Pseudocetherinae (Hemiptera: Reduviidae) revisited: phylogeny and taxonomy of the lobe-headed bugs Author Castillo, Stephanie 802965D7-06BE-4C93-B2B4-1CF8F2C8C2DC University of California, Riverside, 900 University Avenue, Riverside, CA 92521, USA. scast032@ucr.edu Author Rédei, Dávid 74DB4DB4-E632-4CA9-B820-2070E7EB18D4 National Chung Hsing University, 145 Xingda Rd., 40227 Taichung, Taiwan. david.redei@gmail.com Author Weirauch, Christiane 59B6A54F-2044-4E5A-B7F7-AAC5E7D716F0 University of California, Riverside, 900 University Avenue, Riverside, CA 92521, USA. christiane.weirauch@ucr.edu text European Journal of Taxonomy 2022 2022-01-11 788 1 1 95 http://dx.doi.org/10.5852/ejt.2022.788.1625 journal article 2598 10.5852/ejt.2022.788.1625 d6e6119f-4100-4cfc-9cc3-0d6951ee2c28 2118-9773 5846636 46C3CFCA-0CED-4432-AFD8-F4CFC1E0E1E7 Genus Voconia Stål, 1866 Figs 1–4 , 6–14 , 17–20 Voconia Stål, 1866a: 120 . Type species: Voconia pallidipes Stål, 1866 , by subsequent monotypy ( Stål 1866b: 165 ). Gender feminine. Microvarus Jeannel, 1917: 50 . Type species: Microvarus conradti Jeannel, 1917 , by original designation. syn. nov. Kayanocoris Miller, 1954: 5 . Type species: Kayanocoris wegneri Miller, 1954 , by original designation. syn. nov. Paragerbelius Miller, 1958: 66 . Type species: Paragerbelius typicus Miller, 1958 , by original designation. syn. nov. Pseudocethera Villiers, 1963b: 532 . Type species: Pseudocethera monodi Villiers, 1963 , by original designation. syn. nov. Diagnosis Distinguished from the other genus in the subfamily, Gerbelius , by the granulose integument of the head and pronotum ( Fig. 6A, D ), ellipsoid shape of the maxillary plates ( Fig. 6B , blue), short non-adjacent mandibular plates ( Fig. 6B , green), gracile labium with short macrosetae on the morphologically ventral surface of labial segment I ( Fig. 6E ), and the Cu-An 1 cell of the fore wing shorter than the M-Cu cell and triangular ( Fig. 6C ). Redescription BODY LENGTH. 6.7–12.4 mm . BODY. Slender or robust; macropterous or micropterous ( Figs 7–10 ). Fig. 6. Diagnostic characters of Voconia Stål, 1866 shown on Voconia dolichocephala sp. nov. (AMNH_PBI 00168745). A–B . Head, dorsal view; maxillary plates = blue, mandibular plates = green. C . Hemelytra, Cu-An 1 cell = red. D . Head and thorax, lateral view; anteriad directed process = purple; E . Labium. F . Pygophore, dorsal view. G . Paramere. COLORATION.Thorax:scutellum uniform or with contrasting yellow scutellar spine.Hemelytron(abducted): corium with various combinations of yellow spots on the proximal, anteromedial, posteromedial, and distal areas, with medial yellow stripe, or uniformly yellow; membrane uniformly dark or with pale V-shaped marking along R and M veins. INTEGUMENT AND VESTITURE. Head and pronotum: granulose with pubescence and interspersed macrosetae or setigerous tubercles; interocular region with two glabrous lines; antennifer with macroseta, with or without lateral protuberance; morphologically ventral surface of labial segment I with short macrosetae or sparse long macrosetae. Thorax: anterolateral angles of pronotal collar armed with macrosetae or setigerous tubercles. Legs: posterior row of spine-like protuberances on all femora well developed, anterior row poorly developed on ventral surface of mid and hind femora.Abdomen: ventral surface pubescent with long macrosetae. STRUCTURE. Head: globose to elongate; mandibular plates in dorsal view short, not adjacent basally; maxillary plates in dorsal view elongate anteriorly, forming ellipsoid projections; pedicel 0.6–1.9 times length of head width; ventrolateral swelling of buccula with or without lateral protrusion that surpasses buccular margin; labium gracile; morphologically dorsal surface of labial segment II 0.2–0.7 times length of segment I; labial segment III drawn into bent, slender apex. Thorax: scutellum width 0.5–0.8 times length; scutellar spine subhorizontal or raised; proepimeron without or with acute protuberance on posteroventral margin. Hemelytron: Cu-An 1 cell triangular, smaller than M-Cu cell. Legs: fossula spongiosa present on fore and mid legs in males, either present on fore and mid legs or absent on mid legs of females. Distribution This genus currently comprises nine described species and 23 new species with a circumtropical distribution. The lowest known species diversity is in the Neotropics, and the highest species diversity for this genus is in Southeast Asia across India , Indonesia , Laos , Brunei , and the Philippines . Remarks Originally a monotypic genus. It is here expanded to include numerous new species as well as several previously described species formerly placed in Reduviinae. Key to the species of Voconia 1. Corium of macropterous morph entirely yellow [ Kenya ] ................... Voconia chrysoptera sp. nov. – Corium of macropterous morph with various brown and pale patterns (e.g., Fig. 4B–C ) or micropterous ( Fig. 4D ) ..................................................................................................................... 2 2. Micropterus; maxillary plates in lateral view about twice the width of scape [Afrotropical] .......... 3 – Macropterous; maxillary plates in lateral view cylindrical, about the width of scape [Circumtropical] ............................................................................................................................... 5 3. Maxillary plates toothed at apex; legs yellow [West Africa] .............................................................. .......................................................................................... Voconia ifana (Villiers, 1963) comb. nov. – Maxillary plates rounded at apex; legs yellow with distal half of femora brown ............................ 4 4. Paired longitudinal carinae on abdominal tergum II not prominent, not reaching posterior margin of segment; ocelli indistinct [West Africa] ........................ Voconia monodi (Villiers, 1963) comb. nov. – Paired longitudinal carinae on abdominal tergum II prominent, reaching posterior margin of segment ( Fig. 4D ); ocelli distinct [Central Africa] ............. Voconia schoutedeni ( Villiers, 1964 ) comb. nov. 5. Head and pronotum coarsely granulose, with setigerous tubercles ( Fig. 3C–D ) ............................. 6 – Head and pronotum finely granulose, without large setigerous tubercles ( Fig. 3B, H ) .................. 15 6. Antennifer with large spine-like protuberance ( Fig. 3D ) [Afrotropical] .......................................... 7 – Antennifer with setigerous tubercle ( Fig. 3C ) [New World; Australasia] ........................................ 8 7. Head and pronotum almost entirely dark brown; scutellar spine slightly raised; hemelytra blackish with pale coloration at base and apex of corium [Central Africa] ...................................................... ............................................................................... Voconia schoutedeni ( Villiers, 1964 ) comb. nov. – Head and pronotum yellow; scutellar spine subhorizontal; hemelytra light brown with proximal half of corium yellow and distal yellow spot [ Sudan ] ........................................ Voconia tridens sp. nov. 8. Clypeus round [ Mexico ; Honduras ] ........................................................ Voconia mexicana sp. nov. – Clypeus bifid ( Fig. 3C ) [ Australia ; Papua New Guinea ] .................................................................. 9 9. Hemelytron with yellow spots on corium (e.g., Fig. 4B–C ) ........................................................... 10 – Hemelytron with a broad yellow band across corium (e.g., Fig. 7 , V. fasciata sp. nov. and V. grandioculata sp. nov. ) ............................................................................................................... 12 10. Head globose, about as long as wide [ Northern Territory , Australia ; Papua New Guinea ] ................ .......................................................................................................... Voconia brachycephala sp. nov. – Head elongate, 1.4–1.6 times as long as wide .................................................................................11 11. Body dark, almost black; males small, body length 7–8 mm [ Northern Territory , Australia ] ............ ......................................................................................................... Voconia dolichocephala sp. nov. – Body light brown; males larger, body length 8–9 mm [ Northern Territory and Queensland , Australia ] .............................................................................................. Voconia tuberculata sp. nov. 12. Body longer, 9–11 mm ; legs entirely yellow .................................................................................. 13 – Body shorter, 7–9 mm ; femora dark brown, tibiae and tarsi yellowish-brown .............................. 14 13. Eyes as long as ante- and postocular regions in dorsal view; ocelli occupying half-length of postocular region; membranal veins forming Cu-An 1 and M-Cu cells brown; proximal half of corium not entirely yellow; body length 9–10 mm [ Western Australia and Queensland , Australia ] .................... ........................................................................................................... Voconia grandioculata sp. nov. – Eyes shorter than ante- and postocular regions in dorsal view; ocelli occupying less than halflength of postocular region; membranal veins forming Cu-An 1 and M-Cu cells mostly or entirely yellow; proximal half of corium entirely yellow; body length 10.5–12 mm [ New South Wales and Queensland , Australia ] ......................................................................... Voconia pallidipes Stål, 1866 14. Corium band is translucent yellow; body length 8–9 mm [ Queensland , Australia ]............................ ..................................................................................................................... Voconia fasciata sp. nov. – Corium band is opaque yellow; body length approximately 7 mm [ Queensland , Australia ] ............. ........................................................................................................................ Voconia vittata sp. nov. 15. Labial segment I swollen subapically on ventral surface ( Fig. 4E ); labial segment II swollen ventrobasally and narrowed apically ( Fig. 4E ); labial segment III stout, 0.2 times length of segment II ( Fig. 4E ) [Borneo] ...................................................... Voconia wegneri ( Miller, 1954 ) comb. nov. – Labial segments I and II straight; labial segment III elongate, 0.4–0.7 times length of segment II ... ......................................................................................................................................................... 16 16. Head long, about 1.7 times as long as wide; ventral surface of hind femur with four large spines in posterior row ( Fig. 4K ); large pale spot between R and M veins ( Fig. 4C ) ................................... 17 – Head elongate, not more than 1.4 times as long as wide; ventral surface of hind femur with three large and 0–3 small spines in posterior row ( Fig. 4I–J ) .................................................................. 18 17. Mid and hind femora dark, proximal third yellow ( Fig. 4K ); anterior pronotal lobe convex, strongly curved inward on posterolateral margins; body length about 11 mm [Normanby Island] ................. ..................................................................................................................... Voconia bracata sp. nov. – Legs uniformly dark; anterior pronotal lobe gently rounded, posterolateral margins nearly straight; body length 12–13 mm [New Guinea ] ............................. Voconia typica ( Miller, 1958 ) comb. nov. 18. Pronotum dark with contrasting yellow posterior margin; abducted corium dark with antero- and posteroproximal yellow stripes and posteromedial and distal white spots [ Trinidad ] ........................ ............................................................................................................ Voconia trinidadensis sp. nov. – Color patterns variable but combination not as above [Old World] ............................................... 19 19. Proepimeron with posteroventral margin rounded [Afrotropical] .................................................. 20 – Proepimeron with acute protuberance on posteroventral margin [ Oriental ] .................................. 22 20. Fossula spongiosa absent or vestigial on mid leg of males and females; scutellum uniformly brown; abducted corium with an antero- and posteromedial yellow spot; uniformly brown dorsal laterotergites [Central Africa] ............................ Voconia motoensis ( Schouteden, 1929 ) comb. nov. – Fossula spongiosa present on mid leg of male; scutellar spine yellow ( Fig. 4E ); abducted corium with proximal, posteromedial, and distal yellow spots; dorsal laterotergites bicolored ................. 21 21. Body longer, 10–12 mm ; dark dorsal laterotergites with strongly contrasting yellow anterior spots; eyes reaching ventral head margin in lateral view; eyes strongly bulging past postocular lateral margins in dorsal view [West and Central Africa] ...... Voconia conradti ( Jeannel, 1917 ) comb. nov. – Body shorter, 9–10 mm ; light brown dorsal laterotergites with weakly contrasting yellow anterior spots; eyes do not reach ventral head margin in lateral view; eyes do not bulge strongly in dorsal view [Central Africa] .................................................................................. Voconia smithae sp. nov. 22. Membrane of hemelytra dark; membranal veins forming Cu-An 1 and M-Cu cells paler than remainder of membrane ( Fig. 4B ) ................................................................................................................... 23 – Membrane of hemelytra dark; membranal veins dark or with R and M veins proximally pale in V-shape ........................................................................................................................................... 25 23. Body light brown; labium lighter than head, yellow; scutellum dark with contrasting yellow apical spine [Borneo] ........................................................................................... Voconia decorata sp. nov. – Body dark brown, almost black; labium as dark as head; scutellum uniformly dark [ Sumatra ]..... 24 24. Head dark with semicircular yellow stripe encircling both ocelli; body longer, about 11 mm ; anterior pronotal lobe relatively long, posterior lobe about 1.8 times as long as than anterior lobe ............... ................................................................................................................... Voconia coronata sp. nov. – Head dark with yellow spots adjacent to lateral ocellar margin; body shorter, about 9 mm ; anterior pronotal lobe relatively short, posterior lobe about 2.2 times as long as anterior lobe ...................... ................................................................................................................ Voconia lirophleps sp. nov. 25. Legs yellow, distal halves of femora brown [ India ] ................................ Voconia ornata comb. nov. – Femora unicolored [Southeast Asia] ............................................................................................... 26 26. Legs dark brown ............................................................................................................................. 27 – Legs yellow or yellowish-brown .................................................................................................... 29 27. Dorsal laterotergites dark with yellow spots anteriorly; anterior margin of abducted corium with a small pale spot medially; anteriad-directed process of prosternum with small paramedial lobes; stridulitrum elongated anteriorly into a small protuberance [ Philippines ] ......................................... ....................................................................................................................... Voconia bakeri sp. nov. – Dorsal laterotergites uniformly dark; anterior margin of abducted corium with a yellow stripe along proximal half; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes ( Fig. 4J ); stridulitrum not elongated into an anterior protuberance ( Fig. 4J ) [Southeast Asia] ...... 28 28. Body length about 10.0 mm; pronotum dark, posterior margin paler [ Philippines ] ........................... ........................................................................................................................... Voconia nyx sp. nov. – Body length about 8.5 mm ; pronotum uniformly dark [ Philippines ] ......... Voconia minima sp. nov. 29. Anterior margin of abducted corium with medial and distal yellow spots; pronotum and scutellum uniformly dark [ Philippines ] ....................................................................... Voconia hemera sp. nov. – Anterior margin of abducted corium with proximal half yellow and distal yellow spot; pronotum dark, posterior margin pale; scutellum dark with contrasting brown or yellow apical spine ( Fig. 4E ) ... 30 30. Clypeus coloration as rest of head; proximal half of corium entirely yellow, forming a yellow triangle [ Philippines ] .............................................................................................. Voconia isosceles sp. nov. – Clypeus lighter than rest of head, yellow; proximal half of corium not entirely yellow, with brown markings ......................................................................................................................................... 31 31. Labial segment I short, not surpassing posteroventral eye margin in lateral view; Cu-An 1 cell elongate, reaching at least half the length of M-Cu cell [ Laos ] ............... Voconia laosensis sp. nov. – Labial segment I long, surpassing posteroventral eye margin in lateral view; Cu-An 1 cell stout, less than half the length of M-Cu cell .................................................................................................... 32 32. Abducted corium mostly dark with anteroproximal reddish-brown stripe and posteromedial and distal yellow spots; pale posterior margin of pronotum wide, a third of posterior pronotal lobe [Borneo] ................................................................................................. Voconia lasiosoma sp. nov. – Abducted corium with proximal half mostly yellow, distal half dark with distal yellow spot; pale posterior margin of pronotum narrow, a fifth of posterior pronotal lobe [Borneo] ............................ ........................................................................................................................... Voconia loki sp. nov.