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Taxonomic Revision of Neotropical Downeshelea Wirth and Grogan Predaceous Midges (Diptera: Ceratopogonidae)

The genus Downeshelea was described by Wirth and Grogan based on the diagnostic characters of the Monohelea multilineata species group. The first descriptions of species were based on body coloration, which resulted in confusion and misunderstanding of their identification. The aim of this study was to provide an updated diagnosis and description of Downeshelea, describe 18 new species, and redescribe 10 previously poorly described species. New records of species, a key for identification of all New World species, and a table with important morphometric data to distinguish both males and females of the various species are provided along with distribution maps of the 46 known New World species.


Introduction
The Ceratopogonidae are a diverse family of Diptera with 6207 species [1] distributed worldwide, of which 1282 are known from the Neotropical Region [2]. They are quite numerous in almost all aquatic and semi-aquatic habitats but, due to their small size, have still been inadequately collected and studied [3].
As a result of the study of the New World species of Monohelea s. lat., Kieffer by Wirth [4] and Wirth and Williams' [5] study of the Nearctic species, Lane and Wirth [6] divided this genus into groups based on characters from male genitalia and coloration patterns of wings and legs. Lane and Wirth [6] recognized four distinct species groups: fairchildi, hyeroglyphica, multilineata, and tesselata. In 1988, Wirth and Grogan [7] placed the species of the multilineata group into a new genus Downeshelea based on several features of both males and females. The authors designated Downeshelea stonei (Wirth) as type species and maintained its position in Ceratopogonini.
The females of species of Downeshelea are predaceous midges that feed primarily on adult Chironomidae [8]. Because of their feeding habit, they may be important in the biological control of pests, as Yasumatsu [9] indicated took place in rice plantations in Thailand.
Currently, 42 species are included in Downeshelea, with 28 recorded from the Neotropical region (with two of these extending into the Nearctic) [1,10]. With the addition of 18 new species here, there are now 60 species known. The early descriptions were based, primarily, on authors' interpretations of body coloration. This practice resulted in inconsistent species descriptions and misunderstandings Some specimens deposited in the FSP were pinned, which prevented further interpretation of some diagnostic characters. They could not be slide mounted, due to institutional policies of retaining the original preservation and the risk of damaging the ancient specimens fixed in celluloid strips during the mountage process.
Diagnostic characters were illustrated or photomicrographed using a Nikon Eclipse E 200 (Nikon, NY, USA) and a Zeiss AX10 microscope (Zeiss, Germany), except those from FSP that were taken with a cellphone attached to the microscope. Photomicrography and illustrations were prepared using Combine Z (www.combinezp.software.informer.com) and Photoshop GIMP Portable (www.gimp.org), respectively. The aedeagus and parameres of some species were drawn separately from the remainder of the male terminalia for better visualization. The distribution maps were created using QGis 2.18.19 (https://qgis.org) based on slide label coordinates or as approximate coordinates obtained in FallinGrain (www.fallingrain.com). The ISO Alpha03 code was used to identify the countries. Species distributions are presented on maps to better visualize their distribution, using an alphabetical arrangement, when possible. Morphological terms are from Borkent et al. [14]. The antennal ratio was obtained by dividing the combined lengths of the distal five flagellomeres by the combined lengths of the preceding eight flagellomeres for females and the distal four by the preceding nine flagellomeres for males, as was done in our earlier species description of Downeshelea [10,[15][16][17][18][19][20][21]. The distribution of spines on each of tarsomeres 2-4 of all legs are separated by hyphens. Measurement of spermathecae are given in micrometers and those of the wings in millimeters. Meristic information is given as range, following by the mean and number of specimens examined. A table including the measurements important for identification of all Neotropical species is provided. Some females can be diagnosed and are distinctive while others can only be identified when associated with their respective males. The features described in the key are all figured when possible. When the feature is present in several species, we selected only one to represent it.
Neary all leg figures were those of males and, due to the similarity between both sexes, the figures from Downeshelea lanei Felippe-Bauer and Borkent  Diagnosis. Male: only species group of Downeshelea in the Americas with the following combination of character states: wing with three grayish areas: first near apex of M 1 , second sigmoid-shaped extending from m 1 to wing margin in m 2 , third over CuA 2 , (also in CuA 1 in some species); legs brown or pale brown, without bands; apical spines of hind tarsomeres 2-4: 1-1-1; distal portion of paramere abruptly bent ventrally, directed anteromesally, tapering to pointed tip ( Figure  6b), overlapping in some species. Female: only species group of Downeshelea in the Americas with the following combination of character states: wing and legs as described for male; antenna brown, basal portion of flagellomeres 1-7 pale.
Distribution and bionomics. This species is known only from coastal areas in Panama occurring at sea level ( Figure 2).
Taxonomic discussion. This species is included in the multilineata species group because of its wing and leg pigmentation patterns and having a paramere with its distal portion long and abruptly bent ventrally, directed anteromesally. It most closely resembles D. moravia in leg pigmentation pattern and general aspect of parameres. Downeshelea blantoni can be distinguished from D. moravia by its smaller size (1.37 mm in D. moravia) (Table 1), the aedeagus rectangular without elliptical sclerotized anterior areas and distal portion non-serrate (subtriangular with elliptical sclerotized anterior areas, distal portion serrate in D. moravia) and sternite 9 with a large convex median lobe (poorly developed in D. moravia). Specimens of Downeshelea blantoni are only recorded from coastal areas in Panama at lower altitudes while D. moravia is restricted to higher altitudes (1600 m) in forested areas of Costa Rica [10] ( Figure 2). Unfortunately, only one female wing was available for study from the paratypes listed herein. We therefore cannot make further comparisons with females of other species. The holotype and allotype of D. blantoni are pinned specimens (USNM #66436) and no further observations were made of them. Distribution. This species is known from forested, coastal, and mangrove areas in Costa Rica (Alajuela, Heredia, Puntarenas, San José, Cartago, and Limón), Colombia and Brazil (Pará) ( Figure  2). It occurs in Costa Rica up to 1850 m above sea level. In other countries it has been found up to 15 m above sea level.  Distribution. This species is known from forested, coastal, and mangrove areas in Costa Rica (Alajuela, Heredia, Puntarenas, San José, Cartago, and Limón), Colombia and Brazil (Pará) (Figure 2). It occurs in Costa Rica up to 1850 m above sea level. In other countries it has been found up to 15 m above sea level. Distribution. This species is known from Mexico (Yucatan), Bahamas, Grand Cayman, Jamaica, Dominica, Costa Rica (Limón) and Brazil (Rio de Janeiro) ( Figure 2). It has been found in forests near coastal areas, occurring up to 35 m above sea level. Distribution. This species is known only from coastal areas in Brazil (Rio de Janeiro) occurring at sea level ( Figure 2).
Distribution. This species is known only from forested areas in Costa Rica (San José) at 1600 m above sea level ( Figure 2).
Etymology. This species name refers to its similarity to D. stonei but being another species (Latin-alia = other).
Distribution and bionomics. This species is known only from the Brazilian state of Pará. It has been found in forested areas ( Figure 10). The locality of Tracuateua is up to 20 m above sea level.
Etymology. This species name refers to Gregório Aviz, owner of the property where it was collected. Taxonomic discussion. Downeshelea avizi most closely resembles D. chiapasi, D. colombiae, D. gladius, and D. spatha, in the pattern of dark bands on the hind legs and by the absence of a grayish spot in the distal portion of r 3 . The females of D. avizi can be easily distinguished by the pale CuA 2 (CuA 2 with grayish spot in the other aforementioned species), presence of basal dark band on hind femur (absent in D. chiapasi, D. colombiae, and D. gladius), the unequal spermathecae (equal-sized in D. chiapasi, subequal in D. spatha) and by the long hind leg claw (shorter in the other species). The male can be distinguished by the fused parameres (parameres separated in D. chiapasi), apical process of the paramere straight (nearly straight and subapical in D. chiapasi, deeply curved and subapical in D. colombiae, sinuous and apical in D. gladius) and the paramere longer than aedeagus (nearly similar in length D. colombiae, D. gladius, and D. spatha).   Figure 5a) with three grayish areas: first near apex of M 1 , not reaching wing margin; second extending from m 1 to wing margin in m 2 ; third over CuA 1 , CuA 2 extending from mediocubital fork into cua 1, anal cell, reaching wing margin; 2nd radial cell 1.8 length of 1st; length 1.22 mm; width 0.40 mm; costal ratio 0.73. Halter brown. Legs (Figure 5c) brown, hind leg darker; hind tibial comb with 7 spines. Foretarsomere 1 with one basal, one apical spine; midtarsomere 1 with three ventral spines; apical spines of tarsomeres 2-4 of fore-, mid-, hind legs: 2-2-3, 2-2-3, 1-1-2; foretarsal ratio 2.32, midtarsal ratio 2.48, hind tarsal ratio 2.25; claws 0.38 length of their respective tarsomere 5. Abdomen. Brown. Terminalia (Figure 6g): tergite 9 with quadrate apex, apicolateral process long, slender; sternite 9 slightly concave anteriorly, posterior margin with moderately convex median lobe bearing two long setae. Gonocoxite brown, moderately stout, 2.03 times longer than basal width; gonostylus brown, curved, broad basally, 0.71 length of gonocoxite. Parameres (Figure 6h) 1.00 length of aedeagus, fused on basal portion for 0.19 of total length, each with basal arm trilobed, knob slender; stem expanded distally, subapical portion ( Figure 6h) with long hyaline process bent, slender basally, expanded on midportion, abruptly tapering to long pointed tip, posteromesally directed, 0.48 of total length. Aedeagus (Figure 6g) subtriangular, expanded mesally, tapering distally; basal arch U-shaped, heavily sclerotized, extending to 0.25 of total length; distal portion with deep mesal notch and two sclerotized pointed processes. like processes equally sized; aedeagus rectangular, distal portion with two longs, narrow sclerotized lobes (Figure 9e). Female unknown.
Distribution. This species is known only from Brazil (Rio de Janeiro) occurring in a humid area at 60 m above sea level ( Figure 10).    Distribution and bionomics. This species is known only from Brazilian state of Bahia ( Figure 10). It has been found in forested areas up to 150 m above sea level.
Etymology. This species name refers to Bahia, the Brazilian state where it was collected. Taxonomic discussion. The shape of the aedeagus of this species is unique in the genus. The wing and leg patterns resemble those in the multilineata species group, but in D. bahiana the paramere has a subapical process posteriorly directed, not a distal one anteriorly directed as in the multilineata group.   Distribution and bionomics. This species is known only from coastal areas in Panama ( Figure  10), occurring up to 30 m above sea level.
Taxonomic Discussion. This species has a relatively simple male genitalia that readily distinguishes it from other species in the genus. Downeshelea carioca, D. fluminensis and D. quasidentica also have legs without bands and separate parameres, but in these three species the CuA 1 is grayish. Downeshelea balboa has a rectangular aedeagus (triangular in D. fluminensis and D. carioca, quadrangular in D. quasidentica) and simple paramere (paramere with median processes in D. fluminensis and D. carioca). We studied the three paratypes deposited in Faculdade de Saúde Pública de São Paulo. The holotype and allotype of D. balboa are pinned specimens in the USNM (#66441) and no further observations were made of them. Some slide mounted specimens from Belize, El Salvador and Honduras, previously labeled as D. balboa by Wirth, are here identified as D. fluminensis.
Distribution and bionomics. This species is known from Costa Rica (Alajuela) and Colombia ( Figure 10). It has been found in coastal areas, occurring in Colombia at sea level and in Costa Rica from 50 to 100 m above sea level.
Etymology. This species name refers to the bifid distal portion of paramere of the male (Latin-bifida = bifid).
Distribution and bionomics. This species is known from Belize and Costa Rica (Guanacaste) ( Figure 10). It has been found in coastal and forested areas, occurring from 5 m (Belize) to 40 m (Costa Rica) above sea level.
Etymology. This species name refers to the similarity of the distal portion of the paramere to the horn of a goat (Capra).  Figure 14d-f, and Figure 21; Table 1 Figure 11f).
Distribution and bionomics. This species is known only from Brazil (Rio de Janeiro) in forested areas occurring from 20 to 300 m above sea level ( Figure 10).
Taxonomic discussion. The male of D. carioca is distinctive within the genus, with male genitalia which has parameres separated, with sinuous stem flattened apically. Its coloration resembles those in multilineata species group, but D. carioca has a triangular aedeagus and paramere with a median process as in D. fluminensis. It can be separated from this last species by the number of ventral spines on midtarsomere 1 (3-5 in D. carioca; 1-2 in D. fluminensis), the median process of the paramere not beak-shaped and the absence of median horn-like process on the aedeagus.  Figure 14d-f, and Figure 21; Table 1.   Diagnosis. Male: only species of Downeshelea in the Americas with the following combination of character states: r 3 with apical grayish spot ( Figure 12a); legs brown, fore-, midtibia darker apically, hind femur darker subapically ( Figure 12b Figure 12f).
Distribution and bionomics. This species is known only from Brazil (Rio de Janeiro) in a forested area at 300 m above sea level ( Figure 21).    Diagnosis. Male: only species of Downeshelea in the Americas with the following combination of character states: r 3 with apical grayish spot; legs pale brown, hind femur darker subapically, hind tibia darker subbasally (Figure 13c Distribution. This species is known from coastal and forested humid areas in Costa Rica (Puntarenas, Cartago and Limón) and Panama. It occurs from 15 m in Panamá to 1450 m above sea level in Costa Rica ( Figure 21).
Diagnosis. Male: only species of Downeshelea in the Americas with the following combination of character states: r 3 with apical grayish spot ( Figure 13f); legs pale brown, mid-, hind femur darker subapically, hind tibia darker subbasally (Figure 13h), hind tibial comb with 8 spines; male parameres separated, straight, mesally curved apically ( Figure 14l); aedeagus Y-shaped, distal portion with blunt apex (Figure 14k). Female: only species of Downeshelea in the Americas with the following combination of character states: wing and legs as described for male; hind leg claw 1.40 as long as tarsomere 5; unequal spermathecae ( Figure 13g); 3rd rudimentary spermatheca long (Figure 13g).
Distribution. This species is known only from Uruguay at 50 m above sea level ( Figure 21).
Diagnosis. Male: only species of Downeshelea in the Americas with the following combination of character states: r 3 without apical grayish spot; legs yellowish brown, hind femur with subapical, hind tibia with subbasal, apical dark band ( Figure 15a Redescription. Male (Figure 15a). Thorax. Yellowish brown; scutum with inconspicuous brown spots. Wing (as female in Figure 15c) with four grayish areas: first, small, near apex of M 1 (absent in some specimens); second L-shaped, extending from m 1 to wing margin in m 2 ; third, small, in cua 1 near CuA 1 , not reaching wing margin; fourth in distal 1 2 of CuA 2 extending into cua 1, anal cell, not reaching wing margin. Halter knob darkened. Legs (as female in Figure 15d) yellowish brown, hind femur with subapical, hind tibia with subbasal, apical dark band. Abdomen. Yellowish with ventrolateral brown marks on segments 1-7. Terminalia (Figure 19a): tergite 9 with quadrate apex, apicolateral process short; sternite 9 slightly concave anteriorly, posterior margin with moderately convex median lobe bearing 3-4 long setae. Gonocoxite brown, moderately stout, 2.00 (n = 3) times longer than basal width; gonostylus yellowish, distal 1  Distribution and bionomics. This species is known from Nicaragua, Costa Rica (Puntarenas), and Panama ( Figure 21). It has been found in coastal and humid forested areas, occurring at sea level in Panama, from 100 to 200 m in Costa Rica and in Nicaragua at 300 m above sea level.
Taxonomic discussion. We have studied five paratypes from Nicaragua and Panama indicated by Lane and Wirth [6] as deposited in FSP. Some data on the specimen labels did not match that cited in the original description and which is corrected here. The male holotype and female allotype from Nicaragua are pinned specimens (#66348 USNM) and no further observations were made of them. One male paratype from Colombia (IV.1963) was studied and is here described as the new species D. spatha. Other specimens from Colombia, labeled by Wirth as D. chiapasi, are also misidentified and belong to the new species D. gladius and D. spatha. As such, D. chiapasi is no longer recorded from Colombia. Downeshelea chiapasi is easily distinguished from other similar species by the strong dark bands on the hind leg (weak in D. gladius) and the absence of basal dark band on the hind femur (present in D. spatha). Downeshelea chiapasi also has wing and legs pattern similar to D. avizi and D. colombiae. Characters for distinguishing these five related species are in the discussion section of D. avizi.  Redescription. Male. Head (Figure 17c). Eyes narrowly contiguous in lower portion ( Figure  17c). Antenna pale, except basal portion of flagellomere 1, distal portion of flagellomere 10, flagellomeres 11-13 brown; antennal ratio 0.99-1.05 (1.02, n = 9). Palpus with segment 3 slightly swollen on midportion with broad, deep sensory pit; palpal ratio 2.14-2.60 (2.30, n = 8). Thorax. Scutum yellowish brown, without definite pattern in slide mounted specimens; pleura pale brown. Wing (Figure 17a) with three grayish areas: first near apex of M1 not abutting wing margin; second extending from m1 to wing margin in m2; third over CuA1, CuA2 extending into cua1, anal cell, reaching wing margin (Figure 19a   Distribution and bionomics. This species is known from Mexico (Veracruz), Nicaragua (?), Costa Rica (Alajuela, Heredia, Puntarenas, San José, Cartago and Limón) and Panamá in coastal and humid forested areas (Figure 21). It has been found at sea level in Panama, from 20 to 1210 m in Costa Rica and from 300 to 400 m above sea level in Mexico and Nicaragua.
Taxonomic discussion. The analysis of the type series deposited in the FSP and USNM collections showed the presence of pale and dark morphotypes. The holotype of D. chirusi (#66439 USNM) is represented by the pale form and the allotype (USNM) by the dark one that we describe below as D. pulla sp. nov. Both species are also found in male and female paratypes from Panama. Those from Nicaragua (1 male, 1 female) were not located in any of the studied collections but have been described as yellowish, suggesting they may indeed be D. chirusi. We present further comments in the taxonomic discussion of D. pulla sp. nov. below.

Distribution and bionomics.
This species is known only from Colombia ( Figure 21). It has been found in mangrove areas.
Taxonomic discussion. Downeshelea colombiae is similar to D. avizi, D. chiapasi, D. gladius and D. spatha in having a pattern of grayish spots on the wing and legs with dark bands. Characters for distinguishing these species are in the discussion section of D. avizi. We could not study the holotype of D. colombiae (#67563 USNM), which is a slide mounted male specimen. However, we have studied here some male and female specimens from the type locality, designated as D. colombiae by Wirth, that match its description. Based on this material, we redescribe the male and describe the females for the first time.   Figure 19h,i, and Figure 21; Table 1.

Diagnosis.
Male: only species of Downeshelea in the Americas with the following combination of character states: r 3 with apical grayish spot ( Figure 18a); legs brown ( Figure 18c); tergite 9 short ( Figure 19h); parameres fused on basal portion by pointed membrane (Figure 19i), stem with hook-like median process (Figure 19i) and apex curved externally (Figure 19i). Female: only species of Downeshelea in the Americas with the following combination of character states: wing (Figure 18d) and legs as described for male; hind leg claw 1.14-1.31 as long as tarsomere 5 ( Figure 18g); unequal spermathecae ( Figure 18f).
Description. Male. Head (Figure 18b). Eyes separated dorsomedially by 2× width of an ommatidium (Figure 18b Distribution and bionomics. This species is known from forested areas in Costa Rica (Alajuela and Puntarenas), occurring from 80 to 745 m above the sea (Figure 21).
Etymology. This species name refers to its short ninth tergite. (Latin-curta = short). Taxonomic discussion. This species closely resembles D. jurgeni in having a wing pattern with grayish spots and a dark brown body. It can be distinguished by its smaller wing length of 0.85-0.90 mm (1.10-1.22 mm in D. jurgeni). Female specimens can be also distinguished by the mandible with 7-8 teeth (11 teeth in D. jurgeni) and unequal sized spermathecae (equal-sized in D. jurgeni). The male genitalia of both species can be easily distinguished by the form of the median process of the paramere (long hook-like process in D. curta; short, pointed process in D. jurgeni), the broad distal portion of paramere (slender in D. jurgeni) and the gonostylus slightly curved at midlength (straight in D. jurgeni).     Distribution. This species is known only from Trinidad (Figure 21), occurring in coastal and humid forested areas from sea level to 60 m above sea level.

Downeshelea deanei Felippe-Bauer and Quintelas
Taxonomic discussion. This species was described by Felippe-Bauer et al. [18] based on a previous misidentification of a male specimen of D. guianae (Wirth) [6]. In that paper, the authors described a single male specimen from Trinidad and did not consider the description of the female by Lane and Wirth [6], because they did not have in hand the female specimens to confirm the identification. Herein, we describe the female specimens from Trinidad (Port of Spain, Nariva Swamp and Vega de Oropouche) as D. deanei, previously misidentified as D. guianae by Lane and Wirth [6]. The female from D. deanei can be easily distinguished from D. guianae by the presence of ventral spines on midtarsomere 1 (absent in D. guianae).    Figure 24i,j, and Figure 29; Table 1   Description. Male. Head (Figure 22b). Eyes separated dorsomedially by 2.5× width of an ommatidium (not visible in Figure 22b). Antenna brown; antennal ratio 1.14. Palpus with segment 3 slightly swollen on midportion with broad, deep sensory pit; palpal ratio 2.40. Thorax. Scutum brown, postscutellum, pleura pale brown. Wing (Figure 22a) with three grayish area: first near apex of M 1 , not reaching wing margin; second, sigmoid-shaped, extending from m 1 to wing margin in m 2 ; third over CuA 1 , CuA 2 extending into cua 1, anal cell, reaching wing margin; 2nd radial cell 2.6 length of 1st; length 1.15 mm; width 0.40 mm; costal ratio 0.76. Halter brown. Legs (Figure 22c) brown, hind leg darker, all femora slightly darker subapically; hind tibial comb with seven spines. Foretarsomere 1 with one basal, one apical spine; midtarsomere 1 without ventral spines; apical spines of tarsomeres 2-4 of fore-, mid-, hind legs: 2-2-2, 2-2-2, 1-1-1; foretarsal ratio 2.68, midtarsal ratio 2.65, hind tarsal ratio 2.23; claws 0.42 length of their respective tarsomere 5. Abdomen. Brown. Terminalia ( Figure  24d): tergite 9 nearly triangular, abruptly tapering distally, with quadrate apex, apicolateral process long, broad; sternite 9 straight anteriorly, posterior margin with prominent median lobe bearing two long setae. Gonocoxite brown, slender, 2.74 times longer than basal width; gonostylus brown, nearly straight, 0.71 length of gonocoxite. Parameres (Figure 24e Distribution and bionomics. This species is known only from the Brazilian state of Amazonas ( Figure 29). It has been found in river environments up to 20 m above sea level.
Etymology. This species name refers to the divergent aspect of distal half of its paramere. (Latin-divergentis = divergent).
Taxonomic discussion. The divergent paramere with bifid distal portion is similar to that of males of D. bifida and D. quechua. Characters for distinguishing these three species are in the discussion section of D. bifida. Distribution. This species is known from Mexico (Oaxaca, Veracruz), Belize, El Salvador, Honduras and Brazil (Mato Grosso do Sul, Espírito Santo, Rio de Janeiro) ( Figure 29). It has been found in humid areas occurring from 10 m in Brazil (Rio de Janeiro) to 1100 m in Mexico (Oaxaca).

Downeshelea fluminensis Felippe-Bauer and Quintelas
Remarks. Santarém et al. [10] studied material previously identified as D. multilineata deposited in FSP and concluded that the Brazilian specimen from Mato Grosso was misidentified and is actually a specimen of D. fluminensis. The specimens from Belize, El Salvador and Honduras were originally labeled by Wirth as D. balboa. Characters for distinguishing both species and other related species are in the discussion section of D. balboa.
Taxonomic discussion. This species was described from Colombia based on only one female specimen (holotype #67564 USNM). In addition, other male and female specimens from Colombia were labeled by Wirth as D. fuscipennis. Considering that these specimens match the original description, we describe herein the first male of this species and redescribe the female based on these and other specimens from Brazil. The male genitalia of Downeshelea fuscipennis most closely resembles that of D. rodriguezi, but can be distinguished by the paramere with slender median horn-like process (broad horn-like process in D. rodriguezi), by the distal portion greatly bent externally (bent externally to a short, sharp point in D. rodriguezi). These two species also have body coloration similar to that of D. kuna and D. wirthiana, but these two species do not have a paramere with a median process. Also, the distal portion of paramere in D. kuna is a short, slender, spiral, anteriorly directed and in D. wirthiana it is very long, slender with flattened point, and directed posteriorly.    Distribution and bionomics. This species is known from Costa Rica (Alajuela, Puntarenas, Cartago and Limón) and Colombia ( Figure 29). It has been found in forested and coastal areas occurring at sea level in Colombia and in Costa Rica from 15 m to 860 m above sea level.
Etymology. This species name refers to the similarity of its distal portion of the paramere to a sword that was an ancient Roman weapon (Latin-gladius = sword).
Taxonomic discussion. Downeshelea gladius most closely resembles D. spatha in the yellowish pattern of the legs with dark bands, the wing pattern of grayish spots, male terminalia and distal portion of paramere similar to a sword. It can be distinguished by the pale base of the hind femur (brown in D. spatha), the gonostylus yellowish with distal 1 2 brown (entirely brown in D. spatha), and parameres anteromedial fusion rounded basally and distal portion more sinuous and slender (anteromedial fusion and distal portion nearly straight in D. spatha). Characters for distinguishing both species from other species with similar wing and dark bands on the hind legs pattern are in the discussion section of D. avizi.   Distribution. This species is known from Trinidad, Guyana and Brazil (Pará) (Figure 29) in coastal and humid forested areas. It has been found at sea level in Trinidad, at 120 m in Guyana and from 5 to 300 m above sea level in Brazil.
Etymology. This species name refers to the grandchild of Jorge Arturo Lizano, who so generously allowed this species to be collected as part of a detailed inventory of the Diptera on his property in Costa Rica [36,37].
Etymology. This species name refers to the Kuna, Amerindian people from the Choco Department, Colombia, where it was collected.
Taxonomic discussion. The pattern of wing and leg coloration in D. kuna is similar to that of D. fuscipennis, D. rodriguezi sp. nov. and D. wirthiana sp. nov. Characters for distinguishing these species are in the discussion section of D. fuscipennis.  Figure 33g,h, and Figure 39; Table 1.
Etymology. This species name refers to its large size, a feature that is unique in Downeshelea. (Latin-magna = great).
Distribution. This species is known only from Brazil (Pará and Rondônia) in humid forested areas, occurring from 15 to 350 m above sea level (Figure 39).

Redescription. Male. Head
Etymology. This species name refers to its dark body coloration that differs from the similar but yellowish species D. chirusi. (Latin-pulla = dark).
Taxonomic discussion. This species is similar to D. chirusi by virtue of the pattern of dark spots on the wing, leg coloration and the general aspect of male genitalia. However, D. chirusi has a yellowish terminalia, only with distal halves of gonocoxite and gonostylus dark and pale spermathecae, while D. pulla sp. nov. has a uniformly dark brown terminalia and dark spermathecae. In both species, the male paramere are expanded medially, but in D. chirusi they are broader than in D. pulla. Furthermore, there are slight variations of some meristic data as follows: male antennal ratio, length/width of gonocoxite and female hind claws (Table 1) (Figure 36a) with five grayish area: three in apical portion of r3, m1, M1, reaching wing margin in m1; one sigmoid-shaped extending from m1 to wing margin in m2; one over CuA1, CuA2 extending from mediocubital fork into cua1, anal cell, reaching wing margin in CuA2; 2nd radial cell twice length of 1st; length 0.90-0.97 (0.94, n = 2) mm; width 0.32-0.35 (0.34, n = 2) mm; costal ratio 0.72 (n = 2). Halter dark brown. Legs (Figure 36c) brown, hind leg darker, hind tibia slightly darker subbasally, apically; hind tibial comb with six spines. Foretarsomere 1 with one basal, one apical spine; midtarsomere 1 with 1-2 ventral spines; apical  Distribution and bionomics. This species is known only from Bolivia ( Figure 39) in mature rainforest at 484 m above sea level.
Etymology. This species name refers to the Quechua language that is spoken by indigenous Quechua peoples in Bolivia.

Description. Male. Head
Distribution and bionomics. This species is known only from Bolivia ( Figure 39). It has been found in secondary rainforest at 517 m above sea level. One male (uncertain whether holotype or paratype) had a pollinium attached to its body, indicating the possibility that this species may pollinate a species of orchid.
Etymology. This species name refers to Dr. Jaime Ivan Rodriguez Fernandez in recognition of his entomological contributions, particularly in Bolivia.
Taxonomic discussion. This species most closely resembles D. fuscipennis in the general body coloration, wing pattern and male paramere with a median horn-like process. Characters for distinguishing both species are in the discussion section of that species.   Figure 43c-e, and Figure 46; Table 1 Diagnosis. Male: only species of Downeshelea in the Americas with the following combination of character states: r3 without apical grayish spot (Figure 40a); legs brown, fore-, midtibia darker  we maintain the records of D. stonei for these countries with a question mark. The discussion about similarities and differences between D. stonei and D. alia are in the taxonomic discussion of the new species. The male specimen from Trinidad identified by W.W. Wirth [4] had two labels: the first one identifying it as D. stonei, the second, with the same handwriting, as D. guianae. The specimen is actually D. guianae. We presume that Wirth had corrected the misidentification but had not published it.    (Figure 40c). Eyes separate (unable to measure; not visible in Figure 40c). Antenna pale, except basal portion of flagellomere 1, distal portion of 10, flagellomeres 11-13 brown; antennal ratio 1.02 (n = 1) (Figure). Palpus with segment 3 slightly swollen on mid portion with palpal pit not visible in slide mounted specimen; palpal ratio 2.60 (n = 1). Thorax. Scutum brown, without definite pattern in slide mounted and pinned specimens; pleura pale brown. Wing (Figure 40a) with two grayish areas: one sigmoid-shaped extending from m 1 to wing margin in m 2 ; one over CuA 2 extending into cua 1 , anal cell, from mediocubital fork to wing margin; faint grayish area in M 1 not reaching wing margin in some specimens; 2nd radial cell twice length of 1st; length 1.38 (n = 1) mm, width 0.48 (n = 1) mm; costal ratio 0.74 (n = 1). Halter pale, distal portion of knob darker. Legs (as female in Figure 40g) brown, hind leg darker, fore-, midtibia darker apically, hind femur darker subapically, hind tibia darker subbasally; hind tibial comb with six spines; foretarsomere 1 with one basal, one apical spine; midtarsomere 1 with two ventral spines; apical spines of tarsomeres 2-4 of fore-, mid legs 2-2-3, 2-2-3; apical spines of tarsomeres 2-3 of hind leg 1-1, tarsomere 4 absent; fore-, mid-, hind tarsal ratios 2.11 (n = 1), 2.35 (n = 1), 2.42 (n = 1); claws 0.40 (n = 1) length of their respective tarsomere 5. Abdomen. Brown. Terminalia (Figure 43c): tergite 9 with truncated apex, apicolateral processes short (not illustrated); sternite 9 with anterior margin not visible in slide mounted specimens, posterior margin with conspicuous convex median lobe bearing two long setae. Gonocoxite brown, stout, 1.  [4] had two labels: the first one identifying it as D. stonei, the second, with the same handwriting, as D. guianae. The specimen is actually D. guianae. We presume that Wirth had corrected the misidentification but had not published it.

Description. Male. Head
Etymology. This species name refers to Dr. Willis Wirth, for his significant contribution to the taxonomic studies of Worldwide Ceratopogonidae fauna.
Taxonomic discussion.Downeshelea wirthiana closely resembles D. fuscipennis, D. kuna sp. nov. and D. rodriguezi sp. nov. in having a wing pattern with grayish spots and a dark brown body. Characters for distinguishing these species are in the discussion section of D. fuscipennis. D. wirthiana also resembles D. cebacoi in aspects of the male genitalia, but it differs from this species by the dark brown legs and the very long distal portion of the paramere, slender with flattened point.

Conclusions
In this paper, we redescribe Downeshelea, providing new and updated features to better characterize the genus, based on Neotropical specimens. Some new records are noted, 10 species are redescribed, and 18 new species are described, increasing the number of species known to 60 Remarks. The specimens were previously identified as D. colombiae by Wirth, based on the wing spots and the leg coloration pattern. However, the presence of unequal claws on the fore-and mid legs, with the foreleg claws smaller than those of the midleg, make this morphotype unique within the genus.

Conclusions
In this paper, we redescribe Downeshelea, providing new and updated features to better characterize the genus, based on Neotropical specimens. Some new records are noted, 10 species are redescribed, and 18 new species are described, increasing the number of species known to 60 worldwide, with 46 of these present in the Neotropical region. Most species are known from a few specimens and only one locality and large areas in the Neotropical Region have never been sampled for Downeshelea. This strongly suggests that more species are yet to be discovered and strongly indicates the need for more studies in this region. Species from other biogeographic regions, where few studies have been done and descriptions are dated, are also generally poorly known.
Species of Downeshelea are unknown from the Palearctic Region. One species, D. notialisinica Yu and Hao, is known from China [40] but from the province Guangxi, which belongs to the Oriental Region [41]. The remaining species are restricted to the Afrotropical, Oriental, and Australasian Regions. As such, Downeshelea is largely a pantropical group. It seems likely that more species will also be found in the Old World tropics.
Based on distributions, some bionomic features of Neotropical Downeshelea species can be provided. They are generally found in humid areas often associated with the rainy season in each locality. Many are found in coastal forested areas, with mangrove, and otherwise are primarily in riparian and humid forested areas. The majority of species are restricted to localities below 500 m in elevation Only three species-D. jurgeni, D. magna and D. moravia-were found exclusively at high altitudes, from 1600 to 1850 m in Costa Rica, indicating the importance of studying habitats in high altitudes there and elsewhere.
In this paper, we included some important morphological features that can be used to identify some females, but we strongly recommended the association with males, which present more diagnostic characters, primarily in their genitalia. Two female morphotypes are described but remain unnamed, indicating the need of further studies of characters within the genus. This taxonomic study will provide the basis for future phylogenetic approaches to elucidate the probable synapomorphies that determine whether Downeshelea is monophyletic and its relationship to other Ceratopogonidae genera. This analysis will contribute to a better understanding of the phylogeny of the Ceratopogonidae family, whose relationships between many genera widely distributed in the Neotropical Region are still unknown. We emphasize, as well, the importance of body and leg coloration for species recognition, being crucial aspects for the separation of some very close and sympatric species (as in D. chirusi and D. pulla, for example).
Finally, the biology of these species is poorly known and the immature stages of species of Downeshelea are unknown. Downes [8] showed that female adults feed on adult Chironomidae, which can also explain the association of Downeshelea with aquatic habitats.