Brachysandalus Stål, 1867

Liu, Yingqi & Cai, Wanzhi, 2024, Revision of Australian Brachysandalus with the description of nine new species including one cavernicolous species (Hemiptera: Heteroptera: Reduviidae), and observations on male extragenital structure and leg teratology, Zootaxa 5490 (1), pp. 1-112 : 6-8

Genus Brachysandalus Stål, 1867

Brachysandalus Stål, 1867: 260 ; Maldonado Capriles (1990: 345).

Type species: Brachysandalus lurco Stål, 1867 , by subsequent designation ( Jeannel, 1919: 253).

Brachysandalus : Stål (1874: 59), as subgenus of Pirates .

Brachysandalus : Villiers (1968: 1175), reinstated as genus.

Brachysandalus : Cassis & Gross (1995: 339) and ABRS (accessed August 2023), as subgenus of Peirates .

Diagnosis. Hemelytron usually with a pale (variants of yellow / white / orange) oval or variable shaped spot involving apical area of clavus and adjoining area of corium between veins Pcu+1A and Cu (e.g., Fig. 1A ) (note, only rarely this pale spot is indistinguishable), and an inconspicuous yellowish, thin, curved stripe near base of costal margin on membrane (e.g., Fig. 1A ).

Head with postocular region ellipsoidal, almost rounded to neck, neck with lateral tubercles tiny (e.g., Fig. 1A ). Pronotum with collar lateral process distinct and rounded apically, lateral margins of anterior pronotal lobe arcuate (e.g., Fig. 1A ); ventral surface of fore femur with bristly setae and / or tubercles of varying size and shape, or teeth; fore and mid tibiae gradually thickened and expanded to apex, fore tibia with fossula spongiosa occupying over 1/3 but no more than 1/2 tibial length, mid tibia with fossula spongiosa occupying about 1/3 tibial length (e.g., Figs.1B&C ). Brachypterous form with anterior pronotal lobe spherical, posterior lobe distinctly abbreviated; hemelytron rarely surpassing second abdominal tergite (e.g., Fig. 4A ). Males with sternum VII on left side with (e.g., Figs. 55 , 56 ) or less commonly without an extragenital process. Pygophore oval in ventral view (e.g., Fig. 2A ), paramere curved and usually subtriangular (e.g., 2D&E), dorsal phallothecal sclerite, strongly sclerotized and broad, apex obtuse or rounded (e.g., Fig. 2F ); base of venter of phallosoma with a pair of slender sclerites (e.g., Fig. 2G ).

Redescription. Head: Sub-fusiform in dorsal view, anteocular region elongate triangular, postocular region ellipsoidal, anteocular region distinctly longer than postocular, frontoclypeal sulcus distinct; postocular region almost rounded to neck, neck with lateral tubercles not developed; ventral surface of head tumid before eyes; maxillary plate developed. Antennae with all segments cylindrical, scape thickest and shortest, apical three segments slender and of subequal length. Eye reniform, usually not reaching ventral margin in lateral view, width of eye usually shorter than width of interocular space in dorsal view, sometimes wider than width of interocular space in males. Ocelli well developed and located on tubercles in macropterous form, reduced, obsolete and not raised or slightly raised in brachypterous form. Labium with second visible segment longest, tumid on ventral side.

Thorax: Pronotum with collar lateral process distinct, apex rounded, anterior pronotal lobe subtrapezoidal with lateral margins arcuate, posterior pronotal lobe arcuately quadrate, humeri rounded, length of anterior pronotal lobe less than twice length of posterior lobe in macropterous form; anterior pronotal lobe nearly spherical, posterior lobe distinctly shortened, much shorter than anterior lobe in brachypterous form; integument of stripes on anterior lobe pilose and rough. Propleuron with integument often finely granulate, set off from dorsal surface by a carina, stridulitrum long total-striate type. Scutellum triangular, slightly wider than long, disc flat, “Y” shaped ridges distinct. Pleural and sternal integument finely granulate, metapleural sulcus curved with median part often nearly straight in macropterous form and entirely arcuate in brachypterous form; mesosternum with disc raised, metasternum raised with disc mostly flat. Hemelytra fully developed in macropterous or reduced in brachypterous forms.

Legs: Fore coxa elongate, mid and hind coxae globular, mid coxae separated from each other by less than width of one coxa. Fore femur highly thickened, much thicker than mid and hind femora, ventral surface of fore femur usually with tubercles or teeth of varying size and shape, usually arranged in inner and outer rows separated by a shallow median furrow along length of femur, in addition with bristly setae, ventral surface of mid femur also often armed with tubercles or teeth in addition to bristly setae. Fore and mid tibiae gradually thickened to apex, apex expanded, fore tibia with fossula spongiosa occupying over 1/3 but no more than 1/2 tibial length, mid tibia with fossula spongiosa occupying about 1/3 tibial length, hind tibia slender and straight, slightly incrassate at apex. Tarsi three-segmented.

Abdomen: Elongate parallel sided in male and nearly fusiform in female; venter of abdomen in male usually with a median ridge running from posterior portion of second abdominal sternite to posterior portion of seventh abdominal sternite and curved slightly to right side of body. Male sternum VII on left side with, or less commonly without, a visible extragenital process. Pygophore oval in ventral view with median pygophore process oblique to right side; paramere curved and usually subtriangular; dorsal, and lateral phallothecal sclerites strongly sclerotized, dorsal phallothecal sclerite broad, apex obtuse or rounded; base of venter of phallosoma with a pair of slender sclerites.

Comparative notes. In addition to Brachysandalus , there are three peiratine genera distributed in Australia, viz., Ectomocoris Mayr, 1865 , Microsandalus Stål, 1867 and Sirthenea Spinola, 1837 . Brachysandalus can be easily separated from Sirthenea by anteocular part of head less than three times as long as postocular part and antennal insertion near anterior margin of eye (vs. anteocular part of head more than three times as long as postocular part and antennal insertion approximately at middle of anteocular part of head in Sirthenea ). Brachysandalus is more similar in general body shape with Ectomocoris and Microsandalus , and most Australian species of these three genera have wing dimorphism. But Brachysandalus differs from the other two in fore tibia with fossula spongiosa occupying over 1/3 but no more than 1/2 tibial length (vs. fore tibia with fossula spongiosa occupying more than 1/2 of tibial length in Ectomocoris and fore tibia with fossula spongiosa occupying slightly less than 1/3 of tibial length in Microsandalus ). Also, in Brachysandalus , ventral surface of fore femur with bristly setae and / or tubercles of varying size and shape, or teeth, but in Ectomocoris , ventral surface of fore femur with bristly setae only, without any tubercle or tooth, and in Brachysandalus , anterior pronotal lobe subtrapezoidal with collar lateral process distinct, while in Microsandalus , anterior pronotal lobe subspherical and plump with collar process not developed.

Brachysandalus was treated as subgenus of Peirates by Stål (1874), but it could be distinguished from Peirates by the armature on ventral surface of fore femur mentioned above and the common colour pattern that hemelytron usually with a pale spot involving apical area of clavus and adjoining area of corium between veins Pcu+1A and Cu (vs. ventral surface of fore femur without any tubercle or tooth and colour pattern of hemelytron varies among different species in Peirates ).

Key to species of Australian Brachysandalus

1. Macropterous—submacropterous......................................................................... 2

-. Brachypterous....................................................................................... 21

2. Hemelytron with pale (variants of yellow / orange / white), usually oval shaped patch involving only parts of clavus and adjoining area of corium between veins Pcu+1A and Cu (e.g., Fig. 1A ) (except this patch almost indistinguishable on corium in B. fuliginosus ( Erichson, 1842) ( Figs. 14A , 15A ))......................................................... 3

-. Hemelytron with pale (variants of yellow / orange / white) area involving parts of clavus and adjoining area of corium between veins Pcu+1A and Cu as well as an extensive area of most of corium and basal area of membrane (e.g., Figs. 10 , 11A ); (male extragenital process absent or only weakly developed, no males of B. erythromelas ( Walker, 1873) examined)........... 17

3. Membrane with broad subapical pale brownish yellow area contrasting with dark brown remainder area ( Fig. 12A )............................................................................... B. flavidus Malipatil & Liu , sp. nov.

-. Membrane without such subapical brownish yellow area (e.g., Fig. 1A ).......................................... 4

4. Scape long, subequal to head length ( Fig. 28 ); hemelytron very narrow, width of both hemelytra in situ covering about 1/2 abdominal width ( Fig. 28A ); (cave dwelling species)............................ B. howarthi Malipatil & Liu , sp. nov.

-. Scape short, usually much shorter than head length (e.g., Fig. 1 ); hemelytron not narrow as above (e.g., Fig. 1A )......... 5

5. Fore femora long and slender, about 4X as long as maximum width ( Fig. 34 ).... B. longifemoratus Malipatil & Liu , sp. nov.

-. Fore femora short and robust, about 3X as long as maximum width (e.g., Fig. 1 )................................... 6

6. Body large, ca 20 mm or larger.......................................................................... 7

-. Body small to medium, much shorter than 18 mm ........................................................... 9

7. Legs and antennae coloured pale / yellow in contrast to dark brown to black pronotum and hemelytron ( Fig. 8 ).......................................................................................... B. ephippiger (White, 1843)

-. Legs and antennae similarly coloured dark as pronotum and hemelytron ( Figs. 26 , 46 )............................... 8

8. Male extragenital process flat and broad triangular shaped ( Figs. 55A , 56A ); tip of scutellum bluntly upturned as seen in lateral view ( Fig. 46C ); body usually jet black, particularly pronotum, abdomen and legs smooth ( Fig. 46 )................................................................................................. B. sepulchralis ( Distant, 1902)

-. Male extragenital process long, flat spine-like ( Figs. 55D , 56D ); tip of scutellum bluntly projected backwards as seen in lateral view; body dark brown, pronotum, abdomen and legs not smooth, distinctly covered with hairs of varying lengths ( Fig. 26 )...................................................................................... B. helluo Stål, 1867

9. Legs yellowish like connexivum ( Fig. 6 )............................................ B. crassifemur ( Reuter, 1881)

-. Legs brown, dark brown or black like connexivum (e.g., Fig. 1 )............................................... 10

10. Body <10 mm; pronotum black and shiny................................................................ 11

-. Body much more than 10 mm; pronotum dark brown to black and not shiny...................................... 12

11. Venter of abdomen mostly black or dark brown ( Fig. 44B )................................... B. punctorius Stål, 1867

-. Venter of abdomen orangish brown except sternite VII and posterior end dark brown ( Fig. 3B ).............................................................................................. B. ayyammae Malipatil & Liu , sp. nov.

12. Body from above almost uniformly black or brownish black, pale area involving clavus and corium indistinct or almost absent ( Figs. 14A , 15A ); (southern Australian distribution).................................. B. fuliginosus ( Erichson, 1842)

-. Body from above brown or dark brown, pale area involving clavus and corium large and conspicuous (e.g., Fig. 1A )..... 13

13. Membrane with a vague yellow patch near confluence of veins Cu and M ( Fig. 38A )...................................................................................................... B. maculatus Malipatil & Liu , sp. nov.

-. Membrane without vague yellow patch near confluence of veins Cu and M (e.g., Fig. 1A )........................... 14

14. Posterior pronotal lobe narrow, less than 1.5X as wide as anterior lobe width ( Fig. 1A )......... B. alutaceus ( Reuter, 1881)

-. Posterior pronotal lobe broad, about or more than 1.5X as wide as anterior lobe width (e.g., Fig. 24A )................. 15

15. Legs dark brown except apex of fore femur and basal portion of fore tibia yellowish brown ( Fig. 24 ); male extragenital process well developed, in the form of a minute tooth ( Figs. 55L , 56L )................. B. gunbalanyae Malipatil & Liu , sp. nov.

-. Legs generally unicolor ( Figs. 36 , 53 ); male extragenital process absent ( Fig. 36C ), or rarely present as a greatly reduced remnant process ( Figs. 55M–P , 56M–P ); (females brachypterous).............................................. 16

16. Eyes large, eye width subequal to interocular space ( Fig. 36A ); widely distributed except in Western Australia ................................................................................................. B. lurco Stål, 1867

-. Eyes small, eye width about 2/3 interocular space ( Fig. 53A ); distributed only in Western Australia ........................................................................................ B. westraliensis Malipatil & Liu , sp. nov.

17. Legs dark or black coloured contrasting with pale (yellow/ white) corium (e.g., Fig. 11 )............................ 18

-. Legs pale coloured like most of corium (e.g., Fig. 21 )....................................................... 19

18. Abdominal venter generally pale, except sternite VII dark ( Fig. 11B ).................... B. erythromelas ( Walker, 1873)

-. Abdominal venter uniformly dark brown or black like legs ( Figs. 17B , 18B )................ B. fulvipennis ( Walker, 1873)

19. Body size 12–14 mm; fore and mid femora ventrally with two or three conspicuous forwardly directed spinelike teeth in addition to minute tubercles ( Fig. 32B&C )............................................. B. limbatus ( Reuter, 1881)

-. Body size <10 mm; fore and mid femora ventrally with only minute tubercles ( Figs. 21 , 40 )........................ 20

20. Posterior pronotal lobe integument densely and finely pitted or punctured ( Fig. 40A ); hemelytron with apical 2/3 of clavus and adjoining area of corium between veins PCu+1A and Cu yellowish white ( Fig. 40A )........................................................................................................ B. pallidus Malipatil & Liu , sp. nov.

-. Posterior pronotal lobe integument not densely and finely pitted or punctured ( Fig. 21A ); hemelytron with apical 2/3 of clavus and adjoining area of corium between veins PCu+1A and Cu yellow ( Fig. 21A )........ B. fulvipes Malipatil & Liu , sp. nov.

21. Body including wing pads uniformly dark / black, wing pads with posterior margin truncate ( Fig. 14D )............................................................................................. B. fuliginosus ( Erichson, 1842)

-. Body including wing pads dark and pale, wing pads with posterior margin gradually rounded (e.g., Figs. 19A&B )....... 22

22. Legs and connexivum similarly uniformly coloured orange-yellow ( Fig. 31 ).................. B. limbatus ( Reuter, 1881)

-. Legs and connexivum coloured otherwise (e.g., Fig. 4 )...................................................... 23

23. Legs contrastingly coloured lighter than abdomen ( Fig. 22 ); (wing pads with basal 1/3 dark brown, apical 2/3 yellow with lateral areas yellowish brown)............................................... B. fulvipes Malipatil & Liu , sp. nov.

-. Legs similarly coloured dark or black like most of abdomen (e.g., Fig. 4 )........................................ 24

24. Body particularly pronotum and legs shiny smooth black (e.g., Fig. 4 ); body <10 mm .............................. 25

-. Body particularly pronotum and legs dark brown to black, not shiny and smooth (e.g., Fig. 19A&B ); body> 10 mm ...... 26

25. Venter of abdomen uniformly black or dark brown ( Fig. 43C&F )............................. B. punctorius Stål, 1867

-. Venter of abdomen with connexivum and medial areas of sternites with orangish brown or pale markings ( Fig. 4B ).............................................................................. B. ayyammae Malipatil & Liu , sp. nov.

26. Abdominal tergites VI and VII densely covered with golden setae in addition to dark setae but not on other tergites (better seen in posterior slanting view of specimen) (e.g., Fig. 48 )........................................ B. setosus ( Stål, 1874)

-. Abdominal tergites VI and VII not densely covered with golden setae but only dark setae like other tergites ( Fig. 19A&B ). 27

27. Wing pad almost entirely yellow except dark extreme basal area ( Fig. 19A&B ).............. B. fulvipennis ( Walker, 1873)

-. Wing pad basal 1/2–2/3 including narrow costal band on corium black or dark, other area pale coloured................ 28

28. Distribution in Western Australia only; male extragenital process absent or occasionally present as a greatly reduced structure ( Figs. 55M–P , 56M–P )................................................. B. westraliensis Malipatil & Liu , sp. nov.

-. Distribution in Australia except Western Australia; male extragenital process absent.................. B. lurco Stål, 1867

Figures

FIGURE 1. Brachysandalus alutaceus (Reuter, 1881), lectotype, macropterous female, habitus.A, dorsal view; B, ventral view; C, lateral view. Scale bar = 3.00 mm.

FIGURE 2. Brachysandalus alutaceus (Reuter, 1881), male genitalia.A–C, pygophore; D, left paramere; E, right paramere; F– I, phallus.A, G, ventral view; B, caudal view; C, H, I, lateral view; D, E, outer ventrolateral view; F, dorsal view. Abbreviations: bp, basal plate; bpb, basal plate bridge; dps, dorsal phallothecal sclerite; end, endosoma; lps, lateral phallothecal sclerite; mpp, median pygophore process; ped, pedicel; st, struts. Scale bar = 0.50 mm (for A–C); 0.40 mm (for D–I).

FIGURE 3. Brachysandalus ayyammae Malipatil & Liu, sp. nov., holotype, macropterous male, habitus. A, dorsal view; B, ventral view; C, lateral view. Red arrow indicates median carina on abdomen, blue arrow indicates extragenital process. Scale bar = 2.00 mm.

FIGURE 4. Brachysandalus ayyammae Malipatil & Liu, sp. nov., paratype, brachypterous female, habitus. A, dorsal view; B, ventral view; C, lateral view. Scale bar = 2.00 mm.

FIGURE 6. Brachysandalus crassifemur (Reuter, 1881), lectotype, macropterous female, habitus. A, dorsal view; B, ventral view; C, lateral view. Red arrow indicates median carina on abdomen. Scale bar = 3.00 mm.

FIGURE 8. Brachysandalus ephippiger (White, 1843), lectotype, macropterous male, habitus. A, dorsal view; B, ventral view; C, lateral view. Red arrow indicates median carina on abdomen, blue arrow indicates extragenital process. Scale bar = 3.00 mm.

FIGURE 10. Brachysandalus erythromelas (Walker, 1873), holotype, macropterous male, habitus, dorsal view.

FIGURE 11. Brachysandalus erythromelas (Walker, 1873), lectotype of Pirates (Brachysandalus) brevicoxis Stål, 1874, macropterous female, habitus. A, dorsal view; B, ventral view; C, lateral view. Scale bar = 3.00 mm.

FIGURE 12. Brachysandalus flavidus Malipatil & Liu, sp. nov., holotype, macropterous male, habitus. A, dorsal view; B, ventral view; C, lateral view. Red arrow indicates median carina on abdomen, blue arrow indicates extragenital process. Scale bar = 3.00 mm.

FIGURE 14. Brachysandalus fuliginosus (Erichson, 1842), habitus. A–C, lectotype, macropterous male; D–F, paralectotype, brachypterous female. A, D, dorsal view; B, E, ventral view; C, F, lateral view. Red arrow indicates median carina on abdomen.

FIGURE 15. Brachysandalus fuliginosus (Erichson, 1842), lectotype of Pirates (Brachysandalus) melanolestoides Stål, 1874, macropterous male, habitus. A, dorsal view; B, ventral view; C, lateral view. Red arrow indicates median carina on abdomen. Scale bar = 3.00 mm.

FIGURE 17. Brachysandalus fulvipennis (Walker, 1873), lectotype, macropterous male, habitus. A, dorsal view; B, ventral view; C, lateral view. Scale bar = 3.00 mm.

FIGURE 18. Brachysandalus fulvipennis (Walker, 1873), lectotype of Pirates (Brachysandalus) flavo-pictus Stål, 1874, macropterous male, habitus. A, dorsal view; B, ventral view; C, lateral view. Red arrow indicates median carina on abdomen. Scale bar = 3.00 mm.

FIGURE 19. Brachysandalus fulvipennis (Walker, 1873), habitus, dorsal view.A, brachypterous male; B, brachypterous female; C, macropterous male. Scale bar = 3.00 mm.

FIGURE 21. Brachysandalus fulvipes Malipatil & Liu, sp. nov., holotype, macropterous female, habitus. A, dorsal view; B, lateral view; C, ventral view. Scale bar = 2.00 mm.

FIGURE 22. Brachysandalus fulvipes Malipatil & Liu, sp. nov., paratype, brachypterous male, habitus. A, dorsal view; B, ventral view; C, lateral view. Red arrow indicates median carina on abdomen, blue arrow indicates extragenital process. Scale bar = 2.00 mm.

FIGURE 24. Brachysandalus gunbalanyae Malipatil & Liu, sp. nov., holotype, macropterous male, habitus. A, dorsal view; B, lateral view; C, ventral view. Red arrow indicates median carina on abdomen, blue arrow indicates extragenital process. Scale bar = 2.00 mm.

FIGURE 26. Brachysandalus helluo Stål, 1866, lectotype, macropterous female, habitus. A, dorsal view; B, lateral view; C, ventral view. Scale bar = 5.00 mm.

FIGURE 28. Brachysandalus howarthi Malipatil & Liu, sp. nov., holotype, submacropterous male, habitus. A, dorsal view; B, ventral view; C, lateral view. Red arrow indicates median carina on abdomen, blue arrow indicates extragenital process. Scale bar = 4.00 mm.

FIGURE 31. Brachysandalus limbatus (Reuter, 1881), lectotype, brachypterous female, habitus. A, dorsal view; B, ventral view; C, lateral view. Red arrow indicates median carina on abdomen. Scale bar = 2.00 mm.

FIGURE 32. Brachysandalus limbatus (Reuter, 1881), macropterous male, habitus. A, dorsal view; B, ventral view; C, lateral view. Red arrow indicates median carina on abdomen. Scale bar = 3.00 mm.

FIGURE 34. Brachysandalus longifemoratus Malipatil & Liu, sp. nov., holotype, macropterous male, habitus. A, dorsal view; B, lateral view; C, ventral view. Red arrow indicates median carina on abdomen. Scale bar = 3.00 mm.

FIGURE 36. Brachysandalus lurco Stål, 1866, lectotype, macropterous male, habitus. A, dorsal view; B, lateral view; C, ventral view. Red arrow indicates median carina on abdomen. Scale bar = 4.00 mm.

FIGURE 38. Brachysandalus maculatus Malipatil & Liu, sp. nov., holotype, macropterous male, habitus. A, dorsal view; B, ventral view; C, lateral view. Red arrow indicates median carina on abdomen, blue arrow indicates extragenital process, yellow arrow indicates vague yellowish patch on membrane near confluence of veins Cu and M. Scale bar = 3.00 mm.

FIGURE 40. Brachysandalus pallidus Malipatil & Liu, sp. nov., holotype, macropterous male. A, habitus, dorsal view; B, habitus, lateral view; C, habitus, ventral view; D, posterior part of abdomen, ventral view; E, close-up of extragenital process. Red arrow indicates median carina on abdomen, blue arrow indicates extragenital process.

FIGURE 43. Brachysandalus punctorius Stål, 1866, habitus.A–C, brachypterous male; D–F, brachypterous female.A, D, dorsal view; B, E, lateral view; C, F, ventral view. Red arrow indicates median carina on abdomen, blue arrow indicates extragenital process. Scale bar = 2.00 mm.

FIGURE 44. Brachysandalus punctorius Stål, 1866, macropterous male, habitus. A, dorsal view; B, lateral view. Scale bar = 2.00 mm.

FIGURE 46. Brachysandalus sepulchralis (Distant, 1902), lectotype, macropterous female, habitus. A, dorsal view; B, ventral view; C, lateral view. Scale bar = 5.00 mm.

FIGURE 48. Brachysandalus setosus (Stål, 1874), live brachypterous male, Brisbane, Queensland, September 2022 (Photo: Campbell Paine).

FIGURE 53. Brachysandalus westraliensis Malipatil & Liu, sp. nov., holotype, macropterous male, habitus.A, dorsal view; B, ventral view; C, lateral view. Red arrow indicates median carina on abdomen. Scale bar = 2.00 mm.

FIGURE 55. Close-up photos of extragenital process of Australian Brachysandalus species.A, B. sepulchralis (Distant, 1902); B, B. howarthi Malipatil & Liu, sp. nov.; C, B. alutaceus (Reuter, 1881); D, B. helluo Stål, 1866; E, B. ephippiger (White, 1843); F, B. punctorius Stål, 1866; G, B. flavidus Malipatil & Liu, sp. nov.; H, B. crassifemur (Reuter, 1881); I, B. maculatus Malipatil & Liu, sp. nov.; J. B. ayyammae Malipatil & Liu, sp. nov.; K, B. fulvipes Malipatil & Liu, sp. nov.; L, B. gunbalanyae Malipatil & Liu, sp. nov.; M–P, B. westraliensis Malipatil & Liu, sp. nov. Blue arrow indicates extragenital process. Scale bar = 0.50 mm.

FIGURE 56. Illustrations of extragenital process of Australian Brachysandalus species. A, B. sepulchralis (Distant, 1902); B, B. howarthi Malipatil & Liu, sp. nov.; C, B. alutaceus (Reuter, 1881); D, B. helluo Stål, 1866; E, B. ephippiger (White, 1843); F, B. punctorius Stål, 1866; G, B. flavidus Malipatil & Liu, sp. nov.; H, B. crassifemur (Reuter, 1881); I, B. maculatus Malipatil & Liu, sp. nov.; J. B. ayyammae Malipatil & Liu, sp. nov.; K, B. fulvipes Malipatil & Liu, sp. nov.; L, B. gunbalanyae Malipatil & Liu, sp. nov.; M–P, B. westraliensis Malipatil & Liu, sp. nov. Scale bar = 0.50 mm.