Cephalosphaera Enderlein and Neocephalosphaera De Meyer (Diptera: Pipunculidae) of Colombia, with description of nine new species and an updated key to their Neotropical species Author Ramos-Pastrana, Yardany 0000-0002-3193-6659 ya.ramos@udla.edu.co Author Marques, Dayse W. A. 0000-0002-3193-6659 willkenia @ gmail. com; https: // orcid. org / 0000 - 0001 - 7260 - 5760 ya.ramos@udla.edu.co Author Rafael, José Albertino 0000-0002-3193-6659 jarafael @ inpa. gov. br; https: // orcid. org / 0000 - 0002 - 0170 - 0514 ya.ramos@udla.edu.co text Zootaxa 2022 2022-08-29 5178 4 301 333 journal article 133177 10.11646/zootaxa.5178.4.1 15752cb9-d773-46c9-8ed4-930f04bc1d0c 1175-5326 7031688 6E706C39-8F42-4050-8792-0423F4267D2B Key to males of the Neotropical species of Cephalosphaera and Neocephalosphaera This key is a result of the modification of those by Rafael (1992) and Souza & Ale-Rocha (2009) . The initial for genera were adapted from Skevington & Yeates (2001) and Rafael & Skevington (2010) . 1 Hind tibia with a comb of longer setae apically; male abdominal syntergosternite 8 with the membranous area not reaching epandrium ( Fig. 8 )...................................................... Cephalosphaera Enderlein, 1936 ... 2 - Hind tibia without comb of longer setae apically; male abdominal syntergosternite 8 with the membranous area reaching epandrium and almost reaching tergite 5 ( Figs 19, 21 , 32, 34 , 48, 50 , 64, 66 , 77, 79 , 90, 92 , 103, 105 , 118, 120 )................................................................................. Neocephalosphaera De Meyer, 1994 . 13 2 All femora predominantly yellow, sometimes with light brown to brown spots apically or dorsally..................... 3 - All femora predominantly brown to black, sometimes with bases and apices yellow................................ 9 3 All femora completely yellow, apex of phallic guide with a small spine laterally [see figure 3F in Souza & Ale-Rocha (2009) ] or with a rigid lobe laterally [see figure 3N, presented by Souza & Ale-Rocha (2009) ]............................... 4 - All femora yellow, with bases and apices or at least in the middle of the femur with a small spotlight brown to brown dorsoapically; apex of phallic guide with a strong lobe apically [see figure 3H in Souza & Ale-Rocha (2009) ] or a row or tuft of strong spines ventroapically [see figures 3M, 4D, in Souza & Ale-Rocha (2009) ] or a tuft of fine setae ventrally [ Fig. 12 and see figure 3O in Souza & Ale-Rocha (2009) ]........................................................................ 5 4 Both surstyli with tips acute inward directed, with inner margins straight and outer margins curved, with short and fine setae marginally [see figure 2F in Souza & Ale-Rocha (2009) ]; apex of phallic guide with acute lobe apically downward directed and one short spine laterally (see figure 3F in Souza & Ale-Rocha (2009) ]; phallus trifid with two lateral projections apically, giving the false impression of being five ejaculatory ducts [see figure 3F in Souza & Ale-Rocha (2009) ]........................................................................................ C . guanacastensis Rafael & Menezes, 1999 - Both surstyli with subrounded tips, slightly downwards directed, with inner and outer margins curved, without marginal setae [see figure 2N in Souza & Ale-Rocha (2009) ]; apex of phallic guide with a rigid and acute lobe laterally, giving the false impression of being bifid [see figure 3N in Souza & Ale-Rocha (2009) ]; phallus trifid, with one long projection dorsally, forward directed and ejaculatory ducts slightly coiled [see figure 3N in Souza & Ale-Rocha (2009) ]... C . panamensis ( Hardy, 1948 ) 5 Surstyli asymmetrical [see figure 2M in Souza & Ale-Rocha (2009) ]; apex of phallic guide with a tuft of strong and short spine apically (see figure 3M in Souza & Ale-Rocha (2009) ]............................. C . pacaraima Rafael & Rosa, 1991 - Surstyli subsymmetrical [ Fig. 9 and see figures 2H, 2O, 4C in Souza & Ale-Rocha (2009) ]; apex of phallic guide with a strong lobe apically [see figure 3H in Souza & Ale-Rocha (2009) ] or with a row of strong and long spines ventroapically [see figure 4D in Souza & Ale-Rocha (2009) ] or with a tuft of short and fine setae ventroapically ( Fig. 12 and see figure 3O in Souza & Ale-Rocha (2009) ].................................................................................... 6 6 Wing with membrane hyaline; phallus long (see figures 3O, 4.D in Souza & Ale-Rocha (2009) ]....................... 7 - Wing with membrane faintly light brown to brown infuscated; phallus short [ Figs 12 , see figure 3H in Souza & Ale-Rocha (2009) ]............................................................................................. 8 7 Surstyli about equal in length to epandrium, both surstyli equal in length [see figure 4C in Souza & Ale-Rocha (2009) ]; apex of phallic guide with a row of strong and long spines ventroapically (see figure 4D in Souza & Ale-Rocha (2009) ]; phallus with ejaculatory ducts stout, with distal 1/3 distinctly striated (see figure 4D in Souza & Ale-Rocha (2009) ].......................................................................................... C . aurata Souza & Ale-Rocha, 2009 - Surstyli distinctly shorter than the length of epandrium, left surstylus shorter than right (see figure 2O in Souza & Ale-Rocha (2009) ]; apex of phallic guide with a tuft of fine setae ventroapically (see figure 3O in Souza & Ale-Rocha (2009) ]; phallus with ejaculatory ducts thin, with tips slightly coiled (see figure 3O in Souza & Ale-Rocha (2009) ]................................................................................................. C . procera Rafael & Menezes, 1999 8 Postpronotal lobe brown ( Fig. 1 ) both surstyli stout, thickened basally and medially, thin apically, with inner and outer margins sinuous ( Fig. 9 ); apex of phallic guide with a tuft of fine setae ventrally ( Fig. 12 ); phallus somewhat triangle-shaped in lateral view, with ejaculatory ducts slightly coiled ( Fig. 12 ).................................... C . munchiquensis sp. nov. - Postpronotal lobe yellow; both surstyli slender, thickened basally, thin medially and apically, inner and outer margins slightly curved [see figure 2H in Souza & Ale-Rocha (2009) ]; apex of phallic guide with strong lobe apically [see figure 3H in Souza & Ale-Rocha (2009) ]; phallus stem-shaped, with ejaculatory ducts simple, not coiled, backwards directed [see figure 3H in Souza & Ale-Rocha (2009) ]............................................................... C . insularis Rafael, 1996 9 Section between cell dm and vein M 2 equal than vein dm-m ( Fig. 102 ); all femora entirely black; both surstyli with apical sinus on outer margins, forming acute tips, right surstylus shorter than left [see figure 2G in Souza & Ale-Rocha (2009) ]; phallus thin [see figure 3G in Souza & Ale-Rocha (2009) ]......................................... C . incomitata (Hardy, 1965) - Section between cell dm and vein M 2 greater than vein dm-m ( Figs 5 , 18 , 31 , 47 , 63 , 76 , 89 ); all femora brown to black, with bases and apices yellow; both surstyli without apical sinus on outer margins, not forming acute tips, both surstyli equal in length [see figures 2E, 2J, 2L, 2Q in Souza & Ale-Rocha (2009) ]; phallus thickened [see figures 3E, 3J, 3L, 3R in Souza & Ale-Rocha (2009) ]............................................................................................ 10 10 Mesopleuron and mediotergite brown, brown pruinose; surstyli asymmetrical [see figure 2E in Souza & Ale-Rocha (2009) ]; apex of phallic guide triangular shaped [see figure 3E in Souza & Ale-Rocha (2009) ]; phallus stouter apically, one ejaculatory duct with two small protuberances dorsally [see figure 3E in Souza & Ale-Rocha (2009) ]........ C . fairchildi Rafael, 1992 - Mesopleuron and mediotergite dark brown to black, gray pruinose; surstyli subsymmetrical [see figures 2J, 2L, 2Q in Souza & Ale-Rocha (2009) ]; apex of phallic guide rectangular or slightly rectangular shaped [see figures 3J, 3L, 3R in Souza & AleRocha (2009)]; phallus not stouter apically, with ejaculatory ducts simple, without protuberances dorsally (see figures 3J, 3L, 3R in Souza & Ale-Rocha (2009) ]....................................................................... 11 11 Postpronotal lobe yellow; all tibiae brown to black; both surstyli distinctly narrowing in the pre-apical portion, with short and fine setae in the inner and outer margins [see figure 2Q in Souza & Ale-Rocha (2009) ]; apex of phallic guide with tip truncated and one lobe dorsoapically [see figure 3R in Souza & Ale-Rocha (2009) ]; phallus with a dorsal lobe [see figure 3R in Souza & Ale-Rocha (2009) ]...................................................... C . zumbadoi Rafael & Menezes, 1999 - Postpronotal lobe brown to black; all tibiae yellow; both surstyli without distinctly narrowing in the pre-apical portion, without setae in the inner and outer margins [see figures 2J, 2L in Souza & Ale-Rocha (2009) ]; apex of phallic guide with acute tip and without a lobe dorsally apically [see figures 3J, 3L in Souza & Ale-Rocha (2009) ]; phallus simple, without lobe [see figures 3J, 3L in Souza & Ale-Rocha (2009) ]....................................................................... 12 12 Postpedicel yellow, concolor with scape and pedicel; postpronotal lobe brown, gray pruinose; scutellum brown, grayish pruinose; third section costal longer than the length of fourth ( Figs 31 , 76 ); vein dm-m slightly curved ( Figs 5 , 18 , 47 , 76 , 89 , 102 , 117 ); hind femora with ctenidia not prominent; both surstyli with inner margin sinuous and tips truncated inward directed [see figure 2L in Souza & Ale-Rocha (2009) ]; phallus with ejaculatory ducts stout (see figure 3L in Souza & Ale-Rocha (2009) ]................................................................................ C . mocaensis ( Hardy, 1948 ) - Pospedicel yellow, scape, and pedicel brown; postpronotal lobe shining black, brown pruinose; scutellum shining black, brown pruinose; third section costal shorter than the length of fourth ( Figs 5 , 18 , 47 , 63 , 89 , 102 , 117 ); vein dm-m straight ( Figs 31 , 63 ); hind femur with ctenidia very prominent; both surstyli with inner margin slightly curved [see figure 3J in Souza & AleRocha (2009)]; phallus with ejaculatory ducts thin [see figure 3J in Souza & Ale-Rocha (2009) ]..................................................................................................... C . macroctenia Rafael, 1992 13 All femora predominantly yellow to dark yellow, sometimes with distal half or spotlight brown to brown dorsally; both surstyli conical shaped, thickened basally, narrowing medially and apically and inner and outer margins sinuous [ Figs 93 , 121 , and see figures 2K in Souza & Ale-Rocha (2009) ]................................................................. 14 - All femora predominantly brownish yellow to black, sometimes with bases and apices yellow to brown or with brown spot or band dorsally; both surstyli not conical shaped, at least one margin straight or slightly curved ( Figs 22 , 35 , 51 , 67 , 106 , and see figures 2A, 2B, 2C, 2D, 2I, 2P in Souza & Ale-Rocha (2009) ]................................................. 17 14 Postpronotal lobe yellow; all coxae black or at least mid coxa black; fore and hind trochanters yellow, mid trochanter black 15 - Postpronotal lobe whitish yellow or light brown; fore and hind coxae whitish yellow, mid coxae brown; fore and hind trochanter whitish yellow, mid trochanter dark yellow to brown........................................................ 16 15 Postpedicel light yellow, discrepant with scape and pedicel yellow; scutellum yellow; vein r-m located before the basal third of the upper section of the cell dm; section between cell dm and vein M 2 greater than vein dm-m; fore and hind coxae yellow, mid coxa black; all femora with spot brown dorsally; left surstylus with two ventral lobes when seen in lateral view [see figures 9–10 in Rafael & Rosa (1991) ].............................................. N . semispiralis Rafael & Rosa, 1991 - Postpedicel yellow, concolor with scape and pedicel; scutellum dark brown to black; vein r-m located after the basal third of the upper section of the cell dm [see figures 42.22 in Rafael (1992)]; section between cell dm and vein M 2 equal than vein dm-m [see figure 42.22 in Rafael (1992)]; all coxae black; fore femur completely yellow, mid and hind femora with brown discoloration on distal half; left surstylus with subapical sinus when seen in lateral view [see figure 42.58 in Rafael (1992)]....................................................................................... N . miriamae Rafael, 1992 16 Postpronotal lobe whitish yellow ( Fig. 85 ); wing almost hyaline ( Fig. 89 ); section between cell dm and vein M 2 greater than vein dm-m ( Fig. 89 ); both surstyli with tips outward directed, right surstylus slightly shorter and thinner than left ( Fig. 93 ); both surstyli with a small lobe in the proximal third when seen in lateral view ( Figs 94–95 )............... N . spinifera sp. nov. - Postpronotal lobe light brown ( Fig. 114 ); wing faintly brown infuscated ( Fig. 117 ); section between cell dm and vein M 2 smaller than vein dm-m ( Fig. 114 ); both surstyli with tips downward directed, right surstylus slightly longer than left ( Fig. 121 ); both surstyli with a small lobe in the middle third when seen in lateral view ( Figs 122–123 ).......... N . sumapazensis sp. nov. 17 Pospronotal lobe yellow; all femora completely brown, only hind femur with base yellow; fore and mid tibiae completely yellow, hind tibia brown, with white to light yellow ring medially; both surstyli very thickened, with apices thickener than bases [see figure 2D in Souza & Ale-Rocha (2009) ]............................................. N . cristata Rafael, 1992 - Postpronotal lobe brown to black; all femora brownish yellow to black, sometimes with bases dark brown and band dark brown dorsally or bases and apices yellow to light brown or yellow on proximal half; all tibiae completely yellow or fore and hind tibiae completely yellow and hind tibia with brown spot medially or posteriorly; both surstyli not very thickened as above, with apices thinner than bases [ Figs 22 , 35 , 51 , 67 , 80 , 106 , see figures 2A, 2B, 2C, 2I, 2P in Souza & Ale-Rocha (2009) ]..... 18 18 Both surstyli with carinae on apical inner edges [ Figs 22 , 51 , 67 , 106 , see Fig. 2.I , presented by Souza & Ale-Rocha (2009) ]; phallus completely spiralized ( Figs 25 , 109 ) or only from distal 3/4 [ Figs 54 , 70 , and see figure 3I in Souza & Ale-Rocha (2009) ] not forming a knot at the apex.................................................................... 19 - Both surstyli without carinae on apical inner edges [ Figs 35 , 80 and see figures 2A, 2B, 2C, 2P in Souza & Ale-Rocha (2009) ]; phallus slightly spiralized only in distal 1/4 [see figures 3A, 3B, 3P in Souza & Ale-Rocha (2009) ] or forming a knot at the apex [ Fig. 38 , 83 and see figure 3D in Souza & Ale-Rocha (2009) ]................................................. 23 19 Antenna with short acuminate apex [see figure 42.6 in Rafael (1992)]; all trochanters yellow; apex of phallic guide with ventral and dorsal margins slightly straight, without lobes ventrally or dorsally [see figure 3I in Souza & Ale-Rocha (2009) ].................................................................................. N . jamaicensis ( Johnson, 1919 ) - Antenna with long acuminate apex ( Figs 6 , 45 , 61 , 100 ); all trochanters brown to dark brown or at least mid trochanter brown; apex of phallic guide with ventral or dorsal margins sinuous, with a lobe ventrally or dorsally ( Figs 25 , 54 , 70 , 109 )...... 20 20 Both surstyli with acute apices when seen in lateral view ( Figs 23–24 , 68–69 ); apex of phallic guide with a rigid lobe dorsomedially ( Figs 25 , 70 ); ejaculatory apodeme with apex thickened ( Figs 26 , 71 )...................................... 21 - Both surstyli with subrounded apex when seen in lateral view ( Figs 52–53 , 107–108 ); apex of phallic guide with a translucent lobe dorsoapically ( Figs 54 , 109 ); ejaculatory apodeme with apex narrowed ( Figs 55 , 110 ).......................... 22 21 Frons brown pruinose; vein dm-m slightly curved ( Fig. 18 ); syntergosternite 8 without prominent crestlike membranous area ( Figs 15, 19, 21 ); both surstyli with carinae, forming a straight line ventrally when they join ( Fig. 22 ); phallus with ejaculatory ducts completely spiralized ( Fig. 25 )........................................................ N . carinae sp. nov. - Frons gray pruinose; vein dm-m slightly straight ( Fig. 63 ); syntergosternite 8 with prominent crestlike membranous area ( Figs 59–60, 64, 66 ); both surstyli with carinae, not forming a straight line ventrally when they join ( Fig. 67 ); phallus with ejaculatory ducts spiralized, only in distal 3/4 ( Fig. 70 )................................................... N . muisca sp. nov. 22 Frons yellow pruinose; postcranium brown, gray-brown pruinose dorsally ( Fig. 44 ); notopleuron brown, yellow pruinose ( Fig. 43 ); section between cell dm and vein M 2 greater than vein dm-m ( Fig. 47 ); all femora light brown ( Fig. 43 ); syntergosternite 8 with prominent crestlike membranous area ( Figs 43–44, 48, 50 ); phallus with ejaculatory ducts spiralized, only in distal 3/4 ( Fig. 54 ); ejaculatory apodeme somewhat nail-shaped ( Fig. 55 )............................... N . iguaquensis sp. nov. - Frons light brown pruinose; postcranium dark brown to black, brown pruinose dorsally ( Fig. 99 ); notopleuron black, gray pruinose; section between cell dm and vein M 2 equal than vein dm-m ( Fig. 102 ); all femora dark brown to black ( Fig. 98 ); syntergosternite 8 without prominent crestlike membranous area ( Figs 98–99, 103, 105 ); phallus with ejaculatory ducts completely spiralized ( Fig. 109 ); ejaculatory apodeme somewhat funnel-shaped narrowed ( Fig. 110 ).............. N . spiralis sp. nov. 23 Postpronotal lobe brown ( Fig. 27 ); scutellum brown ( Figs 28, 30 , 73, 75 ); third section costal longer than the length of fourth ( Figs 31 , 76 )........................................................................................ 24 - Postpronotal lobe black; scutellum black; third section costal shorter or equal than the length of fourth ( Figs 5 , 18 , 47 , 63 , 89 , 102 , 117 )........................................................................................... 25 24 Frons yellow pruinose; vein dm-m straight ( Fig. 31 ); all trochanters light brown; all femora brownish yellow, with dark brown bases and dark brown band dorsally ( Figs 27–28 ); both surstyli with apices forming a distinctive rounded lobe, tips downward directed, never converging ( Fig. 35 ); apex of phallic guide without lobes dorsally and ventrally, tip upward directed ( Fig. 38 ); ejaculatory apodeme fan-shaped ( Fig. 39 ); phallus with ejaculatory ducts spiralized, only in distal 3/4 ( Fig. 38 ).................................................................................................. N . grisea sp. nov. - Frons gray pruinose; vein dm-m slightly curved ( Fig. 76 ); all trochanters yellow, all femora brown, yellow on proximal half ( Figs 72–73 ); both surstyli with apices forming a distinctive acute lobe, tips inward directed that converging apically ( Fig. 80 ); apex of phallic guide with distinct translucent lobes dorsally and ventrally, tip downward directed ( Fig. 83 ); ejaculatory apodeme funnel-shaped ( Fig. 84 ); phallus with ejaculatory ducts spiralized, only in distal 1/4 ( Fig. 83 )... N . paolae sp. nov. 25 Section between cell dm and vein M 2 equal than vein dm-m [ Figs 102 , and see figures 42.14, 42.25 in Rafael (1992)]; all trochanters brown to black; both surstyli with apices acute or slightly acute [see figures 2A, 2P in Souza & Ale-Rocha (2009) ]25 - Section between cell dm and vein M 2 smaller or greater than vein dm-m ( Figs 5 , 18 , 31 , 47 , 63 , 76 , 89 ); all trochanters yellow; both surstyli with apex subrounded [see figures 2B, 2C in Souza & Ale-Rocha (2009) ]............................. 26 25 Vein r-m located before the basal third of the upper section of the cell dm [see figure 42.14 in Rafael (1992)]; all trochanters black; all femora black with bases and apices yellow; all tibiae yellow; both surstyli equal in length [see figure 2A in Souza & Ale–Rocha (2009)]; apex of phallic guide with a rigid lobe and a tuft of short setae ventrally [see figure 3A in Souza & Ale–Rocha (2009)]................................................................. N. aequatorialis ( Becker, 1919 ) - Vein r-m located after the basal third of the upper section of the cell dm [see figure 42.25 in Rafael (1992)]; all trochanters brown; all femora completely black; fore and mid tibiae yellow, hind tibia with a distinctly black spot medially; left surstylus slightly thinner and longer than right [see figure 2P in Souza & Ale-Rocha (2009) ]; apex of phallic guide without a rigid lobe and with a tuft of long setae ventrally [see figure 3A in Souza & Ale-Rocha (2009) ]........ N . santiagonensis Rafael, 1992 26 Postpedicel black concolor with scape and pedicel, with short acuminate apex [see figure 42.2 in Rafael (1992)]; third section costal shorter than the length of fourth [see figure 42.15 in Rafael (1992)]; section between cell dm and vein M 2 smaller than vein dm-m [see figure 42.15 in Rafael (1992)]; vein dm-m slightly curved [see figure 42.15 in Rafael (1992)]; all coxae black; all femora black with bases and apices yellow; hind tibia with erect setae medially; left surstylus slightly longer than right [see figure 3B in Souza & Ale-Rocha (2009) ]; phallus with ejaculatory ducts slightly coiled at apex, not forming knot [see figure 3B in Souza & Ale-Rocha (2009) ]........................................................ N . arnaudi Rafael, 1992 - Postpedicel yellow, discrepant with scape and pedicel brown, with long acuminate apex [see figure 42.3 in Rafael (1992)]; third section costal equal in length to fourth [see figure 42.16 in Rafael (1992)]; section between cell dm and vein M 2 greater than vein dm-m [see figure 42.16 in Rafael (1992)]; vein dm-m straight [see figure 42.16 in Rafael (1992)]; all coxae brown; all femora brown with bases and apices yellow; hind tibiae without erect setae medially; both susrtyli equal in length [see figure 3C in Souza & Ale-Rocha (2009) ]; phallus with ejaculatory ducts forming a knot at the apex [see figure 3D in Souza & Ale-Rocha (2009) ]....................................................................... N . boutropis ( Hardy, 1965a )