Alluaudomyia Kieffer

(Figs. 13E, 19L–M, 24E, 29P, 32J, 35D, 43S–T, 44A–B, 48E, 58C, 73L)

DIAGNOSIS: Only pupa of Ceratopogonidae with the respiratory organ with two closely appressed or approximated rows of pores and with each pore oblong in shape (Figs. 43S–T, 44A–B). Also only pupae with the abdominal segment 4 sensilla D-4-IV and D-7-IV placed on either side of D-8-IV (Fig. 58C) (in some species D-7- IV is placed more laterally, closer to D-9-IV). Also only pupa with the prothoracic extension abutting the antenna but not extending to the palpus (Fig. 24E) and the halter extending well posterior of the anterior margin of tergite 2 (Fig. 32J).

DESCRIPTION: Total length = 1.81–2.75 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, in some against wing (as in Figs. 15D, 32J). Ecdysial tear medial to base of antenna or to posterior to base (as in Figs. 15B, D, 79D, E); along prothoracic extension. Head: Dorsal apotome (Figs. 19L–M), with ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (Fig. 13E) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 24E) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 35D) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Figs. 19L–M)—1 short to moderate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 24E)—absent; oculars (Fig. 24E)—2 setae, 2 campaniform sensilla. Thorax: Prothoracic extension (Fig. 24E) short, present only dorsolaterally abutting antenna, not extending to palpus; mesonotum with very short tubercles, not extending posteromedially or with slight protuberance, not dividing metathorax medially (Fig. 48E); respiratory organ (Figs. 43S–T, 44A–B) length/width = 3.56–7.20, elongate, slender to moderately thick, somewhat flattened dorsoventrally, with pores closely abutting at apex of respiratory organ, arranged in two closely appressed rows, with or without additional, more basal pores, outer surface with (nearly all species) or without spicules, spines or pointed plates, without pedicel, base without posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, distally with plate; wing (Fig. 35D) with apical tubercle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 32J) broadly abutting; halter apex extending posteriorly to 1/3 length of tergite 2; legs (Fig. 35D) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 32J); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg slightly dorsal to, partially abutting apex of midleg laterally; sensilla: anteromedials—2 setae; anterolaterals—3 setae; dorsal setae (Fig. 29P)—D-1-T, D-2-T, D-4-T, D-5-T setae, D-3-T campaniform sensillum, D-3-T posteromedial to D-4-T; supraalar 2—campaniform sensillum; metathoracics (Fig. 48E)—1 seta, 2 campaniform sensilla; M-3-T near anterior margin of metathorax. Abdomen: without pigmentation pattern or anteromedial margin of tergites 1–4 (fading posteriorly) or tergites 1–5 (tergite 1 with 4 medial), just medial on tergite 6, fading posteriorly, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to pointed, short tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 73L) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 48E) with 7–8 setae, 2 campaniform sensilla, including 3 lateral sensilla, D-2-I, D-3-I closely approximated, D-7-I situated posteriorly near D-8-I; segment 4 (Fig. 58C)—D-2-IV, D-3-IV moderately elongate setae on separate tubercles; D-5-IV, D-8-IV, D-9-IV short setae, on short to moderately elongate, separate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D-5-IV, D-4-IV, D-8-IV, D-7- IV, D-9-IV; L-1-IV short seta on pointed tubercle, well anterior of posterior lateral setae; L-2-IV, L-3-IV, L-4-IV short setae on pointed tubercles, V-5-IV, V-6-IV, V-7-IV short to moderately elongate setae on pointed tubercles; segment 8 without D-3-VIII, without L-1-VIII; segment 9 (Fig. 73L)—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Alluaudomyia is known from 203 species from every Region worldwide ( Borkent 2014). Immatures have been sampled from ponds, bogs, fens, swamps, treeholes and margins of streams, creeks, rivers and lakes. Some species have been reared from wet leaves and mud associated with the aforementioned habitats.

TAXONOMIC DISCUSSION: There are 23 species of Alluaudomyia known as pupae (Tables 2–3). Considering the good visibility of the larvae, which swim on the surface film, there are good prospects of rearing many additional species of this genus.

The dorsal apotomes of A. bella, A. caribbeana, A. distispinulosa, A. footei, A. megaparamera, A. paraspina, A. parv a, A. reyei and A. unguistyla have been illustrated without campaniform sensillum DA-2-H (Elson-Harris & Kettle 1985b, Grogan & Bystrak 1976, Spinelli 1997, Spinelli & Wirth 1984, Wirth & Grogan 1981), but examination of original material of A. bella, A. distispinulosa, A. footei, A. megaparamera, A. paraspina and A. parv a indicates that it is actually present in at least these species. The campaniform sensillum has likely been overlooked in the other species considering that all species of Alluaudomyia I examined had the sensillum present among the similarly sized spicules and that the sensillum is present in most other Ceratopogonidae .

De Meillon (1939) illustrated abdominal segment 5 of A. maculithorax but misidentified the dorsal and ventral surfaces.

The distinctive placement of fourth abdominal segment sensilla D-7-IV lateral to D-8-IV has not been reported or illustrated by those previous authors describing the sensilla of the abdominal segments, likely because this sensillum coeloconica is small, difficult to discern and was overlooked (de Meillon 1939, Debenham 1971, Elson-Harris & Kettle 1985b); this is certainly the case for the three species described by Debenham (1971) and which were reexamined here.

MATERIAL EXAMINED: A. appendiculata: 2 pupal exuviae (of paratype), Hornsby, New South Wales, Australia, 9-X-1956 (ANIC); 1 pupal exuviae, as previous locality, 18-XI-1969 (ANIC); 2 pupal exuviae, Nattai River, Mittagong, New South Wales, Australia, 25-X-1968 (ANIC). A. bella: 1 pupal exuviae, 6.5 km NW of Enderby, British Columbia, Canada, 27-VI-1990 (CNCI); 1 pupal exuviae, 20 km E of Anola, Manitoba, Canada, 16-VI-1990 (CNCI); 1 pupal exuviae (in glycerin), Charleston Provincial Park, Ontario, Canada, 18-V-1986 (CNCI); 1 pupal exuviae, Algonquin Park, Ontario, Canada, 7-VI-1960 (USNM); 6 pupal exuviae, Bat Lake, Algonquin Park, Ontario, Canada, 7-VI-1960 (USNM); 1 pupal exuviae, Notre Dame de Laus, Lac Serpent, Quebec, Canada, 22-VI-1967 (USNM); 1 pupal exuviae, as previous locality, 11-VII-1967 (USNM); 1 pupal exuviae, Patuxent Wildlife Rescue Center, Prince George’s County, Maryland, USA, 19-IV-1976 (WLGC); 2 pupal exuviae, as previous locality, 8-V-1958 (USNM); 1 pupal exuviae, as previous locality, 28-VI-1976 (WLGC); 1 pupal exuviae, as previous locality, 17-V-1976 (WLGC); 3 pupal exuviae, Lake Jimmy, Tahawus, New York, USA, 24-VI-1959 (NYSM); 1 pupal exuviae, Hamilton-Essex, Newcomb, New York, USA, 23-VII-1959 (NYSM); 1 pupal exuviae, Huntington Swamp, Newcomb, New York, USA, 4-VIII-1959 (NYSM); 1 pupal exuviae, Salmon River, Blue Mountain Lake, New York, USA, 23-VI-1959 (NYSM); 1 pupal exuviae, Allegany State Park, New York, USA, 28-V-1963 (USNM); 1 pupal exuviae, Upper Pansapoulas Pond, Montgomery County, Virginia, USA, 3-IX-1976 (VPIC); 2 pupal exuviae, Upper Pandapas Pond, Montgomery County, Virginia, USA, 2-VIII-1976 (VPIC). A. bicornis: 1 pupal exuviae, Manton River, Northern Territory, Australia, 6-VI-1958 (ANIC). A. biestroi: 1 pupal exuviae (of paratype), Artigas A “Lenguazo” y Ruta 3, Uruguay, 8-XI-1984 (MLPA). A. claudia: 1 pupal exuviae (of paratype) Salisbury, Southern Rhodesia, South Africa, 24-II-1949 (SAIM). A. distispinulosa: 1 pupal exuviae, Bulo Bulo, Cochabamba, Bolivia, 12-I-1995 (MLPA). A. fragmentum: 1 pupal exuviae (of paratype), Manton River, Northern Territory, Australia, 6-VI-1958 (ANIC); 1 pupal exuviae (of paratype), as previous locality, 1-VI-1958 (ANIC). A. latipennis: 1 pupal exuviae, Hornsby, New South Wales, Australia, 9-X-1956 (ANIC); 1 pupal exuviae, Nattai River, Mittagong, New South Wales, Australia, 4-XI-1964 (ANIC); 1 pupal exuviae, as previous locality, 15-XI-1968 (ANIC). A. megaparamera: 1 pupal exuviae, Blue Mountain Lake, New York, USA, 29-VI-1959 (NYSM); 1 pupal exuviae, Salmon River, Blue Mountain Lake, New York, USA, 8-VI- 1959 (NYSM); 1 pupal exuviae, Hamilton-Essex, Newcomb, New York, USA, 1-VI-1959 (NYSM); 1 pupal exuviae, as previous locality, 8-VI-? (NYSM). A. natalensis: 1 pupal exuviae, Empangeni, Zululand, South Africa, 16-VII-1938 (SAIM); 1 pupal exuviae, 15 mi. SE of Potgietersrus, Transvaal, South Africa, 3-I-1974 (NMSA). A. needhami: 1 pupal exuviae, Salmon Arm, British Columbia, Canada, 11-VII-1990 (CNCI); 3 pupal exuviae, 30 km N of Nakusp, British Columbia, Canada, 5-VII-1990 (CNCI); 2 pupal exuviae, Patuxent Wildlife Refuge, Prince George’s County, Maryland, USA, 1-VIII-1979 (USNM); 1 pupal exuviae, Upper Pandapas Pond, W.A., Montgomery County, Virginia, USA (VPIC); 1 pupal exuviae, 1.7 mi E of Lunter’s gate, Rt. 785, Montgomery County, Virginia, USA, 22-IV-1976 (VPIC); 1 pupal exuviae, no locality, 10-VI-1953 (USNM); 2 pupal exuviae, no locality, 20-II-1957 (USNM). A. needhami or A. megaparamera: 2 pupal exuviae, Montvale Wayside, Bedford County, Virginia, USA, 16-VI-1977 (VPIC)); 1 pupal exuviae, Beaver Lake Reservoir, Pocahontas State Park, Chesterfield County, Virginia, USA, 30-IV-1977 (VPIC). A. paraspina: 2 pupal exuviae, Lakeland Pond, College Park, Prince George’s County, Maryland, USA, 26-V-1975 (WLGC); 2 pupal exuviae, as previous locality, 26-V- 1975 (WLGC); 5 pupal exuviae, as previous locality, 30-V-1975 (WLGC); 1 pupal exuviae, Mountain Lake Biological Station, Giles County, Virginia, USA (VPIC). A. parva: 1 pupal exuviae, Blue Mountain Lake, New York, USA, 30-VI-1958 (NYSM); 1 pupal exuviae, Salmon River, Blue Mountain Lake, New York, USA, 14-V- 1959 (NYSM); 1 pupal exuviae, Allegany State Park, New York, USA, 28-V-1963 (USNM); 7 pupal exuviae, Alexandria, Virginia, USA, 6-VI-1951 (USNM). A. quadripunctata: 2 pupal exuviae, Tobolki lake, Alol’, Pskov Province, Russia, 30-VI-1969 (ZIN). A. schnacki: 2 pupal exuviae (of paratype), Los Talas (Berisso), Buenos Aires, Argentina, 29-XI-1979 (MLPA). A. splendida: 1 pupal exuviae, Ussuri Nature Reserve, Primorskii Territory, Russia, 12-VI-1973 (ZIN). A. variegata: 1 pupal exuviae, Lake Muson, Leon County, Florida, USA, 22-VIII-1987 (JHEC). A. sp.: 1 pupal exuviae, Bolean Lake, 6 km NE of Falkland, British Columbia, Canada, 1-VII-2009 (CNCI); 1 pupal exuviae, 30 km N of Nakusp, British Columbia, Canada, 5-VII-1990 (CNCI); 1 pupal exuviae, 5 km S of Cottonwood Lake, Nelson, British Columbia, Canada, 6-VII-2008 (CNCI); 1 pupal exuviae, Meachum Lake, New York, USA, 21-VI-1986 (CNCI); 1 pupal exuviae, Ackerman Swamp, Newcomb, New York, USA, 7- VII-1958 (NYSM); 1 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 19-II-1967 (BPBM); 1 pupal exuviae, as previous locality, 2-XII-1967 (BPBM); 2 pupal exuviae, as previous locality, 22-II-1967 (BPBM); 1 pupal exuviae, Houay La stream, 20 kilometers NE of Moung Sayaboury, Sayaboury Province, Laos, 25-XI-1967 (BPBM); 1 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 9-XII-1967 (BPBM); 1 pupal exuviae, as previous locality, 7-I-1968 (BPBM); 1 pupal exuviae, Nam Houng River, Moung Sayaboury, Sayaboury Province, Laos, 17-XII-1967 (BPBM); 1 pupal exuviae, Gnieraoora Pool, Robe River, Pilbara, Western Australia, Australia (ANIC).