Amerohelea Grogan & WirthBezzia KiefferClastrieromyia Spinelli & GroganPachyhelea WirthPalpomyia MeigenPhaenobezzia HaeselbarthParyphoconus Enderlein

Borkent, Art, 2014, The Pupae of the Biting Midges of the World (Diptera: Ceratopogonidae), With a Generic Key and Analysis of the Phylogenetic Relationships Between Genera, Zootaxa 3879 (1), pp. 1-327: 100-113

publication ID

http://dx.doi.org/10.11646/zootaxa.3879.1.1

publication LSID

lsid:zoobank.org:pub:6423894B-97D9-4286-ABB9-D4AF072B57FD

DOI

http://doi.org/10.5281/zenodo.5593061

persistent identifier

http://treatment.plazi.org/id/027587C9-BD4D-3015-FD9B-18C24FB0E49C

treatment provided by

Felipe

scientific name

Amerohelea Grogan & Wirth Bezzia Kieffer Clastrieromyia Spinelli & Grogan Pachyhelea Wirth Palpomyia Meigen Phaenobezzia Haeselbarth Paryphoconus Enderlein
status

 

Amerohelea Grogan & Wirth  

( Figs. 22B View FIGURE 22 , 46G View FIGURE 46 )

DIAGNOSIS: known only from one limited description by Lane et al. (1955) and not diagnosable.

DESCRIPTION: Known only from one limited description by Lane et al. (1955), with the following character states discernable: Head: Dorsal apotome ( Fig. 22B View FIGURE 22 ), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome. Thorax: Respiratory organ ( Fig. 46G View FIGURE 46 ) length/width = 4.00, elongate, moderately slender, with pores closely abutting at apex of respiratory organ, arranged in single row. Abdomen: tergites or sternites entire, each without membranous disc; segment 9 not strongly modified, terminal processes separated basally, each projecting posterodorsolaterally, tapering to pointed apex.

DISTRIBUTION AND HABITAT: The genus Amerohelea   is known from 13 species in the New World ( Borkent   2014). The only specimen known was collected from a reservoir with abundant vegetation and open, sunny exposure ( Lane et al. 1955).

TAXONOMIC DISCUSSION: No specimens of this genus were examined and the pupa is known only from the brief description of A. sordidipes   by Lane et al. (1955). Lane et al. (1955) included drawings of the dorsal apotome, respiratory organ and abdominal tergites 3–9. The dorsal apotome is of a unique outline, no sensilla were illustrated ( Fig. 22B View FIGURE 22 ) and it is almost certainly poorly drawn. The respiratory organ was drawn in an anterior/ posterior perspective ( Fig. 46G View FIGURE 46 ) and so the length/width noted here as 4.00 (taken from the drawing) is only approximate. The drawing of the abdomen did not provide enough resolution to determine the identification of most sensilla (only D-2-IV and D-3-IV can be identified with any confidence) and is not replicated here.

The few features of the pupa previously reported means this genus could neither be keyed nor interpreted phylogenetically. The presence of abdominal tergal apodemes of adult females indicates the genus is a member of the Palpomyiini   .

The previously described pupa of A. sordidipes   was not in the collections of the Faculdade de Saude Publica, Universidade de Sao Paulo (Anice Sallum, pers. comm.) or at the Museu de Zoologia, Universidade de São Paulo (Carlos José Einicker Lamas, pers. comm.) both of which otherwise house some Lane material.

MATERIAL EXAMINED: None available.

Bezzia Kieffer  

( Figs. 12F View FIGURE 12 , 13I View FIGURE 13 , 17D View FIGURE 17 , 22C–D View FIGURE 22 , 28D View FIGURE 28 , 31E View FIGURE 31 , 33J View FIGURE 33 , 41A View FIGURE 41 , 46H–K View FIGURE 46 , 53D View FIGURE 53 , 69C View FIGURE 69 , 77E–H View FIGURE 77 )

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. 53D View FIGURE 53 ), apex of the halter extending posteriorly to about 1/6 length of tergite 2 (as in Fig. 33L View FIGURE 33 ), abdominal segment 4 with V-5-IV, V-6-IV and V-7-IV closely approximated (as in Fig. 70C View FIGURE 70 ) 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 ( Fig. 69C View FIGURE 69 ) (not as in Fig. 70B View FIGURE 70 ), 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 well-developed (not minute), and abdominal segment 8 is without L-1-VIII (not diagnosable as different from Palpomyia   and Phaenobezzia   ); however, most species of Bezzia   have two or more campaniform sensilla on the dorsal apotome ( Figs. 22C–D View FIGURE 22 ), a nearly unique condition found only in two species of Palpomyia   , one of which, P. jonesi   , distinctively has two setae and two campaniform sensilla ( Fig. 22J View FIGURE 22 ).

DESCRIPTION: Habitus as in Fig. 12F View FIGURE 12 . Total length = 2.00– 6.16 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing ( Figs. 17D View FIGURE 17 , 33J View FIGURE 33 ). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17C View FIGURE 17 , 79H View FIGURE 79 ) or with eye appressed to antenna ( Figs. 17D View FIGURE 17 ). Head: Dorsal apotome ( Figs. 22C–D View FIGURE 22 ), with partial ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite ( Fig. 13I View FIGURE 13 ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 28D View FIGURE 28 ) 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. 41A View FIGURE 41 ) anterior to posterior to, posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Figs. 22C–D View FIGURE 22 )—1 moderate to elongate seta, 1–3 campaniform sensilla; dorsolateral cephalic sclerite sensilla—1 seta, 0–1 campaniform sensillum; clypeal-labrals ( Fig. 28D View FIGURE 28 )—1 slender or 2 slender or thick setae; oculars ( Fig. 28D View FIGURE 28 )—1–2 seta, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 28D View FIGURE 28 ) wide, well-developed, extending from palpus to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 53D View FIGURE 53 ); respiratory organ ( Figs. 46H–K View FIGURE 46 ) length/width = 3.73–6.60, moderately elongate to elongate, apical portion swollen in some, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single straight to curved row, outer surface with some wrinkles, with short, wide pedicel, base with short to moderately 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 or more; wing ( Fig. 41A View FIGURE 41 ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 33J View FIGURE 33 ) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs ( Fig. 41A View FIGURE 41 ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 33J View FIGURE 33 ); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally or small gap between the two; sensilla: anteromedials—2 elongate setae, 1 campaniform sensillum (as in Figs. 31L–M View FIGURE 31 ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 31E View FIGURE 31 )—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. 53D View FIGURE 53 )—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: pigmentation light brown, with tergite 1 with 3 medial spots, tergites 2–7 with medial area with stripe, 2 anterolateral spots, sternites 3–7 with medial stripe, anterolateral spot (light brown), 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. 77E–H View FIGURE 77 ) not strongly modified, terminal processes closely approximated to separated basally, each projecting posterodorsolaterally, tapering to pointed apex, in some very slender; sensilla: tergite 1 ( Fig. 53D View FIGURE 53 ) with 8 setae, 2 campaniform sensilla, including 3–4 lateral sensilla, D-2-I, D-3-I closely approximated, D-7-I situated anterolaterally near L-1-I to anteriorly near D-3-I; segment 4 ( Fig. 69C View FIGURE 69 )—D-2-IV, D-3-IV short setae on short tubercles; D-5-IV, D-8-IV, D-9-IV short to moderately elongate setae, D-7-IV present or absent; D-5-IV on single, 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 moderately elongate seta on rounded tubercle, well anterior of posterior lateral setae; L-2- IV, L-3-IV, L-4-IV short to moderately elongate setae on rounded tubercles, L-2-IV, L-3-IV on single tubercle in some, V-5-IV, V-6-IV, V-7-IV short setate 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; with V-5-VIII, V-6-VIII on single tubercle, V-5-VIII tiny, V-6- VIII elongate; segment 9 ( Figs. 77E–H View FIGURE 77 )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Bezzia   is known from 319 species from every Region worldwide ( Borkent 2014). Immatures have been recorded from a wide array of habitats including rice fields, springs, streams, river margins, wet moss, various phytotelmata, pools, marshes, bogs, fens, algal mats in lentic habitats, lakes, and reservoirs.

TAXONOMIC DISCUSSION: The pupae of 51 species of Bezzia   are known ( Tables 2–3 View TABLE 2 ). Goeze (1780) described but did not name a species which is probably a Bezzia   , providing the first morphological study describing the chaetotaxy of the larva and pupa ( Fig. 6B View FIGURE 6 ) of a Ceratopogonidae   . It was later named Tipula goezii   by Schrank (1803). I have placed it in Bezzia   as a new combination as follows but the name is so out of date that it must be considered a nomen nudum. The long slender terminal process and somewhat clavate apex of the respiratory organ ( Figs. 46H, 46J View FIGURE 46 ) is a distinctive combination of many species of Bezzia   .

Bezzia goezii (Schrank)   , 1803: 72 ( Tipula   ). Quedlinburg, Germany. new combination.

Some species have been described more than once by the same author ( Table 2 View TABLE 2 ) but with different names (now synonyms) (e.g. B. circumdata   , B. leucogaster   , B. nobilis   ), suggesting either misidentifications or the possibility of more than one species actually present.

Thomsen (1937) gave a key to three species known to her at that time but one of these, P. flavitarsis   , is now recognized as a Palpomyia   . Thomsen (1937) drew the dorsal apotomes of species of Bezzia   with more than one seta (per side) but likely misinterpreted the multiple campaniform sensilla as the broken bases of setae.

Thienemann (1928) and Mayer (1934a) provided keys to European species groups. Lenz (1934) provided a key to European species but primarily used habitats to distinguish these and, for most, this almost certainly results in inaccurate identifications. Harris (1981) described and keyed the pupae of six unnamed species of Bezzia   from Australia.

Wirth (1983a, 1983b, 1983c) diagnosed the pupae of some Nearctic and Neotropical Bezzia   in the cockerelli   , bicolor   , and nobilis   species groups but did not do so for the remaining species groups of Bezzia   . At present, pupae of Bezzia   cannot be diagnosed as a genus and therefore providing a key to the species in a given region is superfluous. Unless a diagnosis becomes available all Bezzia   should be keyed with species of Palpomyia   and Phaenobezzia   . Alternately, it would be possible to key all species with more than one campaniform sensillum on the dorsal apotome, which would include nearly all Bezzia   and a few Palpomyia   , depending on the area being covered.

Nearly all Bezzia   have more than one campaniform sensillum on the dorsal apotome (see character 6). The only exceptions seen are of B. brevicornis   , B. bromeliae   and B. dorsasetula   , each with one seta and one campaniform sensillum.

Bezzia xanthogaster (Kieffer)   , 1919: 130 ( Probezzia   ) is a junior homonym of Probezzia xanthogaster (Kieffer)   , 1917: 329 ( Bezzia   ) and is here given the new name Bezzia gilvigaster   (a similar specific name also meaning yellow stomach (abdomen for the adult this species).

MATERIAL EXAMINED: B. africana   : 1 pupal exuviae, Burgershall , Hazyview, Transvaal, South Africa, 3- XII-1973 ( NMSA)   ; 1 pupal exuviae, Magaliesberg Agricultural School , Transvaal, South Africa, 13-XI-1973 ( NMSA)   . B. albicornis   : 1 pupal exuviae, Kaushut , Tedjen Province, Turkmenistan, 2-IV-1972 ( ZIN)   ; 1 pupal exuviae, as previous locality, 30-V-1972 ( ZIN)   . B. amana   : 1 pupal exuviae (of holotype), Loutpan , Transvaal, South Africa, 10-I-1974 ( NMSA)   . B. annulipes   : 2 pupal exuviae, Lunzer Untersee between boat house and canal, Lower Austria, Austria, 6-VI-1942 ( ZSMC)   ; 1 pupal exuviae, Germany ( ZSMC)   ; 3 pupal exuviae, Germany ( ZSMC)   ; 12 pupal exuviae, Anninskoe lake , Pskov Province, Russia, 24-VI-1995 ( ZIN)   . B. biannulata   : 1 pupal exuviae (of paratype), 1 pupal exuviae (of allotype), Oceano Beach, San Luis Obispo County, California, USA, 20- VIII-1948 ( USNM)   . B. bicolor   : 6 pupal exuviae, Gievenbeck , Germany, 9-V-1912 ( ZSMC)   ; 4 pupal exuviae, Lunz , Austria ( USNM)   ; 4 pupal exuviae, Shushary , Leningrad Province, Russia, 29-V-1997 ( ZIN)   ; 2 pupal exuviae, Strelna , Leningrad Province, Russia, 12-V-2002 ( ZIN)   . B. bivittata   : 3 pupal exuviae, Salisbury , Wicomico County, Maryland, USA, 4-V-1981 ( WLGC)   ; 1 pupal exuviae, as previous locality, 13-IV-1981 ( WLGC)   . B. blantoni   : 2 pupal exuviae (of paratype), Escobar , Buenos Aires, Argentina, 10-I-1982 ( MPLA)   . B. brevicornis   : 1 pupal exuviae, Magdalena , Buenos Aires, Argentina, 24-XII-1981 ( MPLA)   . B. bromeliae   : 2 pupal exuviae (of paratype), Bayano Field Station , Panama Province, Panama, VI-1976 ( MPLA)   . B. circumdata   : 3 pupal exuviae, Black Lake , Stanleyville, Ontario, Canada, 24-VI-1975 ( USNM)   ; 1 pupal exuviae, as previous locality, 25-VI-1975 ( USNM)   ; 4 pupal exuviae, Rotmoos on Mittersee , Lower Austria, Austria, 10-VI-1942 ( ZSMC)   ; 1 pupa, 5 pupal exuviae, Grosser Plöner See , Slesvig-Holstein, Germany, summer 1952 ( ZSMC)   ; 2 pupa, 5 pupal exuviae, kerosene port on Dortmund-Ems canal, Dortmund, North Rhine-Westphalia, Germany, 28-VII-1908 ( ZSMC)   ; 1 pupa, 3 pupal exuviae, Germany ( ZSMC)   ; 4 pupal exuviae, Vyborg , Leningrad Province, Russia, 27-VII-1998 ( ZIN)   . B. cockerelli   : 1 pupal exuviae, Valdez , Alaska, USA, 1948 ( USNM)   ; 1 pupal exuviae, Bainville , Roosevelt County, Montana, USA, 9-VI-1969 ( USNM)   ; 1 pupal exuviae, Trails Pond , Latah County, Idaho, USA, 22-VII-1969 ( USNM)   ; 5 pupal exuviae, 4 mi N of Upham , McHenry County, North Dakota, USA, 5-VI-1969 (4 USNM, 1 VPIC)   . B. collessi   : 1 pupal exuviae (of paratype), Singapore, 10-IX-1952 ( USNM)   ; 3 pupal exuviae (of paratypes), as previous locality, 28-VIII-1952 ( USNM)   . B. dorsasetula   : 2 pupal exuviae, College Park , Prince George’s County, Maryland, USA, 30-V-1975 ( USNM)   , 1 pupal exuviae, no locality, 6-V-1977 ( VPIC)   . B. fascispinosa   : 1 pupal exuviae, Lakeland Pond , College Park, Prince George’s County, Maryland, USA, 30-V-1975 ( USNM)   ; 1 pupal exuviae, Beaver Lake Reservoir, Pocahontas State Park, Chesterfield County, Virginia, USA, 30-IV-1977 ( VPIC)   ; 1 pupal exuviae, Iron Spring , Elliot Know, Augusta County, Virginia, USA, 22-17-1977 ( VPIC)   . B. flavicornis   : 1 pupal exuviae, Priluki , Belarus, 17-V-1967 ( ZIN)   . B. flavicorporis   : 1 pupal exuviae, Loutpan , Transvaal, South Africa, 10-I-1974 ( NMSA)   . B. gibbera   : 1 pupal exuviae, Dismal Swamp , Camden County, North Carolina, USA, 25-III-1976 ( VPIC)   . B. glabra   : 2 pupal exuviae, Morgan Arboretum , St. Anne de Bellevue, Quebec, Canada, 1964 ( USNM)   ; 1 pupal exuviae, 1 mi. S of Corapeake, Dismal Swamp , Camden County, North Carolina, USA, 29-VII-1976 ( VPIC)   ; 2 pupal exuviae, Keowee Reserve , Seneca , Oconee County, South Colorado, USA, 2-V-1975 ( VPIC)   . B. japonica   : 1 pupal exuviae, Suputinka river , Ussuri Nature Reserve, Primorskii Territory, Russia, 30-V-1973 ( ZIN)   . B. kuhetiensis   : 1 pupal exuviae, Issyk-Kul lake , Kyrgyzstan, 16-VI-1971 ( ZIN)   . B. laciniastyla   : 1 pupal exuviae, Jackson River , approx. 10 mi. upriver from Covington, Natural Well, Allegheny County, Virginia, USA, 4-VI-1977 ( VPIC)   . B. leucogaster   : 4 pupal exuviae, shore of Lottsee SE of Mölln, Slesvig-Holstein, Germany, 16-V-1926 ( ZSMC)   , 2 pupal exuviae, shore of Kirchsee , Preetz (between Kiel and Plön), Slesvig-Holstein, Germany, 11-VII1918 ( ZSMC)   ; 1 pupal exuviae, Severskii Donets River , Donetsk Province, Ukraine, 5-V-1970 ( ZIN)   . B. narynica   : 1 pupal exuviae, Issyk-Kul lake , Kyrgyzstan, 24-V-1971 ( ZIN)   . B. nigrita   : 1 pupal exuviae, Severskii Donets River , Donetsk Province, Ukraine, 5-V-1970 ( ZIN)   ; 1 pupal exuviae, Malinovka , Donetsk Province, Ukraine, 8-V-1970 ( ZIN)   . B. nobilis   : 1 pupal exuviae, 7 km S of Hope , BC, Canada, 8–9-VII-1985 ( CNCI)   ; 1 pupal exuviae, Beaver Creek , Lawrence County, South Dakota, USA, 15-VI-1969 ( USNM)   ; 1 pupal exuviae, Cow Neck Salt Marsh , North Sea, New York, USA, 19-IV-1956 ( NYSM)   ; 1 pupal exuviae, as previous locality, 17-IV-1956 ( NYSM)   ; 4 pupal exuviae, Fishing Creek , Newcomb, New York, USA, 28-V-1958 ( NYSM)   ; 1 pupal exuviae, Port Leyden , New York, USA, 4-VII-1959 ( USNM)   ; 2 pupal exuviae, Letchworth State Park , New York, USA, 13-VI-1963 ( USNM)   ; 1 pupal exuviae, Patuxent Refuge , Maryland, USA, 8-V-1958 ( USNM)   ; 1 pupal exuviae, Vepco Property , Louisa County, Virginia, USA, 12-IX-1976 ( VPIC)   ; 1 pupal exuviae, Washburn County, Wisconsin, USA, 1-IX-1951 ( VPIC)   ; 2 pupal exuviae, Vepco Property , Louisa County, Virginia, USA, 12-IX-1976 ( VPIC)   ; 2 pupal exuviae, Great Swamp , Isle of Wight County, Virginia, USA, 2-VIII- 1976 ( VPIC)   ; 1 pupal exuviae, Cranberry Glade , Pocahontas County, Virginia, USA, 8-IX-1976 ( VPIC)   ; 1 pupal exuviae, Norfolk Gardens , Norfolk County, Virginia, USA, 1-VIII-1976 ( VPIC)   . 1 pupal exuviae, no locality/date ( VPIC); 1 pupal exuviae, Pecan Springs , Devils River, Juno, Texas, USA ( USNM)   ; 1 pupal exuviae, Baton Rouge , Louisiana, USA, 4-III-1947 ( USNM)   ; 1 pupal exuviae, Beloe lake , Alol’, Pskov Province, Russia, 27-VI-1969 ( ZIN)   ; 1 pupal exuviae, Beloe lake , Alol’, Pskov Province, Russia, 5-VII-1969 ( ZIN)   . B. obelisca   : 2 pupal exuviae, Mer Bleue , Ottawa, Ontario, Canada, 27-V-1960 ( USNM)   ; 2 pupal exuviae, McLean Reserve , Tompkins County, New York, USA, 18-VI-1963 ( USNM)   ; 1 pupal exuviae, edge of Big Glade , Cranberry Glades Botanical Area, Pocahontas County, Virginia, USA, 6-IX-1976 ( VPIC)   . B. perplexa   : 1 pupal exuviae, Millpond Creek , North Anna Resrvoir, Louisa County, Virginia, USA, 1-III-1977 ( VPIC)   . B. pulchripes   : 2 pupal exuviae, Mission Rio Cururu Igarape , Brazil, 4-V-1941 ( SDEI)   ; 1 pupal exuviae, Mocoreta , Corrientes, Argentina, 21-III-1985 ( MPLA)   . B. roldani   : 2 pupal exuviae, no locality, 20-VIII-1979 ( MPLA)   . B. saileri   : 1 pupal exuviae (of paratype), Fire Lake , Anchorage, Alaska, USA, 29-V-1948 ( USNM)   . B. signata   : 1 pupal exuviae, Luchevoi , Republic of Karelia, Russia, 14-VI-1966 ( ZIN)   . B. sordida   : 1 pupal exuviae, Olerna , Marin County, California, USA, 22-II-1947 ( USNM)   . B. turkmenica   : 1 pupal exuviae, Hodzhaz-Kala bay , Kyzyl-Arvat Province, Turkmenistan, 18-IV-1972 ( ZIN)   . B. uncistyla   : 2 pupal exuviae, 5 km E of Danby , Vermont, USA, 25-VI-1986 ( CNCI)   ; 1 pupal exuviae, as previous locality, 25–26-VI-1986 ( CNCI)   . B. varicolor   : 1 pupal exuviae, Rideau River , Ottawa, Ontario, Canada, 29-V-1960 ( USNM)   ; 1 pupal exuviae, Lake Temescal , Berkeley, California, USA, 1-V-1948 ( USNM)   ; 1 pupal exuviae, Lake Tomahawk at Kemp Biological Station, Oneida County, Wisconsin, USA, 20-VII-1978 ( USNM)   ; 2 pupal exuviae, Bar Lake , Benzie County, Michigan, USA, 8-VIII-1975 ( VPIC)   ; 1 pupal exuviae (of topotype), Mill Creek Brook, Noyack, New York, USA, 6-V-1057 ( USNM)   ; 1 pupal exuviae, Slaterville Spr. , New York, USA, 12- VI-1964 ( VPIC)   ; 1 pupal exuviae, Lakeland Pond , College Park, Prince George’s County, Maryland, USA, 24-IV- 1977 ( VPIC)   . B. nr. bicolor   : 1 pupal exuviae, no locality, 30-V-1953 ( USNM)   . B. nr. nobilis   : 1 pupal exuviae, Back Bay National Wildlife Refuge , Virginia Beach, Virginia, USA, 30-VIII-1975 ( VPIC)   . B. nr. obelisca   : 1 pupal exuviae, Mer Bleue , Ottawa, Ontario, Canada, 27-V-1960 ( USNM)   ; 2 pupal exuviae, no data ( USNM); 1 pupal exuviae, Mer Bleue , Ottawa, Ontario, Canada, 27-V-1960 ( VPIC)   ; 1 pupal exuviae, Big Glade , Cranberry Glade Botanical Area, Pocahontas County, Virginia, USA, 8-IX-1976 ( VPIC)   . B. nr. pruinosa   : 1 pupal exuviae, no locality, 12-V-1953 ( USNM)   . B. nr. varicolor   : 1 pupal exuviae, W flank of Salt Pond Mountain , Giles County, Virginia, USA, 14-VII-1977 ( VPIC)   . B. sp.: 2 pupal exuviae (in glycerin), 12 pupal exuviae, Bolean Lake , 6 km NE of Falkland, British Columbia, Canada, 12-13-VII-1989 ( CNCI)   ; 1 pupal exuviae (in glycerin), Spanish Lake , 6 km E of Falkland, 50°29.12N 119°28.07W, British Columbia, Canada, 27–28-V-2008 ( CNCI) GoogleMaps   ; 4 pupal exuviae, 3 km E of Salmon Arm , British Columbia, Canada, 9-VI-1988 ( CNCI)   ; 9 pupal exuviae, 6 km E of Salmon Arm , British Columbia, Canada, 6-VI-1990 ( CNCI)   ; 5 pupal exuviae, 9 km S of Salmon Arm , British Columbia, Canada, 19-VII-1988 ( CNCI)   ; 2 pupal exuviae, 8 km E of Sicamous , British Columbia, Canada, 1-VI-1992 ( CNCI)   ; 2 pupal exuviae, 6.5 km NW of Enderby , British Columbia, Canada, 6-V-1992 ( CNCI)   ; 4 pupal exuviae, 30 km N of Nakusp , British Columbia, Canada, 5-VII-1990 ( CNCI)   ; 4 pupal exuviae, 20 km E of Anola , Manitoba, Canada, 16-VI-1990 ( CNCI)   ; 3 pupal exuviae, Trail Bay , Manitoba, Canada ( CNCI)   ; 1 pupal exuviae (in glycerin), 1 mi. E of Orleans , Ontario, Canada, 8-VII-1971 ( CNCI)   ; 1 pupal exuviae, 6 km N of Eardlay , Quebec, Canada, 15-V-1986 ( CNCI)   ; 2 pupal exuviae, Fire Lake , Anchorage, Alaska, USA, 29-V-1948 ( USNM)   ; 5 pupal exuviae, 9 km W of Okanogan , Washington, USA, 13-VI-2008 ( CNCI)   ; 9 pupal exuviae, High Creek Fen Preserve , 9 mi S of Fairplay Park, Colorado, USA, 18-VI-1995 ( CNCI)   ; 1 pupal exuviae, Blue Mountain Lake , New York, USA, 30-VI-1958 ( WLGC)   ; 2 pupal exuviae, Ringwood Pond , Dryden, New York, USA, 7-III-1933 ( USNM)   ; 1 pupal exuviae, Keowee Reserve , Seneca, Ocouee County, South Carolina, USA, 16-V-1974 ( USNM)   ; 1 pupal exuviae, as previous locality, USA, 16-VII-1974 ( USNM)   ; 1 pupal exuviae, 8 km W of Atenas , Costa Rica, 19-VIII-1993 ( CNCI)   ; 2 pupal exuviae, carreterra entre Santa Cecilia y Upala, 1.6 km N.O. de BVirmania, Dos Rios, Upala , Alejuela, Costa Rica, 16-X-2006 ( INBC)   ; 1 pupal exuviae, Barro Colorado Island , Canal Zone, Panama, 24-VI- 1996 ( CNCI)   ; 2 pupal exuviae, as previous locality, 31-X-1995 ( CNCI)   ; 1 pupal exuviae, as previous locality, 30- VI-1996 ( CNCI)   ; 1 pupal exuviae, as previous locality, 10-IX-1995 ( CNCI)   ; 3 pupae, Lerchenauer See , Germany, 26-VII-1990, 27-VII-1990, 2-VIII-22-IX-1990 ( CNCI)   ; 1 pupa, Chiemsee , Germany, 13-VIII-1990   ; 1 pupa, 1 pupal exuviae, Bottsand nr. Stein, Germany, 5-VI-1932 ( ZSMC)   ; 2 pupal exuviae, Darban , Western Australia, Australia, 28-X-1986 ( ANIC)   ; 9 pupal exuviae, as previous locality, 28-X-1985 ( ANIC)   ; 2 pupal exuviae, 5 km N of Darban , Western Australia, Australia, 29-X-1985 ( ANIC)   ; 3 pupal exuviae, Roper River Mission, Northern Territory, Australia, 8-XI-1956 ( ANIC)   ; 3 pupal exuviae, Humpty Doo , Northern Territory, Australia, V-1957 ( ANIC)   ; 1 pupal exuviae, Dayboro , Queensland, Australia, 10-VII-1951 ( ANIC)   ; 10 pupal exuviae, Nattai River , Mittagong, New South Wales, Australia, 4-XI-1964 ( ANIC)   ; 1 pupal exuviae, as previous locality, 15-XI-1968 ( ANIC)   ; 1 pupal exuviae, Gap Creek , The Crags, Mittagong, New South Wales, Australia, 10-II-1966 ( ANIC)   ; 5 pupal exuviae, Damin clearing, The Crags, Mittagong, New South Wales, Australia, 9-II-1966 ( ANIC)   ; 2 pupal exuviae, Deep Creek , Nanasheen, New South Wales, Australia, 7-XII-1956 ( ANIC)   ; 2 pupal exuviae, as previous locality, 25-XI-1956 ( ANIC)   ; 21 pupal exuviae, Middle Creek , New South Wales, Australia, 8-II-1966 ( ANIC)   ; 1 pupal exuviae, Colo Vale , New South Wales, Australia, 16-X-1956 ( ANIC)   ; 2 pupal exuviae, Hornsby , New South Wales, Australia, 18-XI-1969 ( ANIC)   ; 2 pupal exuviae, as previous locality, 25-X-1956 ( ANIC)   ; 2 pupal exuviae, Nepean River , Menangle, New South Wales, Australia, 9-XII-1968 ( ANIC)   ; 1 pupal exuviae, Griffith , New South Wales, Australia, 16-XI-1956 ( ANIC)   ; 1 pupal exuviae, Moruya River , Merricumbene Creek, New South Wales, Australia, 2-III-1964 ( ANIC)   ; 1 pupal exuviae, McCarrs Creek , New South Wales, Australia, 14-I-1969 ( ANIC)   ; 1 pupal exuviae, no locality, 14-I-1937 ( SAIM)   .

Clastrieromyia Spinelli & Grogan  

( Figs. 22F View FIGURE 22 , 28E View FIGURE 28 , 31F View FIGURE 31 , 41B View FIGURE 41 , 46L View FIGURE 46 , 53E View FIGURE 53 , 70A View FIGURE 70 , 77I–J View FIGURE 77 )

DIAGNOSIS: Only pupa of Ceratopogonidae   with the metathorax with a single campaniform sensillum (M-3-T) situated at least ⅓ the length of the metathorax from its anterior margin ( Fig. 53E View FIGURE 53 ), apex of the halter extending posteriorly to about 1/6 length of tergite 2 (as in Fig. 33L View FIGURE 33 ), D-7-I near D-3-I ( Fig. 53E View FIGURE 53 ), abdominal segment 4 with each setae on a rounded tubercle ( Fig. 70A View FIGURE 70 ) and abdominal segment 8 without L-1-VIII.

DESCRIPTION: Total length = 3.20–3.80 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16B View FIGURE 16 , 33B View FIGURE 33 ). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17C View FIGURE 17 , 79H View FIGURE 79 ). Head: Dorsal apotome ( Fig. 22F View FIGURE 22 ), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13H View FIGURE 13 ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 28E View FIGURE 28 ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna ( Fig. 41B View FIGURE 41 ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Fig. 22F View FIGURE 22 )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum on very short tubercle; clypeal-labrals ( Fig. 28E View FIGURE 28 )—2 slender setae; oculars ( Fig. 28E View FIGURE 28 )—2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 28E View FIGURE 28 ) wide, well-developed but very narrow dorsolaterally, extending from palpus to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 53E View FIGURE 53 ); respiratory organ ( Fig. 46L View FIGURE 46 ) length/width = 3.13–4.08, elongate, moderately slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single 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. 41B View FIGURE 41 ) with short tubercle at apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33J View FIGURE 33 ) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs ( Fig. 41B View FIGURE 41 ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33J View FIGURE 33 ); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials—1 long, 1 medium setae, 1 campaniform sensillum; anterolaterals—1 moderately long seta; dorsal setae ( Fig. 31F View FIGURE 31 )—D-1-T, D-2-T, D-4-T, D- 5-T setae, D-3-T campaniform sensillum, D-3-T anterior to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 53E View FIGURE 53 )—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: pigmentation light brown, with tergite 1 with 3 medial spots, tergites 2-7 with medial area with stripe, 2 anterolateral spots, sternites 3-7 with medial stripe, anterolateral spot (light brown), segment 2 as wide or slightly wider than segment 3, segments with undivided, peg-like or thin to thick setae, with rounded, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Fig. 77 View FIGURE 77 I-J) not strongly modified, terminal processes separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 53E View FIGURE 53 ) with 7 setae, 2 campaniform sensilla, including 3 lateral sensilla, D-2-I, D-3-I closely approximated, D-7-I situated anteriorly near D-3-I; segment 4 ( Fig. 70A View FIGURE 70 )—D-2-IV, D-3-IV short to moderately elongate setae on short tubercles; D-5-IV peg-like seta, D-8-IV, D-9-IV short to moderately elongate setae; D-5-IV on single tubercle, D-8-IV, D-9-IV on 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 near D-3-IV; L- 1-IV short seta on rounded tubercle, well anterior of posterior lateral setae; L-2-IV, L-4-IV short setae, L-3-IV moderately elongate seta on rounded tubercles, V-5-IV, V-6-IV short setae, V-7-IV moderately elongate, on moderately elongate tubercles, V-5-IV, V-6-IV moderately closely approximated; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Fig. 77I–J View FIGURE 77 )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Clastrieromyia   is known from four species in the Neotropical Region ( Borkent   2014). Immatures of the only known species have been collected from a cattle-trodden bog.

TAXONOMIC DISCUSSION: Only one species is known as a pupa ( Tables 2–3 View TABLE 2 ).

MATERIAL EXAMINED: C. dycei   : 2 pupal exuviae (of paratypes), Tacuarembo, Uruguay, 29-IX-1980 ( MLPA)   .

Pachyhelea Wirth  

( Figs. 22G View FIGURE 22 , 28F View FIGURE 28 , 31G View FIGURE 31 , 41C View FIGURE 41 , 46M View FIGURE 46 , 53F View FIGURE 53 , 70B View FIGURE 70 , 77K–L View FIGURE 77 )

DIAGNOSIS: Only pupa of Ceratopogonidae   with dorsal apotome with only one campaniform sensillum ( Fig. 22G View FIGURE 22 ), metathorax with only one campaniform sensillum (M-3-T) situated at least ⅓ the length of the metathorax from its anterior margin ( Fig. 53F View FIGURE 53 ), apex of the halter extending posteriorly to about 1/6 length of tergite 2 (as in Fig. 33L View FIGURE 33 ), L-1-IV and L-3-IV short setae on separate tubercles, L-2-IV well separated from L-3-IV, V-7-IV is closer to L-4-IV than to V-5-IV and V-6-IV, which are closely abutting ( Fig. 70B View FIGURE 70 ).

DESCRIPTION: Total length = 7.22 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16B View FIGURE 16 , 33B View FIGURE 33 ). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17C View FIGURE 17 , 79H View FIGURE 79 ). Head: Dorsal apotome ( Fig. 22G View FIGURE 22 ), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13H View FIGURE 13 ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 28F View FIGURE 28 ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 41C View FIGURE 41 ) barely anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Fig. 22G View FIGURE 22 )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 28F View FIGURE 28 )—2 slender setae; oculars ( Fig. 28F View FIGURE 28 )—1 seta, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 28F View FIGURE 28 ) wide, well-developed but very narrow dorsolaterally, extending from palpus to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 53F View FIGURE 53 ); respiratory organ ( Fig. 46M View FIGURE 46 ) length/width = 3.96–5.00, elongate, moderately slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single 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, wrinkles to half length; wing ( Fig. 41C View FIGURE 41 ) with short tubercle at apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33J View FIGURE 33 ) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs ( Fig. 41C View FIGURE 41 ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33J View FIGURE 33 ); 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 View FIGURE 31 ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 31G View FIGURE 31 )—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. 53F View FIGURE 53 )—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, 2 anterolateral spots, 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 tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Figs. 77K–L View FIGURE 77 ) not strongly modified, terminal processes separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 53F View FIGURE 53 ) with 7 setae, 3 campaniform sensilla, including 3 lateral sensilla, D-2- I, D-3-I closely approximated, D-7-I situated anteriorly near D-3-I; segment 4 ( Fig. 70B View FIGURE 70 )—D-2-IV, D-3-IV short to moderately elongate setae, D-2-IV on short tubercle, D-5-IV very short seta, D-8-IV, D-9-IV short setae; D-5-IV without tubercle, D-8-IV, D-9-IV on 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 near D-3-IV; L-1-IV short seta on rounded tubercle, near L-3-IV; L-2-IV, L-3-IV, L-4-IV short setae, L-2-IV, L-3-IV on very short tubercles, L-4-IV on pointed, elongate tubercle, V-5-IV, V-7-IV short setae, V-6-IV moderately elongate, on short tubercles, V-5-IV, V-6-IV closely approximated, V-7-IV closer to L-4-IV; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Figs. 77 View FIGURE 77 K-L)—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Pachyhelea   is known from two species in the Nearctic and Neotropical Regions ( Borkent   2014). The pupa of the only known species was collected in shallow water among filamentous algae ( Spirogyra majuscula   ).

TAXONOMIC DISCUSSION: One species of Pachyhelea   is known as a pupa ( Tables 2–3 View TABLE 2 ).

MATERIAL EXAMINED: P. pachymera   : 1 pupal exuviae, Arroyo Zapata, Partido de Magdalena , Provincia de Buenos Aires, Argentina, 23-XI-1981 ( MLPA)   ; 1 pupal exuviae, as previous locality, 24-XII-1981 ( MLPA)   ; 1 pupal exuviae, Santa Ana, Entre Rios, Argentina, 9-XI-1984 ( MLPA)   .

Palpomyia Meigen  

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

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 View FIGURE 54 ), apex of the halter extending posteriorly to about 1/6 length of tergite 2 (as in Fig. 33L View FIGURE 33 ), abdominal segment 4 with V-5-IV, V-6-IV and V-7-IV closely approximated ( Fig. 70C View FIGURE 70 ) 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 View FIGURE 69 ) (not as in Fig. 70B View FIGURE 70 ), 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 View FIGURE 22 ), a nearly unique condition found otherwise only in P. flavipes   and P. jonesi   (the latter distinctively with two setae ( Fig. 22J View FIGURE 22 ).

DESCRIPTION: Habitus as in Fig. 12G View FIGURE 12 . 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 View FIGURE 17 , 33K View FIGURE 33 ). Ecdysial tear around base of antenna, along lateral margin of face to palpus ( Figs. 17E View FIGURE 17 , 79H View FIGURE 79 ). Head: Dorsal apotome ( Figs. 22H–J View FIGURE 22 ), with partial ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13H View FIGURE 13 ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 28G View FIGURE 28 ) 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 View FIGURE 41 ) well anterior to posterior to, posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Figs. 22H–J View FIGURE 22 )—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 View FIGURE 28 )—2 slender setae; oculars ( Fig. 28G View FIGURE 28 )—1–2 setae, 1 campaniform sensillum or 1 seta, 2 campaniform sensilla. Thorax: Prothoracic extension ( Fig. 28G View FIGURE 28 ) 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 View FIGURE 54 ); respiratory organ ( Figs. 46N–R View FIGURE 46 ) 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 View FIGURE 41 ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 33K View FIGURE 33 ) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs ( Fig. 41D View FIGURE 41 ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 33K View FIGURE 33 ); 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 View FIGURE 31 ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 31H View FIGURE 31 )—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 View FIGURE 54 )—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 View FIGURE 78 ) not strongly modified, terminal processes closely approximated to separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 54A View FIGURE 54 ) 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 View FIGURE 70 , 71A View FIGURE 71 )—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 View FIGURE 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 View TABLE 2 ). 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 View 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 View FIGURE 17 , 22K View FIGURE 22 , 28H View FIGURE 28 , 31I View FIGURE 31 , 33L View FIGURE 33 , 41E View FIGURE 41 , 46S View FIGURE 46 , 54B View FIGURE 54 , 71B View FIGURE 71 , 78I–K View FIGURE 78 )

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 View FIGURE 78 ); 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 View FIGURE 54 ), apex of the halter extending posteriorly to about 1/6 length of tergite 2 ( Fig. 33L View FIGURE 33 ), abdominal segment 4 with V-5-IV, V-6-IV and V-7-IV closely approximated ( Fig. 71B View FIGURE 71 ), 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 View FIGURE 22 ), a nearly unique condition found otherwise only in P. flavipes   and P. jonesi   ( Fig. 22J View FIGURE 22 ).

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 View FIGURE 17 , 33L View FIGURE 33 ). Ecdysial tear around base of antenna, along lateral margin of face to palpus ( Figs. 17F View FIGURE 17 , 79H View FIGURE 79 ). Head: Dorsal apotome ( Fig. 22K View FIGURE 22 ), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13H View FIGURE 13 ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 28H View FIGURE 28 ) 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 View FIGURE 41 ) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Fig. 22K View FIGURE 22 )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeallabrals ( Fig. 28H View FIGURE 28 )—2 slender setae; oculars ( Fig. 28H View FIGURE 28 )—1 seta, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 28H View FIGURE 28 ) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 54B View FIGURE 54 ); respiratory organ ( Fig. 46S View FIGURE 46 ) 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 View FIGURE 41 ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 33L View FIGURE 33 ) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs ( Fig. 41E View FIGURE 41 ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 33L View FIGURE 33 ); 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 View FIGURE 31 ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 31I View FIGURE 31 )—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 View FIGURE 54 )—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 View FIGURE 78 ) 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 View FIGURE 78 ); sensilla: tergite 1 ( Fig. 54B View FIGURE 54 ) 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 View FIGURE 71 )—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 View FIGURE 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 View TABLE 2 ).

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 View FIGURE 71 ).

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 View FIGURE 12 , 22L View FIGURE 22 , 28I View FIGURE 28 , 31J View FIGURE 31 , 41F View FIGURE 41 , 46T View FIGURE 46 , 54C View FIGURE 54 , 71C View FIGURE 71 , 78L View FIGURE 78 )

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

DESCRIPTION: Habitus (female) as in Fig. 12H View FIGURE 12 . 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 View FIGURE 16 , 33B View FIGURE 33 ). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17C View FIGURE 17 , 79H View FIGURE 79 ). Head: Dorsal apotome ( Fig. 22L View FIGURE 22 ), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13H View FIGURE 13 ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 28I View FIGURE 28 ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna ( Fig. 41F View FIGURE 41 ) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), not narrowed sharply posteriorly; sensilla: dorsal apotomals ( Fig. 22L View FIGURE 22 )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 28I View FIGURE 28 )—2 thick setae; oculars ( Fig. 28I View FIGURE 28 )—1 seta, 1 campaniform sensillum? or possibly merely an indentation? Thorax: Prothoracic extension ( Fig. 28I View FIGURE 28 ) wide, well-developed but narrow dorsolaterally, extending from palpus to antenna; mesonotum yes tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 54C View FIGURE 54 ); respiratory organ ( Fig. 46T View FIGURE 46 ) 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 View FIGURE 41 ) 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 View FIGURE 33 ) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs ( Fig. 41F View FIGURE 41 ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33K View FIGURE 33 ); male with apex of foreleg moderately anterior to apex of midleg (as in Fig. 41E View FIGURE 41 ), 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 View FIGURE 31 ); anterolaterals—1 elongate seta; dorsal setae ( Fig. 31J View FIGURE 31 )—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 View FIGURE 54 )—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 View FIGURE 78 ) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 54C View FIGURE 54 ) 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 View FIGURE 71 )—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 View FIGURE 78 )—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 View TABLE 2 ). 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)   .

Bezzia Kieffer  

( Figs. 12F View FIGURE 12 , 13I View FIGURE 13 , 17D View FIGURE 17 , 22C–D View FIGURE 22 , 28D View FIGURE 28 , 31E View FIGURE 31 , 33J View FIGURE 33 , 41A View FIGURE 41 , 46H–K View FIGURE 46 , 53D View FIGURE 53 , 69C View FIGURE 69 , 77E–H View FIGURE 77 )

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. 53D View FIGURE 53 ), apex of the halter extending posteriorly to about 1/6 length of tergite 2 (as in Fig. 33L View FIGURE 33 ), abdominal segment 4 with V-5-IV, V-6-IV and V-7-IV closely approximated (as in Fig. 70C View FIGURE 70 ) 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 ( Fig. 69C View FIGURE 69 ) (not as in Fig. 70B View FIGURE 70 ), 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 well-developed (not minute), and abdominal segment 8 is without L-1-VIII (not diagnosable as different from Palpomyia   and Phaenobezzia   ); however, most species of Bezzia   have two or more campaniform sensilla on the dorsal apotome ( Figs. 22C–D View FIGURE 22 ), a nearly unique condition found only in two species of Palpomyia   , one of which, P. jonesi   , distinctively has two setae and two campaniform sensilla ( Fig. 22J View FIGURE 22 ).

DESCRIPTION: Habitus as in Fig. 12F View FIGURE 12 . Total length = 2.00– 6.16 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing ( Figs. 17D View FIGURE 17 , 33J View FIGURE 33 ). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17C View FIGURE 17 , 79H View FIGURE 79 ) or with eye appressed to antenna ( Figs. 17D View FIGURE 17 ). Head: Dorsal apotome ( Figs. 22C–D View FIGURE 22 ), with partial ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite ( Fig. 13I View FIGURE 13 ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 28D View FIGURE 28 ) 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. 41A View FIGURE 41 ) anterior to posterior to, posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Figs. 22C–D View FIGURE 22 )—1 moderate to elongate seta, 1–3 campaniform sensilla; dorsolateral cephalic sclerite sensilla—1 seta, 0–1 campaniform sensillum; clypeal-labrals ( Fig. 28D View FIGURE 28 )—1 slender or 2 slender or thick setae; oculars ( Fig. 28D View FIGURE 28 )—1–2 seta, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 28D View FIGURE 28 ) wide, well-developed, extending from palpus to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 53D View FIGURE 53 ); respiratory organ ( Figs. 46H–K View FIGURE 46 ) length/width = 3.73–6.60, moderately elongate to elongate, apical portion swollen in some, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single straight to curved row, outer surface with some wrinkles, with short, wide pedicel, base with short to moderately 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 or more; wing ( Fig. 41A View FIGURE 41 ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 33J View FIGURE 33 ) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs ( Fig. 41A View FIGURE 41 ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 33J View FIGURE 33 ); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally or small gap between the two; sensilla: anteromedials—2 elongate setae, 1 campaniform sensillum (as in Figs. 31L–M View FIGURE 31 ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 31E View FIGURE 31 )—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. 53D View FIGURE 53 )—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: pigmentation light brown, with tergite 1 with 3 medial spots, tergites 2–7 with medial area with stripe, 2 anterolateral spots, sternites 3–7 with medial stripe, anterolateral spot (light brown), 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. 77E–H View FIGURE 77 ) not strongly modified, terminal processes closely approximated to separated basally, each projecting posterodorsolaterally, tapering to pointed apex, in some very slender; sensilla: tergite 1 ( Fig. 53D View FIGURE 53 ) with 8 setae, 2 campaniform sensilla, including 3–4 lateral sensilla, D-2-I, D-3-I closely approximated, D-7-I situated anterolaterally near L-1-I to anteriorly near D-3-I; segment 4 ( Fig. 69C View FIGURE 69 )—D-2-IV, D-3-IV short setae on short tubercles; D-5-IV, D-8-IV, D-9-IV short to moderately elongate setae, D-7-IV present or absent; D-5-IV on single, 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 moderately elongate seta on rounded tubercle, well anterior of posterior lateral setae; L-2- IV, L-3-IV, L-4-IV short to moderately elongate setae on rounded tubercles, L-2-IV, L-3-IV on single tubercle in some, V-5-IV, V-6-IV, V-7-IV short setate 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; with V-5-VIII, V-6-VIII on single tubercle, V-5-VIII tiny, V-6- VIII elongate; segment 9 ( Figs. 77E–H View FIGURE 77 )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Bezzia   is known from 319 species from every Region worldwide ( Borkent 2014). Immatures have been recorded from a wide array of habitats including rice fields, springs, streams, river margins, wet moss, various phytotelmata, pools, marshes, bogs, fens, algal mats in lentic habitats, lakes, and reservoirs.

TAXONOMIC DISCUSSION: The pupae of 51 species of Bezzia   are known ( Tables 2–3 View TABLE 2 ). Goeze (1780) described but did not name a species which is probably a Bezzia   , providing the first morphological study describing the chaetotaxy of the larva and pupa ( Fig. 6B View FIGURE 6 ) of a Ceratopogonidae   . It was later named Tipula goezii   by Schrank (1803). I have placed it in Bezzia   as a new combination as follows but the name is so out of date that it must be considered a nomen nudum. The long slender terminal process and somewhat clavate apex of the respiratory organ ( Figs. 46H, 46J View FIGURE 46 ) is a distinctive combination of many species of Bezzia   .

Bezzia goezii (Schrank)   , 1803: 72 ( Tipula   ). Quedlinburg, Germany. new combination.

Some species have been described more than once by the same author ( Table 2 View TABLE 2 ) but with different names (now synonyms) (e.g. B. circumdata   , B. leucogaster   , B. nobilis   ), suggesting either misidentifications or the possibility of more than one species actually present.

Thomsen (1937) gave a key to three species known to her at that time but one of these, P. flavitarsis   , is now recognized as a Palpomyia   . Thomsen (1937) drew the dorsal apotomes of species of Bezzia   with more than one seta (per side) but likely misinterpreted the multiple campaniform sensilla as the broken bases of setae.

Thienemann (1928) and Mayer (1934a) provided keys to European species groups. Lenz (1934) provided a key to European species but primarily used habitats to distinguish these and, for most, this almost certainly results in inaccurate identifications. Harris (1981) described and keyed the pupae of six unnamed species of Bezzia   from Australia.

Wirth (1983a, 1983b, 1983c) diagnosed the pupae of some Nearctic and Neotropical Bezzia   in the cockerelli   , bicolor   , and nobilis   species groups but did not do so for the remaining species groups of Bezzia   . At present, pupae of Bezzia   cannot be diagnosed as a genus and therefore providing a key to the species in a given region is superfluous. Unless a diagnosis becomes available all Bezzia   should be keyed with species of Palpomyia   and Phaenobezzia   . Alternately, it would be possible to key all species with more than one campaniform sensillum on the dorsal apotome, which would include nearly all Bezzia   and a few Palpomyia   , depending on the area being covered.

Nearly all Bezzia   have more than one campaniform sensillum on the dorsal apotome (see character 6). The only exceptions seen are of B. brevicornis   , B. bromeliae   and B. dorsasetula   , each with one seta and one campaniform sensillum.

Bezzia xanthogaster (Kieffer)   , 1919: 130 ( Probezzia   ) is a junior homonym of Probezzia xanthogaster (Kieffer)   , 1917: 329 ( Bezzia   ) and is here given the new name Bezzia gilvigaster   (a similar specific name also meaning yellow stomach (abdomen for the adult this species).

MATERIAL EXAMINED: B. africana   : 1 pupal exuviae, Burgershall , Hazyview, Transvaal, South Africa, 3- XII-1973 ( NMSA)   ; 1 pupal exuviae, Magaliesberg Agricultural School , Transvaal, South Africa, 13-XI-1973 ( NMSA)   . B. albicornis   : 1 pupal exuviae, Kaushut , Tedjen Province, Turkmenistan, 2-IV-1972 ( ZIN)   ; 1 pupal exuviae, as previous locality, 30-V-1972 ( ZIN)   . B. amana   : 1 pupal exuviae (of holotype), Loutpan , Transvaal, South Africa, 10-I-1974 ( NMSA)   . B. annulipes   : 2 pupal exuviae, Lunzer Untersee between boat house and canal, Lower Austria, Austria, 6-VI-1942 ( ZSMC)   ; 1 pupal exuviae, Germany ( ZSMC)   ; 3 pupal exuviae, Germany ( ZSMC)   ; 12 pupal exuviae, Anninskoe lake , Pskov Province, Russia, 24-VI-1995 ( ZIN)   . B. biannulata   : 1 pupal exuviae (of paratype), 1 pupal exuviae (of allotype), Oceano Beach, San Luis Obispo County, California, USA, 20- VIII-1948 ( USNM)   . B. bicolor   : 6 pupal exuviae, Gievenbeck , Germany, 9-V-1912 ( ZSMC)   ; 4 pupal exuviae, Lunz , Austria ( USNM)   ; 4 pupal exuviae, Shushary , Leningrad Province, Russia, 29-V-1997 ( ZIN)   ; 2 pupal exuviae, Strelna , Leningrad Province, Russia, 12-V-2002 ( ZIN)   . B. bivittata   : 3 pupal exuviae, Salisbury , Wicomico County, Maryland, USA, 4-V-1981 ( WLGC)   ; 1 pupal exuviae, as previous locality, 13-IV-1981 ( WLGC)   . B. blantoni   : 2 pupal exuviae (of paratype), Escobar , Buenos Aires, Argentina, 10-I-1982 ( MPLA)   . B. brevicornis   : 1 pupal exuviae, Magdalena , Buenos Aires, Argentina, 24-XII-1981 ( MPLA)   . B. bromeliae   : 2 pupal exuviae (of paratype), Bayano Field Station , Panama Province, Panama, VI-1976 ( MPLA)   . B. circumdata   : 3 pupal exuviae, Black Lake , Stanleyville, Ontario, Canada, 24-VI-1975 ( USNM)   ; 1 pupal exuviae, as previous locality, 25-VI-1975 ( USNM)   ; 4 pupal exuviae, Rotmoos on Mittersee , Lower Austria, Austria, 10-VI-1942 ( ZSMC)   ; 1 pupa, 5 pupal exuviae, Grosser Plöner See , Slesvig-Holstein, Germany, summer 1952 ( ZSMC)   ; 2 pupa, 5 pupal exuviae, kerosene port on Dortmund-Ems canal, Dortmund, North Rhine-Westphalia, Germany, 28-VII-1908 ( ZSMC)   ; 1 pupa, 3 pupal exuviae, Germany ( ZSMC)   ; 4 pupal exuviae, Vyborg , Leningrad Province, Russia, 27-VII-1998 ( ZIN)   . B. cockerelli   : 1 pupal exuviae, Valdez , Alaska, USA, 1948 ( USNM)   ; 1 pupal exuviae, Bainville , Roosevelt County, Montana, USA, 9-VI-1969 ( USNM)   ; 1 pupal exuviae, Trails Pond , Latah County, Idaho, USA, 22-VII-1969 ( USNM)   ; 5 pupal exuviae, 4 mi N of Upham , McHenry County, North Dakota, USA, 5-VI-1969 (4 USNM, 1 VPIC)   . B. collessi   : 1 pupal exuviae (of paratype), Singapore, 10-IX-1952 ( USNM)   ; 3 pupal exuviae (of paratypes), as previous locality, 28-VIII-1952 ( USNM)   . B. dorsasetula   : 2 pupal exuviae, College Park , Prince George’s County, Maryland, USA, 30-V-1975 ( USNM)   , 1 pupal exuviae, no locality, 6-V-1977 ( VPIC)   . B. fascispinosa   : 1 pupal exuviae, Lakeland Pond , College Park, Prince George’s County, Maryland, USA, 30-V-1975 ( USNM)   ; 1 pupal exuviae, Beaver Lake Reservoir, Pocahontas State Park, Chesterfield County, Virginia, USA, 30-IV-1977 ( VPIC)   ; 1 pupal exuviae, Iron Spring , Elliot Know, Augusta County, Virginia, USA, 22-17-1977 ( VPIC)   . B. flavicornis   : 1 pupal exuviae, Priluki , Belarus, 17-V-1967 ( ZIN)   . B. flavicorporis   : 1 pupal exuviae, Loutpan , Transvaal, South Africa, 10-I-1974 ( NMSA)   . B. gibbera   : 1 pupal exuviae, Dismal Swamp , Camden County, North Carolina, USA, 25-III-1976 ( VPIC)   . B. glabra   : 2 pupal exuviae, Morgan Arboretum , St. Anne de Bellevue, Quebec, Canada, 1964 ( USNM)   ; 1 pupal exuviae, 1 mi. S of Corapeake, Dismal Swamp , Camden County, North Carolina, USA, 29-VII-1976 ( VPIC)   ; 2 pupal exuviae, Keowee Reserve , Seneca , Oconee County, South Colorado, USA, 2-V-1975 ( VPIC)   . B. japonica   : 1 pupal exuviae, Suputinka river , Ussuri Nature Reserve, Primorskii Territory, Russia, 30-V-1973 ( ZIN)   . B. kuhetiensis   : 1 pupal exuviae, Issyk-Kul lake , Kyrgyzstan, 16-VI-1971 ( ZIN)   . B. laciniastyla   : 1 pupal exuviae, Jackson River , approx. 10 mi. upriver from Covington, Natural Well, Allegheny County, Virginia, USA, 4-VI-1977 ( VPIC)   . B. leucogaster   : 4 pupal exuviae, shore of Lottsee SE of Mölln, Slesvig-Holstein, Germany, 16-V-1926 ( ZSMC)   , 2 pupal exuviae, shore of Kirchsee , Preetz (between Kiel and Plön), Slesvig-Holstein, Germany, 11-VII1918 ( ZSMC)   ; 1 pupal exuviae, Severskii Donets River , Donetsk Province, Ukraine, 5-V-1970 ( ZIN)   . B. narynica   : 1 pupal exuviae, Issyk-Kul lake , Kyrgyzstan, 24-V-1971 ( ZIN)   . B. nigrita   : 1 pupal exuviae, Severskii Donets River , Donetsk Province, Ukraine, 5-V-1970 ( ZIN)   ; 1 pupal exuviae, Malinovka , Donetsk Province, Ukraine, 8-V-1970 ( ZIN)   . B. nobilis   : 1 pupal exuviae, 7 km S of Hope , BC, Canada, 8–9-VII-1985 ( CNCI)   ; 1 pupal exuviae, Beaver Creek , Lawrence County, South Dakota, USA, 15-VI-1969 ( USNM)   ; 1 pupal exuviae, Cow Neck Salt Marsh , North Sea, New York, USA, 19-IV-1956 ( NYSM)   ; 1 pupal exuviae, as previous locality, 17-IV-1956 ( NYSM)   ; 4 pupal exuviae, Fishing Creek , Newcomb, New York, USA, 28-V-1958 ( NYSM)   ; 1 pupal exuviae, Port Leyden , New York, USA, 4-VII-1959 ( USNM)   ; 2 pupal exuviae, Letchworth State Park , New York, USA, 13-VI-1963 ( USNM)   ; 1 pupal exuviae, Patuxent Refuge , Maryland, USA, 8-V-1958 ( USNM)   ; 1 pupal exuviae, Vepco Property , Louisa County, Virginia, USA, 12-IX-1976 ( VPIC)   ; 1 pupal exuviae, Washburn County, Wisconsin, USA, 1-IX-1951 ( VPIC)   ; 2 pupal exuviae, Vepco Property , Louisa County, Virginia, USA, 12-IX-1976 ( VPIC)   ; 2 pupal exuviae, Great Swamp , Isle of Wight County, Virginia, USA, 2-VIII- 1976 ( VPIC)   ; 1 pupal exuviae, Cranberry Glade , Pocahontas County, Virginia, USA, 8-IX-1976 ( VPIC)   ; 1 pupal exuviae, Norfolk Gardens , Norfolk County, Virginia, USA, 1-VIII-1976 ( VPIC)   . 1 pupal exuviae, no locality/date ( VPIC); 1 pupal exuviae, Pecan Springs , Devils River, Juno, Texas, USA ( USNM)   ; 1 pupal exuviae, Baton Rouge , Louisiana, USA, 4-III-1947 ( USNM)   ; 1 pupal exuviae, Beloe lake , Alol’, Pskov Province, Russia, 27-VI-1969 ( ZIN)   ; 1 pupal exuviae, Beloe lake , Alol’, Pskov Province, Russia, 5-VII-1969 ( ZIN)   . B. obelisca   : 2 pupal exuviae, Mer Bleue , Ottawa, Ontario, Canada, 27-V-1960 ( USNM)   ; 2 pupal exuviae, McLean Reserve , Tompkins County, New York, USA, 18-VI-1963 ( USNM)   ; 1 pupal exuviae, edge of Big Glade , Cranberry Glades Botanical Area, Pocahontas County, Virginia, USA, 6-IX-1976 ( VPIC)   . B. perplexa   : 1 pupal exuviae, Millpond Creek , North Anna Resrvoir, Louisa County, Virginia, USA, 1-III-1977 ( VPIC)   . B. pulchripes   : 2 pupal exuviae, Mission Rio Cururu Igarape , Brazil, 4-V-1941 ( SDEI)   ; 1 pupal exuviae, Mocoreta , Corrientes, Argentina, 21-III-1985 ( MPLA)   . B. roldani   : 2 pupal exuviae, no locality, 20-VIII-1979 ( MPLA)   . B. saileri   : 1 pupal exuviae (of paratype), Fire Lake , Anchorage, Alaska, USA, 29-V-1948 ( USNM)   . B. signata   : 1 pupal exuviae, Luchevoi , Republic of Karelia, Russia, 14-VI-1966 ( ZIN)   . B. sordida   : 1 pupal exuviae, Olerna , Marin County, California, USA, 22-II-1947 ( USNM)   . B. turkmenica   : 1 pupal exuviae, Hodzhaz-Kala bay , Kyzyl-Arvat Province, Turkmenistan, 18-IV-1972 ( ZIN)   . B. uncistyla   : 2 pupal exuviae, 5 km E of Danby , Vermont, USA, 25-VI-1986 ( CNCI)   ; 1 pupal exuviae, as previous locality, 25–26-VI-1986 ( CNCI)   . B. varicolor   : 1 pupal exuviae, Rideau River , Ottawa, Ontario, Canada, 29-V-1960 ( USNM)   ; 1 pupal exuviae, Lake Temescal , Berkeley, California, USA, 1-V-1948 ( USNM)   ; 1 pupal exuviae, Lake Tomahawk at Kemp Biological Station, Oneida County, Wisconsin, USA, 20-VII-1978 ( USNM)   ; 2 pupal exuviae, Bar Lake , Benzie County, Michigan, USA, 8-VIII-1975 ( VPIC)   ; 1 pupal exuviae (of topotype), Mill Creek Brook, Noyack, New York, USA, 6-V-1057 ( USNM)   ; 1 pupal exuviae, Slaterville Spr. , New York, USA, 12- VI-1964 ( VPIC)   ; 1 pupal exuviae, Lakeland Pond , College Park, Prince George’s County, Maryland, USA, 24-IV- 1977 ( VPIC)   . B. nr. bicolor   : 1 pupal exuviae, no locality, 30-V-1953 ( USNM)   . B. nr. nobilis   : 1 pupal exuviae, Back Bay National Wildlife Refuge , Virginia Beach, Virginia, USA, 30-VIII-1975 ( VPIC)   . B. nr. obelisca   : 1 pupal exuviae, Mer Bleue , Ottawa, Ontario, Canada, 27-V-1960 ( USNM)   ; 2 pupal exuviae, no data ( USNM); 1 pupal exuviae, Mer Bleue , Ottawa, Ontario, Canada, 27-V-1960 ( VPIC)   ; 1 pupal exuviae, Big Glade , Cranberry Glade Botanical Area, Pocahontas County, Virginia, USA, 8-IX-1976 ( VPIC)   . B. nr. pruinosa   : 1 pupal exuviae, no locality, 12-V-1953 ( USNM)   . B. nr. varicolor   : 1 pupal exuviae, W flank of Salt Pond Mountain , Giles County, Virginia, USA, 14-VII-1977 ( VPIC)   . B. sp.: 2 pupal exuviae (in glycerin), 12 pupal exuviae, Bolean Lake , 6 km NE of Falkland, British Columbia, Canada, 12-13-VII-1989 ( CNCI)   ; 1 pupal exuviae (in glycerin), Spanish Lake , 6 km E of Falkland, 50°29.12N 119°28.07W, British Columbia, Canada, 27–28-V-2008 ( CNCI) GoogleMaps   ; 4 pupal exuviae, 3 km E of Salmon Arm , British Columbia, Canada, 9-VI-1988 ( CNCI)   ; 9 pupal exuviae, 6 km E of Salmon Arm , British Columbia, Canada, 6-VI-1990 ( CNCI)   ; 5 pupal exuviae, 9 km S of Salmon Arm , British Columbia, Canada, 19-VII-1988 ( CNCI)   ; 2 pupal exuviae, 8 km E of Sicamous , British Columbia, Canada, 1-VI-1992 ( CNCI)   ; 2 pupal exuviae, 6.5 km NW of Enderby , British Columbia, Canada, 6-V-1992 ( CNCI)   ; 4 pupal exuviae, 30 km N of Nakusp , British Columbia, Canada, 5-VII-1990 ( CNCI)   ; 4 pupal exuviae, 20 km E of Anola , Manitoba, Canada, 16-VI-1990 ( CNCI)   ; 3 pupal exuviae, Trail Bay , Manitoba, Canada ( CNCI)   ; 1 pupal exuviae (in glycerin), 1 mi. E of Orleans , Ontario, Canada, 8-VII-1971 ( CNCI)   ; 1 pupal exuviae, 6 km N of Eardlay , Quebec, Canada, 15-V-1986 ( CNCI)   ; 2 pupal exuviae, Fire Lake , Anchorage, Alaska, USA, 29-V-1948 ( USNM)   ; 5 pupal exuviae, 9 km W of Okanogan , Washington, USA, 13-VI-2008 ( CNCI)   ; 9 pupal exuviae, High Creek Fen Preserve , 9 mi S of Fairplay Park, Colorado, USA, 18-VI-1995 ( CNCI)   ; 1 pupal exuviae, Blue Mountain Lake , New York, USA, 30-VI-1958 ( WLGC)   ; 2 pupal exuviae, Ringwood Pond , Dryden, New York, USA, 7-III-1933 ( USNM)   ; 1 pupal exuviae, Keowee Reserve , Seneca, Ocouee County, South Carolina, USA, 16-V-1974 ( USNM)   ; 1 pupal exuviae, as previous locality, USA, 16-VII-1974 ( USNM)   ; 1 pupal exuviae, 8 km W of Atenas , Costa Rica, 19-VIII-1993 ( CNCI)   ; 2 pupal exuviae, carreterra entre Santa Cecilia y Upala, 1.6 km N.O. de BVirmania, Dos Rios, Upala , Alejuela, Costa Rica, 16-X-2006 ( INBC)   ; 1 pupal exuviae, Barro Colorado Island , Canal Zone, Panama, 24-VI- 1996 ( CNCI)   ; 2 pupal exuviae, as previous locality, 31-X-1995 ( CNCI)   ; 1 pupal exuviae, as previous locality, 30- VI-1996 ( CNCI)   ; 1 pupal exuviae, as previous locality, 10-IX-1995 ( CNCI)   ; 3 pupae, Lerchenauer See , Germany, 26-VII-1990, 27-VII-1990, 2-VIII-22-IX-1990 ( CNCI)   ; 1 pupa, Chiemsee , Germany, 13-VIII-1990   ; 1 pupa, 1 pupal exuviae, Bottsand nr. Stein, Germany, 5-VI-1932 ( ZSMC)   ; 2 pupal exuviae, Darban , Western Australia, Australia, 28-X-1986 ( ANIC)   ; 9 pupal exuviae, as previous locality, 28-X-1985 ( ANIC)   ; 2 pupal exuviae, 5 km N of Darban , Western Australia, Australia, 29-X-1985 ( ANIC)   ; 3 pupal exuviae, Roper River Mission, Northern Territory, Australia, 8-XI-1956 ( ANIC)   ; 3 pupal exuviae, Humpty Doo , Northern Territory, Australia, V-1957 ( ANIC)   ; 1 pupal exuviae, Dayboro , Queensland, Australia, 10-VII-1951 ( ANIC)   ; 10 pupal exuviae, Nattai River , Mittagong, New South Wales, Australia, 4-XI-1964 ( ANIC)   ; 1 pupal exuviae, as previous locality, 15-XI-1968 ( ANIC)   ; 1 pupal exuviae, Gap Creek , The Crags, Mittagong, New South Wales, Australia, 10-II-1966 ( ANIC)   ; 5 pupal exuviae, Damin clearing, The Crags, Mittagong, New South Wales, Australia, 9-II-1966 ( ANIC)   ; 2 pupal exuviae, Deep Creek , Nanasheen, New South Wales, Australia, 7-XII-1956 ( ANIC)   ; 2 pupal exuviae, as previous locality, 25-XI-1956 ( ANIC)   ; 21 pupal exuviae, Middle Creek , New South Wales, Australia, 8-II-1966 ( ANIC)   ; 1 pupal exuviae, Colo Vale , New South Wales, Australia, 16-X-1956 ( ANIC)   ; 2 pupal exuviae, Hornsby , New South Wales, Australia, 18-XI-1969 ( ANIC)   ; 2 pupal exuviae, as previous locality, 25-X-1956 ( ANIC)   ; 2 pupal exuviae, Nepean River , Menangle, New South Wales, Australia, 9-XII-1968 ( ANIC)   ; 1 pupal exuviae, Griffith , New South Wales, Australia, 16-XI-1956 ( ANIC)   ; 1 pupal exuviae, Moruya River , Merricumbene Creek, New South Wales, Australia, 2-III-1964 ( ANIC)   ; 1 pupal exuviae, McCarrs Creek , New South Wales, Australia, 14-I-1969 ( ANIC)   ; 1 pupal exuviae, no locality, 14-I-1937 ( SAIM)   .

Clastrieromyia Spinelli & Grogan  

( Figs. 22F View FIGURE 22 , 28E View FIGURE 28 , 31F View FIGURE 31 , 41B View FIGURE 41 , 46L View FIGURE 46 , 53E View FIGURE 53 , 70A View FIGURE 70 , 77I–J View FIGURE 77 )

DIAGNOSIS: Only pupa of Ceratopogonidae   with the metathorax with a single campaniform sensillum (M-3-T) situated at least ⅓ the length of the metathorax from its anterior margin ( Fig. 53E View FIGURE 53 ), apex of the halter extending posteriorly to about 1/6 length of tergite 2 (as in Fig. 33L View FIGURE 33 ), D-7-I near D-3-I ( Fig. 53E View FIGURE 53 ), abdominal segment 4 with each setae on a rounded tubercle ( Fig. 70A View FIGURE 70 ) and abdominal segment 8 without L-1-VIII.

DESCRIPTION: Total length = 3.20–3.80 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16B View FIGURE 16 , 33B View FIGURE 33 ). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17C View FIGURE 17 , 79H View FIGURE 79 ). Head: Dorsal apotome ( Fig. 22F View FIGURE 22 ), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13H View FIGURE 13 ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 28E View FIGURE 28 ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna ( Fig. 41B View FIGURE 41 ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Fig. 22F View FIGURE 22 )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum on very short tubercle; clypeal-labrals ( Fig. 28E View FIGURE 28 )—2 slender setae; oculars ( Fig. 28E View FIGURE 28 )—2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 28E View FIGURE 28 ) wide, well-developed but very narrow dorsolaterally, extending from palpus to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 53E View FIGURE 53 ); respiratory organ ( Fig. 46L View FIGURE 46 ) length/width = 3.13–4.08, elongate, moderately slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single 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. 41B View FIGURE 41 ) with short tubercle at apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33J View FIGURE 33 ) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs ( Fig. 41B View FIGURE 41 ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33J View FIGURE 33 ); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials—1 long, 1 medium setae, 1 campaniform sensillum; anterolaterals—1 moderately long seta; dorsal setae ( Fig. 31F View FIGURE 31 )—D-1-T, D-2-T, D-4-T, D- 5-T setae, D-3-T campaniform sensillum, D-3-T anterior to D-4-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 53E View FIGURE 53 )—1 campaniform sensillum; M-3-T distant from margin of metathorax (at least 1/3 length of metathorax). Abdomen: pigmentation light brown, with tergite 1 with 3 medial spots, tergites 2-7 with medial area with stripe, 2 anterolateral spots, sternites 3-7 with medial stripe, anterolateral spot (light brown), segment 2 as wide or slightly wider than segment 3, segments with undivided, peg-like or thin to thick setae, with rounded, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Fig. 77 View FIGURE 77 I-J) not strongly modified, terminal processes separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 53E View FIGURE 53 ) with 7 setae, 2 campaniform sensilla, including 3 lateral sensilla, D-2-I, D-3-I closely approximated, D-7-I situated anteriorly near D-3-I; segment 4 ( Fig. 70A View FIGURE 70 )—D-2-IV, D-3-IV short to moderately elongate setae on short tubercles; D-5-IV peg-like seta, D-8-IV, D-9-IV short to moderately elongate setae; D-5-IV on single tubercle, D-8-IV, D-9-IV on 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 near D-3-IV; L- 1-IV short seta on rounded tubercle, well anterior of posterior lateral setae; L-2-IV, L-4-IV short setae, L-3-IV moderately elongate seta on rounded tubercles, V-5-IV, V-6-IV short setae, V-7-IV moderately elongate, on moderately elongate tubercles, V-5-IV, V-6-IV moderately closely approximated; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Fig. 77I–J View FIGURE 77 )—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Clastrieromyia   is known from four species in the Neotropical Region ( Borkent   2014). Immatures of the only known species have been collected from a cattle-trodden bog.

TAXONOMIC DISCUSSION: Only one species is known as a pupa ( Tables 2–3 View TABLE 2 ).

MATERIAL EXAMINED: C. dycei   : 2 pupal exuviae (of paratypes), Tacuarembo, Uruguay, 29-IX-1980 ( MLPA)   .

Pachyhelea Wirth  

( Figs. 22G View FIGURE 22 , 28F View FIGURE 28 , 31G View FIGURE 31 , 41C View FIGURE 41 , 46M View FIGURE 46 , 53F View FIGURE 53 , 70B View FIGURE 70 , 77K–L View FIGURE 77 )

DIAGNOSIS: Only pupa of Ceratopogonidae   with dorsal apotome with only one campaniform sensillum ( Fig. 22G View FIGURE 22 ), metathorax with only one campaniform sensillum (M-3-T) situated at least ⅓ the length of the metathorax from its anterior margin ( Fig. 53F View FIGURE 53 ), apex of the halter extending posteriorly to about 1/6 length of tergite 2 (as in Fig. 33L View FIGURE 33 ), L-1-IV and L-3-IV short setae on separate tubercles, L-2-IV well separated from L-3-IV, V-7-IV is closer to L-4-IV than to V-5-IV and V-6-IV, which are closely abutting ( Fig. 70B View FIGURE 70 ).

DESCRIPTION: Total length = 7.22 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16B View FIGURE 16 , 33B View FIGURE 33 ). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17C View FIGURE 17 , 79H View FIGURE 79 ). Head: Dorsal apotome ( Fig. 22G View FIGURE 22 ), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13H View FIGURE 13 ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 28F View FIGURE 28 ) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna ( Fig. 41C View FIGURE 41 ) barely anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Fig. 22G View FIGURE 22 )—1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—1 seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 28F View FIGURE 28 )—2 slender setae; oculars ( Fig. 28F View FIGURE 28 )—1 seta, 1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 28F View FIGURE 28 ) wide, well-developed but very narrow dorsolaterally, extending from palpus to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially ( Fig. 53F View FIGURE 53 ); respiratory organ ( Fig. 46M View FIGURE 46 ) length/width = 3.96–5.00, elongate, moderately slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single 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, wrinkles to half length; wing ( Fig. 41C View FIGURE 41 ) with short tubercle at apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33J View FIGURE 33 ) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs ( Fig. 41C View FIGURE 41 ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33J View FIGURE 33 ); 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 View FIGURE 31 ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 31G View FIGURE 31 )—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. 53F View FIGURE 53 )—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, 2 anterolateral spots, 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 tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Figs. 77K–L View FIGURE 77 ) not strongly modified, terminal processes separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 53F View FIGURE 53 ) with 7 setae, 3 campaniform sensilla, including 3 lateral sensilla, D-2- I, D-3-I closely approximated, D-7-I situated anteriorly near D-3-I; segment 4 ( Fig. 70B View FIGURE 70 )—D-2-IV, D-3-IV short to moderately elongate setae, D-2-IV on short tubercle, D-5-IV very short seta, D-8-IV, D-9-IV short setae; D-5-IV without tubercle, D-8-IV, D-9-IV on 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 near D-3-IV; L-1-IV short seta on rounded tubercle, near L-3-IV; L-2-IV, L-3-IV, L-4-IV short setae, L-2-IV, L-3-IV on very short tubercles, L-4-IV on pointed, elongate tubercle, V-5-IV, V-7-IV short setae, V-6-IV moderately elongate, on short tubercles, V-5-IV, V-6-IV closely approximated, V-7-IV closer to L-4-IV; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Figs. 77 View FIGURE 77 K-L)—with D-5-IX, D-6-IX campaniform sensilla.

DISTRIBUTION AND HABITAT: The genus Pachyhelea   is known from two species in the Nearctic and Neotropical Regions ( Borkent   2014). The pupa of the only known species was collected in shallow water among filamentous algae ( Spirogyra majuscula   ).

TAXONOMIC DISCUSSION: One species of Pachyhelea   is known as a pupa ( Tables 2–3 View TABLE 2 ).

MATERIAL EXAMINED: P. pachymera   : 1 pupal exuviae, Arroyo Zapata, Partido de Magdalena , Provincia de Buenos Aires, Argentina, 23-XI-1981 ( MLPA)   ; 1 pupal exuviae, as previous locality, 24-XII-1981 ( MLPA)   ; 1 pupal exuviae, Santa Ana, Entre Rios, Argentina, 9-XI-1984 ( MLPA)   .

Palpomyia Meigen  

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

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 View FIGURE 54 ), apex of the halter extending posteriorly to about 1/6 length of tergite 2 (as in Fig. 33L View FIGURE 33 ), abdominal segment 4 with V-5-IV, V-6-IV and V-7-IV closely approximated ( Fig. 70C View FIGURE 70 ) 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 View FIGURE 69 ) (not as in Fig. 70B View FIGURE 70 ), 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 View FIGURE 22 ), a nearly unique condition found otherwise only in P. flavipes   and P. jonesi   (the latter distinctively with two setae ( Fig. 22J View FIGURE 22 ).

DESCRIPTION: Habitus as in Fig. 12G View FIGURE 12 . 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 View FIGURE 17 , 33K View FIGURE 33 ). Ecdysial tear around base of antenna, along lateral margin of face to palpus ( Figs. 17E View FIGURE 17 , 79H View FIGURE 79 ). Head: Dorsal apotome ( Figs. 22H–J View FIGURE 22 ), with partial ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13H View FIGURE 13 ) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Fig. 28G View FIGURE 28 ) 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 View FIGURE 41 ) well anterior to posterior to, posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals ( Figs. 22H–J View FIGURE 22 )—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 View FIGURE 28 )—2 slender setae; oculars ( Fig. 28G View FIGURE 28 )—1–2 setae, 1 campaniform sensillum or 1 seta, 2 campaniform sensilla. Thorax: Prothoracic extension ( Fig. 28G View FIGURE 28 ) 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 View FIGURE 54 ); respiratory organ ( Figs. 46N–R View FIGURE 46 ) 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 View FIGURE 41 ) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 33K View FIGURE 33 ) broadly abutting; halter apex extending posteriorly to 1/6 length of tergite 2; legs ( Fig. 41D View FIGURE 41 ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Fig. 33K View FIGURE 33 ); 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 View FIGURE 31 ); anterolaterals—1 moderately long seta; dorsal setae ( Fig. 31H View FIGURE 31 )—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 View FIGURE 54 )—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 View FIGURE 78 ) not strongly modified, terminal processes closely approximated to separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 ( Fig. 54A View FIGURE 54 ) 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 View FIGURE 70 , 71A View FIGURE 71 )—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 View FIGURE 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 View TABLE 2 ). 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 View 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