Epeolus, Latreille, 1802

Onuferko, Thomas M., 2019, A review of the cleptoparasitic bee genus Epeolus Latreille, 1802 (Hymenoptera: Apidae) in the Caribbean, Central America and Mexico, European Journal of Taxonomy 563, pp. 1-69 : 52-62

Key to species of the genus Epeolus in the Caribbean, Central America and Mexico

1. Axilla in dorsal view with tip extending as far back as apex of horizontal dorsal portion of mesoscutellum ( Fig. 9C ). Axilla and mesoscutellum entirely ferruginous ( Fig. 9C ). T1–T4 fasciate, apical fasciae separated into rounded lobes medially ( Fig. 9B ) [Hispaniola] .................................... .................................................................................... E. danieli ( Genaro, 2014) comb. nov. ( Fig. 9 )

– Mesosoma and metasoma without the combination of features listed above [other regions] .......... 2

2. Pronotal collar with anterior margin relatively straight (best seen in dorsal view) ( Fig. 2 A–B). Mesoscutellum with distinct ridge along apex of horizontal dorsal portion overhanging depressed posterior margin (best seen in posterior view) ( Fig. 3 A–B), except in E. pulchellus ( Fig. 3C ). Penis without pair of fleshy lateral lobes ( Fig. 1 A–D, G–I). Frontal area commonly with pair of sparsely punctate to impunctate protrusions, each located near upper mesal margin of compound eye ( Fig. 4A ) ................................................................................................... 3 (‘ Trophocleptria group’)

– Pronotal collar with anterior margin convex (best seen in dorsal view) ( Fig. 2C ). Mesoscutellum without distinct ridge delineating dorsal and posterior portions ( Fig. 3D ). Penis with pair of divergent, fleshy lateral lobes ( Fig. 1 E–F). Frontal area never with protrusions ( Fig. 4 C–F) ............................. ................................................................................................................... 11 (all other Epeolus spp.)

3. Metasomal terga with tomentum entirely black ( Figs 8 A–C, 15A–C). Fore wing dusky throughout or deeply infuscate basally, clear and hyaline apically ( Figs 8 A–B, 15B) ....................................... 4

– At least T2 with yellow apical fascia ( Figs 6 A–C, 10A–C, 11A–C, 16A–C, 17A–C, 19A–C). Fore wing dusky throughout or most deeply infuscate on apical margin ( Figs 6 A–B, 10B, 11A–B, 16A–C, 17A–C, 19C) ................................................................................................................................ 5

4. Mesoscutellum with pair of posteriorly directed teeth ( Fig. 8D ). Mesopleuron coarsely punctate, with punctures in ventrolateral half sparse (many i≥2d) ( Fig. 5A ) .................................................... ..................................................................................................... E. claripennis Friese, 1908 ( Fig. 8 )

– Mesoscutellum without pair of posteriorly directed teeth ( Fig. 15D ). Mesopleuron finely punctate, with punctures in ventrolateral half dense (few i≥2d) ( Fig. 5B ) ........................................................ .................................................................................................... E. niger ( Michener, 1954) ( Fig. 15 )

5. Axilla with lateral margin crenulate, with large tooth near base. Mesoscutellum with pair of posteriorly directed teeth ( Figs 6D , 10D , 16D ) ................................................................................ 6

– Axilla with lateral margin smooth or if crenulate, then without large tooth near base. Mesoscutellum without pair of posteriorly directed teeth ( Figs 11D , 17D , 19D ) ...................................................... 8

6. T1 with only (bright to pale yellow) subapical fascia, which is usually narrower (its breadth sometimes equaling just a few hairs) than the (bright to pale yellow) apical fascia of T2 ( Fig. 6B ) ....................................................................................... E. boliviensis Friese, 1908 ( Fig. 6 )

– T1 with medially narrowed or interrupted (bright to pale yellow) basal fascia, which is as broad as or broader than the (bright to pale yellow) apical fascia of T2 ( Figs 10B , 16B ) .............................. 7

7. T3 and T4 with complete, bright yellow fasciae ( Fig. 10 A–C) .......................................................... ............................................................................................ E. fulvopilosus Cameron, 1902 ( Fig. 10 )

– T3 and T4 without fasciae, although apical impressed areas occasionally with sparse, off-white hairs ( Fig. 16 A–C) ............................................................................... E. nomadiformis sp. nov. ( Fig. 16 )

8. Mesoscutum with well-defined paramedian band ( Figs 2B , 11B ). Axilla in dorsal view with tip extending as far back as or beyond apex of horizontal dorsal portion of mesoscutellum ( Fig. 11D ) ...................................................................................... E. fumipennis Say, 1837 ( Fig. 11 )

– Mesoscutum without paramedian band ( Figs 17B , 19B ). Axilla in dorsal view with tip not extending as far back as apex of horizontal dorsal portion of mesoscutellum ( Figs 17D , 19D ) ....................... 9

9. Mesoscutum with anteromedial ovate patch of bright yellow tomentum as well as a pair of large posterolateral patches ( Fig. 19B ) [ Cuba] ................................ E. pulchellus Cresson, 1865 ( Fig. 19 )

– Mesoscutum with pale tomentum limited to pair of small patches, each between tegula and axilla, and posterior margin narrowly ( Fig. 17B, D ) [Central America and Mexico] ............................... 10

10. Frontal area always with pair of pronounced granulose protrusions, each located near upper mesal margin of compound eye ( Fig. 4A ). Pseudopygidial area of female shorter medially, such that its apex is more than twice as wide as medial length ( Fig. 12B ). T3–T6 of male without fasciae; apical impressed areas occasionally with sparse, off-white hairs, but these do not form distinct bands. Pronotal collar, pronotal lobe, axilla and mesoscutellum entirely ferruginous ( Fig. 20A ) ................................................... E. bifasciatus Cresson, 1864 ( Onuferko 2018a: fig. 26)

– Frontal area with pair of protrusions usually not strongly pronounced, in some cases almost lacking entirely ( Fig. 4B ). Pseudopygidial area of female longer medially, such that its apex is at most twice as wide as medial length ( Fig. 12C ). T3–T6 of male typically with well-developed bright to pale yellow fasciae ( Fig. 17C ). Pronotal collar, pronotal lobe, axilla and mesoscutellum entirely dark brown or black to entirely ferruginous ( Fig. 17 ) ................................................................................ .............................................................................. E. obscuripes Cockerell, 1917 stat. nov. ( Fig. 17 )

11. Axilla in dorsal view with tip extending to or beyond ⅔ the length of mesoscutellum ( Fig. 21 A–C) ......................................................................................................................................................... 12

– Axilla in dorsal view with tip extending to less than ⅔ the length of mesoscutellum ( Figs 13D , 14D , 18D , 21 D–F) ................................................................................................................................... 18

12. Axilla with free portion ~ 2 ∕ 5 of its entire medial length or longer and usually distinctly hooked (i.e., concave, not relatively straight along medial margin) ( Fig. 21A ). Axilla with lateral margin convex ( Fig. 21A ) or relatively straight ( Fig. 22A ) .................................................................................... 13

– Axilla with free portion clearly less than 2 ∕5 of its entire medial length ( Fig. 21 B–C) or if borderline (0.35<x <0.4) then relatively straight (i.e., not concave) along medial margin. Axilla with lateral margin convex ( Fig. 21 B–C) .......................................................................................................... 15

13. Metanotum with distinct posteromedial depression ( Fig. 3E ). T1–T4 with apical fasciae interrupted medially ( Fig. 22A ) ...................................... E. axillaris Onuferko, 2018 ( Onuferko 2018a: fig. 18)

– Metanotum without depression ( Fig. 3F ). T1–T4 with apical fasciae complete ( Fig. 22B ) .......... 14

14. Preoccipital ridge joining hypostomal carina (approximately at 2 ∕ 5 length of proboscidial fossa) ( Fig. 23A ). Mandible simple ( Fig. 24A ) .......... E. ainsliei Crawford, 1932 ( Onuferko 2018a: fig. 4)

– Preoccipital ridge not joining hypostomal carina ( Fig. 23B ). Mandible with small, obtuse preapical (almost submedial) tooth ( Fig. 24B ) .............. E. rufulus Cockerell, 1941 ( Onuferko 2018a: fig. 81)

15. Head dorsally with pair of small protrusions, each located where upper genal area meets vertexal area ( Fig. 4C ). Mesopleuron of male obscured by white tomentum only in upper half (although hypoepimeral area usually with sparser tomentum), with large, sparsely hairy circular area occupying much of ventrolateral half ( Fig. 20B ). Axilla in dorsal view with tip extending to or beyond band of pale tomentum along posterior margin of mesoscutellum ( Fig. 21B ) ................................................ ............................................................................. E. scutellaris Say, 1824 ( Onuferko 2018a: fig. 83)

– Head dorsally without pair of protrusions ( Fig. 4D ). Mesopleuron of male (excluding hypoepimeral area) entirely obscured by white tomentum ( Fig. 20C ). Axilla in dorsal view with tip at most extending to band of pale tomentum along posterior margin of mesoscutellum ( Fig. 21C ) .......... 16

16. T2–T4 each with fascia on or only very slightly separated from apical margin of tergum, more or less evenly broad. T1 with longitudinal extent of discal patch variable, but commonly less than breadth of apical fascia ( Fig. 25A ) ............... E. novomexicanus Cockerell, 1912 ( Onuferko 2018a: fig. 73)

– T2 and T3 (for female) or T2–T4 (for male) each with fascia clearly separated from apical margin of tergum medially, commonly narrowed or narrowly interrupted medially. T1 with longitudinal extent of discal patch no less (and usually greater) than breadth of apical fascia ( Fig. 25B ) ................... 17

17. Flagellum with ventral surface and usually also metasomal sterna same reddish brown or reddish orange color as legs ( Fig. 26A ) ......................... E. basili Onuferko, 2018 ( Onuferko 2018a: fig. 24)

– Entire flagellum, except sometimes F1, and metasomal sterna (excluding apical margins) dark brown or black, clearly not the same reddish-orange color as legs from tibiae to tarsi ( Fig. 26B ) ............... .......................................................................... E. pusillus Cresson, 1864 ( Onuferko 2018a: fig. 79)

18. T1 without apical fascia ( Figs 14B , 18B ) ...................................................................................... 19

– T1 with apical fascia ( Figs 13B , 22 C–L) ........................................................................................ 20

19. Frontal keel without tooth-like process ( Fig. 27A ). Mesopleuron with punctures in ventrolateral half dense (most i<1d) ( Fig. 5C ). T1 with complete or medially interrupted off-white to pale yellow basal fascia ( Fig. 14B ). Antenna, pronotal lobe and legs reddish orange in part ( Fig. 14 A–C) .................. ................................................................................................... E. luteipennis Friese, 1916 ( Fig. 14 )

– Frontal keel with small tooth-like process ( Fig. 27B ). Mesopleuron with punctures in ventrolateral half sparse (most i>1d) ( Fig. 5D ). T1 without fasciae ( Fig. 18B ). Antenna, pronotal lobe and legs entirely dark brown to black ( Fig. 18 A–C) .................................... E. odyneroides sp. nov. ( Fig. 18 )

20. Mesoscutum with anteromedial patch of bright or pale yellow tomentum ( Fig. 22 C–F) .............. 21

– Mesoscutum with gray or bright to pale yellow paramedian band (usually parallel and not joined except sometimes posteriorly) or largely obscured by pale tomentum ( Fig. 22 G–L); if joined posteriorly (i.e., U- or V-shaped), then not distinctly narrowed anterolaterally ............................. 24

21. Dorsum of mesosoma and metasoma with bright yellow short, appressed setae. T1–T4 with apical fasciae complete. T1 with little if any space between off-white basal fascia and bright yellow apical fascia ( Fig. 22C ) ....................................... E. splendidus Onuferko, 2018 ( Onuferko 2018a: fig. 85)

– Dorsum of mesosoma and metasoma with pale yellow short, appressed setae. At least apical fascia of T1 medially interrupted. T1 with elongate discal patch (medially as long as or longer than breadth of apical fascia) between off-white to pale yellow basal and apical fasciae ( Fig. 22 D–F) ............ 22

22. T 1 in dorsal view with apical fascia straight, basal fascia arched and indistinguishable from longitudinal band, such that discal patch is semicircular or triangular ( Fig. 22D ) ............................. .................................................................... E. canadensis Mitchell, 1962 ( Onuferko 2018a: fig. 30)

– T 1 in dorsal view with basal and apical fasciae subparallel, at angles from longitudinal band, such that discal patch is quadrangular, trapezoidal or (if fasciae thickened laterally) diamond-shaped ( Fig. 22 E–F) .................................................................................................................................... 23

23. T1–T4 with apical fasciae broadened submedially and either narrowed or separated into rounded lobes medially ( Fig. 22E ) ........................... E. compactus Cresson, 1878 ( Onuferko 2018a: fig. 36)

– T1–T4 with apical fasciae not broadened submedially; fasciae more or less evenly broad and uninterrupted medially or tapering and thus narrowed or separated medially ( Fig. 22F ) .................. ........................................................................ E. ferrarii Onuferko, 2018 ( Onuferko 2018a: fig. 45)

24. Vertexal area with two pairs of shiny (usually impunctate) protrusions ( Fig. 4E ). T1 apical fascia with pair of small posterolateral patches of black tomentum ( Fig. 22G ) ....................................... 25

– Vertexal area without two pairs of protrusions, at most with single pair of closely punctate dorsal protrusions, each located where upper genal area meets vertexal area ( Fig. 4F ). T1 apical fascia without pair of small posterolateral patches of black tomentum ( Fig. 22 H–L) .............................. 26

25. Mesopleuron with punctures in ventrolateral half sparse (few i≤1d) ( Fig. 5E ) .................................. ......................................................... E. chamaesarachae Onuferko, 2018 ( Onuferko 2018a: fig. 34)

– Mesopleuron with punctures in ventrolateral half dense (many i≤1d) ( Fig. 5F ) ................................ ................................................................. E. diadematus Onuferko, 2018 ( Onuferko 2018a: fig. 41)

26. Metanotum with blunt median process ( Fig. 13D ). T 1 in dorsal view with discal patch in shape of rounded triangle with anterolateral sides concave ( Fig. 13B ) ........................................................ 27

– Metanotum without process ( Fig. 21 D–F). T1 with discal patch variable, but if triangular in dorsal view then with anterolateral sides straight or convex ( Fig. 22H ) ................................................... 28

27. Mesopleuron either with most interspaces between punctures small (i≤1d) throughout or rugose, with punctures ill-defined ( Fig. 5G ) ................................................... E. hanusiae sp. nov. ( Fig. 13 )

– Mesopleuron with punctures in ventrolateral half well separated (i>1d), upper half more densely punctate than ventrolateral half ( Fig. 5H ) ........................................................................................... ................................................................. E. interruptus Robertson, 1900 ( Onuferko 2018a: fig. 61)

28. T 1 in dorsal view with apical fascia straight, basal fascia arched, though in some cases widely interrupted medially, and indistinguishable from longitudinal band, such that discal patch is semicircular or triangular ( Fig. 22H ) ........ E. flavofasciatus Smith, 1879 ( Onuferko 2018a: fig. 47)

– T 1 in dorsal view with basal and apical fasciae subparallel, usually at angles from longitudinal band, such that discal patch is quadrangular, trapezoidal, or elliptical, but sometimes thickened laterally such that discal patch is diamond-shaped or in the shape of a pointed oval ( Fig. 22 I–L) .............. 29

29. T2–T4 with apical fasciae complete, more or less evenly broad ( Fig. 22 I–J) ................................ 30

– T1 and T2 with apical fasciae broken or at least greatly narrowed medially, those of T3 and T4 broken or complete ( Fig. 22 K–L) ................................................................................................... 32

30. Fore wing with two submarginal cells, apically dusky in female, hyaline throughout in male ( Fig. 20D ). Mesopleuron almost entirely obscured by tomentum, at least in male ( Fig. 20D ). Axilla and mesoscutellum black ( Fig. 22I ) ........... E. mesillae (Cockerell, 1895) ( Onuferko 2018a: fig. 67)

– Fore wing with three submarginal cells, subhyaline, apically dusky in both sexes ( Fig. 20E ). Mesopleuron obscured by tomentum only in upper half, with a large, sparsely hairy circle occupying much of ventrolateral half ( Fig. 20E ). At least axilla ferruginous in part ( Fig. 22J ) ...................... 31

31. Frontal keel strongly raised ( Fig. 27C ). Pygidial plate of male narrow (medial length ~1.5 × basal width) ( Fig. 28A ) ........................................... E. australis Mitchell, 1962 ( Onuferko 2018a: fig. 14)

– Frontal keel weakly protuberant ( Fig. 27D ). Pygidial plate of male broad (medial length ≈ basal width) ( Fig. 28B ) ......................................... E. brumleyi Onuferko, 2018 ( Onuferko 2018a: fig. 28)

32. Axilla and mesoscutellum entirely ferruginous ( Fig. 21D ). Mesopleuron with punctures in ventrolateral half sparse (most i>1d) ( Fig. 5I ) .................................................................................... ....................................................................... E. tessieris Onuferko, 2018 ( Onuferko 2018a: fig. 87)

– Axilla (except sometimes tip) and mesoscutellum black ( Fig. 21 E–F). Mesopleuron with punctures in ventrolateral half dense (most i≤1d) ( Fig. 20F ) .......................................................................... 33

33. F2 of female at least 1.2 × as long as wide ( Fig. 29A ). Axilla of both sexes with free portion more than ¼ as long as its entire medial length, its tip distinctly pointed ( Fig. 21E ) .................................. .................................................................. E. minimus (Robertson, 1902) ( Onuferko 2018a: fig. 69)

– F2 of female at most 1.1 × as long as wide ( Fig. 29B ). Axilla of both sexes with free portion at most ¼ as long as its entire medial length, its tip broadly rounded ( Fig. 21F ) ....................................... 34

34. T3 and T4 each with pair of ovate patches of pale tomentum. Legs mostly to entirely dark brown or black ( Fig. 22K ) ......................................... E. asperatus Cockerell, 1910 ( Onuferko 2018a: fig. 10)

– T3 and T4 each with complete or medially and/or laterally interrupted fascia. Legs mostly to entirely reddish orange ( Fig. 22L ) ...................... E. barberiellus Cockerell, 1907 ( Onuferko 2018a: fig. 22)

Figures

Fig. 9. Epeolus danieli (Genaro, 2014) comb. nov., holotype, ♀ (MNHNSD 18.107). A. Habitus, lateral view. B. Habitus, dorsal view. C. Mesosoma, dorsal view. D. Posterior part of metasoma, dorsal view (blue arrow indicates process of S6). Images courtesy of G. de los Santos, MNHNSD. Scale bars: A–B = 3 mm; C–D = 1 mm.

Fig. 2. Pronotal collar of female, dorsal view. A. E. boliviensis Friese, 1908, which is straight along its anterior margin. B. E. fumipennis Say, 1837, which is relatively straight along its anterior margin. C. E. hanusiae sp. nov., paratype (UCBME), which is convex along its anterior margin. Scale bars = 1 mm.

Fig. 3. Mesosoma of female, posterior view. A. E. fulvopilosus Cameron, 1902, showing mesoscutellum with a distinct ridge (blue arrow) overhanging its depressed posterior margin. B. E. fumipennis Say, 1837, showing mesoscutellum with a faint ridge (blue arrow) overhanging its depressed posterior margin. C. E. pulchellus Cresson, 1865, showing mesoscutellum without a distinct ridge delineating its dorsal and posterior portions. D. E. odyneroides sp. nov., paratype (KUNHM SEMC1248301), showing mesoscutellum without a distinct ridge delineating its dorsal and posterior portions. E. E. axillaris Onuferko, 2018, paratype, showing metanotum with a distinct posteromedial depression (blue arrow). F. E. rufulus Cockerell, 1941, holotype, showing flat metanotum. Scale bars = 1 mm.

Fig. 1. Male genitalia of Epeolus spp. A. E. boliviensis Friese, 1908, in which the penis lacks lobes. B. E. claripennis Friese, 1908, in which the penis lacks lobes. C. E. fulvopilosus Cameron, 1902, in which the penis lacks lobes. D. E. fumipennis Say, 1837, in which the penis lacks lobes. E. E. hanusiae sp. nov., paratype (CNC 754086), showing the penis with a short, fleshy lateral lobe (blue arrow) on each side. F. E. luteipennis Friese, 1916, showing the penis with a long, fleshy lateral lobe (blue arrow) on each side. G. E. niger (Michener, 1954), in which the penis lacks lobes. H. E. obscuripes Cockerell, 1917 stat. nov., in which the penis lacks lobes. I. E. pulchellus Cresson, 1865, in which the penis lacks lobes. Scale bars = 0.5 mm.

Fig. 4. Head of female. A. E. bifasciatus Cresson, 1864 showing frontal area with impunctate, granulose protrusion (blue arrow) on each side. B. E. obscuripes Cockerell, 1917 stat. nov. showing frontal area with very weak/inconspicuous protrusion (blue arrow) on each side. C. E. scutellaris Say, 1824 showing dorsal protrusion (blue arrow) on each side where upper genal area meets vertexal area. D. E. pusillus Cresson, 1864 showing frontal and vertexal areas without protrusions. E. E. chamaesarachae Onuferko, 2018, paratype, showing vertexal area with two shiny, impunctate protrusions (blue arrows) on each side. F. E. flavofasciatus Smith, 1879 showing single (closely punctate) dorsal protrusion (blue arrow) on each side where upper genal area meets vertexal area. Scale bars = 1 mm.

Fig. 8. Epeolus claripennis Friese, 1908. A. Female, habitus, lateral view. B. Female, habitus, dorsal view. C. Male, habitus, lateral view. D. Female axillae and mesoscutellum, dorsal view (blue arrow indicates one of two posteriorly directed tooth-like projections of mesoscutellum). Scale bars: A–C = 3 mm; D = 0.5 mm.

Fig. 6. Epeolus boliviensis Friese, 1908. A. Female, habitus, lateral view. B. Female, habitus, dorsal view. C. Male holotype, habitus, lateral view, ZMB. D. Female axillae and mesoscutellum, dorsal view (blue arrow indicates one of two posteriorly directed tooth-like projections of mesoscutellum). Scale bars: A–C = 3 mm; D = 0.5 mm.

Fig. 5. Mesopleuron of female, lateral view. A. E. claripennis Friese, 1908, holotype (ZMB), blue arrow indicates carina delineating the anterior and lateral surfaces of the mesopleuron. B. E. niger (Michener, 1954). C. E. luteipennis Friese, 1916. D. E. odyneroides sp. nov., paratype (KUNHM SEMC1248301). E. E. chamaesarachae Onuferko, 2018, paratype. F. E. diadematus Onuferko, 2018, paratype. G. E. hanusiae sp. nov., paratype (UCBME). H. E. interruptus Robertson, 1900. I. E. tessieris Onuferko, 2018, paratype. Scale bars = 0.5 mm.

Fig. 15. Epeolus niger (Michener, 1954). A. Female, habitus, lateral view. B. Female, habitus, dorsal view. C. Male, habitus, lateral view. D. Female axillae and mesoscutellum, dorsal view. Scale bars: A–C = 3 mm; D = 0.5 mm.

Fig. 10. Epeolus fulvopilosus Cameron, 1902. A. Female, habitus, lateral view. B. Female, habitus, dorsal view. C. Male, habitus, lateral view. D. Female axillae and mesoscutellum, dorsal view (blue arrow indicates one of two posteriorly directed tooth-like projections of mesoscutellum). Scale bars: A–C = 3 mm; D = 0.5 mm.

Fig. 16. Epeolus nomadiformis sp. nov., ♀♀. A–B. Holotype, (KUNHM SEMC1248290). A. Habitus, lateral view. B. Habitus, dorsal view. C–D. Paratype, (CNC 754064). C. Habitus, lateral view. D. Axillae and mesoscutellum, dorsal view (blue arrow indicates one of two posteriorly directed tooth-like projections of mesoscutellum). Scale bars: A–C = 3 mm; D = 0.5 mm.

Fig. 11. Epeolus fumipennis Say, 1837. A. Female, habitus, lateral view. B. Female, habitus, dorsal view. C. Male, habitus, lateral view. D. Female axillae and mesoscutellum, dorsal view (blue lines indicate the posterior extent of the axilla relative to the length of the horizontal dorsal portion of the mesoscutellum). Scale bars: A–C = 3 mm; D = 0.5 mm.

Fig. 17. Epeolus obscuripes Cockerell, 1917 stat. nov. A. Female, habitus, lateral view. B. Female, habitus, dorsal view. C. Male, habitus, lateral view. D. Female axillae and mesoscutellum, dorsal view (blue lines indicate the posterior extent of the axilla relative to the length of the horizontal dorsal portion of the mesoscutellum). Scale bars: A–C = 3 mm; D = 0.5 mm.

Fig. 19. Epeolus pulchellus Cresson, 1865 A. Female, habitus, lateral view. B. Female, habitus, dorsal view. C. Male, habitus, lateral view. D. Female axillae and mesoscutellum, dorsal view (blue lines indicate the posterior extent of the axilla relative to the length of the horizontal dorsal portion of the mesoscutellum). Scale bars: A–C = 3 mm; D = 0.5 mm.

Fig. 12. Pseudopygidial area of female, dorsal view. A. E. fumipennis Say, 1837, which consists of a basal impressed triangular or semicircular portion covered in silvery short, appressed setae differentiated from a transverse band of coppery short, appressed setae along the posterior-facing apical margin of T5. B. E. bifasciatus Cresson, 1864, which is more than twice as wide as long. C. E. obscuripes Cockerell, 1917 stat. nov., which is not more than twice as wide as long. The pseudopygidial area is the apical portion of T5 that changes slope from the rest of the tergum and is covered in short, coppery or silvery hairs uniform in length (posteromesad the light blue lines). Scale bars = 1 mm.

Fig. 20. Epeolus spp., habitus of male, lateral view. A. E. bifasciatus Cresson, 1864. B. E. scutellaris Say, 1824. C. E. basili Onuferko, 2018, paratype. D. E. mesillae (Cockerell, 1895). E. E. australis Mitchell, 1962. F. E. asperatus Cockerell, 1910. Scale bars = 3 mm.

Fig. 21.Axillae and mesoscutellum of female, dorsal view. A. E. ainsliei Crawford, 1932. B. E. scutellaris Say, 1824. C. E. basili Onuferko, 2018, paratype. D. E. tessieris Onuferko, 2018, holotype. E. E. minimus (Robertson, 1902). F. E. asperatus Cockerell, 1910. Blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length. Scale bars = 0.5 mm.

Fig. 13. Epeolus hanusiae sp. nov. A–B, D. Holotype, ♀ (EMEC 1135889). C. Allotype, ♂ (EMEC 1135885). A. Habitus, lateral view. B. Habitus, dorsal view. C. Habitus, lateral view. D. Axillae and mesoscutellum, dorsal view (blue arrow indicates blunt median process of metanotum; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum). Scale bars: A–C = 3 mm; D = 0.5 mm.

Fig. 14. Epeolus luteipennis Friese, 1916. A. Female, habitus, lateral view. B. Female, habitus, dorsal view. C. Male, habitus, lateral view. D. Female axillae and mesoscutellum, dorsal view (blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum). Scale bars: A–C = 3 mm; D = 0.5 mm.

Fig. 18. Epeolus odyneroides sp. nov. A–B, D. Holotype, ♀ (KUNHM SM0253729). C. Allotype, ♂ (KUNHM SM0255860). A. Habitus, lateral view. B. Habitus, dorsal view. C. Habitus, lateral view. D. Axillae and mesoscutellum, dorsal view (blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum). Scale bars: A–C = 3 mm; D = 0.5 mm.

Fig. 22A–F (continued on next page). Epeolus spp. habitus of female, dorsal view. A. E. axillaris Onuferko, 2018, paratype. B. E. ainsliei Crawford, 1932. C. E. splendidus Onuferko, 2018, paratype. D. E. canadensis Mitchell, 1962. E. E. compactus Cresson, 1878. F. E. ferrarii Onuferko, 2018, paratype. Scale bars = 3 mm.

Fig. 23. Head of male, posterior view. A. E. ainsliei Crawford, 1932, in which the preoccipital ridge joins the hypostomal carina (blue arrow). B. E. rufulus Cockerell, 1941, in which the preoccipital ridge does not join the hypostomal carina (blue arrow). Scale bars = 1 mm.

Fig. 24. Mandibles of female A. E. ainsliei Crawford, 1932, each of which lacks a preapical tooth. B. E. rufulus Cockerell, 1941, each of which has a blunt, obtuse preapical (almost submedial) tooth (blue arrow). Scale bars = 0.5 mm.

Fig. 22G–L (continued). Epeolus spp., habitus of female, dorsal view. G. E. chamaesarachae Onuferko, 2018, paratype. H. E. flavofasciatus Smith, 1879. I. E. mesillae (Cockerell, 1895). J. E. brumleyi Onuferko, 2018, paratype. K. E. asperatus Cockerell, 1910. L. E. barberiellus Cockerell, 1907. Scale bars = 3 mm.

Fig. 25. Metasoma of female, dorsal view. A. E. novomexicanus Cockerell, 1912. B. E. basili Onuferko, 2018, paratype. Scale bars = 2 mm.

Fig. 26. Epeolus spp., habitus of female, ventral view. A. E. basili Onuferko, 2018, paratype showing antennae, legs, and metasomal sterna with similar reddish-orange coloration. B. E. pusillus Cresson, 1864 showing color contrast between reddish-orange legs and dark brown antennae and metasomal sterna. Scale bars = 3 mm.

Fig. 27. Head of female, lateral view. A. E. luteipennis Friese, 1916, in which the frontal keel lacks a tooth-like process. B. E. odyneroides sp. nov., holotype (KUNHM SM0253729), in which the frontal keel has a small tooth-like process (blue arrow). C. E. australis Mitchell, 1962, in which the frontal keel (blue arrow) is strongly raised. D. E. brumleyi Onuferko, 2018, paratype, in which the frontal keel (blue arrow) is weakly protuberant by comparison. Scale bars = 1 mm.

Fig. 28. Pygidial plate of male, dorsal view. A. E. australis Mitchell, 1962 (longer than wide and apically narrowed). B. E. brumleyi Onuferko, 2018, allotype (about as long as wide and broadly rounded apically). Scale bars = 1 mm.

Fig. 29. Antenna (basal portion) of female. A. E. minimus (Robertson, 1902), in which F2 is noticeably longer than wide. B. E. asperatus Cockerell, 1910, in which F2 is not noticeably longer than wide. Scale bars = 0.5 mm.