Sphecodes, Latreille, 1804

Astafurova, Yulia & Proshchalykin, Maxim, 2020, New and little-known bees of the genus Sphecodes Latreille, 1804 (Hymenoptera: Apoidea: Halictidae) from the Himalayas, European Journal of Taxonomy 729, pp. 74-120 : 78-85

Key to the species of Sphecodes of the Himalayas

Males are unknown in S. almoransis , S. binghami , S. bluethgeni sp. nov., S. iridipennis , S. perplexus , S. setiger , S. tantalus , and females are unknown in S. dissimilandus , S. invidus , S. shillongensis .

Males

1. Costal margin of hind wing with 7–14 hamuli. Base of gonocoxite dorsally without impression. Usually larger species: total body length 6.0–14.0 mm .................................................................... 2

– Costal margin of hind wing with 5–6 hamuli. Base of gonocoxite dorsally with or without impression. Usually smaller species: total body length 5.0– 7.5 mm ................................................................. 14

2. Gena flat. Preoccipital lateral carina well-developed ....................................................................... 3

– Gena swollen. Preoccipital carina not developed ............................................................................. 8

3. Fore wing with two cubital cells. Tyloids weakly developed, narrowly semicircular across at most ¼ of the basal flagellar surfaces ( Fig. 2H ). Body length 7.0– 8.5 mm .......... S. turneri Cockerell, 1916

– Fore wing with three cubital cells ..................................................................................................... 4

4. Tyloids covering entire ventral flagellar surface ( Fig. 2G ). Body length 7.0– 8.5 mm ....................... ................................................................................................................... S. kershawi Perkins, 1921

– Ventral flagellar surface with medial glabrous spot or tyloids narrowly semicircular across basal flagellar surface ................................................................................................................................. 5

5. T1 with marginal zone impunctate. Body length 7.0– 8.5 mm ...................... S. laticeps Meyer, 1920

– T1 with marginal zone distinctly punctate (at least on a basal half) ................................................. 6

6. Wings with strong brownish darkening. Tyloids weakly developed, covering at most basal ¼ of ventral flagellar surface and narrowly linear across lateral surface ( Fig. 2D ). Large species: total body length more 12.0 mm ................................................................ S. sikkimensis Blüthgen, 1927

– Wings hyaline, without darkening. Tyloids well developed, covering entire lateral flagellar surface and peripheral part of ventral surface (with medial glabrous spot, variable in size). Smaller species: total body length 7.0– 8.5 mm ........................................................................................................... 7

7. Vertex more elevated, distance from top of head to upper margin of lateral ocellus ca two lateral ocellar diameters as seen in frontal view. Ocello-ocular area with confluent punctures (areolatepunctate). Metasomal terga coarser punctate (15–25 μm). Body length 7.5–8.5 mm ........................ ................................................................................................................. S. invidus ( Cameron, 1897)

– Vertex less elevated, distance from top of head to upper margin of lateral ocellus ca 1.5 lateral ocellar diameter as seen in frontal view. Ocello-ocular area with small shiny interspaces between punctures (at most a half puncture diameter). Metasomal terga finer punctate (10–20 μm). Body length 7.0 mm .............................................................................. S. dissimilandus ( Cameron, 1897)

8. Head rounded, approximately as long as wide. Hind wing with basal vein strongly curved, the angle between basal (M) and cubital (Cu) veins ca 90°. T1 finely and sparsely (sometimes indistinctly) punctate. Gonostylus dorsally with small rectangular process directed to penis valve ( Fig. 3B ). Body length 7.0– 10 mm ................................................................................ S. monilicornis ( Kirby, 1802)

– Head transverse, wider than long. Hind wing with basal vein weakly curved, the angle between basal (M) and cubital (Cu) veins 70–80°. T1 coarsely and densely punctate. Gonostylus of another shape. ...............................................................................................................................................................9

9. Mesoscutum densely punctate, with confluent punctures (areolate-punctate) ............................... 10

– Mesoscutum sparser punctate, medially with punctures separated by at least a puncture diameter .. ..........................................................................................................................................................11

10. Head more transverse, 1.25–1.3 times as wide as long. Gonostylus longer, apically elongated into rounded process with long setae ( Fig. 3E ). Body length 9.0–12.0 mm .............................................. ............................................................................................................. S. albilabris ( Fabricius, 1793)

– Head less transverse, ca 1.2 times as wide as long. Gonostylus short, not enlarged apically ( Fig. 3A ). Body length 9.5–12 mm .......................................................................... S. fumipennis Smith, 1853

11. Vertex long, distance from top of head to upper margin of lateral ocellus approximately three lateral ocellar diameters as seen in dorsal view. Tyloids usually well developed, covering at least entire lateral flagellar surface (with medial glabrous spot, variable in size, on ventral surface) .............. 12

– Vertex shorter, distance from top of head to upper margin of lateral ocellus ca two lateral ocellar diameters as seen in dorsal view. Tyloids weakly developed, covering basal 1/7–⅓ of ventral flagellar surface ............................................................................................................................................. 13

12. F3 long, ca 2 times as long as wide ( Fig. 2E ). Gonostylus elongate, with long apical process ( Fig. 3I ), apically less emarginated as seen in lateral view. Body length 7.0–14.0 mm .................................... .................................................................................................................. S. gibbus ( Linnaeus, 1758)

– F3 short, 1.1–1.3 times as long as wide ( Fig. 2F ). Gonostylus without apical process ( Fig. 3H ), apically more emarginate as seen in lateral view. Body length 6.0–8.0 mm ...................................... ................................................................................................................ S. rufiventris (Panzer, 1798)

13. Tyloids less developed, covering basal 1/7–1/5 of ventral flagellar surface ( Fig. 2C ). Body length 7.0–12.0 mm .............................................................................................. S. alternatus Smith, 1853

– Tyloids more developed, covering basal 1/5–⅓ of ventral flagellar surface. Body length 10.0 mm .. .............................................................................................................. S. lasimensis Blüthgen, 1927

14. T1 densely punctate. Gonostylus large, rectangular and weakly S -curved on inner margin ( Fig. 3F ). Body length 5.0– 7.5 mm ................................................................... S. intermedius Blüthgen, 1923

– T1 impunctate or with a few punctures. Gonostylus of another shape ........................................... 15

15. Tyloids well developed, on flagellomeres (at least from F3 onward) covering entire ventral surface ......................................................................................................................................................... 16

– Tyloids weakly developed, covering at most basal ¼ of ventral surface of last flagellomeres ...... 17

16. F2 with tyloids covering entire ventral surface ( Fig. 2B ). Hypoepimeral area entirely reticulaterugose. Base of gonocoxite dorsally without impression, gonostylus leaf-shaped. Body length 5.0– 5.5 mm ............................................................................................... S. cameronii ( Bingham, 1897)

– F2 glabrous, without tyloids ( Fig. 2A ). Hypoepimeral area medially smooth. Base of gonocoxite dorsally with impression, gonostylus trapezoidal ( Fig. 3D ). Body length 4.5–6.0 mm ..................... ........................................................................................................... S. hakkariensis Warncke, 1992

17. Vertex with longitudinal carina ( Fig. 4B ). Lateral preoccipital carina present ( Fig. 4A ). Gonostylus with large rectangular membranous part ( Fig. 3C ). Body length 7.0– 7.5 mm ................................... .................................................................................................................... S. montanus Smith, 1879

– Vertex without longitudinal carina. Lateral preoccipital carina absent. Gonostylus smaller, of another shape ............................................................................................................................................... 18

18. Antennae shorter, with flagellomeres (from F3 onward) ca 1.1–1.2 times as long as wide. Gonostylus with triangular membranous part ( Fig. 3G ). Body length 5.0– 5.5 mm .............................................. .............................................................................................................. S. simlaensis Blüthgen, 1924

– Antennae longer, with flagellomeres (from F3 onward) ca 1.3 times as long as wide. Gonostylus without membranous part. Body length 5.0 mm ............................. S. shillongensis Blüthgen, 1927

Females

1. Costal margin of hind wing with 7–14 hamuli. Usually larger species: total body length 6.5– 15.0 mm ............................................................................................................................................ 2

– Costal margin of hind wing with 5–6 hamuli. Usually smaller species: total body length 4.0– 8.5 mm ............................................................................................................................................ 13

2. Gena flat. Preoccipital lateral carina well-developed ....................................................................... 3

– Gena swollen. Preoccipital carina not developed ............................................................................. 7

3. Fore wing with two cubital cells. Body length 8.0–9.0 mm ..................... S. turneri Cockerell, 1916

– Fore wing with three cubital cells ..................................................................................................... 4

4. Large species: total body length 12.0–14.0 mm. Costal margin of hind wing with 11–15 hamuli .... ............................................................................................................ S. sikkimensis Blüthgen, 1927

– Smaller species: total body length 7.0–9.0 mm. Costal margin of hind wing with 6–9 hamuli ....... 5

5. Head strongly transverse ( Fig. 5B ), ca 1.3 times as wide as long. T1 impunctate. Body length 7.0– 8.0 mm ........................................................................................................... S. laticeps Meyer, 1920

– Head less transverse ( Fig. 5D ), ca 1.2 times as wide as long. T1 distinctly and densely punctate .. 6

6. Mesoscutum with coarse and confluent punctures, but medially with interspaces 0.5–1 puncture diameter. Costal margin of hind wing with 8–9 hamuli. Pygidial plate as wide as metabasitarsus. Body length 8.0–9.0 mm ....................................................................... S. binghami Blüthgen, 1924

– Mesoscutum mostly areolate-punctate. Costal margin of hind wing with 6–7 hamuli. Pygidial plate 0.7 times as wide as metabasitarsus. Body length 7.5–8.0 mm ................ S. kershawi Perkins, 1921

7. Mesoscutum densely punctate, with punctures separated by less than a puncture diameter ( Fig. 4C ) ........................................................................................................................................................... 8

– Mesoscutum sparsely punctate, medially with some punctures separated by at least two puncture diameters ( Fig. 4F ) ............................................................................................................................ 9

8. Paraocular areas with dense plumose pubescence, obscuring integument. Mesepisternum reticulaterugose ( Fig. 4E ). T1 densely punctate, with punctures separated by 0.5–2 puncture diameters. Body length 9.0–15.0 mm ............................................................................ S. albilabris ( Fabricius, 1793)

– Paraocular areas with relatively dense plumose pubescence, not obscuring integument. Mesepisternum areolate ( Fig. 4D ). T1 sparsely punctate, with punctures separated by 2–6 puncture diameters. Body length 9.5–12.0 mm ....................................................................... S. fumipennis Smith, 1853

9. Terga with marginal zones almost impunctate, on T1 only with a few tiny punctures. Body length 7.5–10.0 mm ........................................................................................ S. lasimensis Blüthgen, 1927

– Terga with marginal zones distinctly punctate at least on T1 and T2 ............................................. 10

10. Vertex shorter, distance from top of head to upper margin of lateral ocellus ca two lateral ocellar diameters as seen in dorsal view. Propodeum laterally reticulate-rugose. T3 with marginal zone distinctly punctate. Body length 8.0–11.0 mm .......................................... S. alternatus Smith, 1853

– Vertex longer, distance from top of head to upper margin of lateral ocellus equal to 2.5–3.0 lateral ocellar diameters as seen in dorsal view. Propodeum laterally strigate or reticulate-rugose. T3 with marginal zone impunctate or punctate .............................................................................................11

11. Head rounded-rectangular on upper margin, square-shaped as seen in frontal view ( Fig. 5C ); vertex sparsely punctate, punctures mostly separated by more than а puncture diameter. Hind wing with basal vein strongly curved, the angle between basal (M) and cubital (Cu) veins ca 90º. T1 indistinctly punctate, with a few fine punctures. Pygidial plate 0.9–1.0 times as wide metabasitarsus. Body length 7.0–10.0 mm ............................................................................. S. monilicornis ( Kirby, 1802)

– Head uniformly rounded on upper margin, oval as seen in frontal view; vertex densely punctate, punctures mostly separated by less than a puncture diameter. Hind wing with basal vein weakly curved, the angle between basal (M) and cubital (Cu) veins ca 70–80°. T1 distinctly punctate, with fine and coarser punctures. Pygidial plate 0.5–0.6 times as wide as metabasitarsus ...................... 12

12. Mesepisternum strigate or strigate-rugose at least ventrally, sides of propodeum strigate. T3 with marginal zone coarsely punctate. Body length 6.5–8.5 mm .................. S. rufiventris (Panzer, 1798)

– Mesepisternum reticulate-rugose, sides of propodeum strigate or reticulate-rugose. T3 with marginal zone impunctate. Body length 7.0–15.0 mm ........................................... S. gibbus ( Linnaeus, 1758)

13. Mandible simple (without an inner tooth) ...................................................................................... 14

– Mandible bi-dentate ........................................................................................................................ 15

14. Paraocular and supraclypeal areas with moderately dense pubescence, not obscuring integument. F1 and F2 as long as wide. Mesoscutum with punctures separated by at most a puncture diameter. Metasomal terga (except almost impunctate T1) coarser punctate (10–15 μm). Pygidial plate as wide as metabasitarsus. Body length 5.5 mm ...................................................... S. setiger Blüthgen, 1924

– Paraocular and supraclypeal areas with dense pubescence, obscuring integument ( Fig. 5A ). F1 and F2 0.6–0.7 times as long as wide. Mesoscutum finely and irregularly punctate with punctures separated by 1–5 puncture diameters. Metasomal terga finely punctate (3–10 μm). Pygidial plate 1.4–1.5 times as wide as metabasitarsus. Body length 5.0– 5.5 mm ................................................... ............................................................................................................ S. cameronii ( Bingham, 1897)

15. Vertex with longitudinal carina. Lateral preoccipital carina present, but weakly developed. Pygidial plate wider, 1.1–1.2 times as wide as metabasitarsus. Body length 7.0– 7.5 mm ............................... .................................................................................................................... S. montanus Smith, 1879

– Vertex without longitudinal carina. Lateral preoccipital absent. Pygidial plate narrower, at most as wide as metabasitarsus .................................................................................................................... 16

16. Pronotum rounded between the dorsal and lateral surfaces. Body length 7.0– 7.5 mm ( Fig. 6 ) ......... .......................................................................................................................... S. bluethgeni sp. nov.

– Pronotum between dorsal and lateral surfaces with sharp angle .................................................... 17

17. T1 distinctly punctate. Pygidial plate narrow, 0.4 times as wide as metabasitarsus. Body length 6.5–8.5 mm ........................................................................................ S. intermedius Blüthgen, 1923

– T1 impunctate or with sparse tiny punctures. Pygidial plate wider, at least 0.7 times as wide as metabasitarsus ................................................................................................................................. 18

18. Head strongly transverse, ca 1.3 times as wide as long. Metasomal terga entirely red .................. 19

– Head less transverse, ca 1.2 times as wide as long. T4 and T5 at least partially brownish or black .. ......................................................................................................................................................... 21

19. Smaller species: total body length 4.0 mm. Legs entirely black ........................................................ ..................................................................................................... S. almoransis Gupta & Saini, 2018

– Larger species: total body length 6.5–8.0 mm. Legs entirely red or at least tarsi red or yellow .... 20

20. Mesoscutum denser punctate, with punctures separated by 1–2 puncture diameters. Pygidial plate as wide as metabasitarsus. Body length 6.5 mm ............................................. S. perplexus Nurse, 1903

– Mesoscutum sparser punctate, with punctures separated by 2–5 puncture diameters. Pygidial plate 0.8 times as wide as metabasitarsus. Body length 8.0 mm ........................... S. tantalus Nurse, 1903

21. Mesoscutum sparsely punctate, with punctures separated by 2–6 puncture diameters ( Fig. 4F ). Hypoepimeral area medially smooth. Body length 4.5–5.0 mm ...... S. hakkariensis Warncke, 1992

– Mesoscutum denser punctate, with punctures separated by 1–4 puncture diameters. Hypoepimeral area entirely rugose ......................................................................................................................... 22

22. Head less transverse, 1.1 times as wide as long. Legs entirely red. Body length 4.5 mm ................. ................................................................................................................... S. iridipennis Smith, 1879

– Head more transverse, 1.2 times as wide as long. At least femora black or brownish. Body length 5.0– 6.5 mm .......................................................................................... S. simlaensis Blüthgen, 1924

Figures

Fig. 2. Antennae in lateral view, males. A. Sphecodes hakkariensis Warncke, 1992. B. S. cameronii (Bingham, 1897).C. S. alternatus Smith, 1853.D. S. sikkimensisBlüthgen, 1927. E. S. gibbus (Linnaeus, 1758). F. S. rufiventris (Panzer, 1798). G. S. kershawi Perkins, 1921. H. S. turneri Cockerell, 1916. Scale bars = 0.5 mm.

Fig. 3. Genitalia in dorsal view, males. A. Sphecodes fumipennis Smith, 1853. B. S. monilicornis (Kirby, 1802). C. S. montanus Smith, 1879. D. S. hakkariensis Warncke, 1922. E. S. albilabris (Fabricius, 1793). F. S. intermedius Blüthgen, 1923. G. S. simlaensis Blüthgen, 1924. H. S. rufiventris (Panzer, 1798). I. S. gibbus (Linnaeus, 1758). Scale bars = 0.25 mm.

Fig. 4. Diagnostic characters of species of Sphecodes. A–B. Males. C–F. Females. A. Genal area and lateral preoccipital carina in lateral view. B. Vertex and longitudinal carina in dorso-lateral view. C–F. Mesosoma in dorsal (C, F) and lateral views (D–E). A–B. S. montanus Smith, 1879. C–D. S. fumipennis Smith, 1853. E. S. albilabris (Fabricius, 1793). F. S. hakkariensis Warncke, 1922. Scale bars = 1.0 mm.

Fig. 5. Head in frontal view, females. A. Sphecodes cameronii (Bingham, 1897). B. S. laticeps Meyer, 1920. C. S. monilicornis (Kirby, 1802). D. S. kershawi Perkins, 1921. Scale bars = 1.0 mm.

Fig. 6. Sphecodes bluethgeni sp. nov. Holotype, ♀ (PCMS/OLBL). Habitus in lateral view and labels. Scale bar = 1 mm.