Palpomyia Meigen

Palpomyia Meigen

(Figs. 2D 12G, 17E, 22H–J, 28G, 31H, 31L–M, 33K, 41D, 46N–R, 54A, 70C, 71A, 78A–H)

DIAGNOSIS: Only pupa of Ceratopogonidae with the metathorax with only one campaniform sensillum (M-3-T) situated at least ⅓ the length of the metathorax from its anterior margin (Fig. 54A), apex of the halter extending posteriorly to about 1/6 length of tergite 2 (as in Fig. 33L), abdominal segment 4 with V-5-IV, V-6-IV and V-7-IV closely approximated (Fig. 70C) or, if V-7-IV is closer to L-4-IV then L-3-IV is closer to L-2-IV than to an elongate L-1-IV (as in Fig. 69C) (not as in Fig. 70B), abdominal segment 8 has V-5-VIII and V-6-VIII on separate tubercles or if on partially to completely fused tubercles, then V-5-VIII is well-developed (not minute), and segment 8 is without L-1-VIII (not diagnosable as different from Bezzia and Phaenobezzia); however, most species of Bezzia have two or more campaniform sensilla on the dorsal apotome (Figs. 22C–D), a nearly unique condition found otherwise only in P. flavipes and P. jonesi (the latter distinctively with two setae (Fig. 22J).

DESCRIPTION: Habitus as in Fig. 12G. Total length = 2.25–7.03 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (Figs. 17E, 33K). Ecdysial tear around base of antenna, along lateral margin of face to palpus (Figs. 17E, 79H). Head: Dorsal apotome (Figs. 22H–J), with partial ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 28G) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire or separated medially by labrum, hypopharynx; apex of antenna (Fig. 41D) well anterior to posterior to, posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Figs. 22H–J)—1 elongate seta, 1 campaniform sensillum or 2 elongate setae, 2 campaniform sensilla; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeallabrals (Fig. 28G)—2 slender setae; oculars (Fig. 28G)—1–2 setae, 1 campaniform sensillum or 1 seta, 2 campaniform sensilla. Thorax: Prothoracic extension (Fig. 28G) wide, well-developed but in some narrow dorsolaterally, extending from palpus to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 54A); respiratory organ (Figs. 46N–R) length/width = 2.46–4.73, moderately elongate to elongate, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single straight to curved row, outer surface smooth or with some wrinkles, with short, wide pedicel, base with short posteromedial apodeme, membranous base of respiratory organ short, annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length or more; wing (Fig. 41D) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33K) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs (Fig. 41D) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33K); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials—2 elongate setae, 1 campaniform sensillum (Figs. 31L–M); anterolaterals—1 moderately long seta; dorsal setae (Fig. 31H)—D-1-T, D-2-T, D-5-T, D-4-T setae, D-3-T campaniform sensillum, D-3-T lateral to anterolateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics (Fig. 54A)—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: pigmentation light to dark brown, tergites 1–7 with medial area with stripe, 2 spots, sternites 3–7 with medial stripe, anterolateral spot, segment 2 as wide or slightly wider than segment 3, segments with undivided, peg-like or thin to thick setae, with rounded to pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Figs. 78A–H) not strongly modified, terminal processes closely approximated to separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 54A) with 8 setae, 2 campaniform sensilla, including 4 lateral sensilla, D-2-I, D-3-I closely approximated, D-7-I situated anterolaterally near L-1-I; segment 4 (Figs. 70C, 71A)—D-2-IV, D-3-IV short to moderately elongate setae on short to pointed tubercles; D-5-IV peg-like or slender seta, D-8-IV, D-9-IV short to moderately elongate setae, D-7-IV present or absent; D-5-IV without or on short tubercle, D-8-IV, D-9-IV on basally fused or separate but closely approximated tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D-5-IV, D-4-IV, D-8-IV, D-9- IV; D-7-IV, if present, near D-3-IV; L-1-IV short to elongate seta on rounded tubercle, just anterior of base of tubercle with L-2-IV, L-3-IV; L-2-IV, L-3-IV, L-4-IV elongate setae on pointed tubercles, L-2-IV, L-3-IV on single tubercle in some, V-5-IV, V-6-IV, V-7-IV short to moderately elongate setae on short tubercles, all closely approximated or with V-7-IV closer to L-4-IV; segment 8 without D-3-VIII, without L-1-VIII; segment 9 (Figs. 78 A-H)—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Palpomyia is known from 270 species from every Region worldwide (Borkent 2014, additional species below). Immatures have been collected from streams, hot springs, river margins, marshes, bogs, fens, ponds, lakes and reservoirs.

TAXONOMIC DISCUSSION: There are 41 species of Palpomyia known as pupae (Tables 2–3). Thienemann (1928), Mayer (1934a) and Lenz (1934) provided similar keys to a few European species groups known at that time. Grogan & Wirth (1979), in their masterful revision of Nearctic Palpomyia, provide a key to the pupae of 16 species known from the Nearctic. However, it is unfortunate that Palpomyia pupae cannot be diagnosed at the generic level so that we cannot yet identify a Palpomyia pupa as such. The key therefore requires foreknowledge that the specimen is a member of the genus (e.g. reared to adult).

Liu & Yu (1991) described the monotypic genus Nemoromyia Liu & Yu based on their newly described N. nemorosa Liu & Yu. They stated that the genus was a member of the Heteromyiini. Borkent (1998) indicated that the species was actually a species of Palpomyia based on various described features but that the type should be reexamined before nomenclatural changes are made. Yu et al. (2005) continued to place the genus and species in the Heteromyiini, although the evidence for this was unclear (it is not noted in the English abstract, but I have not translated the longer Chinese text). I have reexamined the female holotype and associated pupal exuviae and it is clearly a member of the Palpomyia distincta species group and, within this group, is very similar to P. rufa Loew (Grogan & Wirth 1979). The female has the characteristic synapomorphy of posteromedially directed lobes on sternite 8 of the P. distincta group. Furthermore, tergite 8 is very short (much shorter than tergite 7), a feature which is a synapomorphy of the Palpomyiini (the feature is unique in the Ceratopogonidae), further supporting its position as a species of Palpomyia. The pupa of N. nemorosa is very similar to a number of other species of Palpomyia (as indicated in the key). Liu & Yu (1991) noted that the adult female lacked the abdominal tergal apodemes which is a synapomorphy of the Palpomyiini. However, as shown by Borkent & Craig (1994), newly emerged female adults of this group do not show the apodemes which sclerotize and darken after emergence. It is not surprising, therefore, that this reared specimen was reported as lacking the apodemes. I therefore recognize Nemoromyia as a new synonym of Palpomyia and the species as a new combination as follows:

Palpomyia nemorosa (Liu & Yu), 1991: 26 (Nemoromyia). Raohe, Heilongjiang Province, China. new combination.

The pupa of P. lineata has been described more than once by the same authors (Table 2) but with different names (now synonyms), suggesting either misidentifications or the possibility of more than one species actually being present in Europe.

Ronderos et al. (2004) gave a detailed description of the pupa of P. guarani. Their figure of the cephalothorax (their fig. 14) shows the apex of the halter extending only barely past the anterior margin of tergite 2. This would be unique within the Palpomyia + Bezzia + Phaenobezzia + Clastrieromyia + Stenoxenini clade, where the halter extends to about 1/6 the length of tergite 2. Paul et al. (2014) recently thoroughly described the pupa of a species from India, the first known from the Oriental Region, but misidentified some of the sensilla (e.g. SA-2-T, L-3-IV, L-4-IV).

MATERIAL EXAMINED: P. aldrichi: 2 pupal exuviae, Agoura, Los Angeles County, California, USA, 7- IV-1954 (USNM); 2 pupal exuviae, Hopland, Mendocino County, California, USA, 19-V-1964 (USNM). P. altispina: 9 pupal exuviae (of paratypes), Taughannock Falls, Tompkins County, New York, USA, 15-VI-1963 (USNM). P. armatipes: 1 pupal exuviae, Victoria, British Columbia, Canada, VII-1965 (USNM); 7 pupal exuviae, Beaver Creek, Rio Grande County, Colorado, USA, 21-VI-1972 (USNM); 2 pupal exuviae, Cross Creek margin, Rio Grande County, Colorado, USA, 24-VI-1972 (USNM); 3 pupal exuviae, South Fork, Rio Grande County, Colorado, USA, 21-VI-1972 (USNM). P. basilis: 1 pupal exuviae, Allegany State Park, New York, USA, 28-V- 1963 (USNM); 1 pupal exuviae, Blue Ridge, New York, USA, 25-V-1959 (USNM); 1 pupal exuviae, as previous locality, 26-V-1959 (USNM); 2 pupal exuviae, Lake Ravenel, Highlands, Macon County, North Carolina, USA, 10-VI-1986 (USNM); 1 pupal exuviae, as previous locality, 11-VI-1986 (USNM). P. belkini: 1 pupal exuviae (of paratype): Los Angeles River, Reseda, Los Angeles County, California, USA, 6-IV-1955 (USNM); 1 pupal exuviae (of paratype), Agoura, Los Angeles County, California, USA, 7-IV-1954 (USNM). P. distincta: 1 pupa, 2 pupal exuviae, springs at Die, Schleswig-Holstein, Malente, Germany, (ZSMC). P. flaviceps: 1 pupal exuviae, Mud Creek, Freeville, Tompkins County, New York, USA, 19-VI-1963 (USNM); 2 pupal exuviae, Lunzer Untersee, lower Austria, Austria, 1940 (ZSMC). P. flavipes: 1 pupal exuviae, Suputinka river, Ussuri Nature Reserve, Primorskii Territory, Russia, 21-VI-1973 (ZIN). P. hastata: 1 pupal exuviae (of paratype), Blue Mountain Lake, Hamilton County, New York, USA, 10-VI-1960 (USNM); 1 pupal exuviae, Mud Creek, Freeville, Tompkins County, New York, USA, 19-VI-1963 (USNM); 1 pupal exuviae, Letchworth State Park, New York, USA, 13-VI- 1963 (USNM); 1 pupal exuviae, Mount Solon, Virginia, USA, 4-VII-1951 (USNM). P. jamnbacki: 1 pupal exuviae (of holotype), Mud Pond outlet, Blue Mountain Lake, New York, USA, 14-V-1958 (USNM); 1 pupal exuviae (of paratype), Blue Ridge, Essex County, New York, USA, 19-V-1959 (USNM). P. jonesi: 1 pupal exuviae, Black Lake, North Burgess Township, Ontario, Canada, 4-VI-1967 (CNCI); 1 pupal exuviae (of paratype), Washburn County, Wisconsin, USA, 18-V-1953 (USNM); 1 pupal exuviae (of paratype), as previous locality, V-1953 (USNM); 2 pupal exuviae (of paratypes), as previous locality, 22-V-1953 (USNM); 4 pupal exuviae, Fishing Creek Pond, Newcomb, New York, USA, 28-V-1958 (3 NYSM, 1 USNM); 1 pupal exuviae, Blue Mountain Lake, New York, USA, 14-V-1959 (NYSM); 4 pupal exuviae (of paratypes), no locality, 20-V-1953 (USNM); 2 pupal exuviae, no locality, 30-V-1953 (USNM); 1 pupal exuviae (of paratype), no locality, 16-V-1953 (USNM); 4 pupal exuviae (of paratypes), no locality, 22-V-1953 (USNM). P. lineata: 1 pupal exuviae, Algonquin Park, Ontario, Canada, 8- VI-1960 (USNM); 1 pupal exuviae, Morgan Arboretum, St. Anne de Bellevue, Quebec, Canada, 1964 (USNM); 1 pupal exuviae, Snow Hill, Maryland, USA, 19-V-1968 (USNM); 1 pupal exuviae, Lakeland Pond, College Park, Prince George’s County, Maryland, USA, 23-V-1975 (USNM); 1 pupal exuviae, as previous locality, 27-IV-1977 (VPIC); 1 pupal exuviae, Greensport, Long Island, New York, USA, 24-V-1963 (USNM); 1 pupal exuviae, Montauk, Long Island, New York, USA, 24-V-1963 (USNM); 1 pupal exuviae, Fishing Creek, Newcomb, New York, USA, 28-V-1958 (NYSM); 1 pupal exuviae, Beaver Lake Reservoir, Pocahontas State Park, Virginia, USA, 9-V-1977 (VPIC); 1 pupa, Eppendorfer Moor near Hamburg, Germany (ZSMC); 1 pupal exuviae, Schöhsee, Plön, Slesvig-Holstein, Germany (ZSMC); 2 pupae, locality uncertain (ZSMC); 1 pupal exuviae, no locality, 16-V-1953 (USNM); 6 pupal exuviae, Shushary, Leningrad Province, Russia, 19-V-1997 (ZIN). P. magali: 1 pupal exuviae (of holotype), Magaliesberg Agricultural School, Transvaal, South Africa, 13-XI-1973 (NMSA). P. melacheira: 1 pupal exuviae, Suputinka river, Ussuri Nature Reserve, Primorskii Territory, Russia, 30-V-1973 (ZIN). P. nemorosa: 1 pupal exuviae (of holotype), Raohe, Heilongjiang Province, China, VI-1983 (IMBC); P. novitibilialis: 1 pupal exuviae (of paratype), Allegany State Park, New York, USA, 28-V–3-VI-1963 (USNM); 1 pupal exuviae (of paratype), as previous locality, 28-V-1963 (USNM); 2 pupal exuviae (of paratypes), Ivory, Chautauqua County, New York, USA, 31-V-1963 (USNM); 1 pupal exuviae (of paratype), Sinclairville, Chautauqua County, New York, USA, 31-V-1963 (USNM). P. occidentalis: 1 pupal exuviae, 17 km N. Sedona, Arizona, USA, 11-V-1987 (CNCI); 18 pupal exuviae, Rio Penasco approx. 2 mi W of Dunken, New Mexico, USA, 22-V-1973 (WLGC); 1 pupal exuviae, Spanish Queen Mine, Mt. Jemez Springs, Sandoval, New Mexico, USA, 19-VIII-1972 (WLGC); 9 pupal exuviae (of paratypes), Beaver Creek, Lawrence County, South Dakota, USA, 15-VI-1969 (USNM); 1 pupal exuviae (of paratype), Spearfish Creek, Lawrence County, South Dakota, USA, 14-VI-1969 (USNM); 1 pupal exuviae (of paratype), Little White River, Mellette County, South Dakota, USA, 4-VI-1969 (USNM). P. plebeja: 4 pupal exuviae, Holmes Run, Falls Church, Fairfax County, Virginia, USA, 22-VII-1951 (USNM). P. reversa: 1 pupal exuviae, Issyk-Kul lake, Kyrgyzstan, 16-VI-1971 (ZIN). P. rubiginosa: 9 pupal exuviae (of paratypes), Algonquin Park, Ontario, Canada, 7-VI-1960 (USNM). P. rufa: 1 pupal exuviae (of paratype), Whetstone Gulf, Lewis County, New York, USA, 20-VI-1963 (USNM); 1 pupal exuviae (of paratype), 2 pupal exuviae, Mud Creek, Freeville, Tompkins County, New York, USA, 19-VI-1963 (USNM); 1 pupal exuviae (of paratype), Blue Mountain Lake, Hamilton County, New York, USA, 10-VI-1960 (USNM); 1 pupal exuviae, Patuxent Rescue Center, Prince George’s County, Maryland, USA, 6-V-1976 (USNM); 3 pupal exuviae, as previous locality, 13-V-1976 (USNM). P. rufipes: 1 pupal exuviae, Priluki, Belarus, V-1967 (ZIN). P. scalpellifera: 3 pupal exuviae, Letchworth State Park, New York, USA, 13-VI-1963 (USNM). P. serripes: 1 pupal exuviae, Kellersee, Malente, Slesvig-Holstein, Germany (ZSMC). P. spinipes: 1 pupal exuviae, Zaklin'ye, Luga District, Leningrad Province, Russia, 30-VI-1972 (ZIN). P. stonei: 2 pupal exuviae, Allegany State Park, New York, USA, 28-V-1963 (USNM). P. subaspera: 2 pupal exuviae, Shafter Water Reservoir, Kern County, California, USA, 5-VI-1946 (USNM); 2 pupal exuviae, Shafter, Kern County, California, USA, 5-VI-1946 (USNM); 1 pupal exuviae, Fellsmere, Florida, USA, 7-V-1973 (USNM). P. tibialis: 2 pupal exuviae, B. Brod, Luga District, Leningrad Province, Russia, 29-VI-1969 (ZIN). P. tuvae: 1 pupal exuviae, Cholpon-Atinka river, Tien-Shan, Kyrgyzstan, 7-VI-1971 (ZIN). P. nr. lineata: 1 pupal exuviae, Bolean Lake, 6 km NE of Falkland, British Columbia, Canada, 1-VII-2009 (CNCI); 1 pupal exuviae, 11 km S of Patagonia, Arizona, USA, 1-V-1987 (CNCI). P. nr. novitibialis: 2 pupal exuviae, Beaver Lake, Pocahontas State Forest, Chesterfield County, Virginia, USA, 30-IV-1977 (VPIC); 1 pupal exuviae, Hoot Owl Barn, VPI & SU campus, Montgomery County, Virginia, USA, 15-IV-1976 (VPIC). P. nr. plebeia: 1 pupal exuviae, Huntington Road, Newcomb, New York, USA, 14-V-1959 (WLGC). P. nr. rubiginosa: 1 pupal exuviae, Lake Norman, North Carolina, USA, 28-VII-1978 (VPIC). P. sp.: 1 pupal exuviae, 6 km E of Salmon Arm, British Columbia, Canada, 6- VI-1990 (CNCI); 1 pupal exuviae, 9 km S of Salmon Arm, British Columbia, Canada, 19-VII-1988 (CNCI); 4 pupal exuviae, 8 km E of Sicamous, British Columbia, Canada, 1-VI-1992 (CNCI); 1 pupal exuviae, 24 km E of Enderby, British Columbia, Canada, 8-VI-1991 (CNCI); 1 pupal exuviae, Gibson Lake, Kokanee Glacier Provincial Park, British Columbia, Canada, 9-VII-2008 (CNCI); 2 pupal exuviae, 20 km E of Anola, Manitoba, Canada, 16-VI-1990 (CNCI); 1 pupal exuviae, Trail Bay, Manitoba, Canada (CNCI); 1 pupal exuviae (in glycerin), Black Lake, Gatineau, Quebec, Canada, 1-VI-1972, (CNCI); 5 pupal exuviae, 17 km N of Sedona, Arizona, USA, 11-V-1987 (CNCI); 1 pupal exuviae, Quoque, Long Island, New York, USA, 9-V-1957 (WLGC); 1 pupal exuviae, Hoot Owl Barn, VPI & SU campus, Montgomery County, Virginia, USA, 15-IV-1976 (VPIC); P. sp.: 1 pupal exuviae, 5 km NE of Tarcoles, Costa Rica, 26-VII-1993 (CNCI); 2 pupal exuviae, Iquitos, Loreto, Peru, 28-III- 2003 (CNCI); 1 pupal exuviae, as previous locality, 26-IV-2003 (CNCI); 3 pupal exuviae, as previous locality, 28- IV-2003 (CNCI); 1 pupal exuviae, as previous locality, 15-I-2003 (CNCI); 3 pupal exuviae (identified by Mayer as Bezzia annulipes), Rotmoos on Mittersee, Lower Austria, Austria, 10-VI-1942 (ZSMC); 1 pupal exuviae, Krzeszna, Poland, 19-V-1993 (IZUG); 1 pupal exuviae, Gdańsk Osowa, Poland, 18-V-1993 (IZUG); 10 pupal exuviae, Yanchep Ponds, Western Australia, Australia (ANIC); 5 pupal exuviae, as previous locality, 25-X-1985 (ANIC); 1 pupal exuviae, Yeerongpilly, Brisbane, Queensland, Australia (ANIC); 5 pupal exuviae, Griffith, New South Wales, Australia, 16-XI-1956 (ANIC); 2 pupal exuviae, Hornsby, New South Wales, Australia, 25-X-1956 (ANIC); 1 pupal exuviae, as previous locality, 18-XI-1969 (ANIC); 1 pupal exuviae, Middle Creek, Narrabeen, New South Wales, Australia, 4-X-1986 (ANIC); 3 pupal exuviae, Deniliquin, New South Wales, Australia, 19-XI- 1956 (ANIC); 1 pupal exuviae, Breakfast Creek, Spencer, New South Wales, Australia, 3-III-1969 (ANIC); 1 pupal exuviae, Oxford Falls, New South Wales, Australia, 18-I-1967 (ANIC); 5 pupal exuviae, Gap Creek, The Crags, Mittagong, New South Wales, Australia, 10-II-1966 (ANIC); 5 pupal exuviae, Nattai River, Mittagong, New South Wales, Australia, 18-X-1968 (ANIC); 7 pupal exuviae, as previous locality, 6-I-1969 (ANIC); 1 pupal exuviae, as previous locality, 25-X-1968 (ANIC); 2 pupal exuviae, as previous locality, 4-XI-1964 (ANIC); 1 pupal exuviae, as previous locality, 5-II-1969 (ANIC); 3 pupal exuviae, as previous locality, 25-X-1968 (ANIC); 1 pupal exuviae, no location, 30-V-1953 (USNM).

Phaenobezzia Haeselbarth

(Figs. 17F, 22K, 28H, 31I, 33L, 41E, 46S, 54B, 71B, 78I–K)

DIAGNOSIS: Only pupa of Ceratopogonidae with the male with the metathorax with only M-3-T present and situated at least 1/3 length of metathorax and genital lobes very short, extending no more than about 0.8 length of segment 9 (measured medially) (Figs. 78I–J); female (without genital lobes) with the metathorax with only one campaniform sensillum (M-3-T) situated at least ⅓ the length of the metathorax from its anterior margin (Fig. 54B), apex of the halter extending posteriorly to about 1/6 length of tergite 2 (Fig. 33L), abdominal segment 4 with V-5-IV, V-6-IV and V-7-IV closely approximated (Fig. 71B), abdominal segment 8 with V-5-VIII and V-6-VIII on separate tubercles or if on partially to completely fused tubercles, then V-5-VIII well-developed (not minute), without L-1-VIII (not diagnosable as different from Bezzia and Palpomyia); however, most species of Bezzia have two or more campaniform sensilla on the dorsal apotome (Figs. 22C–D), a nearly unique condition found otherwise only in P. flavipes and P. jonesi (Fig. 22J).

DESCRIPTION: Total length = 2.25–4.78 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (Figs. 17F, 33L). Ecdysial tear around base of antenna, along lateral margin of face to palpus (Figs. 17F, 79H). Head: Dorsal apotome (Fig. 22K), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 28H) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 41E) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 22K)—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeallabrals (Fig. 28H)—2 slender setae; oculars (Fig. 28H)—1 seta, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 28H) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 54B); respiratory organ (Fig. 46S) length/width = 3.86–4.93, elongate, moderately slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single curved row, outer surface with some wrinkles, with short, wide pedicel, base with moderate elongate posteromedial apodeme, membranous base of respiratory organ short, annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing (Fig. 41E) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33L) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs (Fig. 41E) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33L); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials—2 elongate setae, 1 campaniform sensillum (as in Figs. 31L–M); anterolaterals—1 moderately long seta; dorsal setae (Fig. 31I)—D-1-T, D-2-T, D-4-T, D-5-T setae, D-3-T campaniform sensillum, D-3-T lateral to D- 4-T; supraalar 2—campaniform sensillum; metathoracics (Fig. 54B)—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: pigmentation light to dark brown, with tergites 1–7 with medial area with stripe, 2 spots or 1 with single medial spot and tergites 2–7 with stripe and 2 spots, medial stripe barely discernable in some, sternites 3–7 with medial stripe, anterolateral spot, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Figs. 78I–K) not strongly modified, terminal processes closely approximated to separated basally, each projecting posterodorsolaterally, tapering to pointed apex; apex of male genital lobe anterior or to level of base of terminal process (Figs. 78I–J); sensilla: tergite 1 (Fig. 54B) with 8 setae, 2 campaniform sensilla, including 4 lateral sensilla, D-2-I, D-3-I closely approximated, D-7-I situated anterolaterally near L-1-I; segment 4 (Fig. 71B)—D-2-IV, D-3- IV moderately elongate setae on moderately developed tubercles; D-5-IV, D-8-IV short setae, D-9-IV elongate seta, D-7-IV present or absent; D-5-IV on short tubercle, D-8-IV, D-9-IV on basally fused, closely approximated tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D-5-IV, D-4-IV, D-8-IV, D-9- IV; D-7-IV, if present, near D-3-IV or L-2-IV; L-1-IV elongate seta on rounded tubercle, just anterior of base of tubercle with L-2-IV, L-3-IV; L-2-IV, L-3-IV, L-4-IV elongate setae on pointed tubercles, L-2-IV, L-3-IV on single tubercle, V-5-IV, V-7-IV moderately elongate setae, V-6-IV elongate seta, on short tubercles, all closely approximated, with tubercles of V-5-IV, V-6-IV fused basally; segment 8 without D-3-VIII, without L-1-VIII; with V-5-VIII, V-6-VIII on single tubercle, V-5-VIII tiny, V-6-VIII elongate. Segment 9 (Figs. 78 I-K)—with D-5-IX, D- 6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Phaenobezzia is known from 34 species from every Region worldwide other than the Australasian Region (Borkent 2014). Immatures have been collected from stream and river margins, seepage pools, rice fields, back waters of rivers, marshes (usually among vegetation), among Pistia stratiotes, lake margins and reservoirs. Knausenberger (1987) provides some further microhabitat information for P. opaca.

TAXONOMIC DISCUSSION: The pupae of nine species of Phaenobezzia are known (Tables 2–3).

Haeselbarth (1965) used the shape of the terminal processes to distinguish the pupae of the four African species he studied and indicated some further character states which may separate some species. Thomsen (1937) drew the dorsal apotome of P. opaca with two setae (and no campaniform sensillum); she likely mistook the campaniform sensillum as the broken base of a seta. Wirth & Grogan (1982) drew abdominal segment 4 of P. opaca with V-7-IV situated laterally, close to L-4-IV and this is significantly different than that observed here for this species and all other Phaenobezzia (Fig. 71B).

MATERIAL EXAMINED: P. cinnae: 2 pupal exuviae, Shaw’s Drift, Eshowe, Zululand, South Africa, 9-IX- 1936 (1 USNM, 1 SAIM); 2 pupal exuviae, no data (ZSMC). P. fulvithorax: 3 pupal exuviae, Bainville, Roosevelt County, Montana, USA, 9-VI-1969 (USNM); 1 pupal exuviae, Cranberry Lake, St. Lawrence County, New York, USA, 25-VI-1963 (USNM); 2 pupal exuviae, Rio Frio, 5 mi NW of Leakey, Real County, Texas, USA, 23-V-1972 (USNM); 1 pupal exuviae, Pedernales River, Gillespie County, Texas, USA, 19-III-1972 (USNM). P. mashonensis: 1 pupal exuviae, Tugela River, Middledrift, South Africa, 6-IX-1935 (USNM); 2 pupal exuviae, Burgershall, Hazyview, East Transvaal, South Africa, 3-XII-1973 (NMSA); 1 pupal exuviae, Sand River between White River and Hazyview, East Transvaal, South Africa, 2-XII-1973 (NMSA); 1 pupal exuviae, Mseleni River, North Zululand, South Africa, 7-II-1936 (1 ZSMC, 1 SAIM); 1 pupal exuviae, N'Kwaleni, Zululand, South Africa, 10- IX-1935 (SAIM). P. opaca: 1 pupal exuviae, Mississippi River, Carleton County, Ontario, Canada, 24-VI-1966 (VPIC); 4 pupal exuviae, Turin, Lewis County, New York, USA, 21-VI-1963 (USNM); 1 pupal exuviae, Independence River, Glenfield, Lewis County, New York, USA, 22-VI-1963 (USNM); 1 pupal exuviae, Meachum Lake, New York, USA, 27-VI-1986 (CNCI); 1 pupal exuviae, Pillsburg State Park, New Hampshire, USA, 26-VI- 1986 (CNCI); 1 pupal exuviae, Dunmore Springs, Pocahontas County, Virginia, USA, 24-VI-1976 (VPIC); 2 pupal exuviae, Bear Creek Lake, Cumberland County, Virginia, USA, 30-IV-1977 (VPIC); 1 pupal exuviae, Jackson River, Allegheny County, Virginia, USA, 4-VI-1977 (VPIC); 1 pupal exuviae, Hamilton County, Florida, USA, 3- VI-1993 (JHEC). P. pistiae: 1 pupal exuviae, Chipinga, Zimbabwe, VIII-1941 (SAIM); 1 pupal exuviae, no location/date (SAIM). P. rubiginosa: 8 pupal exuviae, Anninskoe lake, Pskov Province, Russia, 15-VII-1996 (ZIN). P. vacunae: 4 pupal exuviae, stream nr. Umlalazi River, Eshowe, Zululand, South Africa, 19-IX-1935 (SAIM). P. sp.: 1 pupal exuviae, Canoe, British Columbia, Canada, 7-VII-1990 (CNCI); 1 pupa, 7 pupal exuviae, Dark Canyon Creek, Eddy County, New Mexico, USA, 13-IV-1991 (CNCI); 1 pupal exuviae, 10 mi SE of Middleburg, Lake Dunmore, Vermont, USA, 23-VI-1986 (CNCI); 1 pupal exuviae, Meachum Lake, New York, USA, 21-VI-1986 (CNCI); 1 pupal exuviae, Dunmore Springs, Pocahontas County, West Virginia, USA, 24-VI- 1976 (VPIC); 1 pupal exuviae, Rheocrene, Rio Bento Gomes, Mato Grosso, Brazil, 29-VIII-1994 (CNCI); 1 pupal exuviae (in glycerin), Skulsuza, South Africa, 11-IV-1993 (ANIC); 1 pupal exuviae, Nam Houng river, Moung Sayaboury, Sayaboury Province, Laos, 17-XII-1967 (BPBM); 1 pupal exuviae, as previous locality, 6-I-1968 (BPBM); 2 pupal exuviae, Bamboo Creek, Kimberley, Western Australia, Australia (ANIC); 1 pupal exuviae, Lenard River gorge, Kimberley, Western Australia, Australia, 12-X-1995 (ANIC); 1 pupal exuviae, Mitchell Falls, Kimberley, Western Australia, Australia (ANIC; 1 pupal exuviae, Lower Durack River, Kimberley, Western Australia, Australia (ANIC); 1 pupal exuviae, Crossing Pool, Fortescue River, Pilbara, Western Australia, Australia (ANIC); 2 pupal exuviae, Nattai River, Mittagong, New South Wales, Australia, 15-XI-1968 (ANIC).

STENOXENINI COQUILLETT

Paryphoconus Enderlein

(Figs. 12H, 22L, 28I, 31J, 41F, 46T, 54C, 71C, 78L)

DIAGNOSIS: Only pupa of Ceratopogonidae with abdominal segment 8 with D-3-VIII (Fig. 12H); also, the female is the only Ceratopogonidae with wings abutting ventromedially (Figs. 12H, 41F).

DESCRIPTION: Habitus (female) as in Fig. 12H. Total length = 4.50–6.16 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16B, 33B). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17C, 79H). Head: Dorsal apotome (Fig. 22L), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 28I) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna (Fig. 41F) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), not narrowed sharply posteriorly; sensilla: dorsal apotomals (Fig. 22L)—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 28I)—2 thick setae; oculars (Fig. 28I)—1 seta, 1 campaniform sensillum? or possibly merely an indentation? Thorax: Prothoracic extension (Fig. 28I) wide, well-developed but narrow dorsolaterally, extending from palpus to antenna; mesonotum yes tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 54C); respiratory organ (Fig. 46T) length/width = 3.06–5.06, elongate, moderately slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single curved row, outer surface with some wrinkles, with short, wide pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ short, annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, plates to half length; wing (Fig. 41F) without apical tubercle or angle, with males wings separated medially by fore-, midlegs, female wings abutting medially; halter apex and hind leg (as in Fig. 33J) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs (Fig. 41F) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33K); male with apex of foreleg moderately anterior to apex of midleg (as in Fig. 41E), female with apices of fore-, mid leg not externally visible, dorsal to fused wings; apex of hind leg abutting apex of midleg laterally (dorsal to wing in female); sensilla: anteromedials—2 elongate setae, 1 campaniform sensillum (as in Figs. 31L–M); anterolaterals—1 elongate seta; dorsal setae (Fig. 31J)—D-1-T, D-2-T, D-4-T, D-5-T setae, D-3- T campaniform sensillum; D-1-T, D-2-T separate or on single tubercle, D-3-T lateral to D-4-T; supraalar 2—campaniform sensillum; metathoracics (Fig. 54C)—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: with tergite 1 with 3 medial spots, tergites 2–7 with medial area with stripe and 2 anterolateral spots or with bare patches in 3 medial and anterolateral areas, sternites 3–7 with medial stripe, anterolateral spot; sternite 8 with dark posteromedial apodeme, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 78L) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 54C) with 7 setae, 1 campaniform sensillum, including 3 lateral sensilla, D- 2-I, D-3-I closely approximated, D-7-I absent; segment 4 (Fig. 71C)—D-2-IV, D-3-IV elongate setae on elongate tubercles; D-5-IV, D-8-IV, D-9-IV elongate setae; D-5-IV on single elongate tubercle, D-8-IV, D-9-IV on separate but closely approximated elongate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D-5-IV, D-4-IV, D-8-IV, D-9-IV; D-7-IV near D-3-IV; L-1-IV elongate seta on elongate tubercle, just anterior of base of tubercle with L-3-IV; L-2-IV, L-3-IV, L-4-IV elongate setae on pointed tubercles, V-5-IV, V-6- IV, V-7-IV elongate setae, on short or moderately elongate tubercles, V-6-IV, V-7-IV somewhat closely approximated; segment 8 with D-3-VIII, with or without L-1-VIII; segment 9 (Fig. 78L)—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Paryphoconus is known from 41 species in the Nearctic (one species) and Neotropical Regions (Borkent 2014). Pupae have been collected among vegetation along creeks or on the sandy bottom of a shallow stream.

TAXONOMIC DISCUSSION: The pupae of four species of Paryphoconus are known (Tables 2–3). A female pupa of a Mallochohelea in the ZSMC was labeled " Paryphoconus lanei " but on a modern label and with recent handwriting. The specimen was among material studied by Mayer but does not correspond to his description of Paryphoconus flavidus (as lanei) (Mayer 1959b). The Paryphoconus pupae studied by Mayer (1959b) were not located.

MATERIAL EXAMINED: P. grandis: 2 pupal exuviae, Ruta Nacional 12, Corrientes, Argentina, 18-IX- 2010 (MLPA). P. oliveirai: 1 pupal exuviae, Parque das Garcas, Amazonas Igarape, Brazil, 14-X-2005 (INPA).

Phaenobezzia Haeselbarth

(Figs. 17F, 22K, 28H, 31I, 33L, 41E, 46S, 54B, 71B, 78I–K)

TAXONOMIC DISCUSSION: The pupae of nine species of Phaenobezzia are known (Tables 2–3).

STENOXENINI COQUILLETT

Paryphoconus Enderlein

(Figs. 12H, 22L, 28I, 31J, 41F, 46T, 54C, 71C, 78L)

DIAGNOSIS: Only pupa of Ceratopogonidae with abdominal segment 8 with D-3-VIII (Fig. 12H); also, the female is the only Ceratopogonidae with wings abutting ventromedially (Figs. 12H, 41F).

Paryphoconus Enderlein

(Figs. 12H, 22L, 28I, 31J, 41F, 46T, 54C, 71C, 78L)

DIAGNOSIS: Only pupa of Ceratopogonidae with abdominal segment 8 with D-3-VIII (Fig. 12H); also, the female is the only Ceratopogonidae with wings abutting ventromedially (Figs. 12H, 41F).