Revision of the New World Ceratoculicoides Wirth & Ratanaworabhan (Diptera, Ceratopogonidae, Ceratopogonini) Author Fasbender, Andrew Rhithron Associates Inc., 33 Fort Missoula Road, Missoula, MT 59847, USA. afasbender@rhithron.com text European Journal of Taxonomy 2023 2023-06-26 875 159 202 journal article 54709 10.5852/ejt.2023.875.2147 3351d966-cbeb-42a8-ac49-627b5fc261c7 2118-9773 8083773 32FA008C-B35D-483C-9DBE-1DCCD0868FAC Ceratoculicoides Wirth & Ratanaworabhan, 1971 Figs 1–12 Ceratoculicoides Wirth & Ratanaworabhan, 1971: 170 . Ceratoculicoides – Downes & Wirth 1981: 408 (included in genus key). — Knoz 1987: 391 (diagnosis), 388 (species key). — Wirth & Grogan 1988: 6 (included in genus key), 39 (diagnosis). — Borkent 1992: 434 (included in genus key). — Borkent & Wirth 1997: 95 (in catalog). — Huerta & Borkent 2005: 114 (catalog). — Borkent et al. 2009: 413 (included in genus key). — Borkent & Dominiak 2020: 157 (in catalog). Type species Helealongipennis Wirth, 1952: 170 (by original designation). Diagnosis (adult) Anepisternum with 1–3 setae along posterior margin, katepisternum with 2–4 setae. Tarsomere 1 of hind leg with palisade setae. Wing cells r 1 and r 2 reduced, M 2 base obsolete. Description Male HEAD . Cranium brown. Antenna brown with 13 flagellomeres, 7–11 always fused, fusion sometimes including flagellomeres 5 and 6 as well; 2–3 sensilla coeloconica on 1, flagellomeres 1–10 with plume, 11–12 with ring of laterally directed trichoid sensilla at base, 13 with apical trichoid sensillum, AR 0.65–1.02, FR 1.49–1.92. Eyes separated medially by diameter of 3–5 ommatidia, ommatrichia present. Palpus brown, with 5 segments, 3 rd with sensory pit. THORAX . Scutum, scutellum and pleural sclerites dark brown without distinct coloration patterns; scutum without anteromedial tubercle, humeral pits poorly developed, often with punctations present among dorsocentrals. Anepisternum with 1–3 setae along posterior margin, katepisternum with 2–4 setae. WING . Cells r 1 and r 2 reduced, M 2 base obsolete. Membrane with microtrichia, macrotrichia present only on C and R; membrane unpatterned, without macrotrichia. LEGS . Femora, tibiae without spines; spur formula 1-0-1, hindleg spur pectinate. Hind 1 st tarsomere with single row of palisade setae; all tarsal claws equal-sized, without accessory teeth, straight, apex minutely bifid. PREGENITAL ABDOMEN . Brown, without distinct coloration patterns, margins roughly parallel to genitalia. GENITALIA . Epandrium with apicolateral processes present, cerci and proctiger near posterior margin of epandrium directed ventrally. Gonocoxite cylindrical; gonocoxal apodeme quadrate anteriorly, triangular posteriorly; gonostylus simple, subequal in length to gonocoxite, weakly curved at tip. Sternite 9 (hypandrium) slightly tapering anteriorly, length/width ratio 0.33, with medial emargination. Parameres separate or weakly fused at base; articulating with anterior portion of gonocoxal apodeme, apical arms of paramere stylate, directed posteriorly at distinct angle from base, midpoint arched dorsally in lateral view, apex of paramere ventrally directed, extending to apex of aedeagus or beyond. Aedeagus heavily sclerotized, with or without apical hyaline incision medially, with distinct basal arms articulated on anterior portion of gonocoxal apodeme, posterolateral point present, of variable form. Fig. 1. Ceratoculicoides pacificus sp. nov. , male habitus (CNCI). Scale bar = 100 μm. Female HEAD . All flagellomeres separate, without plume, AR 1.08–1.33, FR 1.37–1.83. Mandible with 8 teeth. THORAX . Dorsocentral portion of scutum with numerous punctations. WING . Membrane with scattered macrotrichia along apical margin. LEGS . Fore-, midleg tarsal claws longer than those of hindleg, equal or slightly unequally sized, gently curving along length, apex with simple point. PREGENITAL ABDOMEN . Margins convex, widest at approximately ⅓ length in specimens fully laden with eggs, posteriorly tapering to rounded apex. Fig. 2. Ceratoculicoides pacificus sp. nov. (CNCI), female habitus. Scale bar = 100 μm. GENITALIA . Tergite 9 simple, bandlike. Sternite 8 simple, undivided. Sternite 9 narrow at base, apex crescent-shaped. 1–2 major spermathecae and a very small third spermatheca usually present. Spermathecal necks weakly curved, spermathecal neck ratio 0.14–0.32. Immature stages Currently unknown. Fig. 3. Ceratoculicoides pacificus sp. nov. (CNCI), male thorax. Scale bar = 100 μm. Key to adult male New World Ceratoculicoides The male lifestage of Ceratoculicoides longipennis from California is currently unknown. A poorly preserved male specimen from California that may be conspecific with C.longipennis is included as C . sp. M1 in the key. The distribution of each species is noted in brackets. 1. Apex of paramere broadly rounded, minimally tapering to apex, minute apical point laterally displaced ( Fig. 9b ) [eastern North America ( Canada and USA )] ........ C.virginianus ( Wirth, 1951 ) – Apex of paramere acute, paramere evenly tapers to apical point that is not laterally displaced ( Fig. 9e, h ) ........................................................................................................................................ 2 2. Lateral margin of aedeagus evenly convex, without constriction ( Fig. 8c, f, i ) .............................. 3 – Lateral margin of aedeagus straight ( Fig. 10c ), apically constricted ( Figs 9c , 10e, h ) or concave ( Fig. 9f, i ) ......................................................................................................................................... 4 3. Medial apex of aedeagus with hyaline incision, length of dorsolateral spines>0.5 width of apex of aedeagus ( Fig. 8i ) [Pacific Northwest ( Canada and USA )] ............................... C. pacificus sp. nov. – Medial apex of aedeagus uniformly sclerotized, without hyaline medial incision, length of dorsolateral spines>0.25 width of apex of aedeagus ( Fig. 8c, f ) [eastern North America ( Canada and USA ) and Colombia ].. .......................................................................................................... C.confusus sp. nov. 4. Aedeagus with posteromedial hyaline incision extending nearly to basal arch, anterior apex broad ( Fig. 9f ) [Southwestern United States , Rocky Mountains ( USA ), Vancouver Island ( Canada )] ........ ............................................................................................................................. C.borkenti sp. nov. – Aedeagus hyaline incision tapering or obscure anteriorly ( Figs 9i , 10h, k ) .................................... 5 5. Aedeagus lateral margins narrowed apically, posterior margin of aedeagus with distinct notch ( Fig. 10e ) [ Jalisco ( Mexico )] ...................................................... C.aliciae Huerta & Borkent, 2005 – Aedeagus lateral margins straight, posterior margin an even arc, without notch ( Figs 9i , 10c, k ) [Pacific Northwest and California ( Canada and USA )] ................................................................... 6 6. Posterolateral point of aedeagus an acutely pointed spine, much longer than basal width, apex directed laterally ( Fig. 9i ) [ Oregon and California ( USA )] ................................. C.grogani sp. nov. – Posterolateral point of aedeagus rounded, length subequal to basal width, directed slightly anteriorly ( Fig. 10c, k ) ...................................................................................................................................... 7 7. Accessory points present between apical and posterolateral points of aedeagus ( Fig. 10c ), wing length> 1.1 mm [ California ( USA )] ................................................................................... C. sp. M1 – Accessory points absent between apical and posterolateral points of aedeagus ( Fig. 10k ), wing length < 1.1 mm [ British Columbia ( Canada ) and California ( USA )] ........... C.propinquus sp. nov. Key to the adult female New World Ceratoculicoides Ceratoculicoideslongipennis from California , known only as a female, is not included in the following key. Three female morphotypes are included but not formally named (see taxonomic discussions below). Individual species distributions are noted in brackets. 1. Medial margin of female genital sclerotization on sternite 9 almost straight ( Fig. 11e ) [ Oregon and California ( USA )] ................................................................................................ C.grogani sp. nov. – Median margin of female genital sclerotization on sternite 9 strongly concave, arched ( Figs 11a–d, f , 12a–f ) ............................................................................................................................................... 2 2. Length of largest spermatheca>80 μm ............................................................................................ 3 – Length of largest spermatheca <80 μm ............................................................................................ 4 3. Wing length < 1.5 mm [eastern North America ( Canada and USA )] ................. C.confusus sp. nov. – Wing length> 1.5 mm [Pacific Northwest ( Canada and USA )] .......................... C. pacifica sp. nov. 4. Length of largest spermatheca>70 μm ............................................................................................ 5 – Length of largest spermatheca <70 μm ............................................................................................ 6 5. Posterior branch of 9 th sternite tapering, apex acutely pointed ( Fig. 11a ) [ Jalisco ( Mexico )] ............ ..................................................................................................... C.aliciae Huerta & Borkent, 2005 – Posterior branch of 9 th sternite not tapering, apex obtusely rounded ( Fig. 12e ) [ Colombia ] .............. ............................................................................................................................................... C. sp. F2 6. Flagellum ratio (FR) <1.5 ( Fig. 7c ) [Southwestern United States , Rocky Mountains ( USA ), Vancouver Island ( Canada )] ............................................................................... C.borkenti sp. nov. – Flagellum ratio (FR)>1.5 ( Figs 6a–d , 7d–e ) ................................................................................... 7 7. Anterior branch of 9 th sternite tapering from base, acutely spiniform ( Fig. 12f ) [ Costa Rica ] .......... .............................................................................................................................................. C. sp. F3 – Anterior branch of 9 th sternite tapering only in apical half ( Fig. 12c–d ) ............................................ .......................... C.virginianus [eastern North America ( Canada and USA ), C. sp. F1 ( Colombia )]