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Dasyhelea mesophylla Dominiak and Borkent , sp. nov. Diagnosis The only Dasyhelea in the Neotropical Region with the following characteristics. Male adult: with a single elongate radial cell, sternite 9 straight, paramere sinusoidal, twisted and tapering to a narrow apex, fused basally with gonocoxal apodemes and forming with them an asymmetrical structure, aedeagus symmetrical with dark, well-developed anterolateral club-like projections and 2 well separated and slender posterior projections with apices that are hooked dorsolaterally. Female adult: with 1 elongate radial cell, frontal sclerite broader than long, with sternite 9 elongate anteriorly, conical and with a rounded apex. Pupa: with the respiratory organ curved and coming to a sharp point, with a double row of about 7 circular pores each (so about 14 total) restricted to the apical 0.3 of the respiratory organ, and with a slightly elongate, pointed, gradually tapering terminal process bearing 2–3 pointed tubercles laterally near its base. Larva: not presently diagnosable, but see taxonomic discussion. Description Male adult. Antenna (Figure 1 (a)) dark; antennal flagellum length 0.58–0.60 mm (n = 3), antennal ratio 0.79–0.95 (n = 3). Frontal sclerite broad, with long, slender projection. Clypeus divided into two parts, with 7–11 setae distributed laterally in two rows (n = 4). Palpus (Figure 5 (b)) with third palpal segment relatively stout, 44–53 μm long (n = 4); palpal ratio of the third segment 2.54–3.17 (n = 4); sensilla capitata present on segment 3 only, rather sparse and distributed closer to its outer margin. Thorax (Figure 1 (b)), aside from slightly lighter lateral sclerites, dark. Scutellum yellowish, with 6–8 bristles and 1–5 smaller setae (n = 5). Wing with 1 radial cell; wing length 0.76–0.81 mm (n = 4), costal ratio 0.45–0.48 (n = 4). Halter dark. Legs (Figure 1 (b)) pale with indistinct, irregular darker patches at midlength of fore- and midfemora and tibiae of all legs, and apical half of hind femur. Hind tibial comb with 5–7 spine-like setae (n = 4). Tarsal ratios: foreleg 2.1–2.3 (n = 3), midleg 2.4–2.6 (n = 3), hind leg 2.2–2.5 (n = 4). Abdominal segments 3–8 well sclerotised; sclerotisation on segments 3–5 disjunct medially. Genitalia (Figures 1 (c), 5(a)). Apicolateral process of tergite 9 prominent, finger-like, with single apical seta. Cercus moderately small, with 4 setae (n = 1). Posterior margin of sternite 9 straight or with shallow excavation medially. Gonocoxite without mesoventral hook. Gonostylus nearly straight, with rounded apex, same length as or slightly longer than gonocoxite. Paramere and gonocoxal apodemes fused and forming an asymmetrical structure; paramere sinusoidal, twisted, tapering to narrow apex. Aedeagus symmetrical, with dark, well-developed anterolateral club-like projections, 2 well-separated and slender posterior projections with apices hooked dorsolaterally; anteroventral margin of aedeagus weakly sclerotised, rounded. Female adult. Antenna (Figure 2 (a)) dark; antennal flagellum length 0.52 mm, antennal ratio 0.84. Frontal sclerite broad, with long, slender projection. Clypeus divided into two parts, with about 15 setae distributed laterally in two rows. Third palpal segment relatively stout, 46 μm long; palpal ratio of the third segment 2.67; sensilla capitata as in male. Scutellum yellowish, with 8 bristles and about 8 smaller setae. Wing with 1 radial cell; wing length 0.78 mm, costal ratio 0.50. Legs pale with indistinct, irregular darker patches at midlength of fore-, hind femora, tibiae. Hind tibial comb with 6 spine-like setae. Tarsal ratios: foreleg 2.3, midleg 2.3, hind leg 2.3. Genitalia (Figures 2 (b,c), 5(c)). Abdominal segments (Figure 2 (b)) 3–8 well sclerotised; sclerotisation on segments 3–5 disjunct in the middle. Sternite 9 elongate anteriorly, conical, with rounded apex (Figures 2 (b), 5(c)). Spermatheca (Figure 2 (c)) single, retort-shaped, length 48 μm (distorted). Fourth instar larva (Figures 3 (a–f), 8(b)). Total length 1.93–3.22 mm (n = 8). Colour in life whitish, with head capsule dark brown, length 0.27–0.35 mm (n = 18), L/ W 1.61 –1.92 (n = 7), tapering to apex; chaetotaxy as in Figures 3 (a–c), unlabelled anterior setae are uncertain, j not visible in figures but present. Labrum (Figures 3 (b–c)) narrow, projecting anteriorly; anterolateral margins of head capsule extending anteriorly. Mandible (Figures 3 (a,c,d)) with 3 teeth, apical 2 teeth elongate, pointed, basal tooth broad. Epipharynx (Figure 3 (d)) massive, strongly sclerotised, lateral arms stout, short, dorsal comb with well-developed fringe. Hypopharynx (Figure 3 (d)) well developed. Caudal segment with approximately 5–7 well-developed hooks (on each side). Other details either not visible or similar to most other Dasyhelea (Díaz et al. 2018, 2019). Third instar larva (Figure 8 (a)). Similar to fourth instar. Total length 1.78–2.59 mm (n = 7). Colour in life whitish, with head capsule dark brown, length 0.20–0.23 mm (n = 8), L/ W 1.62 –1.74 (n = 4), tapering to apex. Pupa. Habitus as in Figures 4 (a), 8(c–d). Total length 1.87–2.29 mm (n = 7). Clypeus (Figure 6 (a)) with CL-1-H, CL-2-H apparently absent in most, one specimen with very tiny seta CL-2-H, otherwise the area of these sensilla represented by small section of contorted cuticle; O-2-H absent; dorsal apotome (Figure 6 (b)) smooth, with DA-1-H a tiny seta, DA-2-H a campaniform sensillum. Mesonotum without short tubercles; respiratory organ (Figure 4 (b)) length/width 5.3–6.3 (n = 5), curved, apex pointed, somewhat circular in cross section, with a double row of about 7 circular pores each (so about 14 total), restricted to about apical 0.3 of respiratory organ, closely abutting apically, separated more basally, outer surface with strong annulations on posterior (inner curve) for about basal 0.2–0.5 to about 0.1–0.8, without spicules, tracheal tube curved along length of respiratory organ, somewhat thicker basally, with spirals restricted to base; sensilla: anterolaterals – 1 seta, 2 campaniform sensilla; dorsal setae (Figure 6 (c)) – D-1-T well anterior of D-2-T, D-3-T relatively close to D-2-T. Metathoracics (Figure 6 (d)) – 2 campaniform sensilla; M-2-T, M-3-T separated (not abutting), relatively near anterior margin of metathorax. Abdomen without dark pigmentation, segments 3–8 each with some wide, shelf-like tubercles; segment 9 with terminal processes closely approximated basally, each projecting posteriorly, each with 2–3 lateral, well-developed, pointed projections, with dorsal one bearing campaniform sensillum at its base; sensilla: tergite 1 (Figure 6 (d)) with 5 setae (D-2-I, D-3-I, L-1-I, L-2-I, L-3-I) and 2 campaniform sensilla (D-4-I, D-7-I); segment 4 (Figure 7 (a)) with D-2-IV absent; D-4-IV, D-8-IV each on wide, low, slightly separate tubercle, D-7-IV on slightly pointed tubercle; posterior dorsal sensilla in transverse row, arranged medially to laterally: D-4-IV, D-7-IV each a campaniform sensillum, D-8-IV a short seta on wide, low tubercle, L-1-IV, L-2-IV, L-3-IV, L-4-IV each a short seta on apically pointed tubercle; V-5-IV absent; V-6-IV a campaniform sensillum; V-7-IV a seta on wide, low tubercle; segment 9 (Figures 4 (c), 7(b)) long, terminal process with 3–4 divisions, laterally directed tubercles curved to somewhat hook-like, posteriorly directed portion elongate, tapering to point; with only one of D-5-IX present (a campaniform sensillum), D-6-IX apparently present (difficult to discern amidst shagreen), V-1-IX, V-2-IX elongate and short seta, respectively. Distribution and bionomics Dasyhelea mesophylla is known only from the type locality on the west coast of Costa Rica at an altitude of about 5 m. Adult specimens were reared from larvae and pupae present in S. minima leaves (Figure 8) floating in a lagoon on the northern margin of Carara National Park, about 1.5 km east from highway 34 along the hiking trail. The lagoon from which this species was collected in 1993 is periodically flooded by the abutting and extremely large Tárcoles River. The larvae and pupae were common but no specific data were taken in this regard. Virtually every clump of leaves had at least one leaf being mined (or that had been previously mined). Leaves were generally in clumps of 6–12 leaves. Borkent and Craig (2001) described the pupa of Stilobezzia rabelloi Lane, which have piercing respiratory organs to obtain oxygen from the dangling ̍roots′ (actually modified leaves), from the same habitat as D. mesophylla. Observations of the mining larvae and of pupae in the S. minima leaves were made under the dissecting microscope as follows. Several fourth instar larvae were observed in some detail, although many were otherwise present. The larvae actively mined the leaves, scraping at live fern tissue, had green material in their guts and produced green excrement. The narrow labrum and anterolateral margins of head capsule extending anteriorly likely facilitate the scraping of plant tissue. The mines of at least fourth instar larvae produced a bulge on the upper surface of the leaves. The frass produced by larvae was either in the mine or on the surface of the leaf. At least some larvae periodically leave the mine for the surface and then burrow back into the leaf. A few larvae were observed to move from one leaf to another. Possibly this is a means of ensuring that overcrowding does not limit any given larva which can probably move from one abutting leaf to another. One larva was observed feeding on the interior of the leaf but with about 2/3 of its body on the surface of the leaf. Another larva was seen crawling on the surface, then gnawing through the surface to the interior and, after feeding for some time, drawing the rest of the body into the leaf. Generally, when a leaf was mined, there were only 1–2 fourth instars present (aside from other earlier instar larvae), but one instance of three larvae in a single leaf was observed. Several third instar larvae were observed mining the interior of the leaves, in an identical manner to the fourth instar larvae, but no third instars were seen on the surface of the leaves. One second instar larva was observed in a tunnel just slightly wider than its body, winding through the leaf. The mines of third and fourth instar larvae were more excavated in places, connected by swollen (leaf surface bulging at least dorsally) tunnels about the width of those larvae. First and second instar larvae seemed to be concentrated on the peripheral margins of the leaves, while third and fourth instar larvae were generally in the main body of the leaves. On the other hand, several leaves were observed to be partially mined but without larvae, suggesting the larva had died or, more likely, had moved on to another leaf to mine there. Two larvae were observed to pupate. One pupa was entirely encased within the mine with no direct opening to the surface, other than the insertion of its respiratory organs through the surface cuticle of the leaf. Another pupa had cut an opening near the anterior end of the pupa, but without the respiratory organs protruding. Observed again later, the pupa had stuck one respiratory organ up through the leaf cuticle, indicating the pupa only needs to obtain oxygen periodically from the surface (or can withdraw for some time if disturbed). Both pupae had their fourth instar exuviae present nearby (Figures 4 (a), 8(c–d)). A number of other pupae were observed with either one or both respiratory organs protruding from the leaf. Two emerging adults in the laboratory had free-floating pupal exuviae. The one pupa noted above that had been entirely encased by the surface of the leaf had made a jagged exit hole in the leaf, and the other appeared to escape through the hole already present. Some leaves were found with pupal exuviae with part of their abdomens still in the leaves. It is worth noting that Forno and Bourne (1984) previously recognised an unnamed Dasyhelea as ̍phytophagous′ on Salvinia molesta D.S. Mitch. in Brazil. Another species, called Dasyhelea sp. 3 (grisea group), was reported by Torreias et al. (2013) from southeastern Brazil, where its ̍immature stages were collected in S. auriculata ′ Aubl (6). However, considering the sampling method used, the immatures (pupae only?) were most probably found among rhizomes and submerged, root-like leaves of Salvinia. Three males and one female were obtained but no details regarding their morphology are given by the authors (Torreias et al. 2013). Pelli and Barbosa (1998) mentioned D. paulistana Forattini and Rabello as present on S. molesta in Brazil, but they did not assign this species to taxa feeding on Salvinia or to taxa causing harm to it. The latter Dasyhelea species together with D. pseudopollinosa Díaz and Ronderos were collected from mats of S. auriculata and Azolla filiculoides Lam. in Brazil and Argentina (Díaz et al. 2014). Taxonomic and phylogenetic discussion This new species belongs to the grisea species group or subgenus D. (Dasyhelea) (Dominiak 2012) if following the subgeneric division proposed by Remm (1962, 1979). The presence of the strongly modified pupal respiratory organ of D. mesophylla is shared by a number of species in the grisea species group, and because it is unique within the family (Borkent 2014), we consider this a synapomorphy of these species. As such, from the limited number of Dasyhelea species known as pupae (Borkent 2014), we consider D. mesophylla to form a monophyletic group with D. traverae Thomsen from eastern United States, D. pollinosa Wirth from the western and eastern United States, D. chani Wirth and Linley from Florida (United States), D. paulistana from Argentina and Brazil, D. pseudopollinosa from Brazil, D. caesia Remm (syn. D. lugensis Brodskaya) from Europe, and two unnamed species from Indonesia (Sumatra, listed as Holoconops sp. in Mayer 1934b) and Australia (A. Borkent, pers. obs.), respectively. All of them have virtually identical respiratory organs; these are thick at the base to about midlength and taper to a sharp apex. The respiratory organs of the pupae of D. mesophylla are somewhat more elongate than those of the other abovementioned species of Dasyhelea. Although not known as immatures, male D. unicolour Remm and D. stackelbergi Remm appear very similar to those in this group of species and may be closely related. The pupae of D. mesophylla stick their respiratory organs through the upper surface of the mined S. minima leaves to obtain oxygen from the exposed surface of the leaf, suggesting that these other species also use their respiratory organs in a similar manner. However, because at least the larvae of D. traverae do not appear to mine leaves (Thomsen 1935, 1937; Waugh and Wirth 1976) it is also possible that Dasyhelea species with such respiratory horns obtain air from water plants, similarly to S. rabelloi. The piercing respiratory organs of pupae of a group of Stilobezzia Kieffer is clearly separately evolved, as evidenced by differences in the details of the respiratory organs compared with Dasyhelea as well as the phyletic distance between the two groups (Borkent and Craig 2001). In spite of the fact that species of Dasyhelea are common throughout much of the world, only 62 out of 628 known species have been described as larvae (Díaz et al. 2013; Borkent 2014, 2016, 2018, 2019; Duan et al. 2019; Borkent and Dominiak 2020; Lu et al. 2020; Borkent et al. 2022). One reason is probably that larvae of Dasyhelea are generally morphologically conservative. However, the very narrow protruding labrum and corresponding anterolateral projections of the head capsule, likely important for scraping leaf material free to ingest, may be distinctive among those larvae which have been described so far, keeping in mind that, of the species with a piercing respiratory organ, only the larvae of D. caesia (Brodskaya 1995), D. chani (Wirth and Linley 1990) and D. traverae (Thomsen 1937) have been described. It is important to recognise that most descriptions of Dasyhelea larvae are rather superficial. Detailed comparative studies are needed of the significant differences in larvae between the numerous species of this genus. There are at least some marked differences exhibited by some taxa (eg those species living in Nepenthes pitcher plants in Southeast Asia which have extremely long, narrow head capsules; Tokunaga 1961; Wirth and Beaver 1979). None of the Dasyhelea species described from larvae or pupae other than D. mesophylla are known to be leaf-miners. Types Holotype: male adult with associated larval and pupal exuviae, on microscope slide, labelled ̍HOLOTYPE Dasyhelea mesophylla Dominiak and Borkent′, ̍ Costa Rica, 5 km NE, Tarcoles, C.R. 26-vii-1993, A. Borkent CD1490, CD 1490′, ̍Reared from larva mining Salvinia leaf′ (CNCI). Allotype: female adult with associated pupal exuviae labelled as for holotype (CNCI). Paratypes: 3 males each with associated pupal exuviae, labelled as for holotype except ̍Reared from pupa embedded in Salvinia leaf′ (1, MNCR; 2, CNCI); 1 pupa, 19 fourth instars, 6 third instars, from type locality, 3 August 1993, in Salvinia leaves, CD1502 (4 fourth instars, MNCR; remainder, CNCI); 2 pupae and their associated fourth instar exuviae in Salvinia leaves, 2 fourth instars, 1 third instar from type locality, 3 August 1993, CD1502 (CNCI). Derivation of specific epithet The name mesophylla refers to the parenchyma between the epidermal layers of a leaf, the tissue where the larva and pupa of the new species have been found., Published as part of Dominiak, Patrycja & Borkent, Art, 2023, A new species of Dasyhelea (Diptera: Ceratopogonidae), mining the leaves of the floating fern Salvinia minima Baker, pp. 665-684 in Journal of Natural History (Oxford, England) (Oxford, England) 57 (9 - 12) on pages 668-680, DOI: 10.1080/00222933.2023.2203336, http://zenodo.org/record/8070228, {"references":["Diaz F, Spinelli GR, Ronderos MM. 2018. Two new species of Dasyhelea Kieffer and the immature of D. azteca Huerta & Grogan from Northwestern Argentina (Diptera: Ceratopogonidae). Zool Anz. 272: 6 - 19. doi: 10.1016 / j. jcz. 2017.11.008.","Borkent A, Craig DA. 2001. Submerged Stilobezzia rabelloi LANE (Diptera: Ceratopogonidae) pupae obtain oxygen from the aquatic fern Salvinia minima BAKER. Proc Entomol Soc Wash. 103: 655 - 665.","Forno IW, Bourne AS. 1984. Studies in South America of arthropods on the Salvinia auriculata complex of floating ferns and their effects on S. molesta. Bull Entomol Res. 74: 609 - 621. doi: 10. 1017 / S 0007485300013985.","Torreias SRS, Ferreira-Keppler RL, Ronderos MM. 2013. Biting midges (Ceratopogonidae: Diptera) present in aquatic macrophytes from wetlands of Marchantaria Island, Iranduba, Central Amazonia, Brazil. J Nat Hist. 48 (2014): 109 - 122. doi: 10.1080 / 00222933.2013.791934.","Pelli A, Barbosa FAR. 1998. Insetos coletados em Salvinia molesta Mitchell (Salviniaceae), com especial referencia as especies que causam dano a planta, na lagoa Olhos d'Agua, Minas Gerais, Brasil. Revista Brasileira de Entomologia. 42 (1 - 2): 9 - 12.","Diaz F, Torreias SRS, Spinelli GR, Donato M, Ronderos MM. 2014. A new species of Dasyhelea from Brazilian Amazonas and the description of the male of D. paulistana (Diptera: Ceratopogonidae). Acta Entomologica Musei Nationalis Pragae. 54: 715 - 728.","Dominiak P. 2012. Biting midges of the genus Dasyhelea Kieffer (Diptera: Ceratopogonidae) in Poland. Pol J Entomol. 81: 211 - 304.","Remm H. 1962. The genus Dasyhelea Kieffer in Estonia (Diptera, Heleidae). Tartu Riikliku Ulikooli Toimetised. 120: 108 - 133.","Remm H. 1979. A catalogue of the Ceratopogonidae (Diptera) of the Estonian S. S. R. Tartu: Eesti NSV Teaduste Akadeemia Tartu; p. 40 - 60.","Borkent A. 2014. The pupae of the Biting Midges of the World (Diptera: Ceratopogonidae), with a generic key and analysis of the phylogenetic relationships between genera. Zootaxa. 3879: 1 - 327. doi: 10.11646 / zootaxa. 3879.1.1.","Mayer K. 1934 b. Ceratopogoniden-Metamorphosen der Deutschen Limnologischen Sunda- Expedition. Archiv fur Hydrobiologie Suppl. 13 (1): 166 - 202.","Thomsen LC. 1935. New species of New York State Ceratopogonidae. J N Y Entomol Soc. 43: 283 - 297.","Thomsen LC. 1937. Aquatic Diptera. Part V. Ceratopogonidae. Memoirs Cornell Univ Agric Exp Station. 210: 57 - 80.","Wirth WW, Waugh WT. 1976. Five new Neotropical Dasyhelea midges (Diptera: Ceratopogonidae) associated with culture of cocoa. Studia Entomologica. 19: 223 - 236.","Diaz F, Ronderos MM, Spinelli GR, Ferreira-Keppler RL, Torreias SRS. 2013. A new species of Dasyhelea Kieffer (Diptera: Ceratopogonidae) from Brazilian Amazonia. Zootaxa. 3686: 85 - 93. doi: 10.11646 / zootaxa. 3686.1.5.","Duan C, Jiang XH, Chang QQ, Hou XH. 2019. First description of the immature stages of Dasyhelea alula and a redescription of adults from China (Diptera, Ceratopogonidae). ZooKeys. 824: 135 - 145. doi: 10.3897 / zookeys. 824.3172","Borkent A, Dominiak P. 2020. Catalog of the biting midges of the world (Diptera: Ceratopogonidae). Zootaxa. 4787 (1): 1 - 377. doi: 10.11646 / zootaxa. 4787.1.1.","Lu X, Duan C, Ning Y, Jiang XH, Hou XH. 2020. Morphology of the immature stages of Dasyhelea silvatica Wang, Zhang & Yu with redescriptions of adults (Diptera, Ceratopogonidae). ZooKeys. 961: 119 - 127. doi: 10.3897 / zookeys. 961.53882.","Borkent A, Dominiak P, Diaz F. 2022. An update and errata for the catalog of the biting midges of the world (Diptera: Ceratopogonidae). Zootaxa. 5120 (1): 53 - 64. doi: 10.11646 / zootaxa. 5120.1.3.","Brodskaya NK. 1995. A new species of biting midge of the genus Dasyhelea Kieff. (Diptera, Ceratopogonidae) from Leningrad region. Int J Dipterol Res. 6: 9 - 12.","Wirth WW, Linley JR. 1990. Description of Dasyhelea chani new species (Diptera: Ceratopogonidae) from leaves of the water lettuce (Pistia stratiotes) in Florida. Fla Entomol. 73: 273 - 279. doi: 10.2307 / 3494811.","Tokunaga M. 1961. Notes on biting midges II. Kontyu. 29: 180 - 185.","Wirth WW, Beaver RA. 1979. The Dasyhelea biting midges living in pitchers of Nepenthes in Southwest Asia (Diptera: Ceratopogonidae). Annales de la Societe Entomologique de France (N S). 15: 41 - 52."]}

spathicercus Wirth Dasyhelea spathicercus Wirth, 1969b: 577. Wirth 1974: 18 [catalogue]; Linsley 1977: 38 [checklist]; Borkent 1991: 105 [review]; Borkent & Wirth 1997: 58 [catalogue]; Borkent & Dominiak 2020: 53 [catalogue]. Distribution. Endemic. Galápagos: Fernandina, Santa Cruz. Remarks. This is an intertidal species (Wirth 1969b)., Published as part of Sinclair, Bradley J., 2023, An annotated checklist of the Diptera of the Galápagos Archipelago (Ecuador), pp. 1-102 in Zootaxa 5283 (1) on page 24, DOI: 10.11646/zootaxa.5283.1.1, http://zenodo.org/record/7912667, {"references":["Wirth, W. W. (1969 b) New species and records for Galapagos Diptera. Proceedings of the California Academy of Sciences, Series 4, 36 (20), 571 - 594.","Wirth, W. W. (1974) Family Ceratopogonidae. In: Papavero, N. (Ed.), A Catalogue of the Diptera of the Americas, south of the United States. Museu de Zoologia, Universidade de S \" o Paulo, 14, 1 - 89.","Linsley, E. G. (1977) Insects of the Galapagos (Supplement). Occasional Papers of the California Academy of Sciences, 125, 1 - 55.","Borkent, A. (1991) The Ceratopogonidae (Diptera) of the Galapagos Islands, Ecuador with a discussion of their phylogenetic relationships and zoogeographic origins. Entomologica scandinavica, 22 (1), 97 - 122. https: // doi. org / 10.1163 / 187631291 X 00336","Borkent, A. & Wirth, W. W. (1997) World species of biting midges (Diptera: Ceratopogonidae). Bulletin of the American Museum of Natural History, 233, 1 - 257. http: // hdl. handle. net / 2246 / 1622","Borkent, A. & Dominiak, P. (2020) Catalog of the biting midges of the world (Diptera: Ceratopogonidae). Zootaxa, 4787 (1), 1 - 377. https: // doi. org / 10.11646 / zootaxa. 4787.1.1"]}

sinclairi Evenhuis Pieza sinclairi Evenhuis, 2002a: 22. Evenhuis 2002b: 56 [catalogue]; Causton et al. 2006: 134 [introduced species]. Mythenteles species: Evenhuis 1990: 57 [distribution]; McMullen 1993: 98 [biology]; McMullen & Naranjo 1994: 27 [biology]; Peck 1996: 122 [mode of introduction]; Peck et al. 1998: 228 [introduced species]. Distribution. Introduced. Neotropical; Galápagos: Española, Floreana, Isabela, Marchena (ICCDRS), Pinta (ICCDRS), Rábida (ICCDRS), San Cristóbal, Santa Cruz, Santa Fé. Remarks. Evenhuis (1990, 2002a) believed that this species was introduced to the islands since there are no records prior to the 1980s, massive numbers in pan traps of recent collectors, records are primarily from inhabited islands and the genus is not known for its dispersal ability. The occurrence of this species on many uninhabited islands, including islands not visited by tourist boats, suggest that it can naturally disperse between islands. Adults of this species were collected from flowers of Scalesia on Pinta (McMullen 1993; McMullen & Naranjo 1994)., Published as part of Sinclair, Bradley J., 2023, An annotated checklist of the Diptera of the Galápagos Archipelago (Ecuador), pp. 1-102 in Zootaxa 5283 (1) on page 29, DOI: 10.11646/zootaxa.5283.1.1, http://zenodo.org/record/7912667, {"references":["Evenhuis, N. L. (2002 a) Pieza, a new genus of microbombyliids from the New World (Diptera: Mythicomyiidae). Zootaxa, 36 (1), 1 - 28. https: // doi. org / 10.11646 / zootaxa. 36.1.1","Evenhuis, N. L. (2002 b) Catalog of the Mythicomyiidae of the world (Insecta: Diptera). Bishop Museum Bulletin in Entomology, 10, 1 - 85.","Causton, C. E., Peck, S. B., Sinclair, B. J., Roque-Albelo, L., Hodgson, C. J. & Landry, B. (2006) Alien insects: threats and implications for conservation of Galapagos Islands. Annals of Entomological Society of America, 99 (1), 121 - 143. https: // doi. org / 10.1603 / 0013 - 8746 (2006) 099 [0121: AITAIF] 2.0. CO; 2","Evenhuis, N. L. (1990) Distributions of genera of microbombyliids (Bombyliodea: Mythicomyiinae) and their zoogeographical implications. Second International Congress of Dipterology, Abstract Volume, 57.","McMullen, C. K. (1993) Flower - visiting insets of the Galapagos Islands. Pan-Pacific Entomologist, 69 (1), 95 - 106.","McMullen, C. K. & Naranjo, S. J. (1994) Pollination of Scalesia baurii spp. hopkinsii (Asteraceae) on Pinta Island. Nocticias de Galapagos, 53, 25 - 28.","Peck, S. B. (1996) Origin and development of an insect fauna on a remote archipelago: The Galapagos Islands, Ecuador. In: Keast, A. & Miller, S. E. (Eds.), The origin and evolution of Pacific Island biotas, New Guinea to Eastern Polynesia: patterns and processes. Academic Publishing, Amsterdam, pp. 91 - 122.","Peck, S. B., Heraty, J., Landry, B. & Sinclair, B. J. (1998) Introduced insect fauna of an oceanic archipelago: the Galapagos Islands, Ecuador. American Entomologist, 44 (4), 218 - 237. https: // doi. org / 10.1093 / ae / 44.4.218"]}

ancora (Coquillett) Ceratopogon ancora Coquillett, 1902a: 87. Dasyhelea ancora: Borkent 1991: 111 [review]. Distribution. Cryptogenic. Nearctic; Galápagos: Isabela, Santa Cruz. Remarks. This species is recorded from both arid and humid zones (Borkent 1991)., Published as part of Sinclair, Bradley J., 2023, An annotated checklist of the Diptera of the Galápagos Archipelago (Ecuador), pp. 1-102 in Zootaxa 5283 (1) on page 23, DOI: 10.11646/zootaxa.5283.1.1, http://zenodo.org/record/7912667, {"references":["Coquillett, D. W. (1902 a) New Diptera from North America. Proceedings of the United States National Museum, 25 (1280), 83 - 126. https: // doi. org / 10.5479 / si. 00963801.25 - 1280.83","Borkent, A. (1991) The Ceratopogonidae (Diptera) of the Galapagos Islands, Ecuador with a discussion of their phylogenetic relationships and zoogeographic origins. Entomologica scandinavica, 22 (1), 97 - 122. https: // doi. org / 10.1163 / 187631291 X 00336"]}

atlantis Wirth & Williams Dasyhelea atlantis Wirth & Williams, 1957: 11. Borkent 1991: 108 [review]. Distribution. Native. Nearctic; Galápagos: Isabela, Santa Cruz. Remarks. Restricted to habitats immediately adjacent to the coast. Immatures were found in salty lagoons and brackish coastal tide pools (Borkent 1991)., Published as part of Sinclair, Bradley J., 2023, An annotated checklist of the Diptera of the Galápagos Archipelago (Ecuador), pp. 1-102 in Zootaxa 5283 (1) on page 23, DOI: 10.11646/zootaxa.5283.1.1, http://zenodo.org/record/7912667, {"references":["Wirth, W. W. & Williams, R. W. (1957) The biting midges of the Bermuda Islands, with descriptions of five new species (Diptera, Heleidae). Proceedings of the Entomological Society of Washington, 59 (1), 5 - 14.","Borkent, A. (1991) The Ceratopogonidae (Diptera) of the Galapagos Islands, Ecuador with a discussion of their phylogenetic relationships and zoogeographic origins. Entomologica scandinavica, 22 (1), 97 - 122. https: // doi. org / 10.1163 / 187631291 X 00336"]}

borgmeieri Wirth & Waugh Dasyhelea borgmeieri Wirth & Waugh, 1976: 225. Borkent 1991: 112 [review]. Distribution. Cryptogenic. Neotropical; Galápagos: Santa Cruz. Remarks. This species is recorded from middle elevations in more humid zones and it has been reared from a variety of rotting vegetation (Borkent 1991)., Published as part of Sinclair, Bradley J., 2023, An annotated checklist of the Diptera of the Galápagos Archipelago (Ecuador), pp. 1-102 in Zootaxa 5283 (1) on page 23, DOI: 10.11646/zootaxa.5283.1.1, http://zenodo.org/record/7912667, {"references":["Wirth, W. W. & Waugh, W. T. (1976) Five new Neotropical Dasyhelea midges (Diptera: Ceratopogonidae) associated with culture of Cocoa. Studia Entomologica, 19 (1 - 4), 223 - 236.","Borkent, A. (1991) The Ceratopogonidae (Diptera) of the Galapagos Islands, Ecuador with a discussion of their phylogenetic relationships and zoogeographic origins. Entomologica scandinavica, 22 (1), 97 - 122. https: // doi. org / 10.1163 / 187631291 X 00336"]}

paracincta Wirth Dasyhelea paracincta Wirth, 1969b: 576. Wirth 1974: 17 [catalogue]; Linsley 1977: 38 [checklist]; Borkent 1991: 107 [review]; Borkent & Wirth 1997: 57 [catalogue]; Borkent & Dominiak 2020: 51 [catalogue]. Distribution. Endemic. Galápagos: Floreana, Isabela, San Cristóbal, Santa Cruz. Remarks. Adults were collected from a variety of habitats from the dry coastal zone to 600 m, and immatures were found in brackish coastal pools, brackish seeps on water tanks and inland freshwater pools (Borkent 1991)., Published as part of Sinclair, Bradley J., 2023, An annotated checklist of the Diptera of the Galápagos Archipelago (Ecuador), pp. 1-102 in Zootaxa 5283 (1) on page 23, DOI: 10.11646/zootaxa.5283.1.1, http://zenodo.org/record/7912667, {"references":["Wirth, W. W. (1969 b) New species and records for Galapagos Diptera. Proceedings of the California Academy of Sciences, Series 4, 36 (20), 571 - 594.","Wirth, W. W. (1974) Family Ceratopogonidae. In: Papavero, N. (Ed.), A Catalogue of the Diptera of the Americas, south of the United States. Museu de Zoologia, Universidade de S \" o Paulo, 14, 1 - 89.","Linsley, E. G. (1977) Insects of the Galapagos (Supplement). Occasional Papers of the California Academy of Sciences, 125, 1 - 55.","Borkent, A. (1991) The Ceratopogonidae (Diptera) of the Galapagos Islands, Ecuador with a discussion of their phylogenetic relationships and zoogeographic origins. Entomologica scandinavica, 22 (1), 97 - 122. https: // doi. org / 10.1163 / 187631291 X 00336","Borkent, A. & Wirth, W. W. (1997) World species of biting midges (Diptera: Ceratopogonidae). Bulletin of the American Museum of Natural History, 233, 1 - 257. http: // hdl. handle. net / 2246 / 1622","Borkent, A. & Dominiak, P. (2020) Catalog of the biting midges of the world (Diptera: Ceratopogonidae). Zootaxa, 4787 (1), 1 - 377. https: // doi. org / 10.11646 / zootaxa. 4787.1.1"]}

mutabilis (Coquillett) Ceratopogon mutabilis Coquillett, 1901b: 602. Dasyhelea mutabilis: Borkent 1991: 109 [review]. Distribution. Native. Nearctic; Galápagos: Isabela, San Cristóbal, Santa Cruz. Remarks. Adults were collected only from habitats immediately adjacent to the coast, among mangroves and coastal pools (Borkent 1991)., Published as part of Sinclair, Bradley J., 2023, An annotated checklist of the Diptera of the Galápagos Archipelago (Ecuador), pp. 1-102 in Zootaxa 5283 (1) on page 23, DOI: 10.11646/zootaxa.5283.1.1, http://zenodo.org/record/7912667, {"references":["Coquillett, D. W. (1901 b) New Diptera in the U. S. National Museum. Proceedings of the United States National Museum, 23 (1225), 593 - 618. https: // doi. org / 10.5479 / si. 00963801.23 - 1225.593","Borkent, A. (1991) The Ceratopogonidae (Diptera) of the Galapagos Islands, Ecuador with a discussion of their phylogenetic relationships and zoogeographic origins. Entomologica scandinavica, 22 (1), 97 - 122. https: // doi. org / 10.1163 / 187631291 X 00336"]}

Three new species of biting midges belonging to the genus Dasyhelea Kieffer are described as a result of the extensive entomological surveys in the Deltaic Proper of Gangetic West Bengal, India. Adult and pupal stages of Dasyhelea (Dasyhelea) multiforamina sp. nov. and D. (Sebessia) bulbosa sp. nov., and all the life stages of D. (Pseudoculicoides) aprojecta sp. nov. are described. Dasyhelea (Prokempia) flaviformis Carter, Ingram and Macfie is also recorded first time from the Oriental region as only adult stage. They are described, illustrated and photomicrographed. A short note on bionomics of the four species is provided. The subgeneric placement of seven species previously described from India and keys to the Indian species of Dasyhelea based on male and female are also furnished. Key words: Dasyhelea, new species, new record, immature, Sebessia, Prokempia, Pseudoculicoides, Key, Oriental Region

Genus DASYHELEA Kieffer aliciae Grogan, Díaz, Spinelli and Ronderos, 2019: 308. Netherlands (Curaçao). aprojecta Brahma, Chatterjee and Hazra, 2020: 69. India. bulbosa Brahma, Chatterjee and Hazra, 2020: 69. India. curacaoensis Grogan, Díaz, Spinelli and Ronderos, 2019: 317. Netherlands (Curaçao). cyrstostyla Grogan, Díaz, Spinelli and Ronderos, 2019: 314. Netherlands (Curaçao). jinshajangensis Chen, Qian and Yu, in Han et al. 2020: 463. China (Sichuan). latiala Grogan, Díaz, Spinelli and Ronderos, 2019: 311. Netherlands (Curaçao). multiforamina Brahma, Chatterjee and Hazra, 2020: 56. India. patiae Szadziewski and Gwizdalska-Kentzer, 2020: 595. United Arab Emirates. recurva Grogan, Díaz, Spinelli and Ronderos, 2019: 310. Netherlands (Curaçao). rhopaloparamera Grogan, Díaz, Spinelli and Ronderos, 2019: 319. Netherlands (Curaçao)., Published as part of Borkent, Art, Dominiak, Patrycja & Díaz, Florentina, 2022, An Update and Errata for the Catalog of the Biting Midges of the World (Diptera: Ceratopogonidae), pp. 53-64 in Zootaxa 5120 (1) on page 56, DOI: 10.11646/zootaxa.5120.1.3, http://zenodo.org/record/6388819, {"references":["Grogan, W. L., Diaz, F, Spinelli, G. R. & Ronderos, M. M. (2019) The biting midges of the Caribbean island Curacao (Diptera: Ceratopogonidae). I. Species in the genus Dasyhelea Kieffer. Zootaxa, 4700 (3), 301 - 325. https: // doi. org / 10.11646 / zootaxa. 4700.3.1","Chatterjee, S., Brahma, S. & Hazra, N. (2020) Two new species of Culicoides Latreille (Diptera: Ceratopogonidae) from the Gangetic Plains of West Bengal, India with a key to the Indian species. Oriental Insects, 55, 1 - 41. https: // doi. org / 10.1080 / 00305316.2020.1775718","Han, S. - K., Bian, S. - Y., Wang, S. - L, Chen, J. - P. Liu, Y., Wang, P. - Z., Deng, Y., Qian, W. - P. & Yu, Y. - X. (2020) A new species and a new record of midges in Sichuan province, China (Diptera: Ceratopogonidae). Chinese Journal of Vector Biology & Control, 31 (4), 462 - 464. [in Chinese, English summary] https: // doi. org / 10.11853 / j. issn. 1003.8280.2020.04.017","Szadziewski, R. & Gwizdalska-Kentzer, M. (2020) New records of biting midges (Diptera: Ceratopogonidae) from the United Arab Emirates, with a description of a new species. Zootaxa, 4894 (4), 594 - 597. https: // doi. org / 10.11646 / zootaxa. 4894.4.6"]}

Most plant species depend upon insect pollination services, including many cash and subsistence crops. Plants compete to attract those insects using visual cues and floral odor which pollinators associate with a reward. The cacao tree, Theobroma cacao, has a highly specialized floral morphology permitting pollination primarily by Ceratopogonid midges. However, these insects do not depend upon cacao flowers for their life cycle, and can use other sugar sources. To understand how floral cues mediate pollination in cacao we developed a method for rearing Ceratopogonidae through several complete lifecycles to provide material for bioassays. We carried out collection and analysis of cacao floral volatiles, and identified a bouquet made up exclusively of saturated and unsaturated, straight-chain hydrocarbons, which is unusual among floral odors. The most abundant components were tridecane, pentadecane, (Z)-7-pentadecene and (Z)-8-heptadecene with a heptadecadiene and heptadecatriene as minor components. We presented adult midges, Forcipomyia sp. (subgen. Forcipomyia), Culicoides paraensis and Dasyhelea borgmeieri, with natural and synthetic cacao flower odors in choice assays. Midges showed weak attraction to the complete natural floral odor in the assay, with no significant evidence of interspecific differences. This suggests that cacao floral volatiles play a role in pollinator behavior. Midges were not attracted to a synthetic blend of the above four major components of cacao flower odor, indicating that a more complete blend is required for attraction. Our findings indicate that cacao pollination is likely facilitated by the volatile blend released by flowers, and that the system involves a generalized odor response common to different species of Ceratopogonidae.

D. (Ps.) sandrageorgei Dominiak, 2013 (Figs. 1C, D) Dasyhelea sandrageorgei Dominiak, 2013, in Dominiak & Alwin 2013: 139 (male, Lebanon). Material examined. Houbara Protected Area 1��� 28.02.2017, 15.03.2017, Malaise trap, 6 males, leg. A. Saji & A. van Harten; Jebel Hafeet National Park, 20.03.2017, Malaise trap, 1 male, leg. A. Saji & A. van Harten. Diagnosis. Males have a pair of slightly convergent apical aedeagal processes, which are membranous on their lateral parts, and a slender curved posterior process on the right paramere, that is distinctly bent distally. The gonostylus is evenly arched, with a short, ventral, finger-like projection on the basal portion (Figs. 1C, D). Females unknown. Distribution. Lebanon, UAE, February���March, May. This is the first record from the UAE., Published as part of Szadziewski, Ryszard & Gwizdalska-Kentzer, Marta, 2020, New records of biting midges (Diptera: Ceratopogonidae) from the United Arab Emirates, with a description of a new species, pp. 594-597 in Zootaxa 4894 (4) on page 594, DOI: 10.11646/zootaxa.4894.4.6, http://zenodo.org/record/4316811, {"references":["Dominiak, P. & Alwin, A. (2013) Five new species and new records of biting midges of the genus Dasyhelea Kieffer from the Near East (Diptera: Ceratopogonidae). Zootaxa, 3683 (2), 133 - 144. https: // doi. org / 10.11646 / zootaxa. 3683.2.3"]}

storai Borkent, 1997b: 14. New name for canariensis Storå. canariensis Storå, 1936: 36 (as variety of flavoscutellata Zetterstedt, preoccupied by Dasyhelea canariensis (Santos Abreu, 1918)). Canary Islands (Spain)., Published as part of Borkent, Art & Dominiak, Patrycja, 2020, Catalog of the Biting Midges of the World (Diptera: Ceratopogonidae), pp. 1-377 in Zootaxa 4787 (1) on page 54, DOI: 10.11646/zootaxa.4787.1.1, http://zenodo.org/record/4280910, {"references":["Borkent, A. (1997 b) The Ceratopogonidae (Diptera) described by Santos Abreu from the Canary Islands. Deutsche Entomologische Zeitschrift, 44, 3 - 18. https: // doi. org / 10.1002 / mmnd. 4800440102","Stora, R. (1936) Fam. Ceratopogonidae. In R. Frey (ed.). Die Dipterenfauna der Kanarischen Inseln und ihre Probleme: 31 - 38, pl. 6. Societas Scientiarum Fennica, Commentationes Biologicae 6 (1), 1 - 237.","Santos Abreu, E. (1918) Ensayo de una Monografia de los Tendipedidos de las Islas Canarias. Memorias de la Real Academia de Ciencias y Artes de Barcelona 14 (2), 159 - 326, 1 pl. https: // doi. org / 10.5962 / bhl. title. 8537"]}

sternalis Remm, 1980: 110 . Tajikistan., Published as part of Borkent, Art & Dominiak, Patrycja, 2020, Catalog of the Biting Midges of the World (Diptera: Ceratopogonidae), pp. 1-377 in Zootaxa 4787 (1) on page 54, DOI: 10.11646/zootaxa.4787.1.1, http://zenodo.org/record/4280910, {"references":["Remm, H. (1980) New species of the family Ceratopogonidae (Diptera) from the Middle Asia [in Russian, English summary]. Tartu Riikliku Ulikooli Toimetised 516, 85 - 128."]}

The ceratopogonid fauna of northern Sardinia (Italy) was investigated using black light traps. The study wascarried out in six localities from June to August 2015. A total of 3,879 ceratopogonids were collected and 40 speciesbelonging to 8 genera were identified. The most abundant genus was Culicoides(96%) followed by Forcipomyia(1.5%), Bezzia(1.2%) and Dasyhelea, Atrichopogon, Alluaudomyia, Stilobezziaand Brachypogon(each

Se presenta el primer relevamiento de Ceratopogonidae de la Reserva Natural Integral Punta Lara, en la provincia de Buenos Aires. Este área protegida representa uno de los relictos más australes de la selva en galería, donde el conocimiento de la fauna de insectos es aún limitado. Las capturas se realizaron en ambientes de pastizal y selva en galería, con trampas Malaise y trampas de luz, tipo CDC y embudo. Un total de 4.195 especímenes pertenecientes a nueve géneros y 39 especies de Ceratopogonidae fueron identificados. Se brinda un listado de las especies presentes en el área, y se proveen datos de su distribución geográfica. Veintitrés especies se citan en la Reserva por primera vez, mientras que Dasyhelea mediomunda Minaya, Forcipomyia (Forcipomyia) taragui Marino, Spinelli & Cazorla, Forcipomyia (Microhelea) fuliginosa (Meigen), Stilobezzia (Eukraiohelea) elegantula (Johannsen) y Stilobezzia (Stilobezzia) punctulata Lane se registran por primera vez en la provincia de Buenos Aires., We present the results of the first survey on the Ceratopogonidae from the “Reserva Natural Integral Punta Lara”, in Buenos Aires province. This protected area represents the southernmost relict of gallery forest where the knowledge of the insect fauna is still limited. Catches were conducted in areas of grassland and marginal forest with Malaise traps, CDC light traps and funnel light traps. A total of 4,195 specimens belonging to nine genera and 39 species of Ceratopogonidae were identified. A list of the species present in the protected area is provided, along with their distributional data. Twenty-three species are recorded from the reserve for the first time, while Dasyhelea mediomunda Minaya, Forcipomyia (Forcipomyia) taragui Marino, Spinelli & Cazorla, Forcipomyia (Microhelea) fuliginosa (Meigen), Stilobezzia (Eukraiohelea) elegantula (Johannsen) and Stilobezzia (Stilobezzia) punctulata Lane are recorded for the first time in Buenos Aires province., Facultad de Ciencias Naturales y Museo, Centro de Estudios Parasitológicos y de Vectores, Instituto de Limnología "Dr. Raúl A. Ringuelet"

Two new species of Dasyhelea Kieffer, D. yunga and D. allegrae, placed in the grisea and mutabilis speciesgroups respectively, are described, illustrated and photographed based on pupae and male and femaleadults. In addition, the immatures of D. azteca are described for the first time. The specimens werecollected in the southernmost area of the Argentinean Yungas. The pupae of D. yunga and D. allegraewere examined using a Compound Microscope (CM), while the larva and pupa of D. azteca by a ScanningElectron Microscope (SEM) and CM. Fil: Diaz, Maria Florentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Centro de Estudios Parasitológicos y de Vectores. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Centro de Estudios Parasitológicos y de Vectores; Argentina Fil: Spinelli, Gustavo Ricardo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Limnología "Dr. Raúl A. Ringuelet". Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Instituto de Limnología; Argentina Fil: Ronderos, Maria Marcela. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Centro de Estudios Parasitológicos y de Vectores. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Centro de Estudios Parasitológicos y de Vectores; Argentina

The immatures of the biting midge Dasyhelea silvatica are described and illustrated for the first time and a complete description of the adult male and female are provided using scanning electron and compound microscopes. The specimens were collected from flooded soil near a pond in Guizhou Province, China, and reared in the laboratory.

Dasyhelea cyrtostyla n. sp. (Figs. 22���30, 61���63) Diagnosis. The only Neotropical species of the mutabilis group with the following combination of characters. Males with apical portion of gonostylus very slender, recurved with a sharply pointed tip; gonocoxal apodemes broadly fused with paramere; paramere with apical section abruptly recurved and adpressed to ventral surface of paramere; and aedeagus with straight basal arch, posterolateral arms with mesal short, slender, subapical toothlike process. Females with moderately short antennal flagellum, flagellomeres 2���8 vasiform, slightly longer than broad, 9���12 slightly longer than 2���8, 13 longest with abruptly tapered apex; subgenital plate with semi-circular anterior portion with broad, ovoid lumen, posterior portion with 2 small posterior sub-basal triangular points; spermatheca large (57.5 x 32.5 ��m), ovoid, very heavily sclerotized, neck short, moderately stout, curved. Male. Head (Fig. 22). Dark brown. Eyes contiguous for a distance equal to the width of 3 ommatidia. Antennal flagellum moderately long; flagellomeres 2���4 very short, 5���9 rhomboid, 10���13 elongate, 10 longest, 13 broadest; antennal ratio 0.77���0.84 (0.80, n=2); flagellum length 0.43���0.47 (n=2) mm. Frontal sclerite elliptical, with elongate posterior extension. Clypeus with 4 pairs of setae. Palpus (Fig. 23) light brown; segment 3 as long as 4+5 combined with scattered capitate sensillae on mesal surface; palpal ratio 2.75���3.14 (2.94, n=2). Thorax. Dark brown; scutellum yellowish, with 6 large, 4 smaller setae. Wing (Fig. 24) moderately slender; membrane hyaline with moderately dense macrotrichiae; veins brown; radial cells without lumen; apices of costa, radius at perpendicular angle with wing margin; fork of CuA 1 just beyond level of apex of costa; without a well-defined anal lobe; wing length 0.62��� 0.70 (0.66, n=2) mm, width 0.22���0.27 (0.25, n=2) mm; costal ratio 0.40���0.46 (0.43, n=2). Abdomen. Dark brown. Genitalia (Figs. 25, 61���63). Tergite 9 broad basally, tapering progressively distally to rounded apex that extends to apices of gonocoxites; apicolateral process slender, with stout subapical seta; cercus conspicuous, located at base of apicolateral process with 4 thin setae. Sternite 9 (Figs. 25, 61) moderately short, 0.40 length of greatest width; posteromedian portion convex, extending to basal arch of aedeagus. Gonocoxite moderately slender, 1.9 x longer than broad; gonostylus as long as gonocoxite, broadest basally, tapering gradually distally, apical 1/5 very slender, recurved with a sharply pointed tip. Paramere and gonocoxal apodemes (Figs. 25, 62) heavily sclerotized, nearly symmetrical; gonocoxal apodemes stout, curved, nearly symmetrical, broadly fused with paramere; paramere stout on proximal 2/3, progressively tapering distally, distal 1/3 slender abruptly bent ventrally and adpressed to ventral surface of paramere, tip pointed. Aedeagus (Figs. 25, 63) massive, heavily sclerotized; 0.45 length of greatest width; basal arch very shallow, straight; basal arms short, rounded; posterolateral arms stout, heavily sclerotized, convergent with mesal short, slender, subapical toothlike process; a hyaline, semicircular structure lies between distal margin of basal arch and posterolateral arms. Female. Head (Fig. 26). Brown. Eyes contiguous for a distance equal to the width of 3 ommatidia. Antennal flagellum with flagellomeres 2���8 vasiform, slightly longer than broad, 9���12 subequal, slightly longer than 2���8, 13 longest with tapered apex; antennal ratio 0.82 (n=2); flagellum length 0.35���0.36 (n=2) mm. Frontal sclerite elliptical, with elongate ventral extension. Clypeus (Fig. 27) with 5 pairs of setae. Palpus (Fig. 28) light brown; segment 3 about as long as 4+5 combined with 3���4 capitate sensillae on mesal surface; palpal ratio 2.57���2.62 (2.59, n=2). Thorax. Brown; scutellum with 7 large, 3 smaller setae. Legs uniformly brown; tarsi pale, tarsomeres 5 infuscated; hind tibial comb with 7 spines; prothoracic tarsal ratio 1.83���2-08 (1.95, n=2), mesothoracic tarsal ratio 2.07���2.15 (2.11, n=2), metathoracic tarsal ratio 2.20-2.31 (2.25, n=2). Wing (Fig. 29) moderately broad; membrane hyaline with dense macrotrichiae; radial cells without lumen; apices of costa, radius at perpendicular angle with wing margin; fork of CuA 1 at level anterior to base of 2 nd radial cell; wing length 0.62 mm, width 0.30 mm; costal ratio 0.44. Halter pale. Abdomen brown. Subgenital plate (Fig. 30) sub-circular with ovoid central lumen; posteromedian margin rounded; posterolateral arms stout, heavily sclerotized, nearly straight, posterior margin with 2 small posterior sub-basal triangular points. Spermatheca (Fig. 30) large (57.5 x 32.5 ��m), ovoid, very heavily sclerotized; neck short, moderately stout, curved, length 7.5 ��m. Distribution. Cura��ao. Type material. Holotype male, allotype female, labeled ��� Cura��ao, Christoffel Nat. Pk., N car route, Thorn scrub area, 13-XI-2014, M. C. Thomas, Backlight trap (deposited in FSCA). Paratypes, 1 male, 1 female: 1 female with same data as holotype; 1 male with same data except Weg Narr, Playa Kanao, 12��9��7.82���� N, 68��52��9.13���� W, 10-XI-2014. Derivation of specific epithet. The specific epithet is a combination of two Greek terms, cyrto (curved) and styl (a pointed instrument), in reference to the slender, recurved apical section of the male gonostylus. Discussion. Males of D. curacaoensis n. sp. have a somewhat similar aedeagus to that of D. cyrtostyla n. sp., but in that species the basal arch is convex, moderately deep and the posterolateral arms are shorter with tapered sharply pointed apices. Males of D. curacaoensis n. sp. also differ from D. cyrtostyla n. sp. in having a paramere that has a J-shaped apical section, and much shorter gonostyli with broader, rounded apices. Females of D. curacaoensis n. sp. readily differ from those of D. cyrtostyla n. sp. in having a very short antennal flagellum with moniliform flagellomeres 2���12 and only 13 longer than broad, a broader frontal sclerite, a subgenital plate with a narrow slit-like lumen, and a much smaller (35 x 28 ��m vs. 57.5 x 32.5 ��m in D. curacaoensis) ovoid spermatheca with a broader neck. Males of Dasyhelea guadeloupensis Del��colle & Rieb (1994) also have a wing with a poorly developed anal lobe and an aedeagus that is somewhat similar to D. cyrtostyla n. sp., but in D. guadeloupensis the posterolateral arms are slender with sharply pointed tips and the basal arch is convex and very deep. Males of D. guadeloupensis also differ from D. cyrtostyla n. sp. in having an elongate sternite 9 that overlies most of the aedeagus with a trifurcate apex, a much shorter paramere with a broadly rounded apex, more elongate gonocoxites with a broad elongate, mesal subapical lobe, and gonostyli that are narrower basally, curved distally with broader, rounded apices. Females of Dasyhelea guadeloupensis also have more elongate flagellomeres 9���13 than in females of D. cyrtostyla n. sp., but females of that Guadeloupe species also have a much broader, apically rounded clypeus, a wing with a more poorly developed anal lobe, a longer more distally rounded subgenital plate without 2 small sub-basal triangular points, with a narrower lumen, and a smaller (44 x 37 ��m vs. 57.5 x 32.5 ��m in D. curacaoensis) globular spermatheca with a longer, retort-like neck., Published as part of Grogan, William L., D��az, Florentina, Spinelli, Gustavo R. & Ronderos, Maria M., 2019, The Biting Midges of the Caribbean island Cura��ao (Diptera: Ceratopogonidae) I. Species in the genus Dasyhelea Kieffer, pp. 301-325 in Zootaxa 4700 (3) on pages 314-315, DOI: 10.11646/zootaxa.4700.3.1, http://zenodo.org/record/3548509, {"references":["Delecolle, J. C. & Rieb, J. P. (1994) Contribution a l'etude des Ceratopogonides de la Guadeloupe. Description de trois especes nouvelles, appartenant aux genres Dasyhelea et Atrichopogon (Diptera, Nematocera). Bulletin de la Societe Entomologique de France, 99, 267 - 279."]}

Dasyhelea flavifrons (Gu��rin-M��neville) Ceratopogon flavifrons Gu��rin-M��neville, 1833: 165 (France). Dasyhelea flavifrons: Kieffer 1919: 51 (combination); Szadziewski & Dominiak 2006: 142 (in review of European synonyms in Ceratopogonidae); Borkent & Grogan 2009: 11 (in Nearctic catalog; distribution); Borkent, 2016: 65 (in online World catalog). Dasyhelea oppressa Thomsen, 1935: 285 (New York); Wirth 1965: 127 (in Nearctic catalog; distribution); Waugh & Wirth 1976: 230 (in revision of eastern USA Dasyhelea; distribution); Wilkening et al. 1985: 519 (Florida county records); Hribar & Grogan 2005: 231 (records from Monroe County, Florida); Szadziewski & Dominiak 2006: 142 [as synonym of D. flavifrons (Gu��rin-M��neville)]; Borkent & Grogan 2009: 11 (in Nearctic catalog; distribution; as synonym of D. flavifrons); Grogan et al. 2016: 208 (Guadeloupe records). Diagnosis. The only Neotropical species of the grisea group with the following combination of characters. Males with slender, distally recurved asymmetrical gonocoxal apodemes; paramere usually fused with right apodeme, the distal portion elongate, curved, apex usually bent ventrolaterally, tip pointed; gonocoxite with or without small basomesal spur; gonostylus straight to slightly curved, surface pubescent except for apex, tip rounded or pointed; aedeagus with heavily sclerotized basal portion and tapered posterolateral arms; and tergite 9 elongate, tapered distally with moderately narrow straight posterior margin and long cylindrical apicolateral processes. Females with subgenital plate bearing a pair of basolateral pointed projections and distal portion more or less arrowhead-shaped; and spermatheca subspherical with long neck region that is often recurved, retort-like. Discussion. This common Holarctic species was previously known in North America as D. oppressa Thomsen. Szadziewski & Dominiak (2006) regarded D. oppressa as a junior synonym of D. flavifrons and they listed 11 other specific synonyms in the Palearctic region. Dasyhelea flavifrons was recently recorded from the Neotropics in the Caribbean region from Guadeloupe (Grogan et al. 2016). We provide the first records of D. flavifrons from Cura��ao. Material examined. CURA��AO, Christoffel, Nat. Pk., Weg Narr, Playa Kanoa, 12��9��7.82���� N, 68��52��9.13���� W, 10-XI-2014, M. C. Thomas, blacklight trap, 1 female; same data except Weg Naar, Playa Kanoa nr. cement Plant, 6-XII-2015, 1 female; same data except Piscadera, 0.5 km N of Carmbi, 12��7��36.88���� N, 69��58��14���� W, 6-XI-2014, 1 female; same data except Piscadera Baai, 5-XII-2015, Blacklight trap in mangroves, 4 females; same data except Christoffel Nat. Pk., 12��20��40.29���� N, 69��06��29.59���� W, 9-X 1I-2015, R. Turnbow, Thorn scrub area, 13-XI- 2014,1 female. New Cura��ao record., Published as part of Grogan, William L., D��az, Florentina, Spinelli, Gustavo R. & Ronderos, Maria M., 2019, The Biting Midges of the Caribbean island Cura��ao (Diptera: Ceratopogonidae) I. Species in the genus Dasyhelea Kieffer, pp. 301-325 in Zootaxa 4700 (3) on pages 306-307, DOI: 10.11646/zootaxa.4700.3.1, http://zenodo.org/record/3548509, {"references":["Guerin-Meneville, F. E. (1833) Notice sur les metamorphoses des Ceratopogons et description de deux especes nouvelles de ce genre, decouvertes aux environs de Paris. Annales de la Societe de Entomologique de France, 2, 161 - 167.","Kieffer, J. J. (1919) Chironomides d'Europe conserves au Musee National Hongrois de Budapest. Annales Historico-Naturales Musei Nationalis Hungarici, 17, 1 - 160. https: // doi. org / 10.5962 / bhl. part. 29619","Szadziewski, R. & Dominiak, P. (2006) New synonyms of European Ceratopogonidae (Diptera). Annales Zoologici, Warszawa, 56, 139 - 146.","Borkent, A. & Grogan, W. L. Jr. (2009) Catalog of the New World Biting Midges North of Mexico (Diptera: Ceratopogonidae). Zootaxa, 2273 (1), 1 - 48. https: // doi. org / 10.11646 / zootaxa. 2273.1.1","Borkent, A. (2016) World Species of Biting Midges (Diptera: Ceratopogonidae). Available from: http: // wwx. inhs. illinois. edu / files / 4514 / 6410 / 0252 / CeratopogonidaeCatalog. pdf (accessed 28 January 2019)","Thomsen, L. (1935). New species of New York State Ceratopogonidae. Journal of the New York Entomological Society, 43, 283 - 297.","Wirth, W. W. (1965) Family Ceratopogonidae. In: Stone, A., Sabrosky, C. W., Wirth, W. W., Foote, R. H. & Coulson, J. R. (Eds.), A Catalog of the Diptera of America North of Mexico. United States Department of Agriculture, Agricultural Research Service, Agriculture Handbook 276. U. S. Government Printing Office, Washington, D. C., pp. 121 - 142.","Wirth, W. W. & Waugh, W. T. (1976) Five new Neotropical Dasyhelea midges (Diptera: Ceratopogonidae) associated with culture of Cocoa. Studia Entomologica, 19, 223 - 236.","Wilkening, A. J., Kline, D. L. & Wirth, W. W. (1985) An annotated checklist of the Ceratopogonidae (Diptera) of Florida with a new synonymy. Florida Entomologist, 68, 511 - 537. https: // doi. org / 10.2307 / 3494854","Hribar, L. J. & Grogan, W. L. Jr. (2005) New records of biting midges (Diptera: Ceratopogonidae) and frog-biting midges (Diptera: Corethrellidae) from the Florida Keys. Florida Scientist, 68, 227 - 235.","Grogan, W. L. Jr., Diaz, F., Spinelli, G. R. & Ronderos, M. M. (2016) The biting and predaceous midges of Guadeloupe (Diptera: Ceratopogonidae). II. Species of the subfamily Dasyheleinae. Zootaxa, 4184 (2), 201 - 254."]}

Dasyhelea Kieffer Dasyhelea Kieffer, 1911: 5. Type species, Dasyhelea halophila Kieffer, by monotypy. Prokempia Kieffer, 1913a: 163 (as subgenus of Dasyhelea). Type species, Dasyhelea ornaticornis Kieffer, 1913a: 180, desig- nated by Wirth 1973: 358. Pseudoculicoides Malloch, 1915: 309. Type species, Ceratopogon mutabilis Coquillett, 1901: 602, by original designation. Tetrahelea Kieffer, 1925: 423. Type species, Dasyhelea insignicornis Kieffer, 1913b: 9, by original designation. Dicryptoscena Enderlein, 1936: 51. Type species, Dasyhelea inclusa Kieffer, 1918: 188, by original designation. Sebessia Remm, 1979: 55 (as subgenus of Dasyhelea). Type species, Dasyhelea flavopyga Zilahi-Sebess, 1940: 49, by original designation. Borkentomyia Yu et al., 2006: 321. (as subgenus of Dasyhelea; systematic position unclear). Type species, Dasyhelea forsteri Grogan & Wirth, 1981: 97, by original designation., Published as part of Grogan, William L., D��az, Florentina, Spinelli, Gustavo R. & Ronderos, Maria M., 2019, The Biting Midges of the Caribbean island Cura��ao (Diptera: Ceratopogonidae) I. Species in the genus Dasyhelea Kieffer, pp. 301-325 in Zootaxa 4700 (3) on pages 302-303, DOI: 10.11646/zootaxa.4700.3.1, http://zenodo.org/record/3548509, {"references":["Kieffer, J. J. (1911) Nouvelles descriptions de chironomides obtenus d'eclosion. Bulletin de la Societe de Moselle, Metz, 27, 1 - 60. https: // doi. org / 10.5962 / bhl. part. 2983","Kieffer, J. J. (1913 a) Nouvelle etude sur les chironomides de l'Indian Museum de Calcutta. Records of the Indian Museum, 9, 119 - 197, pls. 11 - 12. https: // doi. org / 10.5962 / bhl. part. 16290","Wirth, W. W. (1973) Family Ceratopogonidae. In: Delfinado, M. D. & Hardy, D. E. (Eds.), A Catalogue of the Oriental Region. Vol. 1. Suborder Nematocera. University Press of Hawaii, Honolulu, pp. 346 - 388.","Malloch, J. R. (1915) The Chironomidae, or midges, of Illinois, with particular reference to the species occurring in the Illinois River. Bulletin of the Illinois State Laboratory of Natural History, 10, 275 - 543, 24 pls. https: // doi. org / 10.5962 / bhl. title. 9255","Coquillett, D. W. (1901) New Diptera in the U. S. National Museum. Proceedings of the United States National Museum, 23, 593 - 618. https: // doi. org / 10.5479 / si. 00963801.23 - 1225.593","Kieffer, J. J. (1925) Nouveaux genres et nouvelles especes de chironomides piqueurs. Archives de l'Institut Pasteur Algerie, 3, 405 - 430.","Kieffer, J. J. (1913 b) Chironomidae et Cecidomyidae. In: Alluaud, C. A. & Jeannel, R. (Eds.), Voyage de Ch. Alluaud et R. Jeannel en Afrique orientale (1911 - 1912). Resultats Scientigues. Diptera 5. A. Schulz, Paris, pp. 1 - 43. https: // doi. org / 10.5962 / bhl. title. 152165","Enderlein, G. (1936) Ordnung: Zweiflugler, Diptera. Abt. 16. In: Brohmer, P., Ehrmann, P. & Ulmer, G. (Eds.), Die Tierwelt Mitteleuropas 6: Insekten III Teil. Quelle und Meyer, Leipzig, pp. 1 - 259.","Kieffer, J. J. (1918) Beschreibung neuer, auf Lazarettschiffen des ostlichen Kriegsschauplatzes und bei Ignalino in Litauen von Dr. W. Horn gesammelter Chironomiden, mit Uebersichtstabellen einiger Gruppen von palaarktischen Arten (Dipt.). Entomologische Mitteilungen, 7, 35 - 53 + 94 - 110 + 163 - 170 + 177 - 188.","Remm, H. (1979) A catalogue of the Ceratopogonidae (Diptera) of the Estonian S. S. R. Eesti NSV Teaduste Akadeemia Tartu, 40 - 60 pp. [in Russian, Estonian, English summary]","Zilahi-Sebess, G. (1940) Heleiden aus Ungarn. Folia Entomologica Hungarica, 5, 10 - 133, 3 pls.","Yu, Y. - X., Liu, J. - H., Liu, G. - P., Liu, Z. - J., Hao, B. - S., Yan, G. & Zhao, T. - S. (2006) Ceratopogonidae of China, Insecta, Diptera. Vols. 1 - 2. Military Medical Science Press, Beijing, 1699 pp. [in Chinese, English title]","Grogan, W. L. Jr. & Wirth, W. W. (1981) New species of biting midges from the Solomon Islands (Diptera: Ceratopogonidae). Pacific Insects, 23, 93 - 100."]}

Dasyhelea recurva n. sp. (Figs. 6–10, 55–57) Diagnosis. The only Neotropical species of the mutabilis group with the following combination of characters. Males aedeagus heavily sclerotized with broad, stout basal arms, and slender posterolateral arms with recurved, bifid apices; gonocoxal apodemes and paramere asymmetrical, the right gonocoxal apodeme slender and narrowly fused to paramere, the left gonocoxal apodeme stout and broadly fused to paramere. Female unknown. Male. Head (Fig. 6). Dark brown. Eyes contiguous, the distance of contiguity not discernible in holotype, the only known specimen. Antennal flagellum dark brown; flagellomeres 2–9 rhomboidal, 10–13 elongate, only 10–13 sculptured; flagellomeres 2–10 with whorl of long sensillae chaetica comprising the moderately dense plume; antennal ratio 0.95; flagellum length 0.45 mm. Frontal sclerite elliptical, with slender moderately long, pointed, ventral projection. Clypeus with 5 pairs of setae. Palpus (Fig. 7) brown; segment 3 as long as 4+5 combined with 1–2 capitate sensillae on distal 1/4 of mesal surface; palpal ratio 3.60. Thorax (Fig. 8). Dark brown; scutellum with 6 large setae. Femora, tibiae dark brown, tarsi light brown, tarsomeres 5 infuscated; hind tibial comb with 6 spines; prothoracic tarsal ratio 1.91, mesothoracic tarsal ratio 2.25, metathoracic tarsal ratio 2.21. Wing (Fig. 9) membrane hyaline with moderately dense macrotrichiae; radial cells without lumen, 2 nd radial cell very short; apices of costa, radius at perpendicular angle with wing margin; fork of CuA 1 distad to level of apex of 2 nd radial cell; anal lobe moderately developed; wing length 0.62 mm, width 0.20 mm; costal ratio 0.44. Halter dark brown. Abdomen. Dark brown. Genitalia (Figs. 10, 55–57). Tergite 9 tapering gradually distally to rounded apex that extends to or near apices of gonocoxites; apicolateral processes parallel, slender, elongate with 1 medium-sized apical seta; cercus very small, located at base of apicolateral process with 4 setae, the 2 mesal medium-sized, external ones longer. Sternite 9 (Figs. 10, 55) 0.42 length of greatest width; anterior margin nearly straight; posterior portion tapered with broad, rounded apex that extends nearly to basal arch of aedeagus. Gonocoxite stout, 1.7 x longer than broad with 10–12 large setae on apicomesal surface and a slender baso-mesal tubercle; gonostylus as long as gonocoxite, distal portion slightly curved, tapering gradually distally with broader apex and pointed tip. Gonocoxal apodemes and paramere (Figs. 10, 56) asymmetrical; distal-most sections of gonocoxal apodemes broad, foot-shaped, apices rounded; right apodeme nearly straight, proximal portion slender, narrowly fused to paramere; left apodeme much broader than right, proximal portion broadly fused to paramere; paramere moderately stout basally, tapering slightly distally, apex apparently obliquely truncate. Aedeagus (Figs. 10, 57) heavily sclerotized, 0.55 length of greatest width; basal arch concave, extending 0.20 of total length of aedeagus; basal arms short, stout, apices tapered, recurved 60°; medio-posterior margin with 2 short, conical, apicolateral teeth; posterolateral arms convergent proximally, with abruptly recurved apical sections and bifid tips; posteromedian projection short, very lightly sclerotized, with blunt apex. Female. Unknown. Distribution. Curaçao. Type material. Holotype male, labeled “ Curaçao, Christoffel Nat. Pk., Christoffelberg, trailhead parking, 12°20ʹ40.63ʹʹ N, 69°67ʹ11.38ʹʹ W, 9-XI-2014, M. C. Thomas, BL trap ” (deposited in FSCA). Derivation of specific epithet. The specific name is a combination of two Latin terms, re (back, again) and curv (curved), a reference to the distinctive recurved apices of the posterolateral arms of the aedeagus. Discussion. At least two other primarily tropical New World species of Dasyhelea have aedeagal posterolateral arms with recurved apices: D. borgmeieri Wirth & Waugh (1976), that was originally described from Brazil, Grenada, Panama and Trinidad (type locality), then subsequently recorded from the Galápagos Islands (Ecuador) by Borkent (1991), and it was recently discovered in the USA from Florida (WLG; personal observations) and also from Belize (WLG; in FSCA); and, D. cacaoi Wirth & Waugh (1976) that is known only from Panama and Trinidad (type locality). The males of these two species differ from D. recurva n. sp. in having less massive aedeagi with apically pointed posterolateral arms, sternite 9 that extends just beyond the basal arch (D. borgmeieri) or, is very elongate, tapered and extends more than 1/2 the length of the posterolateral arms (D. cacaoi). Males of D. borgmeieri further differ from D. recurva n. sp. in having symmetrical gonocoxal apodemes and a short slender paramere, a greatly curved gonostylus and tergite 9 with very short apicolateral processes. Males of D. cacaoi also differ from D. recurva n. sp. in having more elongate gonocoxites, a longer tergite 9 with a convex apical margin and a longer, slender paramere with pointed curved apex. In addition, this species is also similar to D. curacaoensis n. sp., but the latter species differs from D. recurva in having a very short antennal flagellum, longer gonocoxites, shorter nearly straight gonostyli, a much broader tergite 9, and a longer paramere with curved, J-shaped apical section, and, aedeagus with straight, tapered posterolateral arms with pointed apices.

Dasyhelea latiala n. sp. (Figs. 11���21, 58���60) Diagnosis. The only Neotropical species of the mutabilis group with the following combination of characters. Males 3 rd palpal segment slender, palpal ratio 4.00; gonocoxal apodemes slightly asymmetrical; paramere, slender with bifid apex; aedeagus very complex, composed of two sections, a heavily sclerotized basal portion and a hyaline distal portion; and tergite 9 with rounded apical region that extends considerably beyond gonocoxites and elongate, stout, broadly divergent widely spaced apicolateral processes. Females with moderately long antennal flagellum, flagellomeres 2���8 moniliform to vasiform, 9���13 elongate, much longer than 2���8 and antennal ratio 0.78���0.97; palpus moderately long, segment 5 extending beyond tip of proboscis; wing very broad, length/width ratio 2.03���2.16; subgenital plate with a sub-circular basal portion with an ovoid lumen and a very broad trapezoidal apical portion; spermatheca moderately large (42.5 x 37.5 ��m), ovoid, heavily sclerotized with a short, slightly recurved neck. Male. Head (Fig. 11). Dark brown. Eyes contiguous for a distance equal to the width of 2 ommatidia. Antennal flagellum with flagellomeres 2���9 rhomboid, 10���13 elongate, 13 broader than 10���12; 2���10 with single whorl of long sensillae chaetica comprising the dense plume; antennal ratio 0.97; flagellum length 0.51 mm. Frontal sclerite rhomboidal, with slender elongate ventral extension. Clypeus (Fig. 12) with 4 pairs of setae. Palpus (Fig. 13) brown, elongate, apex extending considerably beyond tip of proboscis; segment 3 slender, as long as 4+5 combined, apex extending near apex of proboscis, with scattered capitate sensillae on inner surface; palpal ratio 4.00. Thorax. Scutum dark brown; scutellum pale brown with 6 large, 2 smaller setae. Wing (Fig. 14) moderately slender; membrane hyaline with moderately dense macrotrichiae; veins brown; 2 nd radial cell very short, open apically; apices of costa, radius perpendicular with wing margin; fork of CuA 1 at level of base of 2 nd radial cell; anal lobe moderately developed; wing length 0.80 mm, width 0.27 mm; costal ratio 0.46. Abdomen. Dark brown. Genitalia (Figs. 15, 58���60). Tergite 9 base broad, tapering progressively distally to rounded apex that extends considerably beyond gonocoxites; apicolateral processes elongate, stout, broadly divergent with subapical seta; cercus conspicuous, located at base of apicolateral process with 3���4 setae. Sternite 9 (Figs. 15, 58) 0.45 length of width; base broadly curved; posteromedian portion broad, tapering slightly distally with notched apex. Gonocoxite stout, 1.7 x longer than broad, distal margin straight, apicomesal surface with group of dense, moderately long setae; gonostylus nearly as long as gonocoxite, proximal 1/2 broad, tapering abruptly at mid-length, slightly curved with pointed tip. Paramere and gonocoxal apodemes (Figs. 15, 59) nearly symmetrical; gonocoxal apodemes slender on proximal 1/2, distal portions broader with small basal spur, apices truncate; paramere slender, nearly straight, apex bifurcate.Aedeagus (Figs. 15, 60) 0.60 length of greatest width, very complex, composed of two sections; basal arch stout, slightly curved, extending nearly 0.50 of total length, distal margin notched; basal arm stout, anterolaterally directed with mid-posterior outer tooth; posterior projection short, stout, with slightly expanded rounded tip; distal portion covered by a hyaline envelope that reaches tips of posteromedian projection and borne on pair of slender, heavily sclerotized processes directed anterolaterally, contiguous to inner margin of basal arms. Female. Head (Fig. 16). Dark brown. Eyes contiguous for a distance equal to the width of 2���3 ommatidia. Antennal flagellum with flagellomeres 2���8 moniliform to vasiform, 9���13 longer than 2���8, 13 longest; antennal ratio 0.78���0.97 (0.87, n=2); flagellum length 0.37���0.40 (n=2) mm. Frontal sclerite rhomboidal, with slender elongate ventral extension. Clypeus (Fig. 17) with 7 pairs of setae; anterior margin concave. Palpus (Fig. 18) light brown; segment 3 nearly as long as 4+5 combined, with scattered capitate sensillae on mesal surface; palpal ratio 3.66 (n=2). Thorax (Fig. 19). Scutum dark brown, paler on humeral angles and prescutellar depressions; scutellum yellowish, with 6 large, 2 smaller setae. Legs light brown, tarsomeres 5 more darkly infuscated; hind tibial comb with 6 spines; prothoracic tarsal ratio 2.14���2.25 (2.19, n=2), mesothoracic tarsal ratio 2.00, metathoracic tarsal ratio 2.18���2.22 (2.20, n=2). Wing (Fig. 20) very broad, length/width ratio 2.03���2.16; membrane hyaline with dense macrotrichiae; veins brown; 2 nd radial cell very short, open distally; apices of costa, radius at slight oblique angle with wing margin; fork of CuA 1 just anterior to base of 2 nd radial cell; anal lobe well developed; wing length 0.65���0.80 (0.72, n=2) mm, width 0.32���0.37 (0.35, n=2) mm; costal ratio 0.50 (n=2). Halter yellowish. Abdomen (Fig. 21). Dark brown. Subgenital plate (Fig. 21) with a sub-circular basal portion with an ovoid lumen and broad trapezoidal apical portion; posteromedian margin rounded with a small bifurcate posteromedian projection; posterolateral arms slender, recurved 30˚ at mid-length. Spermatheca (Fig. 21) small (42.5 x 37.5 ��m), ovoid, heavily sclerotized; neck short, slightly recurved, length 12.5 ��m. Distribution. Cura��ao. Type Material. Holotype male, labeled ��� Cura��ao, Piscadera Baai, 5-XII-2015, M. C. Thomas, Blacklight trap in mangroves���; allotype female with same data except 0.5 km N of Cambi, 12��07��36.88���� N, 69��58��1.14���� W, 6-XI- 2014 (deposited in FSCA). One female paratype with same data as holotype (deposited in MLPA). Derivation of specific epithet. The specific epithet was derived from two Latin terms, lati (broad, wide) and ala (wing), a reference to the very broad wing (Fig. 20) of females of this new species. Discussion. The paramere of the holotype male of D. latiala n. sp. is similar to males of D. yamana D��az & Spinelli (in D��az et al. 2014: Fig. 17E) from Tierra del Fuego province, Argentina which have a slightly shorter, narrower paramere with a more deeply bifid apex. In addition, the gonocoxal apodemes of D. yamana are also symmetrical with a short basolateral spur that is similar to the more elongate basolateral spurs on the larger, nearly symmetrical gonocoxal apodemes of D. latiala. However, males of D. yamana differ from the holotype of D. latiala in having a much longer wing (wing length 1.14���1.23 mm vs. 0.80 mm in D. latiala) that is not as narrow with a more poorly developed anal lobe; a more typical aedeagus with a large, stout posteromedian projection, a broad nearly straight basal arch and slender posterolateral arms with recurved tips; sternite 9 has a longer, more narrow distal extension; stouter gonostyli; and tergite 9 has shorter, less widely divergent apicolateral processes. Females of D. yamana differ from females of D. latiala n. sp. in having a shorter antennal flagellum with only flagellomeres 9���13 slightly longer than broad; a more elongate 3 rd palpal segment with a larger palpal ratio (4.00���4.80 vs. 3.66 in D. latiala); a longer wing (wing length 0.93���1.08 mm vs. 0.65���0.80 mm in D. latiala) with a shorter costa (costal ratio 0.41���0.45 vs. 0.50 in D. latiala) with a more poorly developed anal lobe; a broader frontal sclerite in which the dorsal portion is fused with the narrower ventral portion; a more typical semi-circular subgenital plate but without a broad trapezoidal anterior portion and posteromedian projection; and a spherical spermatheca with a short straight neck. Males of D. williamsi Wirth & Waugh (1976) have a very short broader paramere but the distal 2/3 is deeply bifid and their apices are broad and widely separated; the shape of the aedeagus is much different with long, stout lateral arms that are widely spaced basally, curved distally with broad nearly convergent tips; the distal 1/2 of the gonostyli are much broader; tergite 9 is much shorter with a straight margin and very short apicolateral processes; and sternite 9 is more elongate and extends just below the apex of the aedeagus. In addition, the thorax and much of the abdomen of both sexes of D. williamsi is mostly yellowish. Finally, the very unusual, greatly complex aedeagus of the holotype male of D. latiala does not resemble the aedeagi of other males in the mutabilis group including those in the fasciigera complex. Accordingly, it is possible that we have not correctly associated both sexes of this species. However, we are fairly confident of their association as all three specimens in the type series were collected in the same region and their overall colorations are similar., Published as part of Grogan, William L., D��az, Florentina, Spinelli, Gustavo R. & Ronderos, Maria M., 2019, The Biting Midges of the Caribbean island Cura��ao (Diptera: Ceratopogonidae) I. Species in the genus Dasyhelea Kieffer, pp. 301-325 in Zootaxa 4700 (3) on pages 311-314, DOI: 10.11646/zootaxa.4700.3.1, http://zenodo.org/record/3548509, {"references":["Diaz, F., Spinelli, G. R., Donato, M. & Ronderos, M. M. (2014) A taxonomic revision of the Patagonian species of the Dasyhelea mutabilis group with a phylogenetic analysis of New World species (Diptera: Ceratopogonidae). Journal of Natural History, 48, 2117 - 2175. https: // doi. org / 10.1080 / 00222933.2014.909062","Waugh, W. T. & Wirth, W. W. (1976) A revision of the genus Dasyhelea Kieffer of the eastern United States north of Florida (Diptera: Ceratopogonidae). Annals of the Entomological Society of America, 69, 219 - 247. https: // doi. org / 10.1093 / aesa / 69.2.219"]}

Dasyhelea azteca Huerta & Grogan Dasyhelea azteca Huerta & Grogan, 2006: 893 (Mexico); Borkent 2016: 62 (in online World catalog); D��az et al. 2018: 7 (Argentina; descriptions and figs. 4 th instar larva, male and female pupae). Diagnosis. The only Neotropical species of the grisea group with the following combination of characters. Males aedeagus with stout posterolateral arms, the right arm much shorter than the left arm; and the posterior margin of the sternite 9 with fine spicules. Females with a broad elliptical, bilobate frontal sclerite with a slender, very long ventral projection; flagellomeres 9-12 without sculpturing, only 13 with distinct sculpturing; and spermatheca pyriform, heavily sclerotized with long curved neck and large surface punctations. Discussion. This recently described species was based on specimens from the states of Morelos (type locality) and Jalisco, Mexico (Huerta & Grogan 2006). It was also recently discovered in the Neotropics from Argentina (D��az et al. 2018), and in the USA from Florida (Grogan; personal observation). Males are unique in having an aedeagus with unequal length posterolateral arms, but females are nearly identical with females of D. grisea (Coquillett). We provide the first records of D. azteca in the Caribbean region from Cura��ao. Material examined. CURA��AO, Christoffel, Nat. Pk., Weg Narr, Playa Kanoa, 12��9��7.82���� N, 68��52��9.13���� W, 10-XI-2014, M. C. Thomas, blacklight trap, 1 male, 1 female; same data except Playa Kanoa Rd., 12��09��33.02���� N, 68��52��50.36���� W, 6-XII-2015, R. Turnbow, 3 males; same data except The Woods, 12��21��2.78���� N, 69��06��13.82���� W, 7-XII-2015, R. Turnbow, BL trap, 1 female. New Cura��ao record., Published as part of Grogan, William L., D��az, Florentina, Spinelli, Gustavo R. & Ronderos, Maria M., 2019, The Biting Midges of the Caribbean island Cura��ao (Diptera: Ceratopogonidae) I. Species in the genus Dasyhelea Kieffer, pp. 301-325 in Zootaxa 4700 (3) on page 306, DOI: 10.11646/zootaxa.4700.3.1, http://zenodo.org/record/3548509, {"references":["Huerta, H. & Grogan, W. L. Jr. (2006) A new species and new record of biting midges of the genus Dasyhelea Kieffer (Diptera: Ceratopogonidae) from Morelos and Jalisco, Mexico. Proceedings of the Entomological Society of Washington, 108, 892 - 898.","Borkent, A. (2016) World Species of Biting Midges (Diptera: Ceratopogonidae). Available from: http: // wwx. inhs. illinois. edu / files / 4514 / 6410 / 0252 / CeratopogonidaeCatalog. pdf (accessed 28 January 2019)","Diaz, F., Spinelli, G. R. & Ronderos, M. M. (2018) Two new species of Dasyhelea Kieffer and the immature of D. azteca Huerta & Grogan from northwestern Argentina (Diptera: Ceratopogonidae). Zoologischer Anzeiger, 272, 6 - 19."]}

Dasyhelea aliciae n. sp. (Figs. 1���5, 52���54) Diagnosis. The only Neotropical species of the leptobranchia group with the following combination of characters. Males wing with a short costa (costal ratio 0.42); aedeagus stout, heavily sclerotized, H-shaped; gonocoxal apodemes and paramere asymmetrical, left gonocoxal apodeme barely contacting base of paramere; paramere sinuate with slender elongate recurved distal portion, the basal section of which is adpressed to the apex of the distal portion; sternite 9 moderately elongate, extending just below basal arch of aedeagus, distal margin slightly curved; and apex of tergite 9 broad, curved with very widely spaced elongate, divergent apicolateral processes. Female unknown. Male. Head (Fig. 1). Dark brown. Eyes contiguous for a distance equal to the width of 4���5 ommatidia. Antennal flagellum dark brown; flagellomeres 2���9 sculptured, rhomboidal, 10���13 elongate; flagellomeres 2���10 with whorl of long sensillae chaetica comprising the moderately dense plume; antennal ratio 0.88; flagellum length 0.47 mm. Frontal sclerite elliptical, with short pointed ventral projection, with folded lower marginal ear-like lobes, difficult to see. Clypeus with 5 pairs of setae. Palpus (Fig. 2) brown; segment 3 as long as 4+5 combined with 1���2 scattered hyaline capitate sensillae on distal 2/3 of mesal surface; palpal ratio 3.28. Thorax (Fig. 3). Dark brown; scutellum with 6 large, 3 smaller setae. Femora, tibiae dark brown, extreme bases slightly paler; tarsi with tarsomeres 1���4 light brown, tarsomeres 5 infuscated; hind tibial comb with 8 spines; prothoracic tarsal ratio 2.14, mesothoracic tarsal ratio 2.28, metathoracic tarsal ratio 2.11. Wing (Fig. 4) membrane hyaline with moderately dense macrotrichiae; radial cells without lumen, 2 nd radial cell very short; apices of costa, radius at perpendicular angle with wing margin; fork of CuA 1 slightly distal to level of apex of 2 nd radial cell; anal lobe moderately developed; wing length 0.65 mm, width 0.22 mm; costal ratio 0.42. Halter dark brown. Abdomen brown. Genitalia (Figs. 5, 52���54). Tergite 9 tapering progressively distally to rounded apex that extends to apices of gonocoxites; apicolateral processes very widely spaced, slender, elongate, cylindrical with medium-sized subapical seta; cercus very small, located at base of apicolateral process, with 2 small setae. Sternite 9 (Figs. 5, 52) 0.30 length of greatest width; anterior margin curved; distal portion moderately long, sub-triangular, tapered to broad apex that extends just below basal arch of aedeagus. Gonocoxite straight, moderately slender, 2.2 x longer than broad, with a few large setae on apicomesal side; gonostylus 0.80 length of gonocoxite, distal 1/2 greatly curved, tapering to narrow pointed tip. Gonocoxal apodemes and paramere (Figs. 5, 53) asymmetrical; proximal portions of apodemes stout; basal arm of right apodeme recurved 120��, proximal portion broadly fused to paramere; basal arm of left apodeme broad with bluntly rounded apex,��� distal portion slender, slightly recurved, barely contacting base of paramere; paramere stout, broad basally, tapering and curved distally, distal portion slender, abruptly recurved anteroventrally 180��, and adpressed to apex of proximal portion. Aedeagus (Figs. 5, 54) stout, H-shaped, very heavily sclerotized, 0.71 length of greatest width; basal arch broad, extending 0.30 of total length of aedeagus, posterior margin nearly straight; basal arms short, moderately divergent; posterolateral arms slightly convergent, tips broadly rounded. Female. Unknown. Distribution. Cura��ao. Type material. Holotype male, labeled ��� Cura��ao, Christoffel Nat. Pk., Christoffelberg trailhead parking, 12��20��40.63���� N, 69��67��11.38���� W, 9-XI-2014, M. C. Thomas, BL trap ��� (deposited in FSCA). Derivation of specific epithet. This species is named for Alicia D��az, sister of the 2nd author, Florentina D��az. Discussion. This new species is similar to the holotype male of the recently described D. waughi Grogan et al. (2016) from Guadeloupe which also has a stout H-shaped aedeagus; asymmetrical gonocoxal apodemes and paramere; dark brown scutum, femora and tibiae; wing with a moderately developed anal lobe; and a clypeus with 5 pairs of setae. However, the male of D. waughi differs from this new species in having a wing with a longer costa (costal ratio 0.50 vs. 0.42 in D. aliciae n. sp.), a longer sternite 9 that is narrowly tapered distally, an aedeagus with a broadly concave basal arch and the medio-distal margin has 3 small teeth, and, the apical margin of tergite 9 is more narrowly tapered and the apicolateral processes are parallel, not divergent. In addition, in the holotype of D. waughi, the right gonocoxal apodeme is only narrowly fused with the base of the paramere, but the basal portion of the paramere is much narrower, and the recurved apical section is shaped like a willow leaf and is much broader than in this new species., Published as part of Grogan, William L., D��az, Florentina, Spinelli, Gustavo R. & Ronderos, Maria M., 2019, The Biting Midges of the Caribbean island Cura��ao (Diptera: Ceratopogonidae) I. Species in the genus Dasyhelea Kieffer, pp. 301-325 in Zootaxa 4700 (3) on pages 308-310, DOI: 10.11646/zootaxa.4700.3.1, http://zenodo.org/record/3548509, {"references":["Grogan, W. L. Jr., Diaz, F., Spinelli, G. R. & Ronderos, M. M. (2016) The biting and predaceous midges of Guadeloupe (Diptera: Ceratopogonidae). II. Species of the subfamily Dasyheleinae. Zootaxa, 4184 (2), 201 - 254."]}

Dasyhelea corinneae Gosseries Ceratopogon scutellata Meigen, 1830: 262 (Europe; preoccupied by Say, 1829). Dasyhelea scutellata: Kieffer 1919: 50 (combination); Wirth 1965: 127 (in Nearctic catalog; distribution); Waugh & Wirth 1976: 236 (in revision of eastern USA Dasyhelea; distribution); Wilkening et al. 1985: 519 (Florida county records). Dasyhelea corinneae Gosseries, 1991: 42 (new replacement name for C. scutellata Meigen); Borkent & Grogan 2009: 11 (in Nearctic catalog; distribution); Grogan et al. 2010: 22 (records from Santa Rosa and Walton counties, Florida); Borkent, 2016: 64 (in online World catalog). Diagnosis. The only Neotropical species of the leptobranchia group with the following combination of characters. Males sternite 9 with elongate mid portion that extends beyond the apex of aedeagus and apical section expanded laterally; paramere with relatively short, distal portion, base broadly fused to right gonocoxal apodeme, distal section curved, apex usually not recurved ventrally; tergite 9 with moderately long, parallel apicolateral processes. Females with spermatheca ovoid to subspherical with a short slender straight or curved neck; subgenital plate reduced to a narrow transverse bridge with lateral arms; and abdominal sternite 8 with mid lateral row of 4���8 long, stout setae. Discussion. This common Holarctic species inhabits the eastern United States and Canada from Kansas, New York and New Brunswick south to Louisiana and Florida (Borkent & Grogan 2009). For over 70 years, it was known in North America as Dasyhelea scutellata (Meigen). Gosseries (1991) noted that Ceratopogon scutellata Meigen (1830) was preoccupied by Say (1829), and therefore; he proposed the new replacement name Dasyhelea corinneae, for this wide-ranging, very common species. We provide the first Neotropical records of D. corinneae in the Caribbean region from Cura��ao. Dominiak (2012) assigned D. corinneae to the subgenus Pseudoculicoides, but, we are doubtful it belongs in that subgenus because females lack a circular or ovoidal subgenital plate with a central lumen and male gonocoxal apodemes are narrower than in most males in that subgenus. Material examined. CURA��AO, Playa Kanoa Rd., 12��09��33.02���� N, 68��52��50.36���� W, 6-XII-2015, R. Turn- bow, BL trap, 3 females. New Cura��ao record., Published as part of Grogan, William L., D��az, Florentina, Spinelli, Gustavo R. & Ronderos, Maria M., 2019, The Biting Midges of the Caribbean island Cura��ao (Diptera: Ceratopogonidae) I. Species in the genus Dasyhelea Kieffer, pp. 301-325 in Zootaxa 4700 (3) on pages 307-308, DOI: 10.11646/zootaxa.4700.3.1, http://zenodo.org/record/3548509, {"references":["Meigen, J. W. (1830) Systematische Beschreibung der bekannten europaischen zweiflugeligen Insekten. Vol. 6. bei Friedrich Wilhelm Forftmann, Aachen, iv + 401 pp., 55 - 66 pls.","Say, T. (1829) Descriptions of North American dipterous insects. Journal of the Academy of Natural Sciences of Philadelphia (1829 - 1830), 6, 149 - 178, pls. 19 - 36.","Kieffer, J. J. (1919) Chironomides d'Europe conserves au Musee National Hongrois de Budapest. Annales Historico-Naturales Musei Nationalis Hungarici, 17, 1 - 160. https: // doi. org / 10.5962 / bhl. part. 29619","Wirth, W. W. (1965) Family Ceratopogonidae. In: Stone, A., Sabrosky, C. W., Wirth, W. W., Foote, R. H. & Coulson, J. R. (Eds.), A Catalog of the Diptera of America North of Mexico. United States Department of Agriculture, Agricultural Research Service, Agriculture Handbook 276. U. S. Government Printing Office, Washington, D. C., pp. 121 - 142.","Wirth, W. W. & Waugh, W. T. (1976) Five new Neotropical Dasyhelea midges (Diptera: Ceratopogonidae) associated with culture of Cocoa. Studia Entomologica, 19, 223 - 236.","Wilkening, A. J., Kline, D. L. & Wirth, W. W. (1985) An annotated checklist of the Ceratopogonidae (Diptera) of Florida with a new synonymy. Florida Entomologist, 68, 511 - 537. https: // doi. org / 10.2307 / 3494854","Gosseries, J. (1991) Ceratopogonidae. In: Grootaert, P., de Bruyn, L. & de Meyer, M. (Eds.), Catalogue of the Diptera of Belgium. Institut Royal des Sciences Naturelles de Belgique Documents de Travail, 70, pp. 41 - 45.","Borkent, A. & Grogan, W. L. Jr. (2009) Catalog of the New World Biting Midges North of Mexico (Diptera: Ceratopogonidae). Zootaxa, 2273 (1), 1 - 48. https: // doi. org / 10.11646 / zootaxa. 2273.1.1","Grogan, W. L. Jr., Hribar, L. J., Murphree, C. S. & Cilek, J. E. (2010) New records of biting and predaceous midges from Florida, including species new to the fauna of the United States (Diptera: Ceratopogonidae). Insecta Mundi, 0147, 1 - 59.","Borkent, A. (2016) World Species of Biting Midges (Diptera: Ceratopogonidae). Available from: http: // wwx. inhs. illinois. edu / files / 4514 / 6410 / 0252 / CeratopogonidaeCatalog. pdf (accessed 28 January 2019)","Dominiak, P. (2012) Biting midges of the genus Dasyhelea Kieffer (Diptera: Ceratopogonidae) in Poland. Polish Journal of Entomology, 81, 211 - 304. https: // doi. org / 10.2478 / v 10200 - 012 - 0009 - 8"]}

Key to adult Dasyhelea of Curaçao (females unknown for D. aliciae n. sp. & D. recurva n. sp.) 1 Males.............................................................................................. 2 - Females............................................................................................ 13 2 Aedeagus nearly covered with, or, only distal portion covered with thin hyaline envelope (Fig. 69); gonocoxal apodemes and paramere symmetrical (Fig. 68)......................................................................... 3 - Aedeagus not covered with thin hyaline envelope (Figs. 54, 57, 60, 63, 66); gonocoxal apodemes and paramere usually asymmetrical, rarely symmetrical (Figs. 53, 56, 59, 62, 65)........................................................ 5 3 Small species; wing length 0.85 mm, wing very broad, membrane without macrotrichia; paramere straight, club-shaped; aedeagus very complex, composed of two sections, a heavily sclerotized H-shaped basal portion and a broad, hyaline membranous quadrate posterior portion attached to the distal section of the basal portion, the apex of which is broad with short, blunt apicolateral extensions and an apicocentral lobe (Figs. 45, 67–69) (unplaced to species group)... D. rhopaloparamera n. sp. - Large species; wing length 1.25–1.50 mm, wing narrow, membrane with long macrotrichia, densest apically; paramere straight, tapered distally to pointed or rounded tip; aedeagus nearly entirely covered with thin hyaline envelope (cincta group)...... 4 4 Paramere elongate, slender with sharply pointed tip and base fused to both gonocoxal apodemes; bases of gonocoxal apodemes broad, foot-shaped; distal margin of hyaline envelope of aedeagus without crenulations.............. D. cincta (Coquillett) - Paramere relatively short, rod-like and not usually fused to gonocoxal apodemes; bases of gonocoxal apodemes slender, recurved; distal margin of hyaline envelope of aedeagus with crenulations...................... D. bahamensis (Johnson) 5 Frontal sclerite broad, elliptical, bilobate or heart-shaped; antennal flagellomeres 2–13 heavily sculptured, 13 elongate with a tapered slender apex; gonocoxal apodemes and paramere asymmetrical; gonocoxite often with basomesal hook or spur (grisea group)............................................................................................. 6 - Frontal sclerite slender, elliptical with or without a pair of lower marginal ear-like lobes; antennal flagellomeres 2–12 usually not sculptured, 13 short to moderately long with rounded apex; gonocoxal apodemes and paramere usually asymmetrical, rarely symmetrical; gonocoxite without basomesal hook or spur..................................................... 8 6 Aedeagus with unequal length posterolateral arms, right arm shorter than left arm; sternite 9 mid-portion margin with numerous small spicules................................................................... D. azteca Huerta & Grogan - Aedeagus with equal length posterolateral arms; sternite 9 mid-portion margin without numerous small spicules......... 7 7 Gonocoxite with large basomesal hook; paramere fused to both gonocoxal apodemes, apex curved or straight, not bent ventrad................................................................................ D. grisea (Coquillett) - Gonocoxite with or without small basomesal spur; paramere sometimes fused to only base of right gonocoxal apodeme, apex bent ventrad................................................................ D. flavifrons (Guérin-Méneville) 8 Sternite 9 distal margin extending to just below basal arch or as far as beyond apex of aedeagus (Fig. 5); frontal sclerite usually with pair of lower marginal ear-like lobes; aedeagus usually without a posteromedian projection (Fig. 54); distal margin of tergite 9 usually with short conical apicolateral processes (leptobranchia group)................................... 9 - Sternite 9 distal margin concave, straight or extending to base of or onto aedeagus but not beyond apex (Figs. 10, 15, 25, 34); frontal sclerite elliptical, without lower marginal ear-like lobes; aedeagus with a posteromedian projection (Figs. 57, 60, 63, 66); distal margin of tergite 9 usually with elongate conical or cylindrical apicolateral processes (mutabilis group)....... 10 9 Sternite 9 mid-portion greatly elongate, extending beyond apex of aedeagus, distal section expanded laterally; paramere relatively short, distal portion not recurved ventrally; tergite 9 apicolateral processes parallel.......... D. corinneae Gosseries - Sternite 9 mid-portion moderately elongate, extending just below basal arch of aedeagus, distal margin slightly curved; paramere long, distal portion elongate, slender, recurved ventrally; tergite 9 apicolateral processes divergent (Figs. 5, 52–53)............................................................................................. D. aliciae n. sp. 10 Paramere without tapered, recurved apical section (Fig. 59).................................................. 11 - Paramere with tapered, recurved apical section (Figs. 62, 65)................................................. 12 11 Paramere asymmetrical, broad basally with angled apex; aedeagus broad, heavily sclerotized, apical portion bifurcate with bifid, recurved apices; tergite 9 with parallel apicolateral processes (Figs. 10, 56–57)................... D. recurva n. sp. - Paramere nearly symmetrical, slender basally with bifid apex; aedeagus complex, composed of two sections, a heavily sclerotized basal portion and a hyaline distal portion; tergite 9 with broadly divergent apicolateral processes (Figs. 15, 58–60)............................................................................................ D. latiala n. sp. 12 Gonostylus elongate with very slender, recurved apical section; gonocoxal apodemes broadly fused with paramere; apical section of paramere abruptly recurved, adpressed to ventral side of paramere; aedeagus with very shallow basal arch, posterolateral arms with mesal short, slender, subapical toothlike process (Figs. 25, 62–63)........................ D. cyrtostyla n. sp. - Gonostylus moderately short with slender rounded apex; only right gonocoxal apodeme fused with paramere; apical section of paramere recurved, J-shaped, not adpressed to paramere; aedeagus with deep, curved basal arch, posterolateral arms with pointed apices, without mesal subapical toothlike process (Figs. 34, 65–66)...................... D. curacaoensis n. sp. 13 Two spermathecae (cincta group)....................................................................... 14 - One spermatheca.................................................................................... 15 14 Antennal flagellomeres 9–13 elongate, much longer than broad….. D. cincta (Coquillett) - Antennal flagellomeres 9–12 very short, only 13 slightly elongate, longer than broad............ D. bahamensis (Johnson) 15 Frontal sclerite broad, elliptical, bilobate or heart-shaped; antennal flagellomeres 2–13 usually heavily sculptured, 13 elongate with a tapered slender apex; subgenital plate usually with triangular, arrowhead-shaped or slender distal portion (grisea group)............................................................................................ 16 - Frontal sclerite slender, elliptical, with or without lower marginal ear-like lobes; antennal flagellomeres usually not heavily sculptured, flagellomere 13 short to moderately long with rounded apex; subgenital plate circular with a central lumen, disjunct, or, slender bridge-like................................................................................. 18 16 Subgenital plate with a pair of basolateral pointed projections and distal portion arrowhead-shaped; spermatheca subspherical, often retort-shaped with a long neck that is usually recurved.......................... D. flavifrons (Guérin-Méneville) - Subgenital plate without basolateral pointed projections, distal portion arrowhead-shaped or slender; spermatheca pyriform, with or without long slender neck....................................................................... 17 17 Subgenital plate lightly to moderately sclerotized, distal portion slender or arrowhead-shaped; spermatheca pyriform with large well-developed surface punctations, distal portion curved with long slender neck; flagellomeres 9-12 without sculpturing, only 13 with distinct sculpturing........................................................ D. azteca Huerta & Grogan - Subgenital plate heavily sclerotized with distinct arrowhead-shaped distal portion; spermatheca pyriform without numerous large, well-developed surface punctations, without a long neck; flagellomeres 9-13 with distinct sculpturing....... D. grisea (Coquillett) 18 Subgenital plate slender, bridge-like, without central lumen, or, arched at mid-length with poorly developed central lumen (Fig. 51); antennal flagellomeres 3–13 elongate or only 13 elongate (Fig. 46)......................................... 19 - Subgenital plate circular or ovoidal with central lumen (Figs. 21, 30, 41); antennal flagellomeres 3–12 usually relatively short, 9–13 sometimes longer than broad (Figs. 26, 35) (mutabilis group)............................................. 20 19 Antennal flagellomeres 2–12 vasiform, slightly longer than broad, only 13 elongate, much longer than broad; subgenital plate slender, abruptly elevated at mid length with a narrow disjunct curved sclerite and quadrate central lumen; wing membrane nearly devoid of macrotrichia, a few in margins of cells r 3 and m 1; spermatheca spheroidal, large (67 x 60 µm) with long, slender neck; sternite 8 with 2 separated lengthwise groups of 3–6 long setae (Figs. 46, 50–51) (unplaced to species group)...................................................................................... D. rhopaloparamera n. sp. - Antennal flagellomeres 2–13 elongate, 9-13 much longer than broad; subgenital plate slender, bridge-like, without disjunct anterior sclerite; wing membrane with dense macrotrichia in all cells; sternite 8 with mid lateral row of 4–8 long setae; spermatheca ovoid, small (45 x 30 µm) with short slender neck (leptobranchia group)…………. D. corinneae Gosseries - 20 Antennal flagellum very short (total length 0.31-0.33 mm), flagellomeres 2–12 very short, moniliform, only 13 longer than broad; wing with very short costa, costal ratio 0.23–25; subgenital plate with small narrow lumen; spermatheca small (35 x 28 µm) (Figs. 35, 40–41)................................................................ D. curacaoensis n. sp. - Antennal flagellum elongate to moderately long (total length 0.36–0.48 mm), flagellomeres 9–13 slightly to moderately elongate, longer than 2–8; wing with moderately long costa, costal ratio 0.44–0.50; subgenital plate with large broad lumen; spermatheca medium-size to large (42.5 x 37.5 µm–57.5 x 32.5 µm)............................................... 21 21 Antennal flagellomeres 9–13 elongate, much longer than 2–8; wing very broad with moderately long costa, costal ratio 0.50; subgenital plate with a sub-circular basal portion with a large ovoid lumen and a very broad trapezoidal apical portion; spermatheca medium-size (42.5 x 37.5 µm), ovoid, heavily sclerotized, neck short, slightly recurved (Figs. 16, 20–21)................................................................................................... D. latiala n. sp. - Antennal flagellomeres 2–12 slightly longer than broad, only 13 longer than 9–12; wing broad with moderately short costa, costal ratio 0.44; subgenital plate anterior portion semi-circular with broad, ovoid lumen, posterior portion of main body as broad as anterior portion with 2 small sub-basal triangular points; spermatheca large (57.5 x 32.5 µm), ovoid, very heavily sclerotized, neck short, moderately stout, curved (Figs. 26, 29–30)......................................... D. cyrtostyla n. sp.

Dasyhelea cincta (Coquillett) Ceratopogon cinctus Coquillett, 1901: 605 (Florida). Culicoides cinctus: Kieffer 1906: 54 (combination). Pseudoculicoides cinctus: Malloch 1915: 311 (combination). Dasyhelea cincta: Johannsen 1943: 778 (combination); Wirth 1952: 150 (California; distribution); Wirth & Williams 1957: 9 (Bermuda); Wirth 1965: 127 (in Nearctic catalog; distribution); Waugh & Wirth 1976: 222 (in revision of eastern USA Dasyhelea; distribution); Davies & Giglioli 1979: 592 (Grand Cayman); Spinelli & Wirth 1984: 597 (Jamaica, Haiti, Mexico, Panama); Huerta et al. 1999: 493 (records from Mexico); Borkent & Spinelli 2000: 25 (in New World catalog south of the USA; distribution); Borkent & Spinelli 2007: 60 (in Neotropical catalog; distribution); Borkent & Grogan 2009: 11 (in Nearctic catalog; distribution); Grogan et al. 2016: 208 (Guadeloupe); Borkent, 2016: 63 (in online World catalog). Dasyhelea albopicta Ingram & Macfie, 1931: 187 (Argentina); D��az et al. 2009: 150 [as synonym of D. cincta (Coquillett)] Dasyhelea penthesileae Macfie, 1935: 55 (Brazil), Spinelli, 1983: 407 (redescriptions of male, female, description of pupa; Argentina record). Diagnosis. The only Neotropical species of the cincta group with the following combination of characters. Males with elongate, slender paramere with sharply pointed tip and the base narrowly fused to both gonocoxal apodemes; bases of gonocoxal apodemes broad, foot-shaped; and aedeagus with distal margin of hyaline envelope of aedeagus without crenulations. Females with long antennal flagellum with flagellomeres 9���13 elongate, much longer than broad; spermathecae small (48 x 42 ��m), spherical to ovoid with numerous hyaline punctations and short, slender, oblique necks; and sternite 8 with heavily sclerotized distal portion that overlies the long quadrate subgenital plate which has a deep posterior excavation. Discussion. This widespread New World species inhabits the USA from California to New York south to Florida (Borkent & Grogan 2009), in the Caribbean region it is known from the Bahamas, Bermuda, Jamaica, Haiti, Puerto Rico, Dominica, Grenada, Guadeloupe, and from Mexico to South America in Ecuador, Trinidad, and Brazil south to Argentina (Spinelli & Wirth 1984; Borkent & Spinelli 2000, 2007; D��az et al. 2009; Grogan et al. 2016). We provide the first record of D. cincta from Cura��ao. Material examined. CURA��AO, Piscadera Baai, 5-XII-2015, M. C. Thomas, Blacklight trap in mangroves, 1 male. New Cura��ao record. Grisea Group The grisea group is equivalent to the subgenus Dasyhelea (Dominiak 2012), and species differ from all other groups/subgenera of Dasyhelea by their broad elliptical, bilobate or heart-shaped frontal sclerite, their antennal flagellomeres are heavily sculptured in both sexes and flagellomere 13 is elongate with a tapered slender apex. Males tergite 9 usually with long, cylindrical apicolateral processes, rarely with short, triangular apicolateral processes; sternite 9 relatively short with posterior margin concave, straight or convex; gonocoxal apodemes and paramere asymmetrical; gonocoxite often with basomesal hook or tubercle. Females subgenital plate entire and usually with triangular, arrowhead-shaped or slender distal section; and spermatheca pyriform, heavily sclerotized, and sometime with large, distinct surface punctations., Published as part of Grogan, William L., D��az, Florentina, Spinelli, Gustavo R. & Ronderos, Maria M., 2019, The Biting Midges of the Caribbean island Cura��ao (Diptera: Ceratopogonidae) I. Species in the genus Dasyhelea Kieffer, pp. 301-325 in Zootaxa 4700 (3) on pages 305-306, DOI: 10.11646/zootaxa.4700.3.1, http://zenodo.org/record/3548509, {"references":["Coquillett, D. W. (1901) New Diptera in the U. S. National Museum. Proceedings of the United States National Museum, 23, 593 - 618. https: // doi. org / 10.5479 / si. 00963801.23 - 1225.593","Kieffer, J. J. (1906) Diptera. Fam. Chironomidae. In: Wytsman, P. (Ed.), Genera Insectorum, Fasc., 42, pp. 1 - 78, 4 pls.","Malloch, J. R. (1915) The Chironomidae, or midges, of Illinois, with particular reference to the species occurring in the Illinois River. Bulletin of the Illinois State Laboratory of Natural History, 10, 275 - 543, 24 pls. https: // doi. org / 10.5962 / bhl. title. 9255","Johannsen, O. A. (1943) A generic synopsis of the Ceratopogonidae (Heleidae) of the Americas, a bibliography, and a list of the North American species. Annals of the Entomological Society of America, 36, 763 - 791. https: // doi. org / 10.1093 / aesa / 36.4.763","Wirth, W. W. (1952) The Heleidae of California. University of California Publications in Entomology, 9, 95 - 266.","Wirth, W. W. & Williams, R. W. (1957) The biting midges of the Bermuda Islands, with descriptions of five new species (Diptera, Heleidae). Proceedings of the Entomological Society of Washington, 59, 5 - 14. https: // doi. org / 10.1016 / S 0272 - 2712 (18) 30394 - 9","Wirth, W. W. (1965) Family Ceratopogonidae. In: Stone, A., Sabrosky, C. W., Wirth, W. W., Foote, R. H. & Coulson, J. R. (Eds.), A Catalog of the Diptera of America North of Mexico. United States Department of Agriculture, Agricultural Research Service, Agriculture Handbook 276. U. S. Government Printing Office, Washington, D. C., pp. 121 - 142.","Wirth, W. W. & Waugh, W. T. (1976) Five new Neotropical Dasyhelea midges (Diptera: Ceratopogonidae) associated with culture of Cocoa. Studia Entomologica, 19, 223 - 236.","Davies, J. E. & Giglioli, M. E. C. (1979) The Ceratopogonidae (Diptera) of Grand Cayman, West Indies: species and ecological notes. Mosquito News, 39, 586 - 594.","Spinelli, G. R. & Wirth, W. W. (1984) Las especies Neotropicales del genero Dasyhelea, grupo cincta (Diptera: Ceratopogonidae). Limnobios, 2, 586 - 608","Huerta, H., Ibanez-Bernal, S. & Felippe-Bauer, M. L. (1999) New records of biting midges (Diptera: Ceratopogonidae) in Mexico. Entomologia y Vectores, 6, 491 - 499.","Borkent, A. & Spinelli, G. R. (2000) Catalog of the New World biting midges south of the United States of America (Diptera: Ceratopogonidae). Contributions on Entomology, International, 4, 1 - 107.","Borkent, A. & Spinelli, G. R. (2007) Neotropical Ceratopogonidae (Diptera: Insecta). In: Adis, J., Arias, J. R., Rueda-Delgado, G. & Wantzen, K. M. (Eds.), Aquatic biodiversity in Latin America (ABLA). Vol. 4. Pensoft, Sofia-Moscow, pp. 1 - 198.","Borkent, A. & Grogan, W. L. Jr. (2009) Catalog of the New World Biting Midges North of Mexico (Diptera: Ceratopogonidae). Zootaxa, 2273 (1), 1 - 48. https: // doi. org / 10.11646 / zootaxa. 2273.1.1","Grogan, W. L. Jr., Diaz, F., Spinelli, G. R. & Ronderos, M. M. (2016) The biting and predaceous midges of Guadeloupe (Diptera: Ceratopogonidae). II. Species of the subfamily Dasyheleinae. Zootaxa, 4184 (2), 201 - 254.","Borkent, A. (2016) World Species of Biting Midges (Diptera: Ceratopogonidae). Available from: http: // wwx. inhs. illinois. edu / files / 4514 / 6410 / 0252 / CeratopogonidaeCatalog. pdf (accessed 28 January 2019)","Ingram, A. & Macfie, J. W. S. (1931) Ceratopogonidae. In: Diptera of Patagonia and South Chile. Part II. Fascicle 4. Nematocera. British Museum (Natural History), London, pp. 155 - 232.","Diaz, F., Spinelli, G. R. & Ronderos, M. M. (2009) Biting midges of the Dasyhelea cincta group from Patagonia (Diptera: Ceratopogonidae). Deutsche Entomologische Zeitschrift, 56, 149 - 156. https: // doi. org / 10.1002 / mmnd. 200900013","Macfie, J. W. S. (1935) Ceratopogonidae (Dipt.) from the river Amazon. Stylops, 4, 49 - 56. https: // doi. org / 10.1111 / j. 1365 - 3113.1935. tb 00555. x","Spinelli, G. R. (1983) Notas sobre Ceratopogonidae (Diptera Nematocera) de la Republica Argentina. I. Una nueva especie del genero Alluaudomyia Kieffer, redescripcion de Dasyhelea penthesileae Macfie, y nuevas citas para el genero Stilobezzia Kieffer. Limnmobios, 2, 403 - 411.","Dominiak, P. (2012) Biting midges of the genus Dasyhelea Kieffer (Diptera: Ceratopogonidae) in Poland. Polish Journal of Entomology, 81, 211 - 304. https: // doi. org / 10.2478 / v 10200 - 012 - 0009 - 8"]}

We document the first records of biting midges in the genus Dasyhelea Kieffer (Diptera: Ceratopogonidae) from Curaçao: Dasyhelea azteca Huerta & Grogan, D. bahamensis (Johnson), D. cincta (Coquillett), D. corinneae Gosseries, D. flavifrons (Guérin-Méneville) and D. grisea (Coquillett). The following six new species of Dasyhelea are described from this Caribbean island and their key features are presented in color photographs and illustrations: D. aliciae n. sp., D. recurva n. sp., D. latiala n. sp., D. rhopaloparamera n. sp., D. cyrtostyla n. sp., and D. curacaoensis n. sp. We also provide a key to males and females of all known species of Dasyhelea from Curaçao. Fil: Grogan, William L.. Florida State University; Estados Unidos Fil: Díaz, Florentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Centro de Estudios Parasitológicos y de Vectores. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Centro de Estudios Parasitológicos y de Vectores; Argentina Fil: Spinelli, Gustavo Ricardo. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División Entomología; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Limnología "Dr. Raúl A. Ringuelet". Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Instituto de Limnología; Argentina Fil: Ronderos, Maria Marcela. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Centro de Estudios Parasitológicos y de Vectores. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Centro de Estudios Parasitológicos y de Vectores; Argentina

SummaryPupa, male and female adults of Dasyhelea (Prokempia) flava Carter, Ingram & Macfie, pupa and male adult of D. (Pseudoculicoides) acuta n. sp., and male adult of D. (Pseudoculicoides) comosa n. sp. are described and illustrated.

Synanthropy and community structure of Ceratopogonidae in temperate Argentina were compared across an urbanization gradient to assess the impact of urbanization on species diversity and abundance. Adult ceratopogonids were collected from one natural, one Peri-urban, and two urban sites monthly for 12 months using CDC light traps. Nine-hundred one individuals belonging to nine genera and 59 species were collected. Forcipomyia Meigen was the genus with the highest species richness (n = 20), followed by Atrichopogon Kieffer (14), Dasyhelea Kieffer (8), Stilobezzia Kieffer (7), Culicoides Latreille (5), Bezzia Kieffer (2), Alluaudomyia Kieffer, Monohelea Kieffer, and Palpomyia Meigen (1). Only 15 species were present at the three different areas. Indices of species abundance (ISA) values were statistically similar between urban and Peri-urban areas, but these were significantly lower than those from the natural area. Atrichopogon balseiroi Spinelli, Dasyhelea sp. C., Forcipomyia poulaineae Ingram, and Macfie and F. nana (Macfie) were positively associated with rain fall, while A. albinensis, A. delpontei, D. suarezi, Forcipomyia sp. B, F. sp. I, F. pinamarensis, and F. calchaqui were associated to relative humidity. Species diversity was reduced in urbanized areas and these areas favored ubiquitous species: Forcipomyia taragui Marino, Spinelli and Cazorla, F. poulaineae, and Culicoides venezuelensis Ortiz and Mirsa. Fil: Cazorla, Carla Gisela. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División Entomología; Argentina Fil: Campos, Raul Ernesto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Instituto de Limnología "Dr. Raúl A. Ringuelet". Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Instituto de Limnología; Argentina

The fourth instar larva and pupa of Dasyhelea flavifrons Guerin-Meneville are redescribed, illustrated, and photomicrographed using binocular, phase-contrast, and scanning electron microscopy. Comparisons with the American species of the grisea group were made. The immatures were collected by using a siphon bottle in tree-holes and from water collected in dead snail shells in Salta Province, Argentina, transported to the laboratory and there bred to the emergence of the adults. Details on larval habitats are given. These are the first records from Argentina and in gastrotelmata., Centro de Estudios Parasitológicos y de Vectores

Two new species of the dipteran genus Dasyhelea Kieffer, viz. Dasyhelea silvatica sp.nov., Wang, Zhang and Yu and Dasyhelea arrhena sp.nov., Wang, Zhang and Yu, are described from China. The type specimens are deposited with the Medical Entomology Collection Gallery, Beijing, China. http://zoobank.org/urn:lsid:zoobank.org:pub:2D82EBD2-03B0-4A05-891B-9A548F32DA26.

The water-filled tree hole fauna of biting midges (Diptera: Ceratopogonidae) in Slovakia is documented for the first time. Three Culicoides Latreille and two Dasyhelea Kieffer species were reared from these habitats. Epidemiological significance of C. obsoletus (Meigen, 1818) is briefly discussed.

Dasyhelea nelidae new species (Figs. 25���26) Diagnosis. The only Neotropical species of the fasciigera complex with males having an elongate sub-triangular sternite 9 with a greatly tapered distal portion with irregular margins and slender apex, an inner gonostylus with a short subapical prong and an aedeagus covered with a thin hyaline envelope. Females have an infuscated wing membrane, a small (length 34 ��m) ovoid spermatheca with a short broad neck, and a subgenital plate with a circular, lightly sclerotized anterior portion, a slightly rectangular heavily sclerotized posterior portion and a small lumen. Male. Head (Fig. 25 A). Dark brown. Eyes contiguous for a distance equal to width of 3 ommatidia. Frontal sclerite rhomboid with slender ventral projection extending to base of clypeus. Antennal flagellum 2���9 rhomboid with flagellomeres 9���13 elongate, 13 broad, much shorter than 9���12; flagellomeres 2���10 with whorl of long sensilla chaetica forming the dense plume that extends to or beyond base of flagellomere 13; antennal ratio 0.84��� 0.94 (0.88, n=7). Palpus (Fig. 25 B) light brown; segment 3 moderately long with scattered mesal capitate sensilla; palpal ratio 3.28���3.42 (3.36, n=5). Thorax. Dark brown. Scutellum golden brown with 6 large, 1 smaller setae. Prothoracic tarsal ratio 2.17���2.35 (2.47, n=6), mesothoracic tarsal ratio 2.68���2.70 (2.74, n=6), metathoracic tarsal ratio 2.21���2.38 (2.32, n=6). Wing (Fig. 25 C) slender; membrane slightly infuscated with dense macrotrichia; veins dark brown; 2nd radial cell moderately well developed; junction of costa, radius forming a poorly developed acute angle; fork of CuA1 at level just beyond apex of costa, radius; wing length 0.80���0.88 (0.84, n=5) mm, width 0.26��� 0.30 (0.28, n=5) mm; costal ratio 0.45���0.47 (0.46, n=5). Genitalia (Figs. 25 D, 26A). Tergite 9 tapering gradually distally to moderately broad, truncate apex that extends to or beyond apices of gonocoxites; apicolateral process heavily sclerotized, elongate, cylindrical with moderately short apical seta; cercus apparently represented by 2 shorter lobes arising from basal portion of apicolateral process with short apical setae. Sternite 9 (Figs. 25 D, 26B) 0.50 length of greatest width; distal portion sub-triangular, greatly tapered with irregular margins and slender apex. Gonocoxite straight, stout, 1.55 x longer than greatest width; distal 2/3 becoming increasingly broader, with subapicomesal pointed triangular process, distal margin rounded. Gonostylus (Fig 26 C) bifurcate; outer gonostylus broad on proximal 1/4, distal portion slightly curved, slender with pointed apex; inner gonostylus half as long as outer gonostylus, proximal 2/3 broad, distal 1/3 tapered, curved, distal portion slender with short subapical prong and sharply pointed tip. Gonocoxal apodemes and paramere (Figs. 25 D, 26D) in form of asymmetrical structure; gonocoxal apodemes heavily sclerotized, right apodeme short, curved, broadly fused to paramere, left apodeme much longer, basal portion moderately broad, recurved, distal portion slender, slightly sinuate, extreme apical section narrowly fused to paramere; paramere broad proximally, distal portion curved, slightly tapered with acute, pointed tip. Aedeagus (Figs. 25 D, 26E) 0.50 length of greatest width, covered with thin hyaline envelope which extends from near apices of basal arms to apex of sternite 9; basal arch heavily sclerotized, moderately shallow, extending 0.33 of total aedeagus length, distal margin dentate; basal arm slender, slightly recurved; posterolateral arm heavily sclerotized, apically divided, each half crossing each other. Female. Head (Fig. 25 E). Dark brown. Eyes contiguous for a distance equal to the width of 2 ommatidia. Antennal flagellum with flagellomeres 2���13 vasiform, 9���13 slightly longer than 2���8; antennal ratio 0.65���0.76 (0.70, n=2). Clypeus (Fig. 25 F) with broad, detached wing-like proximal portion; distal portion narrow with 4 pairs of setae. Palpus (Fig. 25 G) pale brown; segment 3 moderately short with 3���4 capitate sensillae on proximal 2/3 of mesal surface; palpal ratio 2.87 (n=3). Thorax. Scutum dark brown; scutellum paler with 6 large, 2 smaller setae. Femora, tibiae dark brown, tarsi slightly paler; hind tibial comb with 5 spines; prothoracic tarsal ratio 2.00���2.22 (2.07, n=3), mesothoracic tarsal ratio 2.22���2.55 (2.39, n=3), metathoracic tarsal ratio 2.08���2.09 (2.08, n=3). Wing (Fig. 25 H) moderately broad; membrane infuscated, covered with dense macrotrichia except for a narrow area posterior of radial cells; veins dark brown; 2nd radial cell reduced to a short suture; fork of CuA1 at level of apex of costa, radius; anal lobe moderately well developed; wing length 0.70���0.74 (0.72, n=3) mm, width 0.30���0.32 (0.31, n=3) mm; costal ratio 0.42���0.46 (0.44, n=3). Halter dark brown. Abdomen (Fig. 25 I). Dark brown. Subgenital plate with a circular, lightly sclerotized anterior portion, a slightly rectangular heavily sclerotized posterior portion and a small lumen; posterior margin straight; posterolateral arms slender, recurved. Spermatheca small (34 x 26 ��m), ovoid, heavily sclerotized, neck short, length 4 ��m. Distribution. Guadeloupe. Type material. Holotype male labeled ��� Guadeloupe, Basse Terre, NE Pigeon (16.14404�� N, 61.74977 �� W), 18-V-2012, RH Turnbow, BL trap ��� (deposited in FSCA); allotype female same data as holotype except 23-V-2012. Paratypes 6 males, 3 females, as follows: same data as holotype, 3 males, 1 female; same data except 17-V-2012, 1 male; same data as allotype 3 males, 1 female; same data except Pointe a Lezard, 17-V-2012, 1 female. Derivation of specific epithet. This species is named after Mrs. N��lida Caligaris, artistic technician of the Divisi��n Entomolog��a, Museo de La Plata. Discussion. This new species is a member of the Dasyhelea fasciigera complex as defined by Remm (1962) and Borkent & Forster (1986), which Dominiak (2012) included in the subgenus Pseudoculicoides (= mutabilis group). Males in the Fasciigera complex differ from males of all other species in the mutabilis group by their bifurcate gonostyli. This complex is composed of the following six species that inhabit the Americas (Borkent & Forster 1986; Dominiak & Szadziewski 2010; Dominiak 2012): Dasyhelea bifida Zilahi-Sebess (1936), D. crassiseta Borkent & Forster, D. fasciigera Kieffer (1925b), D. furva Remm (1967), D. pollex Borkent & Forster, and D. tristyla Wirth. Males of D. bifida differ from this new species by their triangular tergite 9 with a pointed tip and their bifurcate aedeagal posterolateral arms. Males of D. crassiseta differ from this new species by their inner gonostylus that is as long as the outer gonostylus and the paramere is produced beyond the aedeagus. Males of D. fasciigera differ from this new species by their tergite 9 with a deeply notched distal margin and their single aedeagal posterolateral arms that are stout with blunt tips. Males of D. pollex differ from this new species by their trefoil-shaped tergite 9 and their bifurcate aedeagal posterolateral arms. Males of D. tristyla differ from this new species by their U-shaped inner gonostylus and their bifurcate aedeagal posterolateral arms, each of which is recurved ventrolaterally. It is likely that Dasyhelea pumila Macfie (1939), which is known only from Santa Catarina, Brazil, is also a member of the fasciigera complex because it has a bifurcate gonostylus. However, in this species the shape of the male aedeagus is simple and lacks posterolateral arms as in other species in the fasciigera complex, and, the spermatheca has an elongate oblique neck. Regardless, males of D. pumila differ from males of D. nelidae by their sternite 9 with a rounded posteromedian projection and their nearly straight paramere., Published as part of Grogan, Willliam L., D��az, Florentina, Spinelli, Gustavo R. & Ronderos, Maria M., 2016, The biting and predaceous midges of Guadeloupe (Diptera: Ceratopogonidae). II. Species of the subfamily Dasyheleinae, pp. 201-254 in Zootaxa 4184 (2) on pages 244-247, DOI: 10.11646/zootaxa.4184.2.1, http://zenodo.org/record/164718, {"references":["Remm, H. (1962) The genus Dasyhelea Kieffer in Estonia (Diptera, Heleidae). Tartu Riikliku Ulikodi Teimetised, 120, 108 - 131 [in Russian]; English version, 259 - 280.","Borkent, A. & Forster, L. (1986) Review of the Dasyhelea fasciigera species group (Diptera: Ceratopogonidae) with a revision of the Nearctic species. Canadian Journal of Zoology, 64, 1280 - 1287. http: // dx. doi. org / 10.1139 / z 86 - 190","Dominiak, P. (2012) Biting midges of the genus Dasyhelea Kieffer (Diptera: Ceratopogonidae) in Poland. Polish Journal of Entomology, 81, 211 - 304.","Dominiak, P. & Szadziewski, R. (2010) Distribution and new synonymy in European biting midges of the genus Dasyhelea Kieffer (Diptera: Ceratopogonidae). Zootaxa, 2437, 1 - 37.","Zilahi-Sebess, G. (1936) Die Heleiden-Fauna von Szeged und Umgebund. Acta Biologica Szeged Universitat, 4, 39 - 45.","Kieffer, J. J. (1925 b) Chironomiden der Hochmoore Nordeuropas und des ostlichen Mitteleuropas. Beitrage zur Kunde Estlands, 10, 145 - 163.","Remm, H. (1967) On the fauna of Ceratopogonidae (Diptera) in the Caucasus. Tartu Riikliku Ulikodi Teimetised, 194, 3 - 37. [in Russian, English summary]","Macfie, J. W. S. (1939) A report on a collection of Brazilian Ceratopogonidae. Revista de Entomologia, 10, 137 - 219."]}

Dasyhelea turnbowi new species (Figs. 29���30) Diagnosis. The only Neotropical species of Dasyhelea with the following combination of characters: Male with a heart-shaped apex of posteromedian projection of sternite 9 and an aedeagus with a hyaline membrane extending from the distal mesal protuberance of the basal arch to the tips of the posterolateral arms. Female with chaliceshaped subgenital plate and a very small spherical spermatheca with a moderately long, curved, oblique neck. Male. Head (Fig. 29 A). Dark brown. Eyes contiguous for a distance equal to the width of 5���6 ommatidia. Frontal sclerite broad, elliptical, apparently without lower marginal ear-like lobes. Antennal flagellum with flagellomeres 2���9 rhomboid, 10���13 elongate, 10���11 longest, 12 slightly shorter than 10���11, 13 slightly shorter than 12, slightly broader than 10���12 with tapered slender apex. Clypeus (Fig. 29 B) with 5 pairs of setae; ventral margin with small, narrow posterior notch. Palpus (Fig. 29 C) light brown, moderately long; segment 3 as long as 4+5 combined with scattered capitate sensilla on basomesal surface; palpal ratio 2.90. Thorax. Dark brown. Scutellum paler with 9 large, 6 smaller setae. Wing (Fig. 29 D) slender; membrane infuscated, covered with moderately dense macrotrichia; 2nd radial cell long, with very narrow lumen; apex of costa, radius at slight oblique angle with wing margin; fork of CuA1 just anterior of level of mid-portion of 2nd radial cell; anal lobe slightly developed; wing length 1.20 mm, width 0.38 mm; costal ratio 0.51. Abdomen. Dark brown. Genitalia (Figs. 29 E, 30A). Tergite 9 tapering progressively distally, with broad, rounded apex that extends to apices of gonocoxites; apicolateral process cylindrical, moderately slender with long stout apical seta; cercus near outer base of apicolateral process, reduced to a closely approximated group of 4 cylindrical lobes with apical setae. Sternite 9 (Figs. 29 E, 30B) 0.60 length of greatest width, anterior margin rounded; posterior portion with nearly straight posterolateral margins, distal portion sub-triangular with stout, heart-shaped posteromedian projection, tip pointed. Gonocoxite short, stout, 1.40 x longer than broad with short, pointed mesobasal process, distal 2/3 of medial margin with scattered long setae; gonostylus 1.10 x longer than gonocoxite, slender, nearly straight with narrow recurved apex and pointed tip. Gonocoxal apodemes and paramere (Fig. 30 C) forming a symmetrical structure; gonocoxal apodemes broad proximally, bases greatly recurved, with slender distal portions that are narrowly fused to lateral bases of paramere; paramere short, quadrate, lightly sclerotized with truncate tip. Aedeagus (Figs. 29 E, 30D) heavily sclerotized, 0.50 length of greatest width; basal arch extending 0.20 of total aedeagus length, anterior margin concave, distal margin convex with short, blunt, mesal protuberance; basal arm broad, apex recurved 75��, tip broadly pointed; posterolateral arms moderately long, apices slightly curved with blunt, ventrally recurved tip; a hyaline membrane extends from tips of posterolateral arms to distomesal protuberance of basal arch. Female. Head (Fig. 29 F). Dark brown. Eyes contiguous for a distance equal to the width of 5 ommatidia. Frontal sclerite (Fig. 29 G) elliptical, with lower marginal ear-like lobes (barely visible in photograph of allotype). Antennal flagellum dark brown; flagellomeres 2���8 longer than broad, vasiform, 9���12 elongate, vasiform, 13 elongate, slightly broader than 9���12 with greatly tapered apex; antennal ratio 0.89 (n=2). Clypeus (Fig. 29 G) with 4 pairs of setae. Palpus (Fig. 29 H) pale brown, slender, moderately long; segment 3 slender, as long as 4+5 combined with 3���4 capitate sensillae on proximal half of mesal surface; palpal ratio 3.00���3.70 (3.35, n=2). Thorax (Fig. 29 I). Brown. Scutum darker mesally; scutellum pale brown with 8 large, 3 smaller setae. Femora, tibiae pale brown, femorotibial joints dark; tarsomeres 5 infuscated; hind tibial comb with 6 spines; prothoracic tarsal ratio 2.07���2.25 (2.16, n=2), mesothoracic tarsal ratio 2.06���2.25 (2.15, n=2), metathoracic tarsal ratio 2.05���2.11 (2.08, n=2). Wing (Fig. 29 J) moderately broad; membrane hyaline with dense macrotrichia; 2nd radial cell elongate, reduced to a suture; fork of CuA1 at level just anterior to base of 2nd radial cell; anal lobe well developed; wing length 1.02���1.06 (1.04, n=2) mm, width 0.40���0.42 (0.41 n =2) mm; costal ratio 0.53���0.55 (0.54, n=2). Halter uniformly pale brown. Abdomen (Fig. 29 K). Dark brown. Subgenital plate chalice-shaped, posterolateral arms stout, slightly curved. Spermatheca very small, spherical, heavily sclerotized, diameter 42 ��m, neck moderately long, curved, oblique, length 12 ��m. Distribution. Guadeloupe. Type material. Holotype male labeled ��� Guadeloupe, Basse Terre, NE Pigeon, (16.14404�� N, 61.74977�� W), 23-V-2012, RH Turnbow, Blacklight trap ���; allotype female, 1 paratype female, same data as holotype except 18-V- 2012 (holotype, allotype deposited in FSCA; paratype female deposited in MLPA). Derivation of specific epithet. We are pleased to name this unusual new species after the collector, Robert H. Turnbow, in recognition of his important contributions to Coleoptera in the Caribbean Region. Discussion. The holotype male of this new species is very similar to males of D. meloae D��az & Spinelli, in D��az et al. (2014) and D. pailemanensis D��az & Spinelli, in D��az et al. (2014) from the steppes of Patagonia, by their rounded genitalia, the inner distal margin of gonocoxites with a patch of long, slender setae and symmetrical gonocoxal apodemes and paramere. However, males of D. meloae differ from the holotype male of this new species by their sternite 9 which has a very shallow caudomedian excavation, the gonocoxite has a heavily sclerotized, small anteromesal process that is doubly recurved, and broad, thin, doubly recurved parameres with a very short, slender posterior process. The holotype male of D. pailemanensis differs from this new species by its sternite 9 with a convex caudal margin, much thinner parameres with broadly rounded bases and a much shorter, slender posterior apex, and by the longer, mesally curved posterolateral arms of its aedeagus., Published as part of Grogan, Willliam L., D��az, Florentina, Spinelli, Gustavo R. & Ronderos, Maria M., 2016, The biting and predaceous midges of Guadeloupe (Diptera: Ceratopogonidae). II. Species of the subfamily Dasyheleinae, pp. 201-254 in Zootaxa 4184 (2) on pages 249-252, DOI: 10.11646/zootaxa.4184.2.1, http://zenodo.org/record/164718, {"references":["Diaz, F., Spinelli, G. R., Donato, M. & Ronderos, M. M. (2014) A taxonomic revision of the Patagonian species in the Dasyhelea mutabilis group with a phylogenetic analysis of New World species (Diptera: Ceratopogonidae). Journal of Natural History, 48, 2117 - 2175. http: // dx. doi. org / 10.1080 / 00222933.2014.909062"]}

Dasyhelea quadrifurca new species (Figs. 5–6) Diagnosis. The only Neotropical species of the grisea group with males having an aedeagus with a four-pronged fork-like distal portion, a broad heart-shaped frontal sclerite, and moderately long cylindrical apicolateral processes with 1–2 apical setae. Female unknown. Male. Head (Fig. 5 A). Dark brown. Eyes contiguous for a distance equal to the width of 3–4 ommatidia. Frontal sclerite (Fig. 5 B) broad, heart-shaped, with long, slender ventral projection that extends to dorsum of clypeus. Antennal flagellum dark brown; flagellomeres 2–9 rhomboid, 10–13 elongate, 13 longest; flagellomeres 2–10 with whorl of long sensilla chaetica that comprise the dense plume that extends to apex of flagellomere 12; antennal ratio 0.96. Clypeus with 11 pairs of setae. Palpus (Fig. 5 C) brown; segment 3 short, with 1–2 capitate sensilla on distolateral surface; palpal ratio 2.50. Thorax. Scutum dark brown; scutellum slightly paler with 9 large, 5 smaller setae. Legs brown, tarsi paler except tarsomeres 5 infuscated; hind tibial comb with 7 spines; prothoracic tarsal ratio 2.13, mesothoracic tarsal ratio 2.47, metathoracic tarsal ratio 2.16. Wing (Fig. 5 D) membrane hyaline, covered with dense macrotrichia; 2nd radial cell well developed; apices of costa, radius at slight oblique angle; fork of CuA1 anterior to level of base of 2nd radial cell; anal lobe well developed; wing length 1.22 mm, width 0.42 mm; costal ratio 0.50. Halter brownish. Abdomen. Dark brown. Genitalia (Figs. 5 E, 6A). Tergite 9 with broad, rounded caudal margin that extends beyond apices of gonocoxites; apicolateral process slender, elongate, with 1–2 moderately long apical setae; cercus conspicuous, located just anterior of base of apicolateral process, with 4–6 thin setae. Sternite 9 (Figs. 5 E, 6B) 0.33 length of greatest width, tapered abruptly distally, posteromedian margin narrow with shallow excavation. Gonocoxite stout, 1.43 x longer than greatest width, broadest at mid-portion with short, mesal process; gonostylus as long as gonocoxite, nearly straight, proximal 1/3 moderately broad, distal 2/3 slightly tapered, apex slightly broadened, bluntly rounded. Gonocoxal apodemes and paramere (Figs. 5 E, 6C) forming an asymmetrical structure; proximal portions of gonocoxal apodemes stout, recurved; right apodeme 0.60 length of left apodeme, broadly contacting paramere; left apodeme with slender, straight distal portion, tapering slightly distally to pointed tip that does not contact paramere; paramere slender basally, broader mesally, distal portion curved at base nearly 90°, tapering to pointed tip. Aedeagus (Figs. 5 E, 6D) nearly 0.80 length of greatest width; basal arch heavily sclerotized, extending 0.25 of total length; basal arms slender distally, apex recurved; posterolateral arms with inner straight, sub-parallel portions with rounded tips, and stout, straight outer portion with pointed tip that extend 3/4 length of inner posterolateral arm. Female. Unknown. Distribution. Guadeloupe. Type material. Holotype male labeled “ Guadeloupe, Basse Terre, Pointe a Lezard, 17-V-2012, R. H. Turnbow, BL trap ” (deposited in FSCA). Derivation of specific epithet. The specific epithet is from the Latin, quadri (four) and furca (fork), in reference to the four-pronged fork-like distal portion of the male aedeagus. Discussion. The holotype male of this new species is similar to males of three primarily Nearctic species: D. griseola Wirth, D. pseudoincisurata and D. stemlerae Waugh & Wirth (1976) in that its posteromedian margin of sternite 9 has a shallow excavation and aedeagal posterolateral arms bearing an outer process. However, males of D. griseola differ from this new species by their very short apicolateral processes, the hyaline outer process of the aedeagal posterolateral arms are mesally recurved, and their wing has a longer costa (costal ratio 0.57). Males of D. pseudoincisurata also differ from this new species by their straight distal margin of tergite 9 with longer greatly divergent apicolateral processes, outer aedeagal posterolateral arms have broader tips, and the tip of their gonostylus is usually notched. Finally, males of D. stemlerae differ from this new species by their tergite 9 with a much narrower truncate apex and the inner and outer processes of their aedeagal posterolateral arms are of nearly equal length.

Dasyhelea scalpela new species (Figs. 9���10) Diagnosis. The only Neotropical species of the grisea group with males having the apex of the paramere scalpedshaped. Females with fore, hind femora with mesal dark spot, fore tibia with mesal dark ring, the subgenital plate arrow-shaped and the spermatheca with neck elongate, stout and oblique. Male. Head (Fig. 9 A). Dark brown. Eyes broadly contiguous. Frontal sclerite heart-shaped. Antennal flagellum with flagellomeres 2���9 rhomboidal, 10���13 elongate, 13 longest; antennal ratio 0.83���0.94 (0.89, n=10). Palpus (Fig. 9 B) moderately short; segment 3 short with scattered capitate sensilla; palpal ratio 1.90���2.18 (1.98, n=10). Scutellum with 6 large, 2 smaller setae. Wing (Fig. 9 C) moderately slender; membrane hyaline, covered with moderately dense macrotrichia; 2nd radial cell short, well developed; apex of costa, radius perpendicular to wing margin; fork of CuA1 at level of anterior portion of 2nd radial cell; wing length 0.92���1.06 (0.96, n=10) mm, width 0.32���0.34 (0.33, n=10) mm; costal ratio 0.49���0.53 (0.50, n=10). Genitalia (Figs. 9 D, 10A). Tergite 9 tapering distally, apex truncate, extending beyond apices of gonocoxites; apicolateral process stout, conical with medium-size apical seta; cercus small with 3 setae, 1 medium-size, other 2 shorter. Sternite 9 (Figs. 9 D, 10B) 0.83 length of greatest width, with narrow deep posteromedian excavation. Gonocoxite stout, 1.48 x longer than greatest width, broadest mesally; gonostylus as long as gonocoxite, very slightly curved, apex bluntly pointed. Gonocoxal apodemes and paramere (Figs. 9 D, 10C) forming an asymmetrical structure; gonocoxal apodemes moderately slender, bases curved; left apodeme narrowly contacting paramere or separate, right broadly contacting paramere; paramere slender on basal 2/3, distal 1/3 broadened, apex scalped-shaped with recurved, pointed tip. Aedeagus (Figs. 9 D, 10D) with anterior margin nearly straight, 0.72 length of greatest width; basal arms stout, directed laterally; posterolateral arms slightly divergent, tapering distally, with blunt tips. Female. Head (Fig. 9 E). Brown. Eyes contiguous for a distance equal to width of 4 ommatidia. Frontal sclerite broad, bilobed with long, slender ventral projection. Antennal flagellum dark brown; flagellomeres 2���8 vasiform, 9���12 longer vasiform, 13 longest; antennal ratio 0.81���0.92 (0.87, n=5). Clypeus with 5 pairs of setae. Palpus (Fig. 9 F) moderately short, light brown; segment 3 short with 2 mesal capitate sensilla; palpal ratio 1.63���2.27 (2.03, n=5). Thorax. Scutum dark brown, humeral areas paler; scutellum slightly paler with 7 large, 4 smaller setae. Legs pale brown; fore, hind femora with mesal dark spot, fore tibia with mesal dark ring; femorotibial joints dark; tarsomeres 5 infuscated; hind tibial comb with 6 spines; prothoracic tarsal ratio 2.10���2.54 (2.31, =6), mesothoracic tarsal ratio 2.21���2.54 (2.36, =5), metathoracic tarsal ratio 2.14���2.66 (2.35, =5). Wing (Fig. 9 G) moderately broad; membrane hyaline with moderately dense macrotrichia; apex of costa, radius oblique; fork of CuA1 at level anterior to base of 2nd radial cell; wing length 0.84���1.02 (0.92, =6) mm, width 0.34���0.44 (0.39, n=6) mm; costal ratio 0.52��� 0.58 (0.54, n=6). Halter brown. Abdomen (Fig. 9 H). Brown. Subgenital plate with anterior portion somewhat arrowhead-shaped, elongate, posterior portion U-shaped, continuous with posterolateral arms; posterolateral arms heavily sclerotized, curved, connected distally by slender sclerite. Spermatheca ovoid, diameter 60 ��m, neck elongate, stout, oblique, measuring 46 ��m. Distribution. Guadeloupe. Type material. Holotype male, allotype female labeled ��� Guadeloupe, Basse Terre, Trace des Cretes (D-14), 22-V-2012, R. H. Turnbow, BL trap ��� (deposited in FSCA). Paratypes 20 males, 5 females, as follows: same data as holotype 16 males, 4 females; same data except NE Pigeon, (16.14404�� N, 61.74977�� W), 18-V-2012, 3 males; same data except 23-V-2012, 1 male, 1 female. Derivation of specific epithet. The species name refers to the unusual scalpel-shaped apex of the male paramere. Discussion. This new species is very similar to D. flavifrons, but in males of that species, the distal portion of the paramere is bent sharply ventrolaterad and tapers to a blunt point; and females differ by their more broadly elliptical frontal sclerite, yellow halter knob, the subgenital plate that is triangular anteriorly with a lateral posteriorly directed projection, and the neck of the spermatheca is long and straight., Published as part of Grogan, Willliam L., D��az, Florentina, Spinelli, Gustavo R. & Ronderos, Maria M., 2016, The biting and predaceous midges of Guadeloupe (Diptera: Ceratopogonidae). II. Species of the subfamily Dasyheleinae, pp. 201-254 in Zootaxa 4184 (2) on pages 219-222, DOI: 10.11646/zootaxa.4184.2.1, http://zenodo.org/record/164718

Grisea Group The grisea group is equivalent to the subgenus Dasyhelea (Dominiak 2012). Species are distinguished from all other groups/subgenera of Dasyhelea by their broad elliptical, bilobate or heart-shaped frontal sclerite, heavily sculptured antennal flagellomeres and elongate flagellomere 13 with tapered slender apex. Male tergite 9 usually with long apicolateral processes; sternite 9 relatively short with posterior margin concave, straight or convex; gonocoxal apodemes and paramere forming an asymmetrical structure; gonocoxite often with mesal hook or tubercle. Female subgenital plate usually with triangular, arrowhead-shaped or slender distal portion., Published as part of Grogan, Willliam L., D��az, Florentina, Spinelli, Gustavo R. & Ronderos, Maria M., 2016, The biting and predaceous midges of Guadeloupe (Diptera: Ceratopogonidae). II. Species of the subfamily Dasyheleinae, pp. 201-254 in Zootaxa 4184 (2) on page 208, DOI: 10.11646/zootaxa.4184.2.1, http://zenodo.org/record/164718, {"references":["Dominiak, P. (2012) Biting midges of the genus Dasyhelea Kieffer (Diptera: Ceratopogonidae) in Poland. Polish Journal of Entomology, 81, 211 - 304."]}

Dasyhelea pseudoincisurata Waugh & Wirth Dasyhelea pseudoincisurata Waugh & Wirth, 1976: 233 (Georgia; New Hampshire south to Florida); Borkent & Grogan 2009: 11 (in Nearctic catalog; distribution). Diagnosis. This is a large dark brown species, male wing length 1.10–1.50 mm, female wing length 1.20–1.47 mm; femora and tibiae usually with broad dark brown bands. Males with posterior margin of sternite 9 straight to slightly concave; tergite 9 tapered distally with straight posterior margin and elongate, cylindrical, divergent apicolateral processes; gonocoxite usually with short basomesal spur; gonostylus slightly curved, pubescent except apical portion, and tip usually notched. Female subgenital plate with slender, elongate anterior portion, posterior portion broader with curved lateral arms; and spermatheca subspherical with retort-shaped neck. Discussion. Waugh & Wirth (1976) described this species from the eastern United States based on specimens from Florida, Georgia (type locality), Maryland, New Hampshire, Pennsylvania and Virginia. In their Nearctic catalog, Borkent & Grogan (2009) also listed it from Utah and Ontario, Canada. We identified a male from Guadeloupe as this species that we compared with a male paratype from Georgia and several other males from Florida and Maryland. We provide the first record of this species in the Caribbean region from Guadeloupe. Material examined. Guadeloupe, Basse Terre, La Trace du Petit-Malendure, 21-V-2012, R. H. Turnbow, BL trap, 1 male. New Guadeloupe record.

Dasyhelea caribbeana Spinelli & Wirth (Figs. 1���2) Dasyhelea caribbeana Spinelli & Wirth, 1984: 596 (Jamaica; Haiti, Mexico, Panama); Borkent & Spinelli 2000: 25 (in New World catalog south of the USA; distribution); Borkent & Spinelli 2007: 60 (in Neotropical catalog; distribution). Diagnosis. A moderate size species, male wing length 0.98���1.06 mm, female wing length 0.99���1.18 mm. Males are distinguished from males of other Neotropical species in the cincta group by their elongate Y-shaped paramere with a long, slender posterior extension. Females differ from other Neotropical species in the cincta group by the combination of their large (68 x 48 ��m) elongate ovoid spermathecae with short, narrow necks; a short quadrangular subgenital plate; and, abdominal pleurae without dark streaks. Male. Head (Fig. 1 A). Dark brown. Eyes contiguous for a distance equal to the width of 2���3 ommatidia. Antennal flagellum brown; flagellomeres 2���3 globular, 4���9 rhomboid, 10���13 elongate; antennal ratio 0.94���1.06 (1.00, n=3). Frontal sclerite slender, elliptical. Clypeus with 3 pairs of setae. Palpus (Fig. 1 B) brown; segments 3���5 elongate; segment 3 moderately slender with 6 capitate sensillae on mesobasal half; palpal ratio 2.37���2.55 (2.45, n=3). Thorax (Fig. 1 C). Scutum dark brown; scutellum (Fig. 1 D) dark brown on narrow midportion and distal portions beyond setal bases, subcentral portions lighter brown, dorsum with 8 large, 2���3 smaller setae. Legs light brown, femorotibial joints dark brown, tarsomeres 5 slightly infuscated; hind tibial comb with 6 spines; prothoracic tarsal ratio 1.57���1.69 (1.64, n=3), mesothoracic tarsal ratio 1.58���1.62 (1.60, n=3), metathoracic tarsal ratio 1.35��� 1.44 (1.41, n=2). Wing (Fig. 1 E) membrane slightly infuscated, covered with dense long macrotrichia; fork of CuA1 at level of junction of radial cells; wing length 0.98���1.06 (1.01, n=3) mm, width 0.35 (n=3) mm; costal ratio 0.48���0.51 (0.50, n=3). Halter uniformly brown. Abdomen (Fig. 1 C) dark brown. Genitalia (Figs. 1 F, 2A). Tergite 9 tapering moderately distally to rounded apex, extending below apices of gonocoxites; apicolateral process very small, short, with single long apical seta; cercus very short, with 4���5 short setae. Sternite 9 (Figs. 1 F, 2B) 0.40 length of greatest width, posterior margin moderately concave with short, pointed ventrally directed apicolateral processes. Gonocoxite stout, 2 x longer than broad, with short rounded mesobasal lobe; gonostylus (Fig. 2 C) 0.56 length of gonocoxite, slender; proximal 2/3 nearly straight with short sub-basal tubercle, distal 1/3 curved, tapering distally to greatly curved apex, tip pointed. Gonocoxal apodemes and paramere (Figs. 1 F, 2D) narrowly separated, symmetrical; apodemes S-shaped, proximal portions slender, distal portions broad; proximal portion of paramere Y-shaped, apex bulbous, heavily sclerotized, distal portion very slender with pointed tip that extends just beyond apex of tergite 9. Aedeagus (Figs. 1 F, 2E) heavily sclerotized, 1.20 x longer than broad; basal arch extending 0.30 of total length; hyaline envelope extending to midlength of gonocoxite, with thick lateral margins, posterior margin straight; basal arms short, stout, recurved, mesally directed; posterolateral arms slender, extending just beyond posterior margin of hyaline envelope with a bulbous apex and a short, slender, recurved sub-apicolateral pointed process. Discussion. This Neotropical species has been recorded in the Caribbean region from Jamaica (type locality) and Haiti, and, also from Mexico and Panama (Spinelli & Wirth 1984; Borkent & Spinelli 2000, 2007). We provide the first records of D. caribbeana from Guadeloupe as well as the first description and illustrations of the previously unknown male. Material examined. Guadeloupe, Basse Terre, Pigeon, 9-IX-2011, MC Thomas & RH Turnbow, backlight trap, 1 male, 1 female; same data except NE Pigeon, 16.14404�� N, 61.74977�� W, 23-V-2012, RH Turnbow, 2 males, 1 female. New Guadeloupe record., Published as part of Grogan, Willliam L., D��az, Florentina, Spinelli, Gustavo R. & Ronderos, Maria M., 2016, The biting and predaceous midges of Guadeloupe (Diptera: Ceratopogonidae). II. Species of the subfamily Dasyheleinae, pp. 201-254 in Zootaxa 4184 (2) on pages 205-208, DOI: 10.11646/zootaxa.4184.2.1, http://zenodo.org/record/164718, {"references":["Spinelli, G. R. & Wirth, W. W. (1984) Las especies Neotropicales del genero Dasyhelea, grupo cincta (Diptera: Ceratopogonidae). Limnobios, 2, 586 - 608.","Borkent, A. & Spinelli, G. R. (2000) Catalog of the New World biting midges south of the United States of America (Diptera: Ceratopogonidae), Contributions on Entomology, International. 4, 1 - 107.","Borkent, A. & Spinelli, G. R. (2007) Neotropical Ceratopogonidae (Diptera: Insecta). In: Adis, J., Arias, R., Rueda-Delgado, G. & Wantzen, K. M. (Eds.), Aquatic Biodiversity in Latin America (ABLA). Vol. 4. Pensoft; Sofia-Moscow, 198 pp."]}

Dasyhelea waughi new species (Figs. 15–16) Diagnosis. The only Neotropical species of the leptobranchia group with males having an elongate triangular distal portion of sternite 9 with a truncate apex and an H-shaped aedeagus. Females with a uniformly dark brown halter, a triangular disjunct anterior portion of the subgenital plate and a small ovoid spermatheca with a short, stout neck. Male. Head (Fig. 15A). Dark brown. Eyes contiguous for a distance equal to the length of 3–4 ommatidia. Antennal flagellum with flagellomeres 2–9 rhomboid, 10–13 elongate, 11 longest; flagellomeres 2–10 with whorl of long sensilla chaetica comprising the dense plume that extends to midportion of flagellomere 13; antennal ratio 1.11. Palpus moderately short; segment 3 short with scattered captitate sensilla on anteromesal surface; palpal ratio not obtainable due to flattened, distorted segment 3. Thorax. Dark brown. Scutellum with 6 large setae. Prothoracic tarsal ratio 2.28, mesothoracic tarsal ratio 2.33, metathoracic tarsal ratio 2.33. Wing (Fig. 15B) moderately slender; membrane hyaline with sparse macrotrichia, devoid of macrotrichia below wing base and base of r-m crossvein; 2nd radial short with narrow lumen; apex of costa, radius perpendicular to wing margin; fork of CuA1 just beyond level of apex of costa, radius; anal lobe moderately well developed; wing length 0.74 mm, width 0.26 mm; costal ratio 0.50. Abdomen very dark brown. Genitalia (Figs. 15C, 16 A). Tergite 9 tapering abruptly distally, apex moderately narrow, rounded, extending 2/3 length of gonocoxites; apicolateral process slender, with large apical seta; cercus conspicuous, slender, located at base of apicolateral process with 4 moderately long setae. Sternite 9 (Figs. 15C, 16 B) 0.45 length of greatest width; distal portion elongate, triangular, tip truncate, folded ventrally; posterolateral margins irregular. Gonocoxite straight, 1.80 x longer than broad, broadest at mid-length; gonostylus 0.88 length of gonocoxite, proximal half broad, pilose, tapered distally, distal half slender, smooth, curved, apex narrowed, tip pointed. Gonocoxal apodemes and paramere (Figs. 15C, 16 C) forming an asymmetrical structure; proximal portions of apodemes broad, extreme bases recurved 60º; distal portions more slender with mesal rounded tubercle; right apodeme barely contacting pointed base of paramere; left apodeme broadly contacting paramere; proximal 2/3 of paramere moderately broad at mid-length, extreme base slender, slightly recurved, distal 1/3 bent sharply ventrolaterad, tapered distally, more or less leaf–shaped, tip pointed. Aedeagus (Figs. 15C, 16 D) heavily sclerotized, H-shaped, 0.80 length of greatest width; basal arch broadly concave, extending to 0.40 of total aedeagus length; basal arms slightly divergent; distal margin with several small median teeth; posterolateral arms stout, slightly curved, apices mesally directed, tips rounded. Female. Head (Fig. 15D). Dark brown. Eyes contiguous for a distance equal to the width of 3 ommatidia. Frontal sclerite (Fig. 15E) elongate, ellipsoid with lower marginal ear-like lobes. Antennal flagellum dark brown; flagellomeres 2–8 vasiform, progressively increasing in length, 9–12 more elongate vasiform, 13 longest, with slender tapered apex; antennal ratio 0.86. Clypeus (Fig. 15F) with 5 pairs of setae. Palpus (Fig. 15G) dark brown; segment 3 slender, moderately long with 5–6 capitate sensillae on basal half of mesal surface; palpal ratio 4.00. Thorax. Uniformly dark brown. Scutellum with 6 large setae. Femora, tibiae dark brown, tarsomeres 1–4 slightly paler, tarsomeres 5 infuscated; hind tibial comb with 6 spines; prothoracic tarsal ratio 2.13, mesothoracic tarsal ratio 2.43, metathoracic tarsal ratio 2.07. Wing (Fig. 15H) moderately broad; membrane infuscated, covered with dense macrotrichia; 2nd radial cell reduced to short suture; fork of CuA1 just beyond level of apex of costa, radius; anal lobe moderately well developed; wing length 0.86 mm, width 0.38 mm; costa short, costal ratio 0.39. Halter uniformly dark brown. Abdomen (Fig. 15I). Dark brown. Subgenital plate disjunct; anterior portion triangular; posterior portion somewhat rectangular with deep posterior arch; posterolateral arms slender, slightly curved. Spermatheca heavily sclerotized, slightly ovoid, length/width 46 x 40 µm, neck short, moderately broad, length 6 µm. Distribution. Guadeloupe. Type material. Holotype male, allotype female labeled “ Guadeloupe, Basse Terre, NE Pigeon, 16.14404º N, 61.74977º W, 23-V-2012, RH Turnbow, Backlight trap ” (deposited in FSCA). Derivation of specific epithet. This new species is dedicated to William T. Waugh, who shared an office with the senior author during graduate school at the University of Maryland, and in recognition of his important contributions to the taxonomy of Dasyhelea in North America (Waugh & Wirth 1976) and the Neotropics (Wirth & Waugh 1976). Discussion. Characters for distinguishing D. waughi n. sp. from D. juanae n. sp. are provided in the discussion section of the latter species. Dasyhelea waughi is also similar to D. messersmithi by its distal 1/3 of the paramere which is folded ventrolaterad and leaf-shaped, the single posterolateral arms of the aedeagus and the ovoid spermatheca with a straight neck. However, males of D. messersmithi differ from the holotype male of D. waughi by their convex sternite 9 and straight gonostylus. Whereas females of D. messersmithi differ from the allotype female of D. waughi by their yellow scutellum and the club-shaped anterior portion of their subgenital plate.

Dasyhelea patrycjae new species (Figs. 13–14) Diagnosis. The only Neotropical species of the leptobranchia group with males having an extremely elongate palpal segment 3 (palpal ratio 7.13), conical apicolateral processes, the posterolateral angles of posteromedian projection of the sternite 9 bifid and the posterolateral arms of the aedeagus single, stout and sub-triangular. Female unknown. Male. Head (Fig. 13 A). Dark brown. Eyes apparently contiguous for a distance equal to the width of 3 ommatidia. Frontal sclerite distorted and damaged during slide-mounting, shape not discernible. Antennal flagellum (Fig. 13 B) with flagellomeres 2–8 rhomboid, 10–12 elongate, slender, 13 shorter, broader than 10–12 with narrow pointed apex; flagellar plume lost during collection and preservation; antennal ratio 1.05. Clypeus (Fig. 13 C) with 5 pairs of setae. Palpus (Fig. 13 D) slender, elongate; segment 3 greatly elongate, 1.50 x longer than combined lengths of segments 4+5 with 2–3 capitate sensilla on mesobasal portion of distal half; palpal ratio 7.13. Thorax. Scutum dark brown; scutellum pale brown with 7 large, 2 smaller setae. Femora, tibiae brown, tarsi slightly paler except tarsomeres 5 infuscated; hind tibial comb with 6 spines; prothoracic tarsal ratio 1.94, mesothoracic tarsal ratio 1.84, metathoracic tarsal ratio not available. Wing (Fig. 13 E) membrane slightly infuscated, densely covered with macrotrichia; 2nd radial cell moderately well developed with slender lumen; apex of costa, radius at slight oblique angle; fork of CuA1 at level of apex of costa; wing length 0.96 mm, width 0.30 mm; costal ratio 0.48. Halter brown. Abdomen. Dark brown. Genitalia (Figs. 13 F, 14A). Tergite 9 distorted, due to slight twisting during slide-mounting, tapering abruptly distally with narrow, rounded apex; apicolateral process short, conical with stout apical seta; cercus small with 2 medium-size, 2 shorter setae. Sternite 9 (Figs. 13 F, 14B) 0.70 length of greatest width, anterior margin slightly curved; tapering distally, posteromedian margin with deep rounded excavation, posterolateral lobes bifid. Gonocoxite stout, twice as long as broad; gonostylus as long as gonocoxite, proximal 1/3 moderately broad at base, tapering gradually distally, distal 2/3 slender, nearly straight, apex slightly curved with sharply pointed tip. Paramere and gonocoxal apodemes (Figs. 13 F, 14C) forming an asymmetrical structure; gonocoxal apodemes broadest at bases, apices recurved anterad; right apodeme broadly connected to base of paramere, left apodeme not connected to paramere; paramere long, slender, proximal 2/3 moderately broad, nearly straight, distal 1/3 elongate, slender, abruptly recurved anteroventrally nearly 180º, apex tapering to very slender pointed tip. Aedeagus (Figs. 13 F, 14D) 0.60 length of greatest width; basal arch heavily sclerotized, slender with broad, shallow concave basal arch that extends 0.25 of total aedeagus length; basal arm very slender, apex slightly recurved; posterolateral arms single, sub-triangular, stout proximally, tapering gradually distally, more abruptly tapered at distal 1/4 with short apices that are slightly mesally directed. Female. Unknown. Type material. Holotype male labeled “ Guadeloupe, Basse Terre, Pointe a Lezard, 20-V-2012, RH Turnbow, BL trap ” (deposited in FSCA). Derivation of specific epithet. We dedicate this new species to our good friend and colleague, Patrycja Dominiak, in recognition of her important contributions on Dasyhelea in the Palearctic region. Discussion. This new species differs radically from all other males in the leptobranchia group by its greatly elongate, slender palpal segment 3 (palpal ratio 7.13). For example, the palpal ratios of the other three species in the leptobranchia group from Guadeloupe range from 2.13 for D. waughi n. sp.; 2.25–3.85 for D. thomasi n. sp.; and 3.37–4.85 for D. juanae n. sp. In addition, D. pabloi from Argentina and Chile, has a moderately elongate palpal segment 3 with a palpal ratio of 4.22–5.25, but, the wing has an extremely short costa (costal ratio 0.33–0.39 vs. 0.48 in D. patrycjae n. sp.), the aedeagus has much longer posterolateral arms, and the distal portion of the paramere is shorter with only the very short tip recurved. Dasyhelea waughi n. sp., is the only other Neotropical species with a paramere that has a greatly elongate, recurved distal portion, but, it is much broader than in D. patrycjae n. sp.

Dasyhelea guadeloupensis Delécolle & Rieb Dasyhelea guadeloupensis Delécolle & Rieb, 1994: 272 (Guadeloupe); Borkent & Spinelli 2000: 25 (in New World catalog south of the United States; distribution); Borkent & Spinelli 2007: 60 (in Neotropical catalog; distribution). Diagnosis. A small species that differs from other Neotropical species in the mutabilis group by the following combination of characters: Male with short palpal segment 3 (palpal ratio 3.00); sternite 9 with an elongate convex posteromedian projection with triangular posterolateral lobes; gonocoxite moderately long with proximal portion slender and distal portion with broad elongate subapical lobe; gonostylus long, slender, slightly curved with rounded apex; aedeagus heavily sclerotized with crescent-shaped basal arch and moderately long, curved, widely separated posterolateral arms with medially directed apices; and gonocoxal apodemes and paramere asymmetrical with rod-like paramere that extends just below apices of aedeagal posterolateral arms. Females with circular subgenital plate with very broad anterior portion and a small (length 44 µm) spherical spermatheca with a slender retort-shaped neck. Discussion. Delécolle & Rieb (1994) described this species from four males and three females collected by colored light trap during March 1993 from Petit Bourg, Guadeloupe. However, we did not find any specimens of this species in our material from Basse Terre, Guadeloupe.