Dasyhelea Kieffer

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 : 45-47

Dasyhelea Kieffer

( Figs. 11G , 14F , 18F–L , 23F–G , 29H–J , 32E–F , 34E , 43A–G , 47F , 56C , 72K–L , 73A–G )

DIAGNOSIS: Only pupa of Ceratopogonidae with the labium clearly split medially by an elongate suture (or overlapping halves) ( Figs. 23F–G ); most species have uniquely complex terminal processes each of which is bifid or forked and bearing two setae; also most species have the posterior sensilla of abdominal segments 3–8 arranged on broad shelf-like tubercles, forming a partial and unique ring around each segment.

DESCRIPTION: Habitus as in Fig. 11G . Total length = 1.63–4.16 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face ( Fig. 14F ). Ecdysial tear not extending at all ( Figs. 14F , 79C ); posterolateral portion of eye. Head: Dorsal apotome ( Figs. 18F–L ), with ventral line of weakness, with or without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13B ) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts ( Figs. 23F–G ) with mandible, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium divided medially by longitudinal suture or two halves slightly overlapping; apex of antenna ( Fig. 34E ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals ( Figs. 18F–L )—1 short to elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla—2 small setae, 1 campaniform sensillum; clypeal-labrals ( Figs. 23F–G )—2 minute setae; oculars ( Figs. 23F–G )—0–1 seta, 1 campaniform sensillum. Thorax: Prothoracic extension ( Figs. 23F–G ) absent; mesonotum with or without short tubercles, not extending posteromedially to extending posteromedially, completely dividing metathorax medially ( Fig. 47F ); respiratory organ ( Figs. 43A–G ) length/ width = 2.81–29.14, highly variable, moderately to very elongate, apex pointed to truncate, somewhat flattened or circular in cross-section, with pores closely abutting or slightly separated at apex of respiratory organ, arranged in single line of varying shapes, with or without additional, more basal pores, outer surface with or without annulations, with or without spicules, without pedicel, base with short, strung out posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, smooth or with spirals restricted to base; wing ( Fig. 34E ) with apical tubercle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg ( Figs. 32E–F ) just separate; halter apex abutting anterolateral knob-like extension of tergite 2; legs ( Fig. 34E ) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing ( Figs. 32E–F ); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg slightly dorsal to, partially abutting apex of midleg laterally; sensilla: anteromedials—2 setae; anterolaterals—1 seta, 1 campaniform sensillum; dorsal setae ( Figs. 29H–J )—D-1-T, D-2-T seta, D-3-T campaniform sensillum, D-1-T well anterior of D-2-T; supraalar 2—campaniform sensillum; metathoracics ( Fig. 47F )—2 campaniform sensilla; M-3-T near anterior margin of metathorax. Abdomen: without pigmentation pattern or some with two light brown pigment spots near middle of tergites 1–7, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to pointed, short, wide, shelf-like tubercles, rarely with elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 ( Figs. 72K–L , 73A–G ) highly variable in shape, terminal processes widely separated to closely approximated basally, each projecting posteriorly to laterally, generally shelf-like with two apices or undivided but modified and tapering to pointed apex, in some elongate and slender; sensilla: tergite 1 ( Fig. 47F ) with 4 setae, 2 campaniform sensilla, including 3 lateral sensilla, D-2-I well posterior from anterior margin of tergite 1, D-3-I absent, D-7-I situated posteriorly; segment 4 ( Fig. 56C )—D-2-IV short to elongate seta, without tubercle, D-3-IV absent; D-8-IV short to (rarely) elongate seta, D-5-IV, D-9-IV absent; D-8-IV without or on wide, low tubercle, D-4-IV-D-7-IV on wide, low, slightly separate tubercles, rarely all on elongate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D-4-IV, D-8-IV, D-7-IV; L-1-IV short to moderately elongate seta, in transverse row with other lateral setae, arranged L-2-IV, L-1-IV, L-3-IV, L-4-IV; L-2-IV, L-3-IV, L-4-IV short setae on low, shelflike tubercles or (rarely) on elongate tubercles, V-6-IV, V-7-IV short setae on wide, low tubercles or (rarely) on elongate tubercles, V-5-IV absent; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Figs. 72K–L , 73A–G )—with D-5-IX, D-6-IX campaniform sensilla, V-1-IX, V-2-IX setae, in some one seta or one campaniform sensillum likely absent.

DISTRIBUTION AND HABITAT: The genus Dasyhelea is known from 609 species from every Region worldwide ( Borkent 2014, Díaz, et al. 2014). Immatures are known from a wide array of small aquatic habitats, primarily from algal growth on damp soil or other wet substrates, wet moss, algae or moss at the margins of standing or running water and amongst floating vegetation (including algal mats). As one would expect with a species rich genus, immatures occur in a wide array of other microhabitats but all of these are small aquatic (or soaking wet) habitats such as from Pistia , wet decomposing vegetation, ant refuse piles (of Atta ), mining floating leaves of Salvinia , sap from tree wounds, wet woodland leaf packs, small ground or rock pools, phytotelmata, margins of extreme saline pools, hot springs, intertidal zone, marine splash-zone, salt marshes, small artificial habitats such as rain gutters, and attacking tobacco seedlings grown hydroponically. Larvae appear to require water on their bodies to survive, while pupae either remain immersed in media or crawl upward to drier substrate.

Münchberg (1961) studied the function of the respiratory organs of D. flavifrons (as D. versicolor ) under experimental conditions, removing either one or both of the respiratory organs or applying sealant to the pores. His questionable approach resulted in death for those with both respiratory organs removed and very little survivorship in those with one removed. He suggested that some respiration may occur through the terminal processes but this is clearly not possible (there is no opening posteriorly). Some young pupae could remain submerged for substantial periods of time indicating they likely extract oxygen from the surrounding water.

TAXONOMIC DISCUSSION: There are 90 species of Dasyhelea known as pupae ( Tables 2–3 ) and there is a high level of morphological differentiation between many of them. Clearly, there are numerous synapomorphies waiting to be interpreted. Furthermore, this genus is extremely species rich, with many undescribed species, and pupal morphology will certainly assist in interpreting these.

Even though there are a goodly number of named species with described pupae, it is clear that the identity of some of them is uncertain. For example, Thienemann (1925) recognized the pupae of D. flavifrons and five other species ( D. brevitibialis , D. lignicola , D. obscura , D. sensualis , D. versicolor ) now all considered to be one species (the latter as synonyms of D. flavifrons ). Similarly, Lenz (1934) noted differences between the pupae of D. flavifrons and some of these same taxa. Some of the differences noted are likely valid indicators of species differences, although it is now uncertain what adults were actually reared and associated. Mayer (1934a) also describes differences between species now considered synonyms (e.g. D. modesta with its synonyms D. inclusa and D. longipalpis ).

Mayer (1934a) and Lenz (1934) provided a key to the 19 European species known at that time and Thienemann (1925) has a key to some species from northern Germany. The pupae of 11 Nearctic species were described and illustrated by Waugh & Wirth (1976), showing significant differences between them but they did not provide a key. The campaniform sensilla DA-2-H is present in these species but were not illustrated. Mayer (1934c) described and keyed 13 species from Indonesia indicating and illustrating detailed differences between their dorsal apotomes, respiratory organs, and the shape and chaetotaxy of segment 9. Mayer (1934c) showed the dorsal apotome of D. insons with DA-2-H as a seta (so there are two setae on a single tubercle), which would be unique within the genus. All other species I studied had one seta and one campaniform sensillum. Wirth & Beaver (1979) described and keyed the unusual pupae of five species of Dasyhelea found in Nepenthes (monkey cup) in southeast Asia.

Fossil pupae of six species were described (or redescribed) by Pierce (1966). Although the specimens were not reexamined here, based on the form of the terminal processes (divided), they are clearly correctly assigned to genus. Grimaldi & Engel (2005:45) include a photograph of two silicified pupae identified as 'ceratopogonid midges' and which are likely members of Dasyhelea , based on their size and the presence of distinctive shelf-like tubercles nearly encircling the abdominal segments.

Scanning electron photomicrographs of pupae were published by Ronderos et al. (2004), Dominiak (2012) and Díaz et al. (2013). Park & Downing (2001) published an scanning electron photomicrograph of the terminal abdominal segments of the fossil Miocene species D. australis , showing how a similar approach could be used to better understand other fossil Dasyhelea species.

Pupae of Dasyhelea have a reduced number of thoracic sensilla and the naming of the two setae as D-1-T and D-2-T is arbitrary as they may not be homologous to those specific setae in other ceratopogonids.

Most species of Dasyhelea have abdominal segments with a typical and characteristic transverse arrangement of lower, broad tubercles ( Fig. 56C ). A few species found in Nepenthes in southeast Asia, however, have sensilla on somewhat to markedly elongate tubercles ( Wirth & Beaver 1979). An undescribed species from Australia (Running Creek) also had abdominal sensilla D-4 IV, D-8 IV, L-2 IV, L-3 IV, L-4 IV, V-7 IV on markedly elongate tubercles. D-4-IV in these taxa was represented by a seta (rather than a campaniform sensillum present in other Dasyhelea ).

MATERIAL EXAMINED: D. ampullariae : 2 pupal exuviae, Singapore, 22-VIII-1952 (USNM). D. atlantis : 1 pupal exuviae, Oak Island, Long Island, New York, USA, 15-V-1956 (NYSM); 1 pupal exuviae, Orient Beach, Long Island, New York, USA, 25-V-1963 (USNM); 1 pupal exuviae, 2 km E of Isla Santa Cruz, Galapagos Islands, Ecuador, 18-I-1989 (CNCI); 1 pupal exuviae, Pto. Isla Isabela, Galapagos Islands, Ecuador, 6-III-1989 (CNCI). D. bilineata : 3 pupae, Harz, Germany, 4-IV-1931 (ZSMC); 3 pupal exuviae, (Lunz collection), 1941-1943 (ZSMC). D. biseriata : 1 pupal exuviae (of paratype), Singapore, 6-VIII-1952 (USNM). D. bistriata : 6 pupal exuviae, probably Thuringia, from potash works at Heringen on Werra River, Germany, 23-III-1914 (ZSMC). D. brevicornis : 1 pupal exuviae (of paratype), Salmon River, Blue Mountain Lake, New York, USA, 6-VIII-1959 (USNM). D. chani : 1 pupal exuviae (of paratype), Chinese Farm, Ft. Pierce, Florida, USA, 24-VII-1987 (USNM). D. cincta : 8 pupal exuviae, Poncha Hot Springs, Chaffee County Colorado, USA, 21-IX-1991 (CNCI); 1 pupal exuviae, Los Talas, Argentina, 25-IX-1981 (MLPA). D. contigua : 1 pupal exuviae, Sumatra, Toba area, creek north of Balige, in moss of a waterfall, Indonesia, 1-IV-1929. (ZSMC). D. diplosis : 3 pupae, ditches at Sassendorf saltworks (E of Soest), North Rhine-Westphalia, Germany, mid-April to late May, 1912 (ZSMC). D. flava : 1 pupal exuviae, Burgeshall, Hazyview, East Transvaal, South Africa, 3-XII-1973 (NMSA). D. flavicauda : 1 pupal exuviae, Laguna “La Brava”, Corrientes, Argentina (MLPA). D. flavifrons : 2 pupal exuviae (in glycerin), locality uncertain (CNCI); 4 pupal exuviae, Dolina Radości, Gdańsk, Poland, 3-IV-1989 ( IZUG); 2 pupal exuviae, Loiterau river valley nr Taarstedt, Slesvig-Holstein, NE of Schleswig, Germany, 27-IV-1939 (ZSMC); 4 pupal exuviae, Bavaria, Munich, Germany, XII-1922 (ZSMC); 1 pupal exuviae, Buda Park, Budapest, Hungary, VIII-IX-1986 (CNCI). D. fontana : 1 pupal exuviae, 1 pupal exuviae (of holotype), 25 m. E of Middelburg, Sewefontein, Transvaal, South Africa, 5-XII-1973 (NMSA). D. fusca : 1 pupal exuviae, Zoutpan, Transvaal, South Africa (NMSA); 1 pupal exuviae, 15 mi SE of Potgietersrus, Transvaal, South Africa, 3-I-1974 (NMSA); 1 pupal exuviae, Lodwigslust, Hectorspruit, East Transvaal, South Africa, 29-XI-1973 (NMSA); 2 pupal exuviae, Doornpan, Bulge River IV., Transvaal, South Africa, 6-XI-1973 (NMSA); 1 pupal exuviae, Mamba River, Zululand, South Africa, 5- I-1937 (SAIM). D. gigantosalpinx : 1 pupal exuviae, Jam Tin Creek, Maklane, East Transvaal, South Africa, 2-XII- 1973 (NMSA); 1 pupal exuviae (of paratype), Umlalazi River, Eshowe, Zululand, South Africa, 19-XII-1936 (SAIM). D. grisea : 4 pupal exuviae, California (USNM); 1 pupal exuviae, Hamilton-Essex Line, Newcomb, New York, USA, 24-V-1960 (NYSM); 1 pupal exuviae, Grandview Park Natural Preserve, Hampton City, Virginia, USA, 27-XI-1976 (VPIC). D. flavifrons : 1 pupal exuviae, Vancouver, British Columbia, Canada, 15-V-1979 (CNCI). D. halobia : 8 pupal exuviae, Brenner Moor on Trave river NW of Bad Oldesloe, Slesvig-Holstein, Germany, 15-VII-1922 (ZSMC). D. lacustris : 1 pupal exuviae, Lago Ramos Mexia, Neuquen, Argentina, 2-XII- 1983 (MLPA). D. leptobranchia : 1 pupal exuviae, Fishing Creek, Newcomb, New York, USA, 28-V-1958 (USNM). D. messersmithi : 1 pupal exuviae, Res. Sta., Newcomb, New York, USA, 26-VI-1958 (NYSM); 1 pupa, 1 pupal exuviae, Quogue, Long Island, New York, USA, 9-V-1957 (NYSM); 2 pupal exuviae (of paratypes), Kerr County, Texas, USA, coll. 12-VII-1956, emerged 3-IX-1956 (USNM). D. modesta : 6 pupae, at least four localities in or NW of Bad Oldesloe, Slesvig-Holstein, Germany, late April to June (ZSMC); 11 pupae, northern Italy, 24-V- 1950 (ZSMC). D. mutabilis : 3 pupal exuviae, Rusk County, Wisconsin, USA, 17-X-1953 (USNM); 8 pupal exuviae, High Cr. Fen Preserve, 9 mi S of Fairplay Park, Colorado, USA, 18-VI-1995 (CNCI); 1 pupal exuviae, Hamilton-Essex Line, Newcomb, New York, USA, 29-V-1959 (NSYM); 1 pupal exuviae, Cow Neck Salt Marsh, North Sea, Long Island, New York, USA, 9-VII-1957 (NSYM); 2 pupal exuviae, Dyke Marsh, Alexandria, Virginia, USA, 18-V-1977 (USNM); 1 pupal exuviae, 7 km W of Luray, Page County, Virginia, USA, 6-III-1977 (VPIC). D. nepenthicola : 3 pupal exuviae, Telok Bahang, Penang, Malaysia, I-1977 (USNM); 1 pupal exuviae, as previous locality, 4-XII-1976 (USNM); 1 pupal exuviae, as previous locality, XI-1977 (USNM); 1 pupal exuviae, Ayer Itam, Penang, Malaysia, XI-1976 (USNM). D. nigella : 7 pupal exuviae, Berowra Waters, New South Wales, Australia, 18-IX-1956 (ANIC). D. pallidihalter : 2 pupal exuviae, Lodwig’s Folly, Hectorspruit, East Transvaal, South Africa, 21-I-1974 (NMSA). D. paracincta : 1 pupal exuviae, Isla Isabella, Galapagos Islands, Ecuador, 9-III- 1989 (CNCI); 4 pupal exuviae, Pto. Isla Isabela, Galapagos Islands, Ecuador, 6-III-1989 (CNCI). D. perfida : 2 pupal exuviae, Ranoe Bedali, Java, Indonesia (ZSMC). D. pollinosa : 2 pupal exuviae, 1 pupal exuviae (of paratype), Kern County, California, USA (USNM). D. pseudocincta : 3 pupal exuviae, Hotsprings, Thermopolis, Wyoming, USA, 13-VI-1960 (USNM); 3 pupal exuviae, Oak Island, Long Island, New York, USA, 15-V-1956 (1 USNM, 2 NYSM); 2 pupal exuviae (of paratype), Orient Beach, Long Island, New York, USA, 25-V-1963 (USNM); 1 pupal exuviae, Cow Neck Salt Marsh, Long Island, USA, 1-V-1956 (NYSM). D. pseudoincisurata : 3 pupal exuviae, Iroquois Falls, Ontario, Canada, 21-VI-1987 (CNCI); 5 pupal exuviae, Plummers Island, Maryland, USA, 23-VI-1977 (USNM); 1 pupal exuviae, Mt. Solon, Virginia, USA, 22-VIII-1951 (USNM). D. spiniforma : 1 pupal exuviae, Towd Point, Long Island, New York, USA, 30-VI-1955 (NYSM); 2 pupal exuviae, as previous locality, 28-VI-1955 (NYSM). D. tersa : 1 pupal exuviae, between Tjibodas and the Gedeh, Kuripan, Java, Indonesia, 13-VII-1929 (ZSMC). D. thompsoni : 1 pupal exuviae, Louis Fichardt, North Transvaal, South Africa, 2- I-1974 (NMSA); 1 pupal exuviae, Sabi River, East Transvaal, South Africa 3-XII-1973 (NMSA); 1 pupal exuviae, Doornpan, Bulge River, North Transvaal, South Africa, 6-XI-1973 (NMSA); 1 pupal exuviae, Die Tuine, Transvaal, South Africa, 6-I-1983 (SAIM). D. tugelae : 3 pupal exuviae, 15 m SE of Potgietersrus, Transvaal, South Africa, 3-I-1974 (NMSA); 4 pupal exuviae, 7 km S of Messina, North Transvaal, South Africa, 26-XII-1973 (NMSA); 2 pupal exuviae, Onderstepoort, Transvaal, South Africa, 8-XII-1973 (NMSA); 1 pupal exuviae, Glen Allen Farm, Tzaneen, South Africa, Jan–April, 1974 (SAIM). D. nr. brookmani : 6 pupae, High Cr. Fen Preserve, 9 mi S of Fairplay Park, Colorado, USA, 17-VII-1995 (CNCI). D. major (?): 1 pupal exuviae, Horsepen Creek (Poanoke R. trib.), Charlotte County, Virginia, USA, 15-VI-975 (VPIC). D. nr. mutabilis : 1 pupal exuviae, Res. Sta. Stream, Newcomb, New York, USA, 6-VI-1958 (NYSM); 1 pupal exuviae, Salmon River, Blue Mountain Lake, New York USA, 14-V-1959 (NYSM). D. sp.: 4 pupal exuviae (in glycerin), Herman Lake, 4 km SE Eagle Bay, British Columbia, Canada, 26-V-2013 (CNCI); 5 pupal exuviae, Bolean Lake, 6 km NE of Falkland, British Columbia, Canada, 12-VII-1989 (CNCI); 5 pupal exuviae, as previous locality, 12–13-VII-1989 (CNCI); 1 pupal exuviae, 3 km E of Salmon Arm, British Columbia, Canada, 9-VI-1988 (CNIC); 3 pupal exuviae, Salmon Arm, British Columbia, Canada, 19-VII-1988 (CNIC); 5 pupal exuviae, as previous locality, 2-X-2006 (CNCI); 2 pupal exuviae, Gibson Lake, Kokanee Glacier Provincial Park, British Columbia, Canada, 9-VII-2008 (CNCI); 18 pupal exuviae, Dark Canyon Creek, Eddy County, New Mexico, USA, 13-VI-1991 (CNCI); 1 pupal exuviae, Chosa Draw, Eddy County, New Mexico, USA, 26-VIII-1997 (CNCI); 1 pupal exuviae, Midfork Gila River, Grant County, New Mexico, USA, 15-VIII-1991 (CNCI); 2 pupal exuviae, Gila River above Gila, Grant County, New Mexico, USA, 16-VIII-1991 (CNCI); 2 pupal exuviae, Towd Point, Southampton, Long Island, New York, USA, 29-VII-1954 (NYSM); 2 pupal exuviae, Salt Marsh, Towd Point, Southampton, Long Island, New York, USA, 10- VIII-1954 (NYSM); 1 pupal exuviae, Dune Rd. Salt Marsh, H. Bays, Long Island, New York, USA, 4-VI-1956 (NYSM); 3 pupal exuviae, Cow Neck, Long Island, New York, USA, 28-V-1956 (NYSM); 1 pupal exuviae, New Salem, New York, USA, 13-V-1963 (NYSM); 1 pupal exuviae, Dunaley Bog, 7 mi S of Cashiers, Jackson County, North Carolina, USA, 13-VII-1987 (USNM); 1 pupal exuviae, Montgomery County, Virginia, USA, 22-VI-1975 (VPIC); 3 pupal exuviae, Assateague Is., Chincoteague NWR, Accomack County, Virginia, USA, 1-VIII-1975 (VPIC); 1 pupal exuviae, Waycross, Georgia, USA, 26-VI-1972 (USNM); 1 pupal exuviae, Chiemsee, Germany, 13-VIII-1990 (CNCI); 1 pupal exuviae, Grimswald, Germany (CNCI); 1 pupal exuviae, Brunnsee, Germany, 12- VII-1990 (CNCI); 3 pupal exuviae, as previous locality, 13-VII-1990 (CNCI); 1 pupal exuviae, Lustsee, Germany, 1-VI-1990 (CNCI); 2 pupal exuviae, as previous locality, 11-VIII-1990 (CNCI); 1 pupal exuviae, as previous locality, 10-VI-1990 (CNCI); 2 pupal exuviae, La Gomera, Canary Islands, 8-II-1993 (CNCI); 5 pupal exuviae, Skukusa, Kruger National Park, South Africa, II, III-1993 (ANIC); 1 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 16-IV-1967 (BPBM); 1 pupal exuviae, as previous locality, 14-I-1968 (BPBM); 2 pupal exuviae, as previous locality, 22-I-1967 (BPBM); 1 pupal exuviae, as previous locality, 13-I-1968 (BPBM); 2 pupal exuviae, Dumoga Bone National Park, Sulawesi, Indonesia, 28-I-1985 (ANIC); 1 pupal exuviae, as previous locality, 30-I-1985 (ANIC); 1 pupal exuviae, as previous locality, 1-II-1985 (ANIC); 1 pupa, Darwin Harbour, Darwin, Northern Territory, Australia, 12-V-2006 (CNCI); 2 pupal exuviae, Collie Rd., W. Darban, Western Australia, Australia (ANIC); 5 pupal exuviae, Darban, Western Australia, Australia, 29-X-1985 (ANIC); 1 pupal exuviae, Gnieraoora Pool, Robe River, Pilbara, Western Australia, Australia, 30-X-1995 (ANIC); 12 pupal exuviae, as previous locality (ANIC); 1 pupal exuviae, Upper Durack River, Kimberley, Western Australia, Australia (ANIC); 1 pupal exuviae, Richenda Gorge, Kimberley, Western Australia, Australia, 10-V-1995 (ANIC); 1 pupal exuviae, Palm Springs, Fitzroy, Kimberley, Western Australia, Australia, 12-V-1995 (ANIC); 1 pupal exuviae, King Edward River, Kalumburu, Kimberley, Western Australia, Australia (ANIC); 9 pupal exuviae, Cangan Pool, Pilbara, Western Australia, Australia (ANIC); 1 pupal exuviae, as previous locality, 15-X-1995 (ANIC); 2 pupal exuviae, Whale Back Creek, Fortescu River, Pilbara, Western Australia, Australia, 16-V-1995 (ANIC); 1 pupal exuviae, Erewallana Pool, Pilbara, Western Australia, Australia, 1-X-1995 (ANIC); 1 pupal exuviae, as previous locality, 1-V-1995 (ANIC); 2 pupal exuviae, as previous locality, 9-V-1995 (ANIC); 5 pupal exuviae, Whiskey Pool, Ashburton River, Pilbara, Western Australia, Australia, 28-IV-1995 (ANIC); 1 pupal exuviae, Mooka Ruins, Gascoyne River, Western Australia, Australia, 15-VIII-1995 (ANIC); 3 pupal exuviae, Yanchep National Park, Western Australia, Australia, 24-X-1985 (ANIC); 1 pupal exuviae, Receptably Cemetery, Townsville, Queensland, Australia, 31-V-1950 (ANIC); 1 pupal exuviae, Southport, Queensland, Australia, 21-V- 1953 (ANIC); 3 pupal exuviae, Basin, Southport, Queensland, Australia, 31-XII-1950 (ANIC); 1 pupal exuviae, Gladstone, Queensland, Australia, 28-I-1947 (ANIC); 1 pupal exuviae, as previous locality, 27-I-1947 (ANIC); 1 pupal exuviae, FNQS Running Creek, Weipa, Queensland, Australia, 9-X-1985 (ANIC); 6 pupal exuviae, Hornsby, New South Wales, Australia, 22-II-1966 (ANIC); 1 pupal exuviae, Careel Bay, New South Wales, Australia, 16-II- 1966 (ANIC); 1 pupal exuviae, as previous locality, 29-X-1956 (ANIC); 1 pupal exuviae, as previous locality, 20- IX-1965 (ANIC); 2 pupal exuviae, Colo Vale, New South Wales, Australia, 24-II-1958 (ANIC); 6 pupal exuviae, as previous locality, 18-X-1956 (ANIC); 2 pupal exuviae, Stockyard Creek, Colo Vale, New South Wales, Australia, 10-III-1957 (ANIC); 2 pupal exuviae, Nattai River, Mittagong, New South Wales, Australia, 4-XI-1964 (ANIC); 1 pupal exuviae, as previous locality, 18-X-1968 (ANIC); 3 pupal exuviae, as previous locality, 25-X-1968 (ANIC); 1 pupal exuviae, as previous locality, 15-XI-1968 (ANIC); 2 pupal exuviae, as previous locality, 18-II-1969 (ANIC); 8 pupal exuviae, Nattai River, The Crags, Mittagong, New South Wales, Australia, 9-II-1966 (ANIC); 1 pupal exuviae, Picton Lakes, New South Wales, Australia, 2-II-1966 (ANIC); 4 pupal exuviae, Mona Vale, New South Wales, Australia, 10-X-1956 (ANIC); 1 pupal exuviae, Oxford Falls, New South Wales, Australia, 6-XI- 1947 (ANIC); 4 pupal exuviae, as previous locality, 18-I-1967 (ANIC); 3 pupal exuviae, Wattamolla, New South Wales, Australia, 24-II-1969 (ANIC); 3 pupal exuviae, McCarrs Creek, New South Wales, Australia, 28-II-1971 (ANIC); 2 pupal exuviae, as previous locality, 4-III-1971 (ANIC); 2 pupal exuviae, as previous locality, 14-I-1969 (ANIC); 6 pupal exuviae, as previous locality, 20-IX-1956 (ANIC); 1 pupal exuviae, Anson Bay, Norfolk Island, Australia, 14-IV-1972 (ANIC); 9 pupal exuviae, Norfolk Island, Australia, 19-IV-1972 (ANIC); 3 pupal exuviae, no data (ANIC); 1 pupal exuviae, Australia, 9-VI-1971 (ANIC).

Figures

FIGURE 11. Habitus. A. Forcipomyia (Thyridomyia) rugosa, male, in dorsal view (from Chan & LeRoux 1970). B. Forcipomyia (Pterobosca) crinume, female, in ventral view (from Tokunaga 1932). C. Forcipomyia (Lasiohelea) rugosa, male, in dorsal view (from Saunders 1964). D. Atrichopogon hexastichus, male, in dorsal view (from Nielsen 1951). E. Atrichopogon wirthi, female, in dorsal view (from Chan & Linley 1988). F. Atrichopogon muelleri, sex unknown, in dorsal view, with larval exuviae on posterior of abdomen (from Szadziewski et al. 1997). G. Dasyhelea vericolor, sex unknown, in dorsal view (from Szadziewski et al. 1997). H. Schizonyxhelea bulla, sex unknown, in dorsal view (from Thomsen 1937).

FIGURE 13. Anterior portion of heads and thoraces, in anterior or anterodorsal view. A. Leptoconops kerteszi, ecdysial sutures of head enhanced to show position. B. Culicoides denticulatus. C. Ceratopogon nr. abstrusus. D. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). E. Alluaudomyia bella, dorsal apotome absent. F. Clinohelea curriei. G. Pellucidomyia leei. H. Mallochohelea caudellii. I. Bezzia sp. (from Bolean Lake, BC, Canada).

FIGURE 14. Anterior portion of heads. A. Austroconops mcmillani, in ventral view, dorsal apotome not present. B. Leptoconops bequaerti, in anterior view (distorted slide specimen). C. Forcipomyia (Phytohelea) sp. in anterior view (Florida, USA). D. Forcipomyia (Phytohelea) sp. in anteroventral view (Florida, USA). E. Atrichopogon sp., in anterior view (from 6.5 km NW Enderby, BC, Canada). F. Dasyhelea sp., in anteroventral view (Herman Lake, 4 km SE Eagle Bay, BC, Canada).

FIGURE 18. Dorsal apotomes, anterior view. A. Austroconops mcmillani (from Borkent & Craig 2004). B. Leptoconops bequaerti. C. Forcipomyia (Phytohelea) n. sp. near F. antiguensis. D. Forcipomyia (Euprojoannisia) sp. (undescribed, from Peru). E. Atrichopogon sp. (undescribed, from Costa Rica). F. Dasyhelea insons (from Mayer 1934c). G. Dasyhelea tersa (from Mayer 1934c). H. Dasyhelea assimilis (from Mayer 1934c). I. Dasyhelea pseudocincta (from Waugh & Wirth 1976, modified). J. Dasyhelea pseudoincisurata (from Waugh & Wirth 1976, modified). K. Dasyhelea messersmithi (from Waugh & Wirth 1976, modified). L. Dasyhelea atlantis (from Waugh & Wirth 1976, modified).

FIGURE 23. Posterior portion of heads, in ventral view. A. Austroconops mcmillani. B. Leptoconops kerteszi. C. Forcipomyia (Euprojoannisia) sp. (from Peru). D. Forcipomyia (Phytohelea) n. sp. near antiguensis. E. Atrichopogon sp. (from St. Pierre de Wakefield, Quebec, Canada). F. Dasyhelea paracincta. G. Dasyhelea pseudoincisurata.

FIGURE 29. Dorsal setae from left side of thoraces. A. Austroconops mcmillani. B. Leptoconops kerteszi. C. Leptoconops linleyi. D. Forcipomyia (Euprojoannisia) sp. (from Peru). E. Forcipomyia (Forcipomyia) bipunctata. F. Forcipomyia (Phytohelea) n. sp. nr. antiguensis. G. Atrichopogon sp. (from Lake Opinicon, Ontario, Canada). H. Dasyhelea paracincta. I. Dasyhelea atlantis. J. Dasyhelea pseudoincisurata. K. Culicoides haematopotus. L. Culicoides phlebotomus. M. Paradasyhelea albipunctata. N. Ceratopogon boomerangus. O. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). P. Alluaudomyia needhami. Q. Baeodasymyia christopheri (from Borkent & Craig 1999). R. Austrohelea campbellensis. S. Stilobezzia antennalis. T. Schizonyxhelea forattinii (from Borkent 2000a).

FIGURE 32. Cephalothoraces and abdominal segments 1–3, in lateral view. A. Austroconops mcmillani. B. Leptoconops linleyi. C. Forcipomyia (Phytohelea) sp. (Florida, USA). D. Atrichopogon sp. (from 6.5 km NW Enderby, BC, Canada). E. Dasyhelea sp., in anteroventral view (Herman Lake, 4 km SE Eagle Bay, BC, Canada). F. Dasyhelea oppresa. G. Culicoides denticulatus. H. Ceratopogon nr. abstrusus. I. Brachypogon sp. (from 6 km E. Falkland, BC, Canada). J. Alluaudomyia bella. K. Baeodasymyia michaeli. L. Stilobezzia antennalis.

FIGURE 34. Legs, wings and apices of the antennae, in ventral view. A. Austroconops mcmillani. B. Leptoconops kerteszi. C. Forcipomyia (Euprojoannisia) sp. (from Peru). D. Atrichopogon sp. (from Lake Opinicon, Ontario, Canada). E. Dasyhelea paracincta. F. Culicoides sp. (from 3 km E. Carp, Ontario, Canada).

FIGURE 43. Respiratory organs, most in dorsal view, Fig. 43T in lateral view, pattern on tracheal tubes shown only in some. A. Dasyhelea pseudoincisurata (from Waugh & Wirth 1976). B. Dasyhelea grisea (from Waugh & Wirth 1976). C. Dasyhelea traverae (from Waugh & Wirth 1976). D. Dasyhelea pseudocincta (from Waugh & Wirth 1976). E. Dasyhelea leptobranchia (from Waugh & Wirth 1976). F. Dasyhelea brevicornis (from Waugh & Wirth 1976). G. Dasyhelea hitchcocki (from Wirth 1976a). H. Culicoides chiopterus (from Jamnback 1965). I. Culicoides furens (from Jamnback 1965). J. Culicoides guttipennis (from Jamnback 1965). K. Culicoides dewulfi (from Kettle & Lawson 1952, as C. pseudochiopterus). L. Culicoides punctatus (from Kettle & Lawson 1952). M. Culicoides sanguisuga (from Jamnback 1965). N. Culicoides variipennis (from Jamnback 1965). O. Paradasyhelea albipunctata (from Elson-Harris & Kettle 1985a). P. Ceratopogon inverecundus (from Borkent & Grogan 1995). Q. Ceratopogon willisi (from Borkent & Grogan 1995). R. Brachypogon sp. (from Virginia, USA). S. Alluaudomyia needhami (from Wirth & Grogan 1981). T. Alluaudomyia fumosipennis (from Debenham 1971).

FIGURE 47. Metathoraces and tergites 1 (in dorsal view). A. Austroconops mcmillani. B. Leptoconops kerteszi. C. Forcipomyia (Forcipomyia) bipunctata. D. Forcipomyia (Phytohelea) n. sp. nr. antiguensis. E. Atrichopogon sp. (from Lake Opinicon, Ontario, Canada). F. Dasyhelea pseudoincisuata.

FIGURE 56. Abdominal segment four, in dorsal and ventral view (shagreen not shown). A. Forcipomyia dominicana. B. Atrichopogon sp. (from Lake Opinicon, Ontario, Canada). C. Dasyhelea paracincta.

FIGURE 72. Abdominal segments 9. A. Austroconops mcmillani, male, in dorsal and ventral view (from Borkent & Craig 2004). B. Leptoconops whitseli, male, in dorsal and ventral view (from Clastrier & Wirth 1978). C. Leptoconops whitseli, female, in dorsal and ventral view (from Clastrier & Wirth 1978). D. Leptoconops asilomar, male, in dorsal and ventral view (modified from Clastrier & Wirth 1978). E. Leptoconops asilomar, female, in dorsal and ventral view (modified from Clastrier & Wirth 1978). F. Forcipomyia anabaenae, female, in dorsal view (from Chan & Saunders 1965). G. Forcipomyia eques, male, in ventral view (from Chan & LeRoux 1965). H. Forcipomyia rugosa, female, in dorsal view (from Chan & LeRoux 1970). I. Forcipomyia brevis, female, in dorsal view (from Mayer 1934a). J. Atrichopogon wirthi, male, in dorsal view, arrow indicating anteriorly directed spines (from Chan & Linley 1988). K. Dasyhelea contigua, female, in ventral view (from Mayer 1934c). L. Dasyhelea grata, male, in ventral view (from Mayer 1934c).

FIGURE 73. Abdominal segments 9. A. Dasyhelea laeta, female, in ventral view (from Mayer 1934c). B. Dasyhelea assimilis, male, in lateral view with ventral to the right (from Mayer 1934c). C. Dasyhelea assimilis, male, in lateral view with ventral to the right (from Mayer 1934c). D. Dasyhelea grisea, male, in dorsal view (from Waugh & Wirth 1976). E. Dasyhelea chani, female, in dorsal view (from Wirth & Linley 1990). F. Dasyhelea atlantis, female, in ventral view (from Waugh & Wirth 1976). G. Dasyhelea gluchovae, female, in ventral view (from Brodskaya 1996). H. Culicoides haematopotus, female, in dorsal and ventral view (shagreen not shown). I. Paradasyhelea albipunctata, female, in dorsal view (Elson-Harris & Kettle 1985a). J. Ceratopogon boomerangus, female, in ventral view (modified from Borkent & Grogan 1995). K. Brachypogon artemis, female, in dorsal view (from Debenham 1991). L. Alluaudomyia appendiculata, female, in dorsal view (from Elson-Harris & Kettle 1985b).

FIGURE 79. Evolution of ecdysial sutures and splits in pupal exuviae of Ceratopogonidae, head and anterior portion of thorax in ventral view. Numbers refer to character states of characters 2 and 8 in text. Red indicates ecdysial splits. A. Leptoconopinae. B. Forcipomyiinae. C. Dasyheleinae. D. Most Ceratopogonini. E. Most Ceratopogonini, Clinohelea. F. Allohelea, Atyphohelea, Parabezzia. G. Heteromyia, Pellucidomyia, Hebetula. H. most Sphaeromiini s. lat., most Palpomyiini, Paryphoconus; some Bezzia are nearly completed fused (as in Fig. 79I), with only the dorsal apotome separate, along with an ecdysial split along the base of the antenna and the dorsal thoracic split. I. Jenkinshelea, Macropeza, Mallochohelea, Probezzia, Neobezzia.