Brontostoma basalis
Creators: name: Gil-Santana, Hélcio R.
Publisher: name: Zenodo
Year: 2020
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Hemiptera
Reduviidae
Brontostoma
Brontostoma basalis

description: Brontostoma basalis (Stål, 1859)
(Figs. 1–70)
Daraxa basalis Stål, 1859: 181 (original description).
Mindarus sanguinosus Stål, 1872: 102 (original description). New subjective synonym.
Ectrichodia pallitarsis Walker, 1873: 59–60 (original description). New subjective synonym.
Mindarus basalis: Stål (1872: 102) (checklist), Lethierry & Severin (1896: 134) (catalog), Haviland (1931: 133, 145) (citation, comments about general coloration).
Ectrichodia basalis: Walker (1873: 61) (catalog).
Brontostoma basalis: Wygodzinsky (1949: 22) (catalog), Wygodzinsky (1951: 40, 51) (key, remarks about coloration of a male and a female), Maldonado (1990: 28) (catalog), Dougherty (1995: 203) (citation, geographical distribution), Gil-Santana et al. (2004: 127) (citation), Gil-Santana et al. (2005: 78) (citation), Gil-Santana (2008: 44, 45) (new record, Figs. 1–2), Gil-Santana & Baena (2009: 42–44) (historical remarks, short description of male genitalia, Figs. 1–12), Gil-Santana et al. (2013: 62) (citation).
Mindarus sanguinosus: Lethierry & Severin (1896: 134) (catalog).
Brontostoma sanguinosum: Wygodzinsky (1949: 22) (catalog), Wygodzinsky (1951: 39, 45–47) (key, redescription, Figs. 17–21), Maldonado (1990: 30) (catalog), Dougherty (1995: 204) (citation, geographical distribution), Carpintero & Maldonado (1996: 132) (citation, geographical distribution), Gil-Santana et al. (2004: 127,128) (citations), Gil-Santana et al. (2005: 78) (citation), Forero (2006: 5, 6–7) (Fig. 4, historical remarks, additional records from Colombia), Gil-Santana et al. (2013: 62) (citation).
Ectrichodia pallitarsis: Lethierry & Severin (1896: 142) (catalog, as “Ectrichodidarum subfam. species incerti generis”).
Brontostoma pallitarsis: Wygodzinsky (1949: 22) (catalog), Wygodzinsky (1951: 40, 51) (key, citation of specimens examined), Maldonado (1990: 30) (catalog), Gil-Santana et al. (2004: 127) (citations), Gil-Santana et al. (2005: 78) (citations).
Distribution. Panama, Colombia, Ecuador, Peru, Guyana, Suriname, French Guiana (new record), and Brazil.
Diagnostic characters and their variability. Male. Body length 21–28 mm.
VESTITURE. Antennae covered by long setae (except on dorsal portion of the scape, which is glabrous), particularly numerous on the first four segments and even longer on the lateral and ventral portions of the scape and pedicel.
COLORATION (Figs. 1–2, 4–5, 11, 13, 15–16, 18–34, 61): head; thorax, including scutellum and transverse furrow of pronotum, but excepting hind lobe of pronotum; legs, except tarsi, generally blackish to dark brown. Abdomen mostly darkened.
Intraspecific variability in color. Antennae mostly dark, distiflagellomeres variable in color, from grayish, paler to whitish; in latter case with last segment commonly somewhat pale grayish, sometimes only at apex; some segments of labium sometimes paler, more commonly the last one. Hind lobe of pronotum highly variable, ranging from completely reddish (Fig. 27–28, 30, 32) to completely darkened (Figs. 11, 13), with intermediates decorated with various dark markings of brownish to blackish coloration, ranging from small dark areas restricted to basal median portion, progressively occupying approximately its basal third, basal half, or basal two-thirds (Figs. 1, 4, 15, 18–19, 22) of posterior lobe of pronotum, with only humeral angles reddish (Figs. 20, 24, 61). Hemelytra: approximately distal half to distal two thirds of clavus and adjacent area on corium pale yellowish or yellowish, veins between pale areas more commonly concolorous with ground color of corium or sometimes with the adjacent pale portions; corium completely reddish (Figs. 27, 30, 32) or with different shades of red, with a central dark spot of variable size (Figs. 1, 4, 15, 18, 28, 61), or with a dark longitudinal stripe of variable width (Fig. 19, 22, 26), or corium almost entirely darkened, with only base and lateral margin variably paler (Fig. 11, 13, 20, 24); membrane translucent, frequently with yellowish or golden tones; portions between veins, including cells may be slightly or more intensely darkened, in latter case only on basal portion of membrane or variably more extensively to its distal portion; veins either concolorous with adjacent surface of membrane or of different color (paler or darker, yellowish or whitish). A few specimens with paler portions or markings on approximately midportion of tibiae, small (Fig. 18–19) or larger (Fig. 24–25). Tarsi darkened or with some to all segments completely or partially pale. Connexivum with variable tones (pale to dark) of red, orange, yellowish, gray, or even light brownish (Figs. 1, 4, 11, 13, 15–16, 18–33, 61); connexival dark markings on segments II and/or VII present or absent, dorsally either restricted to inner portion of segment (e.g. Figs. 18–19) or extended to their external margin, in extreme cases forming complete dark markings resulting in an alternating pale and dark colored pattern on connexival segments (e.g. Figs. 4, 11, 13, 15, 61); these complete dark markings variably narrower towards external margin; laterally they occupy approximately either one third (e.g. Fig. 15), or somewhat less than a half (e.g. Fig. 4) or a half of the lengths of the respective segments (e.g. Figs. 11, 13). Sternites with a median pair of transverse, regularly bordered, pale markings, approximately located at basal portion of sternites IV–VI (e.g. Figs. 21, 23) or at almost all sternites (IV–VII) (e.g. Figs. 2, 31), (III– VI) (e.g. Fig. 29), (III–VII) (e.g. Fig. 25), except the first visible; these markings might occasionally be united at midportion in some or all segments in which they are present (e.g. Fig. 29), forming a band in each of them; varying in size among specimens and among segments, but not reaching posterior margin of respective segments; when present on sternites III and VII, marking(s) may be smaller than those on others segments (e.g. Figs. 25, 29 and 2, 31, respectively); their color vary from pale yellowish, yellow-reddish, reddish, orange or a darkened ill-defined tone. Some median markings and pale lateral margin or lateral markings may be faint in some segments therefore appear partially or completely absent, possibly because due to chemical processes inside the abdomen (e.g. Figs. 16, 33), inferred by the asymmetry of this feature on a given segment; the aforementioned ill-defined darkened tones observed or reported in some specimens (e.g. Fig. 2) might be attributed to similar processes. Lateral borders of sternites II–VII generally paler, almost always concolorous with median markings or ventral connexivum. A lateral pale narrow band of variable width, formed by pale areas on sternites II–VI or III–VI, frequently variably narrower approximately on the subdistal portion or distal half of each segment, resulting in a subtly undulate medial outline; in other specimens dark coloration at subdistal portion or distal halves of sternites II–VI or III–VI reaches lateral margin of each segment, resulting in separate pale lateral spots of variable size; sternite VII more extensively and progressively paler laterally and towards apex. Visible distal borders of segment VIII and genital capsule partially or variably pale to mostly darkened.
STRUCTURE. Wing polymorphism. Hemelytra not reaching, reaching or slightly surpassing apex of the abdomen.
Terminalia. Abdominal segment VIII (Figs. 34–37): almost completely concealed, except its postero-ventral margin, which is somewhat elevated and larger at lateral portions; sclerotized only on ventral face, becomes wider towards posterior margin; posterior and anterior margins curved; dorsal portion entirely membranous and narrower; spiracles on dorsal margin of ventral portion. Male genitalia. Genital capsule: medial process of pygophore (mpp) not visible in ventral view (Figs. 34, 38); exposed portion of pygophore subhexagonal, anterior margin rounded, integument glabrous, mostly smooth and shiny, with some thin transverse incomplete and irregular shallow linear impressions; less pigmented in the non-exposed portion; parameres (pa) mildly exposed in situ, their apices very close or in contact in resting position (Figs. 34, 38); a narrow dorsal (transverse) somewhat curved bridge (br) between anterior and posterior genital openings in dorsal view (Figs. 39). Medial process of pygophore (mpp) moderately sclerotized, apical margin rounded (Figs. 39, 41–42). Parameres symmetrical, elongated, curved just above middle third; apex rounded, with a very sclerotized subapical tooth; mostly glabrous, with a somewhat dense group of setae on inner surface, below tooth (Figs. 39–40). Subapical tooth somewhat curved, with an acute apex (Figs. 39–40). Phallus: articulatory apparatus with basal plate extension (bpe) much shorter than basal plate, latter with curved and thick basal plate arms (bpa), connected by a relatively narrow basal plate bridge (bpb) (Figs. 43–44, 46–55). Dorsal phallothecal sclerite (dps) symmetrical, sclerotized only in its apical half, middle of anterior margin deeply sinuate, enlarged apically, more pronouncedly sinuate laterally to anterior margin, with lateral margins forming a ventral curved fold (vf), with an acute apex (vfa), projecting laterally (Figs. 46, 49, 54, 56). Endosomal struts (es) larger at their basal portion, largely united at base and even more extensively at apex (Figs. 46, 48, 56). Phallus with its membranous wall smooth and longitudinally striated on basal portion ventrally (Fig. 45). Basoventral portion of endosoma wall formed by a thin membrane, smooth and longitudinally striated, very frequently partially or sometimes almost completely detached from the underlying endosoma wall as an external layer (el) (Figs. 51–53, 57) (observable when the endosoma is expanded): inflated endosoma more or less tubular; in some specimens it forms a large ventral fold (evf) attached to apex of external ventral layer (el) (Figs. 51–53, 57); in one specimen, this fold was directed dorsally, its apex attached to the basal portion of the wall, therefore forming a dorsal fold (edf) (Figs. 54–55). Upper half of median portion of endosomal struts (es) with conspicuous medial sinuous longitudinal borders (slb) (Fig. 56A, B). Two endosoma processes: a basal large dense subtriangular process (stp) and a median process (mp) that is attached by its midportion to the apex of the former (astp) (Figs. 48–50, 52–55, 57–59). Median process (mp) flat, finely striated, faintly sclerotized, somewhat more at mid portion; shaped as a pair of arched portions (Figs. 46, 48, 57–60).
Female: Body length 21–30 mm. With the exception of the submacropterous syntype of Mindarus sanguinosus (Fig. 7), all other examined females were extreme brachypterous, with a distinctly wider fore lobe and a shorter hind lobe of pronotum (Figs. 62, 64, 66, 68, 70). First antennal segment glabrous, base of second antennal segment almost glabrous, with a few sparse, short, adpressed setae; remaining portion of second segment and other antennal segments covered by numerous short setae and scattered longer straight setae. General coloration similar to that of males, apparently less variable, although a smaller number of specimens were examined. In extreme brachypterous individuals completely exposed tergites entirely dark to blackish (Figs. 62, 70), one specimen had pairs of sublateral pale reddish spots on anterior halves of segments III–VI (Fig. 66), other two females had similar markings but smaller on segment III, becoming progressively slightly larger until those on segment VI, while on segment VII, a transverse median pale reddish marking on the anterior half of segment VII (Fig. 64); another female with pairs of similar faintly, almost indistinguishable, paler areas on tergites IV–V, (Fig. 68). In this latter specimen the markings of the tergites as well as those on sternites, which also seem darkened (Fig. 69) when compared with those similar of the other female (Fig. 67), may possibly have become fainted by some chemical reaction as commented above. In females in which median transverse pale markings of sternites were evident, they were all united at midportion forming bands variably narrower at midportion (Figs. 8, 67, 69); lateral border of sternite VII similarly narrowly colored as preceding segments (Figs. 8, 67, 69).
Type material examined. Daraxa basalis Stål, 1859: [SURINAME]: male syntype: [handwritten labels:] Daraxa / Stål // basalis / Stål // [green label:] Surin. Cordua // [printed labels:] 3041 // Zool. Mus. / Berlin // [red label:] Typus; male syntype (?): [handwritten labels:] x Daraxa ♂ / basalis / Stål / Paratypus // [blue label:] Surinam / Cordŭa // 3041 // [printed labels:] Zool. Mus. / Berlin // [red label]: Paratypus (ZMHB).
Ectrichodia pallitarsis Walker, 1873: male type: [printed label:] 79. ECTRICHODIA PALLITARSIS. // [printed rounded label with light green margin:] Type // [handwritten small label:] 535 / a (BMNH).
Additional specimens examined. ECUADOR: Napo, Coca, 250 m, Amazonian rain forest, iii-iv.1982, leg. G. Onore, Brit. Mus. 1982-246, “ Brontosoma sanguinosum Stal ” (1 male, BMNH); same but xi-xii.1982, B.M. 1983-255 (1 female, BMNH). FRENCH GUIANA: Bélizon, iii–iv, ix–xii, 1997, leg. H. Gaspard (7 males, MNRJ); Piste de Coralie, 19.iii.1997, leg. H. Gaspard (1 male, MNRJ). PERU: Iquitos, x.1988, xi.1989 (2 females, MNRJ); same but xi–xii.1989 (3 males, MNRJ). SURINAME: Paramaribo, 28.xii.1957, leg. P. H. v. Doesburg Jr., “ Brontostoma basalis (Stal) ”, Det. V. Dougherty, RMNH.INS.722820 [QR code] (1 male + 1 female “in copula”, NBC); Paramaribo, 10.xii.1957, leg. P. H. v. Doesburg Jr. (1 male, NBC); Surin Ben, Dicrene, 8.xi.1963, leg. P. H. v. Doesburg Jr., “ Brontostoma basalis (Stal) ”, Det. V. Dougherty (1 female, NBC); Paramaribo, Geijersvlijt, 21.iv.1940, leg. Geijskes, “ Brontostoma basalis (Stål) ”, Wygodzinsky det., [ex] Coll. Wygodzinsky (1 male, MNRJ); Lelydorp, 9.xii.1938, km 20, leg. Geijskes,“ Brontostoma basalis (Stål) ”, Wygodzinsky det., [ex] Coll. Wygodzinsky (1 female, MNRJ); Langaman Kondre, Marowijne Dist., viii.1965, leg. B. Malkin (2 males, MNRJ); Rust en Werk, 1937, leg. de Kraker, “ Brontostoma pallitarsis (Walker) ”, Wygodzinsky det., [ex] Coll. Inst. Med. Reg. (1 male, MNRJ).
Discussion. Brontostoma basalis was described (as Daraxa basalis) based on an unspecified number of male(s) (syntypes) from Suriname, without mentioning a precise locality (Stål 1859, 1872). A male (Figs. 1–2) bearing Stål’s handwritten identification label (Fig. 3) and another male individual (Figs. 4–6) are deposited in ZMHB. Both specimens bear red labels attached subsequently as a matter of curatorial routine, the first with the text “Typus” (Fig. 3), the second “ Paratypus ” (Fig. 6). As no holotype was designated in the original description, the latter specimen cannot be considered as paratype. Both specimens match the original description, and both bear a locality label “ Surinam ” (or “Surin.”), “Cordua”. According with Papavero (1971), the German dipterologist Christian Rudolph Wilhelm Wiedemann (1770–1840) had many specimens of flies from Suriname, part of them obtained “from a collector named Cordua (?), about whom he was not able to obtain any detail” (Papavero 1971). This historical record makes probable that both male specimens were collected by the same collector and they were examined by Stål. Therefore, in the contrary of the opinion expressed by Gil-Santana & Baena (2009), both specimens are regarded as syntypes here (ICZN 1999, Art. 73.2).
In both specimens the hind lobe of pronotum is as dark as the fore lobe on approximately its basal third (Fig. 4) to its basal two thirds (Fig. 1), and the corium of hemelytra is pale reddish with a medium sized central dark spot, while the approximately distal half of clavus and adjacent area on corium are pale yellowish (Figs. 1, 4). Haviland (1931) commented that the specimens of Kartabo (Guyana) agreed fairly well with the original description of this species, though the latter had described the color as testaceous, while the specimens observed by Haviland were “of a fine coral pink”. It is noteworthy, however, that Stål (1859) described the general color of D. basalis as “nigricans” (=blackish) with some portions such as the posterior margin of the pronotum, clavus, the corium of hemelytra (with exception of the black spot on disc), pale markings of the connexivum and a paired series of abdominal ventral markings as testaceous. Therefore, Haviland (1931) probably wanted to highlight the difference in color of these latter parts, possibly because they became remarkable when colored as “fine coral pink”. Subsequently, Wygodzinsky (1951) diagnosed B. basalis as a species bearing a yellowish corium of hemelytra with a small dark spot in it. He based his statements on a male from Suriname reexamined here (Figs. 15–17), which he argued to have matched very well with the original description of the species, except for its smaller length (19 mm). He also furnished a short diagnosis of the micropterous female of this species, also based on a single specimen from Suriname (Figs. 64–65), highlighting that its pale parts of the body were red instead of yellowish. He stated that the general coloration of this female and the reddish rudimentary hemelytra with a central dark small spot had made him “put the specimen provisionally in this species” (B. basalis) [translated from Spanish, not literally]. A short description and some figures of the male genitalia of B. basalis were furnished by Gil-Santana & Baena (2009). Brontostoma basalis has already been recorded from Suriname (Stål 1859, 1872; Walker 1873; Wygodzinsky 1949, 1951; Maldonado 1990; Dougherty 1995), Guyana (Lethierry & Severin 1896; Haviland 1931; Dougherty 1995) and Brazil (Gil-Santana 2008).
Brontostoma sanguinosum was originally described as Mindarus sanguinosus Stål 1872, based on females from Brasilia borealis (northern Brazil) and Bogotá in Nova Granada (now Colombia) (Stål 1872). A female syntype from Bogotá (Figs. 7–10) is deposited at NHRS. Forero (2006) argued that this specimen was probably collected in a lowland area of Colombia rather than in Bogotá, because this species is not found above 1500 meters of altitude, while Bogotá is at 2600 meters; the specimen may have been labeled as “ Bogota ” just because it was the shipping denomination for the commercial trade (Forero 2006). The above specimen was redescribed by Wygodzinsky (1951); some of its characteristics are noteworthy, such as the hind lobe of pronotum and the corium of hemelytra being completely reddish, and its submacroptery, with the hemelytra reaching the distal portion of the penultimate tergite (VI) (Fig. 7). Wygodzinsky (1951) recorded the total length of the type specimen as being 27 mm; the same measurement given by Stål (1872) as “ 7 mm ” was evidently a typographic error. A short-winged (apparently [extreme] brachypterous) female syntype from northern Brazil, mentioned in the original description of M. sanguinosus (Stål 1872), is currently of unknown depository. Forero (2006) presented a photograph in dorsal view of a male from Colombia; the coloration of this individual was very similar to that of the female type, while the hemelytra were longer, approaching the tip of the abdomen. Brontostoma sanguinosum has so far been recorded from northern Brazil (“Brasilia borealis”) and Colombia (Bogotá in Nova Granada, but see remarks above) (Stål 1872), other localities in the latter country (Forero 2006), Peru (Maldonado 1990), Ecuador (Dougherty 1995) and Panama (Carpintero & Maldonado 1996).
Brontostoma pallitarsis was described (as Ectrichodia pallitarsis) based on a single male (the holotype) without locality (Walker 1873), the specimen is deposited now in BMNH (Figs. 11–12). This specimen is generally darkened, including the entire pronotum and corium of hemelytra, while the approximately distal two thirds of the clavus and the adjacent area on the corium are paler (Walker 1873; Fig. 11). The total length of the holotype is approximately 23 mm. Wygodzinsky (1951) recorded three males from Suriname; one of them, very similar to the holotype, was studied in course of the present study (Figs. 13–14). Maldonado (1990) also recorded B. pallitarsis from Suriname, while Dougherty (1995) did not include this species in her list of species of Brontostoma.
The general structure and vestiture of all taxa studied here are the same as described for Brontostoma by Dougherty (1995) and Carpintero & Maldonado (1996). Among the males examined, a study of a broad range of color varieties, including all the patterns previously considered as belonging to separated species (Figs. 1, 4, 11, 13, 18, 27, 30), in addition to several intermediate forms (Figs. 19–20, 22, 24, 26, 28), all of them presenting the same characteristics in the male genitalia (Figs. 34–60), supports the synonymies proposed here (Brontostoma basalis = B. sanguinosum, syn. nov. = B. pallitarsis, syn. nov.).
It is noteworthy that, because the male examined by Wygodzinsky (1951) had the pale portions yellowish (Fig. 15), he stated that the corium of hemelytra of B. basalis would be of this color in his key. However, as described above, none of the other specimens examined, including the type specimens, possessed a yellowish corium. It might be one of the numerous color variations recorded to this species, but taking into account that no other specimen with such a coloration has been recorded, the possibility that the specimen examined by him was faded is considered here as the most probable. Therefore, the eventual yellowish color of the corium was not included in the range of variation described above.
Wygodzinsky (1951) considered the identification of the extreme brachypterous female of B. basalis examined by him (Figs. 64–65) as “provisional”. A male and a female conserved “in copula” (therefore almost certainly conspecific), deposited in NBC (Figs. 61–63), were examined in the course of the present study; the male (Fig. 61) was generally very similar to the syntypes of B. basalis (Figs. 1, 4), whilst the female (Fig. 62) to the above mentioned individual described by Wygodzinsky (1951) (Fig. 64). These specimens help to corroborate the identification proposed by the present author. It is interesting to note that in extreme brachypterous females, the coloration of the small coria of the reduced hemelytra resembles the alleged corresponding pattern of a male of the same population or color form, as suggested by Wygodzinsky (1951). Therefore, the central dark small spot in the coria of the female examined by him (Fig. 64) and of the female conserved “in copula”, deposited in NBC (Fig. 62) would correspond to the males with a median dark marking on their coria (e.g. Figs. 1, 4, 15, 18, 28, 61). Similarly, in BMNH, there is a male and a female from the same locality (Napo, Coca), in Ecuador (Figs. 27, 70). Both specimens have the coria of hemelytra completely reddish. On the other hand, while a syntype of Mindarus sanguinosus (Fig. 7) is submacropterous, the other female syntype of this taxon described by Stål (1872) was apparently [extreme] brachypterous, as the other females of B. basalis examined in this study were (Figs. 62, 64, 66, 68, 70), agreeing with the statement of Forthman & Weirauch (2017) that females of some species of Brontostoma exhibit wing polymorphism. In regard to sexual dimorphism, despite the small number of females examined, it was possible to record their reduced antennal setation, with a glabrous first segment, and a basal portion of the second segment almost glabrous, with only some sparse adpressed short setae. The extreme brachypterous females clearly had a wider anterior lobe and a shorter posterior lobe of pronotum (Figs. 62, 64, 66, 68, 70). In relation to the coloration, in almost all males the pale median transverse markings of sternites were recorded as a pair of separated markings (e.g. Figs. 2, 5, 16, 21, 23, 25, 31), while in the females in which these markings were evident, they were all united at midportion forming bands (Figs. 8, 67, 69). The lateral border of sternite VII in females was recorded similarly narrowly colored as the preceding segments (Figs. 8, 67, 69) and not as more extensively and progressively paler colored laterally and towards apex as in males (e.g. Figs. 21, 23, 25, 29, 34). In respect to the characteristics of continuous variability, among the specimens examined here it was not possible to confirm or disregard differences with certainty. The general body length was very similar between males and females examined (21–28 mm and 21–30 mm, respectively). A female clearly seemed to have a wider abdomen (Figs. 66–67). However, in order to clarify whether the total length and width of the abdomen, the median markings of the sternites, the coloration of the lateral border of sternite VII, and other potentially dimorphic characteristics, such as the thickness of the fore femora, the size of the eyes and ocelli, indeed vary significantly between males and females, it would be necessary to examine more specimens preferably of the same populations.
Regarding previous records of the length of males of B. basalis and the results obtained here, it is noteworthy to mention that the syntype specimen of B. basalis (Fig. 1) labeled as “Typus” and the other syntype (labeled as “ Paratypus ”) (Fig. 4) measured 21.5 mm and 21.0 mm, respectively, which are very close to the measurement recorded to this species by Stål (1859; 22 mm). However, the male specimen for which a length of 19 mm was recorded by Wygodzinsky (1951) actually measured 21.5 mm. The difference was possibly caused by the fact that the head of the latter individual was strongly tilted down (Fig. 15), which may have caused the underestimated measurement provided by Wygodzinsky (1951).
The presence and position of a large fold (ef) on the endosoma wall (e.g. Figs. 51–53; 54–55) was considered as subject of intraspecific variation, which apparently also occurs in B. infensum (see below), and therefore a character without diagnostic value at species level.
description: Published as part of Gil-Santana, Hélcio R., 2020, New synonymies among species of Brontostoma (Hemiptera: Heteroptera: Reduviidae), pp. 301-325 in Zootaxa 4869 (3) on pages 303-319, DOI: 10.11646/zootaxa.4869.3.1, http://zenodo.org/record/4418623