Atrichopogon 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 : 43-44

Atrichopogon Kieffer

( Figs. 11D–F , 14E , 18E , 23E , 29G , 32D , 34D , 42S–T , 47E , 56B , 72J )

DIAGNOSIS: Only pupa of Ceratopogonidae with terminal process with at least some spicules directed anteriorly.

DESCRIPTION: Habitus as in Figs. 11D–F . Total length = 1.41–3.44 mm. With ( Fig. 11F ) larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face ( Fig. 14E ). Ecdysial tear anterior or slightly medial to base of antenna ( Figs. 14E , 79B ); along posterolateral portion of eye. Head: Dorsal apotome ( Fig. 18E ), without ventral line of weakness, with 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 ( Fig. 23E ) with mandible, lacinia well-developed, not overlapping apically; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, in some by hypopharynx; apex of antenna ( Fig. 34D ) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals ( Fig. 18E )—1 moderate to elongate seta; dorsolateral cephalic sclerite sensilla—1 short seta or 1 very short or elongate seta, 1 campaniform sensillum; clypeal-labrals ( Fig. 23E )—1 seta, 1 campaniform sensillum; oculars ( Fig. 23E )—1 campaniform sensillum. Thorax: Prothoracic extension ( Fig. 23E ) absent; mesonotum with long, bifurcating tubercles, extending posteromedially, completely dividing metathorax medially ( Fig. 47E ); respiratory organ ( Figs. 42 S-T) length/width = 1.25–3.50, variable, knob-like to elongate, with posterior basal swelling or lobe, somewhat flattened laterally, with pores closely abutting or slightly separated at apex of respiratory organ, arranged in single straight or curved row or two widely spaced rows, outer surface smooth or with a few wrinkles, with welldeveloped, flattened pedicel, base without posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, surface smooth; wing ( Fig. 34D ) without apical tubercle, separated medially by fore-, midlegs; halter apex and hind leg ( Fig. 32D ) broadly separate; halter apex abutting anterolateral edge of tergite 2; legs ( Fig. 34D ) with lateral margin of foreleg near midlength of wing slightly sinuous to evenly curved; hind leg visible at lateral margin of wing ( Fig. 32D ); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials—1 campaniform sensillum; anterolaterals—1 short seta, 1 campaniform sensillum; dorsal setae ( Fig. 29G )—D-1-T, D-2-T setae, D-3-T campaniform sensillum; D-2-T, D-3-T on single tubercle in some, or closely associated; supraalar 2—campaniform sensillum present or absent; metathoracics ( Fig. 47E )—2 campaniform sensilla; M-3-T near anterior margin of metathorax. Abdomen: without pigmentation pattern, segment 2 as wide or somewhat wider than segment 3, segments with undivided, thin to thick setae, without tubercles or segments 1–5 to 1–8 with branched or setaceous elongate tubercles, tubercles rounded to pointed, tergites or sternites entire, each without membranous disc; segment 9 ( Fig. 72J ) variable in shape, terminal processes widely separated basally, each projecting posterolaterally, tapering near apex to point, with at least some spicules directed anteriorly; sensilla: tergite 1 ( Fig. 47E ) with 3 setae, 2 campaniform sensilla, including 2 lateral sensilla, D-3-I absent, D-7-I absent; segment 4 ( Fig. 56B )—D-2-IV moderately elongate seta, without tubercle, D-3-IV absent; D-4-IV present; D-5-IV, D-7-IV, D-8-IV, D-9-IV absent; D-4-IV without tubercle, L-1-IV slender or stout, short to elongate seta present or absent, V-5-IV, V-6-IV present or absent, V-7-IV small seta, none on tubercles; segment 8 without D-3-VIII, without L-1-VIII; segment 9 ( Fig. 72J )—D-5-IX campaniform sensillum, possibly D-6-IX present but inadequate specimens examined.

DISTRIBUTION AND HABITAT: The genus Atrichopogon is known from 521 species from every Region worldwide ( Borkent 2014 ). Immatures occur in aquatic to terrestrial habitats, generally in shaded areas on algae or mosses present on or at the edge of ponds, marshes and streams, or over wet soil, rock surfaces and wet wood. Larvae and pupae are also found under bark of logs, on rotting leaves, and other moist microhabitats in forests where fungi and microorganisms are available for food.

TAXONOMIC DISCUSSION: The pupae of only 39 species of Atrichopogon have been described ( Tables 2–3 ) and these different significantly in numbers of details. Indeed, the morphology of both larvae and pupae are so diverse that previous workers have suggested that the immatures would provide a better means of distinguishing the species ( Nielsen 1951, Ewen & Saunders 1958, Chan & Linley 1988).

Szadziewski et al. (1995) provided a key to European subgenera and Lenz (1934) gave a key to the few European species previously described as pupae. Ewen & Saunders (1958) described 19 species from the New World but did not include a key (in spite of obvious differences between taxa). Nielsen (1951) described six species from Denmark in well-illustrated and detailed study, including their behaviour and biology. His phylogenetic interpretation of the genus included pupal structures. Thomsen (1937) gave a brief key to the three Nearctic species she studied.

The description of A. websteri by Thomsen (1937) suggests it was misidentified. As illustrated, the elongate respiratory organ, the lack of lateral abdominal tubercles and the terminal processes closely approximated medially and with posteriorly directed spicules suggest it is actually a Forcipomyia .

Pupae of Atrichopogon have a reduced number of thoracic sensilla, making the exact naming of these unsure. D-3-T, as a unique campaniform sensillum, is likely homologous to that in other Ceratopogonidae . The abdominal chaetotaxy is also strongly reduced and the remaining sensilla vary in position, making identification of specific sensilla (as in Fig. 56B ) often uncertain. Some species have one or two more sensilla on abdominal segment four (e.g. A. maculosus has two lateral setae) and the presence of heavy shagreen in many species makes it particularly difficult to discern sensilla when these are small. In A. jacobsoni , sensillum D-4-IV identified here appears to be homologous to an elongate seta. Furthermore, homologies are further confused by the failure of some descriptions to distinguish tubercles with or without sensilla (e.g. Ewen & Saunders 1958).

MATERIAL EXAMINED: A. bifidus : 1 pupa (of paratype), Nictheroy, Brazil, 31-VII-1923 (CNCI). A. caribbeanus : 2 pupal exuviae (of paratypes) Tobago, Tobago and Trinidad, 31-V-1953 (CNCI). A. corpulentus : 1 pupal exuviae (of paratype), Nanaimo, BC, Canada, 22-VII-1926 (CNCI). A. crinitus : 2 pupal exuviae (of paratypes), Nanaimo, BC, Canada, 22-VII-1926 (CNCI). A. flavus : 2 pupal exuviae (of paratypes), Beaver Creek, Saskatoon, Saskatchewan, Canada, 8-X-1955 (CNCI). A. fusculus : 2 pupal exuviae, Pike Lake, Saskatoon, Saskatchewan, Canada, IX-1956 (CNCI). A. fuscus : 5 pupal exuviae, no locality, coll. by Goetghebuer (so likely western Europe) (CNCI). A. geminus (as A. levis ): 3 pupal exuviae, Put-in-Bay, Ohio, USA (CNCI). A. humicolus : 1 pupal exuviae (of paratype), Saskatoon, Saskatchewan, Canada, 9-IX-1955 (CNCI). A. inconspicuus : 4 pupal exuviae (of paratypes), Saskatoon, Saskatchewan, Canada, 17-24-IX-1955 (CNCI). A. incultus : 1 pupal exuviae (of paratype), Siquirres, Costa Rica, 18-VI-1956 (CNCI). A. jacobsoni : 1 pupal exuviae, Batu Caves, Cavern A, Kuala Lumpur, Selanger, Malaysia, 27-XII-1960 (ANIC); 1 pupa, 1 pupal exuviae, as previous locality, 27-XII-1960 (USNM). A. maculosus : 4 pupae, 5 pupal exuviae (of paratypes), Beaver Creek, Saskatoon, Saskatchewan, Canada, 25-28-VIII-1955 (CNCI); 1 pupal exuviae, Patuxent Wildlife Refuge, Prince George’s County, Maryland, USA, 1- VIII-1979 (USNM); 2 pupal exuviae, as previous locality, 10-VIII-1979 (USNM); 1 pupal exuviae, 5 mi NW of Davidsburg, York County, Pennsylvania, USA, 4-VII-1961 (USNM). A. minutus : 4 pupal exuviae, Lit. Abington, Cambs., England, Great Britain, 27-VIII-1924 (CNCI); 2 pupal exuviae, Truro, Canada, 10-VIII-1925 (CNCI); 8 pupal exuviae, Nanaimo, BC, Canada, 22-VII-1926 (CNCI); 11 pupal exuviae, Victoria, BC, Canada, 26-VIII-1947 (CNCI); 14 pupal exuviae, previous locality, 20-VII-1948 (CNCI); 11 pupal exuviae, previous locality, 12-VII- 1948 (CNCI). A. obscurus : 1 pupa, 1 pupal exuviae (of paratype), Mayaguez, Puerto Rico, 25-II-1953 (CNCI). A. remigatus : 1 pupal exuviae (of paratype), Petropolis, Brazil, 27-VII-1923 (CNCI). A. saundersi : 4 pupal exuviae (of paratypes), Mayaguez, Puerto Rico, 27-III-1953 (CNCI). A. tuberculatus : 2 pupal exuviae (of paratypes), Macaras, Trinidad, 14-VII-1957 (CNCI). A. winnertzi (as A. meloesugans ): 12 pupal exuviae, Strelly, Notts., England, Great Britain, 3-XI-1922 (CNCI). A. wirthi : 2 pupal exuviae, Chinese Farm, Ft. Pierce, Florida, USA, VII-1987 (CNCI). A. nr. humicolus : no locality given, V-1953 (USNM). A. (Lophomyidium) sp.: 1 pupal exuviae, orilla Rio Dantes, Parque Nacional Barbilla, Cartago, Costa Rica, 10-II-2006 (CNCI). A. sp.: 4 pupal exuviae, 2 pupal exuviae (in glycerin), 6.5 km NW of Enderby, British Columbia, Canada, 27-VI-1990 (CNCI); 1 pupal exuviae, Lake Opinicon, Ontario, Canada, coll. 11-VII-1966, emerged 29-VIII-1966 (CNCI); 1 pupal exuviae, as previous locality, 12-XII-1966 (CNCI); 1 pupal exuviae, as previous locality, 25-VIII-1966 (CNCI); 2 pupal exuviae, St. Pierre de Wakefield, Quebec, Canada, 25-VI-1964 (CNCI); 5 pupal exuviae, orilla Rio Dantes, Parque Nacional Barbilla, Cartago, Costa Rica, 10-II-2006 (CNCI).

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 42. Respiratory organs, most in dorsal view, Fig. 42B in lateral view, pattern on tracheal tubes shown only in some. A. Austroconops mcmillani (from Borkent & Craig 2004). B. Austroconops mcmillani (from Borkent & Craig 2004). C. Leptoconops americanus (from Clastrier & Wirth 1978). D. Leptoconops asilomar (from Clastrier & Wirth 1978). E. Forcipomyia (Euprojoannisia) wirthi (from Saunders 1957). F. Forcipomyia (Euprojoannisia) longispina (from Saunders 1957). G. Forcipomyia (Warmkea) malayae (from Saunders 1957). H. Forcipomyia (Caloforcipomyia) caerulea (from Saunders 1957). I. Forcipomyia (Caloforcipomyia) varicolor (from Saunders 1957). J. Forcipomyia (Metaforcipomyia) cerifera (from Saunders 1957). K. Forcipomyia (Forcipomyia) genualis (from Saunders 1957, as raleighi). L. Forcipomyia (Forcipomyia) nigra (from Saunders 1924). M. Forcipomyia (Forcipomyia) ciliata (from Saunders 1924). N. Forcipomyia (Phytohelea) bromelicola (from Saunders 1925, (as an Apelma)). O. Forcipomyia (Synthyridomyia) acidicola (Synthyridomyia) (from Saunders 1924). P. Forcipomyia (Trichohelea) crinita (from Saunders 1964). Q. Forcipomyia (Lasiohelea) quasicornuta (from Saunders 1964). R. Forcipomyia (Lasiohelea) cornuta (from Saunders 1964). S. Atrichopogon wirthi (from Chan & Linley 1988). T. Atrichopogon thienemanni (from Lenz 1934).

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 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.