Five new species of Culicoides Latreille described from Colombia, yielding a new species list and country records (Diptera: Ceratopogonidae)

The following five new species of Culicoides from Colombia are described, illustrated and placed to subgenus or species group: Culicoides antioquiensis, Culicoides gabrieli, Culicoides inermis, Culicoides micayensis and Culicoides nigrifemur. C. gabrieli is also known from Peru. When possible, their position in previously published keys is indicated and their features discussed in light of the most recent revisions. A list of 180 Culicoides species known (114) or suspected of being in Colombia (66) is given in a Table. Of these, 12 including the new species are recorded from Colombia for the first time.

Species in the genus Culicoides Latreille are by far the most notorious members of the 109 currently recognized extant genera of Ceratopogonidae. In the Neotropical region, Borkent and Spinelli (2007) recorded 266 species and Spinelli et al. (2007) described another species from Brazilian Amazonia. Seventy of these species are miserable pests of humans and domestic animals and seven of them serve as vectors of a variety of diseases, summarized by Borkent and Spinelli (2007).
There is no reliable, modern key to the subgenera and species groups of neotropical Culicoides. The only review of the genus for the region is the monography by Forattini (1957), but due to the numerous descriptions of new species and nomenclatorial actions by subsequent authors, this revision has not been updated. The easiest guide to species identification is that of Wirth et al. (1988), in which some meristic characters states and photographs of wings are provided. From there readers may refer to keys to subgenera or species groups, or species lists referred to local revisions from Panama (Wirth & Blanton 1959), Amazon Basin (Wirth & Blanton 1973), the Caribbean (Wirth & Blanton 1974), Trinidad and Tobago (Aitken et al. 1975), Florida, USA (Blanton & Wirth 1979), Colombia (Barreto 1986), South Amazon Basin (Spinelli & Wirth 1986), Argentina (Spinelli et al. 2005) and Costa Rica (Spinelli & Borkent 2004), among others.
In his catalog of the Culicoides from Colombia, Barreto (1986) recorded 88 species, and since then 14 species have to be added for a total of 102 Culicoides species presently known from Colombia. Although this is a relatively large number of species, at present their sanitary importance in Colombia appear to be restricted to the annoyance caused by the female biting habits. None of the recorded species has been irrefutably incriminated as a vector of pathogens, only Culicoides insignis Lutz is suspected to be involved in the transmissión of the bluetongue virus and the haemorragic fever virus to cattle (Homan et al. 1985), and Culicoides insinuatus Ortiz in the transmission of the filaroid Mansonella ozzardi to man (Tidwell & Tidwell 1982).
A recent study of the collections of Culicoides from the Museo de La Plata, in La Plata, Argentina, and the Instituto Nacional de Salud, in Bogotá, Colombia, revealed the presence of five new species and an additional seven species not recorded yet to Colombia. The purpose of this paper is to provide the descriptions and records of this material, as well as an updated list of the 114 species inhabiting the country with their corresponding distribution. This list includes 66 species not formally recorded and expected to be discovered in Colombia.

MATERIAL AND METHODS
All specimens, mounted on microscope slides in Canada balsam, were examined and measured with a binocular compound microscope at 40-400X and drawings of certain diagnostic characters were prepared with an attached camera lucida. Wing photographs were taken with a Pentax Optio S 40, digital camera through a Leitz Wetzlar SM-LUX, binocular microscope.
Terms for structures follow those used in the Manual of Nearctic Diptera (McAlpine et al. 1981). Wing veins follow the system of the Manual of Nearctic Diptera, with modifications proposed by Szadziewski (1996). Names of veins are always in upper case and those of cells in lower case. Pale areas in cell r 3 posterior to or immediately distal to the 2nd radial cell are called poststigmatic pale spots. Ratios used follow .
Specimens were deposited, as noted, in the collections of the Instituto Nacional de Salud, Bogotá, Colombia (INS), and the Museo de La Plata, La Plata, Argentina (MLP).
Culicoides (Hoffmania) antioquiensis sp. n., 11,25) Diagnosis: Only species in the hylas group with slender third palpal segment, apices of veins M 1 , M 2 broadly pale, apex of CuA 1 with small pale spot and apex of CuA 2 dark, spermathecae without sclerotized necks, male tergite 9 with a distinct distal notch, gonostylus with subapical tooth and with V-shaped base of the separate portion of parameres.
Female: head dark brown. Eyes (Fig. 11) bare, contiguous by distance equal to diameter of two ommatidia. Flagellum (Fig. 3) brown, bases of flagellomeres pale; AR 1.00; sensilla coeloconica on flagellomeres 1, 9-13. Palpus (Fig. 4) dark brown; third segment cylindrical, slender, sensilla scattered on surface; PR 4.15; P/H ratio 1.23. Mandible with 22 teeth. Scutum dak brown, with sublateral yellowish brown patches; scutellum, postscutellum dark brown. Legs dark brown; foreknee blackish with narrow pale ring on each side; midknee broadly yellowish on each side of joint; hindfemur dark to tip, hindtibia with narrow basal and apical pale rings; hind tibial comb with seven spines, second from spur longest. Wing (Fig. 25) length 1.40 mm; width 0.60 mm; CR 0.67; with contrasting pattern; pale spot over crossvein r-m abutting wing margin; second radial cell in pale spot; r 3 with elongate pale spot anterior to base of M 1 , distal pale spot in r 3 transverse, reniform, barely abutting wing margin; M 2 straddled by pale spot nearly its midlength; distal pale spot in m 1 broadly separated from wing margin; distal pale spot in m 2 rounded, broadly abutting wing margin; pale spot in cua 1 small, narrowly connected with narrow pale line bordering lower margin of CuA 1 ; anal cell with two basal, two distal pale spots; apices of M 1 , M 2 broadly pale, apex of CuA 1 with small pale spot, apex of CuA 2 dark; pale spot posterior to medial fork broadly connected with pale spot lying anterior to cubital fork. Macrotrichia sparce on distal half of wing, extending nearly to base of anal cell. Halter pale. Abdomen dark brown. Two ovoid, unequal spermathecae without sclerotized necks (Fig. 5), measuring 57 by 45 μ, and 46 by 39 μ; rudimentary third, ring present.
Distribution -Colombia; known only from the typelocality.
Taxonomic discussion -Culicoides antioquiensis sp. n. is a member of the hylas group of the subgenus Hoffmania Fox. This new species keys out in Wirth and Blanton (1968) to couplet 3 where is distinguished from Culicoides heliconiae Fox and Hoffman by the dark apex of vein CuA 2 and from Culicoides palpalis Macfie by the small pale area in the apex of vein CuA 1 .
The male is very similar to C. palpalis, but in the latter species the tergite IX bears a very small, papilliform caudomedian process, the gonostylus lacks the subapical tooth and the base of the separate portion of parameres is rounded. The female of C. palpalis differs from the new species by the spermathecae with short and slender necks, by the distal pale area in cell r 3 broadly abutting wing margin and by the large pale spot in cua 1 , broadly connected with the pale line bordering lower margin of CuA 1 .
The wing pattern of C. antioquiensis is similar to the one of Culicoides hylas, but the later species has dark mid knee and the third palpal segment bears an irregular sensory pit. The male of C. hylas has a small papilliform on posterior margin of tergite 9 and the fused proximal portion of the parameres is nearly as broad as long.
Etymology -The name of this species refers to Antioquia, the Department of the type-locality.
The male genitalia of C. bambusicola, a species inhabiting Eastern Brazil and Argentina, and Colombia and Venezuela, is similar to the one of C. gabrieli sp. n. Nevertheless, the sternite 9 lacks the mesal notch and has stouter parameres. Regarding the female, the wing of C. bambusicola exhibits a distal rounded pale area in cell r 3 .
Culicoides raposoensis, a species also inhabiting Colombia, differs from C. gabrieli sp.n. by the cell r 3 with a distal pale spot abutting wing margin, by the pale apex of the hindtibia, and by the male genitalia with parameres with well developed ventral lobe and aedeagus lacking lateral pointed processes.
Etymology -We are pleased to name this species after the Colombian writer Gabriel García Márquez, in recognition of his monumental literary work, the delight of several generations of readers around the World.
Distribution -Colombia, known only from the type-locality.
Taxonomic discussion -C. inermis sp. n. belongs in the decor species group of the subgenus Anilomyia, reviewed by for the Neotropics by Wirth and Blanton (1970). The antennal sensillar pattern 1,5-8 is unique in the subgenus. The wing pattern is nearly identical to Culicoides decor (Williston). However, apart from the different distribution of sensilla coeloconica whithin flagellomeres, C. inermis sp. n. is easily distinguished from C. decor by the shorter proboscis (P/H ratio 0.85 in C. decor), unarmed mandibles (armed in C. decor) and by the maxillary palpus bearing only three well developed segments, with apical sensory pit in the third segment (with 5 developed segments in C. decor). These extra alar characteristics also distinguish C. inermis from other species in the group, all of them with palpus 5-segmented, armed mandibles and P/H ratio ranging from 0.68-0.94.
Etymology -This species is named inermis referring the unarmed mandibles.
Distribution -Colombia, known only from the type-locality.
Taxonomic discussion -C. micayensis sp. n. is a member of the eublepharus group and unplaced to subgenus. The following four species of the eublepharus group also have one spermathecae: Culicoides archboldi Wirth and Blanton, Culicoides eublepharus Macfie, Culicoides guadeloupensis Floch and Abonnenc and Culicoides rangeli Ortiz and Mirsa. Of these the most similar is C. archboldi, but it is readily distinguished from C. micayensis by the sensilla coeloconica on flagellomeres 1,9-12, by the absence of pale spots in cells m 1 , m 2 , cua 1 , and anal cell, and by the long and coarse macrotrichia covering most of wing, reaching in two rows to base of cell m 2 . The wing pattern of C. micayensis sp. n. is similar to the one of C. tamboensis, but the later species has two well developed spermathecae and bears sensilla coeloconica on flagellomeres 1,9 -12 Etymology -The name of this species refers to Rio Micay, the type-locality.

Culicoides nigrifemur sp. n. (covagarciai group) (Figs 22-24)
Diagnosis: Only species in the covagarciai species group with third palpal segment slender, head and proboscis equal in length, and entirely dark hindfemur.
Distribution -Colombia, known only from the type-locality.
Taxonomic discussion -C. nigrifemur sp. n. belongs in the covagarciai group of the subgenus Anilomyia, and is readily distinguished from the species included in that group by the entirely dark brown hindfemur and by the equal length of head and proboscis. Wirth and Blanton (1956d) reviewed the Culicoides covagarciai Ortíz species group for the Neotropics, and this new species keys out to Culicoides marshi Wirth and Blanton in couplet 2, with the exception that the proboscis of C. marshi is longer than its head. Besides that, the eyes are broadly contiguous in C. marshi, and only separated by two ommatidia in C. nigrifemur sp. n. The wing pattern of C. nigrifemur is nearly identical to the one of C. covagarciai, but apart from the different length of proboscis and legs coloration, C. covargaciai is easily distinguished from C. nigrifemur sp. n. by its swollen third palpal segment.
Etymology -This species is named nigrifemur referring the uniformly dark coloration of the hindfemur.