Simulium (Trichodagmia) (Diptera, Simuliidae) phylogeny revisited: the Neotropical and Afrotropical connection Author Molina, Óscar S. A05ECC62-2739-4D86-99FE-8ADF624D6FC6 Instituto Oswaldo Cruz - IOC / Fiocruz, Laboratório de Simulídeos e Oncocercose (LSO / IOC- Fiocruz), Coleção de Simulídeos do IOC - CSIOC / Fiocruz, Referência Nacional em Simulídeos, Oncocercose e Mansonelose, Av. Brazil, 4.365, Rio de Janeiro, RJ, Brazil. o.sanchezmolina@gmail.com Author Gil-Azevedo, Leonardo H. 0EC71A47-C1C7-47B0-9D0E-C6BFF6A32C65 Departamento de Entomologia. Museu Nacional-Universidade Federal do Rio de Janeiro, Quinta da Boa Vista s / n, 20940 - 040, São Cristóvão, Rio de Janeiro, RJ, Brazil. lhgazevedo@gmail.com text European Journal of Taxonomy 2021 2021-09-30 773 80 119 journal article 4110 10.5852/ejt.2021.773.1517 a5546d7c-09e9-4dd2-982b-e419c3f047e0 2118-9773 5542069 C69631FE-E71B-4782-8399-1AD84527DA7E Simulium ( Hemicnetha ) Enderlein, 1934 Figs 9–12 , 19–20, 26 , 41–42 , 60–62 , 80–81 , 99–102 , 111, 122–123 , 140–143 , 153–154 Hemicnetha Enderlein, 1934b: 190 (as genus). Type species: Hemicnetha mexicana Enderlein, 1934 [= Simulium paynei ( Vargas , 1942) (subst. name) nom. nov.] Dyarella Vargas , Martínez Palacios & Díaz Nájera, 1946: 105 , figs 8–28 (as subgenus of Simulium ). Type species: Simulium mexicanum Bellardi 1982 [= Simulium tarsatum Macquart, 1846 (1844) ]. Species Simulium brachycladum Lutz & Pinto, 1932 S. bricenoi Vargas , Martínez Palacios & Díaz Nájera, 1946 S. cristalinum Coscarón & Py-Daniel, 1989 S. earlei Vargas , Martínez Palacios & Díaz Nájera, 1946 S. freemani Vargas & Díaz Nájera, 1949 S. guerrerense Vargas & Díaz Nájera, 1956 S. hieroglyphicum Peterson, Vargas & Ramírez-Pérez, 1988 S. hinmani Vargas , Martínez Palacios & Díaz Nájera, 1946 S. hippovorum Malloch, 1914 S. lobatoi Luna Dias, Hernández, Maia-Herzog & Shelley, 2004 S. paynei Vargas , 1942 S. pulverulentum Knab, 1915 S. rubrithorax Lutz, 1909 S. smarti Vargas , 1946 S. solarii Stone, 1948 S. tarsale Williston, 1896 S. tarsatum Macquart, 1846 S. virgatum Coquillett, 1902 S. yepocapense Dalmat, 1949 . Diagnosis Female Scutum black to dark brown (e.g., S . pulverulentum , S . earlei , S . hieroglyphicum , S . bricenoi , S . tarsatum ) or red to light brown (e.g., S . brachycladum , S . cristalinum , S . lobatoi , S . paynei , S . rubrithorax , S . virgatum ); scutal pattern present. Cibarium unpigmented anteromedially, armed with sharp teeth ( Figs 9–10, 12 ) or unarmed ( Fig. 11 ). Tarsal claw with subbasal tooth shorter than a third of claw length (except in S . lobatoi , in which the tooth has about half of claw length). Hypoginial valves fine and pointed ( Fig. 19 ) or subtriangular ( Fig. 20 ), longer than sternite VIII width (except in S. lobatoi , in which are shorter), forwardly directed. Anal lobe subrectangular (except in S . lobatoi , in which is hemispheric with curved distal border), twice as long as cercus width at base. ( Fig. 26 ). Male Ventral plate two times longer than wide or with sub-equal width and length, not notched (except in S . arlei ); lateral shoulders absent ( Fig. 42 ) or present ( Fig. 41 ), in this case, not projected (except in S . lobatoi ); posteromedian process absent ( Fig. 42 ) or present ( Fig. 41 ), in this case, is cylindrical and longer than ventral plate width (except in S . lobatoi , in which is smaller). Gonocoxite longer than wide, with base almost equal in width to gonostylus width at their point of intersection ( Figs 60–62 ). Gonostylus cylindrical, with medial region wider than basal region; distinctly longer than gonocoxite, with blunt apex and apical spicule ( Figs 60–62 ). Parameres with both distinctly large and small accessory spines. Figs 71–90. Pupae. 71–75 . Simulium ( Anasolen ) Enderlein, 1930 . 71 . ambositrae Grenier & Grjebine, 1959 . 72 . dentulosum Roubaud, 1915 . 73 . imerinae Roubaud, 1905 . 74 . iphias De-Meillon, 1951 . 75 . nili Gibbins, 1934 . — 76–77 . S . ( Freemanellum ) Crosskey, 1969 . 76 . debegene De-Meillon, 1934 . 77 . manense Elsen & Escaffre, 1976 . — 78–79 . S. ( Hearlea ) Vargas et al. 1946 . 78 . canadense Hearle, 1932 . 79 . capricorne De León, 1945 . — 80–81 . S . ( Hemicnetha ) Enderlein, 1934. 80 . paynei Vargas, 1942 . 81 . rubrithorax Lutz, 1909 . — 82–83 . S ( Shewellomyia ) Peterson, 1975. 82 . claricentrum Adler, 1990 . 83 . pictipes Hagen, 1880 . — 84–85 . S . ( Obuchovia ) Rubtsov, 1947 . 84 . albellum Rubtsov, 1947 . 85 . galloprovinciale Giudicelli, 1963 . — 86–88 . S . ( Trichodagmia ) Enderlein, 1934. 86 . nigrimanum Macquart, 1838 . 87 . orbitale Lutz, 1910 . 88 . scutistriatum Lutz, 1909 . — 89–90 . S . ( Disculter ) subgen. nov. 89 . rivasi Ramírez-Pérez, 1971 . 90 . oviedoi Ramírez-Pérez, 1971 . Scale bars = 1 mm. Pupa Cocoon aperture upward directed ( Figs 80–81 ),separated from substratum, corbicular ( Fig.81 ) or not. Gill shorter than pupa body, with trunk developed and base enclosed by cocoon anterior margin ( Figs 80–81 ) (except in S . lobatoi , in which the gills are totally free); with 6–90 filaments arranged in different planes (tridimensional); filaments flexible, cylindrical, with apices blunt (e.g., S . hieroglyphicum , S. bricenoi , S. paynei , S. rubrithorax , S. virgatum , S. tarsatum ) or acute (e.g., S. brachycladum , S. cristalinum , S. lobatoi , S . pulverulentum , S. earlei ), concolorous with rest of gill; inferior branch multi-branched ( Figs 99–100 ) or single-branched ( Figs 100–101 ). Thoracic trichomes generally simple. Larva (last instar) Cephalic apotome spots insertion area pigmented ( Figs 122–123 ) (except in S. lobatoi ). Postocciput extended ( Fig. 122 ) over the cervical sclerites, or not ( Fig. 123 ). Antenna ranging from equal in length to or shorter than labral fan stalk (e.g., S. earlei , S. bricenoi , S. paynei , S. rubrithorax , S . tarsatum ), to longer (e.g., S . brachycladum , S. cristalinum , S. lobatoi , S. pulverulentum , S. hieroglyphicum ), without secondary annulations. Mandibles with preapical teeth as long as apical tooth; two mandibular serrations of different sizes. Postgenal cleft triangular; postgenal bridge length less than hypostomal length ( Figs 140–143 ). Hypostoma anterior margin convex ( Fig. 154 ) or straight ( Fig. 153 ); hypostomal teeth extended beyond anterior margin of ventral wall of hypostoma; median tooth distinctly longer than lateral ones ( Fig. 154 ) or at same level ( Fig. 153 ); lateral serrations well-developed, blunt (except in S . pulverulentum , in which are sharp); hypostomal paralateral teeth present (except in S. brachycladum ). Body surface glabrous. Abdomen gradually expanding posteroventrally to last segments, then abruptly contracting to posterior proleg ( Fig. 111 ). Anal sclerite no encircling base of posterior proleg. Posterior portion of abdomen without sclerotized accessory plates. Without 1+1 ventral papillae. Rectal papillae with three multi-branched lobes. Remarks Simulium ( Hemicnetha ) was recovered as monophyletic based on four synapomorphies ( Fig. 1 , Table 2 , node 21), one of them unique: male gonostylus with medial region larger than basal region (20 (0), Figs 60–62 ). We therefore propose the revalidation of the subgenus, which comprises 19 species. Coscarón & Coscarón-Árias (2007) divided S . ( Hemicnetha ) into four species-groups: Paynei, Brachycladum, Mexicanum, and Oviedoi. We included the following representatives in our analysis: Paynei (five species), Brachycladum (four species), Mexicanum (two species), and Oviedoi (two species). Simulium ( Hemicnetha ) sensu Coscarón & Coscarón-Árias (2007) was not recovered, because Oviedoi does not group with the remaining species-groups. Moreover, except for Oviedoi, none of the previously defined species-groups in this subgenus were recovered. We therefore revalidate S. ( Hemicnetha ), but without Oviedoi and without a subdivision into species-groups pending a more comprehensive investigation on the subgenus. Distribution Simulium ( Hemicnetha ) is mostly distributed in Mesoamerica and South America, being recorded from the following countries: Argentina , Belize , Bolivia , Brazil , Canada , Colombia , Costa Rica , Dominica , Ecuador , El Salvador , French Guiana, Guadeloupe, Guatemala , Guyana , Honduras , Mexico , Nicaragua , Panama , Paraguay , Peru , Saint Vincent , Suriname , Trinidad and Tobago and Venezuela . However, S . bricenoi , S . freemani , S . paynei , S . solarii , and S . virgatum are present in the USA , and S . hippovorum occurs in Canada and the USA ( Adler 2020 ).