taxonID	type	description	language	source
027587C9BD393052FE2F1A8248ECE2F7.taxon	discussion	This is the first attempt at keying pupae of all known genera of Ceratopogonidae of the world. Generally, regional keys are more useful for identification but the material I examined was seriously limited and often included reared species representing genera which were otherwise unknown within a given region as immatures. The inclusion of distributions for each keyed genus, based on all stages, will help to exclude some taxa for those keying material from a given area. A key to the known subgenera of Forcipomyia is also included. Each of the families of Culicomorpha was keyed by Borkent (2012). Ceratopogonidae are the only Diptera with a curved S-shaped hind leg lying under the wing, with the terminal process at the apex of the abdomen either lobe-shaped or tapering to a point, without abdominal spiracles on segments 5 – 7 (or elsewhere on the abdomen) and a respiratory organ with at least some apical pores (many with additional subapical pores). Some ceratopogonid pupae are superficially quite similar to those of some Psychodidae but can be distinguished by their S-shaped hind leg. In the following key I have attempted to key out common taxa first and to use features that can be seen in material preserved in alcohol or glycerin. Regrettably, for some genera, slide mounting is required and in such instances, readers are strongly urged to read the materials and methods section — poorly mounted specimens and especially those that are crushed by coverslips, are often miserable to identify. Well prepared material is always of benefit in using keys to insects but is particularly important with ceratopogonid pupae. If pupae, and especially pupal exuviae, are compressed by coverslips, this can make identification hellish but in most instances, yet possible, if the reader is experienced in recognizing morphological features. So too, dirty specimens often obscure important sensilla and can make their detection nearly impossible. Features of segment 4 are generally also present on segments 5 – 7, so that if segment 4 is obscured the specimens may yet be identified using these other segments as surrogates. Readers will see that some genera, such as Palpomyia, key out in more than one place in the key. Some genera could not be diagnosed as separate from all other genera and this may indicate the need for further research. Either some of these genera are paraphyletic, I may have missed some important features, been fooled by features that are markedly homoplastic (i. e. using highly variable features as part of the key), was misled by misassociations and / or misidentifications, or any combination of these potential problems. If there are differences between males and females noted in the key, males can be distinguished from females by the presence of two well-developed genital lobes, nearly always situated ventromedially (some Forcipomyia males have the lobes placed dorsally). The relative development of the male genital lobes reflects the general length of the developing gonocoxites and gonostyli. Readers should be particularly aware that this key is undoubtedly flawed and seriously incomplete, with only 45 of 111 genera known, and of those known, for many genera, only a small percentage have been reared (Table 3). Of these known genera, Amerohelea is too poorly understood to be included. A key to the subgenera of Forcipomyia is included below but only 14 of 35 subgenera are known as pupae. For further challenges, readers are urged to read " Warning to Readers " above. Properly prepared specimens are important and sometimes vital to identification (see Materials and Methods). Overall, pupal exuviae in good condition are easier to identify than whole pupae (which need to be carefully cleared to see sensilla).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD0A3053FD8A18D84EDAE0F4.taxon	description	(Figs. 8 A, 14 A, 18 A, 23 A, 29 A, 32 A, 34 A, 42 A – B, 47 A, 55 A, 72 A)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD0A3053FD8A18D84EDAE0F4.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with pair of elongate dorsocephalic sclerite setae abutting the respiratory organ (Fig. 8 A); also unique with prothoracic extension restricted to lateral margin of palpus as narrow lobe (Fig. 23 A).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD0A3053FD8A18D84EDAE0F4.taxon	description	DESCRIPTION: Habitus as in Fig. 8 A. Total length = 1.84 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum separate from lateral margin of face (Fig. 14 A). Ecdysial tear along anterior of and anteromedial to base of antenna, entire head capsule loose (Figs. 14 A, 79 A). Head: Dorsal apotome (Fig. 18 A), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 A) separated from scutum by thin cuticle, separate from scutum upon emergence, each side broadly meeting medially in whole pupa; mouthparts (Fig. 23 A) with mandible, lacinia well-developed, not overlapping apically; palpus extending anterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna (Fig. 34 A) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Fig. 18 A) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 2 elongate setae with apices laying against respiratory organ, 1 campaniform sensillum (Fig. 8 A); clypeal-labrals (Fig. 23 A) — 1 elongate seta (campaniform sensillum, if present, invisible on wrinkled cuticle); oculars (Fig. 23 A) — 2 elongate setae, one longer than other. Thorax: Prothoracic extension (Fig. 23 A) restricted to lateral margin of palpus as narrow lobe; mesonotum without tubercles, extending posteromedially, nearly dividing metathorax medially (Fig. 47 A); respiratory organ (Figs. 42 A – B) length / width = 3.27, moderately elongate, somewhat flattened laterally, with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface with some wrinkles, with elongate conical pedicel, base without posteromedial apodeme, membranous base of respiratory organ elongate, annulated, tracheal tube slightly sinuous, with spirals extending just over half length, distally smooth; wing (Fig. 34 A) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 32 A) broadly separate; halter apex extending to about half length of tergite 1; legs (Fig. 34 A) with lateral margin of foreleg near midlength of wing somewhat sinuous; hind leg not visible at lateral margin of wing (Fig. 32 A); with apex of foreleg ventral to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 1 very minute seta; anterolaterals — 2 elongate, 1 short setae; dorsal setae (Fig. 29 A) — D- 1 - T, D- 2 - T, D- 4 - T bifurcating setae, D- 3 - T campaniform sensillum, D- 3 - T, if present, posteromedial to D- 4 - T; supraalar 2 — elongate seta (Fig. 8 A); metathoracics (Fig. 47 A) — 2 setae, 1 campaniform sensillum; M- 3 - T near anterior margin of metathorax. Abdomen: without pigmentation pattern, segments 2, 3 equally or nearly equally wide, segments with elongate, bifurcating setae, with rounded to pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 72 A) not strongly modified, terminal processes widely separated basally, each projecting posterodorsolaterally with apex directed dorsally, tapering to pointed apex; sensilla: tergite 1 (Fig. 47 A) with 5 setae, 1 campaniform sensillum, including 1 lateral sensillum, D- 2 - I, D- 3 - I on shared tubercle, D- 7 - I absent; segment 4 (Fig. 55 A) — D- 2 - IV, D- 3 - IV elongate, bifid setae on single tubercle; D- 5 - IV, D- 8 - IV, D- 9 - IV bifurcating elongate setae on short, separate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 8 - IV, D- 7 - IV, D- 9 - IV; D- 4 - IV positioned medial to D- 3 - IV; L- 1 - IV elongate seta on short, pointed tubercle, well anterior of posterior lateral setae, L- 2 - IV, L- 3 - IV, L- 4 - IV bifid setae on short pointed tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV bifid setae, V- 5 - IV, V- 6 - IV on elongate tubercles, V- 7 - IV on short tubercle; segment 8 without D- 3 - VIII, with L- 1 - VIII; segment 9 (Fig. 72 A) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD0A3053FD8A18D84EDAE0F4.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Austroconops, once broadly distributed in the Cretaceous, is now known from two species from southwestern Australia (Borkent & Craig 2004). Although immmatures are unknown in nature, the larvae moved snake-like through very wet substrate in the laboratory, clearly indicating that their natural habitat must be subaquatic or aquatic. It would be valuable to search for larvae and pupae at Yanchep National Park, in Western Australia, where a large adult population of A. mcmillani lives with this in mind. The behaviour of the lethargic pupa was described by Borkent & Craig (2004).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD0A3053FD8A18D84EDAE0F4.taxon	discussion	TAXONOMIC DISCUSSION: Borkent & Craig (2004) described the pupa of A. mcmillani from a single pupal exuviae, reared from an egg, and this is the only known specimen of the genus. Borkent & Craig (2004) missed D- 3 - T and D- 4 - I (both campaniform sensilla), identified the closely approximated sensilla D- 2 - IV and D- 3 - IV as a single seta (as dasm ii) and missed the minute D- 1 - IV.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD0A3053FD8A18D84EDAE0F4.taxon	materials_examined	MATERIAL EXAMINED: A. mcmillani: 1 pupal exuviae, Yanchep National Park, Western Australia, Australia, reared from egg from female collected 20 - XI- 2001 (CNCI).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD0B3051FD5C1951485CE3F1.taxon	description	(Figs. 8 B – C, 13 A, 14 B, 18 B, 23 B, 29 B – C, 32 B, 34 B, 42 C – D, 47 B, 55 B, 72 B – E)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD0B3051FD5C1951485CE3F1.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with apically rounded, lobe-like terminal process (Figs. 72 B – E); also unique without the hind leg visible laterally along the wing margin and the palpus extending beyond the labium (Fig. 23 B); also unique to many species in the presence of thick, recurved setae on abdominal segments (Fig. 55 B).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD0B3051FD5C1951485CE3F1.taxon	description	DESCRIPTION: Habitus as in Figs. 8 B – C. Total length = 1.69 – 2.69 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum separate from lateral margin of face (Fig. 14 B). Ecdysial tear along anterior to base of antenna, entire head capsule loose (Figs. 14 B, 79 A). Head: Dorsal apotome (Fig. 18 B), with ventral bend uncertain (no alcohol pupal exuviae examined), without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (Fig. 13 A) separated from scutum by thin cuticle, separate from scutum upon emergence, each side broadly meeting medially in whole pupa; mouthparts (Fig. 23 B) with mandible, lacinia well-developed, not overlapping apically; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 34 B) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Fig. 18 B) — 1 short seta, at least 2 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 short seta, 1 campaniform sensillum; clypeal-labrals (Fig. 23 B) — 1 very short seta; oculars (Fig. 23 B) — 1 short seta, 1 campaniform sensillum (barely visible). Thorax: Prothoracic extension (Fig. 23 B) absent; mesonotum with moderately sized tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 47 B); respiratory organ (Figs. 42 C – D) length / width = 1.17 – 1.95, short, squat or moderately elongate, somewhat flattened dorsoventrally, with pores closely abutting at apex of respiratory organ, arranged in single or double row, outer surface smooth, with elongate conical pedicel, base without posteromedial apodeme, membranous base of respiratory organ short, tracheal tube somewhat curved, expanded apically, with spirals extending just over half length, distally smooth; wing (Fig. 34 B) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Figs. 8 B, 32 B) broadly separate; halter apex extending to about half length of tergite 1; legs (Fig. 34 B) with lateral margin of foreleg near midlength of wing strongly curved, sinuous; hind leg not visible at lateral margin of wing (Fig. 32 B); with apex of foreleg ventral to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 6 campaniform sensilla; anterolaterals — 1 short seta, 1 campaniform sensillum; dorsal setae (Figs. 29 B – C) — D- 1 - T short or moderately elongate thick seta, D- 2 - T campaniform sensillum, D- 4 - T campaniform sensillum or moderately elongate thick seta, D- 3 - T absent or campaniform sensillum; D 1 - T, D- 4 - T closely approximated or on short tubercle, D- 3 - T posteromedial to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 47 B) — 1 seta, 2 – 3 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: without pigmentation pattern, segments 2, 3 equally or nearly equally wide, segments with undivided, thin and thick, straight or recurved setae, with rounded to pointed, short tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Figs. 72 B – E) elongate, fleshy, with short tubercles, terminal processes widely separated basally, each projecting posterolaterally to nearly laterally, tapering to rounded, lobe-like apex; sensilla: tergite 1 (Fig. 47 B) with 2 setae, 4 campaniform sensilla, including 2 lateral sensilla, D- 2 - I a seta or campaniform sensillum, D- 3 - I absent, D- 7 - I absent; segment 4 (Fig. 55 B) — D- 2 - IV thick spine on short tubercle, D- 3 - IV absent; D- 5 - IV, D- 8 - IV short, thick, straight or recurved spines; D- 5 - IV, D- 7 - IV on separate, short tubercles, D- 4 - IV, D- 8 - IV on single tubercle, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 8 - IV, D- 7 - IV; L- 1 - IV short, thick seta on short tubercle, well anterior of posterior lateral setae, L- 2 - IV, L- 3 - IV thick setae on short tubercles, V- 5 - IV, V- 7 - IV on short tubercles, V- 6 - IV absent; segment 8 without D- 3 - VIII, with L- 1 - VIII; segment 9 (Figs. 72 B-E) — all with D- 1 - IX seta, D- 2 - IX, D- 3 - IX, D- 5 - IX campaniform sensilla, following in at least some: L- 1 - IX, L- 2 - IX, L- 3 - IX, V- 1 - IX, V- 2 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD0B3051FD5C1951485CE3F1.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Leptoconops is known from 150 species from every Region worldwide (Borkent 2014) but are restricted to wet or damp sand or sandy soil (generally freshwater or marine beaches protected from wave action), seepage areas in desert oases, wet margins of salt flats, cracked clay soils, halomorphic, calcareous soil, and the margin of vernal ponds in xeric areas.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD0B3051FD5C1951485CE3F1.taxon	discussion	TAXONOMIC DISCUSSION: Only 14 species of Leptoconops are known as pupae (Tables 2 – 3). Gutsevich & Glukhova (1970) provided a key to three species known from the former USSR. Painter (1927) provided first somewhat detailed description of a Leptoconops (L. bequaerti (as Leptoconops sp. )) from Honduras and Ishigami (1959) provided similar details for L. nipponensis in Japan. Clastrier (1972) in a very detailed morphological study, described and illustrated the pupa of L. gallicus (as L. kerteszi), identifying all parts, describing each sensillum and describing the differences between males and females. His description differs in some details from the generic description here. He shows 1 short seta and 4 campaniform dorsolateral cephalic sclerite sensilla, 1 ocular (campaniform sensillum), 3 anteromedials (campaniform sensilla) and only 1 seta and 1 campaniform sensillum on segment 9. In addition, I record further additional sensilla here. Although some differences may be due to intraspecific variation, it should be noted that the differences are likely due to the difficulty in discerning very small setae or campaniform sensilla, particularly in areas of uneven cuticle. This was the case with Clastrier's (1972) study of L. gallicus, for which I was able to reexamine the species. An SEM study of a species of Leptoconops would be invaluable in better understanding this member of the earliest lineage of Ceratopogonidae, keeping in mind that many of the features of this genus are strongly autapomorphic. Clastrier & Wirth (1978) provided further details of some Nearctic species, showing that at least some features vary within the genus. Leptoconops pupae have reduced numbers of sensilla on the thorax and 3 – 9 sensilla on abdominal segment 9 and naming these was somewhat arbitrary. However, at least L. linleyi had two campaniform sensilla on the terminal process that are likely homologous to the two found on most other Ceratopogonidae. I examined only one, poorly preserved male pupa specimen of Leptoconops (Styloconops) spinosifrons (with a pharate adult) and this species warrants particular attention, especially as one character state, the lack of recurved abdominal spines (see character 49) may indicate the subgenus is the sister group to other members of the genus. As illustrated by Laurence & Mathias (1972), there may be only one dorsal mesothoracic seta but I could not confidently confirm this. If so, it would be uniquely derived within the Culicomorpha. Dzhafarov (1962, 1964) described a pupal exuviae as L. bezzii but it clearly is misidentified to genus. In overall habitus it is somewhat similar to Allohelea but details differ from the only known species in that genus. The numerous thoracic setae illustrated, if accurate, suggest it is not even in the Ceratopogonidae. Dzhafarov (1962) does not mention rearing methods nor provides any other indication that the pupal exuviae was definitely associated with either larvae or adults and Dzhafarov (1964) noted the presence of only one pupal exuviae collected from wet soil. The eggs, larvae, and adults described by Dzhafarov (1962) are certainly correctly identified as Leptoconops.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD0B3051FD5C1951485CE3F1.taxon	materials_examined	MATERIAL EXAMINED: L. americanus: 1 pupa, 4.4 mi. N. Bodie, Mono County, California, USA, 23 - VI- 1965 (USNM); 1 pupa, Saltair, Salt Lake County, Utah, USA, 23 - V- 1969 (USNM). L. asilomar: 1 pupa, 2 pupal exuviae, ¾ mi. S. Asilomar, Monterey County, California, USA, 21 - II- 1964 (USNM); 3 pupae, as previous locality, 12 - III- 1964 (USNM); 1 pupa, as previous locality, 19 - III- 1964 (USNM). L. bequaerti: 1 pupal exuviae, Great Island, Yarmouth, Cape Cod, Massachusetts, USA, 28 - VI- 1961 (USNM); 2 pupal exuviae, Ochos Rios, St. Ann, Jamaica, 1969 – 1970 (USNM); 1 pupal exuviae, Puerto Castilla, Honduras, 1926 (USNM). L. gallicus: 2 pupal exuviae, La Figueirasse, France, 8 - VI- 1972 (USNM). L. kerteszi: 2 pupal exuviae, La Figueirasse, France, V- 1971 (USNM); 6 pupal exuviae, 4 pupal exuviae (in glycerin), Between River Ombrone and Principina a Mare (province of Grosseto), Italy, Spring-Summer, 1999 (CNCI). L. linleyi: 1 pupa, 2 pupal exuviae, Great Island, Yarmouth, Cape Cod, Massachusetts, USA, 28 - VI- 1961 (USNM); 5 pupal exuviae, Sebastian, Indian River County, Florida, USA, X- 1972 (USNM); 3 pupae, as previous locality, 15 - X- 1994 (CNCI); 4 pupae (in glycerin), as previous locality, 15 - X- 1994 (CNCI). L. pugnax: 2 pupal exuviae, La Figueirasse, France, 8 - VI- 1972 (USNM). L. spinosifrons: 1 pupa (with pharate adult), Seychelles Islands, 1970 (USNM). L. whitseli: 9 pupae (of paratypes), 2 pupal exuviae (of paratypes), Pebble Beach, Monterey County, California, USA, 11 - VIII- 1964 (USNM).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD09305CFD62189B4ECFE319.taxon	description	(Figs. 2 A – B, 8 D – H, 9 A – H, 10 A – H, 11 A – C, 14 C – D, 18 C – D, 23 C – D, 29 D – F, 32 C, 34 C, 42 E – R, 47 C – D, 55 C, 56 A, 72 F – I)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD09305CFD62189B4ECFE319.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae without a prothoracic extension (Figs. 23 C – D), with the labium not divided medially by an elongate suture (as is present in Fig. 23 F) and with the terminal processes either closely abutting basally (Figs. 8 D – H, 9 A – H, 10 A – H, 11 A – D, 72 F – H) or the terminal process spicules (if present) directed posteriorly to posterolaterally (not anteriorly); combination of retaining larval exuviae on the abdomen or with very elongate tubercles on the thorax (Figs. 9 A, C, F, 10 A – E) and / or abdomen, and the terminal process with spicules (if present) directed posteriorly to posterolaterally (not anteriorly) is unique to many species.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD09305CFD62189B4ECFE319.taxon	description	DESCRIPTION: Habitus as in Figs. 8 D – H, 9 A – H, 10 A – H, 11 A – C. Total length = 1.69 – 3.03 mm. With (as in Fig. 11 F) or without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face (Fig. 14 D). Ecdysial tear to anterior margin of base of antenna (Figs. 14 C – D, 79 B); along posterolateral portion of eye. Head: Dorsal apotome (Figs. 18 C – D), without ventral line of weakness, with or without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 B) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Figs. 23 C – D) with mandible, lacinia well-developed, not overlapping apically; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 34 C) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Figs. 18 C – D) — absent or 1 very short seta; dorsolateral cephalic sclerite sensilla — absent or 1 short seta, 1 campaniform sensillum; clypeal-labrals (Figs. 23 C – D) — 1 seta, 1 campaniform sensillum; oculars (Figs. 23 C – D) — 1 campaniform sensillum. Thorax: Prothoracic extension (Figs. 23 C – D) absent; mesonotum with or without elongate tubercles, extending posteromedially, completely dividing metathorax medially (Figs. 47 C – D); respiratory organ (Figs. 42 E – R) length / width = 1.05 – 4.75, highly variable, knob-like to elongate, slender to nearly spherical, with or without posterior basal swelling or lobe, somewhat flattened laterally or circular in cross-section, with pores closely abutting at apex of respiratory organ, arranged in single line of varying shapes or two widely spaced rows, outer surface smooth or with spicules, without or with short pedicel, base without 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. 34 C) with apical tubercle or angle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (Figs. 10 C, 32 C) broadly to narrowly separate; halter apex abutting anterolateral edge of tergite 2; legs (Fig. 34 C) with lateral margin of foreleg near midlength of wing slightly sinuous to evenly curved; hind leg visible at lateral margin of wing (Fig. 32 C); with apex of foreleg moderately to well anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 0 – 2 campaniform sensilla; anterolaterals — with or without 1 short or elongate seta or 1 campaniform sensillum; dorsal setae (Figs. 29 D – F) — D- 1 - T seta, D- 2 - T seta present or absent, D- 3 - T campaniform sensillum; D- 2 - T, D- 3 - T on single tubercle in some, or closely associated (if D- 2 - T present); supraalar 2 — campaniform sensillum; metathoracics (Figs. 47 C – D) — 0, 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 2 – 8 with spines or thick tubercles or segments 1 – 5 with branched or thick spined elongate tubercles, tubercles rounded to pointed, tergites or sternites entire, each without membranous disc; segment 9 (Figs. 72 F – I) highly variable in shape, terminal processes widely separated to closely approximated basally, each projecting posteriorly to laterally, tapering to pointed apex or tapering near apex to point; sensilla: tergite 1 (Figs. 47 C – D) with 3 – 5 setae, 1 – 2 campaniform sensilla, including 1 – 2 lateral sensilla, D- 2 - I, D- 3 - I (if present) well separated, D- 7 - I absent or situated posteriorly near D- 8 - I; segment 4 (Figs. 55 C, 56 A) — D- 2 - IV peg-like to slender seta, D- 3 - IV slender seta, without tubercles or very short separate ones; D- 8 - IV, when present, a moderately elongate seta; D- 5 - IV present or absent, D- 7 - IV, D- 9 - IV absent; D- 4 - IV without to on moderately elongate tubercle, posterior dorsal sensilla with D- 4 - IV, D- 8 - IV lateral to each other or D- 8 - IV absent, L- 1 - IV short to elongate seta present on short to long tubercle, close to L- 2 - IV, L- 2 - IV, L- 3 - IV, L- 4 - IV on separate rounded to elongate tubercles, in some with L- 2 - IV, L- 3 - IV closely approximated, V- 5 - IV, V- 6 - IV, V- 7 - IV short to moderately elongate setae, V- 5 - IV present or absent, on separate tubercles or V- 6 - IV, V- 7 - IV on single tubercle; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Figs. 72 F – I) — D- 5 - IX campaniform sensillum, D- 6 - IX campaniform sensillum or short or elongate seta.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD09305CFD62189B4ECFE319.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Forcipomyia is known from 1150 species in 35 subgenera from every Region worldwide (Borkent 2014). The larvae and pupae are found in many semi-terrestrial to moist terrestrial habitats, including under loose bark of logs, under decaying leaves and manure, decomposing vegetable matter and fungi, moss on soil, rotting wood or rocks, sap at tree wounds, in and above the water of various phytotelmata, mud along streams, bogs, and shallow water ponds.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD09305CFD62189B4ECFE319.taxon	discussion	TAXONOMIC DISCUSSION: There are 146 species of Forcipomyia, in 14 subgenera, known as pupae (Tables 2 – 3). The pupae of this genus vary tremendously morphologically and are rich in numerous derived features, including overall body form, chaetotaxy, shape of respiratory organs and more. The group is ripe for a cladistic analysis and combined with their various behaviours and diverse niches, are certain to provide a rich opportunity to examine diversification within the genus. A few features were put into a phylogenetic context by Chan & LeRoux (1971 c). Although earlier work had shown that pupae of Forcipomyia have a variety of interesting modifications (e. g. Long 1902, Speiser 1910, Carter 1919, Rieth 1915), it was not until L. G. Saunders (1924, 1925, 1957, 1959, 1964), in a series of remarkable papers, studied the group that the wealth of features and their importance in their classification became apparent. He described exemplars from the 10 available subgenera (out of 14 for which immatures are known today), showing that the larvae, pupae and adults had congruent and distinctive features, indicating a solid basis for their classification (although derived features were not interpreted). He provided a consistent format for the description of the pupae and laid the groundwork for all future studies. Work by Chan & Saunders (1965), Chan & LeRoux (1965, 1970, 1971 a, 1971 b) and Chan & Linley (1989) added significantly with further detailed descriptions of Forcipomyia pupae. The pupae are distinctive enough that a key to the subgenera is possible (see above). Further differences exist at the species level and early work indicated these also could be keyed (Malloch 1915 — Illinois; Lenz 1934 — Europe; Wirth & Howarth 1982 — Hawaiian F. (Euprojoannisia); Lewańczyk et al. 2010 — European species of F. (Forcipomyia )). Wirth & Howarth (1982), Chen et al. (1980), Spinelli et al. (2995) and Marino et al. (2010) provided SEM photomicrographs of pupae of Forcipomyia. I initially thought that the subgenera F. (Lepidohelea) and F. (Schizoforcipomyia) could be separated on the basis of the relative size of the medial tubercle on the dorsal apotome. Previous descriptions of species of F. (Lepidohelea) show these to be long. The only species of F. (Schizoforcipomyia) to be described for this feature is F. borbonica by Chan & LeRoux (1971 b, as F. petersoni), who illustrate the pupa as having a short, rounded tubercle on the middle of the dorsal apotome, although in the text they indicate this to be " elongate ". I examined the two paratype pupal exuviae of F. petersoni and found both to have an elongate tubercle, indistinguishable from those of species of F. (Lepidohelea). Pupae of Forcipomyia have reduced numbers of thoracic sensilla, as well as fewer on the dorsum of the abdominal segments and the exact naming of these was uncertain. D- 3 - T and D- 4 - IV are likely correct as unique campaniform sensilla (in these areas).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD09305CFD62189B4ECFE319.taxon	materials_examined	MATERIAL EXAMINED: F. aeria: 2 pupal exuviae (of paratype), Mayaquez, Puerto Rico, 18 - III- 1953 (USNM). F. anabaenae: 1 pupal exuviae (of paratype), Singapore, 25 - III- 1962 (CNCI). F. antiguensis: 1 pupal exuviae (of paratype), Falmouth Harbour, Antigua, 6 - V- 1953 (CNCI). F. ashantii: 1 pupal exuviae, Tafo, Ghana, 17 - V- 1963 (ANIC); Berger's Hall, Hazyview, East Transvaal, South Africa, 3 - XII- 1973 (ANIC). F. attenuata: 1 pupal exuviae (of paratype), La Lola, Costa Rica, 2 - VI- 1956 (CNCI). F. bipunctata: 7 pupa, 23 pupal exuviae, 4 mi. W., 3 mi. S Schuyler, Nebraska, USA, various dates, 1990 - 1991 (CNCI). F. blantoni: 2 pupal exuviae, Turrialba, Costa Rica, 2 - IX- 1964 (USNM); 2 pupal exuviae, as previous locality, 26 - XI- 1964 (USNM); 1 pupal exuviae, as previous locality, 21 - XII- 1964 (USNM); 1 pupal exuviae, as previous locality, 29 - XII- 1964 (USNM). F. borbonica: 1 pupal exuviae (labeled as paratype of an undescribed species F. lerouxi), Wallace Way, Singapore, 1961 (LEMQ). F. brasiliensis: 1 pupal exuviae, Maracas, Trinidad, 18 - VII- 1957 (USNM). F. bromeliae: 1 pupal exuviae (of paratype), Rio de Janeiro, Brazil, 23 - VIII- 1923 (CNCI). F. bystraki: 1 pupal exuviae, Calvert, Cecil County, Maryland, USA, 29 - III- 1969 (USNM); 2 pupal exuviae, Headwaters, Highland County, Virginia, USA, VI- 1969 (USNM). F. caerulea: 1 pupal exuviae (of paratype), Rio de Janeiro, Brazil, 8 - VIII- 1923 (CNCI). F. canadensis: 2 pupae (of paratypes), Pike Lake, Saskatchewan, Canada, X- 1957 (USNM). F. caribbeana: 1 pupal exuviae (of paratype), Trinidad, 12 - V- 1953 (CNCI). F. cerifera: 1 pupal exuviae, Rio de Janeiro, Brazil, 1 - VIII- 1923 (CNCI). F. cornuta: 1 pupal exuviae (of paratype), Costa Rica, 4 - VI- 1956 (CNCI). F. crinita: 1 pupal exuviae (of paratype), Emma Lake, Saskatoon, Saskatchewan, Canada, 19 - IV- 1957 (CNCI). F. edmistoni: 1 pupal exuviae (of paratype), Allegany State Park, New York, USA, 28 - V- 3 - VI- 1963 (USNM); 1 pupal exuviae (of paratype), Bittinger 4 - H Campground, Garrett County, Maryland, USA, 5 - V- 1960 (USNM); 1 pupa (of paratype), 1 pupal exuviae (of paratype), Patuxent Wildlife Refuge, Maryland, USA, 28 - VII- 1979 (USNM). F. elegantula: 1 pupal exuviae, Saskatoon, Saskatchewan, Canada, 3 - IX- 1950 (USNM); 1 pupa, 1 pupal exuviae, Plummer's Island, Maryland, USA, 3 - VI- 1976 (USNM); 1 pupal exuviae, as previous locality, 10 - VI- 1976 (USNM); 1 pupa, 2 pupal exuviae (in glycerin), as previous locality, 10 - VI- 1976 (WLGC). F. flavescens: 1 pupal exuviae (of paratype), Luna, P. I., 25 - X- 1961 (CNCI). F. harpegonata: 1 pupa (of paratype), Puerto Rica, 23 - I- 1953 (USNM). F. herediae: 1 pupal exuviae (of paratype), Siquirres, Costa Rica, 1 - VI- 1956 (USNM). F. hutsoni: 1 pupa (of paratype), South Island, Aldabra, 1 - 17 - II- 1968 (USNM). F. hygrophila: 3 pupae, Sidney, British Columbia, Canada, 20 - VIII- 1947 (USNM); 2 pupal exuviae, no locality data (USNM). F. intermedia: 1 pupal exuviae (of paratype), nr. Siquirres, Costa Rica, 15 - V- 1956 (CNCI). F. jocosa: 1 pupal exuviae (of paratype), Mt. St. Benedict, Trinidad, 13 - V- 1953 (CNCI). F. lesliei: 2 pupae, Clark Hall, Dominica, 9 - I- 1965 (USNM); 1 pupa (of paratype of F. bicolor), 2 pupal exuviae (of paratypes of F. bicolor), Mayaquez, Puerto Rico, 3 - II- 1953 (USNM). F. luteigenua: 1 pupa (of paratype), 1 pupal exuviae (of paratype), Finca la Tigra, nr. La Virgen, Heredia Province, Costa Rica, 12 - XI- 1981 (USNM); 1 pupa (of paratype), as previous locality, 17 - XI- 1981 (USNM); 1 pupa (of paratype), as previous locality, 30 - VII- 1982 (USNM). F. malayae: 2 pupal exuviae (of holotype, paratype), Cameron’s Highlands, Malaysia, 29 - I- 1934 (CNCI). F. mortuifolii: 1 pupal exuviae, no locality data (USNM). F. musae: 2 pupal exuviae, km 123 Br 174, Manaus, Amazonas, Brazil, 29 - VII- 2005 (CNCI). F. nodosa: 1 pupal exuviae (of paratype), nr. Siquirres, Costa Rica, 15 - VII- 1956 (CNCI). F. oligarthra: 1 pupal exuviae (of paratype), Carcega Beach, Puerto Rico, 22 - II- 1953 (CNCI). F. pictoni: 1 pupa, 2 pupal exuviae, Siquirres, Costa Rica, 11 - VI- 1956 (USNM). F. pinicola: 2 pupal exuviae, no locality (USNM). F. pulchrithorax: 3 pupae, Cambridge, England, United Kingdom, 31 - VIII- 1922 (USNM). F. quasicornuta: 1 pupal exuviae (of paratype), nr. Siquirres, Costa Rica, 6 - V- 1050 (CNCI). F. seminole: 2 pupal exuviae, Hills, Puerto Rico, 6 - II- 1953 (USNM); 2 pupae, 2 pupal exuviae, Georgetown, Guyana, 20 - V- 1943 (USNM). F. sonora: 2 pupae, 2 pupal exuviae, Death Valley, Saratoga Springs, California, USA, 19 - II- 1955 (USNM). F. spinosa: 2 pupae (of paratypes), 1 pupal exuviae, Mayaquez, Puerto Rico, 3 - II- 1953 (USNM); 1 pupal exuviae (of paratype), as previous locality, 3 - II- 1953 (CNCI); 1 pupal exuviae, Ilheus, Bahia, Brazil, 20 - X- 1977 (USNM). F. terrestris: 1 pupal exuviae (of paratype), St. Augustine, Trinidad, 9 - VII- 1957 (CNCI); 1 pupa (of paratype), 3 pupal exuviae (of paratype), Las Hermanas, Trinidad, 9 - VII- 1957 (USNM). F. trinidadensis: 1 pupal exuviae (of holotype), St. Augustine, Trinidad, 3 - VII- 1957 (CNCI). F. tuberculata: 1 pupa, 2 pupal exuviae, Hacienda Theobroma, Siquirres, Costa Rica, 1 - VI- 1956 (USNM); 1 pupa, Bayano Field Station, Panama Province, Panama, VI- 1976 (USNM); 1 pupal exuviae (of paratype), Trinidad, 8 - V- 1953 (CNCI); 1 pupa (of paratype), 2 pupal exuviae (of paratypes), North Range, Trinidad, 8 - V- 1953 (USNM). F. usingeri: 3 pupae (of paratypes), Berkeley Hills, Alameda County, California, USA, X- 1947 (USNM). F. varicolor: 1 pupal exuviae (of paratype), Rio de Janeiro, Brazil, 1 - VIII- 1923 (CNCI). F. wirthi: 1 pupal exuviae (of paratype), Alum Rock Park, Santa Clara County, California, USA, 8 - IV- 1948 (USNM). F. (Phytohelea) nr. antiguensis: 3 pupal exuviae, 1 pupal exuviae (in glycerin), Riverwoods Field Laboratory, nr. Cornwall, Highlands County, Florida, USA, 13 - I- 1999 (CNCI). F. (Euprojoannisia) sp.: 1 pupal exuviae, 52 km S of airport, Iquitos, Loreto, Peru, 1 - II- 2003 (CNCI). F. (Phytohelea) sp.: 2 pupal exuviae, 2 pupal exuviae (in glycerin), Jardin Botanico, Rio Piedras, San Juan, Puerto Rico, 24 - X- 2006 (CNCI). F. sp.: 1 pupa, 1 pupal exuviae (in glycerin), Nebraska, USA (CNCI); 1 pupal exuviae, Rio Dantes, P. N. Barbilla, Cartago, Costa Rica, 10 - II- 2006 (CNCI); 2 pupal exuviae, Iquitos, Loreto, Peru, 1 - II- 2003 (CNCI); 1 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 19 - II- 1967 (BPBM); 1 pupal exuviae, as previous locality, 9 - 12 - XII- 1967 (BPBM).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD04305DFD6E1B7A4E47E583.taxon	description	(Figs. 11 D – F, 14 E, 18 E, 23 E, 29 G, 32 D, 34 D, 42 S – T, 47 E, 56 B, 72 J)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD04305DFD6E1B7A4E47E583.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with terminal process with at least some spicules directed anteriorly.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD04305DFD6E1B7A4E47E583.taxon	description	DESCRIPTION: Habitus as in Figs. 11 D – F. Total length = 1.41 – 3.44 mm. With (Fig. 11 F) larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face (Fig. 14 E). Ecdysial tear anterior or slightly medial to base of antenna (Figs. 14 E, 79 B); along posterolateral portion of eye. Head: Dorsal apotome (Fig. 18 E), without ventral line of weakness, with dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 B) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 23 E) 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. 34 D) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Fig. 18 E) — 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. 23 E) — 1 seta, 1 campaniform sensillum; oculars (Fig. 23 E) — 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 23 E) absent; mesonotum with long, bifurcating tubercles, extending posteromedially, completely dividing metathorax medially (Fig. 47 E); 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. 34 D) without apical tubercle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 32 D) broadly separate; halter apex abutting anterolateral edge of tergite 2; legs (Fig. 34 D) with lateral margin of foreleg near midlength of wing slightly sinuous to evenly curved; hind leg visible at lateral margin of wing (Fig. 32 D); 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. 29 G) — 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. 47 E) — 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. 72 J) 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. 47 E) with 3 setae, 2 campaniform sensilla, including 2 lateral sensilla, D- 3 - I absent, D- 7 - I absent; segment 4 (Fig. 56 B) — 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. 72 J) — D- 5 - IX campaniform sensillum, possibly D- 6 - IX present but inadequate specimens examined.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD04305DFD6E1B7A4E47E583.taxon	distribution	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.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD04305DFD6E1B7A4E47E583.taxon	discussion	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. 56 B) 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).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD04305DFD6E1B7A4E47E583.taxon	materials_examined	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).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD023046FD521ACB4E15E1CC.taxon	description	(Figs. 11 G, 14 F, 18 F – L, 23 F – G, 29 H – J, 32 E – F, 34 E, 43 A – G, 47 F, 56 C, 72 K – L, 73 A – G)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD023046FD521ACB4E15E1CC.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the labium clearly split medially by an elongate suture (or overlapping halves) (Figs. 23 F – 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.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD023046FD521ACB4E15E1CC.taxon	description	DESCRIPTION: Habitus as in Fig. 11 G. Total length = 1.63 – 4.16 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face (Fig. 14 F). Ecdysial tear not extending at all (Figs. 14 F, 79 C); posterolateral portion of eye. Head: Dorsal apotome (Figs. 18 F – L), with ventral line of weakness, with or without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 B) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Figs. 23 F – 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. 34 E) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Figs. 18 F – L) — 1 short to elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 2 small setae, 1 campaniform sensillum; clypeal-labrals (Figs. 23 F – G) — 2 minute setae; oculars (Figs. 23 F – G) — 0 – 1 seta, 1 campaniform sensillum. Thorax: Prothoracic extension (Figs. 23 F – G) absent; mesonotum with or without short tubercles, not extending posteromedially to extending posteromedially, completely dividing metathorax medially (Fig. 47 F); respiratory organ (Figs. 43 A – 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. 34 E) with apical tubercle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (Figs. 32 E – F) just separate; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 34 E) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Figs. 32 E – 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. 29 H – 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. 47 F) — 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. 72 K – L, 73 A – 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. 47 F) 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. 56 C) — 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. 72 K – L, 73 A – 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.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD023046FD521ACB4E15E1CC.taxon	distribution	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.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD023046FD521ACB4E15E1CC.taxon	discussion	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 (1934 a) also describes differences between species now considered synonyms (e. g. D. modesta with its synonyms D. inclusa and D. longipalpis). Mayer (1934 a) 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 (1934 c) 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 (1934 c) 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. 56 C). 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).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD023046FD521ACB4E15E1CC.taxon	materials_examined	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).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1E3045FD591EBC49C5E4E4.taxon	description	(Figs. 2 F, 3 A – C, 4 A – C, 5 A – B, 13 B, 15 A, 19 A – D, 24 A, 29 K – L, 32 G, 34 F, 43 H – N, 48 A, 57 A, 73 H)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1E3045FD591EBC49C5E4E4.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with a prothoracic extension extending from the palpus to the antenna (Fig. 24 A), the halter and hind leg slightly separated or barely touching (Figs. 3 B, 32 G), and the dorsal apotome relatively flat (Figs. 19 A – D) and without a central dome-like protuberance (bearing the sensilla) (as in Figs. 19 H – J) and with or without spicules but if with a lateral row of stout, pointed spicules then also with further stout spicules more medially.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1E3045FD591EBC49C5E4E4.taxon	description	DESCRIPTION: Habitus as in Figs. 3 A – C. Total length = 1.56 – 3.13 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face (Fig. 15 A). Ecdysial tear medial to base of antenna (Figs. 15 A, 79 D); along prothoracic extension. Head: Dorsal apotome (Figs. 19 A – D), without ventral line of weakness, with or without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (Fig. 13 B) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 24 A) with mandible, lacinia well-developed, overlapping; palpus extending equal to or posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 34 F) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Figs. 19 A – D) — 1 moderate to elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 2 setae, 1 campaniform sensillum; clypeal-labrals (Fig. 24 A) — 0 – 2 setae; oculars (Fig. 24 A) — 2 elongate setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 24 A) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially but with metathorax having medially split to extending posteromedially, completely dividing metathorax medially (Fig. 48 A); respiratory organ (Figs. 43 H – N) length / width = 3.82 – 8.13, elongate, slender, circular in cross-section, with pores closely abutting or slightly separated at apex of respiratory organ, arranged in single straight or curved row or two widely spaced rows, with or without additional, more basal pores, outer surface with or without annulations, with or without spicules, with elongate, slender pedicel, base with short, strung out posteromedial apodeme, membranous base of respiratory organ short to moderately elongate, tracheal tube straight to slightly curved along length, with spirals restricted to base, distally with reticulations or plates; wing (Fig. 34 F) with apical tubercle or angle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 32 G) just separate to just touching; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 34 F) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 32 G); 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 — 1 seta; anterolaterals — 2 setae, 1 campaniform sensillum; dorsal setae (Figs. 29 K – L) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T setae, D- 3 - T campaniform sensillum, D- 3 - T posterior to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 48 A) — 1 seta, 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: without pigmentation pattern or with tergites 1 – 7 with pair medially and pair anterolaterally (fading posteriorly) or tergites 2 – 7 with medial group of 3 and anterolateral pair, sternites 3 – 7 with medial stripe, anterolateral spot, or 2 pairs, fading posteriorly, 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 (Fig. 73 H) not strongly modified, terminal processes closely approximated basally, each projecting posteriorly to posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 48 A) 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- 2 - I, D- 3 - I; segment 4 (Fig. 57 A) — D- 2 - IV, D- 3 - IV short to moderately elongate setae on short tubercles; D- 5 - IV (rarely absent), D- 8 - IV, D- 9 - IV short to moderately elongate setae (D- 9 - IV absent in some), D- 5 - IV, D- 4 - IV, D- 7 - IV, D- 8 - IV, D- 9 - IV on short to moderately elongate, separate tubercles, in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 7 - IV, D- 8 - IV, D- 9 - IV; L- 1 - IV short seta on short to pointed tubercle, well anterior of posterior lateral setae, L- 2 - IV, L- 3 - IV, L- 4 - IV setae of variable length, on short to pointed tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV short to moderately elongate setae on rounded tubercles; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 73 H) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1E3045FD591EBC49C5E4E4.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Culicoides is known from 1343 species from every Region worldwide (Borkent 2014) but is notably absent from New Zealand. Immatures are present in a wide array of habitats but generally are associated with small and limited aquatic or very wet microhabitats. They are diverse and common in mud or decaying substrate on the margins of pools, swamps, ponds, streams and rivers, but are also found in such diverse habitats as damp or wet decomposing vegetation, wet leaf packs, manure, fungi, sap from wounds on trees, phytotelmata, seed husks, dead cacti, crab holes, springs, seeps, bogs, saline pools, fens, mangrove swamps, salt marshes, and river bottom (one species).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1E3045FD591EBC49C5E4E4.taxon	discussion	TAXONOMIC DISCUSSION: There are 226 species of Culicoides known as pupae (Tables 2 – 3). Some of these, however, are poorly described, or are questionably associated (in part, because some adults are difficult to identify accurately). Furthermore, some authors have described differences between species which are now considered synonyms, likely indicating problems either with identification of the adults or the real possibility that more than one species is present under that name (Thienemann 1928, Mayer 1934 a, Lenz 1934, Dzhafarov 1964). Many using this work will primarily want to identify pupae of Culicoides to genus. Aside from the features in the key and the diagnosis above, members of this genus have a nearly unique arrangement of the dorsolateral cephalic sclerite setae, with one long and the other quite small (in most species and difficult to see) to equally long. Pupae of Paradasyhelea also have this arrangement but they are relatively rare taxa restricted to Southern Hemisphere localities as well as the Olympic Peninsula in Washington, USA. In areas of sympatry, the arrangement of the spicules on the dorsal apotome will serve to distinguish the two genera. Nearly all other Ceratopogoninae have only one dorsolateral cephalic sclerite seta. Of the exceptions, Baeodasymyia, rare and restricted to the Neotropical Region, has two setae but also has a respiratory organ with a distinctive separate, short, expanded apical portion bearing pores in a single plane (Fig. 44 C), some Parabezzia (e. g. P. petiolata) have both setae very short and peg-like, and one specimen of Serromyia has both setae very short and slender. Members of Culicoides may also be recognized by a combination of the halter and hind leg slightly separated or barely touching (Figs. 3 B, 32 G) and D- 7 - I placed anteriorly on tergite 1, near D- 2 - I and D- 3 - I. There are various keys to species of pupae of Culicoides which include species from a given area or of a select species group, as follows (arranged geographically, New World to Old World, north to south): - Jones (1955) — Wisconsin, USA. - Jamnback (1965) — New York, USA; described the differences between subgenera and some species groups. - Blanton & Wirth (1979) — Florida, USA; described the differences between subgenera and some species groups. - Jones (1961) — to 13 Nearctic species. - Atchley (1970) — Nearctic species of C. (Selfia). - Jamnback & Wirth (1963) — eastern Nearctic species of obsoletus species group. - Kann (1980) — Nearctic species of guttipennis species group, noting differences of the species group with other groups of Culicoides. - Lamberson et al. (1992) — tree-hole species from eastern North America. - Thienemann (1928) — northern Europe. - Lenz (1934) — Europe. - Mayer (1934 a) — Europe. - Kettle & Lawson (1952) — British. - Gutsevich & Glukhova (1970) — former USSR - Glukhova (1989) — Russia. - Dzhafarov (1964) — Transcaucasus. - Brodskaya (1999) — eastern Russia and northeastern China - Howarth (1985) — Laos. - Kettle & Elson (1976, 1980), Elson-Harris & Murray (1992) — Australia. - Nevill (1969) — seven South African species. - Nevill and Dyce (1994) — similis species group (Africa). - Nevill et al. (2007) — imicola species group (Africa). All the authors above have shown that there are significant differences between species of Culicoides, allowing for their identification at that level. These studies speak to the diverse morphology within this huge genus and provide good evidence that the pupal stage could be an important basis for further understanding the habitats utilized by the immatures. Larvae are far more conservative in their structural divergence and are challenging to identify (e. g. Murphree & Mullen 1991). Dyce (1998) included pupal diagnoses for the subgenera and species groups of Australian Culicoides providing tentative characterizations (results were from a workshop). Nevill et al. (2007) described the variation of some pupal features in the imicola species group and showed how pupae may provide morphological evidence of differences that is more obscure in adults of some species of Culicoides. Geographic variation of pupal features was described by Atchley (1971 a) for three Nearctic species and sexual differences were noted in the shape of the dorsal apotome by Atchley (1971 b). Spătaru (1971) described intraspecific variation of various pupal features of C. stigma. Of the numerous features that vary between species and which were noted or utilized by previous authors, it is worth noting that the relative arrangement of sensilla D- 4 - I, D- 8 - I and D- 9 - I varies considerably within Culicoides, with many species having D- 8 - I and D- 9 - I lateral to D- 4 - I. The diagnosis for pupae of Culicoides as given above is generally more simple than may be initially apparent. The combination of a well-developed prothoracic extension and the halter and hind leg slightly separated or barely touching distinguishes all Culicoides except for the Holarctic and rarely collected Ceratopogon, which have a unique dome-like protuberance on their dorsal apotome, and Paradasyhelea, restricted to the Southern Hemisphere (New Zealand, Australia, New Caledonia, Argentina, Chile) and the Olympic Peninsula of Washington, USA (the latter rare) and which have a lateral row of stout, pointed spicules and further stout spicules more medially on the dorsal apotome. The identification of the larva and pupa of C. punctatus and C. nipponensis respectively in Table 2 from Kitaoka (1998) was provided by Shigeo Kitaoka (pers. comm.).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1E3045FD591EBC49C5E4E4.taxon	materials_examined	MATERIAL EXAMINED: C. annettae: 1 pupal exuviae, 2 km NE Tarcoles, Costa Rica, 17 - XII- 1993 (CNCI). C. annuliductus: 1 pupal exuviae (paratype), Bayano field station, Panama Province, Panama, VI 1976 (USNM). C. arboricola: 1 pupal exuviae (paratype), Gwynns Falls Park, Baltimore, Maryland, USA, VII- 1931 (USNM). C. baueri: 1 pupal exuviae, Tom’s Creek, Montgomery County, Virginia, USA, 27 - V- 1976 (VPIC); 1 pupal exuviae, Cedar Run, Montgomery County, Virginia, USA, 6 - IX- 1971 (VPIC). C. bayano: 1 pupal exuviae (paratype), Bayano field station, Panama Province, Panama, VI 1976 (USNM). C. bergi: 1 pupal exuviae (paratype), Ringwood Game Preserve, Tompkins County, New York, USA, 10 - VII- 1970 (USNM). C. bermudensis: 1 pupal exuviae, Hacks Neck, Acomack County, Virginia, USA, 31 - VII- 1976 (VPIC). C. bickleyi: 1 pupal exuviae (of holotype), Cranesville Swamp, Garrett County, Maryland, USA, 6 - V- 1960 (USNM). C. brookmani: 1 pupal exuviae (paratype), Arroyo Seco R. S., Monterey County, California, USA, 1 - VII- 1948 (USNM). C. californiensis: 1 pupal exuviae (paratype), Bakersfield Kern County, California, USA, V- 1954 (USNM). C. chaverrii: 13 pupal exuviae (of holotype, paratypes), Costa Rica, various localities, dates as recorded by Spinelli and Borkent (2004), (CNCI, INBC). C. crepuscularis: 1 pupal exuviae, Pondapas Road, Montgomery County, Virginia, USA, 3 - IX- 1976 (VPIC). C. defoliarti: 1 pupal exuviae (paratype), Southwestern Research Station, Portal, Arizona, USA, 5 - 9 - VI- 1972 (USNM). C. denningi: 1 pupal exuviae (paratype), Saskatoon, Saskatchewan, Canada, VIII- 1947 (USNM). C. denticulatus: 24 pupal exuviae (in glycerin), Spanish Lake, 6 km E of Falkland, 50 ° 29.12 N 119 ° 28.07 W, BC, Canada, 27 – 28 - V- 2008 (CNCI). C. dicrourus: 2 pupal exuviae, Florida, Siquirres, Limon, Costa Rica, 12 - III- 2006 (INBC). C. filiductus: 1 pupal exuviae (paratype), Bayano field station, Panama Province, Panama, VI 1976 (USNM). C. guttipennis: 4 pupal exuviae, Ottawa, Ontario, Canada, 1973 (CNCI); 1 pupal exuviae, lab colony, W. Knausenberger, 30 - V- 1975 (VPIC). C. haematopotus: 1 pupal exuviae, 10 km W of Old Chelsea, Quebec, Canada, 15 - VII- 1986 (CNCI). C. heliconiae: 2 pupal exuviae, Florida, Siquirres, Limon, Costa Rica, 12 - III- 2006 (INBC). C. jacksoni: 1 pupal exuviae (paratype), Cedar Creek Canyon, Lincoln County, New Mexico, USA, 13 - VI- 1968 (USNM). C. jamnbacki: 1 pupal exuviae (of holotype), Huntinton Marsh, Newcomb, New York, 14 - V- 1959 (USNM). C. kettlei: 1 pupal exuviae (paratype), Big Horn Drive, Riverside County, California, USA, 22 - IX- 1988 (USNM). C. loisae: 1 pupal exuviae, 10 km W of Old Chelsea, Quebec, Canada, 15 - VIII- 1986 (CNCI). C. neofagineus: 1 pupal exuviae (paratype), Pinery Canyon, Chiricahua Mountains, Cochise County, Arizona, USA, 10 - VII- 1958 (USNM). C. nubeculosus: 15 pupae, 20 pupal exuviae, from colony at Institute for Animal Health, Pirbright Laboratory, Great Britain. C. phlebotomus: 8 pupal exuviae, 3 km E of Cahuita, Costa Rica, 30 - X- 1993 (CNCI). C. piliferus: 1 pupal exuviae, Elliot Knob, Augusta County, West Virginia, USA, 5 - VI- 1977 (VPIC); 1 pupal exuviae, Little Stoney Creek, Giles County, Virginia, USA, 3 - VII- 1977 (VPIC). C. scanloni: 1 pupal exuviae (of holotype), Alexandria, Virginia, USA, 2 - VI- 1951 (USNM). C. sonorensis: 10 pupae, 10 pupal exuviae, from colony at Laramie, Wyomying, USA (CNCI). C. spinosus: 1 pupal exuviae (paratype), Roland Park, Baltimore, Maryland, USA, V- 1930 (USNM). C. stellifer: 1 pupal exuviae, Watoga State Park, Pocahontas County, Virginia, USA, 1976 (VPIC). C. variipennis: 4 pupal exuviae, nr. Alfred, Ontario, Canada, 30 - IV- 1985 (CNCI). C. nr. crepuscularis: 3 pupal exuviae, Ohanapecosh campground turnoff, Ranier National Park, Washington, USA, 13 - VI- 2008 (CNCI); 1 pupal exuviae, 9 km W of Okanogan, Washington, USA, 13 - VI- 2008 (CNCI). C. sp.: 1 pupal exuviae, Salmon Arm, British Columbia, Canada, 7 - VI- 1988 (CNCI); 1 pupal exuviae, 3 km E of Salmon Arm, British Columbia, Canada, 9 - VI- 1988 (CNCI); 1 pupal exuviae (in glycerin), 3 km E. Carp, Ontario, Canada, 9 - VI- 1986 (CNCI); 10 pupal exuviae, 17 km N of Sedona, Arizona, USA, 11 - V- 1987 (CNCI); 4 pupal exuviae, 20 km NE of Tuscon, Arizona, USA, 4 - V- 1987 (CNCI); 1 pupal exuviae, Res. Sta., Stream, Newcomb, New York, USA, 6 - VI- 1958 (NYSM); 2 pupal exuviae, Pleasant Lake, Pierce County, North Dakota, USA, VI- 1969 (USNM); 3 pupal exuviae, Iquitos plantation, Loreto, Peru, 1 - II- 2003 (CNCI); 5 pupal exuviae, Chittering Road, Bindoon, Western Australia, Australia, 27 - X- 1985 (ANIC); 34 pupal exuviae, Bindoon, Western Australia, Australia, 27 - X- 1985 (ANIC); 16 pupal exuviae, Bindoon (as Bundoon), Australia, 27 - X (ANIC); 11 pupal exuviae, Darban, Western Australia, Australia, 28 - X- 1985 (ANIC); 5 pupal exuviae, as previous locality, 29 - X- 1985 (ANIC); 3 pupal exuviae, 5 km W of Darban, Western Australia, Australia, 29 - X- 1985 (ANIC); 1 pupal exuviae, Collie Road, W. Darban, Western Australia, Australia (ANIC); 1 pupal exuviae, Gap Creek, tributary of Fitzroy River, Kimberley, Western Australia, Australia (ANIC); 4 pupal exuviae, Lady Carrington Drive, Royal National Park, New South Wales, Australia (ANIC); 1 pupal exuviae, Careel Bay, New South Wales, Australia, 20 - IX- 1966 (ANIC); 1 pupal exuviae, no locality, 9 - VI- 1971 (ANIC).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1D3043FD6A1D2149A7E3D4.taxon	description	(Figs. 19 E – G, 24 B, 29 M, 35 A, 43 O, 48 B, 57 B, 73 I)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1D3043FD6A1D2149A7E3D4.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with a prothoracic extension extending from the palpus to the antenna (Fig. 24 B), the halter and hind leg are slightly separated or barely touching (as in Fig. 32 G), and the dorsal apotome (Figs. 19 E – G) has a lateral row of stout, pointed spicules and without further stout spicules more medially.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1D3043FD6A1D2149A7E3D4.taxon	description	DESCRIPTION: Total length = 1.50 – 2.09 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face (as in Fig. 15 A). Ecdysial tear medial to base of antenna (as in Figs. 15 A, 79 D); along prothoracic extension. Head: Dorsal apotome (Figs. 19 E – G), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 B) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 24 B) with mandible, lacinia well-developed, overlapping; palpus extending equal to or just posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 35 A) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Figs. 19 E – G) — 1 moderate to elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 2 setae, 1 campaniform sensillum; clypeal-labrals (Fig. 24 B) — 2 elongate setae; oculars (Fig. 24 B) — 2 elongate setae. Thorax: Prothoracic extension (Fig. 24 B) wide, welldeveloped, extending from palpus to antenna; mesonotum with short tubercles, extending posteromedially, completely dividing metathorax medially (Fig. 48 B); respiratory organ (Fig. 43 O) length / width = 6.50 – 7.06, elongate, slender, circular in cross-section, with pores closely abutting or slightly separated at apex of respiratory organ, arranged in single row, with additional, more basal pores, outer surface with annulations, without other surface modifications, with elongate, slender pedicel, base with short posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, distally smooth or plates; wing (Fig. 35 A) with apical tubercle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 32 G) just separate; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 35 A) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 32 G); 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 — 1 seta; anterolaterals — 2 setae, 1 campaniform sensillum; dorsal setae (Fig. 29 M) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T setae, D- 3 - T campaniform sensillum, D- 3 - T posterior to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 48 B) — 1 seta, 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: without pigmentation pattern, 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 (Fig. 73 I) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 48 B) with 7 setae, 1 or 2 campaniform sensilla, including 3 lateral sensilla, D- 2 - I, D- 3 - I closely approximated, D- 7 - I situated posteriorly near D- 8 - I; segment 4 (Fig. 57 B) — D- 2 - IV, D- 3 - IV short to moderately elongate setae on short tubercles; D- 5 - IV, D- 8 - IV, D- 9 - IV short setae, D- 5 - IV, D- 4 - IV, D- 7 - IV, D- 8 - IV, D- 9 - IV on short, separate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 7 - IV, D- 8 - IV, D- 9 - IV; L- 1 - IV short seta on short tubercle, well anterior of posterior lateral setae; L- 2 - IV, L- 3 - IV, L- 4 - IV moderately elongate setae, on short to pointed tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV short to moderately elongate setae on rounded tubercles; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 73 I) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1D3043FD6A1D2149A7E3D4.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Paradasyhelea is known from 11 species in the Southern Hemisphere from New Caledonia, Australia, New Zealand, Chile and Argentina and (one species) in the Olympic Peninsula of the northwest USA (Borkent 2014). Immatures have been found in the sandy, loamy or muddy margins of creeks or rivers, bogs, or swamps. There is one record from the muddy margin of a lake (Kettle & Elson 1975 a).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1D3043FD6A1D2149A7E3D4.taxon	discussion	TAXONOMIC DISCUSSION: Three species of Paradasyhelea are known as pupae, all from Australia (Tables 2 – 3). Elson-Harris & Kettle (1985 a) provide a key to these species (republished by Elson-Harris & Murray (1992 )) and suggested some features which would distinguish Paradasyhelea pupae from those of Culicoides but I could not confirm these differences as being consistently different between the two genera.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1D3043FD6A1D2149A7E3D4.taxon	materials_examined	MATERIAL EXAMINED: P. albipunctata: 6 pupal exuviae (of paratypes), Kiandra, New South Wales, Australia, 19 - XII- 1956 (USNM); 1 pupal exuviae (of paratype), Oxford Falls, New South Wales, Australia, 6 - XII- 1956 (USNM), 1 pupal exuviae, as previous locality, 10 - XI- 1956 (USNM); 1 pupal exuviae (of paratype), McCarr's Creek, New South Wales, Australia, 20 - IX- 1956 (USNM); 2 pupal exuviae (of paratype), as previous locality, 11 - XI- 1956 (ANIC); 1 pupal exuviae, Colo Vale, New South Wales, Australia, 17 - I- 1957 (USNM); 1 pupal exuviae (of paratype), Middle Creek, Narrabeen, New South Wales, Australia, 4 - XI- 1956 (ANIC); 1 pupal exuviae (of paratype), as previous locality, 8 - IX- 1956 (ANIC). P. minuta: 1 pupal exuviae (of paratype), South Creek, Deewhy, New South Wales, Australia, 27 - IX- 1956 (USNM); 2 pupal exuviae (of paratypes), Middle Creek, Narrabeen, New South Wales, Australia, 12 - IX- 1956 (USNM); 1 pupal exuviae (of paratype), Galston Gorge, New South Wales, Australia, 6 - IX- 1956 (USNM); 1 pupal exuviae, Colo Vale, New South Wales, Australia, 7 - III- 1957 (ANIC). P. sp: 18 pupal exuviae, S. Manjimup, Western Australia, Australia, 30 - X- 1985 (ANIC); 3 pupal exuviae, Gap Creek, Mittagong, New South Wales, Australia, 10 - II- 1966 (ANIC); 2 pupal exuviae, McCarr's Creek, New South Wales, Australia, 8 - II- 1966 (ANIC); 1 pupa (in glycerin), Borlase Stream, Lake Rotoiti, St. Arnaud, Nelson Lakes National Park, 41 ° 48 ' 52 " S 172 ° 51 ' 55 " E, South Island, New Zealand, 1 – 2 - II- 2000 (CNCI).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1B3040FD61187C4E88E7C4.taxon	description	(Figs. 13 C, 15 B, 19 H – J, 24 C, 29 N, 32 H, 35 B, 43 P – Q, 48 C, 57 C, 58 A, 73 J)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1B3040FD61187C4E88E7C4.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the dorsal apotome with a pronounced central dome from which both dorsal apotome setae arise (Figs. 19 H – J); also unique in having an inner tracheal tube of the respiratory organ strongly hooked apically, appearing as an inverted " J " (Figs. 43 P – Q).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1B3040FD61187C4E88E7C4.taxon	description	DESCRIPTION: Total length = 1.88 – 3.38 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face (Fig. 15 B). Ecdysial tear medial or posteromedial to base of antenna (Figs. 15 B, 79 D); along prothoracic extension. Head: Dorsal apotome (Figs. 19 H – J), without ventral line of weakness, without dorsomedial tubercle, with central dome; dorsolateral cephalic sclerite (Fig. 13 C) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 24 C) with only mandible or mandible ventral to lacinia; palpus extending equal to or just posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 35 B) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Figs. 19 H – J) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 24 C) — 2 short, slender setae; oculars (Fig. 24 C) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 24 C) wide, welldeveloped, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially or with slight protuberance, not dividing metathorax medially (Fig. 48 C); respiratory organ (Figs. 43 P – Q) length / width = 2.74 – 3.68, moderately elongate, apical half somewhat flattened dorsoventrally, with pores closely abutting or slightly separated at apex of respiratory organ, arranged in single straight or curve row, outer surface smooth or with some wrinkles, with or without short, wide pedicel, base with slender, strung out posteromedial apodeme, membranous base of respiratory organ short, tracheal tube apically J-shaped, surface smooth or distally with plates; wing (Fig. 35 B) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 32 H) just separate; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 35 B) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 32 H); with apex of foreleg ventral to or moderately anterior to apex of midleg; apex of hind leg slightly dorsal to, partially abutting apex of midleg laterally; sensilla: anteromedials — 1 seta; anterolaterals — 2 setae; dorsal setae (Fig. 29 N) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T setae, D- 3 - T campaniform sensillum, D- 3 - T posterior to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 48 C) — 1 seta, 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: without pigmentation pattern or pigmentation patterns only evident in alcohol / glycerin material, with tergite 1 with 3 medial spots, tergites 2 – 8 with 3 medial spots, anterolateral spots, sternites 3 – 7 with medial spot, anterolateral spot, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with bilobed, short and triangular to elongate and slender tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 73 J) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally to nearly laterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 48 C) with 6 setae, 2 campaniform sensilla, including 3 lateral sensilla, D- 2 - I, D- 3 - I closely approximated, D- 7 - I situated posteriorly near D- 8 - I; segment 4 (Figs. 57 C, 58 A) — D- 2 - IV, D- 3 - IV setae on either short tubercle, bilobed or bifid elongate tubercles; D- 5 - IV present or absent, D- 8 - IV, D- 9 - IV short to elongate setae; each at base of short bilobed or elongate bifid, separate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 7 - IV, D- 8 - IV, D- 9 - IV or D- 5 - IV absent and D- 7 - IV anterior to D- 9 - IV; L- 1 - IV short seta at base of short bilobed or elongate bifid tubercle, well anterior of posterior lateral setae; L- 2 - IV, L- 3 - IV, L- 4 - IV short setae at base of bilobed elongate bifid tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV on bilobed or at base of bifid, elongate tubercles; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 73 J) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1B3040FD61187C4E88E7C4.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Ceratopogon is known from 43 species in the Holarctic Region with adults emerging in cold environments (Arctic, mountainous, or in early spring) (Borkent & Grogan 1995). Immatures have been found in moss in cold forest pools, at the margins of shallow water bodies, bogs, fens, in moist soil, on flood plains and on lake shores.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1B3040FD61187C4E88E7C4.taxon	discussion	TAXONOMIC DISCUSSION: Six species of Ceratopogon are known as pupae (Tables 2 – 3) and the five Nearctic ones are keyed by Borkent & Grogan (1995) as part of a comprehensive revision of the genus. Ceratopogon boomerangus is the only member of the genus missing the full complement of abdominal sensilla present in most Ceratopogonidae (and all other Ceratopogon), with fourth abdominal segment sensillum D- 5 - IV absent. The figure of the fourth abdominal segment of C. boomerangus shown in Fig. 57 C (from Borkent & Grogan 1995) is distorted by compression and the distribution of the sensilla are broader than is natural (i. e. the dorsal and ventral sensilla are actually more lateral, L- 1 - IV, L- 2 - IV and L- 3 - IV are shown as ventral). This is true for all the species illustrated by Borkent & Grogan (1995: 183). The fourth abdominal segment of C. nr. inverecundus in Fig. 58 A was drawn from an uncompressed specimen in glycerin. Mayer (1934 a: 238) described a pupa referred to as " Genus incertum " from Germany which was reexamined for this study. It is a species of Ceratopogon but cannot be identified to species at present.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD1B3040FD61187C4E88E7C4.taxon	materials_examined	MATERIAL EXAMINED: C. abstrusus: 1 pupal exuviae (of paratype), Sand Pond Br., Blue Ridge, New York, USA, 26 - V- 1959 (NYSM); 1 pupal exuviae (of paratype), The Gulf Brook, Blue Ridge, New York, USA, 25 - V- 1958 (NYSM); 1 pupal exuviae (of paratype), New York (?) (NYSM); 1 pupal exuviae (of paratype), Hamilton-Essex, Newcomb, New York, USA, 28 - V- 1959 (NYSM). C. arcanus: 2 pupal exuviae (of paratypes), Fishing Creek, Newcomb, New York, USA, 5 - V- 1958 (NYSM); 2 pupal exuviae (of paratype), Huntington Rd. Marsh, Newcomb, New York, USA, 14 - V- 1958 (NYSM); 1 pupal exuviae (of paratype), Blue Mountain Lake, New York, USA, 3 - V- 1960 (NYSM); 1 pupal exuviae (of paratype), High Rock Pond outlet, Inlet, New York, USA, 8 - V- 1958 (NYSM); 2 pupal exuviae (of paratypes), Hamilton-Essex, Newcomb, New York, USA, 11 - V- 1959 (NYSM). C. boomerangus: 1 pupal exuviae (of allotype), College Park, Prince George County, Maryland, USA, 14 - IV- 1976 (USNM). C. inverecundus: 2 pupal exuviae (of paratypes), Edge 6 Mile Creek, Long Lake, New York, USA, 18 - V- 1959 (NYSM). C. naccinervis: 1 pupal exuviae, Severskii Donets River, Donetsk Province, Ukraine, 16 - IV- 1970 (ZIN). C. willisi: 1 pupa (paratype), College Park, Prince Georges County, Maryland, 17 - III- 1976 (USNM); 1 pupa (paratype), as previous locality, 26 - III- 1976 (USNM). C. nr. abstrusus: 1 pupal exuviae (in glycerin), 2 pupal exuviae, Bolean Lake, 6 km NE Falkland, BC, Canada, 12 - 13 - VII- 1989 (CNCI). C. nr. inverecundus: 1 pupal exuviae (in glycerin), 1 pupal exuviae, Bolean Lake, 6 km NE Falkland, BC, Canada, 12 - 13 - VII- 1989 (CNCI). C. sp.: 1 pupa, Haspertalsperre Sauerland, Westfalen, Germany 1 - VI-? (ZSMC).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD18304EFD6F1FFA4912E319.taxon	description	(Figs. 13 D, 15 C, 19 K, 24 D, 29 O, 32 I, 35 C, 43 R, 48 D, 58 B, 73 K)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD18304EFD6F1FFA4912E319.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with setae D- 1 - T, D- 2 - T, D- 4 - T, and D- 5 - T tightly appressed and on a single, short tubercle (Fig. 29 O). Also only pupa with prothoracic extension abutting the antenna but not extending to the palpus (Fig. 24 D) and halter extending only to the anterior margin of tergite 2 (Fig. 32 I).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD18304EFD6F1FFA4912E319.taxon	description	DESCRIPTION: Total length = 1.13 – 2.03 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face (Fig. 15 C). Ecdysial tear medial to base of antenna (Figs. 15 C, 79 D); along prothoracic extension. Head: Dorsal apotome (Fig. 19 K), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (Fig. 13 D) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 24 D) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 35 C) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Fig. 19 K) — 1 short seta present or absent, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, no campaniform sensillum; clypeal-labrals (Fig. 24 D) — 0 – 1 seta or 1 seta and 1 small pit; oculars (Fig. 24 D) — 2 setae, 2 campaniform sensilla. Thorax: Prothoracic extension (Fig. 24 D) short, present only dorsolaterally abutting antenna, not extending to palpus; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 48 D); respiratory organ (Fig. 43 R) length / width = 2.37 – 4.77, moderately elongate to elongate, with blunt apex, area with single subbasal spiracle slightly expanded, somewhat flattened dorsoventrally, with pores closely abutting at apex of respiratory organ, arranged in single row, with one additional subbasal pore, outer surface with annulations, without other surface modifications, with or without short, wide pedicel, base without posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with weak spirals restricted to base; wing (Fig. 35 C) with apical tubercle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 32 I) just separate to just touching; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 35 C) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 32 I); with apex of foreleg well anterior to apex of midleg; apex of hind leg slightly dorsal to, partially abutting apex of midleg laterally; sensilla: anteromedials — 1 seta; anterolaterals — 2 setae; dorsal setae (Fig. 29 O) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T setae, all on one tubercle, D- 3 - T campaniform sensillum, D- 3 - T posterolateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 48 D) — 1 seta, 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: without pigmentation pattern, 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 (Fig. 73 K) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally to nearly laterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 48 D) with 8 setae, 2 campaniform sensilla, including 3 lateral sensilla, D- 2 - I, D- 3 - I closely approximated, D- 7 - I situated posteriorly near D- 8 - I; segment 4 (Fig. 58 B) — D- 2 - IV peg-like or slender seta, D- 3 - IV moderately elongate seta on separate tubercles; D- 5 - IV short seta, D- 8 - IV, D- 9 - IV moderately elongate setae; on short, separate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 7 - IV, D- 8 - IV, D- 9 - IV; L- 1 - IV elongate seta on pointed tubercle, well anterior of posterior lateral setae; L- 2 - IV, L- 3 - IV, L- 4 - IV moderately elongate setae on pointed tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV short to moderately elongate setae on rounded tubercles; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 73 K) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD18304EFD6F1FFA4912E319.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Brachypogon is known from 200 species from every Region worldwide (Borkent 2014). Immatures are generally in such small to moderately sized lentic habitats as pools, marshes, bogs and fens as well as the mud, sand or detritus at the margins of streams and creeks. At least within the Nearctic, species are diverse and often common in bogs and fens. At least two species were common as adults in the intertidial zone in western Costa Rica (pers. obs.).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD18304EFD6F1FFA4912E319.taxon	discussion	TAXONOMIC DISCUSSION: There are only seven species of Brachypogon known as pupae (Tables 2 – 3). Some previous descriptions have included illustrations of the respiratory organ with pores only at the apex (Mayer 1940, Kettle & Lawson 1952, Glukhova 1979 b, Harris 1981, Elson-Harris 1990, Szadziewski et al. 1994, 1997). All specimens examined here had an additional pore near the base and in some instances this was difficult to see. If present in all Brachypogon, the arrangement of pores and the squat, rectangular to somewhat funnel-shape of the respiratory organ would be unique in the Ceratopogonidae. Debenham (1991) illustrates three species of Brachypogon from Australia and Mayer (1940) described one species from Sweden with only two posterior lateral sensilla on segment 4 (likely L- 3 - IV, L- 4 - IV) but these should be reexamined for the smaller L- 2 - IV possibly present. Thienemann (1936: 176) described a pupa as " genus incertum " that is almost certainly a species of Brachypogon. It has the characteristic respiratory organ shape and arrangement of pores (one basal) and the distribution of abdominal sensilla matches fairly well (there is an extra anterior lateral sensillum that would be unique to the genus but was likely an error).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD18304EFD6F1FFA4912E319.taxon	materials_examined	MATERIAL EXAMINED: B. nitidulus: 1 pupal exuviae, Zaklin'e, Leningrad Province, Russia, 24 - VI- 1972 (ZIN). B. taivoi: 1 pupal exuviae, Chu river, Kochkorka, Issyk-Kul Province, Kyrgyzstan, 30 - VIII- 1971 (ZIN). B. ussuriensis: 2 pupal exuviae, Ussuri Nature Reserve, Primorskii Territory, Russia, 6 - VI- 1973 (ZIN). B. sp.: 1 pupal exuviae (in glycerin), 6 km E Salmon Arm, BC, Canada, 6 - VI- 1990 (CNCI); 1 pupa, 3 pupal exuviae (in glycerin), Spanish Lake, 6 km E. Falkland, 50 ° 29.12 N 119 ° 28.07 W, BC, Canada, 27 - 28 - V- 2008 (CNCI); 1 pupal exuviae, Blue Mountain Lake, New York, USA, 3 - V- 1960 (NYSM); 2 pupal exuviae, Flag Glade, Pocahontas County, Virginia, USA, 24 - VI- 1976 (VPIC); 5 pupal exuviae, 1 km downstream lower Cascade Falls, Giles County, Virginia, USA, 3 - VII- 1977 (5 VPIC); 1 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 19 - II- 1967 (BPBM); 1 pupal exuviae, Houay La stream, Moung Sayaboury, Sayaboury Province, Laos, 25 - XI- 1967 (BPBM); 1 pupal exuviae, Moung Sayaboury, Sayaboury Muong, Laos, 19 - 23 - II- 1967 (BPBM).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD16304CFD6C1B7A4D85E1E9.taxon	description	(Figs. 13 E, 19 L – M, 24 E, 29 P, 32 J, 35 D, 43 S – T, 44 A – B, 48 E, 58 C, 73 L)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD16304CFD6C1B7A4D85E1E9.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the respiratory organ with two closely appressed or approximated rows of pores and with each pore oblong in shape (Figs. 43 S – T, 44 A – B). Also only pupae with the abdominal segment 4 sensilla D- 4 - IV and D- 7 - IV placed on either side of D- 8 - IV (Fig. 58 C) (in some species D- 7 - IV is placed more laterally, closer to D- 9 - IV). Also only pupa with the prothoracic extension abutting the antenna but not extending to the palpus (Fig. 24 E) and the halter extending well posterior of the anterior margin of tergite 2 (Fig. 32 J).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD16304CFD6C1B7A4D85E1E9.taxon	description	DESCRIPTION: Total length = 1.81 – 2.75 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, in some against wing (as in Figs. 15 D, 32 J). Ecdysial tear medial to base of antenna or to posterior to base (as in Figs. 15 B, D, 79 D, E); along prothoracic extension. Head: Dorsal apotome (Figs. 19 L – M), with ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (Fig. 13 E) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 24 E) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 35 D) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Figs. 19 L – M) — 1 short to moderate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 24 E) — absent; oculars (Fig. 24 E) — 2 setae, 2 campaniform sensilla. Thorax: Prothoracic extension (Fig. 24 E) short, present only dorsolaterally abutting antenna, not extending to palpus; mesonotum with very short tubercles, not extending posteromedially or with slight protuberance, not dividing metathorax medially (Fig. 48 E); respiratory organ (Figs. 43 S – T, 44 A – B) length / width = 3.56 – 7.20, elongate, slender to moderately thick, somewhat flattened dorsoventrally, with pores closely abutting at apex of respiratory organ, arranged in two closely appressed rows, with or without additional, more basal pores, outer surface with (nearly all species) or without spicules, spines or pointed plates, without pedicel, base without posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, distally with plate; wing (Fig. 35 D) with apical tubercle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 32 J) broadly abutting; halter apex extending posteriorly to 1 / 3 length of tergite 2; legs (Fig. 35 D) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 32 J); 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 — 3 setae; dorsal setae (Fig. 29 P) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T setae, D- 3 - T campaniform sensillum, D- 3 - T posteromedial to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 48 E) — 1 seta, 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: without pigmentation pattern or anteromedial margin of tergites 1 – 4 (fading posteriorly) or tergites 1 – 5 (tergite 1 with 4 medial), just medial on tergite 6, fading posteriorly, 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 (Fig. 73 L) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 48 E) with 7 – 8 setae, 2 campaniform sensilla, including 3 lateral sensilla, D- 2 - I, D- 3 - I closely approximated, D- 7 - I situated posteriorly near D- 8 - I; segment 4 (Fig. 58 C) — D- 2 - IV, D- 3 - IV moderately elongate setae on separate tubercles; D- 5 - IV, D- 8 - IV, D- 9 - IV short setae, on short to moderately elongate, separate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 8 - IV, D- 7 - IV, D- 9 - IV; L- 1 - IV short seta on pointed tubercle, well anterior of posterior lateral setae; L- 2 - IV, L- 3 - IV, L- 4 - IV short setae on pointed tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV short to moderately elongate setae on pointed tubercles; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 73 L) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD16304CFD6C1B7A4D85E1E9.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Alluaudomyia is known from 203 species from every Region worldwide (Borkent 2014). Immatures have been sampled from ponds, bogs, fens, swamps, treeholes and margins of streams, creeks, rivers and lakes. Some species have been reared from wet leaves and mud associated with the aforementioned habitats.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD16304CFD6C1B7A4D85E1E9.taxon	discussion	TAXONOMIC DISCUSSION: There are 23 species of Alluaudomyia known as pupae (Tables 2 – 3). Considering the good visibility of the larvae, which swim on the surface film, there are good prospects of rearing many additional species of this genus. The dorsal apotomes of A. bella, A. caribbeana, A. distispinulosa, A. footei, A. megaparamera, A. paraspina, A. parv a, A. reyei and A. unguistyla have been illustrated without campaniform sensillum DA- 2 - H (Elson-Harris & Kettle 1985 b, Grogan & Bystrak 1976, Spinelli 1997, Spinelli & Wirth 1984, Wirth & Grogan 1981), but examination of original material of A. bella, A. distispinulosa, A. footei, A. megaparamera, A. paraspina and A. parv a indicates that it is actually present in at least these species. The campaniform sensillum has likely been overlooked in the other species considering that all species of Alluaudomyia I examined had the sensillum present among the similarly sized spicules and that the sensillum is present in most other Ceratopogonidae. De Meillon (1939) illustrated abdominal segment 5 of A. maculithorax but misidentified the dorsal and ventral surfaces. The distinctive placement of fourth abdominal segment sensilla D- 7 - IV lateral to D- 8 - IV has not been reported or illustrated by those previous authors describing the sensilla of the abdominal segments, likely because this sensillum coeloconica is small, difficult to discern and was overlooked (de Meillon 1939, Debenham 1971, Elson-Harris & Kettle 1985 b); this is certainly the case for the three species described by Debenham (1971) and which were reexamined here.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD16304CFD6C1B7A4D85E1E9.taxon	materials_examined	MATERIAL EXAMINED: A. appendiculata: 2 pupal exuviae (of paratype), Hornsby, New South Wales, Australia, 9 - X- 1956 (ANIC); 1 pupal exuviae, as previous locality, 18 - XI- 1969 (ANIC); 2 pupal exuviae, Nattai River, Mittagong, New South Wales, Australia, 25 - X- 1968 (ANIC). A. bella: 1 pupal exuviae, 6.5 km NW of Enderby, British Columbia, Canada, 27 - VI- 1990 (CNCI); 1 pupal exuviae, 20 km E of Anola, Manitoba, Canada, 16 - VI- 1990 (CNCI); 1 pupal exuviae (in glycerin), Charleston Provincial Park, Ontario, Canada, 18 - V- 1986 (CNCI); 1 pupal exuviae, Algonquin Park, Ontario, Canada, 7 - VI- 1960 (USNM); 6 pupal exuviae, Bat Lake, Algonquin Park, Ontario, Canada, 7 - VI- 1960 (USNM); 1 pupal exuviae, Notre Dame de Laus, Lac Serpent, Quebec, Canada, 22 - VI- 1967 (USNM); 1 pupal exuviae, as previous locality, 11 - VII- 1967 (USNM); 1 pupal exuviae, Patuxent Wildlife Rescue Center, Prince George’s County, Maryland, USA, 19 - IV- 1976 (WLGC); 2 pupal exuviae, as previous locality, 8 - V- 1958 (USNM); 1 pupal exuviae, as previous locality, 28 - VI- 1976 (WLGC); 1 pupal exuviae, as previous locality, 17 - V- 1976 (WLGC); 3 pupal exuviae, Lake Jimmy, Tahawus, New York, USA, 24 - VI- 1959 (NYSM); 1 pupal exuviae, Hamilton-Essex, Newcomb, New York, USA, 23 - VII- 1959 (NYSM); 1 pupal exuviae, Huntington Swamp, Newcomb, New York, USA, 4 - VIII- 1959 (NYSM); 1 pupal exuviae, Salmon River, Blue Mountain Lake, New York, USA, 23 - VI- 1959 (NYSM); 1 pupal exuviae, Allegany State Park, New York, USA, 28 - V- 1963 (USNM); 1 pupal exuviae, Upper Pansapoulas Pond, Montgomery County, Virginia, USA, 3 - IX- 1976 (VPIC); 2 pupal exuviae, Upper Pandapas Pond, Montgomery County, Virginia, USA, 2 - VIII- 1976 (VPIC). A. bicornis: 1 pupal exuviae, Manton River, Northern Territory, Australia, 6 - VI- 1958 (ANIC). A. biestroi: 1 pupal exuviae (of paratype), Artigas A “ Lenguazo ” y Ruta 3, Uruguay, 8 - XI- 1984 (MLPA). A. claudia: 1 pupal exuviae (of paratype) Salisbury, Southern Rhodesia, South Africa, 24 - II- 1949 (SAIM). A. distispinulosa: 1 pupal exuviae, Bulo Bulo, Cochabamba, Bolivia, 12 - I- 1995 (MLPA). A. fragmentum: 1 pupal exuviae (of paratype), Manton River, Northern Territory, Australia, 6 - VI- 1958 (ANIC); 1 pupal exuviae (of paratype), as previous locality, 1 - VI- 1958 (ANIC). A. latipennis: 1 pupal exuviae, Hornsby, New South Wales, Australia, 9 - X- 1956 (ANIC); 1 pupal exuviae, Nattai River, Mittagong, New South Wales, Australia, 4 - XI- 1964 (ANIC); 1 pupal exuviae, as previous locality, 15 - XI- 1968 (ANIC). A. megaparamera: 1 pupal exuviae, Blue Mountain Lake, New York, USA, 29 - VI- 1959 (NYSM); 1 pupal exuviae, Salmon River, Blue Mountain Lake, New York, USA, 8 - VI- 1959 (NYSM); 1 pupal exuviae, Hamilton-Essex, Newcomb, New York, USA, 1 - VI- 1959 (NYSM); 1 pupal exuviae, as previous locality, 8 - VI-? (NYSM). A. natalensis: 1 pupal exuviae, Empangeni, Zululand, South Africa, 16 - VII- 1938 (SAIM); 1 pupal exuviae, 15 mi. SE of Potgietersrus, Transvaal, South Africa, 3 - I- 1974 (NMSA). A. needhami: 1 pupal exuviae, Salmon Arm, British Columbia, Canada, 11 - VII- 1990 (CNCI); 3 pupal exuviae, 30 km N of Nakusp, British Columbia, Canada, 5 - VII- 1990 (CNCI); 2 pupal exuviae, Patuxent Wildlife Refuge, Prince George’s County, Maryland, USA, 1 - VIII- 1979 (USNM); 1 pupal exuviae, Upper Pandapas Pond, W. A., Montgomery County, Virginia, USA (VPIC); 1 pupal exuviae, 1.7 mi E of Lunter’s gate, Rt. 785, Montgomery County, Virginia, USA, 22 - IV- 1976 (VPIC); 1 pupal exuviae, no locality, 10 - VI- 1953 (USNM); 2 pupal exuviae, no locality, 20 - II- 1957 (USNM). A. needhami or A. megaparamera: 2 pupal exuviae, Montvale Wayside, Bedford County, Virginia, USA, 16 - VI- 1977 (VPIC )); 1 pupal exuviae, Beaver Lake Reservoir, Pocahontas State Park, Chesterfield County, Virginia, USA, 30 - IV- 1977 (VPIC). A. paraspina: 2 pupal exuviae, Lakeland Pond, College Park, Prince George’s County, Maryland, USA, 26 - V- 1975 (WLGC); 2 pupal exuviae, as previous locality, 26 - V- 1975 (WLGC); 5 pupal exuviae, as previous locality, 30 - V- 1975 (WLGC); 1 pupal exuviae, Mountain Lake Biological Station, Giles County, Virginia, USA (VPIC). A. parva: 1 pupal exuviae, Blue Mountain Lake, New York, USA, 30 - VI- 1958 (NYSM); 1 pupal exuviae, Salmon River, Blue Mountain Lake, New York, USA, 14 - V- 1959 (NYSM); 1 pupal exuviae, Allegany State Park, New York, USA, 28 - V- 1963 (USNM); 7 pupal exuviae, Alexandria, Virginia, USA, 6 - VI- 1951 (USNM). A. quadripunctata: 2 pupal exuviae, Tobolki lake, Alol’, Pskov Province, Russia, 30 - VI- 1969 (ZIN). A. schnacki: 2 pupal exuviae (of paratype), Los Talas (Berisso), Buenos Aires, Argentina, 29 - XI- 1979 (MLPA). A. splendida: 1 pupal exuviae, Ussuri Nature Reserve, Primorskii Territory, Russia, 12 - VI- 1973 (ZIN). A. variegata: 1 pupal exuviae, Lake Muson, Leon County, Florida, USA, 22 - VIII- 1987 (JHEC). A. sp.: 1 pupal exuviae, Bolean Lake, 6 km NE of Falkland, British Columbia, Canada, 1 - VII- 2009 (CNCI); 1 pupal exuviae, 30 km N of Nakusp, British Columbia, Canada, 5 - VII- 1990 (CNCI); 1 pupal exuviae, 5 km S of Cottonwood Lake, Nelson, British Columbia, Canada, 6 - VII- 2008 (CNCI); 1 pupal exuviae, Meachum Lake, New York, USA, 21 - VI- 1986 (CNCI); 1 pupal exuviae, Ackerman Swamp, Newcomb, New York, USA, 7 - VII- 1958 (NYSM); 1 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 19 - II- 1967 (BPBM); 1 pupal exuviae, as previous locality, 2 - XII- 1967 (BPBM); 2 pupal exuviae, as previous locality, 22 - II- 1967 (BPBM); 1 pupal exuviae, Houay La stream, 20 kilometers NE of Moung Sayaboury, Sayaboury Province, Laos, 25 - XI- 1967 (BPBM); 1 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 9 - XII- 1967 (BPBM); 1 pupal exuviae, as previous locality, 7 - I- 1968 (BPBM); 1 pupal exuviae, Nam Houng River, Moung Sayaboury, Sayaboury Province, Laos, 17 - XII- 1967 (BPBM); 1 pupal exuviae, Gnieraoora Pool, Robe River, Pilbara, Western Australia, Australia (ANIC).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD14304DFDB81E2F4C43E6EC.taxon	description	(Figs. 2 F, 19 N, 24 F, 29 Q, 32 K, 35 E, 44 C, 48 F, 59 A, 74 A)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD14304DFDB81E2F4C43E6EC.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the respiratory organ elongate and with three distinct sections: a cylindrical, smooth basal portion, a cylindrical middle portion with annulations for most of its distal length, and a compressed, rounded or concave apical portion (Fig. 44 C); also unique with a short foreleg, with its apex well anterior of the apex of the wing (Fig. 35 E); also unique in being very small, with total body length 0.91 – 1.19 mm (only Schizonyxhelea are similarly small, with a body length 1.19 – 2.16 mm; however, the unknown pupae of Nannohelea, Baeohelea and some others will certainly be equally tiny).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD14304DFDB81E2F4C43E6EC.taxon	description	DESCRIPTION: Total length = 0.91 – 1.19 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, in some specimens against wing (as in Figs. 15 D, 32 K). Ecdysial tear medial to base of antenna (as in Figs. 15 B, 79 D); along prothoracic extension. Head: Dorsal apotome (Fig. 19 N), without ventral line of weakness, with dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 B) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 24 F) with mandible, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna (Fig. 35 E) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Fig. 19 N) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 2 setae, 1 campaniform sensillum; clypeal-labrals (Fig. 24 F) — not visible; oculars (Fig. 24 F) — 2 elongate setae. Thorax: Prothoracic extension (Fig. 24 F) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, extending posteromedially, completely dividing metathorax medially (Fig. 48 F); respiratory organ (Fig. 44 C) length / width = 3.75 – 6.67, elongate, with three distinct sections: cylindrical basal portion smooth, cylindrical middle portion with annulations for most of distal length, compressed apical portion rounded or concave apically, apical portion somewhat flattened, with pores closely abutting or slightly separated at apex of respiratory organ, arranged in single row, with one additional, more basal pore, outer surface with annulations, without other surface modifications, with moderately elongate pedicel, base without posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base; wing (Fig. 35 E) with slight angle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 32 K) broadly abutting; halter apex extending posteriorly to 1 / 3 length of tergite 2; legs (Fig. 35 E) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 32 K); with apex of foreleg far anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 1 seta; anterolaterals — 2 setae; dorsal setae (Fig. 29 Q) — D- 1 - T, D- 2 - T, D- 4 - T setae, D- 3 - T campaniform sensillum, D- 5 - T absent; D- 1 - T, D- 2 - T on single tubercle, D- 3 - T posteromedial to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 48 F) — 1 seta, 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: without pigmentation pattern, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with slightly rounded to pointed, or serrate short tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 74 A) not strongly modified but with dorsolateral row of spines at near midlength, bordering shallow posteromedial concavity, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 48 F) with 6 setae, 1 campaniform sensillum, including 3 lateral sensilla, D- 2 - I, D- 3 - I closely approximated, D- 7 - I situated posteriorly near D- 8 - I; segment 4 (Fig. 59 A) — D- 2 - IV, D- 3 - IV short setae on serrate tubercles; D- 5 - IV barely visible short seta, D- 8 - IV short seta, D- 9 - IV absent; D- 5 - IV, D- 4 - IV, D- 7 - IV, D- 8 - IV on short, serrate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 7 - IV, D- 8 - IV; L- 1 - IV absent, L- 2 - IV, L- 3 - IV short setae on pointed tubercles, L- 4 - IV thick seta on serrate tubercle, V- 5 - IV, V- 6 - IV, V- 7 - IV short setae on serrate tubercles; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 74 A) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD14304DFDB81E2F4C43E6EC.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Baeodasymyia is known from five species in the Neotropical Region (Borkent 2014). The immatures of two species have been reared from small springs in Costa Rica (Borkent & Craig 1999).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD14304DFDB81E2F4C43E6EC.taxon	discussion	TAXONOMIC DISCUSSION: Borkent & Craig (1999) described the pupae of the only two species known in this stage (Tables 2 – 3). Pupae of Baeodasymyia are lacking thoracic sensillum D- 5 - T and two sensilla on abdominal segment 4: D- 9 - IV and L- 1 - IV.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD14304DFDB81E2F4C43E6EC.taxon	materials_examined	MATERIAL EXAMINED: B. christopheri: 1 pupal exuviae (of paratype), Atenas, Costa Rica, 5 - X- 1993 (as larva, reared) (CNCI); 1 pupa (of paratype), 2 km NE Tarcoles, Costa Rica, 17 - XII- 1993 (as larva, reared) (CNCI). B. michaeli: 3 pupae (of paratypes), 9 pupal exuviae (of paratypes), 1 pupal exuviae, Atenas, Costa Rica, 25 - X- 1993 (as larvae, reared) (CNCI, INBC); 1 pupal exuviae (in glycerin), 2 km NE of Tarcoles, Costa Rica, 17 - XII- 1993 (CNCI).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD15304AFD9B1F294C04E6A4.taxon	description	(Figs. 19 O, 24 G, 29 R, 35 F, 44 D, 49 A, 59 B, 74 B)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD15304AFD9B1F294C04E6A4.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the apex of the hind leg ventral to the apex of the midleg (Fig. 35 F) (only the female is known; perhaps the male, without the elongate hind leg claw of the developing adult will not have this feature).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD15304AFD9B1F294C04E6A4.taxon	description	DESCRIPTION: Total length = 2.50 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face (as in Fig. 15 D). Ecdysial tear around base of antenna (as in Figs. 15 D, 79 E); along prothoracic extension. Head: Dorsal apotome (Fig. 19 O), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 B) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 24 G) with mandible well-developed, lacinia absent; palpus extending equal to or just posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 35 F) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Fig. 19 O) — 1 moderately elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, no campaniform sensillum; clypeal-labrals (Fig. 24 G) — 2 elongate setae; oculars (Fig. 24 G) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 24 G) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, extending posteromedially, completely dividing metathorax medially (Fig. 49 A); respiratory organ (Fig. 44 D) length / width = 6.25 – 6.25, elongate, slender, somewhat flattened dorsoventrally, with pores closely abutting at apex of respiratory organ, arranged in single row, with additional more basal pores, outer surface smooth, with moderately elongate pedicel, base with short posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, distally with plate; wing (Fig. 35 F) with slight angle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 A) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 35 F) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 32 L); with apex of foreleg slightly anterior to apex of midleg; apex of hind leg ventral to apex of midleg, apex of each hind leg nearly abutting apically and abutting apices of foreleg; sensilla: anteromedials — 2 setae; anterolaterals — 3 setae; dorsal setae (Fig. 29 R) — D- 1 - T, D- 2 - T, D- 4 - T setae, D- 3 - T campaniform sensillum, D- 5 - T absent, D- 3 - T posteromedial to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 49 A) — 1 seta, 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: with anterior margins of tergites 3 – 7, anterior margins of sternites 3 – 7 pigmented, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with bilobed or rounded, short tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 74 B) not strongly modified, terminal processes widely separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 49 A) with 6 setae, 2 campaniform sensilla, including 3 lateral sensilla, D- 2 - I, D- 3 - I closely approximated, D- 7 - I situated posteriorly near D- 8 - I; segment 4 (Fig. 59 B) — D- 2 - IV, D- 3 - IV short or moderately elongate setae on separate tubercles; D- 5 - IV, D- 8 - IV short seta, D- 9 - IV absent; on short bilobed tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 7 - IV, D- 8 - IV; L- 1 - IV 1 seta on bilobed tubercle, well anterior of posterior lateral setae; L- 2 - IV, L- 3 - IV, L- 4 - IV short setae on bilobed tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV short setae on bilobed tubercles; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 74 B) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD15304AFD9B1F294C04E6A4.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Austrohelea is known from seven species from Australia, New Zealand and Argentina (Borkent 2014). The only pupa known was reared from the subantarctic Campbell Island but the specific habitat was not recorded.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD15304AFD9B1F294C04E6A4.taxon	discussion	TAXONOMIC DISCUSSION: The unique pupal exuviae was previously described by Sublette & Wirth (1980) and reexamined here (Tables 2 – 3).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD15304AFD9B1F294C04E6A4.taxon	materials_examined	MATERIAL EXAMINED: A. campbellensis: 1 pupal exuviae, Tucser Cove, Campbell Island, New Zealand, 22 - I- 1969 (NZAC).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD123049FD511EF34C91E501.taxon	description	(Figs. 15 D, 19 P, 20 A – D, 25 A – B, 29 S, 32 L, 36 A – B, 44 E – K, 49 B, 59 C, 60 A, 74 C – E)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD123049FD511EF34C91E501.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the thoracic sensilla D- 1 - T, D- 2 - T, D- 3 - T and D- 4 - T all present on flat cuticle (not on tubercles) and close to one another and with D- 5 - T either absent or, at most, a minute pit lacking any seta (Fig. 29 S).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD123049FD511EF34C91E501.taxon	description	DESCRIPTION: Total length = 1.69 – 3.53 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, in some against wing (Figs. 15 D, 32 L). Ecdysial tear posterior to base of antenna (Figs. 15 D, 79 E); along prothoracic extension. Head: Dorsal apotome (Figs. 19 P, 20 A – D), with ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 B) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Figs. 25 A – B) with mandible well-developed, fused to surface, lacinia absent; palpus extending equal to or just posterior to posterolateral margin of labium; labium entire or separated medially by labrum, hypopharynx; apex of antenna (Figs. 36 A – B) just anterior to posterior to, posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Figs. 19 P, 20 A – D) — 1 short to elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Figs. 25 A – B) — absent, with 1 minute seta, 1 campaniform sensillum or 2 setae; oculars (Figs. 25 A – B) — 1 seta, 1 campaniform sensillum or 2 setae or 2 setae, 2 campaniform sensilla. Thorax: Prothoracic extension (Figs. 25 A – B) wide, well-developed, extending from palpus to antenna; mesonotum without tubercles, not extending posteromedially to extending posteromedially, completely dividing metathorax medially (Fig. 49 B); respiratory organ (Figs. 44 E – K) length / width = 4.00 – 9.00, variable, elongate, slender, somewhat flattened or circular in cross-section, with pores closely abutting or slightly separated at apex of respiratory organ, arranged in single row, with or without additional, more basal pores, outer surface smooth, without or with short, wide to elongate, slender pedicel, base without posteromedial apodeme, membranous base of respiratory organ short to moderately elongate, tracheal tube straight to slightly curved along length, with spirals restricted to base, distally with plate; wing (Figs. 36 A – B) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 32 L) broadly abutting; halter apex extending posteriorly to 1 / 4 length of tergite 2; legs (Figs. 36 A – B) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 32 L); with apex of foreleg slightly to moderately anterior to apex of midleg; apex of hind leg slightly ventral to, partially abutting or just abutting apex of midleg laterally; sensilla: anteromedials — 2 setae; anterolaterals — 2 – 3 setae; dorsal setae (Fig. 29 S) — D- 1 - T, D- 2 - T, D- 4 - T setae, D- 3 - T campaniform sensillum, D- 5 - T absent or, at most, a minute pit lacking any seta, with D- 1 - T, D- 2 - T, D- 3 - T, D- 4 - T all present on flat cuticle, closely approximated, D- 3 - T posteromedial to posterolateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 49 B) — 1 very small to long seta, 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: without pigmentation pattern or, segment 2 as wide or slightly wider than segment 3, segments with or without bifurcating setae, with bilobed, bifid or rounded to pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Figs. 74 C – E) not strongly modified, terminal processes closely approximated basally, each projecting posteriorly to laterally, tapering to pointed apex, in some elongate, slender; sensilla: tergite 1 (Fig. 49 B) with 7 – 8 setae, 2 campaniform sensilla, including 3 lateral sensilla (4 in S. enigma), D- 2 - I, D- 3 - I closely approximated, D- 7 - I situated posteriorly near D- 8 - I; segment 4 (Figs. 59 C, 60 A) — D- 2 - IV, D- 3 - IV short or moderately elongate setae or D- 3 - IV absent, without or on low tubercles; D- 5 - IV, D- 8 - IV, D- 9 - IV short to elongate simple or bifurcating setae or D- 5 - IV, D- 9 - IV absent; on short to moderately elongate, separate tubercles or D- 7 - IV, D- 8 - IV on single tubercle, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 7 - IV, D- 8 - IV, D- 9 - IV to only D- 4 - IV, D- 7 - IV, D- 8 - IV present, L- 1 - IV absent or (rarely) short to elongate seta on short to elongate tubercle, in nearly transverse row with other lateral setae, arranged L- 2 - IV, L- 3 - IV, L- 4 - IV; L- 1 - IV, L- 2 - IV, L- 3 - IV, L- 4 - IV short to elongate setae on pointed or bifid tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV short to elongate, simple or bifurcating setae on short, rounded to elongate tubercles, in some with V- 6 - IV, V- 7 - IV on single tubercle; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Figs. 74 C-E) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD123049FD511EF34C91E501.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Stilobezzia is known from 341 species from every Region worldwide (Borkent 2014, minus species here transferred to Schizonyxhelea). Immatures are in a wide variety of aquatic and sem-aquatic habitats including springs, swamps, bogs, fens, rice fields, rock pools, treeholes, soil in a tidal marsh, and the margins of ponds, lakes, streams and rivers.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD123049FD511EF34C91E501.taxon	discussion	TAXONOMIC DISCUSSION: There are only 29 species of Stilobezzia known as pupae (Tables 2 – 3). As indicated below, obtaining the pupae of additional species will likely lead to a better understanding of the cladistic relationships within this group. The earliest description of a Stilobezzia pupa may be that by Packard (1871) who described it as a Tanypus (Chironomidae). Although clearly a ceratopogonid, the taxon cannot be identified with certainty although the overall habitus (especially the lateral direction of the terminal processes) suggests it is a species of Stilobezzia. However, it would be informative if this species (noted as common at that time) was resampled at Clear Lake, California to confirm the identification. The pupae of Stilobezzia species are strikingly diverse, showing considerable variation, for example in the number and distribution of sensilla of abdominal segment 4 (Figs. 59 C, 60 A). The shape of the respiratory organs (Figs. 44 E – K) and terminal processes (Figs. 74 C – E) also vary broadly. The remarkably modified respiratory organs of S. rabelloi (Fig. 44 I) and S. poikiloptera are used to pierce the submerged roots of aquatic plants to obtain oxygen for the pupa (Borkent & Craig 2001). Ronderos et al. (2012) described a peculiar species of Stilobezzia as S. enigma and discussed it's phylogenetic placement, concluding that it likely belongs within the subgenus S. (Stilobezzia), based on the lack of any indication of tubercles on the dorsum of the thorax (otherwise bearing the dorsal sensilla). Within the analysis here, the pupa has synapmorphies 27, 29 and 39 and confirms its placement within the genus. The presence of four lateral sensilla on tergite 1, all of which are setae, is a feature unique with the Ceratopogonidae. The presence of D – 7 - I near D- 2 - I and D- 3 - I is unique within the genus (see character 53). Cazorla et al. (2006) drew the sensilla of abdominal segment 4 of S. fiebrigi with more sensilla than are known in any other Ceratopogonidae and this species should be reexamined to confirm their presence and distribution. Harris (1981) described seven species of Stilobezzia from Australia, six of which were considered new and provided a key to these. However, her species 6 is here recognized as a member of Schizonyxhelea, with its characteristic features (see below under that genus). Based on the conclusion that Schizonyxhelea is clearly monophyletic and most closely related to Baeodasymyia, some species previously recognized as members of Stilobezzia are now placed in Schizonyxhelea as new combinations (see below under that genus). The phylogenetic relationships between species of Stilobezzia are uncertain and the genus warrants a detailed cladistic analysis to test the reality of the current division into four subgenera. Clastrier (1976) recognized that the current subgenera (at that time with three subgenera) were artificial and correctly suggested further character states (primarily of the antennae, wings and thoracic chaetotaxy) to interpret the group. Unfortunately, he did not interpret the newly reported features cladistically and the result was a difficult to interpret new arrangement of the species. Wirth & Grogan (1988) did not use the system suggested by Clastrier (1976) and named an additional subgenus for two distinctive Australian species. Some pupal features are likely important in future phylogenetic analyses of this interesting genus. For example, the dorsal setae arising from the mesonotum of Ceratopogonidae pupae nearly always arise from at least low tubercles. Within Stilobezzia, three species of the subgenus S. (Acanthohelea) (S. gracilis, S. lutea, and S. orientis) have the setae arising from very low tubercles. Those of the subgenus S. (Stilobezzia), however, have the setae arising directly from the cuticle and this unique feature indicates their monophyly (S. antennalis, S. coquillettii, S. diversa, S. flavirostris, S. glauca, S. limnophila, S. navaiae, S. pallidiventris, S. papillata, S. pictipes, S. rabelloi, and S. sybleae). Two species of S. (Acanthohelea), S. papillata (examined here) and S. ochracea (as described by Kettle & Lawson 1952) also have slender setae arising directly from the cuticle, indicating that at least these two species of S. (Acanthohelea) are more closely related to species of the subgenus S. (Stilobezzia) than they are to other S. (Acanthohelea). It is worth noting that S. papillata and S. ochracea have unique elongate spicules on the apices of their respiratory organs, certainly indicating that these two are sister species. Three of the species of S. (Acanthohelea), namely S. gracilis, S. lutea, and S. orientis, have most of the setae on abdominal segments 3 – 7 arising from distinctive bifid tubercles in which each half of the tubercle is very slender and about as long as the seta borne by it. This feature is nearly unique in the Ceratopogonidae, otherwise similar to that in some species of Ceratopogon (where it is clearly independently derived). The feature is, therefore, derived. The two species of S. (Acanthohelea), S. papillata and S. ochracea, which, as indicated above, are likely related to species of the subgenus S. (Stilobezzia) lack this feature. Furthermore, the strongly modified pupa of S. (S.) rabelloi (Borkent & Craig 2001) does have bifid tubercles although they are much larger than those of S. gracilis, S. lutea, and S. orientis. This suggests that S. rabelloi is actually a member of S. (Acanthohelea). Although this species lacks the defining presence of macrotrichia on the apex of the adult wing, it does bear strong setae on veins R and R 3, which are otherwise lacking in the subgenus S. (Stilobezzia) but are present in S. (Acanthohelea).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD123049FD511EF34C91E501.taxon	materials_examined	MATERIAL EXAMINED: S. antennalis: 7 pupal exuviae, Salmon Arm, British Columbia, Canada, 7 - VI- 1988 (CNCI); 1 pupal exuviae, 3 km E of Salmon Arm, British Columbia, Canada, 9 - VI- 1988 (CNCI); 1 pupal exuviae, 30 km N of Nakusp, British Columbia, Canada, 5 - VII- 1990 (CNCI); 1 pupal exuviae, Plummer’s Island, Montgomery County, Maryland, USA, 4 - VI- 1976 (WLGC); 2 pupal exuviae, Patuxent Wildlife Refuge, Prince George’s County, Maryland, USA, 5 - VII- 1977 (USNM); 1 pupal exuviae, as previous locality, 24 - V- 1977 (USNM); 1 pupal exuviae, Highlands Lake Ravenel, Macon County, North Carolina, USA, 10 - VI- 1986 (USNM); 2 pupal exuviae, Grandview Natural Preserve, Hampton, Virginia, USA, 30 - VII- 1976, 25 - VIII- 1976 (VPIC); 1 pupal exuviae (in glycerin), Grandview Natural Preserve, Hampton, Virginia, USA, 30 - VII- 1976 (VPIC); 2 pupal exuviae, Dyke Marsh, Alexandria, Virginia, USA, 3 - VI- 1975 (WLGC); 1 pupal exuviae, no locality, 6 - VIII- 1953 (USNM). S. coquilletti: 1 pupal exuviae, Farmer Agnew’s Property, Montgomery County, Virginia, USA, 22 - VIII- 1975 (VPIC); 2 pupal exuviae, Hasler Farm, Co. Rt. 763, Rockingham County, Virginia, USA, 24 - VII- 1975 (VPIC); 1 pupal exuviae, Balboa, Panama, 3 - IX- 1942 (USNM); 1 pupal exuviae, Balboa, Panama, VIII- 1942 (USNM). S. diversa: 1 pupal exuviae, Patuxent Wildlife Refuge, Prince George’s County, Maryland, USA, 24 - V- 1977 (USNM). S. flavirostris: 1 pupal exuviae, Maikov, Rovno Province, Ukraine, 29 - V- 1974 (ZIN). S. glauca: 1 pupal exuviae, Patuxent Rescue Center, Prince George’s County, Maryland, USA, 13 - V- 1976 (USNM); 3 pupal exuviae, 4.5 mi. E of Corapeake Road from Desert Road, Camden County, North Carolina, USA, 29 - VII- 1976 (VPIC); 1 pupal exuviae, Dyke Marsh, Alexandria, Virginia, USA, 29 - VI- 1952 (WLGC); 1 pupal exuviae, Falls Church, Virginia, USA, 30 - VI- 1951 (WLGC); 1 pupal exuviae, Dyke Marsh, Alexandria, Virginia, USA, 29 - VI- 1952 (WLGC); 1 pupal exuviae, Watoga State Park, Pocahontas County, Virginia, USA, 25 - VI- 1976 (VPIC); 1 pupal exuviae, Blackwater River State Forest Biological Station, Santa Rosa County, Florida, USA, 23 - V- 1973 (WLGC); 1 pupal exuviae, Rock Springs, Orange County, Florida, USA, 21 - IV- 1970 (USNM). S. gracilis: 1 pupal exuviae, no locality / date (ZSMC). S. limnophila: 1 pupal exuviae, Jam Tin Creek, Malelane, East Transvaal, South Africa, 2 - XII- 1973 (NMSA); 2 pupal exuviae, Sterk River, S. W., Potgietersrus, Transvaal, South Africa, 3 - I- 1974 (USNM). S. lutea: 1 pupal exuviae, 10 km W. Old Chelsea, Quebec, Canada, 15 - VII- 1986 (CNCI); 1 pupal exuviae, Patuxent Rescue Center, Prince George’s County, Maryland, USA, 17 - V- 1976 (WLGC); 1 pupal exuviae, Beltsville, Prince George’s County, Maryland, USA, 28 - V- 1975 (WLGC); 2 pupal exuviae, Mud Creek, Tompkins County, Freeville, New York, USA, 19 - VI- 1963 (USNM); 2 pupal exuviae, Hamilton-Essex, Newcomb, New York, USA, 17 - VI- 1959 (NYSM); 1 pupal exuviae, as previous locality, 22 - V- 1959 (USNM); 1 pupal exuviae, as previous locality, 28 - VI- 1958 (NYSM); 1 pupal exuviae, as previous locality, 25 - V- 1959 (NYSM); 1 pupal exuviae, as previous locality, 24 - V- 60 (NYSM); 1 pupal exuviae, as previous locality, 29 - V- 1959 (NYSM); 1 pupal exuviae, as previous locality, 11 - V- 1959 (NYSM); 1 pupal exuviae, Letchworth State Park, New York, USA, 13 - VI- 1963 (USNM); 2 pupal exuviae, 1 km downstream lower Cascade Falls, Giles County, Virginia, USA, 3 - VII- 1977 (VPIC); 1 pupal exuviae, Alexandria, Virginia, USA, 13 - V- 1958 (WLGC). S. navaiae: 13 pupal exuviae (of paratypes), Cranberry Lake, St. Lawrence County, New York, USA, 25 - VI- 1963 (USNM); 4 pupal exuviae, 5 km E of Danby, Vermont, USA, 25 - VI- 1986 (CNCI); 1 pupal exuviae, 5 km E. Danby, Vermont, USA, 25 - 26 - VI- 1986 (CNCI). S. orientis: 1 pupal exuviae (of holotype), Burgeshall, Hazyview, East Transvaal, South Africa, 3 - XII- 1973 (NMSA). S. pallidiventris: 5 pupal exuviae, McLean Reserve, Tompkins County, New York, USA, 18 - VI- 1963 (2 WLGC, 1 CNCI, 2 USNM); 3 pupal exuviae, Allegany State Park, New York, USA, 28 - V- 1963 (1 WLGC, 1 CNCI, 1 USNM); 1 pupal exuviae, Ivory, Chautauqua County, New York, USA, 31 - V- 1963 (USNM); 5 pupal exuviae, Ona, Hardee County, Florida, USA, 6 - XI- 1983 (USNM). S. papillata: 1 pupal exuviae, Tien-Shan, Issyk-Kul Province, Kyrgyzstan, 24 - VI- 1971 (ZIN); 2 pupal exuviae, Cholpon-Atinka river, Tien-Shan, Kyrgyzstan, 24 - VI- 1971 (ZIN). S. pictipes: 3 pupal exuviae, Hillman Rr. Margin, Darban, Western Australia, Australia, 29 - X- 1985 (ANIC); 1 pupal exuviae, Warriewood, New South Wales, Australia, 29 - IX- 1956 (ANIC); 2 pupal exuviae, as previous locality, 4 - XI- 1956 (ANIC); 1 pupal exuviae, Griffith, New South Wales, Australia, 16 - XI- 1956 (ANIC). S. rabelloi: 3 pupal exuviae, 5 km NE of Tarcoles, Costa Rica, 26 - VII- 1993 (CNCI); 1 pupal exuviae, as previous locality, 20 - VII- 1993 (CNCI). S. sybleae: 1 pupal exuviae, McLean Reserve, Tompkins County, New York, USA, 18 - VI- 1963 (WLGC); 1 pupal exuviae, Dismal Swamp, Camden County, North Carolina, USA, 25 - III- 1976 (VPIC); 12 pupal exuviae, Gainsville, Alachua County, Florida, USA, 20 - IV- 1967 (5 USNM, 5 WGLC, 2 CNCI); 1 pupal exuviae, Rock Springs, Orange County, Florida, USA, 21 - IV- 1970 (USNM); 1 pupal exuviae, Dyke Marsh, Alexandria, Virginia, USA, 29 - VI- 1952 (USNM); 1 pupal exuviae, Dyke Swamp, Alexandria, Virginia, USA, 11 - VI- 1952 (USNM). S. thomsenae or S. bulla: 1 pupal exuviae, Montgomery County, Virginia, USA, 23 - VI- 1977 (VPIC). S. sp.: 1 pupal exuviae, Negrito Creek Reserve, Catron County, New Mexico, USA, 21 - IX- 1992 (CNCI); 1 pupal exuviae, Grandview Natural Preserve, Hampton, Virginia, USA, 30 - VII- 1976 (VPIC); 3 pupal exuviae, 3 km N of Caldera, Costa Rica, 17 - XII- 1993 (CNCI); 1 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 20 - VIII- 1967 (BPBM); 1 pupal exuviae, as previous locality, 4 - III- 1968 (BPBM); 1 pupal exuviae, Houng River, Moung Sayaboury, Sayaboury Province, Laos, 30 - V- 1967 (BPBM); 2 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 12 - II- 1967 (BPBM); 1 pupal exuviae, as previous locality, 22 - I- 1967 (BPBM); 1 pupal exuviae, as previous locality, 23 - VIII- 1967 (BPBM); 1 pupal exuviae, Phu Khi Minh Mountain, Moung Sayaboury, Sayaboury Province, Laos, 8 - I- 1967 (BPBM); 1 pupal exuviae, Bindoon, Western Australia, Australia, 27 - X- 1985 (ANIC); 1 pupal exuviae, Weeli Woli Spring, Fortescue River, Pilbara, Western Australia, Australia, 17 - X- 1995 (ANIC); 1 pupal exuviae, Mooka Ruins, Gascoyne River, Western Australia, Australia (ANIC); 1 pupal exuviae, Gap Creek trib. Fitzroy River, Western Australia, Australia (ANIC); 1 pupal exuviae, Lady Carrington Drive, Royal National Park, New South Wales, Australia (ANIC); 1 pupal exuviae, McCarrs Creek, New South Wales, Australia, 14 - I- 1969 (ANIC).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD113037FD871D4248E7E5F0.taxon	description	(Figs. 11 H, 20 E – F, 25 C, 29 T, 36 C, 44 L – M, 49 C, 60 B, 74 F – G)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD113037FD871D4248E7E5F0.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the terminal process with a posterior row of thick spines and no other thick spicules present (Fig. 74 G); also unique in the thoracic D- 1 - T much more stout than the other dorsal seta (Fig. 29 T).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD113037FD871D4248E7E5F0.taxon	description	DESCRIPTION: Habitus as in Fig. 11 H. Total length = 1.19 – 2.16 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face (as in Fig. 15 D). Ecdysial tear around base of antenna (as in Figs. 15 D, 79 E); along prothoracic extension. Head: Dorsal apotome (Figs. 20 E – F), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 B) separated from scutum by thin cuticle, separate from scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 25 C) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 36 C) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Figs. 20 E – F) — 1 moderately elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, no campaniform sensillum; clypeal-labrals (Fig. 25 C) — 2 minute setae; oculars (Fig. 25 C) — 2 elongate setae. Thorax: Prothoracic extension (Fig. 25 C) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, extending posteromedially, completely dividing metathorax medially (Fig. 49 C); respiratory organ (Figs. 44 L – M) length / width = 4.92 – 7.08, elongate, slender, somewhat flattened dorsoventrally, with pores closely abutting at apex of respiratory organ, arranged in single row, with additional more basal pores, outer surface smooth, with moderately elongate pedicel, base with short posteromedial apodeme, membranous base of respiratory organ moderately elongate, tracheal tube straight to slightly curved along length, with spirals restricted to base, distally with plate; wing (Fig. 36 C) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (similar to Fig. 32 L) broadly abutting; halter apex just posterior to knob-like extension of tergite 2; legs (Fig. 36 C) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 32 L); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 2 setae, 1 thicker than other; anterolaterals — 3 setae; dorsal setae (Fig. 29 T) — D- 1 - T, D- 2 - T, D- 4 - T setae with D- 1 - T stouter than others, D- 3 - T campaniform sensillum, D- 5 - T absent, D- 3 - T posteromedial to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 49 C) — 1 seta, 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: without pigmentation pattern, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with slightly rounded to pointed, or serrate short tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Figs. 74 F – G) with two dorsomedial short tubercles, one anterior to other, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex, with posterior row of spines; sensilla: tergite 1 (Fig. 49 C) with 3, 5 setae, 2 – 3 campaniform sensilla, including 1 lateral sensillum, 1 seta close, D- 7 - I situated posteriorly near D- 8 - I; segment 4 (Fig. 60 B) — D- 2 - IV, D- 3 - IV short setae on serrate tubercles; D- 5 - IV barely visible short seta (or perhaps a campaniform sensillum), D- 8 - IV, D- 9 - IV short seta; D- 5 - IV, D- 4 - IV, D- 7 - IV, D- 8 - IV, D- 9 - IV on short, serrate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 7 - IV, D- 8 - IV; L- 1 - IV short seta, on pointed tubercle, well anterior of posterior lateral setae; L- 2 - IV, L- 3 - IV short setae, L- 4 - IV thick seta on serrate tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV moderately elongate setae on serrate tubercles; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Figs. 74 F – G) — with D- 5 - IX campaniform sensillum, D- 6 - IX possibly absent (not discernable).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD113037FD871D4248E7E5F0.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Schizonyxhelea is known from nine species from the New World and Oriental Regions (Borkent 2014, additions below). Schizonyxhelea bulla has been reared from mud from the margins of a bog, from the grassy margin of a marsh and from mud, pond weeds and Sphagnum (Thomsen 1937, Wirth & Grogan 1981). Borkent (2000 a) found a larva of S. forattinii in wet mud in a small seep which flowed into the outflow of a larger spring in Costa Rica.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD113037FD871D4248E7E5F0.taxon	discussion	TAXONOMIC DISCUSSION: This genus previously included only two Neotropical species (Borkent 2014) (Tables 2 – 3). In his description of the larva and pupa of S. forattinii, Borkent (2000 a) pointed out the striking similarities especially between the pupae of Schizonyxhelea forattinii Wirth & Grogan and Stilobezzia bulla Thomsen. A number of these are here interpreted as synapomorphies of the genus (characters 24, 54, 76, 79). Nearly all male Schizonyxhelea have a unique and distinctive transverse band of cuticle ventral to the parameres (at about midlength), that is clearly a synapomorphy of the genus. It is uncertain what this structure is, considering that it is likely that a very reduced set of more ventrally located small sclerites is homologous to the aedeagus of other Ceratopogonidae. The male of S. forattinii lacks the ventral transverse band of cuticle but does have a pair of broad extensions from the midlength of the gonocoxites which extend dorsally to the parameres and overlap medially. A further synapomorphy of all species examined here is the reduced set of small sclerites making up the aedeagus. Some other adult features may be synapomorphies (Borkent 2000 a) but further study is needed. Historically, the shape of the claws of female adults has been used as a primary character to identify and recognize genera. The females of the two species originally placed in Schizonyxhelea (S. forattinii, S. guyana) had equal claws on each leg, a single, bent spermatheca and reduced wing venation. The additional species recognized here have single claws on each leg (with a basal tooth) and are similar to those of Stilobezzia, with two welldeveloped spermathecae and wings with two radial cells. I have examined a female from Costa Rica which has a wing venation identical to those of S. forattinii and S. guyana but has single claws and two spermathecae. I have examined other, non-related, Stilobezzia with equal claws and the same variation occurs, for example, in Serromyia (Borkent & Bissett 1990) and Alluaudomyia (pers. obs. unnamed species). Clearly, single or equal claws are variable within these genera and should be used with caution in distinguishing other genera. On the basis of the distinctive male genitalia and pupal synapomorphies, this genus now includes: Schizonyxhelea brevicostalis (Das Gupta & Wirth), 1968: 28 (Stilobezzia). Malaysia, new combination. Schizonyxhelea bulla (Thomsen), 1935: 289 (Stilobezzia). USA, new combination. Schizonyxhelea caribe (Lane & Forattini), 1958: 208 (Stilobezzia). Panama, new combination. Schizonyxhelea diminuta (Lane & Forattini), 1958: 209 (Stilobezzia). Panama, new combination. Schizonyxhelea forattinii Wirth & Grogan, 1988: 81. Brazil. Schizonyxhelea guyana Clastrier, 1984: 2. French Guiana. Schizonyxhelea obscura (Lane & Forattini), 1958: 216 (Stilobezzia). Panama, new combination. Schizonyxhelea panamensis (Lane & Forattini), 1958: 218 (Stilobezzia). Panama, new combination. Schizonyxhelea thomsenae (Wirth), 1953: 83 (Stilobezzia). USA, new combination. Schizonyxhelea scutata (Lane & Forattini), 1961: 92 (Stilobezzia). Panama, new combination. Harris (1981) described seven species of Stilobezzia from Australia. Her species 6 is a Schizonyxhelea, with its characteristic features (i. e. thoracic sensilla D- 1 - T short and stout, distribution of abdominal 4 sensilla with each on a serrate tubercle, posteromedial margin of terminal process with row of spines). I have also examined a reared series of an unnamed species from Laos and there are a number of unnamed species (based on adults) from Costa Rica. Clearly the genus is more diverse and broadly distributed than was previously recognized. Wirth & Grogan (1981) described the pupa of S. bulla (as a Stilobezzia) but did not describe or illustrate the posterior row of spines which are actually present on the terminal process. Borkent (2000 a) missed CL- 1 - H and CL- 2 - H on the mouthparts of the pupa as well as D- 9 - IV on abdominal segment 4 (the single specimen was dirty). Stilobezzia insolita Das Gupta & Wirth from Malaysia is known only as a female but it has a reduced wing venation and an apically swollen and basally curved spermatheca, all similar to some Schizonyxhelea (e. g. S. forattinii). However, until further study, this species remains in Stilobezzia.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD113037FD871D4248E7E5F0.taxon	materials_examined	MATERIAL EXAMINED: S. bulla: 1 pupal exuviae, Algonquin Park, Ontario, Canada, 8 - VI- 1960 (USNM); 1 pupal exuviae, Patuxent Rescue Center, Prince George’s County, Maryland, USA, 17 - V- 1976 (USNM); 2 pupal exuviae, as previous locality, 4 - VI- 1976 (USNM); 1 pupal exuviae, as previous locality (WLGC); 1 pupal exuviae, Allegany State Park, New York, USA, 28 - V- 1963 (WLGC); 2 pupal exuviae, McLean Reserve, Tompkins County, New York, USA, 18 - VI- 1963 (WLGC); 1 pupal exuviae, Bergen Swamp, Genesee County, New York, USA, 14 - VI- 1963 (WLGC); 2 pupal exuviae, Research Station Pond, Newcomb, New York, USA, 30 - VI- 1958 (NYSM); 1 pupal exuviae, Meadow Hole, E. Berne, New York, USA, 29 - VI- 1963 (NYSM); 1 pupal exuviae, Wet Meadow Hole, E. Berne, New York, USA, 24 - VI- 1963 (NYSM); 1 pupal exuviae, Mitchell’s Meadow Pool, E. Berne, New York, USA, 29 - VII- 1963 (NYSM); 1 pupal exuviae, Lake Jimmy, Tahawus, New York, USA, 24 - VI- 1959 (NYSM); 1 pupal exuviae, Mud Pond Outlet, Blue Mountain Lake, New York, USA, 8 - VI- 1959 (NYSM); 1 pupal exuviae, Sphagnum Bog, Blue Mountain Lake, New York, USA, 7 - VI- 1959 (NYSM); 1 pupal exuviae, Salmon River, Blue Mountain Lake, New York, USA, 8 - VI- 1959 (NYSM); 1 pupal exuviae, Beartown Mt. Bog, Virginia, 16 - VIII- 1977 (VPIC); 1 pupal exuviae, Flag Glade, Pocahontas County, Virginia, USA, 24 - VI- 1976 (VPIC); 1 pupal exuviae, Jefferson Nt. Forest, Little Meadows, Giles County, Virginia, USA, 6 - IX- 1976 (VPIC). S. forattinii: 1 pupal exuviae, 2 km NE Tarcoles, Costa Rica, 17 - XII- 1993 (reared from larva) (CNCI). S. sp.: 1 pupal exuviae, Moung Sayaboury, Sayaboury Province, Laos, 22 - II- 1967 (BPBM); 1 pupal exuviae, as previous locality, 19 - II- 1967 (BPBM); 5 pupal exuviae, Ban Na Tak, Moung Sayaboury, Sayaboury Province, Laos, 9 - XII- 1967 (BPBM).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6C3035FD511A824F24E064.taxon	description	(Figs. 20 G, 25 D, 30 A, 36 D, 44 N, 49 D, 60 C, 74 H)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6C3035FD511A824F24E064.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with abdominal segment 4 sensilla L- 1 - IV, L- 2 - IV and L- 3 - IV close together on a rounded tubercle (Fig. 60 C).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6C3035FD511A824F24E064.taxon	description	DESCRIPTION: Total length = 3.10 – 4.19 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, midleg, wing (as in Figs. 15 D, 33 B). Ecdysial tear medial to base of antenna (as in Figs. 15 D, 79 D). Head: Dorsal apotome (Fig. 20 G), without ventral line of weakness, with short dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 B) separated from scutum by thin cuticle, but partially fused to scutum upon emergence, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 25 D) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna (Fig. 36 D) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 20 G) — 1 short seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 – 2 short setae (1 specimen with one very short, 1 elongate seta), 1 campaniform sensillum; clypeallabrals (Fig. 25 D) — absent; oculars (Fig. 25 D) — 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 25 D) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 49 D); respiratory organ (Fig. 44 N) length / width = 2.64 – 4.09, moderately elongate, wide apically, somewhat flattened laterally or circular in cross-section, with pores closely abutting or slightly separated at apex of respiratory organ, arranged in single row, outer surface smooth, with short, wide pedicel, base without posteromedial apodeme, membranous base of respiratory organ moderately elongate, tracheal tube straight to slightly curved along length, with spirals restricted to base or to half length, distally with plates; wing (Fig. 36 D) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 A) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 36 D) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 32 L); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 1 seta; anterolaterals — 1 seta; dorsal setae (Fig. 30 A) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T setae, D- 3 - T campaniform sensillum; D- 1 - T, D- 2 - T on single tubercle, D- 3 - T posterior to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 49 D) — 1 small seta, 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: without pigmentation pattern, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded short tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 74 H) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 49 D) 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. 60 C) — D- 2 - IV peg-like seta, D- 3 - IV short slender setae, on rounded tubercles; D- 5 - IV, D- 8 - IV short setae, D- 9 - IV absent; D- 5 - IV on short separate tubercle, D- 4 - IV and D- 8 - IV on short separate tubercle, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 8 - IV; D- 7 - IV near L- 1 - IV; L- 1 - IV on rounded tubercle with L- 2 - IV, L- 3 - IV all short setae, L- 4 - IV on short tubercle, V- 5 - IV, V- 6 - IV, V- 7 - IV short setae, on rounded tubercles, V- 5 - IV, V- 6 - IV on single tubercle; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 74 H) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6C3035FD511A824F24E064.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Serromyia is known from 35 species from every Region worldwide other than the Neotropical Region (Borkent & Bissett 1990; Borkent 2014). Species have been collected from bogs, fens, wet meadows, streams or small rivers. At least within the Holarctic Region, species are restricted to wooded areas. Strenzke (1950) and Thienemann (1950) found immatures in mosses at lake margins in Germany and Kettle & Lawson (1952) found larvae in mud associated with marshlands.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6C3035FD511A824F24E064.taxon	discussion	TAXONOMIC DISCUSSION: There are only two species of Serromyia known as pupae (with one of these actually of uncertain identity) (Tables 2 – 3). Of the five specimens of Serromyia available for study, one was of S. atra and previously studied by Borkent & Bissett (1990). It included a pharate adult, ensuring correct identification of this specimen. Of the remaining specimens, three were of pupal exuviae and one of a whole pupae, all previously slide mounted, with old, similar labels (perforated edges), with the same handwriting (likely that of Thienemann), and at least one collected by Strenzke (1950, site 220, Grosser Ukleisee, Germany), with the notation " gez. " (short for gezüchtet, meaning reared). Strenzke (1950: 349) noted that larvae were " not rare in damp mosses [in German] ". All were previously identified as S. femorata (probably by Strenzke), even though no associated adult material was present. The pupa was young, so there were no pharate adult features to be studied. As such, there is a level of uncertainty as to both the species identification (Borkent & Bissett 1990) as well as the generic identity. However, it appeared that all four were conspecific and Strenzke likely correctly identified the emerged adults as Serromyia, considering their distinctively fat and spinose hind femora. The pupae also matched the description of S. femorata by Kettle & Lawson (1952) and were very similar to that of S. atra, further suggesting that the generic identification of these specimens is correct. Borkent and Bissett (1990) missed the sensillum coeloconica D- 7 - IV on abdominal segment 4 of the Serromyia pupa they illustrated.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6C3035FD511A824F24E064.taxon	materials_examined	MATERIAL EXAMINED: S. atra: 1 pupa, Schöhsees, Germany (ZSMC). S. femorata: 1 pupa, 2 pupal exuviae, no locality, date (likely Thienemann collection, ZSMC); 1 pupal exuviae, Grosser Ukleisee, Germany (likely Thienemann collection, ZSMC).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6D3032FD5718A14C41E13C.taxon	description	(Figs. 15 E, 20 H, 25 E, 30 B, 36 E, 44 O, 49 E, 61 A, 74 I)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6D3032FD5718A14C41E13C.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the thorax and abdomen with moderately developed rounded tubercles (Figs. 30 B, 61 A, as in 33 A) and the respiratory organ with a separate pore at midlength, in addition to those at its apex and, in one other species, at its base (Fig. 44 O).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6D3032FD5718A14C41E13C.taxon	description	DESCRIPTION: Total length = 3.25 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, midleg, wing (Fig. 15 E, as in Fig. 33 B). Ecdysial tear extending into eye sheath (Figs. 15 E, 79 F). Head: Dorsal apotome (Fig. 20 H), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 25 E) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 36 E) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 20 H) — 1 short seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, no campaniform sensillum; clypeal-labrals (Fig. 25 E) — 2 minute setae; oculars (Fig. 25 E) — 2 setae, 2 campaniform sensilla. Thorax: Prothoracic extension (Fig. 25 E) wide, well-developed, extending from palpus to antenna; mesonotum with well-developed moderately sized tubercles, not extending posteromedially, with slight protuberance, not dividing metathorax medially (Fig. 49 E); respiratory organ (Fig. 44 O) length / width = 3.42, moderately elongate, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single row, with 3 additional more basal pores, outer surface smooth, with moderately elongate, wide pedicel, base with very slender posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, distally with plate; wing (Fig. 36 E) with slight angle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 A) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 36 E) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 32 L); with apex of foreleg ventral to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 2 setae, 1 thicker than other; anterolaterals — 1 very short seta (or perhaps a campaniform sensillum but with shagreen making identification uncertain); dorsal setae (Fig. 30 B) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T very short setae, D- 3 - T campaniform sensillum; D- 1 - T, D- 2 - T, D- 4 - T on well-developed tubercle, D- 5 - T on separate well-developed tubercle, D- 3 - T posterior to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 49 E) — 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax (difficult to discern among shagreen). Abdomen: without pigmentation pattern, 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. 74 I) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 49 E) with 5 setae, 1 campaniform sensillum (difficult to discern in shagreen), including 2 lateral sensilla, 1 seta?, D- 7 - I absent (or not discernible in shagreen); segment 4 (Fig. 61 A) — D- 2 - IV peg-like seta on elongate tubercle, D- 3 - IV short seta on low tubercle; D- 5 - IV peg-like seta, D- 8 - IV, D- 9 - IV short setae, D- 7 - IV absent or not visible among shagreen; D- 5 - IV and D- 4 - IV on elongate tubercle, D- 8 - IV and D- 9 - IV on elongate tubercle, posterior dorsal sensilla on two tubercles in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 8 - IV, D- 9 - IV; L- 1 - IV short peg on elongate tubercle, well anterior of posterior lateral setae; L- 2 - IV, L- 3 - IV short setae on separate elongate tubercles, L- 4 - IV short seta without tubercle, V- 5 - IV, V- 6 - IV, V- 7 - IV short setae, on rounded tubercles, V- 6 - IV, V- 7 - IV on single tubercle; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 74 I) — with D- 5 - IX campaniform sensillum, D- 6 - IX possibly absent (dense shagreen present).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6D3032FD5718A14C41E13C.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Allohelea is known from 57 species from every Region worldwide (Borkent 2014). The two species known as immatures have been collected among grass on a stream margin or from damp soil in a swampy area (Glukhova 1979 a, de Meillon 1939).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6D3032FD5718A14C41E13C.taxon	discussion	TAXONOMIC DISCUSSION: There are only two species of Allohelea known as pupae (Tables 2 – 3). In A. japonica, the spirals are restricted to the base of the tracheal tube and the remaining portion is composed of distinctive (and unique) plates. Because de Meillon (1939) does not show any modifications to the tracheal tube of A. mimas the modifications in A. japonica should considered an autapomorphy of that species for the present. De Meillon (1939) suggested there were no sensilla on the abdomen of A. mimas but likely the pronounced tubercles and heavy shagreen made these difficult to see (as is the case for A. japonica).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6D3032FD5718A14C41E13C.taxon	materials_examined	MATERIAL EXAMINED: A. japonica: 1 pupal exuviae, Komarovskii Nature Reserve, Primorskii Territory, Russia (ZIN).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6A3030FD5C19994F68E081.taxon	description	(Figs. 20 I, 25 F, 30 C, 36 F, 44 P, 49 F, 61 B – C, 74 H)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6A3030FD5C19994F68E081.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the mesonotum with at least one moderately elongate tubercle bearing some sensilla (Fig. 30 C), with three ventral setae (V- 5 - IV, V- 6 - IV, V- 7 - IV) on very short tubercles (Figs. 61 B-C) and the respiratory organ with pores restricted to its apex (Fig. 44 P).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6A3030FD5C19994F68E081.taxon	description	DESCRIPTION: Total length = 2.09 – 2.38 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, midleg, wing (as in Figs. 15 D, 33 B). Ecdysial tear posterior to base of antenna or just medial to base (as in Figs. 15 D, 79 D, E); along prothoracic extension. Head: Dorsal apotome (Fig. 20 I), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (probably as in Fig. 13 H) probably fused to scutum (inadequate material available), each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 25 F) with mandible welldeveloped, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 36 F) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 20 I) — 1 short seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 very short seta, 1 campaniform sensillum; clypeal-labrals (Fig. 25 F) — 1 slender seta; oculars (Fig. 25 F) — 1 seta, 2 campaniform sensilla. Thorax: Prothoracic extension (Fig. 25 F) wide, well-developed but narrow dorsolaterally, extending from palpus to antenna; mesonotum with short to well-developed moderately sized tubercles, not extending posteromedially, with short protuberance, not dividing metathorax medially (Fig. 49 F); respiratory organ (Fig. 44 P) length / width = 3.71 – 4.22, moderately elongate, somewhat flattened dorsoventrally, with pores closely abutting at apex of respiratory organ, arranged in single row (but see taxonomic discussion), outer surface smooth, with moderately elongate, wide pedicel, base without posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to basal third, distally smooth or with reticulations; wing (Fig. 36 F) with angle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 A) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 36 F) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 32 L); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — absent (or perhaps very short among shagreen); anterolaterals — 1 seta (shagreen making it difficult to interpret if more); dorsal setae (Fig. 30 C) — D- 1 - T, D- 2 - T, D- 4 - T very short setae, D- 5 - T either very short or a campaniform sensillum (difficult to discern difference), D- 3 - T campaniform sensillum; D- 1 - T, D- 2 - T, D- 3 - T, D- 4 - T on single well-developed tubercle; supraalar 2 — campaniform sensillum; metathoracics (Fig. 49 F) — 1 very small seta, 1 campaniform sensillum; 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: without pigmentation pattern, segment 2 as wide or slightly wider than segment 3, segments with undivided, peg-like or thin to thick setae, with rounded, very short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 74 H) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 49 F) with 6 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. 61 B-C) — D- 2 - IV peg-like seta, D- 3 - IV elongate seta, D- 2 - IV on short or elongate tubercle, D- 3 - IV on short tubercle; D- 5 - IV, D- 8 - IV peg-like setae, D- 9 - IV short seta, D- 4 - IV absent or not visible in M. hirsuta; on short or large separate 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 L- 1 - IV; L- 1 - IV short seta, on rounded or elongate tubercle with L- 3 - IV; L- 2 - IV, L- 3 - IV, L- 4 - IV short setae without tubercles (or very short) or well-developed tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV short setae, on rounded tubercles, V- 6 - IV, V- 7 - IV closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 74 H) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6A3030FD5C19994F68E081.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Monohelea is known from 88 species from every Region worldwide (Borkent 2014). Immatures have been reared from Sphagnum moss from lake margins and a swamp, or Sphagnum bogs.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6A3030FD5C19994F68E081.taxon	discussion	TAXONOMIC DISCUSSION: There are only two species of Monohelea known as pupae (Tables 2 – 3). The species differ significantly in that M. hirsuta has well-developed tubercles on the abdomen (Fig. 61 C), very similar to that of Allohelea japonica, while in M. obscura these are very short. The pupal exuviae of M. obscura was on a separate slide without an associated adult. It had been collected by H. A. Jamnback and was slide mounted by W. W. Wirth. Although likely correctly associated (Jamnback certainly had ample experience with rearing Ceratopogonidae), there remains a question of its identity. The fourth abdominal segment of M. hirsuta (Fig. 61 C) is illustrated without sensillum D- 4 - IV but this sensillum coeloconica (in other ceratopogonids) is likely present but hidden among the strong shagreen present over much of the abdomen of this species. Harris (1981) also described the pupae of three species she identified as Monohelea (as sp. n. 1, sp. n. 2, sp. n. 3). Since then Wirth & Grogan (1988) recognized the status of six genera formerly placed in Monohelea, including four genera known from Australia: Monohelea, Allohelea, Austrohelea, and Downeshelea. The reared adults were never described but Harris (1981: 114) noted that " M. sp. n. 2 (a ' picture wing' species) is definitely different from M. sp. n. 1 and M. sp. n. 2 [sic] (plain wing species) at the pupal stage ". Because of this obvious error, it is uncertain which species of these three are actually " picture wing " (i. e. possibly Monohelea, Downeshelea, or Allohelea) and which are most likely Austrohelea (plain wing) as far as adult features are concerned (one possible exception is Monohelea tigrina (Skuse), an Australian species with plain wings). However, named species of Austrohelea are known only from Western Australia and Tasmania, while the material of Harris (1981) came from Queensland. Additionally, her pupal key on pg. 92 identifies the three species but within that key Monohelea sp. n. 2 actually refers to M. sp. n. 1 in the text and figures and her keyed Monohelea sp. n. 1 actually refers to the texts and figures of M. sp. n. 2. In the current study, material from elsewhere was available for all these genera other than Downeshelea. Based on pupal features described by Harris (1981), all three would be members of Monohelea based on the presence of L- 1 - IV directly dorsal to L- 3 - IV, a feature unique to this genus within this group of genera (but based on only two species) in its exact relative placement of these two sensilla (see character 65). In addition all three have a peg-like D- 5 - T on the mesothorax, a feature here recognized here as a synapomorphy of Atyphohelea but not scorable with the material I examined of the closely related Monohelea because the presence of D- 5 - T was uncertain in the two species (and two specimens) available because of the small size of the sensilla amongst heavy shagreen, as well as poor perspective. However, Monohelea and related genera in the present scheme do not have a full complement of posterior dorsal sensilla illustrated as present in M. sp. n. 1 (D- 4 - IV, D- 5 - IV, D- 7 - IV, D- 8 - IV, D- 9 - IV) and the two species I examined have most sensilla on segment 4 as short pegs (in M. sp. n. 1 only V- 5 - IV was apparently peg-like), making the generic identification of M. sp. n. 1 uncertain. Both M. sp. n. 2 and M. sp. n. 3 would fit the diagnosis of Monohelea as presented here. However, all three species had at least one extra, separate subapical pore on the respiratory organ that is not present in the two species I examined. Further to this, if two of these three species did have clear wings (but were correctly identified at the time in the broadly encompassing concept of Monohelea), they could only belong to Austrohelea, a genus here considered more distantly related to Monohelea and without L- 1 - IV directly dorsal to L- 3 - IV for the one species I examined. Unfortunately, the location of the material described by Harris (1981) is uncertain and could not be reexamined (Marlene Elson-Harris, pers. comm. 1993). The respiratory organ of M. hirsuta was drawn by Wirth & Grogan (1981) with an apparent longitudinal ridge or thickening but examination of the original material indicated that this was actually a flattened, possibly partially collapsed portion of the respiratory organ. The pupae of Monohelea hirsuta, Allohelea, some Parabezzia and Atyphohelea are similar to one another, with pronounced tubercles on the thorax and abdomen. However, Monohelea have L- 3 - IV ventral to L- 1 - IV, while in Allohelea and Atyphohelea L- 1 - IV is further anterior than any of the other lateral sensilla. Parabezzia are missing L- 1 - IV.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD6A3030FD5C19994F68E081.taxon	materials_examined	MATERIAL EXAMINED: M. hirsuta: 1 pupal exuviae (of holotype), Patuxent Wildlife Refuge, Prince George County, Maryland, USA, 2 - VIII- 1977 (USNM). M. obscura: 1 pupal exuviae, Blue Mountain Lake, New York, USA, 10 - VI- 1958 (NYSM).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD683031FD6F18C24F43E60E.taxon	description	(Figs. 15 F, 20 J, 25 G, 30 D, 37 A, 44 Q, 50 A, 62 A, 74 K)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD683031FD6F18C24F43E60E.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the thorax and abdomen with moderately developed rounded tubercles (Figs. 30 D, 62 A, as in 33 A), with three ventral setae (V- 5 - IV, V- 6 - IV, V- 7 - IV) on well-developed tubercles (Fig. 62 A) (not as in Fig. 61 A) and the respiratory organ with pores restricted to its apex (Fig. 44 Q).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD683031FD6F18C24F43E60E.taxon	description	DESCRIPTION: Total length = 2.20 – 2.94 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, midleg, wing (Fig. 15 F, as in Fig. 33 B). Ecdysial tear extending into eye sheath (Figs. 15 F, 79 F). Head: Dorsal apotome (Fig. 20 J), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 25 G) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 37 A) anterior to barely posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 20 J) — 1 short seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 very short seta, 1 campaniform sensillum; clypeal-labrals (Fig. 25 G) — 2 short, slender setae; oculars (Fig. 25 G) — 1 seta, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 25 G) wide, well-developed, extending from palpus to antenna; mesonotum with welldeveloped moderately sized tubercles, not extending posteromedially, with slight protuberance, not dividing metathorax medially (Fig. 50 A); respiratory organ (Fig. 44 Q) length / width = 2.63 – 3.45, moderately elongate, somewhat flattened dorsoventrally, 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 moderately elongate, annulated, tracheal tube straight to slightly curved along length, with annulations to 3 / 4 length; wing (Fig. 37 A) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 A) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 37 A) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 32 L); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg slightly ventral to, partially abutting apex of midleg laterally; sensilla: anteromedials — 1 very short seta, 1 campaniform sensillum; anterolaterals — 1 short seta (shagreen making it difficult to interpret if more); dorsal setae (Fig. 30 D) — D- 1 - T, D- 2 - T, D- 5 - T peg-like setae, D- 4 - T seta, D- 3 - T campaniform sensillum; D- 1 - T, D- 2 - T, D- 4 - T on single tubercle, D- 5 - T on separate tubercle, D- 3 - T posterior to D- 4 - T; supraalar 2 — campaniform sensillum (difficult to discern among shagreen); metathoracics (Fig. 50 A) — 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: without pigmentation pattern, 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. 74 K) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 50 A) with 7 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. 62 A) — D- 2 - IV peg-like seta, D- 3 - IV moderately elongate seta, D- 2 - IV on large tubercle, D- 3 - IV on short tubercle; D- 5 - IV, D- 8 - IV peg-like setae, D- 9 - IV short seta, D- 7 - IV absent or not visible among shagreen; D- 5 - IV, D- 8 - IV on elongate separate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 8 - IV, D- 9 - IV; L- 1 - IV elongate seta on rounded tubercle just anterior to L- 3 - IV; L- 2 - IV; L- 3 - IV, L- 4 - IV short setae on rounded tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV moderately elongate setae, on rounded well-developed tubercles, V- 6 - IV, V- 7 - IV closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 74 K) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD683031FD6F18C24F43E60E.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Atyphohelea is known from one Nearctic species and one unnamed species from Taiwan (Borkent 1998). Adults are associated with smaller bodies of water, seeps, bogs, and small marshes. Pupae have been reared from a lake margin in North Carolina, U. S. A.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD683031FD6F18C24F43E60E.taxon	discussion	TAXONOMIC DISCUSSION: Only one species of Atyphohelea is known as a pupa (Tables 2 - 3). The fourth abdominal segment of A. macroneura is illustrated without sensilla D- 4 - IV and D- 7 - IV but these campaniform sensilla (their form in other Ceratopogonidae) are likely present but hidden among the strong shagreen present over much of the abdomen of this species (in addition the limited material was rather dirty, further obscuring observation of campaniform sensilla). In nearly all Ceratopogonidae, the anteroventral margin of the wing forms a near right angle, with the margin completely or nearly perpendicular to the dorsal margin of the antenna. I found only two exceptions in the family. In the somewhat crushed specimens of A. macroneura studied here, the angle appeared less abrupt and more rounded (the wing side of the angle was greater). However, I could not convince myself that it was not a product of distortion. Fresh material would be able to test this idea. If present, the feature would be nearly unique in the family. It was also present in an exuviae of one unnamed species of Jenkinshelea (from Kruger N. P) which also had a greater angle, clearly an instance of homoplasy. Other Jenkinshelea had a nearly right angle anteroventral margin.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD683031FD6F18C24F43E60E.taxon	materials_examined	MATERIAL EXAMINED: A. macroneura: 3 pupal exuviae, Highlands Lake Ravenel, Macon County, North Carolina, USA, 10 - VI- 1986 (CNCI).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD69303FFD581E484F9CE3D4.taxon	description	(Figs. 16 A – B, 20 K – L, 25 H, 30 E, 33 A, 37 B, 44 R – S, 50 B, 62 B – C, 74 L, 75 A)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD69303FFD581E484F9CE3D4.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with a well-developed prothoracic extension (Fig. 25 H) and the mesonotum with only 2 setae (D- 1 - T, D- 2 - T) and D- 3 - T (campaniform sensillum) (Fig. 30 E). Also, only pupa of Ceratopogonidae with a well-developed prothoracic extension (Fig. 25 H) and the abdominal segment 4 with 9 - 10 sensilla (Figs. 62 B-C) (lacking D- 3 - IV, L- 1 - IV, L- 4 - IV, V- 7 - IV and, in some, D- 9 - IV).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD69303FFD581E484F9CE3D4.taxon	description	DESCRIPTION: Total length = 1.50 – 2.59 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, midleg, wing (Figs. 16 A – B, 33 A). Ecdysial tear medial to antennal base (as in Figs. 15 D, 79 D) or tearing into eye, either posteriorly or posterolaterally (Figs. 16 A – B, 79 F). Head: Dorsal apotome (Figs. 20 K – L), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 25 H) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna (Fig. 37 B) equal to posterior to, posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Figs. 20 K – L) — 1 short seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 very short seta, 1 campaniform sensillum or 2 setae (only in P. petiolata); clypeallabrals (Fig. 25 H) — absent or 1 short peg; oculars (Fig. 25 H) — 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 25 H) wide, well-developed, extending from palpus to antenna; mesonotum with or without welldeveloped moderately sized tubercles, not extending posteromedially, with or without slight protuberance, not dividing metathorax medially (Fig. 50 B); respiratory organ (Figs. 44 R – S) length / width = 1.95 – 2.82, short, squat, somewhat flattened dorsoventrally, with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface with some wrinkles, with or without short, wide pedicel, base with short posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with annulations to half length; wing (Fig. 37 B) with apical tubercle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33 A) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 37 B) with lateral margin of foreleg near midlength of wing with abrupt angle or evenly curved; hind leg visible at lateral margin of wing (Fig. 33 A); with apex of foreleg slightly to moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 1 very short seta, 1 campaniform sensillum; anterolaterals — 1 short seta; dorsal setae (Fig. 30 E) — D- 1 - T, D- 2 - T peg-like setae, D- 3 - T campaniform sensillum; D- 2 - T, D- 3 - T on single tubercleor closely approximated tubercles; supraalar 2 — campaniform sensillum; metathoracics (Fig. 50 B) — 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: with 2 spots medially on tergites 1 - 7 or tergites 1 - 3 and fading posteriorly, segment 2 as wide or slightly wider than segment 3, segments with undivided, peg-like or thin to thick setae, with rounded to pointed or apically toothed, short to moderately elongate tubercles, tergites or sternites entire, not membranous or sternites 4 - 7 or 4 - 8 each with narrow membranous disc; segment 9 (Figs. 74 L, 75 A) not strongly modified, terminal processes closely approximated basally, each projecting nearly posteriorly to posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 50 B) with 5 setae, 2 campaniform sensilla or 7 setae, 2 campaniform sensilla, including 4 lateral sensilla, D- 2 - I present, D- 3 absent, D- 7 - I situated anterolaterally near L- 1 - I; segment 4 (Figs. 62 B-C) — D- 2 - IV peg-like seta on short to moderately elongate tubercle, D- 3 - IV absent; D- 5 - IV peg-like seta, D- 8 - IV, D- 9 - IV (when present) short setae; only D- 8 - IV on short tubercle or D- 5 - IV, D- 8 - IV, D- 9 - IV on moderately elongate, separate tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 8 - IV or D- 5 - IV, D- 4 - IV, D- 8 - IV, D- 9 - IV; L- 1 - IV absent, L- 2 - IV, L- 3 - IV short setae on pointed or toothed tubercles, L- 4 - IV absent, V- 5 - IV, V- 6 - IV short setae, V- 5 - IV without tubercle or on short toothed tubercle, V- 6 - IV on short toothed or pointed tubercle, V- 7 - IV absent; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Figs. 74 L, 75 A) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD69303FFD581E484F9CE3D4.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Parabezzia is known from 40 species from nearly every Region worldwide but absent from most of the Palaearctic (only in north Africa) and the Australasian Region (Borkent 2014). Pupae have been collected on the edges of streams, rivers and ponds. Grogan & Wirth (1977 a) reported that P. alexanderi pupae were the most common of all ceratopogonids at a lily pond in Maryland, USA.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD69303FFD581E484F9CE3D4.taxon	discussion	TAXONOMIC DISCUSSION: There are seven species of Parabezzia known as pupae (Tables 2 – 3). There are two quite different types of pupae in this genus. One has numbers of rounded tubercles (Figs. 33 A, 62 B), the other, called the alexanderi group, has sharp tubercles (Fig. 62 C) and abdominal sternites with membranous discs (e. g. P. alexanderi, P. balseiroi). All species of Parabezzia have reduced numbers of sensilla on every part of the body (see diagnosis). Some species have no clypeal-labral setae (P. bystraki, P. stagni) while others have a single peg (P. huberti, P. alexanderi, P. downesi, P. petiolata, P. balseiroi), a condition unique in the family. Grogan & Wirth (1977 a) provided a key to the pupae of the four Nearctic species known at that time and Wirth & Grogan (1981) included a key to three species at Plummer's Island, Maryland, USA, including the newly described pupa of P. bystraki. Harris (1981) and Elson-Harris (1990) described the pupa of " Ceratopogon sp. n. 1 " from Australia which would fit the diagnosis and distinctive description of Parabezzia here (the reared adult was not subsequently described). The described presence of eight sensilla on abdominal segment 4 (she likely missed the small campaniform sensilla D- 4 - IV and D- 7 - IV, hence a total of 10) is unique within the Ceratopogoninae for Parabezzia. However, at present there are no Parabezzia otherwise known from the Australasian Region. One potential candidate may be Heteroceratopogon, a genus related to Parabezzia and present in Australia. Unfortunately, the original material has been lost (Marlene Elson-Harris, pers. comm. 1993). If Harris (1981) correctly associated the larvae with the pupa and the above reidentification is correct, she provides the only larval description of a species of Parabezzia (or the only known larva and pupa of Heteroceratopogon). Grogan & Wirth (1977 a) illustrated the dorsal apotomes of species of Parabezzia with DA- 2 - H being a very short seta but reexamination of original material indicates that these are campaniform sensilla, as in nearly all other Ceratopogonidae. Previous descriptions (Table 2) missed D- 7 - IV (campaniform sensillum) on abdominal segment 4.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD69303FFD581E484F9CE3D4.taxon	materials_examined	MATERIAL EXAMINED: P. alexanderi: 1 pupal exuviae, Lakeland Pond, College Park, Prince George’s County, Maryland, USA, 11 - VI- 1975 (USNM); 3 pupal exuviae, as previous locality, 27 - V- 1975 (2 WLGC, 1 CNCI); 2 pupal exuviae, as previous locality, 30 - V- 1975 (1 USNM, 1 CNCI). P. balseiroi: 2 pupal exuviae (of paratypes), Santa Ana, Entre Rios, Argentina, 9 - XI- 1984 (MLPA). P. bystraki: 1 pupal exuviae, Patuxent Wildlife Rescue Center, Prince George’s County, Maryland, USA, 20 - VII- 1976 (USNM). P. downesi: 9 pupal exuviae (of paratypes), Algonquin Park, Ontario, Canada, 8 - VI- 1960 (3 CNCI, 6 USNM). P. eupetiolata: 1 pupal exuviae (of paratype), 1 pupal exuviae (of allotype), Glenfield, Independence River, Lewis County, New York, USA, 22 - VI- 1963 (USNM). P. huberti: 1 pupal exuviae, Patuxent Wildlife Rescue Center, Prince George’s County, Maryland, USA, 20 - VII- 1977 (USNM); 4 pupal exuviae (of paratypes), Falls Church, Virginia, USA, 22 - VII- 1951 (USNM); 2 pupal exuviae (of paratypes), Mount Solon, Virginia, 4 - VII- 1951 (USNM). P. stagni: 1 pupal exuviae (of holotype), Doornpan, Bulge River, North Transvaal, South Africa, 6 - XI- 1973 (NMSA). P. sp.: 1 pupal exuviae, 5 km E of Dandy, Vermont, USA, 25 - VI- 1986 (CNCI).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD67303CFD6618314C72E0AC.taxon	description	(Figs. 20, 26 A, 30 F, 31 K, 37 C, 44 T, 50 C, 63 A, 75 B)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD67303CFD6618314C72E0AC.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with abdominal segment 4 with only two lateral sensilla (L- 2 - IV, L- 3 - IV) and with most sensilla at the bases of bifid tubercles (Fig. 63 A).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD67303CFD6618314C72E0AC.taxon	description	DESCRIPTION: Total length = 2.91 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of face, midleg, wing (as in Figs. 16 B, 33 B). Ecdysial tear uncertain but not around base of antenna; at least medial to antennal base (as in Figs. 15 B, 79 D). Head: Dorsal apotome missing; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side uncertain in whole pupa; mouthparts (Fig. 26 A) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 37 C) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 20) — uncertain; dorsolateral cephalic sclerite sensilla — 1 elongate seta, 1 campaniform sensillum; clypeallabrals (Fig. 26 A) — 1 small seta; oculars (Fig. 26 A) — 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 26 A) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, extending posteriorly but medial division of metathorax uncertain (Fig. 50 C); respiratory organ (Fig. 44 T) length / width = 4.71, elongate, slender, somewhat flattened, at least apically (perhaps more), with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface smooth, with short, wide pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ moderately elongate, tracheal tube straight to slightly curved along length, surface smooth; wing (Fig. 37 C) with apical tubercle lateral to apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 A) narrowly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 37 C) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 32 L); with apex of foreleg slightly anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 2 elongate setae, 1 campaniform sensillum (Fig. 31 K); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 30 F) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T setae, D- 3 - T campaniform sensillum; D- 1 - T, D- 2 - T on single tubercle, D- 3 - T posteromedial to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 50 C) — 1 seta on one side only, 2 campaniform sensilla on both sides; M- 3 - T near anterior margin of metathorax. Abdomen: without pigmentation pattern, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to bilobed, short to somewhat elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 75 B) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 50 C) 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. 63 A) — D- 2 - IV short seta on bifid tubercle; D- 3 - IV moderately elongate seta without tubercle; D- 5 - IV, D- 8 - IV, D- 9 - IV short setae; at base on bifid, separate 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 situated anterorlaterally near D- 3 - IV; L- 1 - IV absent, L- 2 - IV, L- 3 - IV short setae each at base of bifid tubercle, L- 4 - IV absent, V- 5 - IV, V- 6 - IV, V- 7 - IV moderately elongate setae, at base of bifid, slender tubercles, V- 6 - IV, V- 7 - IV closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 75 B) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD67303CFD6618314C72E0AC.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Echinohelea is known from 27 species from nearly every Region worldwide but is absent from the Palaearctic Region (Borkent 2014). An unpublished record from Borneo is known (pers. obs.). The only known immature, the pupa of E. lanei described here, was collected from Flanders, New York, USA. The Flanders Pond site was scrubby red maple growth flooded with a few inches of clear water (Hugo Jamnback, pers. comm.). Grogan (1975) reported a reared specimen of E. lanei collected from under bark of a rotting log in Maryland, USA and Knausenberger (1987) found this species in damp dark brown mud (his collection No. 29) in a roadside ditch in Virginia, USA. De Meillon & Wirth (1991) noted two African species reared from wet soil and from mud at a pond margin.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD67303CFD6618314C72E0AC.taxon	discussion	TAXONOMIC DISCUSSION: Only one moderately well preserved pupa of Echinohelea is known, that of E. lanei (Tables 2 – 3). Wirth (1994 b) provided an earlier description. Details of the sensilla distribution of abdominal segment 4 differs here because the specimen was distorted by compression and is reconstructed here. The dorsal apotome of the specimen was not present and cannot, therefore, be characterized. Wirth (1994 b) noted that the pupa of Echinohelea was similar to some Heteromyiini (as described by Elson-Harris & Kettle 1986) and this is reflected in the phylogenetic relationship supported here (and Borkent 1995) between Echinohelea and Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD67303CFD6618314C72E0AC.taxon	materials_examined	MATERIAL EXAMINED: E. lanei: 1 pupal exuviae, Flanders Pond, Flanders, New York, USA, 4 - VI- 1963 (NYSM).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD64303DFD5F19514873E799.taxon	description	(Figs. 13 F, 16 C, 20 M – N, 26 B, 30 G, 33 B, 37 D, 45 A – B, 50 D, 63 B, 75 C)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD64303DFD5F19514873E799.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the metathorax with only two campaniform sensilla (M- 2 - T, M- 3 - T) on the anterior margin of the metathorax, D- 7 - I situated anteriorly, about 1 / 2 way between D- 5 - I and D- 2 - I (Fig. 50 D, but see taxonomic discussion) and the dorsal apotome relatively narrow (Figs. 20 M-N; compared to Fig. 20 O).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD64303DFD5F19514873E799.taxon	description	DESCRIPTION: Total length = 2.59 – 4.47 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (Figs. 16 C, 33 B). Ecdysial tear around base of antenna tear posterolaterally (Figs. 16 C, 79 E); along prothoracic extension. Head: Dorsal apotome (Figs. 20 M – N), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (Fig. 13 F) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 26 B) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 37 D) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Figs. 20 M – N) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 26 B) — 2 slender setae; oculars (Fig. 26 B) — 2 elongate setae. Thorax: Prothoracic extension (Fig. 26 B) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially, with or without slight protuberance, not dividing metathorax medially (Fig. 50 D); respiratory organ (Figs. 45 A – B) length / width = 3.59 – 7.65, elongate, slender, gradually somewhat flattened dorsoventrally, with pores closely abutting or slightly separated at apex of respiratory organ, arranged in single row, outer surface smooth, with moderately elongate pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ moderately elongate, plain or partly annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, remainder with wrinkles; wing (Fig. 37 D) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33 B) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 37 D) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33 B); with apex of foreleg slightly to moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally, small ventral lobe; sensilla: anteromedials — 1 elongate or 2 short setae; anterolaterals — 1 short seta; dorsal setae (Fig. 30 G) — D- 1 - T, D- 2 - T, D- 5 - T peg-like, D- 4 - T seta, D- 3 - T campaniform sensillum; D- 1 - T, D- 2 - T on single tubercle, D- 3 - T lateral to slightly posterolateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 50 D) — 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: pigmentation very light brown or just indicated by bare patches, tergite 1 with 3 medial spots, tergites 2 – 7 with medial area with stripe, 2 anterolateral spots, sternites not pigmented or sternites 3 – 7 with bare median patch; anterolateral areas not evident, segment 2 as wide or slightly wider than segment 3, segments with undivided, peg-like or thin to thick setae, with rounded to pointed or bifid, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 75 C) not strongly modified, terminal processes separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 50 D) with 7 – 8 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. 63 B) — D- 2 - IV peg-like seta, D- 3 - IV seta, D- 2 - IV on bifid tubercle, D- 3 - IV on short tubercle; D- 5 - IV peg-like seta, D- 8 - IV, D- 9 - IV short setae; D- 5 - IV on single tubercle, D- 8 - IV, D- 9 - IV on single bifid tubercle or separate but closely approximated, 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 pointed tubercle, well anterior of posterior lateral setae; L- 2 - IV, L- 3 - IV, L- 4 - IV short setae on elongate tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV short setae on bifid tubercles, all closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 75 C) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD64303DFD5F19514873E799.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Clinohelea is known from 40 species from every Region worldwide (Borkent 2014). Immatures are known from the margins of streams, fens, seepage area into a lake, muddy margins of rice paddies, and rock pools (de Meillon & Wirth 1991, Knausenberger 1987).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD64303DFD5F19514873E799.taxon	discussion	TAXONOMIC DISCUSSION: There are seven species of Clinohelea known as pupae (Tables 2 – 3). Spinelli & Duret (1993) included the differences between the two Neotropical species known in their key to the adults. Elson-Harris & Kettle (1986) keyed the two known Australian species. Wirth & Grogan (1979) illustrated the dorsal apotome of C. bimaculata with an excavated dorsal margin but reexamination of this species indicates that this was likely based on a damaged or misdrawn specimen. Elson-Harris & Kettle (1986) illustrated abdominal segment 4 of C. dryas with D- 7 - IV in a posterior position, which would be unique within a broader group, including the Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini (see character 63); the species should be reexamined. They likely overlooked both D- 4 - IV and D- 7 - IV in their illustrations of C. tasmaniensis. Elson-Harris (1990) used the enlarged base upon which DA- 1 - H is situated (on the dorsal apotome) to distinguish Australian Heteromyiini. Within this region (and elsewhere), the feature does not characterize Pellucidomyia and therefore is not inclusive. The base is large and swollen in Clinohelea (Figs. 20 M – N) but I could not consistently use this feature to distinguish this genus from some Sphaeromiini s. lat. (e. g. Fig. 21 D, F). The strongly tapering dorsal portion of the dorsal apotome of Clinohelea pointed out by Elson-Harris (1990) is distinctive within the Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD64303DFD5F19514873E799.taxon	materials_examined	MATERIAL EXAMINED: C. bimaculata: 1 pupal exuviae, Spruce Knob Lane, Randolph County, Virginia, USA, 21 - VII- 1976 (VPIC). C. curriei: 2 pupal exuviae, 1 pupal exuviae (in glycerin), 6 km E of Salmon Arm, BC, Canada, 6 - VI- 1990 (CNCI). C. horacioi: 2 pupal exuviae, Pipeline Road, Gamboa, Canal Zone, Panama, VII- 1967 (MLPA). C. nigripes: 1 pupal exuviae, Pipeline Road, Gamboa, Canal Zone, Panama, VII- 1967 (MPLA). C. unimaculata: 2 pupal exuviae, Severskii Donets River, Donetsk Province, Ukraine, 2 - IV- 1970 (ZIN).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD65303AFD401FFA4935E4E4.taxon	description	(Figs. 16 D – E, 20 O, 26 C, 30 H, 37 E, 45 C – D, 50 E, 63 C, 75 D – F)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD65303AFD401FFA4935E4E4.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with only one campaniform sensillum (DA- 2 - H) on the dorsal apotome (Fig. 20 O), the metathorax with only two campaniform sensilla (M- 2 - T, M- 3 - T) on the anterior margin of the metathorax and with a long slender terminal process, lacking an elongate seta (Figs. 75 D – F).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD65303AFD401FFA4935E4E4.taxon	description	DESCRIPTION: Total length = 5.31 – 7.13 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (Fig. 16 E, as in Fig. 33 B). Ecdysial tear around base of antenna, with narrow connection between face and base of antenna (Figs. 16 D – E, 79 G); along prothoracic extension. Head: Dorsal apotome (Fig. 20 O), with ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 26 C) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna (Fig. 37 E) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Fig. 20 O) — 1 short to moderate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 26 C) — 2 slender setae; oculars (Fig. 26 C) — 2 elongate setae, 0 – 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 26 C) wide, well-developed, narrow dorsolaterally, not extending to antenna; mesonotum without or with 1 spinous tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 50 E); respiratory organ (Figs. 45 C – D) length / width = 3.12 – 5.90, elongate, slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single curved row, outer surface with or without some wrinkles, with short to moderately elongate pedicel, base with moderate elongate posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, remainder with wrinkles to near apex; wing (Fig. 37 E) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 A) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 37 E) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33 B); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally in male, ventral to apex of midleg in female; sensilla: anteromedials — 1 short, 1 long seta; anterolaterals — 1 moderately long seta; dorsal setae (Fig. 30 H) — D- 1 - T, D- 2 - T, D- 5 - T peg-like or elongate setae, D- 4 - T seta, D- 3 - T campaniform sensillum or D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T all elongate, D- 3 - T campaniform sensillum, D- 3 - T lateral to slightly posterolateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 50 E) — 1 - 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: with tergite 1 with 3 medial spots (fused in some), tergites 2 - 7 with medial area with stripe, 2 spots, anterolateral spots, sternites 3 - 7 with medial, anterolateral spot, segment 2 as wide or slightly wider than segment 3, segments with undivided, peglike or thin to thick setae, with pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Figs. 75 D – F) not strongly modified, terminal processes separated basally, each projecting posterodorsolaterally, elongate, slender, tapering to pointed, hooked apex; sensilla: tergite 1 (Fig. 50 E) with 8 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. 63 C) — D- 2 - IV peg-like seta, D- 3 - IV seta, on short tubercles; D- 5 - IV peg-like seta, D- 8 - IV short setae, D- 9 - IV moderately elongate seta; D- 5 - IV on short tubercle, D- 8 - IV, D- 9 - IV on separate but closely approximated short 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 tooth on pointed tubercle, just anterior of L- 3 - IV; L- 2 - IV, L- 4 - IV short setae, L- 3 - IV moderately elongate seta, on pointed tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV on pointed tubercles; V- 5 - IV, V- 6 - IV closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Figs. 75 D-F) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD65303AFD401FFA4935E4E4.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Heteromyia is known from 13 species in the New World (Borkent 2014). Immatures have been found in semi-aquatic or aquatic vegetation, with rearings from Sphagnum and Cabomba. Knausenberger (1987) provides further details of habitats, including swamps and stream and river margins. Adults have been collected from bogs, fens and, in Panama, a hotspring with abundant wet soil and vegetation on its margin.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD65303AFD401FFA4935E4E4.taxon	discussion	TAXONOMIC DISCUSSION: Three species of Heteromyia are known as pupae (Tables 2 – 3). The elongate, slender terminal process (Figs. 75 D – F) is distinctive in the group Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini except for Dibezzia (Fig. 75 I) which has an associated elongate seta (D- 5 - IX) and some species of Bezzia (Fig. 77 E) which have two or more campaniform sensilla on the dorsal apotome (Figs. 22 C – D). Wirth & Grogan (1977) illustrated the dorsal apotome of H. wokei with two setae but reexamination shows the presence of one seta and one campaniform sensillum.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD65303AFD401FFA4935E4E4.taxon	materials_examined	MATERIAL EXAMINED: H. clavata: 1 pupal exuviae, Juan Mina Station, Canal Zone, Panama, 3 - VI- 1939 (USNM). H. pratti: 1 pupal exuviae, Patuxent Rescue Center, Prince George’s County, Maryland, 13 - VI- 1977 (USNM). H. wokei: 4 pupal exuviae, Bluefields, Nicaragua, 25 - VIII- 1943 (USNM).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD623038FD6C1D214D85E38C.taxon	description	(Figs. 12 A, 13 G, 16 F, 20 P, 26 D, 30 I, 33 C, 37 F, 45 E – F, 50 F, 64 A, 75 G – H)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD623038FD6C1D214D85E38C.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the thorax and segment 2 markedly wider than abdominal segment 3 (Fig. 12 A); also unique in having an apically truncate segment 9 and very widely spaced terminal processes (Figs. 12 A, 75 G – H); also unique in having a well-developed prothoracic extension (Fig. 26 D) and a respiratory organ with the pores arranged in a nearly complete circle (Figs. 45 E – F).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD623038FD6C1D214D85E38C.taxon	description	DESCRIPTION: Habitus as in Fig. 12 A. Total length = 2.22 – 2.84 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16 B, 33 C). Ecdysial tear around base of antenna, with narrow connection between face and base of antenna (Figs. 16 F, 79 G). Head: Dorsal apotome (Fig. 20 P), with ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (Fig. 13 G) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 26 D) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 37 F) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Fig. 20 P) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 26 D) — 2 moderately thick setae; oculars (Fig. 26 D) — 1 – 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 26 D) wide, well-developed, narrow dorsolaterally, not extending to antenna; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 50 F); respiratory organ (Figs. 45 E – F) length / width = 2.27 – 2.38, moderately elongate, trumpetshaped, somewhat flattened apically (circular area), with pores closely abutting at apex of respiratory organ, arranged circularly, outer surface smooth, without pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, plates to half length; wing (Fig. 37 F) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33 C) broadly abutting; halter apex extending just anterior of anterolateral margin of tergite 2; legs (Fig. 37 F) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33 C); 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 (1 / 2 distance to anterolaterals) (as in Figs. 31 L – M); anterolaterals — 1 elongate seta; dorsal setae (Fig. 30 I) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T setae, D- 3 - T campaniform sensillum; D- 1 - T, D- 2 - T on single tubercle, D- 3 - T posterior to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 50 F) — 1 - 2 campaniform sensilla; M- 3 - T near anterior margin of metathorax. Abdomen: pigmentation very light brown or just indicated by bare patches, with tergite 1 with 3 medial spots, 2 (medial area with stripe, 2 spots), anterolateral spots, 3 – 7 only light brown medial patch, sternites not pigmented or sternites 3 – 7 with light brown medial patch, segment 2 much wider than segment 3, segments with or without bifurcating setae, with rounded to pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Figs. 75 G – H) widely truncate posteriorly, terminal processes widely separated basally, each projecting posteriorly or slightly posterolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 50 F) with 8 setae, 1 campaniform sensillum or 7 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. 64 A) — D- 2 - IV, D- 3 - IV long setae on moderately elongate tubercles; D- 5 - IV, D- 8 - IV, D- 9 - IV elongate setae, D- 5 - IV, D- 8 - IV simple or bifurcating; D- 5 - IV on 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 moderately elongate seta, close to base of tubercles with L- 2 - IV, L- 3 - IV; L- 2 - IV, L- 3 - IV, L- 4 - IV elongate, simple or bifid setae on elongate tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV elongate simple or bifid setae, on rounded tubercles, V- 6 - IV, V- 7 - IV closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Figs. 75 G- H) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD623038FD6C1D214D85E38C.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Pellucidomyia is known from nine species in the Neotropical, Afrotropical and Australasian Regions (Borkent 2014), including the new combination of P. geari noted below. Pupae have been collected from small streams and creeks, backwaters of a creek, and from the margin of a reservoir.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD623038FD6C1D214D85E38C.taxon	discussion	TAXONOMIC DISCUSSION: Two species of Pellucidomyia are known as pupae (Tables 2 – 3). Pupae of this genus are distinctive (see diagnosis) and easily distinguishable from other Ceratopogonidae. Examination of the pupae of South African species Macropeza geari indicates that it shares the pupal synapomorphies of P. leei and that it belongs within this genus as a new combination as follows: Pellucidomyia geari (de Meillon & Wirth), 1981: 547 (Macropeza). new combination. Pellucidomyia geari is otherwise known only as the male holotype. Because males of Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini are generally poorly characterized in much of the world, a generic misplacement is more than possible. Additionally, the male adult P. geari is very similar to that of P. sambulena (de Meillon) 1942 which likely is a senior synonym. Examination of the types is needed to confirm this possibility.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD623038FD6C1D214D85E38C.taxon	materials_examined	MATERIAL EXAMINED: P. geari: 1 pupal exuviae (of holotype), Burgershall, Hazyview, East Transvaal, South Africa, 3 - XII- 1973 (NMSA). P. leei: 1 pupal exuviae (in glycerin), Mason Creek, Queensland, Australia, no date (CNCI); 4 pupal exuviae, Merricumbene Creek, Moruya River, New South Wales, Australia, 2 - III- 1964 (ANIC).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD603039FD9918374C97E0F4.taxon	description	(Figs. 21 B, 26 F, 30 K, 33 D, 38 B, 45 H, 51 B, 64 C, 75 J)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD603039FD9918374C97E0F4.taxon	diagnosis	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. 51 B) and L- 1 - IV and L- 3 - IV slender, contrasting with the basally thick L- 2 - IV and L- 4 - IV (Fig. 64 C); also unique with the metathorax with only one campaniform sensillum (M- 3 - T) situated posterior from the anterior margin of metathorax (Fig. 51 B), halter extending just anterior of the anterolateral margin of tergite 2 (Fig. 33 D), and abdominal segment 4 with the lateral setae scattered (Fig. 64 C) (not forming a transverse row as in Fig. 67 B).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD603039FD9918374C97E0F4.taxon	description	DESCRIPTION: Total length = 3.59 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16 B, 33 B). Ecdysial tear around base of antenna, with narrow connection between face and base of antenna (as in Figs. 16 E, 79 G). Head: Dorsal apotome (Fig. 21 B), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 26 F) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 38 B) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 21 B) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 26 F) — 2 slender setae; oculars (Fig. 26 F) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 26 F) wide, well-developed, extending from palpus to antenna; mesonotum yes, low ones tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 51 B); respiratory organ (Fig. 45 H) length / width = 3.67 – 3.81, moderately elongate, 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 moderately elongate, annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing (Fig. 38 B) with short tubercle at apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33 D) broadly abutting; halter apex extending just anterior of anterolateral margin of tergite 2; legs (Fig. 38 B) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33 D); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 2 elongate setae (as in Figs. 31 L – M); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 30 K) — D- 1 - T, D- 2 - T, D- 5 - T, setae, D- 4 - T broken seta?, D- 3 - T campaniform sensillum, D- 3 - T anterolateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 51 B) — 1 campaniform sensillum; M- 3 - T distant from margin of metathorax (at least 1 / 3 length of metathorax). Abdomen: with tergite 1 with 1 medial spot, tergites 2 – 7 with medial area with stripe, 2 anterolateral spots, 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, short tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 75 J) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 51 B) 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. 64 C) — D- 2 - IV, D- 3 - IV short to moderately elongate setae on short tubercles; D- 5 - IV, D- 8 - IV, D- 9 - IV 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 moderately elongate seta on short tubercle, just anterior of base of tubercle with L- 2 - IV; L- 2 - IV, L- 3 - IV, L- 4 - IV moderately elongate setae, L- 2 - IV, L- 4 - IV on rounded tubercles, V- 5 - IV, V- 7 - IV moderately elongate, V- 6 - IV short seta, on rounded tubercles, all closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 75 J) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD603039FD9918374C97E0F4.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Hebetula is known from 22 species in the Oriental, Afrotropical and Australasian Regions (Borkent 2014). Pupae have been reared from moss in a creek and from the margin of a river.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD603039FD9918374C97E0F4.taxon	discussion	TAXONOMIC DISCUSSION: Only one species of this genus is known as a pupa (Tables 2 – 3). The dorsal setae on the thorax of the single pupa differed on either side, with D- 3 - T missing on the right side. D- 4 - T was either a sensillum basiconica (as shown here) or the setae were broken off on both sides. If future specimens confirm D- 4 - T as a sensillum basiconica, this would be a distinctive and unique feature within the Ceratopogonidae (D- 4 - T is absent in some taxa). Elson-Harris (1987) illustrated the respiratory organ of H. tonnoiri with a midband of darker pigmentation but reexamination indicates the pigmentation extends to the base of the respiratory organ.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD603039FD9918374C97E0F4.taxon	materials_examined	MATERIAL EXAMINED: H. tonnoiri: 1 pupal exuviae, Nattai River, Mittagong, New South Wales, Australia, 4 - XI- 1964 (ANIC).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD613026FD4E199F4801E6A4.taxon	description	(Figs. 12 B, 21 A, 26 E, 30 J, 38 A, 45 G, 51 A, 64 B, 75 I)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD613026FD4E199F4801E6A4.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with a well-developed prothoracic extension (Fig. 26 E) and an elongate seta (D- 5 - IX) on the base of the terminal process (Figs. 12 B, 75 I).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD613026FD4E199F4801E6A4.taxon	description	DESCRIPTION: Habitus as in Fig. 12 B. Total length = 4.03 mm (5.5 mm from Mayer 1934 c). Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16 B, 33 B). Ecdysial tear uncertain anteriorly; present along prothoracic extension. Head: Dorsal apotome (Fig. 21 A), with partial ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 26 E) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna (Fig. 38 A) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 21 A) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 26 E) — 2 slender setae; oculars (Fig. 26 E) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 26 E) wide, well-developed, extending from palpus but narrow dorsolaterally, extent to antenna uncertain; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 51 A); respiratory organ (Fig. 45 G) length / width = 3.11 – 3.61, moderately elongate, with blunt apex, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface with plates with very fine spicules, without pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, plates to 3 / 4 length; wing (Fig. 38 A) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 A) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 38 A) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33 C); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 2 elongate setae (as in Figs. 31 L – M); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 30 J) — D- 1 - T, D- 2 - T, D- 4 - T setae, D- 5 - T broken or absent, D- 3 - T campaniform sensillum, D- 3 - T lateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 51 A) — 1 campaniform sensillum; M- 3 - T distant from margin of metathorax (at least 1 / 3 length of metathorax). Abdomen: without pigmentation pattern, 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. 75 I) not strongly modified, terminal processes closely approximated basally, each projecting nearly posteriorly, elongate, slender, tapering to pointed apex; sensilla: tergite 1 (Fig. 51 A) with 6 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. 64 B) — D- 2 - IV, D- 3 - IV short to moderately elongate setae, D- 2 - IV on short tubercle; D- 8 - IV, D- 9 - IV moderately elongate setae, D- 5 - IV, D- 7 - IV absent (or perhaps not visible); D- 8 - IV, D- 9 - IV on elongate, single tubercle, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 4 - IV, D- 8 - IV, D- 9 - IV; L- 1 - IV short seta without tubercle, just anterior of base of tubercle with L- 2 - IV; L- 2 - IV, L- 3 - IV, L- 4 - IV moderately elongate setae on elongate tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV moderately elongate setae, without tubercles, all closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 75 I) — with D- 5 - IX elongate seta, D- 6 - IX campaniform sensillum.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD613026FD4E199F4801E6A4.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Dibezzia is known from five species in the Oriental and Afrotropical Regions (Borkent 2014, additional species below). The two species were reared from either " bamboo peat " (Johannsen 1932) in Indonesia or a tree hole in Singapore (Wirth & Ratanaworabhan 1981 a).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD613026FD4E199F4801E6A4.taxon	discussion	TAXONOMIC DISCUSSION: The pupae of this genus are known from two species (Tables 2 – 3). Originally described as Johannsenomyia prominens by Johannsen (1932), Mayer (1934 a: 244, 1934 c) described the pupa of this Indonesian species as (?) Homohelea prominens, based on the pupal exuviae from which the male holotype of this species was reared. Wirth (1973) placed the species in Mallochohelea as a new combination but did not identify it as such or give any justification for doing so and this is where it has remained since. The descriptions by Mayer (1934 a, 1934 c), however, clearly indicate the presence of the elongate seta at the base of the terminal process, a feature of Dibezzia and unique within the Ceratopogoninae. In addition, Mayer (1934 c) noted the presence of " breiten Schuppen " (broad scales) on the surface of the respiratory organ, which is what the fine scales appear as in D. debenhamae. This feature is unique within the Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini. I therefore remove the name from Mallochohelea and consider it a member of Dibezzia as follows: Dibezzia prominens (Johannsen) 1932: 435 (Johannsenomyia). new combination. Dibezzia prominens, as illustrated by Mayer (1934 c), has D- 5 - IV represented as a small campaniform sensillum or perhaps a very small seta and D- 7 - IV located posterolateral to D- 3 - IV (like other members of Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini; see character 64). I could not find either on the single, dirty, and distorted specimen of D. debenhamae I examined.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD613026FD4E199F4801E6A4.taxon	materials_examined	MATERIAL EXAMINED: D. debenhamae: 1 pupal exuviae (of allotype), Singapore, VI- 1959 (USNM).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7E3027FD631EE14F21E4D1.taxon	description	(Figs. 17 A, 21 C, 26 G, 30 L, 33 E, 38 C, 45 I, 51 C, 65 A, 75 K – L)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7E3027FD631EE14F21E4D1.taxon	diagnosis	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. 51 C), abdominal segment 1 with D- 2 - I and D- 3 - I well separated (Fig. 51 C), abdominal segment 4 with L- 1 - IV a short seta (Fig. 65 A), D- 5 - IV on an undivided tubercle, and D- 4 - IV situated medial to D- 8 - IV and D- 9 - IV (not diagnosable as different from Mallochohelea).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7E3027FD631EE14F21E4D1.taxon	description	DESCRIPTION: Total length = 2.22 – 5.63 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (Figs. 17 A, 33 E). Ecdysial tear extending into eye sheath (Figs. 17 A, 79 I). Head: Dorsal apotome (Fig. 21 C), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 26 G) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 38 C) barely anterior to barely posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum); sensilla: dorsal apotomals (Fig. 21 C) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 26 G) — 2 slender or thick setae; oculars (Fig. 26 G) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 26 G) wide, well-developed, extending from palpus to antenna; mesonotum with short to moderately sized tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 51 C); respiratory organ (Fig. 45 I) length / width = 2.38 - 3.72, very short, squat to elongate, 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 short posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing (Fig. 38 C) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33 E) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 38 C) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33 E); 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 — 1 short peg, 1 elongate seta (as in Fig. 31 N); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 30 L) — D- 1 - T, D- 2 - T D- 5 - T peg-like or short, stout setae, D- 4 - T elongate seta, D- 3 - T campaniform sensillum; D- 1 - T, D- 2 - T on single tubercle, D- 3 - T anterolateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 51 C) — 1 campaniform sensillum; M- 3 - T distant from margin of metathorax (at least 1 / 3 length of metathorax). Abdomen: pigmentation light to moderately brown, tergite 1 with 3 medial spots, tergites 2 - 7 with medial area with stripe, 2 anterolateral spots, 2 pairs on tergite 8, sternites not pigmented or sternites 3 - 5 with medial stripe, anterolateral spot (sternites 6 - 7 membranous), 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 tubercles, tergites or sternites entire, not membranous or with sternites 5 - 7 each with membranous disc; segment 9 (Figs. 75 K-L) not strongly modified, terminal processes separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 51 C) with 7 setae, 2 campaniform sensilla, including 4 lateral sensilla, D- 2 - I, D- 3 - I well separated, D- 7 - I situated anterolaterally near L- 1 - I; segment 4 (Fig. 65 A) — D- 2 - IV, D- 3 - IV moderately elongate setae on short tubercles; D- 5 - IV, D- 8 - IV peg-like setae, D- 9 - IV moderately elongate seta; 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 short tubercle, near L- 3 - IV; L- 2 - IV, L- 4 - IV short setae, L- 3 - IV elongate seta, L- 2 - IV on pointed tubercle, L- 3 - IV, L- 4 - IV on short tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV moderately elongate setae, all closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Figs. 75 K – L) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7E3027FD631EE14F21E4D1.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Jenkinshelea is known from 20 species in the Nearctic, Palaearctic, Oriental, and Afrotropical Regions (Borkent 2014). Immatures have been reared from lake, stream, and river margins, rice fields, marshes, ponds and dead leaves in a swamp.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7E3027FD631EE14F21E4D1.taxon	discussion	TAXONOMIC DISCUSSION: Pupae of five species of Jenkinshelea are known (Tables 2 – 3). Male pupae of J. magnipennis from Rideau River, Ottawa, Ontario (females not available) had significantly larger abdominal tubercles bearing D- 2 - IV, D- 5 - IV and D- 8 - IV than did those from nearby Black Lake, Stanleyville, Ontario, likely indicating the presence of more than one species.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7E3027FD631EE14F21E4D1.taxon	materials_examined	MATERIAL EXAMINED: J. albaria: 1 pupal exuviae, Lake Wissota, Chippewa County, Wisconsin, USA, 22 - VII- 1979 (USNM); 1 pupal exuviae, Potomac River, Fairfax County, Virginia, USA, 7 - VI- 1955 (USNM); 9 pupal exuviae, Potomac River at Scott Run, Fairfax County, Virginia, USA, 7 - VI- 1955 (1 USNM, 8 WLGC); 1 pupal exuviae, no locality, VI- 1955 (USNM). J. magnipennis: 9 pupal exuviae, Black Lake, Stanleyville, Ontario, Canada, 29 - VI- 1975 (USNM); 2 pupal exuviae, Rideau River, Ottawa, Ontario, Canada, 29 - V- 1960 (USNM); 1 pupal exuviae, Lac Serpent Notre Dame de Laus, Quebec, Canada, 17 - VI- 1968 (LEMQ). J. polyxenae: 1 pupal exuviae, Bathing Pond, Eshowe, Zululand, South Africa, 24 - IX- 1930 (SAIM). J. rhodesiensis: 1 pupal exuviae, 1 pupal exuviae (of paratype), Msasa Stream, Salisbury, Southern Rhodesia, 30 - III- 1942 (SAIM); 1 pupal exuviae, Sand River between White River and Hazyview, East Transvaal, South Africa, 2 - XII- 1973 (NMSA). J. tokunagai: 1 pupal exuviae, Nam Houng river, Sayaboury Province, Laos, 4 - 7 - III- 1968 (BPBM). J. sp.: 2 pupal exuviae (in glycerin), Ottawa, ON, Canada, 29 - VI- 1975 (CNCI); 1 pupa, 1 pupal exuviae, Lac Serpent Notre Dame de Laus, Quebec, Canada, 17 - VI- 1968 (LEMQ).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7F3025FD5C1D324FBAE01C.taxon	description	(Figs. 12 C, 21 D, 26 H, 30 M, 38 D – E, 45 J, 51 D, 65 B – C, 76 A – B)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7F3025FD5C1D324FBAE01C.taxon	diagnosis	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. 51 D), abdominal segment 4 with L- 1 - IV a short seta (Fig. 65 B) and L- 3 - IV situtated well anterior of L- 2 - IV; also unique with D- 2 - I and D- 3 - I well separated (Fig. 51 D) and L- 1 - VIII present (for all species other than M. natalensis); also unique 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. 51 D) and D- 4 - IV situated between D- 8 - IV and D- 9 - IV (except for M. natalensis but in that species D- 4 - V and D- 4 - VI are between D- 8 and D- 9 of their respective segments).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7F3025FD5C1D324FBAE01C.taxon	description	DESCRIPTION: Habitus as in Fig. 12 C. Total length = 2.59 – 4.88 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16 B, 33 B). Ecdysial tear extending into eye sheath (as in Figs. 17 A, 79 I). Head: Dorsal apotome (Fig. 21 D), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 26 H) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Figs. 38 D – E) anterior to posterior to, posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 21 D) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 26 H) — 2 slender or thick setae; oculars (Fig. 26 H) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 26 H) wide, well-developed, extending from palpus to antenna; mesonotum without or with well-developed moderately sized tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 51 D); respiratory organ (Fig. 45 J) length / width = 2.15 – 4.31, very short, squat to elongate, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface with some wrinkles, with short, wide to moderately elongate pedicel, base with short to elongate posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing (Figs. 38 D – E) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 A) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Figs. 38 D – E) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33 E); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally, small ventral lobe; sensilla: anteromedials — 1 short, 1 elongate seta, 0 – 1 campaniform sensillum (as in Fig. 31 N); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 30 M) — D- 1 - T, D- 2 - T, D- 5 - T peg-like setae or setae, D- 4 - T elongate seta or peg-like seta, D- 3 - T campaniform sensillum; D- 1 - T, D- 2 - T on single tubercle, D- 3 - T lateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 51 D) — 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 (in female only), tergites 2 – 7 with medial area with stripe, 2 anterolateral 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 or bifid, short to moderately elongate tubercles, tergites or sternites entire, not membranous or with sternites 5 – 7 each with membranous disc; segment 9 (Figs. 76 A – B) not strongly modified, terminal processes closely approximated basally, each projecting posterolaterally to nearly laterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 51 D) with 8 setae, 2 campaniform sensilla, including 4 lateral sensilla, D- 2 - I, D- 3 - I well separated, D- 7 - I situated anterolaterally near L- 1 - I; segment 4 (Figs. 65 B-C) — D- 2 - IV, D- 3 - IV elongate setae; D- 2 - IV on elongate or bifid tubercle, D- 3 - IV on bump or bifid tubercle; D- 5 - IV peg-like seta, D- 8 - IV, D- 9 - IV short to moderately elongate setae, D- 7 - IV present or absent; D- 5 - IV on single bifid 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 or D- 5 - IV, D- 8 - IV, D- 4 - IV, D- 9 - IV; D- 7 - IV, if present, near D- 3 - IV; L- 1 - IV short to moderately elongate seta on short tubercle, near L- 3 - IV; L- 2 - IV, L- 4 - IV short setae, L- 3 - IV moderately elongate seta, L- 2 - IV, on pointed or bifid tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV short to moderately elongate setae, on short bilobed tubercles, all closely approximated or with only V- 5 - IV, V- 6 - IV close; segment 8 without D- 3 - VIII, with L- 1 - VIII; segment 9 (Figs. 76 A-B) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7F3025FD5C1D324FBAE01C.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Macropeza is known from 22 species in the Nearctic, Palaearctic, Oriental, and Afrotropical Regions (Borkent 2014). Macropeza geari is transferred to Pellucidomyia as a new combination above. Immatures have been collected from mud and among plants at stream and river margins, reservoirs, and from aquatic plants (Pistia stratiotes).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7F3025FD5C1D324FBAE01C.taxon	discussion	TAXONOMIC DISCUSSION: The pupae of four named species of Macropeza have been described (Tables 2 – 3). All species of Macropeza, other than M. natalensis, are nearly unique in having L- 1 - VIII on abdominal segment 8 (see character 71). In addition, nearly all species have another synapomorphy, with D- 4 - IV between D- 8 - IV and D- 9 - IV. In M. natalensis the sensilla is medial to D- 8 - IV and D- 9 - IV but is between the two comparable sensilla on abdominal segments 5 and 6. These observations may suggest that M. natalensis is the sister group to the remaining Macropeza examined or alternately, that the single specimen available had aberrantly distributed sensilla on segment 4.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7F3025FD5C1D324FBAE01C.taxon	materials_examined	MATERIAL EXAMINED: M. albitarsis: 2 pupal exuviae, Piława river, Szwecja nr Wałcz, Poland, 5 - VII- 2007 (IZUG). M. bayeri: 1 pupal exuviae (of paratype), M’posa River, Empangeni, Zululand, South Africa, 14 - I- 1937 (SAIM). M. natalensis: 1 pupal exuviae, Manypuza River, INYONI, Zululand, South Africa, 19 - VIII- 1936 (SAIM). M. sp.: 1 pupal exuviae, Nam Houng River Margin, Moung Sayaboury, Sayaboury Province, Laos, 17 - XII- 1967 (BPBM); 2 pupal exuviae (different species), as previous locality, 17 - 20 - XII- 1967 (BPBM); 1 pupal exuviae, as previous locality, 6 - 9 - I- 1968 (BPBM).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7D3022FD6D18794C84E7FA.taxon	description	(Figs. 2 C, E, 12 D, 13 H, 17 B, 21 E, 27 A, 30 N, 33 F, 38 F, 45 K, 51 E, 66 A, 76 C – D)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7D3022FD6D18794C84E7FA.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the metathorax with only one campaniform sensillum (M- 3 - T) situated posterior from the anterior margin of the metathorax (Fig. 51 E), abdominal segment 1 with D- 2 - I and D- 3 - I well separated (Fig. 51 E), abdominal segment 4 with L- 1 - IV a short seta (Fig. 66 A), D- 5 - IV on an undivided tubercle, and D- 4 - IV situated medial to D- 8 - IV and D- 9 - IV (not diagnosable as different from Jenkinshelea).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7D3022FD6D18794C84E7FA.taxon	description	DESCRIPTION: Habitus as in Fig. 12 D. Total length = 2.81 – 6.50 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (Figs. 17 B, 33 F). Ecdysial tear extending into eye sheath (Figs. 17 B, 79 I). Head: Dorsal apotome (Fig. 21 E), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 27 A) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 38 F) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 21 E) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 27 A) — 2 thick setae; oculars (Fig. 27 A) — 1 – 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 27 A) wide, well-developed nearly even width to narrow dorsolaterally, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 51 E); respiratory organ (Fig. 45 K) length / width = 2.48 – 2.56, very to moderately short, squat, 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 without (?) or with very short posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing (Fig. 38 F) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33 F) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 38 F) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33 F); 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 — 1 short peg, 1 elongate seta (as in Fig. 31 N); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 30 N) — D- 1 - T, D- 2 - T, D- 5 - T peg-like setae, D- 4 - T elongate seta, D- 3 - T campaniform sensillum; D- 1 - T, D- 2 - T on single tubercle, D- 3 - T anterolateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 51 E) — 1 campaniform sensillum; M- 3 - T distant from margin of metathorax (at least 1 / 3 length of metathorax). Abdomen: pigmentation light brown, poorly defined in some; with tergite 1 with 3 medial spots, tergites 2 – 7 with medial area with stripe, 2 anterolateral spots, some with 1 – 2 pairs on segment tergite 8, sternites 3 – 7 (anterolateral spots on 5 – 7 poorly defined, pale) all light brown, or just very light medial strip on sternites 4, 5 and with 6 – 7 membranous or just very light medial strip on sternites 3, 4 and with sternites 5 – 7 membranous or absent, 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, with sternites 5 – 7 or 6 – 7 each with membranous disc; segment 9 (Figs. 76 C – D) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 51 E) with 8 setae, 2 campaniform sensilla, including 3 lateral sensilla, D- 2 - I, D- 3 - I well separated, D- 7 - I situated anteriorly near D- 3 - I; segment 4 (Fig. 66 A) — D- 2 - IV, D- 3 - IV moderately elongate setae, D- 2 - IV on bifid tubercle, D- 3 - IV on short tubercle; D- 5 - IV peg-like seta, D- 8 - IV short seta, D- 9 - IV moderately elongate seta; 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 pointed tubercle, just anterior of base of tubercle with L- 2 - IV; L- 2 - IV, L- 3 - IV, L- 4 - IV short setae on short tubercles, V- 5 - IV, V- 7 - IV short setae, V- 6 - IV elongate seta, on short or moderately elongate tubercles, all closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Figs. 76 C-D) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7D3022FD6D18794C84E7FA.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Mallochohelea is known from 48 species from every Region worldwide (Borkent 2014, with some moved to Anebomyia and Dibezzia). Immatures have been reared from stream and river margins, thermal stream, reservoirs, ponds, and lakes.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7D3022FD6D18794C84E7FA.taxon	discussion	TAXONOMIC DISCUSSION: The pupae of 12 species of Mallochohelea have been described (Tables 2 – 3). Wirth & Grogan (1979) provide a key to the pupae of the two species of Mallochohelea from the Potomac Valley (Washington DC area, USA), as well as Anebomyia atripes, then called M. atripes. Thienemann (1928), Mayer (1934 a) and Lenz (1934) provided keys to a few European species known at that time (as members of either Dicrobezzia (now a synonym of Probezzia) or Johannsenomyia). However, they all recognized M. dentata as distinct from M. munda (the former now considered a synonym of the latter) and both Thienemann (1928) and Mayer (1934 a) additionally recognized M. breviforceps as distinct and it too is now considered a synonym of M. munda.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7D3022FD6D18794C84E7FA.taxon	materials_examined	MATERIAL EXAMINED: M. albibasis: 1 pupal exuviae, Cottonwood Lake, 5 km S of Nelson, British Columbia, Canada, 6 - VII- 2008 (CNCI); 2 pupal exuviae, 7 km S of Hope, British Columbia, Canada, 8 - VIII- 1985 (CNCI); 1 pupal exuviae, Outegamie County, Wisconsin, USA, 15 - VI- 1954 (USNM). M. albihalter: 1 pupal exuviae, Allegany State Park, New York, USA, 28 - V- 1963 (USNM). M. caudellii: 1 pupal exuviae (in glycerin), Bolean Lake, 6 km E of Falkland, BC, Canada, 12 – 13 - VII- 1989 (CNCI). M. errinae: 1 pupal exuviae (of holotype), Klein Yukskei, Johannesburg, Transvaal, South Africa, V- 1939 (SAIM); 1 pupal exuviae, Mageliesberg Agricultural School, Transvaal, South Africa, 13 - XI- 1973 (NMSA). M. inermis: 2 pupal exuviae, Vrevo lake, Leningrad Province, Russia, 19 - VI- 1968 (ZIN); 1 pupal exuviae, no locality / date (ZSMC). M. satelles: 4 pupal exuviae (of paratype), Nattai River, Mittagong, New South Wales, Australia, 18 - II- 1969 (ANIC). M. setigera: 1 pupal exuviae, River Gruzskii Elanchik, Donetsk Province, Ukraine, 29 - IV- 1970 (ZIN). M. smithi: 4 pupal exuviae, Black Lake, Stanleyville, Ontario, Canada, 29 - VI- 1975 (WLGC); 1 pupal exuviae, Rideau River, Ottawa, Ontario, Canada, 29 - V- 1960 (USNM); 1 pupal exuviae, Snow Hill, Maryland, 19 - V- 1968 (USNM); 1 pupal exuviae, 10 mi SE of Middleberg, Lake Dunmore, Vermont, USA, 23 - VI- 1986 (CNCI). M. sybleae: 1 pupal exuviae (of paratype), Centerville, Humboldt County, California, USA, 13 - VIII- 1948 (USNM). M. termophila: 2 pupal exuviae (of paratype), Caimancito, Jujuy, Argentina, 12 - VIII- 1980 (MLPA). M. nr. caudellii: 1 pupal exuviae, 7 km S of Hope, British Columbia, Canada, 8 - VIII- 1985 (CNCI). M. nr. pullata: 3 pupal exuviae, 6 km NE of Falkland, British Columbia, Canada, 12 - VII- 1989 (CNCI). M. sp.: 1 pupal exuviae, Bear Creek, 0.5 mi SE of Karlsbad Springs, Ontario, Canada, 1967 (CNCI); 1 pupal exuviae, McCarrs Creek, New South Wales, Australia, 11 - XI- 1956 (ANIC).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7A3020FD561C444FD6E799.taxon	description	(Figs. 21 F, 27 B, 30 O, 31 N, 33 G, 39 A, 45 L – M, 51 F, 66 B, 76 E)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7A3020FD561C444FD6E799.taxon	diagnosis	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. 51 F), abdominal segment 1 with D- 2 - I and D- 3 - I well separated (Fig. 51 F), abdominal segment 4 with L- 1 - IV a short seta and D- 5 - IV on a bifid tubercle (Fig. 66 B).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7A3020FD561C444FD6E799.taxon	description	DESCRIPTION: Total length = 2.91 – 5.56 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Fig. 16 B, 33 G). Ecdysial tear extending into eye sheath (as in Figs. 17 A, 79 I). Head: Dorsal apotome (Fig. 21 F), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 27 B) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 39 A) barely anterior to equal to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 21 F) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 27 B) — 2 moderately to very thick setae; oculars (Fig. 27 B) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 27 B) wide, well-developed nearly even width to narrow dorsolaterally, extending from palpus to antenna; mesonotum with short to moderately sized tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 51 F); respiratory organ (Figs. 45 L – M) length / width = 2.61 – 4.00, short to intermediate length, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface with a few to numbers of wrinkles, with short, wide pedicel, base with very short, pale posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing (Fig. 39 A) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33 G) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 39 A) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33 G); 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 — 1 short peg or short thick seta, 1 elongate seta (Fig. 31 N); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 30 O) — D- 1 - T, D- 2 - T, D- 5 - T peg-like setae, D- 4 - T seta, D- 3 - T campaniform sensillum; D- 1 - T, D- 2 - T on single tubercle, D- 3 - T lateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 51 F) — 1 campaniform sensillum; M- 3 - T distant from margin of metathorax (at least 1 / 3 length of metathorax). Abdomen: pigmentation light brown in some; with tergite 1 with 3 medial spots, tergites 2 – 7 with medial area with stripe, 2 anterolateral spots, 2 pairs on tergite 8 in some, sternites 3 – 4 (5 – 7 membranous) or 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 or bifid, short to moderately elongate tubercles, tergites or sternites entire, not membranous or with 5 – 7 or 6 – 7 each with membranous disc; segment 9 (Fig. 76 E) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 51 F) with 8 setae, 2 campaniform sensilla, including 3 lateral sensilla, D- 2 - I, D- 3 - I moderately or well separated, D- 7 - I situated anterolaterally near L- 1 - I; segment 4 (Fig. 66 B) — D- 2 - IV, D- 3 - IV short to moderately elongate setae on short tubercles; D- 5 - IV peg-like seta, D- 8 - IV short seta, D- 9 - IV moderately elongate seta; D- 5 - IV on single, bifid 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 to moderately elongate seta on rounded tubercle, just anterior of base of tubercle with L- 2 - IV; L- 2 - IV, L- 4 - IV short setae, L- 3 - IV moderately elongate seta on pointed tubercles, V- 5 - IV, V- 7 - IV short setae, V- 6 - IV elongate seta, on short or moderately elongate tubercles, all closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 76 E) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7A3020FD561C444FD6E799.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Probezzia is known from 28 species, all restricted to the Holarctic Region (Borkent 2014). Larvae are present in the benthos of lakes, reservoirs, rivers and streams and pupae can be found on their margins. The pupae of some species are particularly common in the flotsam on beaches (Williams 1955, pers. obs.). Knausenberger (1987) provides detailed information on the behaviour, ecology and distribution of the immatures of a number of species in the eastern USA.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7A3020FD561C444FD6E799.taxon	discussion	TAXONOMIC DISCUSSION: The pupae of 14 species of Probezzia have been described (Tables 2 – 3). The abdominal membranous discs are present or absent in different species of Probezzia and it is clear that the feature is homoplastic within the genus, with even otherwise very similar species with or without the discs. Knausenberger (1987) discussed the puzzling presence or absence but could not indicate a related ecological factor to explain this character state distribution. Pupae of P. seminigra have abdominal membranous discs on sternites 5 – 7. However, one male, certainly conspecific with others from the same locality, had discs on only the posterior half of 5 and then full ones on 6 – 7, indicating intraspecific variation. The feature is discussed further under Character 50. Study of a series of Probezzia pallida indicated that at least three species are actually present under this name. Specimens from each of North Dakota, of Ottawa, Ontario and Allegeny, New York and of other localities in New York could be distinguished based on the presence or absence of pupal abdominal membranous discs, shapes and sizes of pupal CL- 1 - H, D- 1 - T and D- 2 - T, size and pigmentation of male adult antenna, size of female adult claws, and pigmentation of female adult postnotum.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD7A3020FD561C444FD6E799.taxon	materials_examined	MATERIAL EXAMINED: P. albitibia: 3 pupal exuviae (of paratypess), Algonquin Park, Ontario, Canada, 8 - VI- 1960 (USNM); 2 pupal exuviae, Meachum Lake, New York, USA, 21 - VI- 1986 (CNCI); 3 pupal exuviae (of paratypes), Blue Mountain Lake, Hamilton County, New York, USA, 10 - VI- 1960 (USNM); 2 pupal exuviae (of paratypes), Piercefield, St. Lawrence County, New York, USA, 26 - VI- 1963 (USNM); 1 pupal exuviae (of paratype), Potomac River at Scott Run, Fairfax County, Virginia, 4 - VI- 1955 (USNM). P. flavonigra: 2 pupal exuviae, 10 km SW of Anahim Lake, British Columbia, Canada, 3 - VII- 1992 (CNCI); 2 pupal exuviae, Canoe, British Columbia, Canada, 15 - VII- 1990 (CNCI); 5 pupal exuviae, Sicamous, British Columbia, Canada, 8 - VII- 1989 (CNCI); 1 pupal exuviae, 28 km E of Enderby, British Columbia, Canada, 13 - VII- 1989 (CNCI). P. fuscipennis: 6 pupal exuviae, 10 mi SE of Middleburg, Lake Dunmore, Vermont, USA, 23 - VI- 1986 (CNCI); 1 pupal exuviae (of paratype), Blue Mountain Lake, Hamilton County, New York, USA, 10 - VI- 1960 (CNCI). P. pallida: 4 pupal exuviae, Rideau River, Ottawa, Ontario, Canada, 29 - May- 1960 (1 CNCI, 3 USNM); 6 pupal exuviae, White Earth River, Mountrail County, North Dakota, USA, 7 - VI- 1969 (USNM); 1 pupal exuviae, Allegany State Park, New York, USA, 28 - V- 1963 (USNM); 1 pupal exuviae, Piercefield, St. Lawrence County, New York, USA, 26 - VI- 1963 (USNM); 1 pupal exuviae, Genesee River, Portageville, New York, USA, 13 - VI- 1963 (USNM); 2 pupal exuviae, no locality, VI- 1953 (USNM). P. rosewalli: 1 pupal exuviae (of paratype), Kilbourne, Louisiana, USA, 10 - V- 1947 (USNM). P. sabroskyi: 1 pupal exuviae, 10 km S of Okanagan Falls, British Columbia, Canada, 14 - V- 1989 (CNCI); 1 pupal exuviae, Meachum Lake, New York, USA, 21 - VI- 1986 (CNCI); 1 pupal exuviae, Allegany State Park, New York, USA, 28 - V- 1963 (USNM). P. seminigra: 12 pupal exuviae, Cottonwood Lake, 5 km S of Nelson, British Columbia, Canada, 6 - VII- 1992 (CNCI); 2 pupal exuviae, Canoe, British Columbia, Canada, 15 - VII- 1990 (CNCI); 3 pupal exuviae (in glycerin), Sicamous, BC, Canada, 8 - VII- 1989 (CNCI); 1 pupal exuviae, Rutka Tartak, nr. Suwałki, Poland 2 - VII- 1973 (IZUG); 3 pupal exuviae, Anninskoe lake, Pskov Province, Russia, 23 - 26 - VI- 1995 (ZIN); 1 pupal exuviae, Lower Austria, Lunzer Untersee (the lowest-lying of the three major lakes in the Lunz area), Spongilla aufwuchs in ' middle zone' near S shore, Austria, 27 - VI- 1942 (ZSMC); 2 pupal exuviae, shore of pond at " Zillertal " inn, North Rhine-Westphalia, Riemke N of Bochum, Germany, 9 - VI- 1916 (ZSMC). P. smithi: 3 pupal exuviae, Wolf River, Outaganie County, Wisconsin, USA, 15 - VI- 1954 (USNM); 4 pupal exuviae, Potomac River at Scott Run, Fairfax County, Virginia, USA, 7 - VI- 1955 (USNM). P. williamsi: 2 pupal exuviae, 10 mi SE of Middleburg, Lake Dunmore, Vermont, USA, 23 - VI- 1986 (CNCI). P. xanthogaster: 6 pupal exuviae, Allegany State Park, New York, USA, 28 - V- 1963 (USNM); 8 pupal exuviae, Taughannock Falls, Tompkins County, New York, USA, 15 - VI- 1963 (USNM); 1 pupal exuviae, Genesee River, Portageville, New York, USA, 13 - VI- 1963 (USNM). P. nr. seminigra: 1 pupal exuviae, 10 km S. Okanagan Falls, British Columbia, Canada, 14 - V- 1989 (CNCI). P. sp.: 2 pupal exuviae, Trail Bay, Manitoba, Canada (CNCI); 2 pupal exuviae (in glycerin), Shirley Bay, Ottawa, Ontario, Canada, 13 - VI- 1972, (CNCI); 4 pupal exuviae, Meachum Lake, New York, USA, 21 - VI- 1986 (CNCI); 1 pupal exuviae, 1 km N of Camden, Maine, USA, 1 - VIII- 1985 (CNCI); 1 pupal exuviae, Potomac River at Scott Run, Fairfax County, Virginia, USA, 7 - VI- 1955 (USNM); 2 pupae, Lerchenauer See, Germany, 27 - VI- 1990, 28 - VII- 8 - VIII- 1990 (CNCI); 5 pupal exuviae, Anninskoe lake, Pskov Province, Russia, 23 - 26 - VI- 1995 (ZIN).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD783021FDCA1FFA4F17E429.taxon	description	(Figs. 21 G, 27 C, 30 P, 39 B, 45 N, 52 A, 66 C, 76 F)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD783021FDCA1FFA4F17E429.taxon	diagnosis	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. 52 A), abdominal segment 1 with D- 2 - I and D- 3 - I well separated (Fig. 52 A), and abdominal segment 4 with L- 1 - IV a long seta (Fig. 66 C).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD783021FDCA1FFA4F17E429.taxon	description	DESCRIPTION: Total length = 3.81 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16 B, 33 B). Ecdysial tear extending into eye sheath (as in Figs. 17 A, 79 I). Head: Dorsal apotome (Fig. 21 G), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 27 C) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 39 B) barely anterior to equal to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 21 G) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 27 C) — 2 thick setae; oculars (Fig. 27 C) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 27 C) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 52 A); respiratory organ (Fig. 45 N) length / width = 3.50, very short, squat, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single curved row, outer surface smooth, with short, wide pedicel, base with short posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to 1 / 3 length; wing (Fig. 39 B) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 A) broadly abutting; halter apex abutting anterolateral knoblike extension of tergite 2; legs (Fig. 39 B) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33 E); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 1 short peg, 1 elongate seta, 1 campaniform sensillum (as in Fig. 31 N); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 30 P) — D- 1 - T, D- 2 - T, D- 5 - T peg-like setae, D- 4 - T seta, D- 3 - T campaniform sensillum; D- 1 - T, D- 2 - T on single, short tubercle, D- 3 - T lateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 52 A) — 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, 2 pairs on tergite 8, at least sternites 3 – 5 with medial stripe, anterolateral spot (sternites 6 – 7 membranous), 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, with sternites 6 – 7 each with membranous disc; segment 9 (Fig. 76 F) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 52 A) with 8 setae, 2 campaniform sensilla, including 3 lateral sensilla, D- 2 - I, D- 3 - I well separated, D- 7 - I situated anteriorly near D- 3 - I; segment 4 (Fig. 66 C) — D- 2 - IV, D- 3 - IV short to moderately elongate setae on short tubercles; D- 5 - IV, D- 8 - IV peg-like setae, D- 9 - IV moderately elongate seta; 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 elongate seta on pointed tubercle, well anterior of posterior lateral setae; L- 2 - IV, L- 4 - IV short setae, L- 3 - IV elongate seta on pointed tubercles, V- 5 - IV, V- 7 - IV short setae, V- 6 - IV elongate seta, on short or moderately elongate tubercles, all closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 76 F) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD783021FDCA1FFA4F17E429.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Neobezzia is known from eight species in the Neotropical Region (Borkent 2014). Immatures were found in mats of the floating aquatic fern Salvinia auriculata in a lagoon or among plants and algal mats in a pond.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD783021FDCA1FFA4F17E429.taxon	discussion	TAXONOMIC DISCUSSION: Only two species of Neobezzia are known as pupae (Tables 2 – 3). Ronderos et al. (2011) provided detailed SEM photomicrographs as well as description of N. fittkau. They reported the total length of male pupae as 4.06 – 4.64 mm and that of females as 4.86 – 4.98 mm but included the length of the terminal process (not included here). Regardless, this genus shows the sexual dimorphism in size present in all Heteromyiini + Sphaeromiini s. lat. + Palpomyiini + Stenoxenini and some genera of Ceratopogonini.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD783021FDCA1FFA4F17E429.taxon	materials_examined	MATERIAL EXAMINED: N. fittkaui: 1 pupal exuviae, Ilha da Marchantaria, Município de Iranduba, Amazonas, Brazil, 27 - VII- 2010 (MLPA).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD79302FFD521C6A4F11E679.taxon	description	(Figs. 21 H – I, 27 D, 30 Q, 33 H, 39 C, 45 O – P, 52 B, 67 A, 76 G)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD79302FFD521C6A4F11E679.taxon	diagnosis	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. 52 B), thorax with D- 5 - T a peg-like seta (as in Fig. 30 P) (or a short, stout seta, only in N. curticornis, then on a well-developed tubercle, Fig. 30 Q), and abdominal segment 1 with D- 2 - I and D- 3 - I closely approximated (Fig. 52 B).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD79302FFD521C6A4F11E679.taxon	description	DESCRIPTION: Total length = 3.22 – 7.16 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16 B, 33 H). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17 C, 79 H). Head: Dorsal apotome (Figs. 21 H – I), with ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 27 D) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 39 C) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Figs. 21 H – I) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 27 D) — 0 – 1 slender or 2 thick setae; oculars (Fig. 27 D) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 27 D) wide, well-developed but narrow dorsolaterally, extending from palpus to antenna; mesonotum with very short tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 52 B); respiratory organ (Figs. 45 O – P) length / width = 2.61 – 3.56, elongate, moderately slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface with a few to numbers of wrinkles, with short, wide pedicel, base with short posteromedial apodeme, membranous base of respiratory organ moderately elongate, annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing (Fig. 39 C) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33 H) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2 or past tergite 2; legs (Fig. 39 C) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33 H); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 1 short peg, 1 elongate seta (as in Fig. 31 N); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 30 Q) — D- 1 - T, D- 2 - T peg-like or setae, D- 4 - T seta, D- 5 - T peg-like seta, D- 3 - T campaniform sensillum, D- 3 - T lateral to slightly posterolateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 52 B) — 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 tergite 1 with 3 medial spots, tergites 2 – 7 with medial area with stripe, 2 anterolateral spots, 2 pairs on tergite 8 in some, sternites not pigmented or sternites 3 – 4 or sternites 3 – 7 (5 – 7 membranous and with single medial spot), 2 spots on sternite 8 in some, 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, not membranous or with sternites 5 – 7 each with membranous disc; segment 9 (Fig. 76 G) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; apex of male genital lobe anterior or to level of base of terminal process (as in Figs. 78 I – J), except for N. schwarzii where it extends to about half length of base of terminal process (as in Fig. 78 E); sensilla: tergite 1 (Fig. 52 B) 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. 67 A) — D- 2 - IV, D- 3 - IV short to moderately elongate setae on short tubercles; D- 5 - IV, 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, if present, 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 on pointed tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV short setae on short tubercles, all closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 76 G) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD79302FFD521C6A4F11E679.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Nilobezzia is known from 74 species from every Region worldwide (Borkent 2014; additional species below). Immatures are present in lakes, grassy wetlands, ponds, salt marshes (Knausenberger 1987), reservoirs, creeks, and rivers.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD79302FFD521C6A4F11E679.taxon	discussion	TAXONOMIC DISCUSSION: There are 17 species of Nilobezzia known as pupae (Tables 2 – 3). Only two species, N. schwarzii and N. mallochi, have abdominal membranous discs (see character 50). Nearly all male pupae have short genital lobes, not extending posteriorly past the base of the terminal processes, reflecting the short, modified male adult genitalia; only those of N. schwarzii have more elongate genital lobes (even though the male adult genitalia doesn't seem disproportionally longer). Nilobezzia theileri (de Meillon & Wirth) was originally described as a Sphaeromias (de Meillon & Wirth 1981) but is here considered a member of Nilobezzia as a new combination as follows: Nilobezzia theileri (de Meillon & Wirth), 1981: 552 (Sphaeromias). new combination. The pupa has the diagnostic features of this genus and reexamination of the adult holotype shows that the basal tooth of the claws of the adult female are actually outer and not inner, as described and inaccurately illustrated by de Meillon & Wirth (1981). The adult actually keys to Nilobezzia in the key to the Afrotropical fauna by de Meillon & Wirth (1991). Elson-Harris (1987) keyed the pupae of six Australian species. Elson-Harris (1990) used the position of L- 3 - IV ventral to L- 1 - IV (her lpm ii " in line with " lasm) to distinguish Australian Nilobezzia from Australian Hebetula and Lanatomyia and this feature does indeed work for known Australian taxa.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD79302FFD521C6A4F11E679.taxon	materials_examined	MATERIAL EXAMINED: N. aranea: 1 pupal exuviae (of holotype), 1 pupal exuviae (of allotype), Breakfast Creek, Spencer, New South Wales, Australia, 4 - II- 1965 (ANIC); 1 pupal exuviae (of paratype), as previous locality, 3 - III- 1969 (ANIC); 1 pupal exuviae, McCarrs Creek, New South Wales, Australia, 22 - I- 1969 (ANIC); 1 pupal exuviae, Oystershell Road, New South Wales, Australia, 4 - II- 1965 (ANIC). N. basispinigera: 1 pupal exuviae, Warregor, Engonia, New South Wales, Australia, 11 - XII- 1963 (ANIC). N. brevicornis: 1 pupal exuviae, Ft. Summer # 1, DeBaca County, New Mexico, USA, 21 - VIII- 1967 (USNM); 2 pupal exuviae, Bitter Lakes, Chaues County, New Mexico, USA, 17 - VIII- 1967 (USNM). N. curticornis (as N. triquetrinotata): 1 pupal exuviae (of holotype of N. triquetrinotata), Griffith, New South Wales, Australia, 15 - XI- 1956 (ANIC); 1 pupal exuviae (of allotype of N. triquetrinotata), 6 pupal exuviae (of paratypes of N. triquetrinotata), previous locality (ANIC). N. formosa: 1 pupal exuviae, Kozennyi Torets river, Raigorodok, Donetsk Province, Ukraine, 19 – 30 - V- 1970 (ZIN); 2 pupal, as previous locality, 1 - IV- 1970 (ZIN). N. fuscitarsus: 1 pupal exuviae, Darban, Western Australia, Australia, 28 - X- 1985 (ANIC); 2 pupal exuviae, Humpty Doo, Northern Territory, Australia, 5 - XII- 1958 (ANIC). N. neotropica: 1 pupal exuviae, Lago Monte Aleque, Sao Paulo State, Brazil, 5 - VII- 2000 (CNCI). N. robusta: 1 pupal exuviae; Douglas, Cape Province, South Africa, 23 - I- 1974 (NMSA); 1 pupal exuviae (of paratype), Berg River, Piquetberg, South Africa, 16 - III- 1953 (SAIM). N. schwarzii: 3 pupal exuviae, Lake Providence, Louisiana, USA, VII- 1953 (USNM); 3 pupal exuviae, Sanibel Island, Lee County, Florida, USA, 11 - V- 1973 (USNM); 1 pupal exuviae, Lake Munson, Leon County, Florida, USA, 17 - VIII- 1987 (JHEC); 1 pupal exuviae, 3 km N of Caldera, Costa Rica, 17 - XII- 1993 (CNCI). N. theileri: 1 pupal exuviae (of holotype; with adult female), Loutpan, Transvaal, South Africa, 10 - I- 1974 (NMSA). N. virago: 2 pupal exuviae, Darwin, Northern Territory, Australia, 7 - VI- 1958 (ANIC); 1 pupal exuviae, 1 pupal exuviae (of allotype), Gove, Northern Territory, Australia, II- 1970 (ANIC). N. whartoni: 1 pupal exuviae, Spencer, New South Wales, Australia, 4 - II- 1965 (ANIC); 8 pupal exuviae, Breakfast Creek, Spencer, New South Wales, Australia, 4 - II- 1965 (ANIC); 1 pupal exuviae, Oystershell Road., New South Wales, Australia, 4 - II- 1965 (ANIC). N. sp.: 1 pupal exuviae, Constance Bay, Ontario, Canada, 8 - VI- 1979 (CNCI); 1 pupal exuviae (in glycerin), Kruger National Park, South Africa (ANIC); 9 pupal exuviae, Bindoon, Western Australia, Australia, 27 - X- 1985 (ANIC).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD77302CFD831EDA49E0E77C.taxon	description	(Figs. 21 J, 27 E, 30 R, 39 D, 45 Q, 52 C, 67 B, 76 H – I)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD77302CFD831EDA49E0E77C.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with abdominal segment 4 with all four lateral setae (L- 1 - IV, L- 2 - IV, L- 3 - IV, L- 4 - IV) arranged transversely, closely approximated and on the anterior half of segment (Fig. 67 B).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD77302CFD831EDA49E0E77C.taxon	description	DESCRIPTION: Total length = 4.03 – 5.38 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16 B, 33 B). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17 C, 79 H). Head: Dorsal apotome (Fig. 21 J), without ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 27 E) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna (Fig. 39 D) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 21 J) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeallabrals (Fig. 27 E) — 2 slender setae; oculars (Fig. 27 E) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 27 E) wide, well-developed but narrow dorsolaterally, not extending to antenna; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 52 C); respiratory organ (Fig. 45 Q) length / width = 2.41 – 3.09, moderately elongate, swollen apically, 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 moderately elongate, tracheal tube straight, swollen curved apically in some, with spirals restricted to base, plates to half length; wing (Fig. 39 D) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 A) broadly abutting; halter apex abutting or just anterior to anterolateral knob-like extension of tergite 2; legs (Fig. 39 D) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33 I); male with apex of foreleg moderately anterior to apex of midleg, female with apex of foreleg slightly anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally, small ventral lobe; sensilla: anteromedials — 2 elongate setae (as in Figs. 31 L – M); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 30 R) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T setae, D- 3 - T campaniform sensillum, D- 3 - T anterolateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 52 C) — 1 campaniform sensillum; M- 3 - T distant from margin of metathorax (at least 1 / 3 length of metathorax). Abdomen: with tergite 1 with 1 medial spot, tergites 2 – 7 with medial area with stripe, 2 anterolateral spots, sternites 3 – 7 with anterolateral spots difficult to discern, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded to slightly pointed, short tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Figs. 76 H – I) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally to somewhat laterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 52 C) with 7 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. 67 B) — D- 2 - IV, D- 3 - IV short to moderately elongate setae on short tubercles; D- 5 - IV, D- 8 - IV, D- 9 - IV 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 short tubercle, near L- 2 - IV, L- 3 - IV, L- 4 - IV on anterior half of segment; L- 2 - IV, L- 3 - IV, L- 4 - IV short setae on short pointed tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV short setae on rounded tubercles; V- 5 - IV, V- 6 - IV closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Figs. 76 H-I) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD77302CFD831EDA49E0E77C.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Johannsenomyia is known from 27 species from every Region worldwide but is absent from most of the Palaearctic (known only from north Africa) (Borkent 2014). Immatures have been reared from river margins, lakes and Knausenberger (1987) additionally recorded a species from a pond. Williams (1955) described how mature larvae of J. argentata burrowed into the sand on a lake beach to pupate.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD77302CFD831EDA49E0E77C.taxon	discussion	TAXONOMIC DISCUSSION: Only one species of Johannsenomyia has been described as a pupa (Tables 2 – 3). Mayer (1957) also described the pupa of J. albidorsata (de Meillon) (as a Dicrohelea) but I have examined the specimen and it is in fact a species of Bezzia. De Meillon (1937) in his description of the adult actually noted that the pupal exuviae of the holotype had been mixed with those of a Palpomyia and a Bezzia and its identification was therefore uncertain. Mayer (1957) however was clearly unaware of the uncertain association and described the purported pupal exuviae as if it was that of J. albidorsata. Debenham (1974) gave a brief synopsis of the pupa of Johannsenomyia but based it on the misidentified description by Mayer (1957).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD77302CFD831EDA49E0E77C.taxon	materials_examined	MATERIAL EXAMINED: J. argentata: 1 pupal exuviae, Storrs, Connecticut, USA, 1 - VI- 1953 (USNM); 1 pupal exuviae, Patuxent Wildlife Refuge, Prince George’s County, Maryland, USA, 24 - V- 1977 (USNM); 6 pupal exuviae, Potomac River at Scott Run, Fairfax County, Virginia, USA, 7 - VI- 1955 (USNM).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD74302DFD721FD94941E471.taxon	description	(Figs. 21 K, 27 F, 30 S, 39 E, 45 R – S, 52 D, 67 C, 76 J)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD74302DFD721FD94941E471.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the prothoracic extension wide medially and not extending to the antenna (Fig. 27 F), the dorsal apotome with only a ventral tubercle (Fig. 21 K), abdominal segment 4 with the L- 1 - IV anterior to remaining lateral sensilla but with all these situated at the segment's midlength (Fig. 67 C) and abdominal segment 8 with the two ventral sensilla (V- 5 - VIII and V- 6 - VIII) on separate tubercles.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD74302DFD721FD94941E471.taxon	description	DESCRIPTION: Total length = 4.78 – 5.19 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16 B, 33 B). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17 C, 79 H). Head: Dorsal apotome (Fig. 21 K), with ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 27 F) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 39 E) anterior to just anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 21 K) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta; clypeal-labrals (Fig. 27 F) — 2 slender setae; oculars (Fig. 27 F) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 27 F) wide, well-developed but narrow dorsolaterally, not extending to antenna; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 52 D); respiratory organ (Figs. 45 R – S) length / width = 2.95 – 3.06, moderately elongate, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single curved row, outer surface with some wrinkles, without pedicel, base with elongate posteromedial apodeme, membranous base of respiratory organ short, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing (Fig. 39 E) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 A) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 39 E) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33 I); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 2 elongate setae (as in Figs. 31 L – M); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 30 S) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T setae, D- 3 - T campaniform sensillum; D- 1 - T, D- 2 - T on single tubercle, D- 3 - T anterolateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 52 D) — 1 campaniform sensillum; M- 3 - T distant from margin of metathorax (at least 1 / 3 length of metathorax). Abdomen: with tergites 1 - 7 with medial area with stripe, 2 anterolateral spots, 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 tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 76 J) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 52 D) with 6 setae, 2 campaniform sensilla, including 2 lateral sensilla, D- 2 - I, D- 3 - I closely approximated, D- 7 - I situated anteriorly near D- 3 - I; segment 4 (Fig. 67 C) — D- 2 - IV, D- 3 - IV short to moderately elongate setae on short tubercles; D- 5 - IV, 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 elongate seta on short tubercle, just anterior of L- 2 - IV, L- 3 - IV, L- 4 - IV on anterior half of segment; L- 2 - IV, L- 3 - IV, L- 4 - IV moderately elongate setae on pointed tubercles, V- 5 - IV, V- 7 - IV short setae, V- 6 - IV elongate seta, on short rounded tubercles, all closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Fig. 76 J) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD74302DFD721FD94941E471.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Lanatomyia is known from three species in the Oriental and Australasian Regions (Borkent 2014). Immatures have been collected from sandy pool edges. Harris (1981) described pupal behaviour, with pupae keeping their abdomens in substrate until just prior to adult emergence.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD74302DFD721FD94941E471.taxon	discussion	TAXONOMIC DISCUSSION: Two species of Lanatomyia are known as pupae (Tables 2 – 3). Debenham (1974) described the lateral sensilla of abdominal segment 6 of L. miles and these are considerably more scattered in an anterior to posterior arrangement than that reported here for segment 4 (Fig. 67 C) (based on original material). Elson-Harris (1987) described the abdominal segment 4 sensilla of L. miles and appears to show the lateral and posterior sensilla as relatively more posterior on the segment than shown here (Fig. 67 C). It appears that she did not draw part of the posterior portion of the segment (as she did do for L. electra). Elson-Harris (1987) keyed the two Australian species.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD74302DFD721FD94941E471.taxon	materials_examined	MATERIAL EXAMINED: L. electra: 1 pupal exuviae, Yellow Water, Australia, 14 - I- 1956 (ANIC). L. miles: 1 pupal exuviae, Mountain Lake, Thurmere Lakes, New South Wales, Australia, 2 - II- 1966 (ANIC); 1 pupal exuviae, Nattai River, Mittagong, New South Wales, Australia, 5 - II- 1965 (ANIC).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD753028FDA21CD24CAEE38C.taxon	description	(Figs. 21 L, 27 G, 30 T, 39 F, 45 T, 52 E, 68 A, 76 K – L)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD753028FDA21CD24CAEE38C.taxon	type_taxon	Type species: Mallochohelea atripes Wirth, 1962: 281, by present designation. Included species and taxonomic discussion: given below.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD753028FDA21CD24CAEE38C.taxon	etymology	Etymology: Anebos-myia (beardless-fly) refers to the lack of sternite 8 tufts on female adults of species in this genus.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD753028FDA21CD24CAEE38C.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the dorsal apotome with two pairs of tubercles (Fig. 21 L).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD753028FDA21CD24CAEE38C.taxon	description	DESCRIPTION: Total length = 3.41 – 4.91 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16 B, 33 B). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17 C, 79 H). Head: Dorsal apotome (Fig. 21 L), with ventral line of weakness, with additional dorsal pair dorsomedial tubercles, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 27 G) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 39 F) barely posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 21 L) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 27 G) — 2 slender setae; oculars (Fig. 27 G) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 27 G) wide, well-developed but narrow dorsolaterally, not extending to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 52 E); respiratory organ (Fig. 45 T) length / width = 2.91 – 5.40, moderately elongate, 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, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing (Fig. 39 F) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 A) broadly abutting; halter apex abutting anterolateral knob-like extension of tergite 2; legs (Fig. 39 F) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33 I); with apex of foreleg moderately anterior to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 2 elongate setae (as in Figs. 31 L – M); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 30 T) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T setae, D- 3 - T campaniform sensillum, D- 3 - T anterolateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 52 E) — 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 tergite 1 with 3 medial spots, tergites 2 – 7 with medial area with stripe, 2 anterolateral spots, sternites 3 – 7 with pale to well defined 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 tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Figs. 76 K – L) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 52 E) with 8 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. 68 A) — D- 2 - IV, D- 3 - IV moderately elongate setae on short tubercles; D- 5 - IV, D- 8 - IV, D- 9 - IV 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 pointed tubercle, near L- 3 - IV; L- 2 - IV, L- 3 - IV, L- 4 - IV moderately elongate setae on pointed tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV moderately elongate setae on short tubercles, all closely approximated; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Figs. 76 K-L) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD753028FDA21CD24CAEE38C.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Anebomyia is known from five species from the Nearctic, Afrotropical and Oriental Regions (all previously in Mallochohelea; Borkent 2014). Immatures are known from stream and river margins, lakes and ponds. Knausenberger (1987) discusses the various habitats in which A. atripes was collected in the eastern USA. At least some pupae were on emergent vegetation (although they do not have abdominal membranous discs).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD753028FDA21CD24CAEE38C.taxon	discussion	TAXONOMIC DISCUSSION: Pupae are known from two species (Tables 2 – 3). All the species placed here in the new genus Anebomyia were previously in Mallochohelea. The new genus is recognized because the newly included species are not related to those now restricted to Mallochohelea, a genus most closely related to Jenkinshelea, Macropeza, Probezzia and Neobezzia (see phylogenetic analysis below, Fig. 81). There are differences between the two genera in both male and female adults and in pupae, with most of the adult features not yet interpreted cladistically (but see character 85). They are summarized as follows: Knausenberger (1987: 267 - 277) provided further differences between the two genera (as albibasis and atripes groups), including a larval feature. His information was based on Nearctic species and the character states need further testing with extraterritorial material. As indicated above, members of Anebomyia differ from true Mallochohelea by the absence of distinctive tufts of elongate setae on sternite 8 of female adults. In A. atripes, elongate setae are absent. In A. fluminea there are elongate setae on sternite 8 but these are not arranged as a tight tuft (as is true in Mallochohelea and other related genera; see character 85). Examination of female adult M. texensis indicates that it too lacks a well defined sternite 8 tuft. Wirth (1962), in erecting Mallochohelea, pointed out that some of the included species such as M. atripes, M. spinipes and M. texensis differed considerably from the genotype (M. albibasis) and " may, eventually need a separate category ". He further pointed out in a strikingly prescient statement " The discovery of the pupa of this group should also serve greatly to clarify its relationships ". The pupae have indeed provided the synapomorphies to clarify the relationship of these species. Although Wirth (1962) considered M. spinipes as closely related to A. atripes, I have not included it in Anebomyia. I have not examined material of this species and know only that the adult female has spines on its femora; the sternite 8 tufts are not described and the male remains unknown. Like this and a number of other species presently in Mallochohelea, reexamination of specimens is needed before the names are potentially transferred to the new genus. Of course, knowing the pupae of further species will certainly clarify their taxonomic position. All European species of Mallochohelea fit the characterization of this genus (Patrycja Dominiak, pers. comm.). The female of the European / Asian species M. tianshanica, however, has spines on the femora and welldeveloped sternite 8 setal tufts (Patrycja Dominiak, pers. comm.) and the male has basally fused parameres (Remm 1980). This suggests that M. tianshanica is a true Mallochohelea but is the only species known with femoral spines. Debenham's (1974) description of Mallochohelea and the included Australian species suggests that they all actually belong within Mallochohelea. The presence of femoral spines on the Afrotropical species M. kirki (de Meillon & Wirth 1981) may indicate they are members of Anebomyia but the presence or absence of the female sternite 8 setal tufts is unknown and males are undescribed, making its placement uncertain. The African species M. siricis has a female with spines on the forefemur and the male has separate parameres; although the condition of the female sternite 8 seta tufts is uncertain, I consider this sufficient evidence to place it in Anebomyia. The Chinese species M. yunnana, known only as a female, has femora spines and lacks the sternite 8 setal tufts, indicating it too is an Anebomyia.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD753028FDA21CD24CAEE38C.taxon	synonymic_list	Anebomyia atripes (Wirth), 1962: 281 (Mallochohelea). Type locality: New Brunswick, New Jersey, USA. new combination. Anebomyia fluminea (de Meillon & Wirth), 1981: 550 (Mallochohelea). Type locality: Sand River between White River and Hazyview, E. Transvaal, South Africa. new combination. Anebomyia siricis (de Meillon), 1961: 50 (Sphaeromias). Type locality: Fenerive, Madagascar. new combination. Anebomyia texensis (Wirth), 1962: 283 (Mallochohelea). Type locality: Kerrville, Texas, USA. new combination. Anebomyia yunnana (Yu & Zou), in Yu et al. 2005: 1503 (Mallochohelea). Malipo, Yunnan Province, China. new combination.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD753028FDA21CD24CAEE38C.taxon	materials_examined	MATERIAL EXAMINED: A. atripes: 1 pupal exuviae, Black Lake, North Burgess Township, Ontario, Canada, 30 - VI- 1971 (CNCI); 1 pupal exuviae, Patuxent Wildlife Research Center, Prince George County, Maryland, USA, 4 - VI- 1976 (USNM); 2 pupal exuviae, as previous locality, 24 - V- 1977 (USNM); 3 pupal exuviae, as previous locality, 13 - VI- 1977 (USNM); 2 pupal exuviae, as previous locality, 1977 (USNM); 1 pupal exuviae, Lower 3 Runs Creek, Savannah R. Plantation, Barnwell County, South Carolina, USA, 16 - II- 1979 (WLGC). A. fluminea: 1 pupal exuviae (of holotype), Sand River between White River and Hazyview, East Transvaal, South Africa, 2 - XII- 1973 (NMSA).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD703029FD5B18314C72E0F4.taxon	description	(Figs. 27 H, 31 A, 40 A – B, 46 A – B, 52 F, 68 B, 77 A)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD703029FD5B18314C72E0F4.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the abdominal tubercles all apically rounded (Fig. 68 B), abdominal segment 4 with D- 8 - IV and D- 9 - IV on separate or, at most, only basally fused tubercles (Fig. 68 B) and abdominal segment 8 with the two ventral sensilla (V- 5 - VIII, V- 6 - VIII) on a single tubercle and V- 5 - VIII tiny and V- 6 - VIII elongate.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD703029FD5B18314C72E0F4.taxon	description	DESCRIPTION: Total length = 4.88 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16 B, 33 B). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17 C, 79 H). Head: Dorsal apotome as described by de Meillon & Wirth (1981), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 27 H) with mandible well-developed, lacinia absent; palpus extending equal to or just posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 40 A – B) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (as described by de Meillon & Wirth 1981) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, campaniform sensillum not visible (dirty specimens); clypeal-labrals (Fig. 27 H) — 2 slender setae; oculars (Fig. 27 H) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 27 H) wide, well-developed but narrow dorsolaterally, not extending to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 52 F); respiratory organ (Figs. 46 A – B) length / width = 4.55 – 5.70, elongate, slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface smooth, 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. 40 A – B) with short tubercle at apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (similar to Fig. 33 J) broadly abutting; halter apex extending posteriorly to 1 / 4 length of tergite 2; legs (Fig. 40 A – B) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33 I); male with apex of foreleg moderately anterior to apex of midleg, female with apex of foreleg ventral to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 2 elongate setae (as in Figs. 31 L – M); anterolaterals — 1 moderate seta; dorsal setae (Fig. 31 A) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T setae, D- 3 - T campaniform sensillum, D- 3 - T lateral to slightly posterolateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 52 F) — 1 campaniform sensillum; M- 3 - T distant from margin of metathorax (at least 1 / 3 length of metathorax). Abdomen: with tergite 1 uncertain, tergites 2 - 7 with medial area with stripe, 2 anterolateral spots, sternites 3 – 7 with anterolateral spots hard to see, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with rounded, short tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 77 A) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 52 F) with 8 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. 68 B) — D- 2 - IV, D- 3 - IV moderately elongate setae, D- 2 - IV on short tubercle; D- 5 - IV, D- 8 - IV, D- 9 - IV moderately elongate setae; D- 5 - IV on single tubercle, D- 8 - IV, D- 9 - IV on basally fused, flattened 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 moderately elongate seta arising from flat surface, moderately anterior of posterior lateral setae; L- 2 - IV, L- 3 - IV, L- 4 - IV short setae on rounded tubercles, those of L- 2 - IV, L- 4 - IV flattened, V- 5 - IV, V- 6 - IV, V- 7 - IV short setae on rounded, flattened tubercles, all closely approximated; 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 (Fig. 77 A) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD703029FD5B18314C72E0F4.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Homohelea is known from 19 species in the Palaearctic, Oriental, and Afrotropical Regions (Borkent 2014). Pupae have been collected from a river margin, a brackish marsh, ponds, and ground pools. Ingram & Macfie (1921) noted pupae were common in " puddles of dirty water near a stand-pipe ".	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD703029FD5B18314C72E0F4.taxon	discussion	TAXONOMIC DISCUSSION: Three species of Homohelea are known as pupae (Tables 2 – 3). The dorsal apotomes of the available specimens were absent and cannot, therefore, be characterized. The dorsal apotome of H. delanoe was described (but not illustrated) by de Meillon & Wirth (1981) as having " a pair of small tubercles each bearing a slender hair ", which is the typical condition in most Ceratopogonidae. In addition to the male specimens examined, photos were made available (Burgert Muller, NMSA) of the holotype female of H. albitudinis and a non-type female specimen of H. delanoe, allowing comparisons to be made of sexually dimorphic features with the male pupae at hand (see character 44).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD703029FD5B18314C72E0F4.taxon	materials_examined	MATERIAL EXAMINED: H. delanoe: 2 pupal exuviae, Zoutpan, Transvaal, South Africa, 10 - I- 1974 (USNM).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD713016FD6419514C2BE776.taxon	description	(Figs. 28 A, 31 B, 40 C, 46 C – D, 53 A, 68 C, 77 B)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD713016FD6419514C2BE776.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the abdominal tubercles all apically pointed (Fig. 68 C), abdominal segment 4 with D- 8 - IV and D- 9 - IV on basally fused tubercles (Fig. 68 C) and abdominal segment 8 with the two ventral sensilla (V- 5 - VIII, V- 6 - VIII) on a single tubercle and V- 5 - VIII tiny and V- 6 - VIII elongate; not diagnosable as different from Sphaeromias.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD713016FD6419514C2BE776.taxon	description	DESCRIPTION: Total length = 6.50 – 6.97 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16 B, 33 B). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17 C, 79 H). Head: Dorsal apotome (as in Fig. 22 A), with ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 28 A) with mandible well-developed, lacinia absent; palpus extending equal to or just posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna (Fig. 40 C) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (as in Fig. 22 A) — uncertain; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 28 A) — 2 slender setae; oculars (Fig. 28 A) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 28 A) wide, well-developed but narrow dorsolaterally, not extending to antenna; mesonotum with very short tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 53 A); respiratory organ (Figs. 46 C – D) length / width = 3.39 – 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 elongate posteromedial apodeme, membranous base of respiratory organ moderately elongate, annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing (Fig. 40 C) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (similar to Fig. 33 J) broadly abutting; halter apex extending posteriorly to 1 / 4 length of tergite 2; legs (Fig. 40 C) with lateral margin of foreleg near midlength of wing slightly angled; hind leg visible at lateral margin of wing (as in Fig. 33 I); female with apex of foreleg ventral to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 2 elongate setae (as in Figs. 31 L – M); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 31 B) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T setae, D- 3 - T campaniform sensillum, D- 3 - T anterolateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 53 A) — 1 campaniform sensillum; M- 3 - T distant from margin of metathorax (at least 1 / 3 length of metathorax). Abdomen: with tergite 1 with 1 medial spot, tergites 2 – 7 with medial area with stripe, 2 anterolateral spots, 2 pairs on tergite 8, sternites 3 – 7 with medial stripe, anterolateral spot, 2 spots on sternite 8, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 77 B) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 53 A) with 8 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. 68 C) — D- 2 - IV, D- 3 - IV short to moderately elongate setae on pointed tubercles; D- 5 - IV, D- 8 - IV, D- 9 - IV short to moderately elongate setae; D- 5 - IV on single 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 near D- 3 - IV; L- 1 - IV elongate seta from surface, moderately anterior of posterior lateral setae; L- 2 - IV, L- 3 - IV, L- 4 - IV short on pointed tubercles, V- 5 - IV, V- 6 - IV, V- 7 - IV short setae on short tubercles, all closely approximated, V- 5 - IV, V- 6 - IV with tubercles 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 (Fig. 77 B) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD713016FD6419514C2BE776.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Leehelea is known from nine species in the Oriental and Australasian Regions (Borkent 2014). Pupae have been found in sand from a river margin and from a lily pond.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD713016FD6419514C2BE776.taxon	discussion	TAXONOMIC DISCUSSION: The pupae of two species of Leehelea are known (Tables 2 – 3). Male and female pupae of Homohelea and Sphaeromias, two genera closely related to Leehelea, are sexually dimorphic in the arrangement of their forelegs. In males, the foreleg is relatively short (Figs. 40 A, D) while in females the foreleg overlaps the midleg entirely (Figs. 40 B, E). Only female pupae of Leehelea are known (Fig. 40 C) and it is likely that the males, once discovered, will share this dimorphism (see character 44). The dorsal apotome of one specimen was present but at too great an angle to illustrate. Seta DA-A- 1 could not be seen but was likely broken off (considering that all Ceratopogonidae other than some Forcipomyia have a seta present). Overall, the shape of the dorsal apotome appeared similar to that of Sphaeromias longipennis. Debenham (1974) noted that the pupae of Leehelea were similar to those of Sphaeromias and neither could I find any differences between the two of any significance. Considering these strong similarities, the validity of recognizing the two genera should be reconsidered. Regarding the differences given by Debenham (1974), Sphaeromias may be paraphyletic in relation to Leehelea. They are separated primarily on the basis of a single difference in the shape of the female claw (presence or absence of an inner tooth and their length) and degree of fusion of the male parameres. Further differences seem to me to be minor (size of leg setae and their bases) or invalid (the brush on the ventral surface of Ta 4 is present in at least S. longipennis). Nandi et al. (2012) lists other differences but at least most apply to both genera. Elson-Harris (1987) keyed the pupae of the two Australian species, L. hispida and L. wasselli.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD713016FD6419514C2BE776.taxon	materials_examined	MATERIAL EXAMINED: L. hispida: 1 pupal exuviae, Nepean River, Menangle, New South Wales, Australia, 9 - XII- 1968 (ANIC). L. wasselli: 1 pupal exuviae, Roper River Mission, Northern Territory, Australia, 8 - XI- 1956 (ANIC).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4E3014FD671FD049E9E2FC.taxon	description	(Figs. 2 F, 12 E, 17 C, 22 A, 28 B, 31 C, 33 I, 40 D – E, 46 E, 53 B, 69 A, 77 C)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4E3014FD671FD049E9E2FC.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with the abdominal tubercles all apically pointed (Fig. 68 C), abdominal segment 4 with D- 8 - IV and D- 8 - IV on only basally fused tubercles (Fig. 68 C) and abdominal segment 8 with the two ventral sensilla (V- 5 - VIII, V- 6 - VIII) on a single tubercle and V- 5 - VIII tiny and V- 6 - VIII elongate; not diagnosable as different from Leehelea, a genus known only from the Oriental and Australasian Regions.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4E3014FD671FD049E9E2FC.taxon	description	DESCRIPTION: Habitus as in Fig. 12 E. Total length = 6.63 – 8.53 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (Figs. 17 C, 33 I). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17 C, 79 H). Head: Dorsal apotome (Fig. 22 A), with ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 28 B) with mandible well-developed, lacinia absent; palpus extending just posterior to well posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna (Figs. 40 D – E) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 22 A) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 28 B) — 2 slender setae; oculars (Fig. 28 B) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 28 B) wide, well-developed but narrow dorsolaterally, not extending to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 53 B); respiratory organ (Fig. 46 E) length / width = 3.20 – 4.06, elongate, moderately slender, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single row, outer surface with a few wrinkles, with moderately elongate pedicel, base with moderate elongate posteromedial apodeme, membranous base of respiratory organ short to moderately elongate and annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles for most of length; wing (Figs. 40 D – E) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33 I) broadly abutting; halter apex extending posteriorly to 1 / 6 length of tergite 2; legs (Figs. 40 D – E) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33 I); male with apex of foreleg moderately anterior to apex of midleg, female with apex of foreleg ventral to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 2 elongate setae (as in Figs. 31 L – M); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 31 C) — 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. 53 B) — 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, 2 spots on sternite 8; sternite 8 with dark posteromedial apodeme, segment 2 as wide or slightly wider than segment 3, segments with undivided, thin to thick setae, with pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 77 C) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 53 B) with 8 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. 69 A) — D- 2 - IV, D- 3 - IV short to moderately elongate 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 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; L- 1 - IV elongate seta on short tubercle, just anterior of base of tubercle with L- 2 - IV, L- 3 - IV; L- 2 - IV, L- 3 - IV, L- 4 - IV moderately elongate setae, L- 2 - IV, L- 3 - IV on single pointed tubercle, L- 4 - IV on elongate tubercle, V- 5 - IV, V- 6 - IV, V- 7 - IV short setae on short tubercles, all closely approximated, V- 5 - IV, V- 6 - IV with tubercles 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 (Fig. 77 C) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4E3014FD671FD049E9E2FC.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Sphaeromias is known from 30 species from every Region worldwide other than the Neotropical Region (Borkent 2014; minus one species, see below). Immatures are known from water hyacinth, Pistia stratiotes, from peat soil in Rhizophora mangroves, shady swamps, a grassy wetland, rivers, and lakes (sometimes in blanket algae). Larvae of Holarctic species are reported from the benthos of lakes. Knausenberger (1987) provides further details of various microhabitats within the genus.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4E3014FD671FD049E9E2FC.taxon	discussion	TAXONOMIC DISCUSSION: The pupae of five species of Sphaeromias are known (Tables 2 – 3). Sphaeromias theileri de Meillon & Wirth (1981) is now considered a species of Nilobezzia (see taxonomic discussion under that genus). Male and female pupae of Sphaeromias and Homohelea are sexually dimorphic in the arrangement of their forelegs. In males, the foreleg is relatively short (Figs. 40 A, D) while in females the foreleg overlaps the midleg entirely (Figs. 40 B, E) (see character 44). The feature is likely present in two related genera, Xenohelea and Leehelea. Some pupae of Sphaeromias are the largest of all Ceratopogonidae pupae (Fig. 2 F).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4E3014FD671FD049E9E2FC.taxon	materials_examined	MATERIAL EXAMINED: S. bifidus: 1 pupal exuviae, Black Lake, North Burgess Township, Ontario, Canada, 21 - VI- 1971 (CNCI). S. fasciatus: 2 pupae, Tegeler See, Germany, 20 - V- 1931 (ZSMC); 7 pupal exuviae, Strelna, Leningrad Province, Russia, 30 - V- 1998 (ZIN). S. longipennis: 1 pupal exuviae, Black Lake, Stanleyville, Ontario, Canada, 24 - VI- 1975 (USNM); 2 pupal exuviae, as previous locality, 25 - VI- 1975 (USNM); 5 pupal exuviae, Rideau River, Ottawa, Ontario, Canada, 29 - V- 1960 (USNM); 1 pupal exuviae (in glycerin), Black Lake, Quebec, Canada, 21 - VI- 1971 (CNCI); 1 pupal exuviae, Bemus Point, Chautauqua Lake, New York, USA, 31 - V- 1963 (USNM); 1 pupal exuviae, Wanakena, St. Lawrence County, New York, USA, 25 - VI- 1963 (USNM); 1 pupal exuviae, Cranberry Lake, St. Lawrence County, New York, USA, 25 - VI- 1963 (USNM). S. pictus: 4 pupal exuviae, Raigorodok, Donetsk Province, Ukraine, 30 - V- 1970 (ZIN). S. sp.: 2 pupal exuviae, White Lake, Ontario, Canada, 29 - V- 1967 (CNCI); 1 pupal exuviae, Rutka Tartak nr Suwałki, Poland, 2 - VII- 1993 (IZUG).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4C3015FD511B5F4C46E039.taxon	description	(Figs. 28 C, 31 D, 40 F, 46 F, 53 C, 69 B, 77 D)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4C3015FD511B5F4C46E039.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with abdominal tergite sensilla D- 8 - IV and D- 9 - IV on the apex of a single lobed, flattened tubercle (Fig. 69 B).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4C3015FD511B5F4C46E039.taxon	description	DESCRIPTION: Total length = 5.59 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (as in Figs. 16 B, 33 B). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17 C, 79 H). Head: Dorsal apotome missing; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 28 C) with mandible well-developed, lacinia absent; palpus extending just posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna (Fig. 40 F) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals — uncertain; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeallabrals (Fig. 28 C) — 2 slender setae; oculars (Fig. 28 C) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 28 C) wide, well-developed but narrow dorsolaterally, not extending to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 53 C); respiratory organ (Fig. 46 F) length / width = 4.83, 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, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length; wing (Fig. 40 F) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (similar to Fig. 33 J) broadly abutting; halter apex extending posteriorly to 1 / 4 length of tergite 2; legs (Fig. 40 F) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33 I); female with apex of foreleg ventral to apex of midleg; apex of hind leg abutting apex of midleg laterally; sensilla: anteromedials — 2 elongate setae (as in Figs. 31 L – M); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 31 D) — D- 1 - T, D- 2 - T, D- 4 - T, D- 5 - T setae, D- 3 - T campaniform sensillum, D- 4 - T uncertain, broken along length, D- 3 - T lateral to D- 4 - T; supraalar 2 — campaniform sensillum; metathoracics (Fig. 53 C) — 1 campaniform sensillum; M- 3 - T distant from margin of metathorax (at least 1 / 3 length of metathorax). Abdomen: with 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, thin to thick setae, with rounded to pointed, short to moderately elongate tubercles, tergites or sternites entire, each without membranous disc; segment 9 (Fig. 77 D) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 53 C) with 7 setae, 3 campaniform sensilla (1 broken seta?), including 3 lateral sensilla, D- 2 - I, D- 3 - I closely approximated, D- 7 - I situated anteriorly near D- 3 - I; segment 4 (Fig. 69 B) — D- 2 - IV, D- 3 - IV moderately elongate setae, D- 2 - IV on short tubercle; D- 5 - IV, D- 8 - IV, D- 9 - IV short to moderately elongate setae; D- 5 - IV on single tubercle, D- 8 - IV, D- 9 - IV on single flattened tubercle, 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 short tubercle, just anterior of base of tubercle with L- 2 - IV, L- 3 - IV; L- 2 - IV, L- 3 - IV, L- 4 - IV short setae on rounded, flattened tubercles, L- 2 - IV, L- 3 - IV on single tubercle, V- 5 - IV, V- 6 - IV, V- 7 - IV short setae on short tubercles, all closely approximated; 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 (Fig. 77 D) — with D- 5 - IX campaniform sensillum, D- 6 - IX possibly absent (not discernable).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4C3015FD511B5F4C46E039.taxon	distribution	DISTRIBUTION AND HABITAT: The genus Xenohelea is known from 14 species in the Afrotropical and Oriental Regions (Borkent 2014). The only known pupa was collected in mud from " aus einem klaren Gewässer " (a clear body of water) (Mayer 1957).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4C3015FD511B5F4C46E039.taxon	discussion	TAXONOMIC DISCUSSION: Only one species of Xenohelea is known as a pupa (Tables 2 – 3). Male and female pupae of Homohelea and Sphaeromias, two genera closely related to Xenohelea, are sexually dimorphic in the arrangement of their forelegs. In males, the foreleg is relatively short (Figs. 40 A, D) while in females the foreleg overlaps the midleg entirely (Figs. 40 B, E). Only one female Xenohelea is known (Fig. 40 F) and it is likely that the male, once discovered, will share this dimorphism (see character 44). Mayer (1957) described the same specimen as I studied here, the only one known for the genus. He shows D- 8 - IV and D- 9 - IV on separate, abutting tubercles but they are clearly located together on a single rounded tubercle. In addition, he showed V- 5 - IV and V- 6 - IV on a single tubercle, which was indeed present on one side but slightly separated on the other (that drawn here). The dorsal apotome of the single available specimen was missing and cannot, therefore, be characterized. The female holotype was remounted in Canada Balsam from drying and damaged Hoyer's, perhaps accounting for some slight differences between the description here and that by Mayer (1957) (e. g. body length).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4C3015FD511B5F4C46E039.taxon	materials_examined	MATERIAL EXAMINED: X. galatea: 1 pupal exuviae (of holotype), Nametil Quelimane, Mozambique, 7 - I- 1940 (SAIM).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4D3015FD9B18C24FB0E49C.taxon	description	(Figs. 22 B, 46 G)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4D3015FD9B18C24FB0E49C.taxon	diagnosis	DIAGNOSIS: known only from one limited description by Lane et al. (1955) and not diagnosable.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4D3015FD9B18C24FB0E49C.taxon	description	DESCRIPTION: Known only from one limited description by Lane et al. (1955), with the following character states discernable: Head: Dorsal apotome (Fig. 22 B), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome. Thorax: Respiratory organ (Fig. 46 G) 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.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4D3015FD9B18C24FB0E49C.taxon	distribution	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).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4D3015FD9B18C24FB0E49C.taxon	discussion	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. 22 B) and it is almost certainly poorly drawn. The respiratory organ was drawn in an anterior / posterior perspective (Fig. 46 G) 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.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4D3015FD9B18C24FB0E49C.taxon	materials_examined	MATERIAL EXAMINED: None available.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4D3011FD3A1CF94FECE426.taxon	description	(Figs. 12 F, 13 I, 17 D, 22 C – D, 28 D, 31 E, 33 J, 41 A, 46 H – K, 53 D, 69 C, 77 E – H)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4D3011FD3A1CF94FECE426.taxon	diagnosis	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. 53 D), apex of the halter extending posteriorly to about 1 / 6 length of tergite 2 (as in Fig. 33 L), abdominal segment 4 with V- 5 - IV, V- 6 - IV and V- 7 - IV closely approximated (as in Fig. 70 C) 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. 69 C) (not as in Fig. 70 B), 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. 22 C – D), 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. 22 J).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4D3011FD3A1CF94FECE426.taxon	description	DESCRIPTION: Habitus as in Fig. 12 F. Total length = 2.00 – 6.16 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (Figs. 17 D, 33 J). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17 C, 79 H) or with eye appressed to antenna (Figs. 17 D). Head: Dorsal apotome (Figs. 22 C – D), with partial ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (Fig. 13 I) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 28 D) 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. 41 A) anterior to posterior to, posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Figs. 22 C – D) — 1 moderate to elongate seta, 1 – 3 campaniform sensilla; dorsolateral cephalic sclerite sensilla — 1 seta, 0 – 1 campaniform sensillum; clypeal-labrals (Fig. 28 D) — 1 slender or 2 slender or thick setae; oculars (Fig. 28 D) — 1 – 2 seta, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 28 D) wide, well-developed, extending from palpus to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 53 D); respiratory organ (Figs. 46 H – K) 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. 41 A) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33 J) broadly abutting; halter apex extending posteriorly to 1 / 6 length of tergite 2; legs (Fig. 41 A) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33 J); 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. 31 L – M); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 31 E) — 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. 53 D) — 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. 77 E – H) 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. 53 D) 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. 69 C) — 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. 77 E – H) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4D3011FD3A1CF94FECE426.taxon	distribution	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.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4D3011FD3A1CF94FECE426.taxon	discussion	TAXONOMIC DISCUSSION: The pupae of 51 species of Bezzia are known (Tables 2 – 3). 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. 6 B) 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. 46 H, 46 J) 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) 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 (1934 a) 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 (1983 a, 1983 b, 1983 c) 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).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD4D3011FD3A1CF94FECE426.taxon	materials_examined	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 - VII 1918 (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 ° 29.12 N 119 ° 28.07 W, British Columbia, Canada, 27 – 28 - V- 2008 (CNCI); 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).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD49301EFDB21C604D8CE40C.taxon	description	(Figs. 22 F, 28 E, 31 F, 41 B, 46 L, 53 E, 70 A, 77 I – J)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD49301EFDB21C604D8CE40C.taxon	diagnosis	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. 53 E), apex of the halter extending posteriorly to about 1 / 6 length of tergite 2 (as in Fig. 33 L), D- 7 - I near D- 3 - I (Fig. 53 E), abdominal segment 4 with each setae on a rounded tubercle (Fig. 70 A) and abdominal segment 8 without L- 1 - VIII.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD49301EFDB21C604D8CE40C.taxon	description	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. 16 B, 33 B). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17 C, 79 H). Head: Dorsal apotome (Fig. 22 F), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 28 E) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna (Fig. 41 B) anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 22 F) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum on very short tubercle; clypeal-labrals (Fig. 28 E) — 2 slender setae; oculars (Fig. 28 E) — 2 setae, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 28 E) wide, well-developed but very narrow dorsolaterally, extending from palpus to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 53 E); respiratory organ (Fig. 46 L) 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. 41 B) with short tubercle at apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 J) broadly abutting; halter apex extending posteriorly to 1 / 6 length of tergite 2; legs (Fig. 41 B) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33 J); 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. 31 F) — 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. 53 E) — 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 I-J) not strongly modified, terminal processes separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 53 E) 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. 70 A) — 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. 77 I – J) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD49301EFDB21C604D8CE40C.taxon	distribution	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.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD49301EFDB21C604D8CE40C.taxon	discussion	TAXONOMIC DISCUSSION: Only one species is known as a pupa (Tables 2 – 3).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD49301EFDB21C604D8CE40C.taxon	materials_examined	MATERIAL EXAMINED: C. dycei: 2 pupal exuviae (of paratypes), Tacuarembo, Uruguay, 29 - IX- 1980 (MLPA).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD46301FFD521C494ED8E46E.taxon	description	(Figs. 22 G, 28 F, 31 G, 41 C, 46 M, 53 F, 70 B, 77 K – L)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD46301FFD521C494ED8E46E.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with dorsal apotome with only one campaniform sensillum (Fig. 22 G), metathorax with only one campaniform sensillum (M- 3 - T) situated at least ⅓ the length of the metathorax from its anterior margin (Fig. 53 F), apex of the halter extending posteriorly to about 1 / 6 length of tergite 2 (as in Fig. 33 L), 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. 70 B).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD46301FFD521C494ED8E46E.taxon	description	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. 16 B, 33 B). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17 C, 79 H). Head: Dorsal apotome (Fig. 22 G), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 28 F) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 41 C) barely anterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 22 G) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 28 F) — 2 slender setae; oculars (Fig. 28 F) — 1 seta, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 28 F) wide, well-developed but very narrow dorsolaterally, extending from palpus to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 53 F); respiratory organ (Fig. 46 M) 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. 41 C) with short tubercle at apex of hind leg, separated medially by fore-, midlegs; halter apex and hind leg (as in Fig. 33 J) broadly abutting; halter apex extending posteriorly to 1 / 6 length of tergite 2; legs (Fig. 41 C) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33 J); 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. 31 L – M); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 31 G) — 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. 53 F) — 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. 77 K – L) not strongly modified, terminal processes separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 53 F) 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. 70 B) — 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 K-L) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD46301FFD521C494ED8E46E.taxon	distribution	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).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD46301FFD521C494ED8E46E.taxon	discussion	TAXONOMIC DISCUSSION: One species of Pachyhelea is known as a pupa (Tables 2 – 3).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD46301FFD521C494ED8E46E.taxon	materials_examined	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).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD47301BFD5F1CA94C9AE13C.taxon	description	(Figs. 2 D 12 G, 17 E, 22 H – J, 28 G, 31 H, 31 L – M, 33 K, 41 D, 46 N – R, 54 A, 70 C, 71 A, 78 A – H)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD47301BFD5F1CA94C9AE13C.taxon	diagnosis	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. 54 A), apex of the halter extending posteriorly to about 1 / 6 length of tergite 2 (as in Fig. 33 L), abdominal segment 4 with V- 5 - IV, V- 6 - IV and V- 7 - IV closely approximated (Fig. 70 C) 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. 69 C) (not as in Fig. 70 B), 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. 22 C – D), a nearly unique condition found otherwise only in P. flavipes and P. jonesi (the latter distinctively with two setae (Fig. 22 J).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD47301BFD5F1CA94C9AE13C.taxon	description	DESCRIPTION: Habitus as in Fig. 12 G. Total length = 2.25 – 7.03 mm. Without larval exuviae retained on abdomen. Exuviae with flagellum appressed against lateral margin of midleg, wing (Figs. 17 E, 33 K). Ecdysial tear around base of antenna, along lateral margin of face to palpus (Figs. 17 E, 79 H). Head: Dorsal apotome (Figs. 22 H – J), with partial ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 28 G) 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. 41 D) well anterior to posterior to, posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Figs. 22 H – J) — 1 elongate seta, 1 campaniform sensillum or 2 elongate setae, 2 campaniform sensilla; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeallabrals (Fig. 28 G) — 2 slender setae; oculars (Fig. 28 G) — 1 – 2 setae, 1 campaniform sensillum or 1 seta, 2 campaniform sensilla. Thorax: Prothoracic extension (Fig. 28 G) wide, well-developed but in some narrow dorsolaterally, extending from palpus to antenna; mesonotum without tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 54 A); respiratory organ (Figs. 46 N – R) length / width = 2.46 – 4.73, moderately elongate to elongate, somewhat flattened apically, with pores closely abutting at apex of respiratory organ, arranged in single straight to curved row, outer surface smooth or with some wrinkles, with short, wide pedicel, base with short posteromedial apodeme, membranous base of respiratory organ short, annulated, tracheal tube straight to slightly curved along length, with spirals restricted to base, wrinkles to half length or more; wing (Fig. 41 D) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33 K) broadly abutting; halter apex extending posteriorly to 1 / 6 length of tergite 2; legs (Fig. 41 D) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33 K); 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. 31 L – M); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 31 H) — 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. 54 A) — 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. 78 A – H) not strongly modified, terminal processes closely approximated to separated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 54 A) 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. 70 C, 71 A) — D- 2 - IV, D- 3 - IV short to moderately elongate setae on short to pointed tubercles; D- 5 - IV peg-like or slender seta, D- 8 - IV, D- 9 - IV short to moderately elongate setae, D- 7 - IV present or absent; D- 5 - IV without or on short tubercle, D- 8 - IV, D- 9 - IV on basally fused or separate but closely approximated tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 8 - IV, D- 9 - IV; D- 7 - IV, if present, near D- 3 - IV; L- 1 - IV short to elongate seta on rounded tubercle, just anterior of base of tubercle with L- 2 - IV, L- 3 - IV; L- 2 - IV, L- 3 - IV, L- 4 - IV elongate setae on pointed tubercles, L- 2 - IV, L- 3 - IV on single tubercle in some, V- 5 - IV, V- 6 - IV, V- 7 - IV short to moderately elongate setae on short tubercles, all closely approximated or with V- 7 - IV closer to L- 4 - IV; segment 8 without D- 3 - VIII, without L- 1 - VIII; segment 9 (Figs. 78 A-H) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD47301BFD5F1CA94C9AE13C.taxon	distribution	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.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD47301BFD5F1CA94C9AE13C.taxon	discussion	TAXONOMIC DISCUSSION: There are 41 species of Palpomyia known as pupae (Tables 2 – 3). Thienemann (1928), Mayer (1934 a) and Lenz (1934) provided similar keys to a few European species groups known at that time. Grogan & Wirth (1979), in their masterful revision of Nearctic Palpomyia, provide a key to the pupae of 16 species known from the Nearctic. However, it is unfortunate that Palpomyia pupae cannot be diagnosed at the generic level so that we cannot yet identify a Palpomyia pupa as such. The key therefore requires foreknowledge that the specimen is a member of the genus (e. g. reared to adult). Liu & Yu (1991) described the monotypic genus Nemoromyia Liu & Yu based on their newly described N. nemorosa Liu & Yu. They stated that the genus was a member of the Heteromyiini. Borkent (1998) indicated that the species was actually a species of Palpomyia based on various described features but that the type should be reexamined before nomenclatural changes are made. Yu et al. (2005) continued to place the genus and species in the Heteromyiini, although the evidence for this was unclear (it is not noted in the English abstract, but I have not translated the longer Chinese text). I have reexamined the female holotype and associated pupal exuviae and it is clearly a member of the Palpomyia distincta species group and, within this group, is very similar to P. rufa Loew (Grogan & Wirth 1979). The female has the characteristic synapomorphy of posteromedially directed lobes on sternite 8 of the P. distincta group. Furthermore, tergite 8 is very short (much shorter than tergite 7), a feature which is a synapomorphy of the Palpomyiini (the feature is unique in the Ceratopogonidae), further supporting its position as a species of Palpomyia. The pupa of N. nemorosa is very similar to a number of other species of Palpomyia (as indicated in the key). Liu & Yu (1991) noted that the adult female lacked the abdominal tergal apodemes which is a synapomorphy of the Palpomyiini. However, as shown by Borkent & Craig (1994), newly emerged female adults of this group do not show the apodemes which sclerotize and darken after emergence. It is not surprising, therefore, that this reared specimen was reported as lacking the apodemes. I therefore recognize Nemoromyia as a new synonym of Palpomyia and the species as a new combination as follows: Palpomyia nemorosa (Liu & Yu), 1991: 26 (Nemoromyia). Raohe, Heilongjiang Province, China. new combination. The pupa of P. lineata has been described more than once by the same authors (Table 2) but with different names (now synonyms), suggesting either misidentifications or the possibility of more than one species actually being present in Europe. Ronderos et al. (2004) gave a detailed description of the pupa of P. guarani. Their figure of the cephalothorax (their fig. 14) shows the apex of the halter extending only barely past the anterior margin of tergite 2. This would be unique within the Palpomyia + Bezzia + Phaenobezzia + Clastrieromyia + Stenoxenini clade, where the halter extends to about 1 / 6 the length of tergite 2. Paul et al. (2014) recently thoroughly described the pupa of a species from India, the first known from the Oriental Region, but misidentified some of the sensilla (e. g. SA- 2 - T, L- 3 - IV, L- 4 - IV).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD47301BFD5F1CA94C9AE13C.taxon	materials_examined	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).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD433019FD8E1999487AE059.taxon	description	(Figs. 17 F, 22 K, 28 H, 31 I, 33 L, 41 E, 46 S, 54 B, 71 B, 78 I – K)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD433019FD8E1999487AE059.taxon	diagnosis	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. 78 I – J); female (without genital lobes) with the metathorax with only one campaniform sensillum (M- 3 - T) situated at least ⅓ the length of the metathorax from its anterior margin (Fig. 54 B), apex of the halter extending posteriorly to about 1 / 6 length of tergite 2 (Fig. 33 L), abdominal segment 4 with V- 5 - IV, V- 6 - IV and V- 7 - IV closely approximated (Fig. 71 B), 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. 22 C – D), a nearly unique condition found otherwise only in P. flavipes and P. jonesi (Fig. 22 J).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD433019FD8E1999487AE059.taxon	description	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. 17 F, 33 L). Ecdysial tear around base of antenna, along lateral margin of face to palpus (Figs. 17 F, 79 H). Head: Dorsal apotome (Fig. 22 K), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 28 H) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium separated medially by labrum, hypopharynx; apex of antenna (Fig. 41 E) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), narrowed posteriorly; sensilla: dorsal apotomals (Fig. 22 K) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeallabrals (Fig. 28 H) — 2 slender setae; oculars (Fig. 28 H) — 1 seta, 1 campaniform sensillum. Thorax: Prothoracic extension (Fig. 28 H) wide, well-developed, extending from palpus to antenna; mesonotum with short tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 54 B); respiratory organ (Fig. 46 S) 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. 41 E) without apical tubercle or angle, separated medially by fore-, midlegs; halter apex and hind leg (Fig. 33 L) broadly abutting; halter apex extending posteriorly to 1 / 6 length of tergite 2; legs (Fig. 41 E) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (Fig. 33 L); 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. 31 L – M); anterolaterals — 1 moderately long seta; dorsal setae (Fig. 31 I) — 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. 54 B) — 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. 78 I – K) not strongly modified, terminal processes closely approximated to separated basally, each projecting posterodorsolaterally, tapering to pointed apex; apex of male genital lobe anterior or to level of base of terminal process (Figs. 78 I – J); sensilla: tergite 1 (Fig. 54 B) 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. 71 B) — D- 2 - IV, D- 3 - IV moderately elongate setae on moderately developed tubercles; D- 5 - IV, D- 8 - IV short setae, D- 9 - IV elongate seta, D- 7 - IV present or absent; D- 5 - IV on short tubercle, D- 8 - IV, D- 9 - IV on basally fused, closely approximated tubercles, posterior dorsal sensilla in transverse row, arranged medially to laterally: D- 5 - IV, D- 4 - IV, D- 8 - IV, D- 9 - IV; D- 7 - IV, if present, near D- 3 - IV or L- 2 - IV; L- 1 - IV elongate seta on rounded tubercle, just anterior of base of tubercle with L- 2 - IV, L- 3 - IV; L- 2 - IV, L- 3 - IV, L- 4 - IV elongate setae on pointed tubercles, L- 2 - IV, L- 3 - IV on single tubercle, V- 5 - IV, V- 7 - IV moderately elongate setae, V- 6 - IV elongate seta, on short tubercles, all closely approximated, with tubercles of V- 5 - IV, V- 6 - IV fused basally; segment 8 without D- 3 - VIII, without L- 1 - VIII; with V- 5 - VIII, V- 6 - VIII on single tubercle, V- 5 - VIII tiny, V- 6 - VIII elongate. Segment 9 (Figs. 78 I-K) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD433019FD8E1999487AE059.taxon	distribution	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.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD433019FD8E1999487AE059.taxon	discussion	TAXONOMIC DISCUSSION: The pupae of nine species of Phaenobezzia are known (Tables 2 – 3). Haeselbarth (1965) used the shape of the terminal processes to distinguish the pupae of the four African species he studied and indicated some further character states which may separate some species. Thomsen (1937) drew the dorsal apotome of P. opaca with two setae (and no campaniform sensillum); she likely mistook the campaniform sensillum as the broken base of a seta. Wirth & Grogan (1982) drew abdominal segment 4 of P. opaca with V- 7 - IV situated laterally, close to L- 4 - IV and this is significantly different than that observed here for this species and all other Phaenobezzia (Fig. 71 B).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD433019FD8E1999487AE059.taxon	materials_examined	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).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD413006FD7918E34803E184.taxon	description	(Figs. 12 H, 22 L, 28 I, 31 J, 41 F, 46 T, 54 C, 71 C, 78 L)	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD413006FD7918E34803E184.taxon	diagnosis	DIAGNOSIS: Only pupa of Ceratopogonidae with abdominal segment 8 with D- 3 - VIII (Fig. 12 H); also, the female is the only Ceratopogonidae with wings abutting ventromedially (Figs. 12 H, 41 F).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD413006FD7918E34803E184.taxon	description	DESCRIPTION: Habitus (female) as in Fig. 12 H. 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. 16 B, 33 B). Ecdysial tear around base of antenna, along lateral margin of face to palpus (as in Figs. 17 C, 79 H). Head: Dorsal apotome (Fig. 22 L), uncertain ventral line of weakness, without dorsomedial tubercle, without central dome; dorsolateral cephalic sclerite (as in Fig. 13 H) fused to scutum, each side separated medially by dorsal apotome in whole pupa; mouthparts (Fig. 28 I) with mandible well-developed, lacinia absent; palpus extending posterior to posterolateral margin of labium; labium entire (not divided medially); apex of antenna (Fig. 41 F) posterior to posterior extent of midlength portion of midleg (portion lateral to mesosternum), not narrowed sharply posteriorly; sensilla: dorsal apotomals (Fig. 22 L) — 1 elongate seta, 1 campaniform sensillum; dorsolateral cephalic sclerite sensilla — 1 seta, 1 campaniform sensillum; clypeal-labrals (Fig. 28 I) — 2 thick setae; oculars (Fig. 28 I) — 1 seta, 1 campaniform sensillum? or possibly merely an indentation? Thorax: Prothoracic extension (Fig. 28 I) wide, well-developed but narrow dorsolaterally, extending from palpus to antenna; mesonotum yes tubercles, not extending posteromedially, not dividing metathorax medially (Fig. 54 C); respiratory organ (Fig. 46 T) 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. 41 F) 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. 33 J) broadly abutting; halter apex extending posteriorly to 1 / 6 length of tergite 2; legs (Fig. 41 F) with lateral margin of foreleg near midlength of wing evenly curved; hind leg visible at lateral margin of wing (as in Fig. 33 K); male with apex of foreleg moderately anterior to apex of midleg (as in Fig. 41 E), 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. 31 L – M); anterolaterals — 1 elongate seta; dorsal setae (Fig. 31 J) — 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. 54 C) — 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. 78 L) not strongly modified, terminal processes closely approximated basally, each projecting posterodorsolaterally, tapering to pointed apex; sensilla: tergite 1 (Fig. 54 C) 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. 71 C) — 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. 78 L) — with D- 5 - IX, D- 6 - IX campaniform sensilla.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD413006FD7918E34803E184.taxon	distribution	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.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD413006FD7918E34803E184.taxon	discussion	TAXONOMIC DISCUSSION: The pupae of four species of Paryphoconus are known (Tables 2 – 3). A female pupa of a Mallochohelea in the ZSMC was labeled " Paryphoconus lanei " but on a modern label and with recent handwriting. The specimen was among material studied by Mayer but does not correspond to his description of Paryphoconus flavidus (as lanei) (Mayer 1959 b). The Paryphoconus pupae studied by Mayer (1959 b) were not located.	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
027587C9BD413006FD7918E34803E184.taxon	materials_examined	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).	en	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): 1-327, DOI: http://dx.doi.org/10.11646/zootaxa.3879.1.1
