Oxypilinae

Schwarz, Christian J. & Shcherbakov, Evgeny, 2017, Revision of Hestiasulini Giglio-Tos, 1915 stat. rev. (Insecta: Mantodea: Hymenopodidae) of Borneo, with description of new taxa and comments on the taxonomy of the tribe, Zootaxa 4291 (2), pp. 243-274 : 268-269

Key to Oxypilinae

(after Beier 1934, Roy & Svenson 2007, Roy 2013, and Svenson et al. 2015, modified); Bornean members are keyed out to species level.

1. Pronotum with 3–4 pairs of conical or spine-like tubercles; dilated part of femur about half as wide as non-dilated part; pos- tero-ventral margin of fore femur not or very slightly denticulate (tribus Oxypilini)................................. 2

- Pronotum smooth or slightly tuberculate, but never with conical or spine-like tubercles; dilated part of femur about as wide as non-dilated part; postero-ventral margin of fore femur distinctly denticulate (tribus Hestiasulini).......................7

2. Prozona with two pairs of conical or spine-like tubercles (Afrotropical)...........................................3

- Prozona with one pair of spine-like tubercles (Oriental).......................................................4

3. Walking leg femora without lobes; females apterous................................. Oxypilus Audinet-Serville, 1831

- Ventral lamina of walking leg femora produced into 3 small lobes; females brachypterous..................................................................................................... Junodia Schulthess-Rechberg, 1899

4. Metazona with two spine-like processes; process on the vertex long (over 75% width of head), bifid, with two pairs of smaller lateral processes; frontal shield with a median keel (genus Ceratomantis Wood-Mason, 1876 ).........................5

- Metazona with four spine-like processes, one paramedian and one lateral set; process on vertex short (less than 25% width of head), simple; frontal shield with two flattened, paramedian keels (genus Pachymantis Saussure, 1871 ).................6

5. Anterior side of fore coxa with a black median stripe extending from base to apex. Pronotum short, about 1.2 times as long as wide.......................................................................... C. kimberlae Svenson, 2007

- Anterior side of fore coxa with a subapical black spot only; pronotum longer, about 1.4 times as long as wide............................................................................................... C. gigliotosi Roy, 2007

6. Anterior side of fore coxa yellowish-brown with black apex and two large round spots; antero-ventral side of fore trochanter black; dilated part of fore femur black, ventral part reddish-brown............................ P. maculicoxa Roy, 2013

- Anterior side of fore coxa yellowish-brown at base, then largely black; antero-ventral side of fore trochanter yellowish-brown; anterior side of fore femur totally black.................................................. P. piceifemur Roy, 2013

7. Male subgenital plate with styli ( Figs 21–22 ); ventral phallomere with membraneous gap along left margin ( Figs 21–22 , 45– 50 ); dorsal lamina of left phallomere not folded longitudinally ( Figs 21–22 , 45–50 ); head with a process or tubercle ( Figs 1–2 , 51–52 ); ventral side of meso femora without lobe; alae of female hyaline or subhyaline ( Fig. 2 ).......................8

- Male subgenital plate without styli ( Figs 24–25 , 41 ); ventral phallomere without membraneous gap along left margin ( Figs 26– 28 , 42–44 ); dorsal lamina of left phallomere folded longitudinally ( Figs 26 , 42–44 ); head without tubercle ( Figs 3–6 , 33–34 , 53–59 ); if with tubercle ( Fig. 37 ) then ventral side of meso femora with a proximal lobe; alae of female (where known) subopaque ( Figs 4, 6 , 33–34 , 56 )........................................................................... 11

8. Discoidal field of tegmen in both sexes iridescent, without false crossveins, hyaline or very slightly infumate, without dark markings ( Figs 8 , 51–52 ); dorsal and ventral laminae of left phallomere fused along left margin ( Figs 22 , 45–47 ); apical process missing (Indo-Malayan) (genus Catestiasula Giglio-Tos, 1915 ).............................................9

- Discoidal field of tegmen in both sexes with false crossveins, slightly infumate to subopaque, with dark markings ( Figs 1–2 ); dorsal and ventral laminae of left phallomere not fused along left margin; apical process broad and short ( Figs 21 , 48–50 ) (Indo-Singhalese)....................................................................................10

9. Anterior side of fore femora with only one small whitish spot in the claw groove, distal spots, if present, not clearly white ( Figs 17–18 )....................................................................... C. moultoni Giglio-Tos, 1915

- Anterior side of fore femora with two to three whitish spots along ventral margin ( Figs 19–20 ).............................................................................................. C. nitida ( Brunner d. Wattenwyl, 1893)

10. Process on vertex with pointed apex in both sexes, and with a pair of accessory conical tubercles at base at least in females; paramedian carinae on frontal shield well-developed; denticulation at postero-ventral margin of fore femur heavier; discoidal field of tegmen at most with numerous dark spots.................................... Ephestiasula Giglio-Tos, 1915

- Process on vertex with bifid apex in females ( Fig. 2 ), without accessory conical tubercles at base; paramedian carinae on frontal shield indistinct; denticulation at postero-ventral margin of fore femur fainter; discoidal field of tegmen with darker apex and with a large smoky patch in proximal half (may be indistinct in males).................... Hestiasula Saussure, 1871

11. Vertex with a tubercle ( Fig. 37 ); ventral side of meso femora with a proximal lobe. Fore coxae dichromatic ( Fig. 13 ); dilated part of fore femur less wide than non-dilated part ( Figs 13 , 39 ) (genus Pseudohestiasula gen. nov.)..... P. borneana sp. nov.

- Vertex without tubercle ( Figs 3–6 , 33 , 53–59 ); meso femora without lobes; dilated part of fore femur much wider than nondilated part ( Figs 9–12, 16 ) (genus Astyliasula gen. nov.).....................................................12

12. Both pairs of wings of males with an extensive dark pattern ( Figs 3 , 57 ); pronotal ratio of female> 1.25............................................................................................... A. phyllopus ( De Haan, 1842)

- Both pairs of wings of males hyaline with dark apices, tegmina with a transverse line of brownish spots, alae with darkened posterior corner ( Figs 5 , 53 ); pronotal ratio of female <1.2............................. A. sarawaca ( Westwood, 1889)

Figures

FIGURES 21 – 22. Subgenital plate and genitalia of 21. H. brunneriana from India (ES) and 22. C. moultoni from Sabah (ES). Arrows indicate weakly sclerotized membrane at base of sclerotized left margin of ventral phallomere (scale bar: 1 mm).

FIGURES 43 – 50. Genitalia of various Hestiasulini. 43. A. phyllopus from Sabah (CS). 44. A. basinigra from Myanmar (SMNK). 45. C. nitida from Myanmar (SMNK). 46. C. nitida from Sumatra (SMNK). 47. C. sp. from Kerala (SMNK). 48. Ephestiasula amoena from Nepal (ES). 49. E. pictipes from India (CS). 50. H. gyldenstolpei from Thailand (ES) (scale bar: 1 mm).

FIGURES 1 – 8. Dorsal habitus of Hestiasulini. 1. Male of Hestiasula brunneriana (Sri Lanka) (CS). 2. Female of H. brunneriana (Sri Lanka) (CS). 3. Male of Astyliasula phyllopus from Sabah (CS). 4. Female of A. phyllopus from Brunei (CS). 5. Male of A. sarawaca from Brunei (SMNK). 6. Female of A. sarawaca from Kalimantan (RMNH). 7. Pseudohestiasula borneana, holotype (SMNK). 8. Male of Catestiasula moultoni from Sarawak (CS) (scale bar: 10 mm).

FIGURES 51 – 52. Life aspect of Catestiasula. 51. Male of C. moultoni, Mt. Trus Madi, Sabah, Malaysia; Photo © Alexey Yakovlev. 52. Male of unidentified Catestiasula specimen from Ulu Gombak NP, Selangor, Malaysia; Photo © Bernhard Jacobi.

FIGURES 23 – 25. Terminalia of A. sarawaca. 23. Supra-anal plate and cerci. 24. Subgenital plate, dorsal view. 25. Subgenital plate, posterior view (scale bar: 1 mm).

FIGURES 40 – 42. Genitalia of P. borneana. 40. Supra-anal plate and cerci (pilosity omitted). 41. Subgenital plate in dorsal view. 42. Genitalia in ventral view. Arrow indicates fold in dorsal lamina (scale bar: 1 mm).

FIGURES 26 – 32. Genitalia of A. sarawaca. 26. Genitalia of Trus Madi male in ventral view. 27 – 28. Ventral processes of two Brunei males. 29 – 30. Phalloid apophysises of two Brunei males. 31. Right phallomere in antero-sinistral view. 32. Left phallomere in dorsal view. Arrow indicates fold in dorsal lamina (scale bar: 1 mm).

FIGURES 33 – 36. Holotype of A. sarawaca; Photos Katherine Child © OUMNH. 33. Dorsal view and labels. 34. Ventral view (scale bar: 10 mm). 35. Pronotum in dorsal view (scale bar: 1 mm). 36. Right femur in anterior view (scale bar: 1 mm).

FIGURES 53 – 57. Life aspect of Bornean Astyliasula. 53. Male of A. sarawaca, Kubah NP, Sarawak, Malaysia; Photo © Fu- Sheng Huang. 54. Female of A. phyllopus in dorsal view, Mulu NP, Borneo, Sarawak, Malaysia; Photo © Paul Bertner. 55. Captive raised female of A. phyllopus in cryptic resting position; Photo © Daniel Döpfner. 56. A. phyllopus, displaying female, Mulu NP, Borneo, Sarawak, Malaysia; Photo © Paul Bertner. 57. A. phyllopus, displaying male, Mt. Trus Madi, Borneo, Sabah, Malaysia; Photo © Evgeny Shcherbakov.

FIGURES 58 – 59. A. phyllopus. 58. Ootheca with hatching nymphs. 59. Displaying first instar nymph. Photos © Jürg Sommerhalder.

FIGURES 37 – 39. P. borneana. 37. Head in anterior view. 38. Pronotum. 39. Left fore leg in anterior view (scale bar: 1 mm).

FIGURES 9 – 20. Fore legs of Hestiasulini treated in this paper, anterior view. 9. A. phyllopus, male from the Malay Peninsula (ES). 10. A. phyllopus, male from Brunei, mirrored (CS). 11. A. phyllopus, female from Brunei (CS). 12. A. sarawaca, male from Brunei (CS). 13. P. borneana, holotype (SMNK). 14. H. brunneriana, male from Sri Lanka (CS). 15. H. brunneriana, female from Sri Lanka (CS). 16. A. sarawaca, female from Kalimantan; Photo C. J. Schwarz © RMNH (RMNH). 17. C. moultoni, male from Sarawak (CS). 18. C. moultoni, female from Brunei, mirrored (CS). 19. C. nitida, male from Myanmar (SMNK). 20. C. nitida, male from Sumatra (SMNK) (not to scale).