List of Culicoides biting midges ( Diptera : Ceratopogonidae ) from the state of Amazonas , Brazil , including new records

Culicoides are vectors of pathogenic agents that infect humans and other animals. Here, we provide a list of Culicoides from the state of Amazonas and also document new records from Presidente Figueiredo Municipality, Amazonas, Brazil. We provide a map of recorded species and a wing atlas for identification. The Culicoides fauna of Amazonas is now known to include 89 known species that belong to seven subgenera, 10 informal species groups, and one ungrouped species. We record nine species of Culicoides (C. aldomari, C. batesi, C. brownie, C. flavivenulus, C. franklini, C. guamai, C. para­ maruim, C. pusilloides and C. tidwelli) for the first time from Amazonas state. Culicoides brownie and C. tidwelli are reported for the first time from Brazil.


INTRODUCTION
Culicoides Latreille, 1809, or biting midges, are known vectors of pathogenic nematodes, protozoa, and viruses that cause infection in humans and other animals.In addition to their vectorial importance, these midges are pernicious due to their bites (Mellor et al. 2000).There are 1,355 species described worldwide (Borkent 2015); 116 species inhabit Brazil and 80 have been recorded in the state of Amazonas (Farias et al. 2016;Santarém and Felippe-Bauer 2016).A survey conducted in Presidente Figueiredo Municipality recorded species previously unknown in the state of Amazonas.Here, we revise the list Culicoides species occurring in Amazonas state and record the nine new species records for the state, and two new records for Brazil.

MATERIAL AND METHODS
We compiled a list of Culicoides species (Table 1), and mapped each species in Amazonas state, based on Borkent and Spinelli (2007), Farias et al. (2016), Castellón and Felippe-Bauer (2015), and Santarém and Felippe-Bauer (2016).The municipalities in Amazonas state where there are records of Culicoides are mapped in Figure 1.We collected additional records during an entomological study in a rural settlement of Rio Pardo, Presidente Figueiredo Municipality, Amazonas, Brazil.This survey was carried out between June and August 2010, using CDC light traps in forest fragments and peridomicilies (shelters for domestic animals and cultivation).Specimens of Culicoides were preserved in 70% ethanol, dissected, and slide-mounted in Canada balsam using the techniques described by Wirth and Marston (1968).We identified our specimens using various identification keys, as follows: Wirth and Blanton (1959); Spinelli et al. (1993); Felippe-Bauer et al. (2013); and Santarém et al. (2015).Subgeneric classification of Culicoides species was based on Borkent and Spinelli (2007) and Borkent (2015).Some voucher specimens are deposited at the Laboratory of Infectious Disease Ecology in Amazon (ILMD/FIOCRUZ, Amazônia; Table 1).Microphotographs of the wings were obtained using a digital system adapted to an optical microscope with a digital camera.

Lists of species
Table 1.Continued.

Continued
Identification: Medium-sized, dark brown; wing as figured; legs brown with pale bands; third segment palpus pit deep; spermathecae subspherical and very unequal.This species is differs from C. pusillus, from which it can be distinguished by second radial cell distinctly pale on the apical half, while in C. pusillus the second radial cell is pale only on extreme distal (Wirth and Blanton 1959).

Culicoides pusillus
Identification: Small size, bare; wing as shown in figure, without marginal pale spots, the very short, entirely dark second radial cell, blackish mesonotum with two faint darker vittae and relatively pale.This species is an exception among the related species of the subgenus Culicoides (Avaritia) due to the apex of the very short second radial cell entirely dark or only very slightly encroached on by the poststigmatic pale spot (Wirth and Blanton 1959).
Identification: A large species; third palpal segment slender with broad shallow pit; wing with distal pale spot in cell m 1 lying far from wing margin, no pale spot present in cell m 2 lying adjacent to midportion of mediocubital stem; aedeagus conspicuously cleft distad with two sharp laterally directed teeth on each side; parameres with simple twisted tips without fringing spines (Vitale et al. 1981).

Culicoides debilipalpis
Identification: Third palpal segment stout, PR 2.00-2.60;flagellomere 8 longer than 9; wing as shown in figure, with distal pale spot in cell m 1 separated from wing margin by a distance aproximate to its length, macrotrichia distributed on distal two-thirds of wing, extending in at least two rows to base of cell m 2 ; halter brown (Spinelli et al. 2005).
Identification: Eyes narrowly separated; third palpal segment moderately swollen, PR 2.0; AR 0.9; wing with distal light spot transverse in r 3 , r-m with light spot, presence of two bright spots in cell m 1 , the second spot not touch the edge of the wing, without clear area on one-third of vein M 2 , apices of veins M 1 and M 2 usually clear; light spot more or less confluent in anal cell.(Waller et al. 1990).

Culicoides glabrior
Identification: Eyes bare, broadly contiguous; third segment broad, with a very broad, shallow sensory pit; fourth segment much broader than fifth; wing with transverse distal spot in cell r 3 , only one distal spot in cell m 2 , pale spots present behind medial fork and in front of mediocubital fork, a single transverse pale spot in apex of anal cell, extending nearly to wing margin; halter brown, the end of the knob slightly paler; spermathecae two, pyriform, subequal (Wirth and Blanton 1959).

Culicoides hoffmani
Identification: Eyes separated, the line of separation broad above but narrowed below, short interfacetal hairs; third segment very short and swollen, with a moderately large and deep sensory pit; distal pale spot in cell r 3 moderately large, rounded; two pale spots in cell m 1 ; one pale spot each in apices of cells m 2 , CuA 1 and anal cell; a Identification: Eyes broadly separated, third segment very short and swollen, with a large, deep sensory pit; wing with pale spot over r-m crossvein about as broad as long, poststigmatic pale spots in cell r 3 small and rounded, well separated, the posterior one slightly larger and located slightly proximad of the anterior one; distal pale spot in cell r 3 moderately large, rounded; two pale spots in cell m 1 , CuA 1 and anal cell; definite pale spots present behind medial fork and in front of mediocubital fork (Wirth and Blanton 1959).
Distribution: Brazil (Acre, Amazonas and Roraima), Costa Rica, and Panama.Culicoides todatangae Wirth & Blanton, 1973 (Figures 3, 10I) Culicoides todatangae Wirth and Blanton (1973): 447. Pará, Brazil.Identification: Eyes narrowly separated, with long interfacetal hairs; third palpal segment long and slender, with a small sensory pit deeper than diameter of pore opening; wing as shown in figure; halter slightly infuscated; spermathecae two, oval with long slender necks, slightly unequal; halter slightly infuscated.This species resembles C. insinuatus both species having the pale sport in cell m 2 in front of mid-portion of the mediocubital stem (Wirth and Blanton 1973).

Culicoides youngi
Identification: Eyes nearly contiguous; third segment short and swollen to tip, with broad, round, shallow, sensory pit opening by a slightly smaller pore; second radial cell dark to tip with moderately broad lumen, slightly tapering distally, pale spot over r-m crossvein transverse, poststigmatic pale spot in cell r 3 not divided, continued and expanded around posterior margin of second radial cell; distal pale spot presente in cell r 3 , cell m 1 with two pale spots, cell m 2 with distinct pale spot lying behind medial fork and indistinct spot in front of mediocubital fork and a pale spot at apex of cell near wing margin, cell CuA 1 with large rounded pale spot in distal portion, anal cell with pale spot in distal portion, dark at base; vein M 1 with distinct pale spot at wing margin (Wirth and Barreto 1978).

Identification:
Wing with three pale spots in cell m 1 , distal pale spot distinctly separated from the subapical pale spot; third segment palpus short and stout (PR 1.7); sclerotized ring short; third rudimentary spermatheca greatly elongate.Male: paramere with definite broad ventral swelling, without elongate lobe (Felippe-Bauer et al. 2003).

Culicoides paraensis
Identification: Third segment palpus longer and more slender (PR 2.1-2.8);wing with three pale spots in cell m 1 , distal pale spot distinctly separated from the subapical pale spot; third rudimentary spermatheca shorter; sclerotized ring long and curved; male with paramere uniformly slender in midportion, with elongate lobe (Felippe-Bauer et al. 2003).

Culicoides quasiparaensis
Identification: Eyes narrowly separated above, contiguous below, the interocular space wedge-shaped; third palpal segment broad (PR 1.7) with pit broad and shallow, rarely deep and opening by a smaller pore; wing as shown in figure with three pale spots in cell m 1 ; distal pale spot distinctly separated from the subapical pale spot; spermathecae very unequal sized (Felippe-Bauer et al. 2003).

Culicoides aitkeni
Identification: Third palpal segment with scattered sensilla; wing length 1.50 mm; r 3 with pale spot present anterior to base of M 1 and in apices of CuA 1 and CuA 2 , a single pale spot crossing second radial cell; mid and hind femur with subapical pale band; spermathecae with short, slender necks.(Felippe-Bauer et al. 2009).
Distribution: Brazil (Amazonas, Pará); Trinidad and Tobago.Identification: This species is easily distinguished from the other species of the hylas group by the presence of double pale spots in r 3 , one crossing second radial cell and the other in distal portion of r 3 , by the absence of pale spot in front of base of M 1 and by the mid femur dark to tip (Felippe-Bauer et al. 2009).

Culicoides bimaculatus
Identification: Eyes contiguous by a distance equal to the diameter of 2 ommatidial facets; wing with r-m crossvein pale, one transverse distal pale spot in cell r 3 broadly reaching wing margin, two distal pale spots in cell m 1 (the second one very faint in two of the available specimens), apices of veins M 1 , M 2 , CuA 1 pale, apex of vein CuA 2 dark; spermathecae small, spherical, subequal with short necks.This species resembles C. foxi by virtue of the presence of a small blackish spot behind the apex of the second radial cell, and by the dark brown halter.Also, it can be distinguished from C. foxi by the stouter third palpal segment with irregular sensory pit, and by the pale r-m crossvein (Spinelli et al. 1993).

Culicoides brasilianum
Identification: Eyes contiguous by a distance equal to the diameter of 1 ommatidial facet; third palpal segment slightly broader just beyond middle, pit irregular; r-m crossvein pale, one transverse distal pale spot in cell r 3 broadly meeting wing margin, two distal pale spots in cell m 1 , apices of veins M 1 , M 2 and CuA 1 pale, apex of vein CuA 2 dark; halter knob dark, pedicel pale; spermathecae pyriform, unequal.Male genitalia with apicolateral processes of ninth tergum small and close together; aedeagus with terminal papilla; parameres connected at bases by a short loop, main bodies stout, apex with minute fringing hairs.This species distinguished from the related species by the wing pattern with extensive pale areas; from C. guttatus it also differs in the pale r-m crossvein and dark halter, and from C. ignacioi by the pale r-m crossvein (Spinelli et al. 1993).

Identification:
Wing with second distal pale spot in cell m 1 sometimes faint or ausent.This species is very similar to C. franklini, from which it can be distinguished by the stouter third palpal segment with irregular pit, eyes contiguous for 2.5-3.0 ommatidial facets (by 4 or 5 facets in C. franklini), and by its dark halter (Spinelli et al. 1993).
Identification: Eyes contiguous by a distance equal to diameter of 1.5-2.5 ommatidial facets; antenna brown, bases of flagellar segments 3-10 pale; third palpal segment slender, slightly broad in middle, pit irregular; wing as shown in figure, with r-m crossvein pale (faintly darkened in some topotypic specimens), one large, transverse, distal pale spot in cell r 3 broadly meeting wing margin, two distal pale spot in cell m 1 , the distal one smaller, apices of veins M 1 and M 2 pale, apices of veins CuA 1 and CuA 2 dark; halter pale; spermathecae subspherical to ovoid, unequal (Spinelli et al. 1993).

Culicoides filariferus
Identification: Eyes contiguous by a distance equal to the diameter of 1.5-2.0ommatidial facets; third palpal segment slightly broader in middle, with a conspicuous distal extension, pit irregular; wing as shown in figure, r-m crossvein pale (a faint darkish line, but not a definite dark spot, sometimes present on anterior half); one distal transverse pale spot in cell r 3 broadly meeting wing margin, two distal pale spot in cell m 1 (the second one faint or absent in some specimens); apices of veins M 1 and M 2 pale, apices of veins CuA 1 and CuA 2 dark (a few specimens with a faint pale area at apex of vein CuA 2 ); halter dark brown (Spinelli et al. 1993).

*Culicoides flavivenulus
Identification: Eyes contiguous by a distance equal to the diameter of 1-2 ommatidial facets; wing as shown in figure, r-m crossvein pale, one transverse distal pale spot in cell r 3 reaching wing margin, only one distal pale spot in cell m 1 , vein M 1 slightlly pale at apex; halter pale; spermathecae subspherical to pyriform, subequal.This species is similar to C. lutzi, from which it can be distinguished by the third palpal segment with a well-defined, round pit (irregular in C. lutzi); apices of veins M 2 and CuA 1 dark, broadly pale in C. lutzi (Spinelli et al. 1993).

Culicoides foxi
Identification: Eyes contiguous for a distance equal to the diameter of 1 ommatidial facet; third palpal segment elongated, pit rounded and shallow, the extension beyond the pit slender; wing as shown in figure, r-m crossvein entirely an deeply infuscated, one large, transverse, distal pale spot in cell r 3 broadly meeting wing margin, two distal pale spots in cell m 1 , apices of veins M 1 , M 2 and CuA 1 broadly pale, apex of vein CuA 2 dark; halter knob deeply infuscated, pedicel pale; spermathecae subspherical to ovoid, unequal (Spinelli et al. 1993).

Culicoides heliconiae
Identification: Third palpal segment with scattered sensilla; apical pale band on mid femur, hind femur dark to tip; wing with distal pale spot in r 3 narrow and transverse, r 3 with pale spot present anterior to base of M 1 , a single pale spot crosses the second radial cell, apices of CuA 1 and CuA 2 pale; spermathecae with short, slender necks (Felippe-Bauer et al. 2009).
Identification: Irregular sensory pit in the third palpal segment; wing as shown, r 3 with pale spot present anterior to base of vein M 1 , a single pale spot crossing the second radial cell, apices of CuA 1 and CuA 2 pale; mid femur with subapical pale band, hind femur dark to tip; spermathecae with short, slender necks (Felippe-Bauer et al. 2009).
Identification: Eyes contiguous by a distance equal to the diameter of 1.5 ommatidial facets; third palpus segment broad in middle, with conspicuous irregular pit; wing as shown in figure; halter knob bark brown, pedicel pale.This species is similar to C. fernandoi, from which it can be distinguished by the mandible with 20-22 teeth (14-15 in C. fernandoi), vein R 3 pale (infuscated on lower portion with a very small dark spot behind apex in C. fernandoi), distal pale spot in cell r 3 large, transverse (crescent-shaped or subdivided in C. fernandoi) (Spinelli et al. 1993).

Culicoides insignis
Identification: Eyes contiguous by a distance equal to the diameter of 1.5 ommatidial facets; third palpal segment with definite, irregular pit; wing as shown in figure, r-m crossvein dark on anterior half; vein CuA 2 dark up to the point where it turns abruptly forward to meet the costa; only one distal pale spot in cell m 1 ; halter dark (Spinelli et al. 1993).

Culicoides ocumarensis
Identification: Eyes contiguous by a distance equal to the diameter of 1.5-2.0ommatidial facets; third palpal segment slightly broader in middle, pit irregular; wing as shown in figure, r-m crossvein pale, one transverse distal pale spot in cell r 3 broadly meeting wing margin, two distal pale spot in cell m 1 , apices of veins M 1 and M 2 pale, apices of veins CuA 1 and CuA 2 dark; halter dark brown (Spinelli et al. 1993).

Culicoides palpalis
Identification: Third palpal segment with scattered sensilla; mid femur with apical pale band, hind femur dark to tip; wing with r 3 shows a pale spot present anterior to base of M 1 , a single pale spot crosses the second radial cell, apices of CuA 1 and CuA 2 dark; spermathecae with short, slender necks; male tergite 9 with very small papilliform process on posterior margin; fused portion of the parameres shows a width to length ratio of 1.75 (Felippe-Bauer et al. 2009).Spinelli, 1993 (Figures 4, 11F) Culicoides paraignacioi Spinelli in Spinelli et al. (1993): 66. Colombia.

Culicoides paraignacioi
Identification: Small wing length 1.14 mm, third palpal segment slender, with a rounded, small, shallow pit; wing as figured.This species is similar to C. ignacioi, which differs from the females of C. paraig nacioi by being significantly larger (wing lenth 1.53 mm), and by having the third palpal segment stouter, with a conspicuous irregular sensory pit.The male genitalia of both species are also very similar, with apicolateral processes widely separated, apex of aedeagus truncated, and parameres broadly fused at base, but they differ in the shape of the ninth tergum, which is rounded distally in C. ignacioi and subquadrangular in C. paraignacioi (Spinelli et al. 1993).

*Culicoides paramaruim
Identification: Medium-sized dark brown; wing as figured, r-m crossvein darkened, especially so on anterior half, vein CuA 1 infuscated, one transverse distal pale sport in cell r 3 not reaching wing margin; only one distal pale spot in cell m 1 ; apices of veins M 1 , M 2 , CuA 1 and CuA 2 dark.This species is differs from C. maruim, from which it can be distinguished by the third palpal segment stouter, while in C. maruim is long and slender (Spinelli et al. 1993).

Culicoides plaumanni
Identification: Medium-sized, eyes contiguous by a distance equal to the diameter of 1.5-2.5 ommatidial facets; third palpal segment with irregular pit; wing as shown in figure, r-m crossvein dark on anterior half, one distal, somewhat crescent-shaped pale sport in cell r 3 narrowly meeting wing margin, two distal pale spot in cell m 1 , apices of veins M 1 , M 2 and CuA 1 pale, apex of vein CuA 2 dark; halter dark brown (Spinelli et al. 1993).

Culicoides polypori
Identification: Third palpal segment with scattered sensilla; wing with r 3 with pale spot present anterior to base of M 1 , a single pale spot crosses the second radial cell, apices of CuA 1 and CuA 2 dark; mid and hind femur with subapical pale band; spermathecae with short, slender necks; male tergite 9 with a bilobed process on posterior margin; fused portion of the parameres slightly longer than basal width (Felippe-Bauer et al. 2009).

Identification:
Wing with contrasting pattern, r-m crossvein dark on anterior half, vein R 3 pale, one large transverse distal pale spot in cell r 3 broadly meeting wing margin, cell m 1 with two distal pale spots, apices of veins M 1 , M 2 and CuA 1 pale, apex of vein CuA 2 dark, scattered macrotrichia present at apices of cells r 3 , m 1 , and m 2 .Several species in the guttatus group are very similar to C. pseudodiabolicus, especially in wing; characteristic subdivided pit, pale halter, distinctly unequal spermathecae, and male aedeagus without terminal papilla (Spinelli et al. 1993).
Distribution: Mexico to Peru and Brazil (Acre, Amapá, Amazonas, Pará, Rondônia, Roraima, and Maranhão).Identification: Pale diffuse wing markings, rather than brightly contrasting; r 3 with pale spot present anterior to base of M 1 , pale spot that crosses the second radial cell subdivided in two separate spots, apices of CuA 1 and CuA 2 dark; third palpal segment with scattered sensilla; mid femur with apical pale band, hind femur dark to tip; spermathecae with short, slender necks (Felippe-Bauer et al. 2009).

Culicoides verecundus
Identification: Third palpal segment contains scattered sensilla; mid femur with apical and hind femur with subapical pale band; wing as shown in figure, r 3 with pale spot present anterior to base of M 1 , a single pale spot crossing second radial cell, apices of CuA 1 and CuA 2 dark; spermathecae with short, slender necks.Male tergite 9 with a prominent papilliform process on posterior margin; fused portion of the parameres longer than width; separate portion long and V-shaped at base (Felippe-Bauer et al. 2009).

Culicoides bricenoi
Identification: Second radial cell long, twice longer than first; r 3 with two large pale spots, the distal one located in distal portion of cell reaching anterodistal wing margin (Felippe-Bauer et al. 2013).

Culicoides discrepans
Identification: A large species; wing with distal pale spot in r 3 single, basal pale spot in m 1 large, slightly connected with the basal pale spot in m 2 , distal pale spot in m 2 reaching wing margin; one distal pale spot in anal cell, pale spot on M 1 restricted to extreme apex, connected with the distal pale spot in r 3 and m 1 (Felippe-Bauer et al. 2013).

Identification:
The only species in the subgenus Mataemyia with one spermatheca; male with stout, sinuate parameres with filiform tip; posteromedial projection of aedeagus tapered to slender, narrow, blunt tip, with a pair of well developed subapical points (Spinelli et al. 2007).
Identification: A large species; wing as shown in figure, first and 2 nd second radial cells nearly similar in length, distal pale spot in r 3 oblique, broadly reaching wing margin; anal cell with two distal small, round, pale spot (Spinelli et al. 2007).
Identification: Eyes narrowly separated, bare; third palpal segment moderately swollen, with moderately large, shallow, sensory pit; wing as figured, pale area at wing base and over r-m crossvein; very faint indications of pale spots in cells on distal part of wing; halter infuscated; spermathecae two, slightly, pyriform, slightly unequal (Wirth and Blanton 1959).

Culicoides acotylus
Identification: Third palpal segment moderately swollen, without sensory pit, the sensoria scattered on surface of segment; wing as figured, second radial cell very dark, r-m crossvein blackish, in the center of a large pale spot which broadly reaches costal margin, cell r 3 with five definite pale spots, apices of veins M 1 and M 2 pale margined a short distance, a pale spot each in apex of cell m 2 and cell CuA 1 , neither attaining wing margin, two pale spots in distal portion of anal cell and two in basal portion of this cell; cell m 2 with a pale spot lying in front of mediocubital fork, a lying behind medial fork and another lying across cell about halfway from these spots and wing base; halter pale; spermathecae two, pyriform, slightly unequal, the bases of the ducts sclerotized a short distance (Wirth and Blanton 1959).

Culicoides belemensis
Identification: Eyes narrowly separated, bare; third segment only slightly swollen with small, round, shallow, sensory pit on distal; wing as figured; halter brownish; spermathecae two, ovoid, unequal; aedeagus with distimedian process moderately slender with parallel sides and blunt, concaved tip; parameres with distinct ventral swelling ending in a distal lobe, apical portion slender, bearing lateral spines and terminating in a filiforme tip (Wirth and Blanton 1973).
Culicoides fairchildi Wirth and Blanton (1955): 102.Panama.Identification: Third palpal segment swollen, with a broad, shallow, irregular sensory pit; second radial cell very dark; pale spot over r-m crossvein broadly reaching costal margin, distal pale spot in cell r 3 rounded, almost reaching anterior wing margin, distal pale spot in cell m 1 elongate and nearly reaching wing margin, two pale spots, which may be more or less fused, in distal part of anal cell; base of anal cell pale; cell m 2 with a small pale spot lying in front of mediocubital fork and a pale area extending from behind medial fork to base of wing; halter yellowish (Wirth and Blanton 1959).
Distribution: Costa Rica, Panama, Colombia, Ecuador, and Brazil (Amazonas).Wirth & Blanton, 1953 (Figure 6) Culicoides carpenteri Wirth and Blanton (1953): 72.Panama Identification: Third palpal segment scarcely swollen, with a broad, shallow, sensory pit; second radial cell very dark, large pale area over r-m crossvein, distal pale spot in cell r 3 extensive, broadly extending across cell near apex from anterior wing margin to vein M 1 ; apices of veins M 1 , M 2 and CuA 1 dark; subapical pale spot in cell m 2 , distal pale spot in cell m 1 broadly meeting wing margin; two pale spots in distal part of cell m 2 , the distal one broadly meeting wing margin; the proximal one connected by a pale area extending to base of cell and including the pale spots lying in front of mediocubital fork and behind medial fork; pale area in cell CuA 1 nearly filling entire cell; anal cell pale except for a large dark area centering on middle of stem of mediocubital vein; halter pale (Wirth and Blanton 1959).
Identification: Eyes narrowly separated above, with long interfacetal hairs; third palpal segments swollen, with a broad, shallow, sensory pit; wing as shown in figure, poststigmatie pale spots in cell r 3 more or less fused, the posterior one located slightly proximad of the anterior one, distal pale spot in cell r 3 small, only one small pale spot in distal part of anal cell and one pale spotin distal part of cell m 2 ; halter whitish; spermatheca one, pyriform (Wirth and Blanton 1956).

Culicoides transferrans
Identification: Eyes narrowly separated, with long interfacetal hairs; third palpal segment moderately swollen, with a broad, shallow, sensory pit; wing as shown in figure, with four pale spots in a rhomboid in cell r 3 , the distal pair usually fused and broadly attaining wing margin anteriorly; two pale spots each in cells m 1 and apices of cells m 2 and anal cell; pale spots present behind medial fork and in front of mediocubital fork; halter yellowish; spermatheca one, oval with a short portion of the duct sclerotized (Wirth and Blanton 1959).

Culicoides propriipennis
Identification: This species is most similar to C. tetra thyris, but C. propriipennis can easily be distinguished by the presence of a third pale spot at the wing margin in cell m 1 , by the second pale spot in the distal portion of cell m 2 , wing as figured (Wirth and Blanton 1959).

Culicoides rangeli
Identification: Eyes nearly contiguous, with long interfacetal hairs; third segment swollen, with a very broad, shallow, sensory pit, open and without regular pore; posterior poststigmatic pale spot in cell r 3 very small or entirely absent, distal spot in cell r 3 transverse and not double, not attaining wing margin; two very small pale spots each in cell m 1 and in apices of cell m 2 and anal cell; a pale spot present behind medial fork and another in front of mediocubital fork; halter pale; spermatheca one, pyriform, with long sclerotized neck (Wirth and Blanton 1959).
Identification: Third palpal segment short and slightly swollen, with a small, deep sensory pit; wing as shown in figure, with second radial cell moderately dark, cell r 3 with two separate round poststigmatic pale spots, the posterior one located behind second radial cell, the distal pale spot hourglass-shaped on wing margin, apices of veins M 1 and M 2 with a small pale spot at wing margin, cell CuA 1 with a round pale spot almost meeting wing margin, only one small round pale spot in distal part of anal cell, cell m 2 with a small pale spot present lying behind medial fork; halter pale; spermathecae two, oval, slightly unequal (Wirth and Blanton 1959).

Culicoides leopoldoi
Identification: Wing as figured, with pale spot on crossvein r-m extensive; pale spot present behind second radial cell, proximal to the poststigmatic pale spots; poststigamtic pale spots longitudinally aligned, posterior one smaller; distal pale spot in cell r 3 large, with narrow proximal extension; one spermatheca (Spinelli et al. 2005).

Culicoides tetrathyris
Identification: Resembling C. propriipennis in wing pattern, mesonotal and leg markings, structural characters, and male genitalia, but C. propriipennis differs by having the sides of the mesonotum brownish instead of bluish pruinose, a third pale spot in cell m 1 at the wing margin, the pale spots in cell r 3 arranged in a rhomboid instead of a trapezoid, distal sensory tufts in falgellomeres 1, 7-12, and the pit on the third palpal segment broader.The male genitalia of the two species are practically identical, but in C. propriipennis the stem of the paramere is stouter and not bent near the base (Wirth and Blanton 1959).
Identification: Wing as figured; separated from C. fieldi by its shorter second palpal segment (9/13 as long as third segment), flagellomeres 9-11 and 12 subequal in length and by the presence of a distinct pale spot in front of the mediocubital fork (Wirth and Blanton 1959.Distribution: Venezuela, Trinidad, and Brazil (Amazonas, Roraima, and Rio de Janeiro).Wirth & Blanton, 1956 (Figure 7) Wirth and Blanton (1956): 50.Honduras.

Culicoides fieldi
Identification: Eyes contiguous, with long interfacetal hairs; third segment moderately swollen, with a small, deep, sensory pit; pale spot over r-m crossvein small, poststigmatic pale spots in cell r 3 more or less separated into two distinct, distal pale spot in cell r 3 small and round in center of cell, two pale spots in cell m 1 , only one pale spot each in apices of cells m 2 , CuA 1 and anal cell, pale spot present behind medial fork; halter infuscated; spermatheca one, pyriform (Wirth and Blanton 1959).

Culicoides glabellus
Identification: General appearance, size, and wing markings practically identical to C. leoni Barbosa, 1952 andC. gabaldoni Ortíz, 1954.There are two spermathecae but in C. gabaldoni and C. leoni differs by having an antennal ratio of only 0.75 and sensoria present in flagellomere 1, 5, and 6-8.The male genitalia of C. glabellus is distinguished by the gradually curving base of the paramere, the swollen stem at the distal part of the straight portion, beyond which the paramere tapers rapidly to a simple, filamentous tip (Wirth and Blanton 1959).

Culicoides leoni
Identification: Sensilla coeloconica on flagellomeres 1, 5-8; wing length 0.63 mm, wing pattern with the caudal extension of the post-stigmatic pale spot not oblique in cell r 3 and usually connected to vein M 1 , smaller pale spot on r-m crossvein, one pale spot in cell M 1 near base, and distal pale spot in anal cell abutting wing margin (Huerta et al. 2012).

Culicoides galindoi
Identification: Eyes contiguous, bare; third segment slightly swollen, with a small, shallow, sensory pit; have the pale distal spot in cell r 3 rounded distally, leaving a small dark area in apex of cell; halter pale; spermathecae two, pyriform, subequal (Wirth and Blanton 1959).

Culicoides limai
Identification: Small; the much broader third palpal segment; wing as shown in figure, with distal pale spot in cell m 1 broadly abutting wing margin, second radial wholly included in a dark spot, no pale spot straddling vein M 2 , this vein usually dark to apex, apices of veins M 1 , M 2 dark; subapical pale band on the hind femur and pale apex of the hind tibia; presence of a small ventral lobe on the male parameres (Wirth and Blanton 1959).
Identification: Eyes narrowly separated, bare; third segment cone-shaped with deep circular sensory pit; sensory pattern in flagelomeres 1, 5-8; halter pale; two spermathecae, subequal, with ducts sclerotized a considerable distance.Antennal proportion, distribution of sensoria, and overall wing pattern place C. santanderi within the limai group.This species is distinguished from others in that group by the divided poststigmatic pale spot, the posterior one larger and extending proximal to the anterior one (Browne 1980).
Identification: Wing as figured, distal pale spot in cell r 3 large, filling apex of cell with dark area boadering tip of vein M 1 very little expanded apically; fore and mid femora with suapical and all tibiae with sub-basal, narrow pale rings.This species has a nearly identical wing pattern to C. limai and C. galindoi, but in both related species the distal pale spot in cell r 3 is rounded distally leaving a sizeable dark area at apex of cell.Culicoides limai has male genitalia nearly identical with C. vernoni, but the hind femur has a distinct subapical pale ring.Culicoi des galindoi has the hind femur dark to tip, but the male parameres lack the ventral lobe (Wirt and Blanton 1973).Distribution: Costa Rica, Colombia, Bolivia, and Brazil (Amazonas and Pará).Fox, 1952 (Figure 8) Culicoides caprilesi Fox (1952): 364.Venezuela.Culicoides kintzi Wirth and Blanton (1953): 72.Panama.

Culicoides caprilesi
Identification: Distinguished in pinned mounts by the greater development of the dark mesonotal striae and the much hairier wing on which the dark color pattern is more extensive, dark mark extending from the base of the second radial cell almost to its apex and extending over into cell r 3 as a small rounded dark spot just behind the tip of the second radial cell.In some specimens, the latter dark spot is narrowly connected to the next distal dark area (Wirth and Blanton 1959).

Culicoides pachymerus
Identification: Smaller than C. uniradialis Wirth & Blanton, 1953 and C. caprilesi, its wing is less hairy than that of C. caprilesi, there is no dark spot or connection immediately posterior to the end of the second radial cell as in C. caprilesi.The male genitália of C. pachymerus closely resemble those of C. uniradialis, with a long ventral lobe present on the parameres, but differing in having a median notch on the ninth tergum, aedeagal arch much broader with slender basal arms and no barbs on the tips of the parameres (Wirth and Blanton 1959).
Distribution: Guatemala to Colombia and Brazil (Amazonas).
Identification: Wing as figured, r 3 with five pale spots in the proximal half of cell; m 1 with two pale spots, the distal one far from wing margin; AR 0.72-0.76;proboscis short, P/H ratio 0.61-0.70;male with sternite 9 with slightly perceptible posteromedial excavation; aedeagus with basal arch extending to 0.3 of total length; parameres without ventral lobes, distal portion curved, apex flattened (Santarém et al. 2015).

Identification:
The only species of the reticulatus group with sensilla coeloconica on flagellomeres 1 and 9-13 and a tibial comb with six spines (sensilla coeloconica 1, 5-8 or 1, 6-8 and tibial comb with 4 or 5 spines in other species of the group).Third palpal segment long (PR 4.2) with a small, rudimentary sensory pit (PR 1.6-3.2, with variable sensory pit, in other species of the group).The female of C. castelloni has similar wing pattern to C. reticulatus, but distinguished by the characters described above and by the presence of two parallel, longitudinal, admedian pale spots, anterior to the prescutellar depression, in C. reticulatus with four median anterior yellowish spots arranged as a leaf clover (Santarém et al. 2015).
Culicoides fittkaui Wirth & Blanton, 1973 (Figures 8, 12J) Culicoides fittkaui Wirth and Blanton 1973: 432. Pará, Brazil.Identification: Eyes nearly contiguous; proboscis short, P/H ratio 0.60-0.63;hind femur dark; wing as shown in figure, with pale spots of the wing interconnected, r 3 with two pale spots, second radial cell with pale lumen, m 1 with two pale spots, the distal one reaching wing margin; aedeagus in male with basal arch extending half of total length; parameres with large ventral lobes, apical portion with lateral fringe of spicules (Santarém et al. 2015).

*Culicoides guamai
Identification: Proboscis short, P/H ratio 0.71-0.79;hind femur dark; wing as figured, with pale spots of the wing interconnected, r 3 with two pale spots, m 1 with two pale spots, the distal one reaching wing margin, second radial cell short, with dark lumen; aedeagus in male with basal arch extending to 0.6 of total length; parameres with a strong ventral swelling on proximal portion of stem, distal portion with filamentous tip with fringing spines (Santarém et al. 2015).

Culicoides hildebrandoi
Identification: Second radial cell in dark spot, r 3 with four sparsely distributed pale spots, r-m crossvein pale; hind femur with subapical pale band; scutum with two anterior submedian clover-leaf shaped spots; 3 rd palpal segment elongate, slightly swollen, cylindrical, without a sensory pit but with capitate sensilla scattered on the surface cuticle; tergite 9 of male with a posteromedial notch, parameres with slightly sinuous stem, swollen on mid-portion and without a ventral lobe and the basal Identification: Third palpus segment with a multiple, shallow, irregular sensory pit, PR 2.0-2.9;moderately long proboscis, P/H ratio 0.80-0.94;mandible with 16 teeth; scutum with four median anterior yellowish spots arranged as a leaf clover; wing as shown in figure, r 3 with four pale spots, m 1 with two pale spots; two unequal ovoid spermathecae (Santarém et al. 2015).
Culicoides kuripako Felippe-Bauer, 2010 (Figure 9) Identification: Pale spots of the wing interconnected; r 3 with two pale spots, postigmatic pale spot inverted L-shaped, isolating a dark spot behind second radial cell; m 1 with two pale spots, the distal one reaching wing margin; CuA 1 and CuA 2 pale.Male: sternite 9 with deep posteromedial excavation; aedeagus with basal arch extending to half of total length; parameres with ventral lobes, distal portion with lateral fringe of spicules (Santarém et al. 2015).
Identification: Proboscis short, P/H ratio 0.63-0.66;AR 0.72-0.78;wing as figured, r 3 with five pale spots, m 1 with three pale spots, the distal one near wing margin; hind femur dark, hind tibia dark apically; aedeagus in male with basal arch extending to half the total length; parameres with blade-like ventral lobes, distal portion with lateral fringe with fine spicules (Santarém et al. 2015).
Identification: Wing as figured, with pale spots of the wing interconnected; r 3 with two pale spots, poststigmatic pale spot inverted L-shaped, isolating a dark spot behind second radial cell; m 1 with two pale spots,  the distal one reaching wing margin; CuA 1 pale and CuA 2 dark (Santarém et al. 2015).
Culicoides profundus Santarém, Felippe-Bauer and & Trindade, 2014 (Figures 9, 12P  Identification: Third palpal segment swollen, with large, deep sensory pit in middle portion, opening in a small, rounded pore, PR 1.8-2.4;moderately long proboscis, P/H ratio 0.86-0.95;mandible with 16 teeth; scutum with four median anterior yellowish spots arranged as a leaf clover; wing as shown in figure, r 3 with four pale spots, m 1 with two pale spots, the distal one far from wing margin (Santarém et al. 2015).
Distribution: Brazil (Amazonas and Pará).Identification: Scutum with four median anterior yellowish spots arranged as a leaf clover; third palpal segment with a shallow, round sensory pit in middle portion, with no well-defined contours on distal half, PR 2.0-2.7;proboscis moderately long, P/H ratio 1.0-1.06;mandible with 19 teeth; wing as figured, r 3 with four pale spots; m 1 with two pale spots, the distal one far from wing margin (Santarém et al. 2015).

Culicoides pseudoreticulatus
Distribution: Brazil (Amazonas and Roraima).Identification: Third palpal segment rhomboid, with a shallow, well-developed irregular sensory pit that reaches the extreme distal portion of segment, PR 2.5-3.0;proboscis long; P/H ratio 1.08-1.28;scutum with four median anterior yellowish spots arranged as a leaf clover; wing as figured, r 3 with four pale spots, m 1 with two pale areas, the distal one far from wing margin (Santarém et al. 2015).
Identification: Eyes narrowly separated, bare; third palpal segment short and swollen, with broad, shallow sensory pit; second radial cell dark, pale spot over r-m crossvein very broad, cell r 3 with three pale spots, a small one at end of second radial cell, an elongate one behind second radial cell sometimes narrowly meeting the pale spot over crossvein and the anterior poststigmatic pale spot, and the third pale spot extensive, nearly filling apex of cell r 3; spermathecae two (Wirth and Blanton 1959).

DISCUSSION
The richness of Culicoides in the state of Amazonas is 6.6% of all species of the genus worldwide and 77% all Culicoides species in Brazil (Borkent 2015;Farias et al. 2016;Santarém and Felippe-Bauer 2016).Catalogs and lists of species are important tools in the study of biodiversity of localities, states, countries, and zoogeographic regions (Santarém and Felippe-Bauer 2016).Our list of species may be helpful in identifying the Culicoides fauna of Amazonas, for it gathers together data on the identification and distribution of species occurring in the state.Moreover, we extend the ranges of nine species to the state of Amazonas, including two species are recorded from Brazil for the first time.
Despite the importance Culicoides to human health, the Culicoides fauna of Amazonas state is still poorly known.Our list of Culicoides of Amazonas adds to the knowledge of the diversity of this genus and may be useful for medical entomology, vector epidemiology, and management of these pernicious insects.

FariasFigure 1 .
Figure 1.Maps showing the position of Amazonas state in Brazil and South America.Large image shows the municipalities where there were records of Culicoides (modified after SimpleMappr, http://www.simplemappr.net).

Farias
et al. | List of Culicoides biting midges from the state of Amazonas, Brazil Farias et al. | List of Culicoides biting midges from the state of Amazonas, Brazil distance equal to the diameter of 2-3 ommatidial facets; third palpal segment stout, with irregular pit; wing as shown in figure, crossvein r-m pale, one transverse distal pale spot in cell r 3 reaching wing margin, only one distal pale spot in cell m 1 , apices of veins M 1 , M 2 and CuA 1 broadly pale, apex of vein CuA 2 dark; halter knob pale

Farias
et al. | List of Culicoides biting midges from the state of Amazonas, Brazil

Farias
et al. | List of Culicoides biting midges from the state of Amazonas, Brazil

Farias
et al. | List of Culicoides biting midges from the state of Amazonas, Brazil

Farias
et al. | List of Culicoides biting midges from the state of Amazonas, Brazil