Pseudoanthidium scapulare (Latreille, 1809) Le Divelec

Litman, Jessica R., Fateryga, Alexander V., Griswold, Terry L., Aubert, Matthieu, Proshchalykin, Maxim Yu., Divelec, Romain Le, Burrows, Skyler & Praz, Christophe J., 2021, Paraphyly and low levels of genetic divergence in morphologically distinct taxa: revision of the Pseudoanthidium scapulare complex of carder bees (Apoidea: Megachilidae: Anthidiini), Zoological Journal of the Linnean Society CLXVI (CLXVI), pp. 1-51 : 43-45

KEYS TO THE PSEUDOANTHIDIUM SCAPULARE COMPLEX

Males

1. Apex of coxa 3 with pronounced, flattened, round-tipped tooth, about as long as third tarsal segment is wide at apex ( Fig. 24A, B ) .............................................................................................................. P. cribratum

— Apex of coxa 3 without tooth, or at most with sharp angle much shorter than width of third tarsal segment at apex ( P. canariense , P. tropicum , P. tenellum , P. rozeni , P. scapulare , P. palestinicum , P. kaspareki , P. nanum , P. stigmaticorne ) .............................................................................................................................. 2

2. Apex of gonostylus without notch ( Fig. 23C, D ) ( P. canariense , P. tropicum ).................................................. 3

— Gonostylus notched apically ( Figs. 10A–C , 18A–F , 23A, E, F ) ( P. nanum , P. stigmaticorne , P. tenellum , P. rozeni , P. scapulare , P. palestinicum , P. kaspareki ) ..................................................................................... 4

3. Posterior margin of T 7 with deep, nearly semi-circular emargination medially ( Fig. 25E ); metasoma black with dark orange-yellow maculations ( Fig. 25C ); comparatively large (~ 9 mm)..................... P. canariense

— Posterior margin of T 7 straight, without emargination ( Fig. 25F ); metasoma brown with pale yellow maculations ( Fig. 25D ); comparatively small (~ 5 mm) ................................................................. P. tropicum

4. Lateral comb on S5 small, with longest teeth shorter than maximal width of hind basitarsus ( Fig. 11A, C ) ( P. nanum , P. stigmaticorne ) ............................................................................................................................. 5

— Lateral comb on S5 large, with longest teeth approximately equal to or greater in length than maximum width of hind basitarsus ( Fig. 11B, D, E, G, H ) ( P. tenellum , P. rozeni , P. scapulare , P. palestinicum , P. kaspareki ) ......................................................................................................................................................6

5. Darkened, raised area on S2 chevron-shaped ( Fig. 13A ); depressed area along apical margin of S2 triangular, shiny and glabrous ( Fig. 13A ); preapical patch of hooked hairs on S3 preceded by shiny, glabrous strip, hooked hair patch broad medially, attenuated laterally ( Fig. 12A ); gonostylus nearly parallel sided from base to apex, only slightly wider apically than at base, apical notch deep and nearly centred at apex of gonostylus ( Fig. 10A ); apical comb of S5 broader than its preceding arm ( Fig. 11A )...................... P. nanum

— Darkened raised area on S2 not chevron-shaped ( Fig. 13C ); depressed area along apical margin of S2 not triangular, with short, proclinate, golden hairs and less shiny ( Fig. 13C ); preapical patch of hooked hairs on S3 not preceded by shiny, glabrous strip, pubescent throughout, hooked hair patch equally wide across width of segment ( Fig. 12C ); gonostylus approximately 1.5 times wider at widest point than at base, with apical notch usually less deep than opening of notch is wide (notch especially shallow in specimens from northern Africa and some specimens from southern Italy, and considerably deeper in specimens from most European and Central Asian populations, up to approximately as deep as wide at opening), notch strongly offset laterally ( Fig. 18A–F ); apical comb of S5 no broader than its preceding arm ( Fig. 11C ) ................................................................................................................................................... P. stigmaticorne

6. Hairs on premarginal zone of S3 mostly straight or somewhat bent apically but not forming a distinctly rounded hook and arranged in a brushy arc, exposing nearly entire length of underlying comb of wavy hairs ( Fig. 12D, H ); S2 with strong, long fringe greatly overhanging margin throughout ( Fig. 13B, E, G ) ( P. tenellum , P. rozeni )....................................................................................................................................... 7

— Hairs on premarginal zone of S3 nearly all forming distinctly rounded hooks and arranged more or less directly on top of underlying comb of wavy hairs so at most only apical half of wavy hairs visible ( Fig. 12B, E, G ), S2 without strong fringe, hairs not reaching beyond the apical margin ( Fig. 13D, H ) ( P. scapulare , P. palestinicum , P. kaspareki ) ........................................................................................................................... 8

7. Lateral comb on S5 mitten shaped ( Fig. 11D ); apical notch of gonostylus weakly V-shaped, wider than deep ( Fig. 10C ) ........................................................................................................................................... P. tenellum

— Lateral comb on S5 mostly straight, although they may have a small nub where the ‘thumb’ would be ( Fig. 11H ); apical notch of gonostylus strongly U-shaped, deeper than wide ( Fig. 23F ) ........................... P. rozeni

8. Punctures on terga relatively small, smaller than or equal to width of punctures on scutellum; S4 with a strong V-shaped emargination medially on posterior margin ...................................................... P. scapulare

— Punctures on terga relatively large, larger than those on scutellum; posterior margin of S4 not emarginate posteriorly ( P. palestinicum , P. kaspareki )....................................................................................................... 9

9. Lateral comb on S5 broad and strongly chevron-shaped ( Fig. 11E ); penis valve flattened and evenly tapering to a pointed tip; curve of both inner and outer margins of gonostylus is strongest about two-thirds of way between base and apex ( Fig. 23A ) ................................................................................. P. palestinicum

— Lateral comb on S5 smaller and more evenly rounded ( Fig. 11G ); penis valve nearly parallel-sided and with a rounded tip ( Fig. 23E ); curve of the inner margin of gonostylus is strongest apically, while curve of outer margin is strongest about two-thirds of way between base and apex ( Fig. 23E )………… P. kaspareki

Females

1. Body length less than 5 mm; dark maculations of head and metasoma predominantly reddish-brown ( Fig. 25B )................................................................................................................................................... P. tropicum

— Body length 5 mm or more; dark parts of head and metasoma predominantly black.................................. 2

2. Punctation on T 1– T 3 comparatively fine, punctures smaller than those on mesonotum; in addition, punctation of vertex conspicuously finer than that of scutellum ( P. scapulare , P. nanum ) ( Fig. 9A–F )....... 3

— Punctation on T 1– T 3 comparatively coarse, punctures as large or larger than those on mesonotum (condition not clear in P. tenellum ) ( Figs 17C–F , 21C–F , 25A , 27C–F ); punctation of vertex variable ( Figs 17A, B , 21A, B , 25A , 27A, B ) ( P. canariense , P. tenellum , P. cribratum , P. stigmaticorne , P. palestinicum , P. kaspareki , P. rozeni ) ......................................................................................................................................4

3. Terga on average densely punctured; interspaces between punctures narrow, less than half a puncture diameter, and glittery but not shiny ( Fig. 9B, D, F ); impunctate posterior margin on T 4 weakly shagreened, glittery but not shiny ( Fig. 9F ); base of F1 often with strongly pronounced tooth; clypeus entirely yellow ......................................................................................................................................................... P. scapulare

— Terga on average less densely punctured ( Fig. 9E ); interspaces between punctures sometimes up to half a puncture diameter or more ( Fig. 9E ); overall impression of terga shiny ( Fig. 9E ); impunctate posterior margin of T 4 very weakly shagreened, thus appearing shiny ( Fig. 9E ); F1 usually at most with small tooth or angle; clypeus most often black but may also be yellow with black markings, or entirely yellow ............................................................................................................................................................. P. nanum

4. Maculations on head, mesosoma and metasoma dark orange; head in frontal view entirely dark, without orange maculations (Canary Islands) ( Fig. 25A )......................................................................... P. canariense

— Maculations on head, mesosoma and metasoma yellow; at least paraocular area with yellow maculations .......................................................................................................................................................................... 5

5. Punctures on vertex between compound eye and lateral ocellus distinct, only slightly finer and denser than punctures on T 1 and T 2 ( Figs 17B, F , 21A, E , 27B, F ). Vertex and mesonotum shiny with distinct smooth, shiny interspaces between punctures ( Figs 17B, D , 21A, C , 27B, D ). Punctures of scutellum as dense and at least as large as largest punctures of T 2 ( Figs 17D, F , 21C, E , 27D, F ) ( P. tenellum , P. cribratum , P. rozeni ) ...........................................................................................................................................................................6

— Diameter of punctures on vertex between lateral ocellus, eye and postoccipital margin distinctly smaller than that of punctures on T 1 or T 2 (although in some P. stigmaticorne , punctation on vertex is only slightly finer than that on terga) ( Figs. 17A, E , 21B, F , 27A, E ). Vertex and mesonotum comparatively matte with narrow carina-shaped interspaces ( Figs. 17A, C , 21B, D , 27A, C ). Punctures of scutellum denser and clearly finer than largest punctures of T 2 (except in P. stigmaticorne , where punctation on scutellum is either as dense or slightly denser and as large or slightly larger than that of T 2) ( Figs. 17C, E , 21D, F , 27C, E ) ( P. palestinicum , P. stigmaticorne , P. kaspareki )......................................................................................... 8

6. Punctation on T 1 medially comparatively finer, punctures visibly smaller than those on black part of scutellum ( Fig. 21C, E ); maculations on European specimens creamy white to pale yellow (but lemonyellow in Central Asian specimens) ( Fig. 21A, C, E ) ....................................................................... P. tenellum

— Punctation on T 1 medially coarse, punctures not visibly different from those on black part of scutellum; maculations always dark yellow ( Figs 17D, F , 27D, F ) ( P. cribratum , P. rozeni )............................................ 7

7. Punctation of T 2 coarser, larger than on black part of scutellum ( Fig. 17D, F ); mesonotum with surface between punctures shagreened ( Fig. 17D ).................................................................................... P. cribratum

— Punctation of T 2 finer, punctures not visibly different from those on black part of scutellum ( Fig. 27D, F ); mesonotum with surface between punctures polished ( Fig. 27D )...................................................... P. rozeni

8. Punctation on vertex between lateral ocellus, eye and postoccipital margin appears rough and sometimes indistinct, narrow spaces between punctures matte ( Fig. 21B ); scutellum and axillae entirely predominantly yellow, with only a reduced dark area medially ( Fig. 21D ); clypeus yellow; large, mostly> 0.9 mm ..................................................................................................................................... P. palestinicum

— Punctation on vertex between lateral ocellus, eye and postoccipital margin small but distinct, with shiny narrow spaces between most punctures ( Figs 17A , 27A ); scutellum often with two separated yellow spots extending to part of axilla, medially mostly with an extended, triangular dark area ( Figs 17C , 27C ); clypeus either yellow, yellow with black markings or black; small, mostly ≤ 0.8 mm ( P. stigmaticorne , P. kaspareki ) ...........................................................................................................................................................................9

9. Terga with finer, more regular punctation ( Fig. 17E ); punctation on disc of T 2 nearly as large as punctation of mesonotum ( Fig. 17C, E ); punctation of T 1– T 4 dense and mostly separated by narrow ridge-like interspaces so integument appears relatively matte ( Fig. 17E ). Punctation of scutellum as dense, or slightly less so, than punctation on disc of T 2 ( Fig. 17C, E ). .................................................. P. stigmaticorne

— Terga with a coarse, irregular punctation ( Fig. 27E ); punctation of T 2 much larger than punctation of mesonotum ( Fig. 27C, E ); punctation of T 1 is distinctly sparser in its basal half where punctures are separated by flat smooth interspaces (distances reaching one-third to half the diameter of a puncture), while apically punctures are separated by narrow ridge-like interspaces ( Fig. 27E ); punctation of T 2– T 4 coarse and sparse on disc where punctures are separated by flat, smooth interspaces (distances reaching one-third to half the diameter of a puncture), so that the integument appears shiny ( Fig. 27E ); punctures of apical depressions are coalescent. Punctation is much denser on scutellum than on disc of T 2, punctures separated only by minute carina-shaped interspaces ( Fig. 27C, E ) ............................................. P. kaspareki

Figures

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).

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 18. Gonostyli, Pseudoanthidium stigmaticorne. A, Cape Lukull, Crimea; B, Bet She’an, Israel; C, Agadir, Morocco; D, Restinclières, France; E, Cacela Velha, Portugal; F, Seui, Sardinia.

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 24. Third coxal tooth (circled by white ring), Pseudoanthidium cribratum, male. A, ventral view (Damascus, Syria); B, lateral view (Yasuj, Iran).

Figure 25. A, Pseudoanthidium canariense, female (Santa Cruz de Tenerife, Canary Islands); B, P. tropicum, female (Bandar Abbas, Iran); C, P. canariense, male (Santa Cruz de Tenerife, Canary Islands); D, P. tropicum, male (Bandar Abbas, Iran); E, T7, P. canariense (Santa Cruz de Tenerife, Canary Islands); F, T7, P. tropicum (Bandar Abbas, Iran).

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).