Taxonomic review of Cerozodus Bigot, 1857 (Diptera: Asilidae: Asilinae) with the description of two new species Author Camargo, Alexssandro 0000-0001-7408-7223 Instituto Nacional de Pesquisas da Amazônia-INPA, Coordenação de Biodiversidade-COBIO, Manaus, Amazonas, Brazil. alexssandroecs @ gmail. com; https: // orcid. org / 0000 - 0001 - 7408 - 7223 alexssandroecs@gmail.com Author Vieira, Rodrigo Secretaria de Estado de Educação do Governo do Amazonas-SEDUC, Manaus, Amazonas, Brazil / Instituto Nacional de Pesquisas da Amazônia-INPA, Coordenação de Biodiversidade- COBIO, Manaus, Amazonas, Brazil. Author Rafael, José Albertino 0000-0002-0170-0514 Instituto Nacional de Pesquisas da Amazônia-INPA, Coordenação de Biodiversidade-COBIO, Manaus, Amazonas, Brazil. jarafael @ inpa. gov. br; https: // orcid. org / 0000 - 0002 - 0170 - 0514 jarafael@inpa.gov.br text Zootaxa 2022 2022-11-16 5209 2 151 186 journal article 188711 10.11646/zootaxa.5209.2.1 40304ff0-808f-4371-a80c-4f0b7c27232f 1175-5326 7325889 710CB5B2-3A7A-491D-A317-CC05419F7A6B Cerozodus nodicornis ( Wiedemann, 1828 ) ( Figs 5–12 , 23 ) Asilus nodicornis Wiedemann, 1828: 469 ; Schiner, 1866: 687 (species list); Williston, 1891: 91 (catalogue). Erax nodicornis ; Macquart, 1838: 224 (species list). Erax ? nodicornis ; Walker, 1855: 638 (key and species list). Cerozodus nodicornis ; Bigot, 1857: 532 , 543 (classification and key); Kertész, 1909: 244 (catalogue); Bromley, 1946: 115 (catalogue); Carrera, 1946: 250 , figs. 1–14 ( neotype designation and redescription); Hull, 1962 (2): 484, figs. 372, 712, 1399, 1408, 2442, 2443 (synopsis of world fauna); Martin & Papavero, 1970: 72 (catalogue); Schumann, 1973: 138 (type catalogue); Artigas & Papavero, 1995: 36 , figs. 1–11 (key and catalogue); Papavero, 2009: 30 (catalogue); Vieira, Rafael & Limeira-de-Oliveira, 2013: 181 , figs. 1–13 (revision); Lamas & Camargo, 2021 (CTFB catalogue). Diagnosis. Face yellowish pruinose; mystacal macrosetae whitish with a few brown macrosetae above ( Figs 5C–D , 7C–D ); scape and pedicel yellow; postpedicel black, discoidal or rounded; second article of stylus with a distinct, ventral, rounded lobe (in males) ( Figs 6A–B , 8A–B ); wings yellowish translucent ( Figs 6C , 8C ); legs dark yellow to brown ( Figs 5A , 7A ); terminalia dark yellow to reddish brown; epandrial arms slender, diminishing and smoothly inwardly curved apically ( Figs 6E–G , 8E–G ); hypandrium rectangular with a tongue-like projection at mid-posterior margin, rounded distally ( Fig. 9D ); gonocoxite with a ventral, sub-apical U-shaped indentation ( Fig. 10D–E ) and laterally with a very short, sub-apical conic projection ( Fig. 10B–C ); gonostylus slender ( Fig. 10F ). Redescription. Holotype male ( Figs 5–6 ). Head ( Fig. 5C–D ): eyes black; face reddish brown with yellowish pruinosity becoming whitish on lower facial margin and lower parafacial area; mystacal macrosetae and setae whitish pale with a few brown macroseta above and on lower parafacial area; gena black, whitish and yellowish pruinose; palpus yellow, darker at tip, dark yellow setose, whitish ventrally; proboscis black with dark reddish tip and yellow at base; labial and proboscial setae whitish; frons black with yellowish pruinosity, apruinose small spot above antennal socket; orbital setae dark yellow to brown; ocellar tubercle black, sparsely yellowish pruinose with two proclinate black setae arising shortly behind posterior ocelli; vertex yellowish pruinose; occiput whitish pruinose; 10 yellowish to dark yellow postocular macrosetae; occipital and lower occipital setae white. Antenna ( Fig. 6A–B ): scape and pedicel yellow, scape with apex dark brown; scape and pedicel brown setose, except scape ventrally yellow setose; scape and pedicel scarcely yellow pruinose; postpedicel slightly shorter than scape, rounded in lateral view, laterally compressed with truncate apex, black, except connection with pedicel dark yellow, scattered brown pruinose, with 4–5 extremely short pale setae mid-dorsally; stylus black, first article tubular, extremely short; second article with a basal rounded, laterally compressed projection, ventrally directed, almost as large as postpedicel, remainder of article thin, only slightly enlarged sub-apically and tapering towards the apex, tip dark yellow. Thorax ( Figs 5A–B , 6D ): brown; paramedian stripes and pre and postsutural spots dark brown to black pruinose; scutum anterolaterally, laterally and following paramedian stripes until scutellum brown pruinose; postalar wall and scutellum greyish brown pruinose ( Fig. 6D ); postpronotal lobe reddish and brown pruinose; mesothoracic spiracle dark yellow; pleura reddish brown, except anterior half of anepisternum, anterior and ventral half of katepisternum and mid-ventral spot on metanepisternum black; pleura greyish brown pruinose. Chaetotaxy:antepronotal macrosetae dark yellow and setae whitish; 2 notopleural; 1 supra alar; 1 postalar and two pairs of postsutural dorsocentral black macrosetae; scutum with short black acrostichal setae and thin pale yellow setae before postalar callus and on posterior scutal margin; scutellum with short, sparse, yellow discal scutellar setae ( Fig. 6D ); katatergite with yellow macrosetae and whitish setae; anepisternum dorsally with short black setae before wing insertion and a few whitish setae posteroventrally; proepisternum whitish setose; remainder of pleura with a few scattered whitish setae. Wings ( Fig. 6C ): yellowish translucent; veins dark brown, except veins Sc , R 1 and veins at base of wing reddish brown; bifurcation of R 4 and R 5 beyond discal cell at a distance equivalent to the length of one crossvein r-m ; crossvein r-m shortly after middle of discal cell; m 3 and cua closed before wing margin. Halteres: pale yellow, base of stem and knob slightly darker. Legs ( Fig. 5A ): dark yellow to brown; fore coxa reddish brown, mid and hind black; coxae greyish brown pruinose; trochanters dark yellow with small black spot at tip; femora dark yellow, brownish anteriorly and dorsally; tibiae dark yellow with apical fourth brown; tarsi brownish with tip of tarsomeres brown, tarsi with last tarsomere almost black on apical half. Chaetotaxy: coxae with pale whitish macrosetae and setae; hind trochanter with 3 pale yellow macrosetae; fore femur with 9–10 pale yellow ventral macrosetae; mid femur with 7–8 pale yellow ventral macrosetae, 4–5 anteroventral, 3 anterior and 1 posterodorsal apical black macrosetae; hind femur with 5 anteroventral, 3 anterodorsal (basal one on right side yellow), 1 dorsal preapical, 3 posterodorsal (1 preapical yellow, 2 apical black), 3 posteroventral (yellow) and 1 ventral (yellow) black macrosetae; fore and mid femora with anterior and dorsal setae black, ventral and posterior pale yellow, on hind femur only anteroventral setae black remainder pale yellow; tibiae with black macrosetae and black setae anteriorly, other setae pale yellow; tarsi with black macrosetae and setae, except by a few sparse pale yellow setae on first tarsomeres; brush setae yellowish; pulvilli dark yellow; empodium reddish yellow; claws black, except basal extremity reddish. Abdomen ( Fig. 5A–B ): brown to dark brown; greyish brown pruinose; T1 with 3 pale yellow lateral marginal macrosetae and pale yellow seta laterally and on posterior margin; T2–7 with 4 pale yellow lateral marginal macrosetae; T8 with 5–6 black lateral marginal macrosetae; T2–8 with short mid-dorsal black setae and laterally with short pale yellow setae; sternites reddish brown with greyish pruinosity, and sparsely whitish to pale yellow setose; S8 with 5–6 pale yellow (inner macrosetae black) macrosetae. FIGURE 5. Cerozodus nodicornis ( Wiedemann, 1828 ) , holotype male. A. Habitus, lateral (insert: labels, not to scale); B. Same, dorsal; C. Head, frontal; D. Same, lateral. Scale bar 1 mm. FIGURE 6. Cerozodus nodicornis ( Wiedemann, 1828 ) , holotype male. A. Postpedicel and stylus, anterior; B. Same, dorsal; C. Wing, dorsal; D. Scutellum, dorsal; E. Terminalia, lateral; F. Same, dorsal; G. Same, ventral. Abbreviations: cerc: cercus; epand: epandrium; f a styl: first article of stylus; goncx: gonocoxite; gonst: gonostylus; hypd: hypandrium; ped: pedicel; pped: postpedicel; s a styl: second article of stylus; st: sternite; subepand scl: subepandrial sclerite; tg: tergite. Scale bar 1 mm. FIGURE 7. Cerozodus nodicornis ( Wiedemann, 1828 ) , Carrera’s neotype male. A. Habitus, lateral (insert: labels, not to scale); B. Same, dorsal; C. Head, frontal; D. Same, lateral. Scale bar 1 mm. FIGURE 8. Cerozodus nodicornis ( Wiedemann, 1828 ) , Carrera’s neotype male. A. Postpedicel and stylus, anterior; B. Same, dorsal; C. Wing, dorsal; D. Scutellum, dorsal; E. Terminalia, lateral; F. Same, dorsal; G. Same, ventral. Abbreviations: cerc: cercus; epand: epandrium; f a styl: first article of stylus; goncx: gonocoxite; gonst: gonostylus; hypd: hypandrium; ped: pedicel; pped: postpedicel; s a styl: second article of stylus; st: sternite; tg: tergite. Scale bar 1 mm. FIGURE 9. Cerozodus nodicornis ( Wiedemann, 1828 ) , additional material, male. A. Terminalia, lateral; B. Same, dorsal; C. Same, ventral; D. Hypandrium, ventral; E. Tergite 8, dorsal; F. Sternite 8, ventral; G. Epandrium, dorsal; H. Same, ventral; I. Same, lateral. Abbreviations: cerc: cercus; epand: epandrium; goncx: gonocoxite; gonst: gonostylus; hypd: hypandrium; ph: phallus; subepand scl: subepandrial sclerite. Scale bar 1 mm. FIGURE 10. Cerozodus nodicornis ( Wiedemann, 1828 ) , additional material, male. A. Terminalia, lateral; B. Gonopods, dorsal; C. Same, ventral; D. Same, lateral; E. Gonocoxite, lateral; F. Gonostylus, lateral; G. Phallus, lateral; H. Cercus and subepandrial sclerite, lateral; I. Same, ventral. Abbreviations: cerc: cercus; ej apod: ejaculatory apodeme; goncx: gonocoxite; goncx apod: gonocoxal apodeme; gonst: gonostylus; ph: phallus; subepand scl: subepandrial sclerite. Scale bar 1 mm. FIGURE 11. Cerozodus nodicornis ( Wiedemann, 1828 ) , Carrera’s allotype female. A. Habitus, lateral (insert: labels, not to scale); B. Same, dorsal. Scale bar 1 mm. FIGURE 12. Cerozodus nodicornis ( Wiedemann, 1828 ) , additional material, female. A. Postpedicel and stylus, anterior; B. Terminalia, dorsal; C. Same, dorsal; D. Same, lateral; E. Same, after dissection, lateral; F. Spermathecae, dorsal; G. Detail of genital fork, dorsal; H. Same, lateral. Abbreviations: acc gl: accessory glang; apod: apodeme of genital fork; cerc: cercus; f a styl: first article of stylus; gen fk: genital fork; lat sc: lateral sclerite of genital fork; med sc: medial sclerite of genital fork; ped: pedicel; pped: postpedicel; s a styl: second article of stylus; spe du: spermathecal duct; spe pu: spermathecal pump; spmth: spermathecae; st: sternite; tg: tergite. Scale bar 1 mm. Terminalia ( Figs 6E–G , 8E–G , 9–10 ): dark yellow to reddish brown; epandrium reddish brown at base and laterally ( Fig. 6E ), apical half and ventrally dark yellow, inner apical margin of epandrial arms with a thin black band ( Fig. 6E–F ); cercus and subepandrial sclerite brown; gonocoxite dark yellow, except anterodorsally reddish brown ( Fig. 6E ); gonostylus and hypandrium dark yellow; epandrium with short dorsal and lateroventral apical black setae, other setae pale yellow; cercus yellowish setose; hypandrium with pale yellow setae, except ventrally and apically with black setae; gonocoxite without setae; epandrial arms slender, diminishing and smoothly inwardly curved apically ( Figs 9B–C, 9G–H ), rounded and with a smaller ventral rounded projection in lateral view ( Figs 9A, 9I ); cercus conical; subepandrial sclerite conical with a small triangular projection at base laterally ( Fig. 10H–I ); hypandrium rectangular with a tongue-like projection at mid-posterior margin, rounded distally ( Fig. 9D ); gonocoxite rounded and slightly inwardly curved apically ( Fig. 10B–C ) with a ventral, sub-apical U-shaped indentation ( Fig. 10D–E ) and laterally with a very short, sub-apical conic projection ( Fig. 10B–C ); gonostylus slender with two very short, comma-like projections, one on external margin sub-apically and other mid-ventrally, sparsely covered with short setae dorsally ( Fig. 10F ). Length. Body: 14.9 mm ; wing: 9.6 mm . Holotype condition. Scutum with a hole posteriorly (after pin) ( Fig. 5B ); right wing with cell m 2 damaged at posterior margin ( Fig. 6C ); specimen dusty, mainly abdomen and terminalia ( Fig. 5–6 ). Remarks. The description of the external characters of the male terminalia is based on the holotype ( Fig. 6E–G ), but characters after dissection are based on additional specimens from Batatais, São Paulo ( Fig. 9–10 ). The specimen designated by Carrera (1946) as a neotype ( Fig. 7–8 ) was not dissected but its external morphological characters were compared with the redescription of the holotype and other specimens and observed differences were commented on in the variation section. Female ( Fig. 11–12 ). Similar to the male ( Fig. 11 ), except that the second article of stylus with only a small dilatation basally, almost indistinguishable in some specimens ( Fig. 12A ), contrasting with the broad basal projection present in males ( Figs 6A–B , 8A–B ); terminalia shining black ( Fig. 12B–D ); T8 and T9+10 dark reddish laterally; S8 reddish basally and apically; T8, T9+10 with few scattered, short yellow setae; cercus with yellow setae; S8 with short yellow setae proximally mixed with some longer black setae; two spermathecae ( Fig. 12F ) reaching T7 ( Fig. 12E ); spermathecal ducts ending separately at genital fork ( Fig. 12G–H ); apodeme of genital fork approximately ¼ of the length of lateral sclerites ( Fig. 12F ), conical with triangular apex proximally ( Fig. 12G ) and a concave indentation distally ( Fig. 12G ), in lateral view with a dorsal keel resembling a Portuguese man o’ war ( Fig. 12H ); medial sclerite present ( Fig. 12F ). Variation. Carrera (1946) mentioned in his redescription that some degree of variation was present on the lobular expansion at the base of the second article of stylus. In some males it is as large as the postpedicel and in others as large as almost half of the postpedicel ( Figs 6A–B , 8A–B ). We also observed small variations on the shape of the second article of the stylus which is reduced and more enlarged in a few males, and also in the shape of the apex of the postpedicel which can be rounded, truncate or with its ventral margin more pronounced in the direction of the lobular expansion of the second article of the stylus; mystax with black macrosetae and only a few white macrosetae restricted to the oral border. Taxonomic discussion. As pointed out by Vieira et al . (2013) , Carrera (1946) wrongly designated a neotype and allotype for C. nodicornis believing that the type material of this species had been lost. However, Schumann (1973) listed the type in the MfN collection. Thus, according to article 75.8 of the ICZN (1999), if the name-bearing type presumed to be lost is found still to exist, the rediscovered material becomes again the name-bearing type and the neotype is set aside. In this work, the redescription of C. nodicornis was based on the holotype described by Wiedemann (1828) . The neotype and additional specimens identified by Carrera (1946) from Batatais, São Paulo state , were also compared and corroborated with the holotype described by Wiedemann (1828) . The male and female specimens dissected in this work are also from Batatais, São Paulo . Distribution. Brazil ( Mato Grosso , Mato Grosso do Sul , Goiás (new record) and São Paulo ) ( Fig. 23 ). Material examined. Holotype . Brasilien [ BRAZIL ] / 2040 / Holotypus / c2ed22 { QR Code } ( Fig. 5A [insert]) ( ♂ MfN ) . Additional material. [ BRAZIL ]: Faz. [enda] Aceiro , Jataí [ 17°52′51″S 51°42′50″W ], Goiás —Brasil, x.1962 Exp. [edição] Dep. [artamento] Zool. [ogia] ( 1 ♀ MZUSP ) ; BRASIL : MS[ Mato Grosso do Sul ] : Rio Verde , S 18°09′38.7″ W 54°09′00.1″ Malaise 39, 30.viii–14.ix.2012 Lamas , Nihei & eq. col. SISBIOTA CNPq/FAPESP ( 1 ♂ MZUSP ) ; Corguinho , Taboco , Reserva Quinta do Sol , S 19°46′36.8″ W 55°14′55.7″ , Malaise 14, 29.viii– 29.ix.2012 ( 4 ♂ & 4 ♀ MZUSP ) ; Same data, 29.ix–12.x.2012 ( 2 ♂ & 1 ♀ MZUSP ) ; 12.x–12.xi.2012 ( 2 ♂ & 2 ♀ MZUSP ) ; 12–29.xi.2012 ( 1 ♂ & 2 ♀ MZUSP ) ; S 19°46′40.8″ W 55°14′59.0″ , Malaise 12, 29.viii–12.ix.2012 ( 2 ♂ & 6 ♀ MZUSP ) ; 12–29.ix.2012 ( 7 ♂ & 6 ♀ MZUSP ) ; 29.ix–12.x.2012 ( 3 ♂ & 4 ♀ MZUSP ) ; S 19°46′45.4″ W 55°14′36.2″ , Malaise 13, 29.viii–12.ix.2012 ( 5 ♂ & 3 ♀ MZUSP ) ; 12.ix–29.ix.2012 ( 7 ♂ & 2 ♀ MZUSP ) ; [ BRASIL ], S.[ão] Paulo , Batatais [ 20°53′27″S 47°35′06″W ], Pde.[Padre] Pereira col. x.[1]945 / Cerozodus nodicornis (Wied., 1828) M. Carrera det. [19]45 / Neotipo [ neotype ] / MZ 001357 ( Fig. 7A [insert]) ( 1 ♂ MZUSP ) ; S.[ão] Paulo , Batatais , Pde. [ Padre ] Pereira col. x.[1]945 / Cerozodus nodicornis (Wied., 1828) M. Carrera det. [19]45 / Alotipo [ allotype ] ( Fig. 11A [insert]) ( 1 ♀ MZUSP ) ; S.[ão] Paulo , Batatais , Pde. [ Padre ] Pereira col.[etor] x.[1]945 ( 14 ♀ MZUSP ) ; Same data, Cerozodus nodicornis (Wied., 1828) M. Carrera det. [19]45 ( 1 ♂ {abdomen damaged}, 2 ♀ & 1 {abdomen damaged, not possible to distinguish the sex} MZUSP ) ; ix.[1]945 ( 1 ♂ & 2 ♀ MZUSP ) ; Batatais, S. [ão] P.[aulo] Brasil ; x.1946 Pe. [ Padre ] Pereira col.[etor] ( 6 ♂ & 7 ♀ MZUSP ) .