Pseudocetherinae (Hemiptera: Reduviidae) revisited: phylogeny and taxonomy of the lobe-headed bugs

The concept of the previously monogeneric subfamily Pseudocetherinae (Hemiptera: Heteroptera: Reduviidae) is revised and expanded. We here transfer Gerbelius Distant, 1903, Kayanocoris Miller, 1954, Microvarus Jeannel, 1917, Paragerbelius Miller, 1958, and Voconia Stål, 1866 from Reduviinae to Pseudocetherinae and treat Kayanocoris, Microvarus, Paragerbelius, and Pseudocethera Villiers, 1963 as junior synonyms of Voconia, resulting in new combinations for Voconia conradti (Jeannel, 1917) comb. nov., V. ifana (Villiers, 1963) comb. nov., V. monodi (Villiers, 1963) comb. nov., V. motoensis (Schouteden, 1929) comb. nov., V. ornata (Distant, 1903) comb. nov., V. schoutedeni (Villiers, 1964) comb. nov., V. typica (Miller, 1958) comb. nov., and V. wegneri (Miller, 1954) comb. nov. We also describe 23 new species of Voconia: V. bakeri sp. nov., V. bracata sp. nov., V. brachycephala sp. nov., V. chrysoptera sp. nov., V. coronata sp. nov., V. decorata sp. nov., V. dolichocephala sp. nov., V. fasciata sp. nov., V. grandioculata sp. nov., V. hemera sp. nov., V. isosceles sp. nov., V. laosensis sp. nov., V. lasiosoma sp. nov., V. lirophleps sp. nov., V. loki sp. nov., V. mexicana sp. nov., V. minima sp. nov., V. nyx sp. nov., V. smithae sp. nov., V. tridens sp. nov., V. trinidadensis sp. nov., V. tuberculata sp. nov., and V. vittata sp. nov. Lectotypes are designated for Gerbelius confluens Distant, 1903, G. typicus Distant, 1903, V. conradti comb. nov., V. ornata comb. nov., and V. pallidipes Stål, 1866. A revised diagnosis and description of Pseudocetherinae are provided along with photographs of the species and of the male genitalia of 13 pseudocetherine and five closely related reduviine species. An identification key to the two genera of Pseudocetherinae as well as a key to species of Voconia are presented. A phylogenetic hypothesis is proposed for the relationships of Pseudocetherinae using parsimony analyses of 77 morphological characters.


Imaging, morphological methods, and measurements
Dorsal and lateral habitus photographs, as well as close-ups of the head and male genitalia were taken on a Leica DFC 450 C Microsystems camera (Leica, Wetzlar, Germany) using a Planapo 2.0× objective and a Leica Application Suite (LAS) ver. 4.3. Stacked images were assembled using LAS. Male genitalia were excised and macerated in heated 10% potassium hydroxide solution for 5-10 minutes and rinsed in distilled water. They were temporarily stored in a 96-well plate containing 100% glycerol for examination and eventually sealed in polyethylene vials containing glycerol and attached to the specimen's pin. The pygophore, eighth abdominal segment, one paramere, and the phallus were removed for examination in glycerol gelatin for imaging for Gerbelius sp., G. typicus, V. bakeri sp. nov., V. conradti, V. grandioculata sp. nov., V. lasiosoma sp. nov., V. laosensis sp. nov., V. mexicana sp. nov., V. nyx sp. nov., V. pallidipes Stål, 1866, V. tridens sp. nov., V. tuberculata sp. nov., V. typica (Miller, 1958) and V. vittata sp. nov. Scanning electron micrographs (SEMs) were acquired on a Hitachi TM-1000 Tabletop Electron Microscope (Institute for Integrative Genome Biology, UCR) for one male V. dolichocephala sp. nov. (AMNH_PBI 00168745) to document details of the head, prothorax, labium, hemelytron, and pygophore. The specimen was not removed from the point or coated making this approach maximally non-destructive.
Measurements of selected morphological features were obtained from photographs using ImageJ2 (Rueden et al. 2017) and are provided in Table 1. Body length is recorded in dorsal view along the midline from the clypeal apex to the posteromedial margin of the abdomen. Dorsal head measurements were obtained from images of the head oriented horizontally for better accuracy. This allowed us to record head length from the maxillary plate apex to the posterior margin of the head (defined by the transition of head granulations to smooth neck) since the maxillary plates in most species are slightly longer than the clypeal apex. Head length recorded from the clypeal apex to the posterior margin of the head is recorded in Supp. file 1. The scape, pedicel, and labium were measured from the proximal and distal midpoints.

Terminology
Terminology generally follows Schuh & Weirauch (2020). When referring to the hemelytra we refer to it in an abducted position, meaning away from the midline of the body or perpendicular to the longitudinal axis of the body. We refer to a seta on a protuberance as a setigerous tubercle, large setae as macrosetae, and short, thin macrosetae as pubescence. Abbreviations for wing cells and veins in descriptions and the cladistic analysis are as follows: Cu-An 1 cell, cell between cubital and first anal veins; M-Cu cell, cell between medial and cubital veins; M, medial vein; R, radial vein. We refer to labial segments as: labial segment I, first visible segment; labial segment II, second visible segment; labial segment III, third visible segment. In reduviids the first abdominal tergum is referred to as syntergum I+II. Here, we refer to syntergum I+II individually as tergum I and tergum II since they are clearly identifiable in Pseudocetherinae.

Species descriptions and redescriptions
Descriptions are based on examination of available specimens for a given species, including type material, type images of Voconia conradti, V. pallidipes, and V. wegneri, and the original descriptions of V. ifana and V. monodi. Descriptions were compiled from a character matrix of 98 discrete characters assembled in Mesquite ver. 3.51 (Maddison & Maddison 2019). This descriptive character matrix was exported as descriptions for each taxon in Mesquite and then punctuation and sentence structure were manually edited. Species descriptions are provided for all species of Voconia, including formerly recognized G. ornatus, as well as species placed in the genera Kayanocoris, Microvarus, Paragerbelius, and Pseudocethera but transferred to Voconia in this study. Because the habitus of species of Gerbelius resembles that of other small-bodied Reduviinae compared to Voconia, we suspect that our survey of  (2022) 8 Gerbelius specimens deposited in various natural history collections has not been comprehensive. We therefore refrain from describing new species of Gerbelius as part of the present study and only provide a redescription of the genus. Data interpreted from labels (e.g., interpretation of coordinates or foreign text) in the material citations are distinguished with square brackets and verbatim citations (e.g., archaic geographic names) are encased in double quotation marks.

Cladistic analysis
Forty taxa comprising 33 ingroup species and seven outgroup species were analyzed (Figs 1-2). Outgroup selection was informed by the molecular phylogenies presented in Hwang & Weirauch (2012) and included two Reduvius clade taxa (Reduvius personatus (Linnaeus, 1758)  The descriptive matrix was used as the framework for this analysis by first removing non-informative (autapomorphic) characters. The 77 morphologically discrete characters (29 binary, 48 non-additive [unordered] multistate) used in the analysis are listed in the results. Characters are from the head (39), thorax (12), hemelytron (6), legs (6), abdomen (2), and male genitalia (12). We selected nine characters that rely on color patterns that are not highly variable within species given our specimen sampling. Inapplicable characters were treated as missing data and intraspecific variation coded as polymorphic. Gil-Santana et al. (2007) was used to code C. tomentosus, and male genitalic characters for Reduvius personatus are based on Weirauch et al. (2015). Since male genitalia across Reduviidae are not welldocumented (Forero & Weirauch 2012), the pygophore and phallus dissections of the remaining outgroup taxa are made available. Characters and character states are indicated in the image plates (e.g., 2-0 in Fig. 3 refers to character 2 and state 0 in the results). The character matrix is provided as a text file in Supp. file 2.
The equal weighting (EW) analysis was conducted in TNT ver. 1.5 (Goloboff & Catalano 2016) using New Technology Search with the following parameters: find minimum length 100 times; ratchet (Nixon 1999) with default settings and 200 iterations; tree-drifting with default settings and 200 cycles; tree fusing with default settings. Clade support was evaluated using jackknife with default settings and 200 replicates. Tree length (L), consistency index (CI), and retention index (RI) were also calculated in TNT. The trees were then imported into WinClada ver. 1.00.08 (Nixon 2002) to estimate the strict consensus tree (Fig. 1).
Implied weighting (IW) (Goloboff 1993) analyses were performed in TNT under the same New Technology Search parameters as mentioned above. Several concavity values, which downweigh homoplasious characters, were explored to compare tree length and tree topology under the different weighting schemes: K = 3, K = 6, K = 9, K = 12 (Fig. 2). The lower values penalize homoplasy more strongly, though a value of K = 12 is still relatively strong considering that a value of K = 20 weighs homoplasies "very far from mild" in certain large datasets (Goloboff et al. 2008). The matrix was resampled with symmetric resampling (Goloboff et al. 2003) under default settings and 200 replicates. The IW tree provided greater resolution for our analysis than the EW tree, with the K = 12 analysis having the best fit. We therefore used the most parsimonious tree derived from the K = 12 analysis to document and discuss character transformations. WinClada was used to illustrate unambiguous changes and treated any additional step as homoplastic (Fig. 2). To avoid local optima, we ran IW analyses with 9 several random seeds (1,10,123,1234,12345) and maintained the above-mentioned settings with a concavity of K = 9.

Cladistic analysis
The EW analysis resulted in eight most parsimonious trees (length = 468; CI = 0.353, RI = 0.625). The strict consensus tree collapsed 12 nodes (Fig. 1). The IW analyses with four different concavities recovered one parsimonious tree each, all with the same topology. Increasing the K-values, which decreases the penalty against homoplasious characters, improved the best score and resulted in higher CI and RI values: K = 3 resulted in best score = 33.53464, CI = 0.348, RI = 0.618; K = 6 resulted in best score = 24.58040, CI = 0.349, RI= 0.619; K = 9 resulted in best score = 19.55168, CI = 0.353, RI = 0.625; K = 12 resulted in best score = 16.26989, CI = 0.353, RI = 0.625. Characters and character optimizations are described European Journal of Taxonomy 788: 1-95 (2022) 10 and discussed in the section below using the most parsimonious tree derived from the K = 12 IW analysis. Exploring different random seeds for the IW analyses with K = 9 did not result in different tree statistics (best score = 19.55168; CI = 0.353; RI = 0.625) and the topology remained the same.

Fig. 2.
Tree resulting from implied weighting analysis using K = 12 with characters unambiguously optimized. Symmetric resampling values over 51 are represented for trees obtained with K = 3, K = 6, K = 9, and K = 12 analyses. Characters and character states are described in pp. 11-22. Specimens not to scale.
Reduviinae to Pseudocetherinae in the Taxonomy section below and provide a revised diagnosis for Pseudocetherinae. Gerbelius typicus and two undescribed species of Gerbelius form a well-supported clade, but G. ornatus is nested among species that would be referred to as Microvarus under the generic concepts prior to this work. Gerbelius ornatus is therefore removed from Gerbelius. Within the non-Gerbelius pseudocetherine clade, Pseudocethera and Paragerbelius are highly supported (resampling values of ≥ 99) monophyletic groups, and Australasian species that would be referred to as Voconia under the current generic concept (Fig. 1, yellow highlight) form a well-supported (resampling values of 70-75) clade. However, Microvarus, as currently diagnosed, is paraphyletic in our analyses. This situation is complicated by the fact that we are here describing 23 new species of Pseudocetherinae, most of which would be classified as Microvarus under traditional diagnoses (Fig. 1, green highlight), but we are reluctant to describe new species in a paraphyletic grade. In addition, while we recovered two small, well-supported clades among these new species (V. laosensis sp. nov. + V. lasiosoma sp. nov. from the Oriental region and V. conradti + V. smithae sp. nov. from the Afrotropical region), most relationships among non-Gerbelius Pseudocetherinae are poorly supported. We therefore decided to synonymize Microvarus, Kayanocoris, Paragerbelius, and Pseudocethera under Voconia, rather than preserving these generic concepts and describing several small or monotypic new genera that would be difficult to diagnose.
While our phylogenetic hypothesis may not be robust enough to meaningfully infer the historical biogeography of Pseudocetherinae, it is worth pointing out that with the expansion of Pseudocetherinae through the new species and taxa treated in new combinations, this subfamily is now known from the Afrotropical, Australasian, Oriental and Neotropical regions.

Description and documentation of morphological characters with comments on character optimizations (Figs 3-15)
1. Head, postocular region, coloration medially in dorsal view: uniform (0), with dark patches adjacent to ocellar margin (1), with red patches adjacent to ocellar margin (2), with dark medial stripe (3). Voconia tridens sp. nov. has red patches adjacent to the ocellar margin, while several other pseudocetherines have dark, blackish markings. Most, however, are uniformly colored.
20. Legs, mid and hind femora, posterior row of spine-like protuberances: with three large spines on distal half and three or fewer small spines ( Fig. 4J) (0), with three large spines on distal half and many small spines ( Fig. 4I) (1), with four or more large spines on distal half ( Fig. 4K) (2). The number of spines on the femur is species-specific, though Pseudocetherinae typically have three large spines on the distal half of the femur with a variable number of small spines interspersed. In several species, namely V. bracata + V. typica, V. dolichocephala + V. tuberculata, V. grandiocula, and V. pallidipes, the distal half of the femur has four or more large spines.
21. Head, shape in dorsal view: globose, about as long as wide (0), elongate, less than 1.5 times as long as wide (Fig. 3D) (1), distinctly elongate, at least 1.5 times as long as wide (2). An elongate head (less than 1.5 times as long as wide) is a homoplastic synapomorphy for Pseudocetherinae. There is intraspecific variation in the head shape of V. conradti from globose to elongate. A globose head shape is a homoplastic synapomorphy for the clade containing V. grandioculata sp. nov., V. fasciata sp. nov., V. brachycephala sp. nov., and V. vittata sp. nov. A distinctly elongate head (at least 1.5 times as long as wide) is a homoplastic synapomorphy for the clade containing V. wegneri, V. typica, and V. bracata sp. nov. as well as the clade containing V. tuberculata sp. nov. and V. dolichocephala sp. nov.
22. Head, anteocular region, length relative to head: about one quarter of head length (0), about one third of head length (1), about half of head length (2), two-fifths of head length (Fig. 3D) (3), less than a fifth of head length (4). The anteocular region is about one third the head length in most pseudocetherines but varies interspecifically. An anteocular region that is about two-fifths of the head length is a non-homoplastic synapomorphy for the clade containing V. tridens sp. nov. and V. schoutedeni.
23. Head, anteocular region, length relative to postocular region: shorter than postocular region (0), about as long as postocular region (1), longer than postocular region ( Fig. 3D) (2). Anteocular length was recorded to the apex of the clypeus. An anteocular region that is shorter than the postocular region is synapomorphic for Gerbelius, though more sampling may challenge our analyses. An anteocular region that is longer than the postocular region is synapomorphic for V. tridens sp. nov. + V. schoutedeni.
24. Head, postocular region, length relative to eye: shorter than eye (0), about as long as eye (1), longer than eye ( Fig. 3D) (0), about as wide as synthlipsis (1), narrower than synthlipsis (Fig. 3B) (2). Reconstructed as eyes narrower than synthlipsis in the most recent common ancestor of Pseudocetherinae with several transitions (Tab. S1).  42. Buccula, anterolateral swelling, shape in lateral view: flat buccular margin ( Fig. 3G) (0), with protrusion surpassing buccular margin (1). The swelling of the buccula is either flat apically or has a protruding bump that overlaps with the labium in lateral view. This protrusion may be a small, rounded bump as in V. decorata sp. nov. In V. tridens sp. nov. and V. schoutedeni this protrusion is large and spine-like with small setigerous tubercles along the margin. In V. chrysoptera sp. nov., V. fasciata sp. nov., V. grandioculata sp. nov., V. mexicana sp. nov., and V. ornata the protrusion is rounded with setigerous tubercles along the apical margin. V. pallidipes has large protruding setigerous tubercles.
46. Labium, morphologically dorsal surface of labial segment II, length relative to segment I: 0.2 times the length of segment I (0), 0.3 times the length of segment I (Fig. 3F) (1), 0.4 times the length of segment I (2), 0.5 times the length of segment I (3), 0.6 times the length of segment I (4), 0.7 times the length of segment I (5), 0.8 times the length of segment I (6), 0.9 times the length of segment I (7), longer than segment I (8). The length of segment I was recorded from the proximal and distal midpoints, while the length of segment II reported here is recorded from the morphologically dorsal surface. Measurements were rounded up for values with 0.05 and up. The morphologically ventral surface of segment II is shorter and reported in Supp. file 1. The length of the morphologically dorsal surface of segment II varies interspecifically. However, the length of segment II among members of Gerbelius ranges from 0.8-0.9 times the length of segment I and ranges between 0.2-0.7 times the length of segment I in Voconia, with V. wegneri having the largest disparity between both segments (0.2 times the length of segment I) and V. motoensis having the smallest disparity between both segments (0.7 times the length of segment I).
48. Labium, labial segment III, shape: gradually narrowing, straight (0), drawn into a bent, slender apex ( Fig. 3E-G) (1). Labial segment III is drawn into a bent, slender apex in all Pseudocetherinae, even in V. wegneri which has an unusually short labial segment III. The orientation of the scutellar spine varies interspecifically, and intraspecifically in V. mexicana sp. nov. where the female has a raised scutellar spine but the scutellar spine is subhorizontal in the males.
57. Stridulitrum, anterior margin, shape: not elongated into protuberance ( Fig. 3J) (0), projected into sharp spine (1), projected into small protuberance ( Fig. 3I) (2). The stridulitrum of most Voconia is not elongated into a protuberance. In Gerbelius, V. grandioculata sp. nov., and V. fasciata sp. nov. the stridulitrum projects anteriorly into a sharp spine. The remaining Australian species in the clade with V. chrysoptera sp. nov. and V. mexicana sp. nov., along with V. lasiosoma sp. nov. and V. bakeri sp. nov., have a stridulitrum that protrudes into a small protuberance. 63. Fore leg, fossula spongiosa (female): present (0), absent or vestigial (1). Due to the scarcity of females, there is a lot of missing data for females, but the fossula spongiosa may be present on the fore legs of all pseudocetherine females.
64. Mid leg, fossula spongiosa (female): present (0), absent or vestigial (1). Due to the scarcity of females, there is a lot of missing data for females, but it appears that the fossula spongiosa is absent on the mid legs of pseudocetherine females, except in V. chrysoptera sp. nov. where it is present.

71.
Posterior pygophore margin, position of macrosetae in caudal view: on flat margin ( Fig. 12; V. mexicana sp. nov.) (0), on protuberance ( Fig. 11; Gerbelius sp. 2) (1), on decline ( Fig. 15; Acanthaspis sp.) (2). Adjacent to the median apical process are clusters of macrosetae. Some are on a protuberance (e.g., Gerbelius sp. 2), a decline (e.g., Nalata nr. spinicollis), or this posterior margin may be flat (e.g., V. mexicana sp. nov.). With the amount of missing data, it appears that the shape of the posterior pygophore might vary interspecifically. 77. Basal plate extension, length: 1.5-3.9 times as long as wide (0), 4.0-5.4 times as long as wide ( Fig. 14; V. mexicana sp. nov.) (1), 5.5-6.4 times as long as wide (2), 6.5-8.0 times as long as wide (3), as long as 8.5 times the width (4). Width is recorded from the midpoint. The range for the length to width ratio of the basal plate extension within Pseudocetherinae is between 4.4 and 5.8, with V. typica as the exception having a ratio of 7.9. A basal plate extension that is 4.0-5.4 times as long as wide is a synapomorphy of G. typicus + Gerbelius sp. 2 as well as V. lasiosoma sp. nov. + V. laosensis sp. nov. A basal plate extension that is 5.5-6.4 times as long as wide is a synapomorphy of the Australian clade containing V. pallidipes and kin.

Diagnosis
Distinguished from other Reduviidae by the combination of the following characters: head and pronotum pubescent with interspersed macrosetae or setigerous tubercles; labial segment II straight, as long as remaining segments, and adpressed to head (

Redescription
Usually macropterous, only micropterous morphs are known in three species and one species includes both micropterous and macropterous morphs.
head. Elongate or globose; pubescent with interspersed macrosetae or setigerous tubercles; interocular region with two glabrous lines; scape and pedicel with dense, short macrosetae; scape reaching or slightly surpassing head apex, 0.2-0.3 times length of pedicel; ocelli present in macropterous morphs and minute or absent in micropterous morphs; buccula with ventrolateral swelling; labial segment III drawn into bent, slender apex.
thorax. Prosternum in lateral view drawn into anteriad-directed process; scutellar spine long, either sub-horizontal or raised.
legs. Fossula spongiosa present on fore tibia, usually present on mid tibia of males; all femora with ventral anterior and posterior rows of spine-like protuberances with macrosetae at base of protuberance (Fig. 4I); tibiae laterally compressed ( Fig. 4G-H).

Male
PygoPhore. Posterior margin with a short median apical process, either straight or bent posteriorly.
aedeagus. Basal plate extension long in relation to phallotheca; endosoma covered in spicules.

Habitat and collecting method
Pseudocetherinae have been collected using light traps, but records on habitat and collecting methods are scarce on specimen labels.

Revised diagnosis
Distinguished from the other genus in the subfamily, Voconia, by the smooth or corrugated surface of the head and pronotum (Fig. 3G); laterally flattened shape of the maxillary plates (Fig. 3A); long, basally adjacent mandibular plates (Fig. 3A); stout labium densely covered with long macrosetae on the morphologically ventral surface of labial segment I (Fig. 3G); morphologically dorsal surface of labial segment II 0.8-0.9 times length of labial segment I (Fig. 3G); and Cu-An 1 cell slender and tapering apically, similar in size to the M-Cu cell (Fig. 4A).
Coloration. Variable color patterns; scutellum uniform or with contrasting yellow scutellar spine. integument and vestiture. Head and pronotum: smooth to corrugated with dense pubescence and sparse, long macrosetae; antennifer with macroseta, without lateral protuberance (Fig. 3A); labium with dense, long macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae. Hemelytron: corium with dense setation. Legs: posterior row of protuberances on mid and hind femora with three large spines on distal half and three or fewer small spines. Abdomen: ventral surface pubescent with long macrosetae. head ( Fig. 3A, G). Elongate, 1.2-1.3 times as long as wide; anteocular region about a third of head length and shorter than postocular region (measured to anterior margin of neck); postocular region in dorsal view globose and longer than eye; pedicel 1.1-1.3 times length of head width; mandibular plates in dorsal view long with basal half adjacent; maxillary plates in dorsal view enlarged laterally forming flattened projections, adjacent to clypeus; clypeus width in dorsal view narrower than maxillary plates; interocular glabrous markings V-shaped, joined medially at interocular sulcus; eye width in dorsal view narrower than synthlipsis, eye not reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labium stout; second (first visible) labial segment in lateral view surpassing posteroventral eye margin, gradually widened apically; dorsal surface of third (second visible) labial segment convex, 0.8-0.9 times length of second segment; fourth (third visible) labial segment drawn into bent, slender apex.
legs. Fossula spongiosa present on fore and mid legs in males, present on fore but absent or vestigial on mid legs of females.

Distribution
This genus currently comprises two described species from India and Southeast Asia, and we are aware of at least four undescribed species distributed across India, Laos, Thailand, and Vietnam. Gerbelius typicus is the most widespread among them.

Remarks
Photographs of type specimens in their unit trays at the BMNH were used to evaluate species concepts for G. typicus and G. confluens. This genus previously had a third described species that is now transferred to Voconia, V. ornata. Molecular phylogenetics recovered a Gerbelius clade (Gerbelius sp. 3 + [Gerbelius sp. 1 + Gerbelius sp. 4]) sister to Voconia wegneri sp. nov. (Hwang & Weirauch, 2012). We also recovered a well-supported Gerbelius clade sister to Voconia (Figs 1-2). The pygophore of two specimens were dissected for this study (Figs 11,14); however, the median apical process of G. typicus (AMNH_IZC 00321200) broke off during dissection (Fig. 11). We decided not to describe new species of Gerbelius as part of this study. Since species of Gerbelius somewhat resemble other small-bodied assassin bugs currently classified as Reduviinae, we suspect that our specimen searches in various natural history collections were not comprehensive for this genus. However, this material should be included in a future revision of this genus. Distant, 1903 Gerbelius confluens Distant, 1903b: 260, pl. 16 Distant, 1903 Gerbelius typicus Distant, 1903a

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).
Body. Slender or robust; macropterous or micropterous (Figs 7-10). 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.  (Villiers, 1964) comb. nov.

Key to the species of Voconia
5. Head and pronotum coarsely granulose, with setigerous tubercles ( (Miller, 1954) (Miller, 1958) -2, 7, 9, 11, 14, 20 Diagnosis This species most closely resembles two other dark-brown Philippine species, V. minima sp. nov. and V. nyx sp. nov., with yellow posteromedial yellow spots on the abducted corium, but is differentiated by the yellow patches on the dorsal laterotergites, the faint yellow spot on the anteromedial portion of the abducted corium, the presence of small paramedial lobes on the anteriad-directed process of the prosternum, and the stridulitrum being elongated anteriorly into a small protuberance. It specifically differs from V. minima sp. nov. by the uniformly dark legs and short labial segment I that does not reach the posteroventral eye margin and from V. nyx sp. nov. by the uniformly dark pronotum.

Etymology
Named after Austin Baker, a former colleague at UCR who provided support and insightful comments on this revision.

Description
Male (Figs 7,9) Body length. About 9.8 mm; macropterous. Coloration. Head: dark brown with paired pale patches adjacent to lateral ocellar margin; labium nearly as dark as head. Thorax: coloration as head; scutellum uniformly dark brown. Hemelytron (abducted): clavus dark reddish-brown, distal half yellow; corium dark reddish-brown with anteromedial, posteromedial, and distal yellow spots; membrane dark with pale V-shaped marking along R and M veins; membranal veins R and M proximally pale, remainder as membrane. Legs: dark brown. Abdomen: dark reddish-brown, almost black; dorsal laterotergites dark with yellow spots anteriorly. integument and vestiture. Head and pronotum: finely granulose with dense, long macrosetae interspersed among dense pubescence; interocular region with two pairs of macrosetae paramedially; antennifer with macroseta, base without lateral protuberance; morphologically ventral surface of labium with sparse, short macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae; scutellar lateral carinae with long setation. Hemelytron: corium with sparse long setation. Legs: posterior row of protuberances on mid and hind femora with three large and two small spines on distal half. Abdomen: ventral surface pubescent.
struCture. Head: elongate, about 1.2 times as long as wide; anteocular region about one quarter of head length, shorter than postocular region (measured to anterior margin of neck); postocular region in dorsal view about as long as eye, lateral margins in dorsal view gently rounded; pedicel about 1.2 times length of head width; maxillary plates ellipsoidal, adjacent to and longer than clypeus; apices of maxillary plates in dorsal view converging; clypeus in dorsal view wider than maxillary plates; clypeal apex round, narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view nearly straight; eye width in dorsal view about as wide as synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, not reaching posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.6 times length of segment I. Thorax: pronotal collar narrow medially with anterolateral angles short and flared laterally; anterior pronotal lobe about 0.7 times length of posterior pronotal lobe, lateral margins rounded; glabrous markings on pronotum thin and not deeply depressed; median apodeme depression of pronotum deep, elongated longitudinally; scutellar spine long, raised; anteriad-directed process of prosternum with paramedial lobes; anterior margin of stridulitrum projected into small protuberance; proepimeron with acute protuberance on posteroventral margin. Legs: fossula spongiosa present on fore and mid legs. Abdomen: anterior margin of terga weakly carinulate; terga II and III with paired prominent longitudinal carinae, reaching about two-thirds of tergum III. Pygophore (Fig. 11): transverse bridge with rounded posterior margin; posterior region of ventral surface of pygophore in lateral view straight, not swollen; lateral pygophore margin with protuberance; posterior pygophore margin with clustered macrosetae on protuberance; parameres sinusoidal, apex tapered into squarely rounded tip. Aedeagus (Fig. 14): endosoma almost entirely covered with spicules; apex of dorsal phallothecal sclerite tongue-shaped in dorsal view; basal plate extension about 4.3 times as long as wide.

Unknown.
Distribution (Fig. 20) This species is only known from the type locality in the Philippines. Based on the record of a Miridae Hahn, 1831 specimen with the same collection event, Mt Pomalihi is presumably in Mt Balatukan Range Natural Park. The type locality is also shared with V. isosceles sp. nov.

Remarks
The median apical process broke off during dissection (Fig. 11). It was presumably short as is the case in all Pseudocetherinae, but it was not possible to determine if it was upright or bent posteriorly.

Diagnosis
Distinguished from other species of the genus Voconia by its long, slender body (about 10.8 mm), long head (about 1.7 times as long as wide), hind leg with four large spines in the posterior row, a large pale spot between R and M veins, and dorsal laterotergites I and II yellow, the remainder dark brown. This species is most similar to V. typica, but is differentiated by the yellow proximal third of the mid and hind femora as well as the anterior pronotal lobe, which is convex, strongly curved inward on the posterolateral margins.

Etymology
The specific epithet is the Latin adjective 'bracatus, -a, -um', meaning 'wearing trousers'. It refers to the yellow coloration of the proximal half of the femora. Coloration. Head: dark brown; postocular region with dark patches adjacent to medial ocellar margin and paired pale patches adjacent to lateral ocellar margin; labial segments II and III lighter than head. Thorax: coloration as head; scutellum dark with contrasting yellow apical spine. Hemelytron (abducted): clavus dark reddish-brown with distal half yellow; corium dark reddish-brown with anteromedial, posteromedial, and distal yellow spots; membrane dark, large pale spot between R and M veins; membranal veins R and M proximally pale, remainder as membrane. Legs (Fig. 4K): dark brown with yellow mid and hind coxae, trochanter, and proximal third of femora. Abdomen (Fig. 4C): dark reddishbrown; dorsal laterotergites I and II yellow. integument and vestiture. Head and pronotum: finely granulose with sparse, short macrosetae interspersed among dense pubescence; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with dense, short macrosetae; anterolateral angles of pronotal collar without macrosetae or setigerous tubercles. Legs (Fig. 4K): posterior row of protuberances on mid and hind femora with four large spines on distal half. Abdomen: ventral surface pubescent with long macrosetae. struCture. Head: elongate, about 1.7 times as long as wide; anteocular region about one third of head length, subequal in length to postocular region (measured to anterior margin of neck); postocular region in dorsal view longer than eye, lateral margins gently rounded; pedicel about 1.8 times length of head width; maxillary plates ellipsoidal, adjacent to and reaching apex of clypeus; apex of maxillary plates in dorsal view directed straight; clypeus in dorsal view slightly narrower than maxillary plates; clypeal apex round, not narrowed; interocular glabrous markings V-shaped, joined medially at interocular sulcus; interocular sulcus in dorsal view slightly bent anteromedially, subtriangular; eye width in dorsal view wider than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, surpassing posteroventral eye margin; morphologically dorsal surface of labial segment II slightly curving apically, about 0.4 times length of segment I. Thorax: pronotal collar narrow medially with anterolateral angles short, projected forward; anterior pronotal lobe subequal in length to posterior pronotal lobe, lateral margins rounded; glabrous markings on pronotum thin and not deeply depressed; median apodeme depression of pronotum deep and elongated transversely; scutellar spine long and subhorizontal; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes (Fig. 3J); anterior margin of stridulitrum not elongated into protuberance (Fig. 3J); proepimeron with smoothly rounded posteroventral margin. Legs: fossula spongiosa present on fore and mid legs.

Unknown.
Distribution (Fig. 18) This species is only known from the type locality of Normanby Island off of Papua New Guinea, where no other Pseudocetherinae species are known.

Remarks
This species forms a well-supported clade with another species from Papua New Guinea, V. typica (Figs 1-2).

Diagnosis
Recognized by its Australasian distribution, small size (about 7.4 mm), globose head (about as long as wide), labial segment II swollen ventromedially, and coarsely granulose, tuberculate head and pronotum. Most closely resembles V. dolichocephala sp. nov. in size and hemelytral coloration but V. dolichocephala sp. nov. has a conspicuously elongate head.

Etymology
The specific epithet is the latinized Greek adjective 'brachycephalus, -a, -um', formed from the prefix 'brachy-' (meaning 'short') and the noun 'cephale' (meaning 'head'), referring to the fact that this species has the shortest head among species of Voconia. Coloration. Head: dark brown, postocular region with dark patches adjacent to medial ocellar margin; labium lighter than head. Thorax: coloration as head; scutellum dark with contrasting yellow apical spine. Hemelytron (abducted): clavus reddish-brown with distal yellow stripe; corium reddish-brown with anteromedial and posteromedial yellow spots; membrane dark with pale V-shaped marking along R and M veins; membranal veins R and M proximally pale, remainder as membrane. Legs: dark yellowishbrown. Abdomen: dark brown; dorsal laterotergites uniformly dark. integument and vestiture. Head and pronotum: coarsely granulose, with dense, short setigerous tubercles interspersed among sparse pubescence; interocular region with two pairs of stout setigerous tubercles paramedially and one medially; antennifer with two adjacent lateral setigerous tubercles; morphologically ventral surface of labium with sparse, long macrosetae. Thorax: anterolateral angles of pronotal collar with setigerous tubercles; scutellar lateral carinae with short macrosetae on setigerous tubercles. Legs: posterior row of protuberances on mid and hind femora with three large spines on distal half and three or fewer small spines. Abdomen: ventral surface pubescent with long macrosetae.
struCture. Head: globose, about as long as wide; anteocular region about one quarter of head length, shorter than postocular region (measured to anterior margin of neck); postocular region in dorsal view shorter than eye, lateral margins gently rounded; pedicel about 1.2 times length of head width; maxillary plates ellipsoidal, adjacent to and shorter than clypeus; apices of maxillary plates in dorsal view diverging; clypeus in dorsal view wider than maxillary plates; clypeal apex bifid, narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view curved and smoothly rounded anteriorly; eye width in dorsal view narrower than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view gradually thickens distally, surpassing posteroventral eye margin; labial segment II swollen ventromedially; morphologically dorsal surface of labial segment II curves ventrally, about 0.3 times length of segment I. Thorax: pronotal collar narrow medially with anterolateral angles short, projected forward; anterior pronotal lobe about 0.7 times length of posterior pronotal lobe, lateral margins almost straight; glabrous markings on pronotum conspicuous and not depressed; median apodeme depression of pronotum shallow, elongated longitudinally; scutellar spine long and raised; anteriad-directed process of prosternum with paramedial lobes; anterior margin of stridulitrum projected into small protuberance; proepimeron with smoothly rounded posteroventral margin. Legs: fossula spongiosa of fore leg present.

Male
Unknown.
Distribution (Fig. 18) This species has a disjunct distribution. It is known from Erima in Papua New Guinea (type locality) and the Northern Territory of Australia (Swanson 2015: 268, as V. pallidipes).

Remarks
There is little doubt that the specimen figured by Swanson (2015: 269, fig. 10) as V. pallidipes pertains to V. brachycephala sp. nov. Although the size of the specimen in concern is unknown, its coloration and the short, tuberculate head do not seem to differ from the condition seen in the new species. Since the specimen could not be re-examined, it is, however, not included in the type material of the new species.

Diagnosis
Recognized from other African Voconia species by the unique coloration of the corium, abdomen, and legs which are entirely yellow. Additionally, the posterior row of spines on the mid and hind femora have three large and many small spines, and the antennifer lacks a lateral protuberance. Females possess a fossula spongiosa on the mid leg.

Etymology
The specific epithet is the Latin adjective 'chrysopterus, -a, -um', meaning 'golden-winged', referring to the entirely yellow corium. Coloration. Head: dark brown, maxillary plates lighter; postocular region dark brown with paired pale patches adjacent to lateral ocellar margin; labium lighter than head, yellow. Thorax: coloration as head, with posterior pronotal lobe lighter than anterior lobe; scutellum dark with contrasting yellow apical spine. Hemelytron (abducted): clavus and corium uniformly yellow; membrane uniformly dark; membranal veins as remainder of membrane. Legs: yellow. Abdomen: ventral surface and dorsal laterotergites yellow. integument and vestiture. Head and pronotum: coarsely granulose with sparse pubescence and sparse, short macrosetae; interocular region with two pairs of macrosetae paramedially; antennifer without lateral projection; labium with dense, short macrosetae. Thorax: anterolateral angles of pronotal collar without macrosetae or setigerous tubercles; scutellar lateral carinae with short macrosetae. Hemelytron: corium with dense short setation. Legs: posterior row of protuberances on mid and hind femora with three large spines on distal half and many small spines. Abdomen: ventral surface pubescent.
struCture. Head: elongate, 1.3 times as long as wide; anteocular region about one third of head length, shorter than postocular region (measured to anterior margin of neck); postocular region in dorsal view longer than eye, lateral margins gently rounded; pedicel about 1.8 times length of head width; maxillary plates ellipsoidal, adjacent to and longer than clypeus; apices of maxillary plates in dorsal view diverging; clypeus in dorsal view wider than maxillary plates, apex rounded, not narrowed; interocular glabrous markings curved anterolaterally, joined at interocular sulcus paramedially; interocular sulcus in dorsal view curved and smoothly rounded anteriorly; eye width in dorsal view narrower than synthlipsis, reaching ventral head margin in lateral view; ventrolateral swelling of buccula with lateral protrusion that surpasses buccular margin, setigerous tubercles along apical margin; labial segment I in lateral view straight, reaching posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.6 times length of segment I. Thorax: with pronotal collar narrow medially with anterolateral angles short, projected forward; anterior pronotal lobe about 0.5 times length of posterior pronotal lobe, lateral margins rounded, glabrous markings not depressed; median apodeme depression of pronotum shallow and elongated longitudinally; scutellar spine long and raised; anteriad-directed process of prosternum smoothly rounded without paramedial lobes; anterior margin of stridulitrum projecting into a small protuberance. Legs: fossula spongiosa present on fore and mid legs.

Male
Unknown.
Distribution (Fig. 17) This species is only known from the type locality in Mwingi, Kenya. It is the easternmost Afrotropical species.

Remarks
This is the only female in the subfamily with fossula spongiosa present on the mid leg.

Diagnosis
Distinguished from other species in Voconia by its Afrotropical distribution and stout body. This species most closely resembles V. motoensis and V. smithae sp. nov. It differs from V. motoensis by its distinct coloration: yellow scutellar spine, corium with a proximal and distal yellow spot, and contrasting yellow anterior spots on dark dorsal laterotergites. Despite the strong superficial resemblance to V. smithae sp. nov. due to its coloration and stoutness, closer examination reveals that V. conradti is distinguishable by the longer body (10.4-11.4 mm), dark dorsal laterotergites with strongly contrasting yellow anterior spots, finely granulose head, antennifer with short or absent laterally projecting setigerous tubercle, lack of paired interocular setigerous tubercles, maxillary plates with medial margins diverging, shorter anteocular region than postocular region (measured to where the posterior margin of the granulations meet the anterior margin of the smooth neck), buccula without lateral protrusion, and strongly bulging eyes that greatly surpass the postocular lateral margin in dorsal view and reach the ventral head margin in lateral view.

Type material
Lectotype (
Coloration. Head: dark brown; postocular region with dark patches adjacent to medial ocellar margin and pale patches adjacent to lateral ocellar margin; labium lighter than head. Thorax: coloration as head; pronotum uniformly dark (Fig. 3K) or posterior pronotal lobe lighter than anterior lobe; scutellum dark with contrasting yellow apical spine. Hemelytron (abducted): clavus dark reddish-brown with distal yellow stripe; corium dark reddish-brown with proximal, anteromedial, posteromedial, and distal yellow spots; membrane dark with pale V-shaped marking along R and M veins; membranal veins R and M proximally pale. Legs: completely yellow or yellow with distal half of femur dark brown. Abdomen: dark brown, dorsal laterotergites with yellow spots anteriorly. integument and vestiture. Head and pronotum: finely granulose with sparse, short macrosetae interspersed among sparse pubescence; interocular region with two pairs of macrosetae paramedially; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with sparse, short macrosetae. Thorax (Fig. 3K): anterolateral angles of pronotal collar without macrosetae or setigerous tubercles. Hemelytron: corium with dense long setation. Legs: posterior row of protuberances on mid and hind femora with three large spines on distal half. Abdomen: ventral surface pubescent.
struCture. Head: elongate, 1.1-1.4 times as long as wide; anteocular region about one quarter of head length, shorter than postocular region (measured to anterior margin of neck); postocular region in dorsal view about as long as eye or shorter, lateral margins subrectangular; pedicel about 1.3 times length of head width; maxillary plates ellipsoidal, adjacent to and longer than clypeus; apices of maxillary plates in dorsal view with medial margins diverging; clypeus in dorsal view narrower than maxillary plates; clypeal apex round, narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view curved and smoothly rounded anteriorly; eye width in dorsal view either wider or narrower than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, reaching posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.5 times length of segment I. Thorax (Fig. 3K): pronotal collar in dorsal view narrow medially, with anterolateral angles short, projected forward; anterior pronotal lobe 0.5-0.6 times length of posterior pronotal lobe, lateral margins almost straight; glabrous markings on pronotum slightly depressed; median apodeme depression of pronotum deep, circular; scutellar spine long, raised; anteriad-directed process of prosternum with paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with smoothly rounded posteroventral margin. Legs: fossula spongiosa present on fore and mid legs. Abdomen: anterior margin of terga weakly carinulate; terga II and III with paired prominent longitudinal carinae reaching about one-third of tergum III. Pygophore (Fig. 11): transverse bridge with triangular posterior margin; posterior region of ventral surface of pygophore in lateral view with slight swelling; median apical process in caudal view short; direction of median apical process in lateral view sharply bent posteriorly; lateral pygophore margin with protuberance; posterior pygophore margin with clustered macrosetae on protuberance; parameres sinusoidal, apex tapered into squarely rounded tip. Fig. 11. Pygophore of species of Gerbelius Distant, 1903 andVoconia Stål, 1866. Median apical process of Gerbelius typicus Distant, 1903 and V. bakeri sp. nov. damaged/missing. Scale bars = 0.5 mm.

Female
Mostly as male but differs from males by the following. Body length: about 11.2 mm. Head: elongate, about 1.4 times as long as wide; pedicel about 1.2 times length of head width; ocelli minute, occupying 0.15 times length of postocular region. Legs: fossula absent or vestigial on mid leg. Distribution (Fig. 17) This species is distributed in Western Africa. The type locality is from Bioko, an island off the west coast, north of Equatorial Guinea.

Remarks
Originally described as Microvarus conradti based on several specimens which are to be treated as syntypes (Jeannel 1917). The type material of M. conradti is deposited at the MNHN under loan restrictions; habitus photographs of one of the syntypes (Fig. 7; USI: MNHN_EH24709) were used to evaluate this species concept and this specimen is designated as the lectotype. The synonymy of M. achteni proposed by Villiers (1948) is accepted here without re-examining the type material of Schouteden (1929). This species is transferred to Voconia based on the cladistic analysis conducted in this study. Based on the phylogeny, V. conradti and V. smithae sp. nov. form a well-supported clade (Figs 1-2).

Diagnosis
Distinguished from most species of Voconia by the semicircular yellow stripe encircling both ocelli, membranal veins forming Cu-An 1 and M-Cu cells paler than remainder of membrane, and proepimeron with protuberance on posteroventral margin. This species is most similar to two other Oriental species, V. decorata sp. nov. and V. lirophleps sp. nov. It is distinguishable from V. decorata sp. nov. by its distribution in Sumatra and the dark brown, almost black coloration of the body. Despite the holotypes of V. coronata sp. nov. and V. lirophleps sp. nov. being collected within about 106 km of each other, there are several differences, particularly in size. Among our larger series, conspecific Voconia males do not vary more than about ± 1 mm from each other, and V. coronata sp. nov. is about 2 mm longer than V. lirophleps sp. nov. It is distinguishable by the slightly more elongate head (about 1.3 times as long as wide), the pedicel is much longer (about 1.9 times length of head width), maxillary plates directed straight in dorsal view, postocular region is about as long as the eye, labial segment I is long, almost reaching posterior margin of head, wide pronotal collar, long anterior pronotal lobe relative to posterior pronotal lobe (posterior lobe about 1.8 times as long as anterior lobe), scutellum dark with brown apical spine, inconspicuous distal yellow spot on the corium, and the presence of three large spines and many small spines on the posterior row of spines of the hind leg.

Etymology
The specific epithet is the Latin adjective 'coronatus, -a, -um', meaning 'crowned' or 'encircled', referring to the yellow ring around the postocular region of the head.
Coloration. Head: dark brown; postocular region dark with a semicircular pale stripe encircling both ocelli; labium nearly as dark as head. Thorax: coloration as head; posterior pronotal lobe lighter than anterior lobe; scutellum dark with brown apical spine. Hemelytron (abducted): clavus dark with distal yellow stripe; corium dark with anteroproximal yellow stripe, posteromedial and small distal yellow spots; membrane uniformly dark; membranal veins forming Cu-An 1 and M-Cu cells paler than remainder of membrane. Legs: dark brown, femora darker. Abdomen: almost black, dorsal laterotergites dark with yellow spots anteriorly. integument and vestiture. Head and pronotum: finely granulose with sparse, short macrosetae interspersed among sparse pubescence; interocular region with two pairs of macrosetae paramedially; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with sparse, long macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae; scutellar lateral carinae with long setation. Hemelytron: corium with sparse long setation. Legs: posterior row of protuberances on mid and hind femora with three large spines on distal half and more than three small spines. Abdomen: ventral surface pubescent.
struCture. Head: elongate, 1.3 times as long as wide; anteocular region about one quarter of head length, shorter than postocular region (measured to anterior margin of neck); postocular region in dorsal view about as long as eye, lateral margins gently rounded; pedicel about 1.9 times length of head width; maxillary plates ellipsoidal, adjacent to and surpassing clypeal apex; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view wider than maxillary plates; clypeal apex round, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view bent anteromedially, subtriangular; eye width in dorsal view wider than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, surpassing posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.5 times length of segment I. Thorax: pronotal collar in dorsal view wide medially with anterolateral angles short and flared laterally; anterior pronotal lobe about 0.6 times length of posterior pronotal lobe, lateral margins rounded; glabrous markings on pronotum deeply depressed; median apodeme depression of pronotum shallow, elongated transversely; scutellar spine long, raised; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with acute protuberance on posteroventral margin. Legs: fossula spongiosa present on fore and mid legs.

Unknown.
Distribution (Fig. 20) This species is only known from the type locality of Ketambe in the northernmost province of Sumatra, Aceh.
Voconia decorata sp. nov. urn:lsid:zoobank.org:act:93BE46E2-210B-45C7-8FD6-2C38EC62B2D9 Figs 1-2, 3B, 4J, 7, 9, 20 Diagnosis Distinguished from most other species of Voconia by the yellow ring around the postocular region, membranal veins forming Cu-An 1 and M-Cu cells paler than remainder of membrane, and proepimeron with protuberance on posteroventral margin. This species is most similar to two other Oriental species, V. coronata sp. nov. and V. lirophleps sp. nov. It is distinguishable from these species by its distribution in the Malaysian state of Sabah in Borneo, the light brown coloration of the body, the labium being lighter than the head, and the dark scutellum with a contrasting yellow apical spine.

Etymology
The specific epithet is the Latin adjective 'decoratus, -a, -um', meaning 'decorated, elegant', referring to its light brown coloration and yellow markings on the hemelytron and dorsal laterotergites that gives it a flashier and more elegant look. Coloration. Head (Fig. 3B): light brown, yellowish; postocular region with dark patches adjacent to medial ocellar margin, semicircular pale stripe encircling both ocelli; labium lighter than head. Thorax: coloration as head; posterior margin of posterior pronotal lobe lighter; scutellum dark with contrasting yellow apical spine. Hemelytron (abducted): clavus reddish-brown with distal yellow stripe; corium reddish-brown with anteroproximal yellow stripe, posteromedial and distal yellow spots; membrane light brown with pale V-shaped marking along R and M veins; membranal veins forming Cu-An 1 and M-Cu cells paler than remainder of membrane. Legs: yellowish-brown, distal half of femora darker. Abdomen: yellowish-brown, dorsal laterotergites with yellow spots anteriorly. integument and vestiture. Head (Fig. 3B) and pronotum: finely granulose with sparse, short macrosetae interspersed among sparse pubescence; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with dense, short macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae. Hemelytron: corium with sparse, long setation. Legs (Fig. 4J): posterior row of protuberances on mid and hind femora with three large spines on distal half and one small spine. Abdomen: ventral surface pubescent.

Holotype
struCture. Head (Fig. 3B): elongate, 1.3 times as long as wide; anteocular region about one third of head length, shorter than postocular region (measured to anterior margin of neck); postocular region in dorsal view shorter than eye, lateral margins gently rounded; pedicel about 1.3 times length of head width; maxillary plates ellipsoidal, adjacent to and as long as clypeus; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view about as wide as maxillary plates; clypeal apex round, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view bent anteromedially, subtriangular; eye width in dorsal view narrower than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula with lateral protrusion that surpasses buccular margin; labial segment I in lateral view straight, surpassing posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.5 times length of segment I. Thorax: pronotal collar in dorsal view wide medially with anterolateral angles short, projected forward; anterior pronotal lobe about 0.7 times length of posterior pronotal lobe, lateral margins almost straight; glabrous markings on pronotum deeply depressed; median apodeme depression of pronotum shallow, elongated longitudinally; scutellar spine long, subhorizontal; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with acute protuberance on posteroventral margin. Legs: fossula spongiosa present on fore and mid legs.

Unknown.
Distribution (Fig. 20) This species is only known from the type locality of Quoin Hill in the Malaysian state of Sabah on the island of Borneo.

Diagnosis
Distinguished from most other species of Voconia by its tuberculate and coarsely granulose head and pronotum. This species most closely resembles another Australian species, V. tuberculata sp. nov. These two species are recognized from other Australian species by the acute protuberance on the posteroventral margin of the proepimeron, long macrosetae of the setigerous tubercles, elongate head (1.4-1.6 times as long as wide), and yellow spots antero-and posteromedially on the abducted corium. Voconia dolichocephala sp. nov. differs from V. tuberculata sp. nov. by its overall dark brown, almost black coloration, dorsal laterotergites uniformly dark or with yellow spots posteriorly, and males are smaller than about 7.5 mm. Females slightly surpass the high-end range for female length of V. tuberculata sp. nov. (8.4-8.8 mm).

Etymology
The specific epithet is the latinized Greek adjective 'dolichocephalus, -a, -um', formed from the prefix 'dolicho-' (meaning 'long') and the noun 'cephale' (meaning 'head'), referring to the long head of the type specimen, which is about a fifth of the body length.
Coloration. Head (Fig. 3F): dark brown; postocular region uniformly dark with pale patches adjacent to lateral ocellar margin; labial segment I nearly as dark as head, segments II and III lighter than head. Thorax: coloration as head; scutellum uniformly dark brown. Hemelytron (abducted): clavus dark brown with distal yellow stripe; corium dark brown with anteromedial, posteromedial, and distal yellow spots; membrane dark with pale V-shaped marking along R and M veins; membranal veins R and M proximally pale. Legs: dark brown, tarsi yellow. Abdomen: dark brown; dorsal laterotergites uniformly dark or dark with yellow spots posteriorly. integument and vestiture. Head (Fig. 3F) and pronotum: coarsely granulose with sparse, long setigerous tubercles interspersed among sparse pubescence; interocular region with two pairs of stout setigerous tubercles paramedially; antennifer with long lateral setigerous tubercle; morphologically ventral surface of labium with sparse, short macrosetae. Thorax: anterolateral angles of pronotal collar with setigerous tubercles; scutellar lateral carinae with setigerous tubercles with long macrosetae. Hemelytron: corium with dense long setation. Legs: posterior row of protuberances on mid and hind femora with four large spines on distal half. Abdomen: ventral surface pubescent, long macrosetae interspersed.
struCture. Head (Fig. 3F): elongate, 1.5-1.6 times as long as wide; anteocular region about one third of head length, shorter than postocular region (measured to anterior margin of neck); postocular region in dorsal view as long as eye, lateral margins gently rounded; pedicel 1.3-1.5 times length of head width; maxillary plates ellipsoidal, adjacent to and as long as clypeus; apices of maxillary plates in dorsal view with medial margins diverging; clypeus in dorsal view wider than maxillary plates; clypeal apex bifid, not narrowed; interocular glabrous markings V-shaped, joined medially at interocular sulcus; interocular sulcus in dorsal view nearly straight; eye width in dorsal view narrower than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, surpassing posteroventral eye margin; morphologically dorsal surface of labial segment II curves ventrally, 0.3-0.4 times length of segment I. Thorax: pronotal collar in dorsal view wide medially with anterolateral angles long, flared laterally; anterior pronotal lobe 0.7-0.8 times length of posterior pronotal lobe, lateral margins almost straight; glabrous markings on pronotum conspicuous and not depressed; median apodeme depression of pronotum shallow, elongated longitudinally; scutellar spine long, subhorizontal; anteriad-directed process of prosternum with paramedial lobes; anterior margin of stridulitrum projected into small protuberance; proepimeron with acute protuberance on posteroventral margin. Legs: fossula spongiosa present on fore and mid legs. Abdomen: anterior margin of terga weakly carinulate, terga II and III with deeper and longer carinules; terga II and III with paired prominent longitudinal carinae, reaching about two-thirds of tergum III.

Female
Mostly as male, distinguished from males by the following. Body length: 8.8-9.0 mm. Head: elongate, 1.5-1.6 times as long as wide; pedicel about 1.5-1.7 times length of head width. Legs: fossula spongiosa absent or vestigial on mid leg. Distribution (Fig. 18) This species is only known from the Northern Territory of Australia. The nearest species of Voconia is V. tuberculata sp. nov., known from about 100 km southeast of the three specimens collected in Tindal.

Remarks
Voconia dolichocephala sp. nov. is nested among the Australian clade. According to the phylogeny (Figs 1-2), it is the sister taxon to V. tuberculata sp. nov.

Diagnosis
Distinguished from most other species of Voconia by its tuberculate and coarsely granulose head and pronotum, lateral protrusion of the buccula, and dorsally projecting mandibular plates. This species most closely resembles three other Australian species, V. grandioculata sp. nov., V. pallidipes, and V. vittata sp. nov. This species, V. grandioculata sp. nov., and V. vittata sp. nov. have a similar yellow stripe across the corium; however, the stripe in V. fasciata sp. nov. is pale, almost translucent and the body is larger, total length is about 8.7 mm long. It is easily distinguished from V. pallidipes by its smaller size and coloration, i.e., the membranal veins are not yellow, the proximal half of the corium is not entirely yellow, and the legs are not completely yellow as described above.

Etymology
The specific epithet is the Latin adjective 'fasciatus, -a, -um', meaning 'banded', referring to the pale yellow band that crosses the middle of the corium and scutellar spine. Coloration. Dark brown, maxillary plates lighter; postocular dark with pale patches adjacent to lateral ocellar margin; labium lighter than head. Thorax: coloration as head; posterior pronotal lobe with yellow posterolateral margins; scutellum dark with contrasting yellow apical spine. Hemelytron (abducted): clavus dark brown with distal yellow stripe; corium dark brown with thick medial yellow stripe; membrane and membranal veins uniformly dark. Legs: femora dark brown, tibiae and tarsi yellowishbrown. Abdomen: ventral surface light brown medially, dark brown laterally; dorsal laterotergites I and II paler than remainder. integument and vestiture. Head and pronotum: coarsely granulose with dense, short setigerous tubercles interspersed among sparse pubescence; interocular region with setigerous tubercles medially and two pairs paramedially; antennifer with long lateral setigerous tubercle; morphologically ventral surface of labium with sparse, short macrosetae. Thorax: anterolateral angles of pronotal collar with setigerous tubercles; scutellar lateral carinae with short macrosetae on setigerous tubercles. Hemelytron: corium with dense, long macrosetae. Legs: posterior row of protuberances on mid and hind femora with four spines on distal half. Abdomen: ventral surface pubescent, long macrosetae interspersed. struCture. Head: globose, about as long as wide; anteocular region about one third of head length, about as long as postocular region (measured to anterior margin of neck); postocular region in dorsal view shorter than eye, lateral margins gently rounded; pedicel about 1.3 times length of head width; mandibular plates in anterior view projecting dorsolaterally; maxillary plates ellipsoidal, adjacent to and reaching apex of clypeus; apices of maxillary plates in dorsal view diverging; clypeus in dorsal view wider than maxillary plates; clypeal apex bifid, narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view nearly straight; eye width in dorsal view narrower than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula with lateral protrusion that surpasses buccular margin, setigerous tubercles along apical margin; labial segment I in lateral view gradually thickens distally, surpassing posteroventral eye margin; morphologically dorsal surface of labial segment II curves ventrally, about 0.3 times length of segment I. Thorax: pronotal collar in dorsal view wide medially with anterolateral angles short, flared laterally; anterior pronotal lobe about 0.7 times length of posterior pronotal lobe, lateral margins almost straight; glabrous markings on pronotum depressed near posterolateral margins; median apodeme depression of pronotum shallow, elongated transversely; scutellar spine long, subhorizontal; anteriad-directed process of prosternum with paramedial lobes; anterior margin of stridulitrum projected into sharp spine; proepimeron with smoothly rounded posteroventral margin. Legs: fossula spongiosa present on fore and mid legs.

Unknown.
Distribution (Fig. 18) This species is only known from the type locality of Einasleigh River in Queensland, Australia.

Remarks
Voconia fasciata sp. nov. is nested within the Australian clade (Figs 1-2). Though poorly supported, it forms the sister taxon to V. brachycephala sp. nov. and V. vittata sp. nov.
Voconia grandioculata sp. nov. urn:lsid:zoobank.org:act:23167BBE-77C8-434F-9574-B9F59130EAAB Figs 1-2, 4E, 7, 9, 11, 14, 18 Diagnosis Distinguished from most other species of Voconia by its tuberculate and coarsely granulose head and pronotum, the lateral protrusion of the buccula, and notably the large ocelli that occupy half the length of the postocular region. Due to the yellow stripe across the corium, this species resembles two other Australian species, V. fasciata sp. nov. and V. vittata sp. nov. However, the yellow stripe of this species is bright and opaque, unlike that of V. fasciata sp. nov. Additionally, the legs are entirely yellow, and the corium has a small proximal yellow spot. It is easily distinguished from V. pallidipes by the brown membranal veins, the proximal half of the corium being not entirely yellow, and its globose head.

Etymology
The specific epithet is the Latin adjective 'grandioculatus, -a, -um', formed from the prefix 'grandi-' (meaning 'large') combined with the adjective 'oculatus, -a, -um' (meaning 'possessing eyes'), referring to the large ocelli of this species, occupying half the length of the postocular region.
Coloration. Head: dark brown; postocular region with pale patches adjacent to lateral ocellar margin; labial segment I yellowish brown, segments II and III yellow. Thorax (Fig. 4E): coloration as head; scutellum dark with contrasting yellow apical spine. Hemelytron (abducted): clavus dark brown with distal yellow stripe; corium proximal half mostly yellow, distal half dark brown with yellow spot at distal apex; membrane dark with pale V-shaped marking along R and M veins; membranal veins R and M proximally pale. Legs: yellow. Abdomen: dark brown; dorsal laterotergites uniformly colored. integument and vestiture. Head and pronotum: coarsely granulose with dense, short setigerous tubercles interspersed among sparse pubescence; interocular region with setigerous tubercles medially and two pairs paramedially; antennifer with long lateral setigerous tubercle; morphologically ventral surface of labium with sparse, short macrosetae. Thorax: anterolateral angles of pronotal collar with setigerous tubercles; scutellar lateral carinae with short macrosetae on setigerous tubercles. Hemelytron: corium with dense, long macrosetae. Legs: posterior row of protuberances on mid and hind femora with four spines on distal half. Abdomen: ventral surface pubescent.
struCture. Head: globose, about as long as wide; anteocular region about one quarter of head length, shorter than postocular region (measured to anterior margin of neck); postocular region in dorsal view shorter than eye, lateral margins gently rounded; pedicel 1.4-1.6 times length of head width; maxillary plates ellipsoidal, adjacent to and as long as clypeus; apices of maxillary plates in dorsal view diverging; clypeus in dorsal view wider than maxillary plates; clypeal apex bifid, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view nearly straight; eye width in dorsal view about as wide as synthlipsis; eye reaching ventral head margin in lateral view; ocelli large, occupying halflength of postocular region; ventrolateral swelling of buccula with lateral protrusion that surpasses buccular margin, setigerous tubercles along apical margin; labial segment I in lateral view gradually thickens distally, reaching posteroventral eye margin; morphologically dorsal surface of labial segment II curves ventrally, about 0.4 times length of segment I. Thorax: pronotal collar in dorsal view narrow medially with anterolateral angles short, flared laterally; anterior pronotal lobe 0.6-0.7 times length of posterior pronotal lobe, lateral margins almost straight; glabrous markings on pronotum depressed near posterolateral margins; median apodeme depression of pronotum shallow, elongated transversely; scutellar spine long, raised (Fig. 4E); anteriad-directed process of prosternum with paramedial lobes; anterior margin of stridulitrum projected into sharp spine; proepimeron with smoothly rounded posteroventral margin. Legs: fossula spongiosa present on fore and mid legs. Abdomen: anterior margin of tergum III weakly carinulate, remaining terga not carinulate; terga II and III with paired prominent longitudinal carinae, reaching about one-third of tergum III. Pygophore (Fig. 11): transverse bridge with rounded posterior margin; posterior region of ventral surface of pygophore in lateral view with slight swelling; median apical process short, sharply bent posteriorly in lateral view; lateral pygophore margin with protuberance; posterior pygophore margin with clustered macrosetae on protuberance; parameres sinusoidal, apex round, not sharply tapered or expanded. Aedeagus (Fig. 14): endosoma almost entirely covered with spicules; apex of dorsal phallothecal sclerite tongue-shaped in dorsal view; basal plate extension about 5.7 times as long as wide.

Unknown.
Distribution (Fig. 18) This species is known from Queensland and Western Australia.

Remarks
The phylogenetic placement of V. grandioculata sp. nov. within the Australian clade is uncertain (Figs 1-2). The abdomen of the holotype was extracted, but we did not get sufficient DNA in our NGS library for Illumina sequencing.

Diagnosis
This species is most similar to other Southeast Asian Voconia species with a finely granulose head and pronotum and yellow spots on the anteromedial, posteromedial, and distal apex of the abducted corium. Voconia hemera sp. nov. is larger than V. minima sp. nov. (9.2-9.3 mm). It is distinguished from V. ornata, V. nyx sp. nov., and V. bakeri sp. nov. by its uniformly yellow legs and antenna as well as its uniformly brown scutellum and dorsal laterotergites.

Etymology
Named after the Greek primordial goddess of the day, Hemera. Refers to the yellow legs and antennae of this species, which differs from a similar yet darker species, V. nyx sp. nov., named after the goddess of night. A proper noun in apposition.
Coloration. Head: dark brown; postocular region with dark patches adjacent to medial ocellar margin, semicircular pale stripe encircling both ocelli; labium lighter than head, brownish yellow. Thorax: as head; scutellum uniformly dark brown. Hemelytron (abducted): clavus reddish-brown with distal yellow stripe; corium reddish-brown with anteromedial, posteromedial, and distal yellow spots; membrane dark with pale V-shaped marking along R and M veins; membranal veins R and M proximally pale. Legs: yellowish-brown. Abdomen: dark brown. integument and vestiture. Head and pronotum: finely granulose with dense, long macrosetae interspersed among dense pubescence; interocular region with two pairs of macrosetae paramedially; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with dense, short macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae; scutellar lateral carinae with long setation. Hemelytron: corium with sparse long setation. Legs: posterior row of protuberances on mid and hind femora with three large and two small spines. Abdomen: ventral surface pubescent.
struCture. Head: elongate, about 1.3 times as long as wide; anteocular region about one third of head length, shorter than postocular region (measured to anterior margin of neck); postocular region in dorsal view about as long as eye, lateral margins gently rounded; pedicel about 1.2 times length of head width; maxillary plates ellipsoidal, adjacent to and as long as clypeus; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view wider than maxillary plates; clypeal apex round, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view bent anteromedially, subtriangular; eye width in dorsal view narrower than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, surpassing posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.5 times length of segment I. Thorax: pronotal collar in dorsal view narrow medially, projected forward; anterior pronotal lobe about 0.8 times length of posterior pronotal lobe, lateral margins rounded; glabrous markings on pronotum thin, gently depressed; median apodeme depression of pronotum shallow, elongated transversely; scutellar spine long, raised; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with acute protuberance on posteroventral margin. Legs: fossula spongiosa present on fore and mid legs.

Distribution
This species is only known from the type locality on the Philippine island of Luzon.

Revised diagnosis
Distinguished from other species of Voconia by the following combination of characters: short body length (about 6.5 mm); coarsely granulose head and pronotum; maxillary plates are not adjacent to and far surpass clypeal apex; maxillary plates in lateral view are wide, about twice the width of scape; antennifer with long lateral spine; eyes do not reach ventral head margin in lateral view. It is recognized from micropterous individuals of V. schoutedeni and V. monodi by the toothed apex of the maxillary plates.
struCture. Head: scape not reaching head apex; antennifer with long lateral spine; postocular region in dorsal view as long as eye, lateral margins subrectangular; maxillary plates in lateral view about twice the width of scape and apex toothed; eye not reaching ventral head margin in lateral view; ventrolateral swelling of buccula with lateral protrusion that surpasses buccular margin; labial segment I in lateral view straight, surpassing posteroventral eye margin. Thorax: scutellar spine reduced, subhorizontal. Abdomen: anterior margin of terga carinulate; tergum II with paired prominent longitudinal carinae reaching posterior margin of segment.

Unknown.
Distribution (Fig. 17) This species is known from Northwest Africa.

Remarks
The holotype was on loan and not available for study at the time we requested it. Consequently, this description is based on the interpretation of the short differential diagnosis and illustration of the head in lateral view provided by Villiers (1963a). Due to the limited data that we were able to obtain, it was excluded from cladistic analyses, but we are confident that it likely forms a clade with V. schoutedeni and V. tridens sp. nov. because of the unique head morphology, microptery, and Afrotropical distribution.
Voconia isosceles sp. nov. urn:lsid:zoobank.org:act:826EFBC8-FA03-42DA-8E5E- E0D4F29A5B66  Figs 1 -2, 7, 9, 20 Diagnosis This species most closely resembles another Southeast Asian species, V. loki sp. nov., due to the finely granulose head and pronotum, yellow legs and antennae, dark pronotum with contrasting yellow posterior margin, and the proximal half of the corium being entirely yellow, distal half dark with small distal yellow spot. Voconia isosceles sp. nov. is slightly larger (about 8.8 mm long), the membranal veins forming the Cu-An 1 and M-Cu cells are paler than the remainder of the membrane, and the postocular region is dark with a semicircular pale stripe encircling both ocelli.

Etymology
The specific epithet is the Latin adjective 'isosceles, -, -', which has the same meaning as its equivalent geometric term used in English for a triangle having at least two sides of equal length (also known as a golden triangle). Refers to the yellow or golden isosceles triangle on the proximal half of the corium. Coloration. Head: dark brown; postocular region with dark patches adjacent to medial ocellar margin, semicircular pale stripe encircling both ocelli; labium lighter than head, yellow. Thorax: as head; posterior pronotal lobe with posterior margin lighter, yellow; scutellum dark with contrasting yellow apical spine. Hemelytron (abducted): clavus reddish-brown with distal yellow stripe; corium proximal half yellow, distal half reddish-brown with yellow spot at distal apex; membrane brown with pale V-shaped marking along R and M veins; membranal veins forming Cu-An 1 and M-Cu cells paler than remainder of membrane. Legs: yellow. Abdomen: dark brown ventrally; laterotergites light brown. integument and vestiture. Head and pronotum: finely granulose with dense, long macrosetae interspersed among dense pubescence; interocular region with two pairs of macrosetae paramedially; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with dense, short macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae; scutellar lateral carinae with long setation. Hemelytron: corium with dense long setation. Legs: posterior row of protuberances on mid and hind femora with three large spines on distal half. Abdomen: ventral surface pubescent, long macrosetae interspersed. struCture. Head: elongate, about 1.3 times as long as wide; anteocular region about one third of head length, about as long as postocular region (measured to anterior margin of neck); postocular region in dorsal view about as long as eye, lateral margins gently rounded; pedicel about 1.4 times length of head width; maxillary plates ellipsoidal, adjacent to and as long as clypeus; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view wider than maxillary plates; clypeal apex round, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view bent anteromedially, subtriangular; eye width in dorsal view narrower than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, surpassing posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.3 times length of segment I. Thorax: pronotal collar in dorsal view narrow medially with anterolateral angles short, projected forward; anterior pronotal lobe about 0.6 times length of posterior pronotal lobe, lateral margins rounded; glabrous markings on pronotum conspicuous and not depressed; median apodeme depression of pronotum shallow, elongated transversely; scutellar spine long, raised; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with acute protuberance on posteroventral margin. Legs: fossula spongiosa present on fore and mid legs. Abdomen: anterior margin of terga prominently carinulate; terga II and III with paired prominent longitudinal carinae, almost reaching posterior margin of tergum III.

Unknown.
Distribution (Fig. 20) This species is only known from the type locality in the Philippines.

Remarks
Based on the record of a Miridae specimen from the same collection event as the holotype, Mt Pomalihi is presumably in Mt Balatukan Range Natural Park. The type locality is also shared with V. bakeri sp. nov. Though V. isosceles sp. nov. and V. loki sp. nov. closely resemble each other, we are treating them as separate species because they were not recovered as closely related taxa in our analyses (Figs 1-2). This hypothesis should be further tested with additional material. sp. nov. urn:lsid:zoobank.org:act:D92E888A-8436-42C2-8316-881775941725 Figs 1-2, 7, 9, 12, 14, 20 Diagnosis This species is distinguishable by the elongate Cu-An 1 cell (reaching at least half the length of M-Cu cell) and short labial segment I (not surpassing ventral eye margin in lateral view). This species most closely resembles three other Southeast Asian species, V. isosceles sp. nov., V. lasiosoma sp. nov., and V. loki sp. nov., due to the finely granulose head and pronotum, long setation of the body, semicircular pale stripe encircling both ocelli, and the dark pronotum with contrasting pale posterior margin. However, in V. laosensis sp. nov. this pale posterior margin occupies almost the entire posterior pronotal lobe.

Etymology
A proper noun with the Latin adjectival suffix '-ensis' meaning 'belonging to', referring to the country of the type locality, Laos. To be treated as an adjective.
Coloration. Head: dark brown, maxillary plates and clypeus lighter; postocular region with dark patches adjacent to medial ocellar margin, semicircular pale stripe encircling both ocelli; labium lighter than head, yellow. Thorax: as head; posterior pronotal lobe lighter than anterior lobe; scutellum dark with brown apical spine. Hemelytron (abducted): clavus dark reddish-brown with proximal and distal yellow stripe; corium dark reddish-brown with anteroproximal yellow stripe, posteromedial and distal yellow spots; membrane uniformly dark; pale V-shaped marking along R and M veins. Legs: yellowish-brown. Abdomen: dark brown, laterotergites light brown. integument and vestiture. Head and pronotum: finely granulose with sparse, long macrosetae interspersed among sparse pubescence; interocular region with two pairs of macrosetae paramedially; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with dense, short macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae; scutellar lateral carinae with long setation. Hemelytron: corium with sparse long setation. Legs: posterior row of protuberances on mid and hind femora with three large and one small spines on distal half. Abdomen: ventral surface pubescent.
struCture. Head: elongate, about 1.2 times as long as wide; anteocular region about one third of head length, about as long as postocular region (measured to anterior margin of neck); postocular region in dorsal view shorter than eye, lateral margins gently rounded; pedicel about 1.2 times length of head width; maxillary plates ellipsoidal, adjacent to and reaching apex of clypeus; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view wider than maxillary plates; clypeal apex round, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view nearly straight; eye width in dorsal view narrower than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, not reaching posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.6 times length of segment I. Thorax: pronotal collar in dorsal view narrow medially, anterolateral angles short, flared laterally; anterior pronotal lobe about 0.5 times length of posterior pronotal lobe, lateral margins rounded; glabrous markings on pronotum thin and not deeply depressed; median apodeme depression of pronotum shallow, elongated longitudinally; scutellar spine long, raised; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum not elongated into protuberance. Hemelytron: Cu-An 1 cell elongate, reaching at least half the length of M-Cu cell. Legs: fossula spongiosa present on fore and mid legs. Pygophore (Fig. 12): transverse bridge with rounded posterior margin; posterior region of ventral surface of pygophore in lateral view straight, not swollen; short median apical process upright in lateral view; lateral pygophore margin with protuberance; posterior pygophore margin with clustered macrosetae on protuberance; parameres sinusoidal, apex tapered into squarely rounded tip. Aedeagus (Fig. 14): endosoma almost entirely covered with spicules; apex of dorsal phallothecal sclerite tongueshaped in dorsal view; basal plate extension 4.4 times as long as wide.

Unknown.
Distribution (Fig. 20) This species is only known from the type locality in Laos. This is the only known species of Voconia on mainland Southeast Asia.

Remarks
The pygophore was dissected from this pinned holotype and used to extract DNA, for which we acquired the lowest DNA concentration (0.124 ng/µL) and did not sequence.
Voconia lasiosoma sp. nov. urn:lsid:zoobank.org:act:C5495703-AEE2-49FC-8B08- 099ED0F398D3  Figs 1 -2, 7, 9, 12, 14, 20 Diagnosis Recognized from other Voconia by the densely long setation on the pronotum, corium, legs, and abdomen. This species most closely resembles two other Southeast Asian species, V. isosceles sp. nov. and V. loki sp. nov. due to the finely granulose head and pronotum and dark pronotum with contrasting pale posterior margin. While V. loki sp. nov. is also hairy, V. lasiosoma sp. nov. is larger (about 9.4 mm) than these European Journal of Taxonomy 788: 1-95 (2022) 58 two species; legs, antennae, and corium are dark yellowish-brown rather than yellow; dark portions of corium and clavus are black rather than reddish-brown; Cu-An 1 cell stout (less than half the length of M-Cu cell); and proximal half of corium not entirely yellow (abducted corium with anteroproximal yellowish-brown stripe and posteromedial yellow spot).

Etymology
Derived from the Latinized Greek prefix 'lasio-' meaning 'hairy' combined with the Latinized Greek noun '-soma' meaning 'body'. Refers to the long macrosetae covering the body and hemelytra. A noun in apposition.
Coloration. Head: dark brown; postocular region with dark patches adjacent to medial ocellar margin, semicircular pale stripe encircling both ocelli; labium lighter than head, yellow. Thorax: anterior pronotal lobe darker than head and posterior pronotal lobe; posterior pronotal lobe with dark yellowish-brown gradient; scutellum dark with contrasting dark yellowish-brown apical spine. Hemelytron (abducted): clavus black with proximal dark yellowish-brown spot and distal yellow stripe; corium black with anteroproximal dark yellowish-brown stripe, posteromedial and distal yellow spots; membrane mostly dark with pale stripe along most of M vein; membranal veins R and M proximally pale. Legs: yellowishbrown. Abdomen: dark brown. integument and vestiture. Head and pronotum: finely granulose with dense, long macrosetae interspersed among sparse pubescence; interocular region with two pairs of macrosetae paramedially; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with dense, short macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae; scutellar lateral carinae with long setation. Hemelytron: corium with dense long setation. Legs: posterior row of protuberances on mid and hind femora with three large and three small spines on distal half. Abdomen: ventral surface pubescent, long macrosetae interspersed.
struCture. Head: elongate, about 1.3 times as long as wide; anteocular region about one third of head length, shorter than postocular region (measured to anterior margin of neck); postocular region in dorsal view shorter than eye, lateral margins gently rounded; length of pedicel subequal to head width; maxillary plates ellipsoidal, adjacent to and shorter than clypeus; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view wider than maxillary plates; clypeal apex round, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view nearly straight; eye width in dorsal view about as wide as synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, surpassing posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.5 times length of segment I. Thorax: pronotal collar in dorsal view narrow medially, with anterolateral angles short, flared laterally; anterior pronotal lobe about 0.7 times length of posterior pronotal lobe, lateral margins rounded; glabrous markings on pronotum conspicuous and not depressed; median apodeme depression of pronotum deep, elongated longitudinally; scutellar spine long, subhorizontal; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum projected into small protuberance; proepimeron with acute protuberance on posteroventral margin. Legs: fossula spongiosa present on fore and mid legs. Abdomen: anterior margin of terga prominently carinulate; terga II and III with paired prominent longitudinal carinae, reaching posterior margin of tergum III. Pygophore (Fig. 12): transverse bridge with rounded posterior margin; posterior region of ventral surface of pygophore in lateral view straight, not swollen; short median apical process, upright in lateral view; lateral pygophore margin with protuberance; posterior pygophore margin with clustered macrosetae on protuberance; parameres sinusoidal, apex tapered into squarely rounded tip. Aedeagus ( Fig. 14): endosoma almost entirely covered with spicules; apex of dorsal phallothecal sclerite tongueshaped in dorsal view; basal plate extension 5.6 times as long as wide.

Unknown.
Distribution (Fig. 20) This species is only known from the type locality in Sarawak, Malaysia.

Remarks
The holotype was preserved in ethanol when the abdomen was dissected for DNA extraction. We acquired sufficient DNA in our NGS library for Illumina sequencing for future phylogenomic studies, referred to as DNA voucher R_CW 5472.

Diagnosis
Distinguished from most species of Voconia by the pale membranal veins. This species is most similar to two other Oriental species, V. decorata sp. nov. and V. coronata sp. nov. It is distinguishable from V. decorata sp. nov. by its distribution in Sumatra; the dark brown, almost black coloration of the body; and uniformly brown scutellum. V. lirophleps sp. nov. is about 2 mm shorter than V. coronata sp. nov. It is also distinguishable by the yellow spots adjacent to the lateral ocellar margin; slightly more globose head (about 1.2 times as long as wide); pedicel much shorter (about 1.1 times length of head width); maxillary plates converge in dorsal view, postocular region is shorter than eye; narrow pronotal collar; short anterior pronotal lobe relative to posterior pronotal lobe (about 2.2 times shorter than posterior lobe); conspicuous distal yellow spot on the corium; and only three large spines and two small spines on the posterior row of spines of the hind leg.

Etymology
Derived from the Latinized Greek prefix 'liro-' meaning 'pale' combined with the Latinized Greek noun 'phleps' meaning 'vein'. Refers to the prominent pale coloration of the membranal veins. To be treated as a noun in apposition, ending not to be changed.
Coloration. Head: dark brown; postocular region uniformly dark with pale patches adjacent to lateral ocellar margin; labium nearly as dark as head. Thorax: coloration as head; scutellum uniformly dark brown. Hemelytron ( Fig. 4B; abducted): clavus dark reddish-brown with distal yellow stripe; corium dark reddishbrown with anteroproximal yellow stripe, posteromedial and distal yellow spots; membrane uniformly dark; membranal veins forming Cu-An 1 and M-Cu cells paler than remainder of membrane. Legs: dark brown, femora darker. Abdomen (Fig. 4B): dark brown; dorsal laterotergites dark with yellow spots anteriorly. integument and vestiture. Head and pronotum: finely granulose with sparse, short macrosetae interspersed among sparse pubescence; interocular region with two pairs of macrosetae paramedially; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with sparse, long macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae; scutellar lateral carinae with long setation. Hemelytron: corium with sparse long setation. Legs: posterior row of protuberances on mid and hind femora with three large and three small spines on distal half. Abdomen: ventral surface pubescent.
struCture. Head: elongate, 1.2 times as long as wide; anteocular region about one quarter of head length, about as long as postocular region (measured to anterior margin of neck); postocular region in dorsal view shorter than eye, lateral margins gently rounded; pedicel about 1.1 times length of head width; maxillary plates ellipsoidal, adjacent to and as long as clypeus; apices of maxillary plates in dorsal view converging; clypeus in dorsal view narrower than maxillary plates; clypeal apex round, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view bent anteromedially, subtriangular; eye width in dorsal view about as wide as synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, reaching posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.5 times length of segment I. Thorax: pronotal collar in dorsal view narrow medially with anterolateral angles short, projected forward; anterior pronotal lobe about 0.5 times length of posterior pronotal lobe, lateral margins rounded; glabrous markings on pronotum deeply depressed; median apodeme depression of pronotum shallow, elongated longitudinally; scutellar spine long, raised; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with acute protuberance on posteroventral margin. Legs: fossula spongiosa present on fore and mid legs.

Unknown.
Distribution (Fig. 20) This species is only known from the type locality in North Sumatra.

Remarks
Despite the holotypes of V. coronata sp. nov. and V. lirophleps sp. nov. being collected within about 106 km of each other, there are several differences, particularly in size. Among our larger series, conspecific Voconia males do not vary much in size, and V. lirophleps sp. nov. is about 2 mm shorter than V. coronata sp. nov.; thus, we are keeping them as separate species.

Diagnosis
This species most closely resembles two other Southeast Asian species, V. isosceles sp. nov. and V. lasiosoma sp. nov., due to the finely granulose head and pronotum; legs and antennae yellow; pronotum dark with contrasting yellow posterior margin; and relatively denser and longer setation. Voconia loki sp. nov. is smaller (about 8.3 mm long) than these two species. It is also recognized from V. isosceles sp. nov. by the stout Cu-An 1 cell (less than half the length of the M-Cu cell), brown membranal veins that form the Cu-An 1 and M-Cu cells, and the postocular region brown having dark patches adjacent to the medial ocellar margin. Apart from its smaller size, this species is distinguished from V. lasiosoma sp. nov. by the almost entirely yellow proximal half of the corium, reddish-brown distal half of the corium instead of black, and yellow legs instead of a darker yellowish-brown.

Etymology
Named after the cunning trickster from Norse mythology and from the Marvel Comics' character, Loki, since this specimen deceived and tricked authors in a previous study (Hwang & Weirauch 2012) who misidentified it as "Kayanocoris wegneri" (V. wegneri comb. nov.). A proper noun in apposition. Coloration. Head (Fig. 3H): light brown; clypeus yellow; postocular region with dark patches adjacent to medial ocellar margin; labium lighter than head, yellow. Thorax: pronotum darker than head; posterior pronotal lobe with posterior margin lighter; scutellum dark with contrasting yellow apical spine. Hemelytron (abducted): clavus reddish-brown with distal yellow stripe; proximal half of corium mostly yellow with small black spot, distal half reddish-brown with distal yellow spot; membrane dark with proximally pale V-shaped marking along R and M veins; membranal veins R and M proximally pale, veins forming Cu-An 1 cell pale. Legs: yellow. Abdomen: dark brown; dorsal laterotergites yellow. integument and vestiture. Head and pronotum (Fig. 3H): finely granulose with dense, long macrosetae interspersed among sparse pubescence; interocular region with two pairs of macrosetae paramedially; antennifer with strong macroseta, base not protruded laterally; morphologically ventral surface of labium with dense, short macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae; scutellar lateral carinae with long setation. Hemelytron: corium with dense long setation. Legs: posterior row of protuberances on mid and hind femora with three large and three small spines on distal half. Abdomen: ventral surface pubescent, long macrosetae interspersed.

Holotype
struCture. Head (Fig. 3H): elongate, about 1.2 times as long as wide; anteocular region about one third of head length, about as long as postocular region (measured to anterior margin of neck); postocular region in dorsal view as long as eye, lateral margins gently rounded; pedicel about 0.9 times length of head width; maxillary plates ellipsoidal, adjacent to and shorter than clypeus; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view wider than maxillary plates; clypeal apex round, narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view bent anteromedially, subtriangular; eye width in dorsal view wider than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, surpassing posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.5 times length of segment I. Thorax: pronotal collar in dorsal view narrow medially with anterolateral angles short, projected forward; anterior pronotal lobe about 0.7 times length of posterior pronotal lobe, lateral margins almost straight; glabrous markings on pronotum conspicuous and not depressed; median apodeme depression of pronotum deep, elongated longitudinally; scutellar spine long, raised; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with acute protuberance on posteroventral margin. Legs: fossula spongiosa present on fore and mid legs.

Unknown.
Distribution (Fig. 20) This species is only known from the type locality in Brunei on the island of Borneo.

Diagnosis
One of two New World species of Voconia. Recognized by the coarsely granulose head and prothorax; wide anteocular region; morphologically ventral surface of labium with sparsely long macrosetae; buccula with lateral tubercles protruding past buccular margin; and yellow antero-and posteromedial spots on the abducted corium.

Etymology
The specific epithet is the Latin adjective 'mexicanus, -a, -um', meaning 'from or pertaining to Mexico', referring to the country of the type locality. Coloration. Head: uniformly dark brown; labium lighter than head, yellow brown. Thorax: as head; posterior pronotal lobe with posterior margin lighter; scutellum dark with brown apical spine. Hemelytron (abducted): clavus dark reddish-brown with distal yellow stripe; corium dark reddish-brown with anteromedial and posteromedial yellow spots; membrane dark with pale V-shaped marking along R and M veins; membranal veins R, M, and An 1 proximally pale. Legs: dark brown. Abdomen: dark brown, laterotergites light brown. integument and vestiture. Head and pronotum: coarsely granulose with sparse pubescence on setigerous tubercles; interocular region unarmed with macrosetae; antennifer with 1-3 short lateral setigerous tubercles; morphologically ventral surface of labium with sparse, long macrosetae. Thorax: anterolateral angles of pronotal collar with setigerous tubercles. Hemelytron: corium with dense, short setation. Legs: posterior row of protuberances on mid and hind femora with three large spines on distal half and two to five small spines. Abdomen: ventral surface pubescent.
struCture. Head: elongate, 1.2-1.3 times as long as wide; anteocular region about one third of head length, longer than postocular region (measured to anterior margin of neck); postocular region in dorsal view about as long as eye, lateral margins gently rounded; pedicel 1.4-1.9 times length of head width; maxillary plates ellipsoidal, adjacent to and longer than clypeus; apices of maxillary plates in dorsal view directed straight or diverging; clypeus in dorsal view narrower than maxillary plates; clypeal apex round, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view curved and smoothly rounded anteriorly; eye width in dorsal view narrower than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula with lateral protrusion that surpasses buccular margin; labial segment I in lateral view straight, reaching or slightly shorter than posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.4 times length of segment I. Thorax: pronotal collar in dorsal view narrow medially with anterolateral angles short, projected forward; anterior pronotal lobe about 0.5-0.6 times length of posterior pronotal lobe, lateral margins rounded; glabrous markings on pronotum thin and gently depressed; median apodeme depression of pronotum deep, circular; scutellar spine long, subhorizontal or raised; anteriad-directed process of prosternum with paramedial lobes (Fig. 3I); anterior margin of stridulitrum projected into small protuberance (Fig. 3I); proepimeron with acute protuberance on posteroventral margin. Legs: fossula spongiosa present on fore and mid legs. Abdomen: anterior margin of terga weakly carinulate; terga II and III with paired prominent longitudinal carinae, reaching about one-third of tergum III. Pygophore (Fig. 12): transverse bridge with rounded posterior margin; posterior region of ventral surface of pygophore in lateral view with large swelling; short median apical process sharply bent posteriorly in lateral view; lateral pygophore margin with protuberance; posterior pygophore margin with clustered macrosetae on flat margin; parameres sinusoidal, apex tapered into squarely rounded tip. Aedeagus (Fig. 14): endosoma almost entirely covered with spicules; apex of dorsal phallothecal sclerite rounded in dorsal view; basal plate extension about 4.3 times as long as wide.

Female
Distinguished from males by the following. Body length: about 12.2 mm. Head: elongate, about 1.2 times as long as wide; anteocular region shorter than postocular region (measured to anterior margin of neck); with sparse, short macrosetae on setigerous tubercles and sparse pubescence on setigerous tubercles; pedicel about 1.6 times length of head width; apices of maxillary plates in dorsal view directed straight; labium lighter than head, brown; labial segment I in lateral view surpassing posteroventral eye margin. Thorax: pronotal collar in dorsal view narrow, almost absent medially; anterolateral angles of pronotum long, projected forward. Hemelytron: corium dark with thick medial yellow stripe; with dense, long macrosetae. Legs: posterior row of protuberances on mid and hind femora with three large spines on distal half and many small spines (Fig. 4I); fossula spongiosa absent or vestigial on mid leg. Distribution (Fig. 19) This species is distributed in Southern Mexico and Honduras.

Remarks
The female from Honduras is considerably different from the males from Mexico as the description states above. We keep them as a single species until this hypothesis can be further tested with additional material. Specimen voucher UCR_ENT 00004570 was mis-labelled as collected "4 mi N of Cardale", though it should be "4 mi N of José Cardel" as recorded by other collectors on the same trip.

Diagnosis
This species is recognized from other Philippine species such as V. bakeri sp. nov. and V. nyx sp. nov. by its short body length (about 8.5 mm long); dark tibiae with proximal and distal ends yellow; and abducted corium with yellow stripe occupying half of the anteroproximal margin and expanding medially to become adjacent to posteromedial yellow spot.

Etymology
The specific epithet is the Latin adjective 'minimus, -a, -um', meaning 'smallest'. Refers to the fact that this is among the smallest species of Voconia. Coloration. Head: dark brown; postocular region with pale patches adjacent to lateral ocellar margin; labium nearly as dark as head. Thorax: as head; scutellum dark, tip of apical spine contrasting yellow. Hemelytron (abducted): clavus dark reddish-brown with distal yellow stripe; corium dark reddishbrown with anteroproximal yellow stripe, posteromedial and distal yellow spots; membrane dark with pale V-shaped marking along R and M veins; membranal veins R and M proximally pale. Legs: dark brown, proximal and distal ends of tibiae and tarsi yellow. Abdomen: dark brown. integument and vestiture. Head and pronotum: finely granulose with sparse, long macrosetae interspersed among dense pubescence; interocular region with two pairs of macrosetae paramedially; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with dense, short macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae; scutellar lateral carinae with long setation. Hemelytron: corium with sparse, long setation. Legs: posterior row of protuberances on mid and hind femora with three large and two small spines on distal half. Abdomen: ventral surface pubescent.
struCture. Head: elongate, about 1.4 times as long as wide; anteocular region about one quarter of head length, shorter than postocular region (measured to anterior margin of neck); postocular region in dorsal view about as long as eye, lateral margins gently rounded; pedicel about 1.2 times length of head width; maxillary plates ellipsoidal, adjacent to and longer than clypeus; apices of maxillary plates in dorsal view converging; clypeus in dorsal view about as wide as maxillary plates; clypeal apex round, narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view nearly straight; eye width in dorsal view about as wide as synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, surpassing posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.5 times length of segment I. Thorax: pronotal collar in dorsal view narrow medially with anterolateral angles short, projected forward; anterior pronotal lobe about 0.6 times length of posterior pronotal lobe, lateral margins rounded; glabrous markings on pronotum thin and not deeply depressed; median apodeme depression of pronotum shallow, elongated transversely; scutellar spine long, raised; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with acute protuberance on posteroventral margin. Legs: fossula spongiosa present on fore and mid legs.

Female
Unknown.
Distribution (Fig. 8) This species is only known from the type locality on the Philippine island of Mindanao.

Revised diagnosis
Distinguished from other species of Voconia by the following combination of characters: short body length (about 7.0 mm long); coarsely granulose head and pronotum; maxillary plates not adjacent to and far surpass clypeal apex; maxillary plates in lateral view wide, about twice the width of the scape and not toothed apically; antennifer with long lateral spine. It is recognized from the micropterous V. schoutedeni by the paired, short, and not prominent longitudinal carinae on tergum II, not reaching the posterior margin of the segment; reliefs of the anterior lobe of the pronotum less strong, effaced in front; and indistinct ocelli. Coloration. Head: yellowish brown. Thorax: Pronotum yellowish in front and on the sides, brown on the rest of the surface; scutellum brown. Micropterous wing: yellowish. Legs: brown with proximal and distal part of femur and tarsus testaceous. Abdomen: dark brown, dorsal laterotergites with anterior half yellow. integument and vestiture. Head and pronotum: coarsely granulose with setigerous tubercles; interocular region with two pairs of large setigerous tubercles paramedially; antennifer with long lateral spine. Legs: posterior row of protuberances on mid and hind femora with three or four spines. struCture. Head: scape not reaching head apex and longer than clypeus; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view narrower than maxillary plates; clypeal apex round, narrowed; eye width in dorsal view narrower than synthlipsis; ocelli absent; postocular region in dorsal view longer than eye, lateral margins subrectangular; labial segment I in lateral view straight, surpassing posteroventral eye margin; pronotal anterolateral angles long, flared laterally; scutellar spine reduced, slightly raised. Abdomen: anterior margin of terga carinulate; tergum II with paired, short, and not prominent longitudinal carinae.

Unknown.
Distribution (Fig. 17) This species is known from Northwest Africa.

Remarks
Specimens were on loan and not available for this study. Consequently, this description and localities were inferred from the French description, illustrations, and key provided by Villiers (Villiers 1963b(Villiers , 1964. Villiers (1963b) includes an additional specimen from Guinea that was not treated as type material that we could not examine and have limited information for. We therefore refrain from including it. A drawer image of two specimens from the BMNH in 2011 suggests that this species may be distributed in Sierra Leone (collection event: 11 May 1926) and Nigeria, but their identification is difficult to confirm without better images and without the holotype. (Schouteden, 1929) comb. nov. Figs 1-2, 8, 10, 17 Microvarus motoensis Schouteden, 1929: 252. Microvarus inermiceps Villiers, 1964. Synonymized by Villiers (1976: 171).

Revised diagnosis
Distinguished from other species in Voconia by its finely granulose head; short labial segment I that does not reach posteroventral eye margin; anterior margin of buccula with dense, long macrosetae; and stout body. This species most closely resembles V. conradti and V. smithae sp. nov. in its stout body plan and Afrotropical distribution. It is easily recognized from these two species by the absent or vestigial fossula spongiosa on the midleg and its distinct coloration: uniformly brown scutellum, abducted corium with an anteromedial yellow spot, and uniformly brown dorsal laterotergites.
Coloration. Head: dark brown; labium nearly as dark as head. Thorax: coloration as head; posterior pronotal lobe with posterior margin lighter; scutellum uniformly dark brown. Hemelytron (abducted): clavus dark reddish-brown with distal yellow stripe; corium dark reddish-brown with anteromedial and posteromedial yellow spots; membrane dark with pale V-shaped marking along R and M veins; membranal veins proximally pale. Legs: yellowish-brown with distal half of femora dark brown. Abdomen: dark brown; dorsal laterotergites uniformly dark brown. integument and vestiture. Head and pronotum: finely granulose with sparse, short macrosetae interspersed among dense pubescence; antennifer with macroseta, base not protruded; morphologically ventral surface of labium with dense, short macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae. Hemelytron: corium with sparse long setation. Legs: posterior row of protuberances on mid and hind femora with three large spines on distal half, middle spine much longer. Abdomen: ventral surface pubescent.
struCture. Head: elongate, about 1.2 times as long as wide; anteocular region about one quarter of head length, shorter than postocular region (measured to anterior margin of neck); postocular region in dorsal view shorter than eye, lateral margins gently rounded; pedicel about 1.5 times length of head width; maxillary plates ellipsoidal, adjacent to and longer than clypeus; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view wider than maxillary plates; clypeal apex round, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view bent anteromedially, subtriangular; eye width in dorsal view wider than synthlipsis, eye not reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, not reaching posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.7 times length of segment I. Thorax: pronotal collar wide medially with anterolateral angles short, projected forward; anterior pronotal lobe about 0.7 times length of posterior pronotal lobe, lateral margins almost straight; glabrous markings on pronotum thin and not deeply depressed; median apodeme depression of pronotum deep, circular; scutellar spine long, raised; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with smoothly rounded posteroventral margin. Legs: fossula spongiosa present on fore leg, absent or vestigial on mid leg.

Unknown.
Distribution (Fig. 17) Only known from Moto and Garamba National Park (Villiers 1964) in the Haut-Uélé province of the Democratic Republic of the Congo.

Diagnosis
This species is most similar to other Southeast Asian species of Voconia with a finely granulose head and pronotum and abducted corium with yellow spots on the posteromedial and distal apex. Despite the similarity in color, V. nyx sp. nov. is larger than V. minima sp. nov. (about 10.0 mm long) and has uniformly brown legs. It is distinguished from V. bakeri sp. nov. by its uniformly brown dorsal laterotergites, pale stripe along the posterior margin of the pronotum, and the abducted corium with an anteroproximal yellowish-brown stripe.

Etymology
Named after the Greek primordial goddess of the night, Nyx. Refers to the dark legs and antennae of this species, which differ from those of a similar yet lighter colored species, Voconia hemera sp. nov., named after the goddess of day. A proper noun in apposition. Coloration. Head: dark brown; postocular region with pale patches adjacent to lateral ocellar margin; labium lighter than head, brown. Thorax (Fig. 4F): as head; posterior pronotal lobe with posterior margin lighter; scutellum uniformly dark brown. Hemelytron (abducted): clavus dark reddish-brown with distal yellow stripe; corium dark reddish-brown with yellowish-brown stripe on anteroproximal margin, posteromedial and distal yellow spots; membrane dark with pale V-shaped marking along R and M veins; membranal veins R and M pale. Legs: dark reddish-brown. Abdomen: dark reddish-brown; dorsal laterotergites dark brown. integument and vestiture. Head and pronotum: finely granulose with sparse, long macrosetae interspersed among dense pubescence; interocular region with two pairs of macrosetae paramedially; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with sparse, long macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae; scutellar lateral carinae with long setation. Legs: posterior row of protuberances on mid and hind femora with three large spines on distal half and three or fewer small spines. Abdomen: ventral surface pubescent. struCture. Head: elongate, about 1.3 times as long as wide; anteocular region about one third of head length, about as long as postocular region (measured to anterior margin of neck); postocular region in dorsal view shorter than eye, lateral margins gently rounded; pedicel about 1.2 times length of head width; maxillary plates ellipsoidal, adjacent to and longer than clypeus; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view about as wide as maxillary plates; clypeal apex round, narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view bent anteromedially, subtriangular; eye width in dorsal view about as wide as synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, not reaching posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.6 times length of segment I. Thorax: pronotal collar in dorsal view narrow medially with anterolateral angles short, projected forward; anterior pronotal lobe about 0.6 times length of posterior pronotal lobe, lateral margins rounded; glabrous markings on pronotum thin and not deeply depressed; median apodeme depression of pronotum shallow, elongated transversely; scutellar spine long, raised (Fig. 4F); anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with acute protuberance on posteroventral margin. Legs (Fig. 4G-H): fossula spongiosa present on fore and mid legs. Pygophore (Fig. 12): transverse bridge with triangular posterior margin; posterior region of ventral surface of pygophore in lateral view with large swelling; short median apical process bent posteriorly in lateral view; lateral pygophore margin with protuberance; posterior pygophore margin with clustered macrosetae on protuberance; parameres sinusoidal, apex tapered into squarely rounded tip. Aedeagus (Fig. 14): endosoma almost entirely covered with spicules; apex of dorsal phallothecal sclerite tongue-shaped in dorsal view; basal plate extension 4.6 times as long as wide.

Unknown.
Distribution (Fig. 17) This species is only known from the type locality on the Philippine island of Luzon.

Remarks
The holotype locality of Mt Maquiling is about 10 km from the holotype locality of V. hemera sp. nov. at Los Baños.

Diagnosis
Distinguished from other species of Voconia by the following combination of characters: slender and short body (~7.8-8.7 mm long); proepimeron with protuberance on posteroventral margin; legs yellow with distal half of femora brown; scutellum dark with contrasting yellow apical spine; and dark abducted corium with anteromedial, posteromedial, and distal yellow spots.

Type material
Lectotype ( Coloration. Head: light brown; postocular region with pale patches adjacent to lateral ocellar margin; labium lighter than head, brown to yellow. Thorax: as head; scutellum dark with contrasting yellow apical spine. Hemelytron (abducted): clavus reddish-brown with distal yellow stripe; corium reddishbrown with anteromedial, posteromedial, and distal yellow spots; membrane dark with pale V-shaped marking along R and M veins; membranal veins R and M proximally pale. Legs: yellow, distal half of femora brown. Abdomen: light brown; dorsal laterotergites dark brown with faint pale bands on posterior margins. integument and vestiture. Head and pronotum: finely granulose with dense, short macrosetae interspersed among dense pubescence; interocular region with two pairs of macrosetae paramedially; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with sparse, short macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae. Hemelytron: corium with dense, long macrosetae. Legs: posterior row of protuberances on mid and hind femora with three large spines on distal half and many small spines. Abdomen: ventral surface pubescent.
struCture. Head: elongate, about 1.3 times as long as wide; anteocular region about one third of head length, about as long as postocular region (measured to anterior margin of neck); postocular region in dorsal view about as long as eye, lateral margins gently rounded; pedicel 1.3-1.4 times length of head width; maxillary plates ellipsoidal, adjacent to and longer than clypeus; apices of maxillary plates in dorsal view directed straight or diverging; clypeus in dorsal view about as wide as maxillary plates; clypeal apex round, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view nearly straight; eye width in dorsal view narrower than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula with lateral protrusion that surpasses buccular margin, setigerous tubercles along apical margin; labial segment I in lateral view straight, reaching posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.5 times length of segment I. Thorax: pronotal collar in dorsal view narrow medially with anterolateral angles short, projected forward; anterior pronotal lobe 0.7-0.8 times length of posterior pronotal lobe, lateral margins almost straight; glabrous markings on pronotum thin and deeply depressed; median apodeme depression of pronotum deep, circular; scutellar spine long, subhorizontal; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with acute protuberance on posteroventral margin. Legs: fossula spongiosa present on fore and mid legs.

Female
Mostly as male, distinguished from males by the following. Body length: 7.8-8.5 mm. Head: elongate, 1.3-1.4 times as long as wide; pedicel 1.3-1.5 times length of head width. Legs: fossula spongiosa absent or vestigial on mid leg. Distribution (Fig. 20) Distributed in India and Sri Lanka.

Remarks
Originally described as Gerbelius ornatus based on an unspecified number of specimens (syntypes) from Sri Lanka (Distant 1903a). Cladistic analyses (Weirauch 2008;present study: Figs 1-2) confirm that this species is nested within Voconia, and hence it is hereby transferred to the latter genus. The abdomen of pinned specimen USI: AMNH_PBI 00170697 was removed for DNA extraction, but insufficient DNA was acquired in the NGS library for Illumina sequencing.
A single male pertaining to the type series (Fig. 8), here designated as the lectotype, was examined from a unit tray image at the BMNH, along with six other specimens for which we can only distinguish the locality for one specimen as Halwaldi, India and is not included in our distribution map nor material examined (Fig. 20). While the locality and date for the lectotype are not specified, we are confident that this specimen was studied by Distant. Kirby (1891) notes that the collector E.E. Green shared his collection of Hemiptera with the BMNH, most of which were collected at Pundaloya and could be inferred as the locality where no locality is mentioned. Less likely localities include Nitagala, Nawalapitya, Kandy, Colombo, or other. Stål, 1866 Figs 1-2, 3C, 8, 10, 13-14, 18 Voconia pallidipes Stål, 1866b: 165.

Diagnosis
Distinguished from most other species of Voconia by the tuberculate and coarsely granulose head and pronotum, tuberculate lateral protrusion of the buccula, the anterolateral angles of the pronotal collar with many (more than two) setigerous tubercles, and the abducted corium with the proximal half entirely yellow and distal half dark with distal yellow spot. This species resembles another Australian species, V. grandioculata sp. nov., in coloration but is easily distinguished by its larger size (10.5-11.3 mm long), smaller eyes and ocelli (ocellus does not occupy half-length of postocular region), M-Cu cell partially or entirely yellow, membranal veins forming Cu-An 1 and M-Cu cells mostly or entirely yellow, and yellow clavus with a dark medial spot.

Type material
Lectotype (present designation) AUSTRALIA • 1 ♀; Queensland, Moreton Bay; Stevens leg.; NHRS-GULI000007827. Coloration. Head (Fig. 3C): dark brown; postocular region with pale patches adjacent to lateral ocellar margin; labium lighter than head, yellow. Thorax: as head; scutellum dark with contrasting yellow apical spine. Hemelytron (abducted): clavus yellow with dark medial spot; corium proximal half yellow, distal half dark reddish-brown with yellow spot at distal apex; membrane dark with pale V-shaped marking along R and M veins, M-Cu cell entirely or partially yellow; membranal veins forming Cu-An 1 and M-Cu cells mostly or entirely yellow. Legs: yellow. Abdomen: light brown, laterotergites uniformly yellow. integument and vestiture. Head (Fig. 3C) and pronotum: coarsely granulose with dense, short setigerous tubercles interspersed among sparse pubescence; antennifer with long and 1-2 adjacent short lateral setigerous tubercles; morphologically ventral surface of labium with sparse, short macrosetae. Thorax: anterolateral angles of pronotal collar with setigerous tubercles; scutellar lateral carinae with short macrosetae on setigerous tubercles. Hemelytron: corium with short macrosetae. Legs: posterior row of protuberances on mid and hind femora with four or more large spines on distal half. Abdomen: ventral surface pubescent.
struCture. Head (Fig. 3C): elongate, 1.2-1.3 times as long as wide; anteocular region about one third of head length, about as long as postocular region (measured to anterior margin of neck); postocular region in dorsal view longer than eye, lateral margins gently rounded; pedicel 1.4-1.6 times length of head width; maxillary plates ellipsoidal, adjacent to and longer than clypeus; apices of maxillary plates in dorsal view diverging; clypeus in dorsal view about as wide as maxillary plates; clypeal apex bifid, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view nearly straight; eye width in dorsal view about as wide as synthlipsis; ventral eye margin surpassed by setigerous tubercles in lateral view; ventrolateral swelling of buccula with large lateral setigerous tubercles that surpass buccular margin; labial segment I in lateral view straight, surpassing posteroventral eye margin; morphologically dorsal surface of labial segment II curves ventrally, about 0.4 times length of segment I. Thorax: pronotal collar in dorsal view narrow medially with anterolateral angles short, flared laterally; anterior pronotal lobe about 0.8 times length of posterior pronotal lobe, lateral margins rounded; glabrous markings on anterior pronotal lobe depressed near posterolateral margins of pronotum; median apodeme depression of pronotum deep, elongated transversely; scutellar spine long, subhorizontal; anteriad-directed process of prosternum with paramedial lobes; anterior margin of stridulitrum projected into small protuberance; proepimeron with smoothly rounded posteroventral margin. Legs: fossula spongiosa present on fore and mid legs. Abdomen: anterior margin of terga weakly carinulate, tergum II without carinules laterally; terga II and III with paired prominent longitudinal carinae, reaching about one-third of tergum III. Pygophore (Fig. 13): transverse bridge with rounded posterior margin; posterior region of ventral surface of pygophore in lateral view with large swelling; short median apical process, bent posteriorly in lateral view; lateral pygophore margin with protuberance; posterior pygophore margin with clustered macrosetae on decline; parameres sinusoidal, apex tapered sharply. Aedeagus (Fig. 14): endosoma almost entirely covered with spicules; apex of dorsal phallothecal sclerite bifid in dorsal view; basal plate extension about 5.8 times as long as wide.

Female
As male. Body length: about 10.7 mm. Head: elongate, about 1.3 times as long as wide; pedicel about 1.5 times length of head width. Legs: fossula spongiosa absent or vestigial on mid leg.

Distribution
Only known from Eastern Australia.

Remarks
We were unable to physically examine the type material of Voconia pallidipes (photographs available on the NHRS website) and we thus based our redescription and character coding on a specimen deposited at the AMNH that was identified as V. pallidipes (USI: AMNH_PBI 00170708) by P. Wygodzinsky and that matches the diagnostic features visible in the type photographs. The specimen in the NHRS photographs is the only specimen in the series and thus designated as the lectotype. Characters of the dorsal part of the abdomen (i.e., carinules and carinae), pygophore, and aedeagus are based on a single specimen (USI: AMNH_PBI 00127093), to limit damage to specimens. This species is nested within the well supported Australian clade (Figs 1-2). The record and illustration of V. pallidipes by Swanson (2015: 268-269, fig. 10) is based on misidentification and pertains to V. brachycephala sp. nov. (see under the latter species).

Diagnosis
Distinguished from other species of Voconia by the following combination of characters: coarsely granulose head and pronotum; maxillary plates not adjacent to and far surpassing clypeal apex; maxillary plates in lateral view wide, about twice the width of the scape and not toothed apically; antennifer with long lateral spine; eyes in lateral view do not reach ventral head margin; pedicel short, about 0.6 times length of head width. The macropterous morph is recognized from V. tridens sp. nov. by the almost entirely dark brown head and pronotum, the scutellar spine is slightly raised, and the hemelytra are blackish with pale coloration at the base and apex of the corium.
struCture. Head (Fig. 3D): elongate, about 1.4 times as long as wide, narrower in macropterous morph; anteocular region two-fifths of head length, longer than postocular region (measured to anterior margin of neck); postocular region in dorsal view longer than eye, lateral margins subrectangular; scape not reaching apex of head, about 0.4 times length of pedicel; pedicel about 0.6 times length of head width; maxillary plates conical, not adjacent to or longer than clypeus; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view narrower than maxillary plates; clypeal apex round, narrowed; interocular glabrous markings V-shaped, joined medially at interocular sulcus; interocular sulcus in Fig. 13. Pygophore of species of Voconia Stål, 1866. Scale bars = 0.5 mm. dorsal view nearly straight; eye width in dorsal view narrower than synthlipsis in micropterous morph, eyes considerably larger in macropterous morph; eye not reaching ventral head margin in lateral view; ocelli minute, occupying about 0.16 times length of postocular region in micropterous morph, ocelli considerably larger in macropterous morph; ventrolateral swelling of buccula with acute protrusion surpassing buccular margin, small setigerous tubercles along margin of protrusion; labial segment I in lateral view straight, not reaching posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.4 times length of segment I. Thorax: pronotal collar in dorsal view wide medially with anterolateral angles long, flared laterally; anterior pronotal lobe about 1.3 times length of posterior pronotal lobe, lateral margins rounded in micropterous morph; anterior pronotal lobe shorter than posterior pronotal lobe in macropterous morph; glabrous markings on pronotum deeply depressed; median apodeme depression of pronotum shallow, elongated longitudinally; scutellar spine reduced and subhorizontal in micropterous morph; scutellum about 2 times as long as wide at base, with long slightly raised scutellar spine in macropterous morph; anteriad-directed process of prosternum with paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with smoothly rounded posteroventral margin. Legs: fossula spongiosa present on fore and mid legs. Abdomen (Fig. 4D): anterior margin of terga prominently carinulate, terga II and III with deeper and longer carinules; terga II with paired prominent longitudinal carinae reaching posterior margin of segment.

Female
As micropterous male. Body length: about 7.0 mm.

Distribution
Distributed in Garamba National Park in the Democratic Republic of the Congo.

Remarks
This description of V. schoutedeni is based on the micropterous male holotype (UCR_ENT 00070524), which was the only type specimen available for examination. Any descriptive information of the micropterous female or macropterous form is taken directly from the original description by Villiers (1964). A lectotype was also designated for the macropterous morph, Pseudocethera schoutedeni forme alata Villiers, 1964. However, this infrasubspecific name is permanently unavailable under ICZN Article 45.6.3, therefore this specimen (referred to as No. 1281 by Villiers) has no type status. A drawer image of specimens at the BMNH in 2011 suggests that the macropterous morph may be distributed as far north as Sierra Leone, but this specimen and the macropterous paratype need to be properly examined.

Diagnosis
Distinguished from other species in Voconia by its Afrotropical distribution and stout body. This species most closely resembles V. motoensis and V. conradti. Voconia smithae sp. nov. differs from V. motoensis by its distinct coloration: yellow scutellar spine; abducted corium with a yellow spot at proximal and distal apex, and brown dorsal laterotergites with contrasting yellow anterior spots. Despite the strong superficial resemblance to V. conradti due its to coloration and stoutness, closer examination reveals that V. smithae sp. nov. is distinguishable by the following: shorter body (9.7-9.8 mm long); light brown dorsal laterotergites with weakly contrasting yellow anterior spots; antennifer with long, laterally projecting setigerous tubercle; paired interocular setigerous tubercles; maxillary plates globular, directed straight anteriorly; longer anteocular region than postocular region (measured to where the posterior margin of the granulations meet the anterior margin of the smooth neck); and eyes do not bulge strongly laterally in dorsal view and do not reach ventral head margin in lateral view.

Etymology
Named after one of the collectors of the holotype and colleague from the Weirauch lab, Samantha Smith.
Coloration. Head: dark brown; postocular region with dark patches adjacent to medial ocellar margin, pale patches adjacent to lateral ocellar margin; labium lighter than head. Thorax: coloration as head, posterior margin of posterior pronotal lobe lighter; scutellum dark with contrasting yellow apical spine. Hemelytron (abducted): clavus brown with distal yellow stripe; corium dark or light brown with proximal, anteromedial, posteromedial, and distal yellow spots; membrane dark with pale V-shaped marking along R and M veins; membranal veins R and M proximally pale. Legs: yellowish-brown. Abdomen: yellowish-brown; dorsal laterotergites light brown with yellow spots anteriorly. integument and vestiture. Head and pronotum: coarsely granulose with sparse, short macrosetae interspersed among sparse pubescence; interocular region with two pairs of stout setigerous tubercles paramedially; antennifer with long lateral setigerous tubercle; morphologically ventral surface of labium with sparse, short macrosetae. Thorax: anterolateral angles of pronotal collar without macrosetae or setigerous tubercles. Hemelytron: corium with dense long setation. Legs: posterior row of protuberances on mid and hind femora with three large spines on distal half and three or fewer small spines. Abdomen: ventral surface pubescent.
struCture. Head: elongate, 1.3-1.4 times as long as wide; anteocular region about one third of head length, longer than postocular region (measured to anterior margin of neck); postocular region in dorsal view about as long as eye, lateral margins gently rounded and subrectangular; pedicel 1.4-1.5 times length of head width; maxillary plates ellipsoidal, adjacent to and longer than clypeus; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view narrower than maxillary plates; clypeal apex round, narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view curved and smoothly rounded anteriorly; eye width in dorsal view narrower than synthlipsis; eye not reaching ventral head margin in lateral view; ventrolateral swelling of buccula with tuberculate protrusion that surpasses buccular margin; labial segment I in lateral view straight, reaching posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.6 times length of segment I. Thorax: pronotal collar narrow medially with anterolateral angles short and flared laterally; anterior pronotal lobe about 0.6 times length of posterior pronotal lobe, lateral margins almost straight; glabrous markings on pronotum slightly depressed; median apodeme depression of pronotum deep, circular; scutellar spine long, subhorizontal; anteriad-directed process of prosternum with paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with smoothly rounded posteroventral margin. Legs: fossula spongiosa present on fore and mid legs. Abdomen: anterior margin of terga weakly carinulate; terga II and III with paired prominent longitudinal carinae, reaching about one-third of tergum III.

Unknown.
Distribution (Fig. 17) Only known from Congo and Gabon in Africa.

Remarks
The male paratype varies from the holotype in the following: head and pronotum finely granulose, postocular region subrectangular, buccula without lateral tuberculate protrusion. The holotype was preserved in ethanol when the mid and hind legs were dissected for DNA extraction but was not Illumina sequenced.

Diagnosis
Distinguished from other species of Voconia by the following combination of characters: coarsely granulose head and pronotum; maxillary plates not adjacent to and far surpass clypeal apex; and antennifers with a long, lateral spine. This species is recognized from the macropterous morph of V. schoutedeni by the yellow head and pronotum; the maxillary plates in lateral view are not wide, about the width of the scape; scutellar spine is subhorizontal; and the hemelytra is not blackish, it is a light brown with pale coloration at the base and apex of the corium.

Etymology
The specific epithet is the Latin adjective 'tridens, -, -', meaning 'three-toothed, three-pronged', derived from the prefix 'tri-' combined with the Latin noun 'dens', referring to the elongated and separated maxillary plates and clypeus that form three prongs. Coloration. Head: yellow; postocular region with red patches adjacent to medial ocellar margin; labium as head. Thorax: uniformly yellow; scutellum yellow. Hemelytron (abducted): clavus uniformly yellow; corium proximal half mostly yellow, distal half brown with yellow spot at distal apex; membrane dark with pale V-shaped marking along R and M veins. Legs: yellow. Abdomen: yellow, dorsal laterotergites with a pale brown stripe on posterior margin. integument and vestiture. Head and pronotum: coarsely granulose with dense, short setigerous tubercles; interocular region with two pairs of stout setigerous tubercles paramedially; antennifer with long lateral spine; morphologically ventral surface of labium with sparse, short macrosetae. Thorax: anterolateral angles of pronotal collar without macrosetae or setigerous tubercles; scutellar lateral carinae with pubescent tubercles. Hemelytron: corium with short macrosetae. Legs: posterior row of protuberances on mid and hind femora with three large spines on distal half. Abdomen: ventral surface pubescent.

Holotype
struCture. Head: elongate, about 1.3 times as long as wide; anteocular region two-fifths of head length, longer than postocular region (measured to anterior margin of neck); postocular region in dorsal view longer than eye, lateral margins subrectangular; scape not reaching head apex; pedicel about 0.9 times length of head width; maxillary plates ellipsoidal, not adjacent to or longer than clypeus; apices of maxillary plates in dorsal view diverging; clypeus in dorsal view narrower than maxillary plates; clypeal apex round, narrowed; interocular glabrous markings V-shaped, joined medially at interocular sulcus; interocular sulcus in dorsal view curved and smoothly rounded anteriorly; eye width in dorsal view narrower than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula with acute protrusion surpassing buccular margin, small setigerous tubercles along margin of protrusion; labial segment I in lateral view straight, surpassing posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.4 times length of segment I. Thorax: pronotal collar in dorsal view wide medially with anterolateral angles long, flared laterally; anterior pronotal lobe about 0.7 times length of posterior pronotal lobe, lateral margins almost straight; glabrous markings on pronotum deeply depressed; median apodeme depression of pronotum shallow, elongated longitudinally; scutellar spine long, subhorizontal; anteriad-directed process of prosternum with paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with smoothly rounded posteroventral margin. Legs: fossula spongiosa present on fore and mid legs. Pygophore (Fig. 13): transverse bridge with triangular posterior margin; posterior region of ventral surface of pygophore in lateral view straight, not swollen; short median apical process sharply bent posteriorly in lateral view; lateral pygophore margin with protuberance; posterior pygophore margin with clustered macrosetae on protuberance; parameres round, apex tapered into squarely rounded tip. Aedeagus: endosoma almost entirely covered with spicules; apex of dorsal phallothecal sclerite tongueshaped in dorsal view; basal plate extension about 4.6 times as long as wide.

Unknown.
Distribution (Fig. 17) Only known from the type locality in Northeastern Africa, Sudan.

Remarks
The antennifer spine on the right side of the head of the holotype is absent. We assume it was damaged since three species, V. schoutedeni, V. monodi, and V. ifana, have similar long spines that are paired. The pygophore of specimen AMNH_PBI 00170712 was used for DNA extraction, but we did not get sufficient DNA in our NGS library for Illumina sequencing.

Diagnosis
One of two New World species of Voconia. Recognized by the finely granulose head and prothorax; proepimeron with protuberance on posteroventral margin; pronotum dark with contrasting yellow posterior margin; corium dark with anteroproximal yellow stripe, posteromedial and distal white spots; labial segment I in lateral view not surpassing posteroventral eye margin.

Etymology
Named after the collecting locality of the holotype in Trinidad. A proper noun with the Latin adjectival suffix '-ensis' meaning 'belonging to'. To be treated as an adjective.
Coloration. Head: dark brown; maxillary plates light brown; postocular region with pale patches adjacent to lateral ocellar margin; labium lighter than head, yellow. Thorax: as head; posterior pronotal lobe with posterior margin lighter; scutellum dark with contrasting yellow apical spine. Hemelytron (abducted): clavus reddish-brown with distal white stripe; corium reddish-brown with anteroproximal yellow stripe, posteromedial and distal white spots; membrane dark with white V-shaped marking along R and M veins; membranal veins R and M proximally white. Legs: yellow. Abdomen: dark brown, dorsal laterotergites brown medially, with black stripe adjacent to yellow lateral margin. integument and vestiture. Head and pronotum: finely granulose with sparse, long macrosetae interspersed among dense pubescence; interocular region with two pairs of macrosetae paramedially; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with dense, short macrosetae. Thorax: anterolateral angles of pronotal collar with macrosetae; scutellar lateral carinae with long setation. Hemelytron: corium with sparse, long setation. Legs: posterior row of protuberances on mid and hind femora with three large spines on distal half and many small spines. Abdomen: ventral surface pubescent.
struCture. Head: elongate, about 1.4 times as long as wide; anteocular region about one third of head length, about as long as postocular region (measured to anterior margin of neck); postocular region in dorsal view about as long as eye, lateral margins gently rounded; pedicel about 1.3 times length of head width; maxillary plates ellipsoidal, adjacent to and longer than clypeus; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view wider than maxillary plates; clypeal apex round, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view curved and smoothly rounded anteriorly; eye width in dorsal view wider than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, reaching posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.6 times length of segment I. Thorax: pronotal collar in dorsal view narrow medially with anterolateral angles short, projected forward; anterior pronotal lobe about 0.5 times length of posterior pronotal lobe, lateral margins almost straight; glabrous markings on pronotum thin, not deeply depressed; median apodeme depression of pronotum deep, elongated transversely; scutellar spine long, subhorizontal; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with acute protuberance on posteroventral margin. Legs: fossula spongiosa of fore leg present, absent or vestigial on mid leg. Abdomen: anterior margin of terga prominently carinulate, posterior margin of terga III-VI weakly carinulate; terga II and III with paired prominent longitudinal carinae, reaching about two-thirds of tergum III; medial region between carinae on tergum II coarsely granulose. Distribution (Fig. 19) This species is only known from the type locality, Trinidad.

Diagnosis
Distinguished from most species of Voconia by its tuberculate and coarsely granulose head and pronotum. This species most closely resembles another Australian species, V. dolichocephala sp. nov. These two species are recognized from other Australian species by the acute protuberance on the posteroventral margin of the proepimeron, the long macrosetae of the setigerous tubercles, the elongate head (1.4-1.6 times as long as wide), and the yellow spots antero-and posteromedially on the abducted corium. Voconia tuberculata sp. nov. differs from V. dolichocephala sp. nov. by its overall light brown coloration and males are as long as about 7.9 mm. Females are smaller than the examined V. dolichocephala sp. nov. females (about 8.8 and 9.0 mm).

Etymology
The specific epithet is the Latin adjective 'tuberculatus, -a, -um', meaning 'warty' or 'tuberculate'. Refers to the densely tuberculate surfaces of the body.
Coloration. Head: dark brown; postocular region with dark patches adjacent to medial ocellar margin and pale patches adjacent to lateral ocellar margin; labium lighter than head. Thorax: coloration as head; scutellum uniformly brown. Hemelytron (abducted): clavus reddish-brown with distal yellow stripe; corium reddish-brown with anteromedial, posteromedial, and distal yellow spots; membrane dark with pale V-shaped marking along R and M veins; membranal veins R and M proximally pale. Legs: yellowish-brown. Abdomen: light brown, dorsal laterotergites I and II paler than remainder. integument and vestiture. Head and pronotum: coarsely granulose with sparse, long setigerous tubercles interspersed among sparse pubescence; interocular region with two pairs of stout setigerous tubercles paramedially; antennifer with long lateral setigerous tubercle; morphologically ventral surface of labium with sparse, short macrosetae. Thorax (Fig. 3L): anterolateral angles of pronotal collar with setigerous tubercles; scutellar lateral carinae with setigerous tubercles with long macrosetae. Hemelytron: corium with dense long setation. Legs: posterior row of protuberances on mid and hind femora with four or more large spines on distal half. Abdomen: ventral surface pubescent, long macrosetae interspersed.
struCture. Head: elongate, 1.4-1.6 times as long as wide; anteocular region about one third of head length, shorter than postocular region (measured to anterior margin of neck); postocular region in dorsal view longer than eye, lateral margins gently rounded; pedicel about 1.5-2.2 times length of head width; maxillary plates ellipsoidal, adjacent to and longer than clypeus; apices of maxillary plates in dorsal view diverging; clypeus in dorsal view about as wide as maxillary plates; clypeal apex bifid, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially or medially forming a V-shaped marking; interocular sulcus in dorsal view nearly straight; eye width in dorsal view narrower than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, surpassing posteroventral eye margin; morphologically dorsal surface of labial segment II curves ventrally, about 0.3 times length of segment I. Thorax (Fig. 3L): pronotal collar in dorsal view wide medially with anterolateral angles long, flared laterally; anterior pronotal lobe 0.6-0.8 times length of posterior pronotal lobe, lateral margins almost straight; glabrous markings on pronotum conspicuous, not depressed; median apodeme depression of pronotum shallow, elongated transversely; scutellar spine long, raised; anteriad-directed process of prosternum with paramedial lobes; anterior margin of stridulitrum projected into small protuberance; proepimeron with acute protuberance on posteroventral margin. Legs: fossula spongiosa present on fore and mid legs. Abdomen: anterior margin of terga weakly carinulate, terga II and III with deeper and longer carinules; terga II and III with paired prominent longitudinal carinae, reaching about two-thirds of tergum III. Pygophore (Fig. 13): transverse bridge with rounded posterior margin; posterior region of ventral surface of pygophore in lateral view with large swelling; short median apical process, sharply bent posteriorly in lateral view; lateral pygophore margin with protuberance; posterior pygophore margin with clustered macrosetae on protuberance; parameres sinusoidal, apex tapered sharply. Aedeagus (Fig. 14): endosoma almost entirely covered with spicules; apex of dorsal phallothecal sclerite serrated; basal plate extension about 5.7 times as long as wide.

Female
Mostly as male, distinguished from males by the following. Body length: 8.4-8.8 mm. Head: elongate, 1.5 times as long as wide; pedicel about 1.5-1.7 times length of head width. Legs: fossula spongiosa present on fore leg, absent or vestigial on mid leg. Distribution (Fig. 18) This species is distributed across Northern Territory and Queensland in Australia. The nearest Voconia species is V. dolichocephala sp. nov., found about 100 km northwest of specimen AMNH_PBI 00168742.

Remarks
The direction of the maxillary plates in dorsal view varies from nearly straight or parallel (e.g., AMNH_ PBI 00168741) to strongly diverging (e.g., AMNH_PBI 00168746). Voconia tuberculata sp. nov. is nested within the Australian clade. According to our phylogeny (Figs 1-2), V. tuberculata sp. nov. was recovered as the sister taxon to V. dolichocephala sp. nov. The abdomen of specimen AMNH_PBI 00127096 was used for DNA extraction, but we did not get sufficient DNA in our NGS library for Illumina sequencing.

Revised diagnosis
This species is most similar to V. bracata sp. nov. They are distinguished from other species of the genus Voconia by the slender body, long head (1.7 times as long as wide), the ventral surface of the hind leg with four large spines in the posterior row, the large pale spot between the R and M veins, and the dorsal laterotergites I and II being yellow, the remainder dark brown. The following differentiate V. typica: largest species of Voconia (12.4-12.7 mm long); uniformly brown legs; anterior pronotal lobe is gently rounded, posterolateral margins nearly straight.

Type material
membranal veins R and M proximally pale, remainder as membrane. Legs: dark brown. Abdomen: dark brown, dorsal laterotergites I and II yellow.
integument and vestiture. Head and pronotum: finely granulose with sparse, short macrosetae interspersed among dense pubescence; interocular region with two paired macrosetae; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with dense, short macrosetae. Thorax: anterolateral angles of pronotal collar without macrosetae or setigerous tubercles. Hemelytron: corium with sparse, long setation. Legs: posterior row of protuberances on mid and hind femora with four large spines on distal half. Abdomen: ventral surface pubescent.
struCture. Head: elongate, about 1.7 times as long as wide; anteocular region about one third of head length, about as long as postocular region (measured to anterior margin of neck); postocular region in dorsal view longer than eye, lateral margins gently rounded; pedicel about 1.8 times length of head width; maxillary plates ellipsoidal, adjacent to and reaching apex of clypeus; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view narrower than maxillary plates; clypeal apex round, not narrowed; interocular glabrous markings V-shaped, joined medially at interocular sulcus; interocular sulcus in dorsal view bent anteromedially, subtriangular; eye width in dorsal view wider than synthlipsis; eye reaching ventral head margin in lateral view; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I in lateral view straight, surpassing posteroventral eye margin; morphologically dorsal surface of labial segment II nearly straight, about 0.4 times length of segment I. Thorax: pronotal collar in dorsal view narrow medially with anterolateral angles short, projected forward; anterior pronotal lobe subequal in length to posterior pronotal lobe, lateral margins almost straight; glabrous markings on pronotum thin and deeply depressed; median apodeme depression of pronotum deep and elongated transversely; scutellar spine long, subhorizontal; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with smoothly rounded posteroventral margin. Legs: fossula spongiosa present on fore and mid legs. Abdomen: anterior margin of terga weakly carinulate, tergum II without carinules laterally; terga II and III with paired prominent longitudinal carinae, almost reaching posterior margin of tergum III. Pygophore (Fig. 13): transverse bridge with rounded posterior margin; posterior region of ventral surface of pygophore in lateral view with large swelling; short median apical process sharply bent posteriorly in lateral view; lateral pygophore margin with protuberance; posterior pygophore margin with clustered macrosetae on decline; parameres sinusoidal, apex tapered into squarely rounded tip. Aedeagus (Fig. 14): endosoma almost entirely covered with spicules; apex of dorsal phallothecal sclerite bifid in dorsal view; basal plate extension about 7.9 times as long as wide. Distribution (Fig. 18) This species is distributed on the island of New Guinea, in Papua.

Remarks
This description is based largely on UCR_ENT 00073815, since the holotype (Fig. 8) and paratype remain at the BMNH. Images of the types in their unit trays at the BMNH were used to evaluate this species concept. The abdomen of specimen UCR_ENT 00073815 was used for DNA extraction, but we did not get sufficient DNA in our NGS library for Illumina sequencing.

Diagnosis
Distinguished from other species of Voconia by the following combination of characters: short body length (about 7.1 mm long); globose head; tuberculate and coarsely granulose head and pronotum; interocular region with two setigerous tubercles medially; apex of clypeus bifid; corium dark with thick medial yellow stripe; and dark brown femora with contrasting yellowish-brown tibiae and tarsi. This species most closely resembles another Australian species, V. fasciata sp. nov., but V. vittata sp. nov. is about 1.6 mm shorter and the yellow medial stripe on the corium is opaque. Coloration. Head: dark brown with a semicircular pale stripe encircling both ocelli; labium lighter than head, brown. Thorax: as head; scutellum dark with brown apical spine. Hemelytron (abducted): clavus dark with distal yellow stripe; corium dark with anteroproximal yellow stripe, posteromedial and distal yellow spots; membrane dark with pale V-shaped marking along R and M veins; membranal veins R and M proximally pale. Legs: dark brown. Abdomen: dark brown; dorsal laterotergites dark brown.
integument and vestiture. Head and pronotum: finely granulose with dense, long macrosetae interspersed among dense pubescence; antennifer with short lateral setigerous tubercle; morphologically ventral surface of labium with dense, short macrosetae. Thorax: anterolateral angles of pronotal collar without macrosetae or setigerous tubercles. Legs: posterior row of protuberances on mid and hind femora with four spines on distal half. Abdomen: ventral surface pubescent.
struCture. Head (Fig. 3E): elongate, about 1.5 times as long as wide; anteocular region about half of head length, longer than postocular region (measured to anterior margin of neck); postocular region in dorsal view longer than eye, lateral margins gently rounded; pedicel about 1.1 times length of head width; maxillary plates ellipsoidal, adjacent to and shorter than clypeus; apices of maxillary plates in dorsal view directed straight; clypeus in dorsal view about as wide as maxillary plates; clypeal apex round, not narrowed; interocular glabrous markings anterolaterally curved, joined at interocular sulcus paramedially; interocular sulcus in dorsal view curved and smoothly rounded anteriorly; eye width in dorsal view narrower than synthlipsis; eye reaching ventral head margin in lateral view; ocelli minute, occupying 0.12 times length of postocular region; ventrolateral swelling of buccula without lateral protrusion that surpasses buccular margin, flat margin; labial segment I swollen subapically on ventral surface, surpassing posteroventral eye margin; labial segment II swollen ventrobasally, narrowed apically; morphologically dorsal surface of labial segment II curves ventrally, ~0.2 times length of segment I; labial segment III short, ~0.2 times length of segment II. Thorax: pronotal collar in dorsal view narrow medially with anterolateral angles short, projected forward; anterior pronotal lobe ~0.8 times length of posterior pronotal lobe; glabrous markings on pronotum thin and not deeply depressed; median apodeme depression of pronotum deep, elongated transversely; scutellar spine long, subhorizontal; anteriad-directed process of prosternum smoothly rounded, without paramedial lobes; anterior margin of stridulitrum not elongated into protuberance; proepimeron with smoothly rounded posteroventral margin. Legs: fossula spongiosa present on fore legs, unknown on mid legs.

Unknown.
Distribution (Fig. 20) This species is only known from the type locality in Eastern Borneo.

Remarks
The holotype remains at the NBCN under loan restrictions; habitus photographs were used to evaluate this species concept. Most closely resembles the two species from New Guinea, V. bracata sp. nov. and V. typica. These three species form a poorly supported clade in our analyses (Figs 1-2). This hypothesis should be further tested with additional material and molecular data.

Discussion
Pseudocetherinae was originally described based on Pseudocethera (3 spp. from Africa) as having a short, tri-lobed head with the antennal pedicel longer than the scape, small or vestigial ocelli, slender labium with labial segment I long and adpressed against the ventral head surface, the close proximity of the coxal cavities of fore and mid legs, and the three-segmented tarsi. We expand this concept of Pseudocetherinae by transferring in five genera from Reduviinae: Gerbelius (three spp. from the Oriental region), Kayanocoris (one sp. from Borneo), Microvarus (two spp. from Africa), Paragerbelius (one sp. from Papua), and Voconia (one spp. from Australia). As part of this revision, Kayanocoris, Microvarus, Paragerbelius, and Pseudocethera are synonymized with Voconia. As pointed out in the results section of the cladistic analysis, Gerbelius in the traditional sense is polyphyletic with respect to G. ornatus. Members of Gerbelius share laterally flattened maxillary plates; long prominent mandibular plates; a stout labium; and the Cu-An 1 cell is slender, tapers apically, and is similar in length to the M-Cu cell. Together, G. confluens and G. typicus, with two undescribed species, are part of the sister group of all remaining Pseudocetherinae (Figs 1-2). In contrast, G. ornatus not only closely resembles species of Voconia (anteriorly elongate maxillary plates, gracile labium, and short, triangular Cu-An 1 cell, among other characters) but based on our analyses is nested among them. Therefore, we transfer G. ornatus to Voconia and maintain G. confluens and G. typicus as the only two valid species of Gerbelius, the second genus of Pseudocetherinae.
Though the transfer of Gerbelius into Pseudocetherinae requires significant expansion of the original description, morphological (present study, Figs 1-2; Weirauch 2008), ribosomal (Weirauch 2008;Hwang & Weirauch 2012), and genomic analyses (Knyshov et al. unpubl. data) strongly support Gerbelius and Voconia as sister taxa. The bent apex of the third visible labial segment, anteriad-directed process of the prosternum, absence of dorsal abdominal glands on terga IV-VI, femora with ventral anterior and posterior rows of spine-like protuberances with macrosetae at the base of the protuberance, and laterally compressed tibiae unite Pseudocetherinae and form part of the new description.