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).