T4 ( Fig. 27E ); T5 with lateral yellow spots. Terga with coarse, irregular punctation ( Fig. 27E ); punctures on discs of T1–T4 as large or larger than those of mesonotum, much larger than punctation on depressions of T1–T4 ( Fig. 27E ); punctation of T1 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 T2–T4 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 T2– T4, with punctures separated only by carina-shaped interspaces, nearly coalescent on T3–T4 ( Fig. 27E ). T5 matte, with dense punctation on disc, punctures separated only by narrow, ridge-like interspaces; on depression punctation coalescent. T6 coriaceous with deep coalescent punctures on entire surface. T1–T4 with broad smooth apical margin, length equals 1.3– 1.4 diameter of largest punctures ( Fig. 27E ); T5 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 T2–T4 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 T2 ( 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 T2–T4 are strongly separated by flat, smooth interspaces, giving the integument a shiny appearance ( Fig. 27E ). In P. palestinicum , the punctures of discs T2–T4 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 T1 is distinctly coarser and sparser than it is apically ( Fig. 27E ), whereas the size and the density of the punctation of T1 is regular across the whole surface of the tergum in P. stigmaticorne ( Fig. 17E ). In P. kaspareki , the punctation of T2–T4 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 T2–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 T2–T4 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 <1 puncture diameter wide; apical margin of clypeus with row of short golden setae, may be obscured by shaggy white pilosity. Supraclypeal area sometimes with yellow ventral and lateral margins. Paraocular area pale yellow, matte; integument reticulate; sparse white pilosity present. Antenna with base dark, with tuft of shaggy, pale yellow to white pilosity; anterior face of flagellum brown-ferruginous, in some specimens with third and fourth flagellar segments entirely light ferruginous; posterior face of flagellum dark brown. Frons with shaggy yellowish hairs. Vertex with pair of slender yellow stripes on posterior margin, nearly meeting at centre. Vertex with short, golden hairs; punctation coarse, honeycomb areolate, punctures only slightly smaller than those of scutum. Gena honeycomb areolate with broad but shallow punctures.

Mesosoma : Integument mostly black. Scutum with punctation dense, coarse, honeycomb areolate, with spaces between punctures <0.25 puncture diameter wide, dull; punctures broader than those on vertex, less broad than those of T1–T3; surface covered with short, sparse golden setae; anterior margin with pair of lateral, rectangular yellow markings (in some case extending slightly down lateral margin of scutum). Pronotal lobe either entirely black, black and yellow, or entirely yellow. Axilla either yellow, black or black with yellow stains. Scutellum yellow laterally and posteriorly, with black portion of variable size medially and anteriorly; punctation of black area dense, coarse, honeycomb areolate, with spaces between punctures <0.25 puncture diameter wide, interspaces dull. Tegula ferruginous to dark ferruginous, except for yellow, crescent-shaped marking anteriorly and, sometimes, posteriorly. Mesepisternum shiny, with coarse, honeycomb areolate punctation anteriorly, becoming shallow and sparser posteriorly; small yellow spot present in some individuals. Metepisternum with shallow, honeycomb areolate punctation. Propodeum with lateral face finely, densely punctate; basal area of posterior face with row of coalescent, shallow punctures slightly interrupted medially; rest of posterior face smooth, shiny. Legs black to dark brown except apices of femora, tibiae and tarsal segments yellow; tibial spurs light brown to yellow-gold; trochanters with white pilosity at base, long and shaggy on fore and midlegs, shorter, of even length on hindleg. Wings infuscate.

Metasoma: Terga with integument shiny. T1–T5 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, T6 entirely black; T7 yellow. Punctures on T1–T3 broad with shiny spaces between punctures <0.5 puncture diameter wide; punctures broader than those of mesonotum; punctures on T4–T7 increasingly finer and denser. T1–T3 with broad shiny impunctate apical margin reaching maximal width of about 1.5 times diameter of broadest punctures. Impunctate apical margin of T4 slightly less broad, approximately 1.3 times diameter of broadest punctures. Impunctate margin of T5 narrower still, from approximately 0.8 to less than 0.5 puncture diameters wide medially. Margin of T6 dull, entirely punctate. T7 narrow, about twothirds the length of T6, with shallow rounded median emargination posteriorly. S2 with short, dense, velvety pubescence anteriorly, shiny, nearly glabrous posteriorly, medially with raised, comparatively dark, chevron-shaped zone, adorned with fringe of long, palecoloured hairs ( Fig. 13G ). S3 with short, dense, velvety pubescence anteriorly, posteriorly with premarginal brush of hairs, hooked at the tips, as well as an underlying comb of thickened, wavy hairs ( Fig. 12G ); medially with shiny, hairless chevron-shaped zone in which tip of chevron extends anteriorly as a carina along midline of sternum ( Fig. 12G ). S4 anteriorly with two wide, lateral patches of dense velvety pubescence, separated by glabrous zone, posteriorly shiny, glabrous, medially with longitudinal shiny furrow that begins at posterior margin and extends anteriorly, midline marked by dark, oblong depression. S5 with lateral comb relatively large, with longest teeth wider than maximal width of hind basitarsus ( Fig. 11G ). Gonostylus widest near apex, about 1.25 times as wide at widest point as at base ( Fig. 23E ); notch at apex weakly U-shaped to V-shaped, as deep as wide at opening and more or less centred on apex ( Fig. 23E ); inner margin mostly strongly curved near apex, outer margin most strongly curved about two-thirds of the way between base and apex ( Fig. 23E ). Penis valves nearly parallel-sided, rounded at tips ( Fig. 23E ).

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 T1–T3, 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.

Figures

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).

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).

Figure 21. Dorsal habitus, females.A,vertex Pseudoanthidiumtenellum (Pestszentimre,Hungary);B, vertex P.palestinicum (Rehovot, Israel); C, mesonotum P. tenellum (Pestszentimre, Hungary); D, mesonotum P. palestinicum (Rehovot, Israel); E, metasoma P. tenellum (Pestszentimre, Hungary); F, metasoma P. palestinicum (Rehovot, Israel).

Figure 23. Gonostyli. A, Pseudoanthidium palestinicum (Nahal Keziv, Israel); B, P. cribratum (Karatau, Kazakhstan); C, P. canariense (Santa Cruz de Tenerife, Canary Islands); D, P. tropicum (Bandar Abbas, Iran); E, P. kaspareki (Side, Turkey); F, P. rozeni (Hanna, Pakistan).

Figure 26. Distribution of A, Pseudoanthidium kaspareki; B, P. rozeni.

Figure 27. Dorsal habitus, females. A, vertex Pseudoanthidium kaspareki (Between Kula and Usak, Turkey); B, vertex P. rozeni (Hanna, Pakistan); C, mesonotum P. kaspareki (Between Kula and Usak, Turkey); D, mesonotum P. rozeni (Hanna, Pakistan); E, metasoma P. kaspareki (Between Kula and Usak, Turkey); F, metasoma P. rozeni (Hanna, Pakistan).