First description of the male of Volesusnigripennis Champion, 1899, with new records from Ecuador and Panama, taxonomical notes, and an updated key to the genera of Sphaeridopinae (Hemiptera, Reduviidae) Author R. Gil-Santana, Helcio Author Oliveira, Jader text ZooKeys 2019 841 97 123 http://dx.doi.org/10.3897/zookeys.841.31153 journal article http://dx.doi.org/10.3897/zookeys.841.31153 1313-2970-841-97 671DC5B5CB4E49BCB653EF1932392C9A Volesus nigripennis Champion, 1899 Figs 1-3, 4-8, 9-13, 14-19, 20-25, 26-28, 29-34, 35-39, 40-44, 45-48, 49-52, 53-57 Volesus nigripennis Champion, 1899: 296 [description], Tab. XVIII [Figure 14]; Costa Lima 1940 : 207 [citation], Wygodzinsky 1949 : 65 [catalog]; Maldonado 1990 : 490 [catalog]; Gil-Santana et al. 1999 : 2 [citation]; Forero 2004 : 164 [citation from Colombia], Figures 5.25, 5.103; Forero 2006 : 36 [new record from Colombia], Figures 56-57; Gil-Santana et al. 2015 : 336 [citation]. Notes. Volesus nigripennis was described based on a female from Costa Rica ( Champion 1899 ). The female holotype is deposited at the Swedish Museum of Natural History (NRM), Stockholm, Sweden, and its photos are available on their website (Figs 1-3). Figures 1-3. Volesus nigripennis , female, holotype deposited in NRM, catalog number NHRS-GULI000000089, photographed by Gunvi Lindberg, © 2018 Naturhistoriska riksmuseet. Made available by the Swedish Museum of Natural History under Creative Commons Attribution 4.0 International Public License, CC-BY 4.0, https://creativecommons.org/licenses/by/4.0/legalcode. 1 dorsal view 2 ventral view 3 labels. Scale bar: 10 mm (1). Forero (2004 , 2006 ) recorded this species from Colombia, based on a unique female. These two females have been the only specimens of V. nigripennis known so far. Forero (2004) argued that the knowledge of the male of the species would be useful to a definition in relation to other members of Sphaeridopinae . Additionally, a female specimen of V. nigripennis from Panama was located in the collection of the NMNH. Upon our request, Dr Silvia A. Justi (WRBU) examined the specimen, sent us photos of it and provided the data on the labels, which are transcribed below. The specimen was identified by the Late P. Wygodzinsky. Although it had been previously coated with metal for electronic microscopy, the identification of the specimen is still possible and represents a new record of this species for Panama. Material examined. Volesus nigripennis . ECUADOR, Esmeraldas, Tundaloma Lodge, near Calderon , 01.18277N, 078.75259W ( 01°10'57"N 78°45'09"W ), 55m a.s.l., 8-9.ii.2014, A. Kury & A. Giupponi leg., 1 male (CEIOC), 1 male (CTJMSB). Additional specimen. Volesus nigripennis . PANAMA: Escobal Road / Atl. Canal Zone / 24 VI [19]74 [handwritten] / Col: D. Engleman // Drake Colln. ex / J. Maldonado C. / Coll 1996 [characters partially cut off at the bottom of the label] // Volesus [handwritten] / nigripennis [handwritten] / Champion [handwritten] / Wygodzinsky [det.], 1 female (NMNH). Description. Male. (Figs 4-57). Measurements: total length to tip of abdomen: 16.9-17.3; to tip of forewings: 16.1-16.5; head (excluding neck, measured in lateral view) length: 2.2; length of anteocular portion (measured in lateral view): 0.5; length of postocular portion (measured in lateral view): 0.7; width across eyes: 1.8; interocular distance, dorsal view: 0.9, ventral view: 0.5-0.6; width of eye, dorsal view: 0.5; ventral view: 0.6; length of eye: 0.6-0.7; distance between external margin of ocelli: 0.7-0.8; distance between ocelli: 0.25; maximum width of ocellus: 0.2-0.25; length of antennifer: 0.7; lengths of antennal segments: I: 2.5; II: 3.8; III: 1.5; IV: 0.9; lengths of labial segments, first visible: 0.3; second visible: 1.7-1.8. Thorax: pronotum: fore lobe, length: 0.8; maximum width: 3.2; hind lobe: length 3.0; maximum width: 5.9; scutellum, length: 2.3; width: 2.7; length of process: 1.1-1.2; length of hemelytra: 12.5. Fore legs: length of femur: 3.8; length of tibia: 4.8-4.7; length of spongy fossa: 0.25; length of tarsus: 1.2-1.3; middle legs, length of femur: 4.5-4.6; length of tibia: 4.8-5.1; length of spongy fossa: 0.25; length of tarsus: 1.2-1.3; hind legs: length of femur: 5.2-5.3; length of tibia: 6.3-6.7; length of tarsus: 1.3. Abdomen, length: 12.5; maximum width: 7.7-7.8. Figures 4-8. Volesus nigripennis , male 4-5 dorsal view 6-8 head 6-7 dorsal view 8 except distal half of second visible labial segment, lateral view. Scale bars: 5.0 mm (4-5); 1.0 mm (6); 0.5 mm (7-8). Coloration: general coloration blackish with reddish markings (Figs 4-5, 14, 20, 26, 28, 35). Head generally blackish; neck mostly reddish; apices of antenniferous tubercles pale; antennal segment II brownish black; antennal segments III-IV brownish; labium brownish (Figs 4-6, 9-11, 14, 20). Thorax blackish, brownish black on meso- and metasternum, with the following reddish thoracic markings: on anterior collar and their projections; on lateral and posterior margins of pronotum; on most of fore lobe of pronotum, except its median portion; on hind lobe of pronotum, a median and a pair of lateral converging bands, which are continuous with reddish posterior margin, ending approximately at mid and anterior thirds of hind lobe, respectively; and on postero-superior portion (approximately) of propleura and process of scutellum (Figs 4-6, 14, 16, 20, 26, 28). Legs generally blackish; spongy fossa on fore and mid tibiae somewhat paler (Figs 4-5, 20, 24-25). Hemelytra black, somewhat paler, brownish, on approximately distal half of clavus, medially and about distal half of the membrane, except veins and area just surrounding them (Figs 4-5, 26). Hind wing generally brownish, with veins darkened (Fig. 27). Abdomen blackish to blackish brown; tergite VI with a median reddish spot just below anterior margin; tergite VII almost completely reddish, blackish on and just below anterior margin and with a pair of rounded blackish spots on mid-lateral portion (Fig. 28). Connexivum reddish on: extreme base of segment II, approximately basal third of segments III-V , and somewhat less than basal half of segment VI; connexival portion of segment VII almost entirely reddish with only posterior border of approximately distal half darkened; ventrally, marking on segment II is a small spot on external margin; on segments III-VI connexival reddish markings are prolonged dorsally to a short distance on lateral portion of respective tergite as a subtriangular marking, and ventrally, as a somewhat curved lateral marking, directed backwards, reaching spiracles, which are surrounded by reddish posterior margin; sternite II with anterior margin and median portion, on approximately distal half reddish to reddish brown; transverse median bands, on sternites III-VII , progressively larger, reddish brown in one specimen and pale brownish in other, joining lateral reddish markings described above in sternites V-VII , the latter almost completely reddish, with dark coloration restricted to anterior margin and adjacent to genital capsule (Figs 4-5, 26, 28, 35). Exposed portion of pygophore and parameres blackish (Fig. 35). Figures 9-13. Volesus nigripennis , male 9 head and left antenna, dorsal view 10 left antenna, ventral view 11 head and fore lobe of pronotum, lateral view (black arrow points to prosternal process; first and second visible labial segments indicated by white arrows) 12, 13 ventral view 12 head, except antenniferous, and most part of prosternum (arrows point to first and second visible labial segments) 13 head, except distal half of second visible labial segment. (1 first 2 second visible labial segments). Scale bars: 1.0 mm (9-11); 0.5 mm (12, 13). Figures 14-19. Volesus nigripennis , male, dorsal view 14 head, pronotum, scutellum and basal portions of hemelytra and connexivum 15-18 pronotum 15 median portion 16-17 fore lobe and basal portion of hind lobe 17 left side and midline 18 lateral right portion, including humeral angle 19 scutellum. Scale bars: 2.0 mm (14); 0.5 mm (15-17, 19); 0.3 mm (18). Figures 20-25. Volesus nigripennis , male 20-23 ventral view 20 head and thorax 21 prothorax, arrow points to prosternal process 22 prothorax and mesosternum 23 metasternum, middle and hind coxae, and median portion of base of abdomen 24, 25 apices of tibiae, arrow points to spongy fossa 24 fore tibia, ventral view 24 middle tibia, lateral view. Scale bars: 1.0 mm (20, 23-25) 0.5 mm (21, 22). Figures 26-28. Volesus nigripennis , male, dorsal view 26 distal portion of pronotum, scutellum, hemelytra and connexivum, arrow points to a small additional cell at approximately apical fourth of the cubital vein 27 left hind wing 28 distal portion of thorax and abdomen (wings moved away). Scale bar: 2.0 mm. Vestiture: body generally covered by sparse short, somewhat curved, adpressed, thin, golden to brownish setae. Head: eyes, ocelli and neck glabrous; region adjacent to insertion of labium with more numerous and somewhat longer setae; ventral surface of first visible labial segment and basal portion of second visible labial segment moderately setose, dorsal surface of correspondent portions with fewer setae; additionally, some sparse setae scattered on the proximal third of second visible segment, remainder glabrous. Antenna: segment I sparsely covered with setae similar to those of general vestiture but slightly longer, more numerous at apex; segments II-IV densely setose, covered with scattered longer, somewhat curved, brownish setae and much more numerous shorter, thinner, whitish setae (Figs 9, 10). Thorax. Some longer straight thin setae on posterior margin of pronotum adjacent to lateral portion of scutellar base; setae are sparser on ventral surface; smooth lateral areas of mesosternum glabrous. Hemelytra: small adpressed setae sparsely scattered on corium, more numerous at its apex; apical two thirds of clavus, respective adjacent area of corium and membrane glabrous. Legs generally with similar vestiture of the body; setae longer and thicker on tibiae, becoming more numerous towards apex; tarsi with stiff, pale, yellowish to golden-yellowish, oblique to curved setae, with variable lengths. Abdomen: tergites I-V almost completely glabrous, with some scattered small darkened or pale setae, almost imperceptible; tergite VI with some more numerous pale setae; tergite VII with scattered longer golden setae. Connexivum: lateral margins with numerous adpressed short curved darkened setae, forming a few irregular rows; these setae become somewhat longer and paler on distal margin of segment VII; segments II-VI dorsally glabrous; some sparse setae on dorsal surface of distal third of segment VII. Sternites generally covered with sparse thin golden to pale setae; somewhat longer and more numerous setae on median portion of segments VI-VII and on pygophore, except its middle portion. Structure: Head. Anteocular portion slightly shorter than postocular portion (in lateral view); ocelli separated by a distance slightly larger than transverse width of each ocellus, positioned medially to level of inner posterior angle of eyes and close to transverse sulcus; antenniferous large; first antennal segment slightly longer than head, stout, somewhat curved, its approximately basal fourth slightly thinner; remaining antennal segments progressively thinner, cylindrical; labium reaching or surpassing the mid third of stridulitrum (Figs 6-14, 20-22). Thorax. Anterior collar inconspicuous; anterolateral angles rounded and small (Figs 15, 16); fore lobe with irregular areas with smooth and whitish integument; a median transverse depression on fore lobe present between medial margins of longer curved smooth areas (Figs 14-17); humeral angles acute, slightly prominent (Figs 14, 18); posterior margin of hind lobe slightly curved on middle third (Figs 14, 15). Scutellum sculptured, median depression shallow, process stout, horizontal, apex rounded (Figs 14, 19). Distance between acute prosternal processes: 0.7. Hemelytra generally dull; on extreme base of dorsal surface, laterally, and on lateral portion, basally, moderately shiny; not reaching tip of abdomen, ending somewhat apically to level of the mid third of seventh tergite (Figs 4-5, 26); in one specimen, the membrane has a small additional cell at approximately apical fourth of cubital vein (Fig. 26). Abdomen. Integument generally also rugose (Figs 28-34), except on median portions of sternites IV-VII , in which it is mostly smooth (Figs 34-38). Connexivum largely exposed, laterally to hemelytra (Figs 4-5); anterior margin of tergite I carinulate (Figs 29-31); tergite II with a mid-longitudinal keel and median third of posterior margin curved backwards (Figs 28-31). Sternites carinulate on anterior margin of segments III-V in one specimen and also on segment VI in the other; on sternite III, canaliculae are somewhat larger and extend more towards lateral portion, occupying approximately two thirds of anterior margin, except midline; on following segments canaliculae become progressively slightly smaller and occupy approximately median third of anterior margin, except midline; a median shallow keel on distal two thirds of segment II and somewhat more elevated in sternites III-VI (Figs 35-38). Segment VIII not visible externally, sclerotized on ventral portion, which becomes somewhat wider towards posterior margin; latter almost straight and with a few short setae; dorsal portion membranous and narrower; spiracles on dorsal margin of ventral portion (Figs 39-41). Figures 29-34. Volesus nigripennis , male 29-33 dorsal view 29 process of scutellum, median portions of tergites I-II and basal half of tergite III, arrow points to midlongitudinal keel of tergite II 30-31 mediolateral portions of tergites I-III 30 arrow points to first (dorsal) abdominal spiracle 31 arrow points to the dag on tergite IV 32 lateral portions of tergites III (distal part), IV-V 33 mediolateral portions of distal part of tergite IV, tergite V and basal part of tergite VI, arrows point to the dag on tergites V and VI 34 abdominal segments III-IV, lateral view, arrows point to the vertical sclerite of connexivum. (dag scar of dorsal abdominal gland opening). Scale bar: 1.0 mm. Figures 35-39. Volesus nigripennis , male 35 abdomen, lateroventral view 36-39 ventral view 36 hind coxa, median portions of sternites II-III and basal portion of sternite IV 37 median portion of distal margin of sternite III, sternites IV-V and anterior margin of sternite VI 38 mediolateral portions of distal half of sternite III and sternites IV-V , except lateroposterior angle of the latter 39 segment VIII and genital capsule detached from abdomen. Scale bars: 2.0 mm (35); 1.0 mm (36-39). Figures 40-44. Volesus nigripennis , male 40 distal half of segment VIII and genital capsule, dorsal view (d dorsal portion of segment VIII; pct proctiger; pdm posterior dorsal margin of pygophore; psp medial prolongation of the socket of the insertion of the paramere; spi spiracle of segment VIII) 41 segment VIII, ventral view 42, 43 genital capsule, lateral view 44 pygophore, ventral view. (mpp median process of pygophore; pa paramere; pm posterior margin of pygophore; sp socket of the insertion of the paramere. Scale bars: 0.5 mm (40, 41); 1.0 mm (42-44). Male genitalia (Figs 35, 39-40, 42-57): genital capsule, in ventral and lateral views: exposed portion of pygophore hemispherical, posterior margin (pm) flattened, integument rugose and setose; non-exposed portion of pygophore less pigmented and less sclerotized, integument smooth and glabrous (Figs 39, 42-44); in dorsal view: between anterior and posterior genital openings, a very well-sclerotized dorsal (transverse) somewhat curved bridge; socket of insertion of paramere (sp) approximately in mid portion of pygophore, with a conspicuous medial prolongation obliquely directed posteriorly (psp); numerous, somewhat long, erect setae inserted on inner surface of basal portion of this prolongation; membranous areas of genital opening smooth; proctiger (pct) somewhat enlarged toward apex, with numerous long setae on distal half; posterior dorsal margin of pygophore (pdm) large, forming a horizontal extension of pygophore wall, with some scattered setae on inner margin and more numerous and somewhat shorter elements on median portion (Fig. 40). Median process of pygophore (mpp) only visible in dorsal and lateral views of pygophore, directed upwards, situated some distance from posterior margin, somewhat enlarged, almost straight and subsquared in dorsal and anterior views, respectively (Figs 40, 42-43, 47). Paramere apices in contact in resting position (Fig. 35); parameres (pa) (Figs 40, 42-43) symmetrical, elongated , with a lateral rounded enlargement just above inserted portion, moderately and strongly curved inwards at mid and apical portions, respectively, narrowing towards tip, which is somewhat rounded (Fig. 45) to acute (Fig. 46); with straight to moderately curved setae, more numerous towards apical portion; setae absent on basal (inserted) portion and on inner surface of approximately basal fourth of the not inserted portion (Figs 40, 43, 45-46). Articulatory apparatus with moderately short basal plate arms (bpa); basal arms and basal plate bridge (bpb) forming a subtriangular set (Fig. 50); basal plate bridge (bpb) slightly bent ventrally (Fig. 50); pedicel (pd) elongated, somewhat enlarged at midportion, curved in lateral view (Figs 48, 49, 51, 52). Before inflation of the endosoma, a lateral oval area (loa) somewhat more sclerotized on endosoma wall is evident (Figs 48-50) as well as a conspicuous dorsal pair of membranous lobes on endosoma (mle), united at their basal median portion which is inserted just above apex of dorsal phallothecal sclerite (dps) (Fig. 50). Each membranous lobe on endosoma (mle) is flattened, elongated, apex rounded, directed outwards, laterally to dorsal phallothecal sclerite (Figs 48, 50-52, 55). Dorsal phallothecal sclerite (dps) elongated, thrice curved in lateral view (Figs 51, 54); in dorsal view , it is narrower at approximately midportion and towards apical portion (Fig. 53); apical margin almost straight (Figs 50, 52, 53, 56); at its subapical enlarged portion there is a pair of symmetrical rounded flat lateral expansions (fle) (Figs 50, 52, 53). After inflation of endosoma, endosoma wall is smooth to longitudinally and transversely finely striated at approximately basal two thirds and coarsely rugose at distal third, with some areas in which the rugosities are more sclerotized (ars) (Figs 56, 57); endosoma wall forming three apical expansions: a median subrounded flat expansion (mfe) and a pair of lateral tubular short expansions (lte), each of the latter with a more sclerotized thin longitudinal line along its length (lsl) (Figs 52, 57). Endosoma with the following processes: a pair of flat, somewhat sclerotized, asymmetrical and striated processes (stp) between apex of dorsal phallothecal sclerite and subapical process (sbp) (Fig. 56). The subapical process (sbp) provided with a pair of sclerotized arms, in which basal halves are shorter, diverge more and are formed by stronger sclerotizations of rugosities of wall, while distal half is somewhat longer, less diverging and formed by linear and aggregate thickenings (Figs 51, 52, 56, 57). Figures 45-48. Volesus nigripennis , male genitalia 45, 46 apical portion of paramere, lateral view 47 apical portion of pygophore, anterior view (setae omitted). (mpp median process of pygophore) 48 phallus not inflated, lateral view. (bpa basal plate arm; dps dorsal phallothecal sclerite; loa lateral oval area; mle membranous lobe on endosoma; pd pedicel). Scale bars: 0.2 mm (45); 0.1 mm (46); 0.5 mm (47, 48). Figures 49-52. Volesus nigripennis , male genitalia, phallus 49, 50 not inflated 49 ventral view 50 dorsal view 51, 52 inflated 51 lateral view 52 laterodorsal view. (adps apex of dorsal phallothecal sclerite; bpa basal plate arm; bpb basal plate bridge; dps dorsal phallothecal sclerite; fle flat lateral expansion; loa lateral oval area; lte lateral tubular short expansion; mle membranous lobe on endosoma; pd pedicel; sbp subapical process). Scale bar: 0.5 mm. Figures 53-57. Volesus nigripennis , male genitalia 53, 55-57 dorsal view 54 lateral view 53, 54 dorsal phallothecal sclerite (dps). (apd apex of pedicel; fle flat lateral expansion). 55 subapical enlarged portion of dorsal phallothecal sclerite (dps) and membranous lobe on endosoma (mle), indicated by arrows 56, 57 endosoma portions 56 subapical 57 apical. (adps apex of dorsal phallothecal sclerite; ars area with rugosities more sclerotized; mfe median subrounded flat expansion; lsl longitudinal sclerotized line; lte lateral tubular short expansion; sbp subapical process; stp flat, somewhat sclerotized, striated processes). Scale bar: 0.2 mm. Distribution. Colombia, Costa Rica, Ecuador (new record), Panama (new record). Comments. Volesus nigripennis is the first Sphaeridopinae recorded for Ecuador and Panama ( Froeschner 1981 , 1999 , Maldonado 1990 ). The male specimens (Figs 4, 5, 20, 35) described here seem to be generally similar to the female of the species in structure and coloration ( Champion 1899 , Forero 2006 ; Figs 1, 2). However, only the examination of more specimens of V. nigripennis will make it possible to ascertain whether there is sexual dimorphism. Smooth areas on the fore lobe of pronotum were recorded here in V. nigripennis (Figs 6, 15-17) but it was not possible to distinguish a paired sensory organ similar to that described in Sphaeridops eulus by Maldonado and Santiago-Blay (1992 : figs 13, 14). These authors emphasized that the nature of the sensory organ of these areas could be seen in their SEM images. However, judging by the SEM images obtained in the present study (Figs 15, 17), it is possible that the supposed sensory organ, also mentioned as present in both species of Veseris ( Gil-Santana et al. 1999 , Gil-Santana and Alencar 2001 ) may be in fact a portion of these smooth areas. Only future studies, preferably employing histological techniques will allow the evaluation of the existence and/or possible sensory functions of such portions in these species. Although Champion (1899) had described that the labium would have the second and third visible labial segments equal in length, our studies, including the SEM images, made it clear that the labium is formed by only two visible segments, with the first visible segment short and enlarged and the other long, thin and straight (Figs 11, 12). It is opportune to mention that, according to our request, Dr Dimitri Forero kindly reexamined the female recorded by him from Colombia, sent us photos and confirmed these same features on the labial segments. Similarly, Dr Silvia A. Justi, when examining the female specimen from Panama, also verified that it had only two visible labial segments, with the same characteristics. Some of the portions of the male genitalia of V. nigripennis , such as the parameres and articulatory apparatus, including a basal plate bridge bent ventrally (Figs 40, 45, 46, 50) seem similar to those recorded for species of Veseris ( Gil-Santana et al. 1999 , Gil-Santana and Alencar 2001 ). Weirauch (2008) recorded the presence of the basal plate bridge (=ponticulus basilaris) bent ventrad and a pair of membranous lobes on endosoma, lateral to the dorsal phallothecal sclerite in Sphaeridops amoenus and Salyavata nigrofasciata Costa Lima, 1935 ( Salyavatinae ). Judging by her drawings, these lobes are smaller in S. amoenus and somewhat larger but shorter in S. nigrofasciata , respectively, than those recorded here in V. nigripennis (Figs 50-52). It is noteworthy that Weirauch (2008) considered both characteristics (a basal bridge bent ventrad and the pair or membranous lobes on the endosoma) as synamoporphies of the clade Salyavatinae + Sphaeridopinae obtained in her cladistic analysis. On the other hand, because all other structures, such as those of phallus and endosoma, were not adequately recorded by the above-mentioned authors, nor by others who included just partial or incomplete descriptions of the male genitalia of species of Sphaeridops (e.g. Maldonado and Santiago-Blay 1992 , Gil-Santana et al. 2000 ), only future comprehensive studies of these structures among Sphaeridopinae will allow useful comparisons with the results obtained here.