Zootaxa 3795 (3): 255–274 ISSN 1175-5326 (print edition)

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Copyright © 2014 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
http://dx.doi.org/10.11646/zootaxa.3795.3.2
http://zoobank.org/urn:lsid:zoobank.org:pub:2A90C789-0A57-4905-9253-F00DD530E340

New Neotropical Culicoides and redescription of Culicoides reticulatus Lutz

(Diptera: Ceratopogonidae)

MARIA CLARA ALVES SANTARÉM1,5, ROSIMEIRE LOPES DA TRINDADE2,

TIAGO DO NASCIMENTO DA SILVA1, ELOY GUILLERMO CASTELLÓN3,
CÁTIA ANTUNES DE MELLO PATIU4 & MARIA LUIZA FELIPPE-BAUER1
1
Coleção de Ceratopogonidae, Laboratório de Diptera, Instituto Oswaldo Cruz, Av. Brasil 4365, CEP 21040-900, Rio de Janeiro, RJ,
Brazil. E-mail: mcalves@ioc.fiocruz.br; tnsilva@ioc.fiocruz.br; mlfbauer@ioc.fiocruz.br
2
Divisão de Zoologia, Instituto de Pesquisas Científicas e Tecnológicas do Estado do Amapá, Rod. Juscelino Kubitschek, km 10,
Fazendinha, CEP 68903-419/68903-197, Macapá, AP, Brazil. E-mail: rositrindade@hotmail.com
3
Laboratório de Insetos Sinantrópicos, Instituto Nacional de Pesquisas da Amazônia, Av. André Araújo, 2936, Aleixo, CEP 69060-
001, Manaus, Amazonas, Brazil. E-mail: eloygcb@gmail.com
4
Departamento de Entomologia, Museu Nacional/UFRJ, Quinta da Boa Vista, s/n., São Cristóvão, CEP 20940-040, Rio de Janeiro,
RJ, Brazil. E-mail: camello@acd.ufrj.br
5
Programa de Pós-Graduação em Zoologia, Museu Nacional/UFRJ, Quinta da Boa Vista, s/n., São Cristóvão, CEP 20940-040, Rio de
Janeiro, RJ, Brazil

Abstract

Seven new species of the Culicoides reticulatus species group are described and illustrated based on specimens from Co-
lombia, Panama and Brazilian states of northern (Amazonas, Pará and Roraima) and southeast region (Rio de Janeiro).
The new species are compared with their similar congener, Culicoides reticulatus Lutz, and a systematic key, table with
numerical characters to the females of the species are provided. Redescription of Culicoides reticulatus Lutz is given
based on type series deposited in the Ceratopogonidae Collection of the Oswaldo Cruz Foundation, Brazil. The distribu-
tion of these species is presented in figure 9.

Key words: Culicoides reticulatus Lutz, bloodsucking midges, Brazil, Colombia, Panama, new species

Resumo

Sete espécies novas espécies de Culicoides do grupo reticulatus são descritas e ilustradas baseadas em espécimes da
Colômbia, Panamá e de estados da região Norte (Amazonas, Pará e Roraima) e do Sudeste (Rio de Janeiro) do Brasil. As
espécies novas são comparadas com a similar congenérica, Culicoides reticulatus Lutz, e são fornecidas chave sistemática
e tabela com os caracteres numéricos das fêmeas. A redescrição de Culicoides reticulatus Lutz é feita baseada na série tipo
depositada na Coleção de Ceratopogonidae da Fundação Oswaldo Cruz, Brasil. A distribuição geográfica das espécies está
presente na figura 9.

Palavras-chave: Culicoides reticulatus Lutz, maruins, Brasil, Colômbia, Panamá, espécies novas

Introduction

The precise identity of Culicoides reticulatus Lutz has been subject of considerable confusion in the literature as
mentioned below. Lutz (1913) described the species from specimens collected in the States of Rio de Janeiro, São
Paulo (Santos) and Bahia. He pointed out that the third palpal segment of the female present a rounded pit in
swollen mid portion and a scutum with a prominent pattern with four median anterior yellowish spots arranged as a
leaf clover. Lutz also cited that the immature stages can be found in crab holes in mangroves areas and the females

Accepted by L. Hernandez-Triana: 21 Mar. 2014; published: 13 May 2014 255

can bite human, horses and cow in the vicinity of these areas. Barbosa (1943) described the male and redescribed
the female from specimens collected in mangrove areas of Recife, Pernambuco, Brazil and, as it was observed by
Lutz, corroborated that the third palpal segment of the female present one distinct palpal pit. Forattini & Galvão
(1955), studying material from Brazil (São Vicente and Caraguatatuba, São Paulo State) and Panama, redescribed
C. reticulatus and observed differences in the number of palpal pits, varying from one to three. Wirth & Blanton
(1959), in their revision of Culicoides from Panama, redescribed this species and cited the presence of a double
sensory pit in female specimens. Posteriorly, Wirth & Blanton (1973) expanded its distribution to Colombia and
Costa Rica and Wirth et al. (1988) mentioned its occurrence in Brazil, Colombia, Panama, Costa Rica and
Honduras, being this distribution considered as valid until now. In order to better known the identity of this species,
we studied specimens identified as C. reticulatus, including the only known type series from Bahia, Brazil,
deposited in Ceratopogonidae Collection of the Oswaldo Cruz Foundation (CCER), Rio de Janeiro, Brazil.
The study revealed that the specimens identified as C. reticulatus from different regions are significantly
different from type specimens and represent seven undescribed species, which are herein described and illustrated:
C. amazonicus Santarém, Felippe-Bauer & Trindade, C. diplus Santarém & Felippe-Bauer, C. fluminensis
Santarém & Felippe-Bauer, C. irregularis Santarém, Felippe-Bauer & Castellón, C. profundus Santarém, Felippe-
Bauer & Trindade, C. pseudoreticulatus Santarém, Felippe-Bauer & Castellón and C. rhombus Santarém, Felippe-
Bauer & Castellón. Culicoides reticulatus Lutz is also redescribed and a key, table with numerical characters for
the identification and map with distribution of these related species is included.

Material and methods

Individual studied specimens are deposited in the following collections:

CCER Collection of Ceratopogonidae, Instituto Oswaldo Cruz, Rio de Janeiro, Brazil.

FSP Entomological Collection, Faculdade de Saúde Pública, USP, São Paulo, Brazil.
ILMD Laboratory of Biodiversity in Health, Instituto Leônidas e Maria Deane, Manaus, Amazonas, Brazil.
MLP Entomological Collection, Museu de La Plata, La Plata, Argentina.
MPEG Invertebrate Collection, Museu Paraense Emílio Goeldi, Pará, Brazil.

The slide mounted specimens deposited in the above collections was take on loan for study. Diagnostic
characters were illustrated or microphotographed using a Nikon Eclipse E 200 and a Zeiss AX10 miscroscope, and
the plates were prepared using GIMP Portable 2.6. The general terminology used is that employed for Culicoides
by Felippe-Bauer (2003). Terms for structures follow the Manual of Central America Diptera (Brown et al., 2009).
The measurements of structures are included in the text, those of the spermathecae are in micrometers, while those
of the wings are in millimeters. Meristic information is given as range, following by the mean and number of
specimens examined.

Results

While studying material identified as C. reticulatus Lutz from diverse localities, we observed differences in the
taxonomic parameters actually used in Culicoides. The only female specimen studied by Forattini & Galvão (1955)
from São Vicente has difference in the number of palpal pits (two well separate pits on left palpus and three on
right palpus). The presence of different number of pits in the third palpal segment in the same specimen is not
common and probably constitutes an anomaly as observed for other Culicoides species by Felippe-Bauer & Silva
(2006). The only female specimen from Caraguatatuba present PR=2.1, P/H ratio =1.25 and one oval palpal pit in
transverse position. The meristic data, the shape of the third palpal segment and the position of the palpal pit are
also different from the type series of C. reticulatus, and we don't consider this diverse specimen in our redescription
of C. reticulatus.
Unfortunately, Wirth & Blanton (1973) and Wirth et al. (1988) not cited the deposit house of the specimens
from Costa Rica and Honduras, studied by them. Asking the Instituto Nacional de Biodiversidad (InBIO Costa

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Rica), we were told that don’t exist material from C. reticulatus deposited there. As the latest revision of C.
reticulatus made by Wirth & Blanton (1959) was based on material from Panama, we are led to think that the
inclusion of specimens from Colombia and Costa Rica and the ones from Honduras are based on this revision.
The description of C. reticulatus in Barbosa (1943) based on material from Pernambuco is identical to our
observations of Lutz's material. The study of specimens identified by C. reticulatus of other localities showed
significantly differences from specimens type and represents seven new species. Following the conclusion reached
in our study C. reticulatus was characterized based on type material from Bahia and the ones from Pernambuco
described by Barbosa.

Key to the seven new species of reticulatus group with similar wing pattern to C. reticulatus Lutz
(primarily for females)

1. Third palpal segment with multiple shallow, irregular sensory pits on ventral, dorsal and lateral sides of segment (Fig.5F); pro-
boscis moderately long, P/H ratio 0.80–0.94; unequal sized spermathecae (Fig.5E) . . . . . . . . . . . . . . . . . C. irregularis sp. nov.
- Third palpal segment with one or two sensory pits; proboscis and spermathecae various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Third palpal segment with two well defined sensory pits on distal 2/3 of segment, the basal one on ventral side and the distal
one on lateral side (Fig.3D); mandible with 21 teeth; [male tergite 9 with very short and blunt apicolateral processes, with dis-
tinctly posteromedial notch; parameres ending in a ventral lobe; basal arch of aedeagus extending to 2/3 of total length
(Fig.3G)] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. diplus sp. nov.
- Third palpal segment with one sensory pit on ventral side; mandible teeth various . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Sensory pit deep, round and opening in a small pore (Fig.6E); proboscis moderately long, P/H ratio 0.86–0.95; mandible with
16 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. profundus sp. nov.
- Sensory pit shallow; proboscis long; P/H ratio 1.0–1.28; mandible with 19–25 teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Large species, wing length 1.30 mm; eyes separated by ½ ommatidium (Fig. 1C); [parameres sinuous without ventral lobe
(Fig. 1K); basal arch of aedeagus extending to ½ of total length (Fig. 1J)] . . . . . . . . . . . . . . . . . . . . . . . . . . C. reticulatus Lutz
- Smaller species, wing length 0.80–1.16; eyes separated by < ½ ommatidium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Third palpal segment rhomboidal, greatly swollen at middle length; sensory pit irregular occupying the distal ½ of segment
(Fig.8F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. rhombus sp. nov.
- Third palpal segment not rhomboidal, slightly swollen; sensory pit regular, round occupying the middle portion of segment . .
................................................................................................... 6
6. Wing length 1.03–1.16; a single pale spot over r-m crossvein (Fig.4A) . . . . . . . . . . . . . . . . . . . . . . . . . C. fluminensis sp. nov.
- Wing length 0.80–0.95; pale spot over r-m cross vein subdivided . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Sensory pit small and round, with no well-defined contours on distal ½ (Fig. 7E); P.R. 2.0–2.7; mandible with 19 teeth; two
slightly unequal ovoid spermathecae (Fig. 7F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . C. pseudoreticulatus sp. nov.
- Sensory pit large and oval, with defined contours (Fig.2G); P.R. 2.6–3.2; mandible with 24 teeth; two unequal ovoid spermath-
ecae (Fig.2E) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .C. amazonicus sp. nov.

Culicoides reticulatus Lutz, redescription

Figs. 1A–K; Fig. 9

Culicoides reticulatus Lutz, 1913: 49 (female; Brazil - Rio de Janeiro, São Paulo and Bahia; fig. wing, male, larva; bionomy,
larva in crab holes in mangrove edge; in part material from Bahia); Rieth, 1915: 417 (bionomics data, based on Lutz
1913); Costa Lima, 1937: 413 (key); Barbosa, 1943: 261 (male, female; Brazil - Pernambuco; figs. wing, palpus, male
terminalia); Vargas, 1945: 43 (in list); Barbosa, 1947: 5, 25 (key, distrib.); Macfie, 1948: 73 (key); Vargas, 1949: 205 (in
list); Ortiz, 1950: 464 (distrib.); Iriarte, 1950: 398 (in list); Fox, 1955: 254 (in list); Forattini, 1957: 432 (in neotropical
catalog; figs.; in part specimens from Bahia and Pernambuco); Spinelli & Wirth, 1986: 56 (key, fig. wing); Wirth et al.,
1988: 42 (in Atlas; fig. wing; distrib.; in part material from Brazil, not Pará); Borkent & Wirth, 1997: 80 (in catalog);
Borkent & Spinelli, 2000: 41 (in catalog; distrib.; in part material from Bahia and Pernambuco); Borkent & Spinelli, 2007:
74 (in catalog; distrib. in part material from Bahia and Pernambuco); Borkent, 2014: 99 (in catalog).

Diagnosis. Female: This species is distinguished by the following combination of characters: large sized species;
eyes separated by diameter of ½ ommatidium; palpus with shallow, rounded sensory pit in middle portion; PR
2.6–3.0 (n=6); proboscis long. Male: parameres stem, slightly sinuous in median portion, without ventral lobe;
basal arch of aedeagus extending ½ of total length.
Female. Head. Brown. Eyes bare (Fig.1C), separated by a distance equal to a diameter of ½ ommatidium.
Pedicel brown, flagellum (Fig.1D) pale brown; AR 1.0–1.4 (1.02, n=5); sensilla coeloconica on flagellomeres

NEW CULICOIDES AND C. RETICULATUS LUTZ Zootaxa 3795 (3) © 2014 Magnolia Press · 257
1,6–8. Palpus (Fig.1F) brown; 3rd segment fusiform, slightly swollen in middle length, with a shallow, rounded
sensory pit in middle portion; PR 2.6–3.0 (2.8, n=6). Proboscis long; P/H ratio 1.14–1.17 (1.16, n=4); mandible
with 23–25 (n=4) teeth.

FIGURE 1. Culicoides reticulatus Lutz. Female: A. Wing; C. Eyes separation, anterior view; D. Antenna; E. Spermathecae; F.
Palpus; G. Femora and tibiae of fore-, mid- and hind legs (left to right), lateral view. Male: B. Wing; H. Palpus; I. Terminalia; J.
Aedeagus; K. Parameres.

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 Thorax. Dark brown. Scutum without distinct pattern in slide mounted specimens. Wing (Fig.1A) with
contrasting pattern: second radial cell including dark spot; pale spot over R-M extending from M1 to costal margin,
subdivided; r3 with four small and separated pale spots: first, rounded, between the second radial cell and M1;
second, poststigmatic extending behind second radial cell, abutting wing margin; third, in the middle of cell,
rounded, smaller than second one; fourth, distal pale spot slightly reaching wing margin; m1 with two pale spots, the
distal one far from wing margin and greater than the proximal one; m2 with four pale spots: one proximal, near
CuA, two between the medial and mediocubital forks and a distal one not reaching the wing margin; cua1 with a
rounded pale spot in the middle of cell; anal cell with a faint basal pale area and one distal pale spot near
mediocubital fork; wing base with a faint pale spot on M; M1, M2 and CuA1 with pale apices; macrotrichia scarcely
distributed on distal half of wing; wing length 1.30 (n=4) mm; breadth 0.57–0.59 (0.55, n=4) mm; CR 0.64 (n=4).
Halter knob brown, stem pale. Legs (Fig.1G) mostly brown; femora with subapical pale bands, tibiae with subbasal
pale bands; hind tibia pale apically; hind tibial comb with four spines, the one nearest the spur longest.
Abdomen. Brown. Two subequal ovoid spermathecae (Fig. 1E), measuring 46.4 µm (n=5) by 32.8 µm (n=4)
and 43.5 µm (n=4) by 31 µm (n=2). Rudimentary third spermatheca present.
Male. Similar to female with usual sexual differences. AR 0.93 (n=1). Palpus (Fig.1H) with 3rd segment short,
two round sensory pits; PR1.6 (n=1). Wing length 1.03 (n=1) mm, breadth 0.43 (n=1) mm, CR 0.63 (n=1), pattern
of pale spots as in figure 1B. Terminalia (Fig.1I): tergite 9 not observable in slide mounted specimen; sternite 9
straight on distal margin; gonocoxite 2x longer than greatest breadth, ventral and dorsal root slender, elongated;
gonostylus tapering distally, distal portion slightly curved. Parameres (Fig.1K) separated, each with a sclerotized
basal knob; stem long, curved near base and slightly sinuous in median portion, without ventral lobe; apical portion
tapered, abruptly bent, without lateral fringe of spicules. Aedeagus (Fig.1J) Y-shaped, lateral arms strongly
sclerotized, basal arch triangular extending ½ of total length; distal portion slender, tapering to blunt tip without
median or distal processes.
Specimens examined. Type series, 1 male, 7 females on one microscope slide, labeled "Culicoides reticulatus
Lutz, 1913, Bahia, A. Lutz prep." (CCER n.2983- Costa Lima collection).
Distribution and bionomics. Culicoides reticulatus is now restricted to Brazilian states of Bahia and
Pernambuco (Fig. 9). It is associated to mangrove areas from these states and cited by Lutz (1913) biting human,
horses and cow in the vicinity of these areas.
Discussion. Forattini et al (1960) captured larva and pupa of Culicoides in mangrove areas and described them
as C. reticulatus without correlation with adult stage. Due to incertitude if this description is related to C.
reticulatus Lutz, we don't consider it herein.
In this study, all material identified as C. reticulatus from the Amazon region (Amazonas, Pará and Roraima
States) present important differences from C. reticulatus Lutz and, constitutes five new species as described below.
Then, we think that the bionomic data cited by Veras & Castellón (1998) for C. reticulatus from Amazonas State,
as well as the records published by Wirth & Blanton (1973) and Castellón et al (1993) for this species in Amazonas
and Para states, cannot be considered due to the misidentification of these species.

Culicoides amazonicus Santarém, Felippe-Bauer & Trindade sp. nov.

Figs. 2A–G; Fig. 9

Diagnosis. Female: This species is easily distinguished by the following combination of characters: palpus 3rd
segment swollen, with large, shallow, oval sensory pit on middle of segment; PR 2.6–3.2; long proboscis; mandible
with 24 teeth; two unequal spermathecae.
Female. Head. Brown. Eyes bare (Fig.2B), narrowly separated, smaller than ½ ommatidium. Pedicel brown,
flagellum pale brown, first eight flagellomeres paler on proximal ½; AR 0.98–1.04 (1.0, n=7) (Fig.2C); sensilla
coeloconica on flagellomeres 1,6–8, two on 1, 6–7 and three on 8. Palpus (Fig.2G) brown; 3rd segment swollen,
with a large, shallow, oval sensory pit in middle of segment; PR 2.6–3.2 (2.8, n=10). Proboscis long; P/H ratio
1.03–1.16 (1.11, n=11); mandible with 24 teeth.
Thorax. Dark brown. Prominent pattern of yellowish patches well defined (Fig. 2F); pale humeral depressions
present. Scutum with two pale areas in the anterior submedian portion arranged as leaf clover and two pale areas in
posterior submedian portion; lateral portion, with anterior and posterior pale areas; pale prescutellar depressions

NEW CULICOIDES AND C. RETICULATUS LUTZ Zootaxa 3795 (3) © 2014 Magnolia Press · 259
FIGURE 2. Culicoides amazonicus sp. nov., female. A. Wing; B. Eyes separation, anterior view; C. Antenna; D. Fore-, mid-
and hind legs (left to right); E. Spermathecae; F. Scutum; G. Palpus.

present; scutellum yellowish laterally; postscutellum brown. Wing (Fig. 2A) with contrasting pattern: second radial
cell including dark spot; pale spot over R-M extending from M1 to the costal margin, subdivided; r3 with four ,
small and separated pale spots: first, rounded, between the second radial cell and M1; second, poststigmatic,
extendind behind second radial cell, abutting wing margin; third, rounded, in the middle of cell similar to the
second one; fourth, distal pale spot slightly reaching wing margin; m1 with two pale spots, the distal one far from

260 · Zootaxa 3795 (3) © 2014 Magnolia Press SANTARÉM ET AL.

wing margin and greater than proximal one; m2 with four pale spots: one proximal, near CuA, two between the
medial and mediocubital forks, a distal one not reaching wing margin; cua1 with a rounded and small pale spot in
the middle of cell; anal cell with a faint, basal pale area, one distal pale spot near mediocubital fork and one very
faint pale spot reaching wing margin, not present in some specimens; wing base with a pale spot on M; M1, M2 and
CuA1 with pale apex; macrotrichia scarcely distributed; wing length 0.87–0.95 (0.92, n=10) mm; breadth 0.42–0.47
(0.46, n=10) mm; CR 0.62–0.66 (0.64, n=10). Halter knob brown laterally, stem pale. Legs (Fig.2D) mostly brown;
femora with subapical pale bands, tibiae with subbasal pale bands; hind tibia pale apically; hind tibial comb with
four spines, the one nearest the spur longest.
Abdomen. Brown. Two unequal ovoid spermathecae (Fig.2E), measuring 49.7 µm by 35.2 µm (n=10) and 38.2
µm by 28.1 µm (n=9), with sclerotized ducts with 7.5 µm (n=9). Long, slender, rudimentary third spermatheca
present (22.5 µm).
Male. Unknown.
Type data and depository. Holotype female, BRAZIL, Pará, Juruti, “Fazenda Rezende, Comunidade do Café
Torrado ", 15-16.XII.2007, Trindade, Guimarães & Quarema cols. (MPEG); Paratypes 10 females, same data
except: two, 17-18.XII.2007 (CCER); two, "curral", 06-07.IV.2008 (MPEG, MNRJ); one "mata", 20-21.VI.2008
(CCER); two, 26-27.VI.2008 (MPEG); three, Pará, Paragominas "Fazenda Dr. Jorge Artur, próximo ao lago", 05-
06.XII.2008 (2 CCER; MPEG).
Distribution and bionomics. This species is restricted to Brazilian state of Pará (Fig. 9). It is associated with
forest areas in farms.
Etymology. This species is named in tribute of the Brazilian Amazon Region, where the specimens were
collected.

Culicoides diplus Santarém & Felippe-Bauer sp. nov.

Figs. 3A–H; Fig. 9

Culicoides reticulatus: Wirth & Blanton, 1959: 393; Wirth & Blanton, 1973: 446 (distrib.; in part, at least specimens from
Colombia and Panama); Wirth et al., 1988: 42 (in Atlas; fig. wing; distrib. in part, at least specimens from Colombia and
Panama); Borkent & Spinelli, 2007: 74 (in catalog; distrib. in part, at least specimens from Colombia and Panama).

Diagnosis. Female: This species can be easily distinguished by 3rd segment of palpus with two well defined
sensory pits on distal 2/3 of the segment, the basal one ventrally and the distal one laterally; PR 2.5-2.8. Male:
tergite 9 long, tapering, with very short and blunt apicolateral processes, with distinctly posteromedial notch;
parameres stem sinuous and swollen in median portion ending in a ventral lobe; basal arch of aedeagus extending
2/3 of total length.
Female. Head. Brown. Eyes bare (Fig. 3E), narrowly separated, smaller than ½ ommatidium. Pedicel brown,
flagellum pale brown, first eight flagellomeres paler on ½ proximal; AR 1.04–1.13 (1.08, n=2) (Fig. 3C); sensilla
coeloconica on flagellomeres 1, 6-8, two on 1, three on 6-8. Palpus (Fig.3D) brown; 3rd segment with two well
defined sensory pits on distal 2/3 of the segment, the basal one ventrally and the distal one laterally; PR 2.5–2.8
(2.7, n=4). Proboscis long; P/H ratio 1.0–1.05 (1.02, n=2); mandible with 21 teeth (n=2).
Thorax. Dark brown. Scutum without distinct pattern in slide mounted specimens. Wing (Fig.3A) with
contrasting pattern: second radial cell including dark spot; pale spot over R-M extending from M1 to the costal
margin, subdivided; r3 with four small and separated pale spots: first, rounded, between the second radial cell and
M1; second, poststigmatic, extending behind second radial cell, abutting wing margin; third, rounded, in middle of
cell, similar to the second one; fourth, distal pale spot slightly reaching wing margin; m1 with two pale spots, the
distal one far from wing margin and greater than the proximal one; m2 with four pale spots: one proximal, near
CuA, two between the medial and mediocubital forks, a distal one not reaching wing margin; cua1 with a rounded
pale spot in the middle of cell; anal cell with a faint, basal pale area and two distal pale spots, one near mediocubital
fork and other reaching wing margin; wing base with a pale spot on M; M1, M2 and CuA1 with pale apex;
macrotrichia scarcely distributed on the distal third of wing; wing length 0.94–0.98 (0.96, n=3) mm; breadth
0.46–0.50 (0.47, n=3) mm; CR 0.66–0.67 (0.66 n=3). Halter knob brown in middle, stem pale. Legs (Fig. 3H)
mostly brown; femora with subapical pale bands, tibiae with subbasal pale bands; hind tibia pale apically; hind
tibial comb with four spines, the one nearest the spur longest.

NEW CULICOIDES AND C. RETICULATUS LUTZ Zootaxa 3795 (3) © 2014 Magnolia Press · 261
FIGURE 3. Culicoides diplus sp. nov. Female: A. Wing; C Antenna; D. Palpus; E. Eyes separation, anterior view; F.
Spermathecae; H. Femora and tibiae of fore-, mid- and hind legs (left to right). Male: B. Wing; G. Terminalia.

Abdomen. Brown. Two subequal ovoid spermathecae (Fig. 3F), measuring 48 µm (n= 2) by 41.3 µm (n=2) and
45 µm (n=2) by 37 µm (n=2); long, slender, rudimentary third spermatheca present (15 µm) in some specimens.
Male. Similar to female with usual sexual differences. Sensilla coeloconica on flagellomeres 1,7–10, one on
1,7–8, two on 9, three on 10; AR 0.84 (n=1); PR 1.5 (n=1); wing length 0.80-0.85 (0.83, n=2) mm, breadth 0.35-
0.38 (0.36, n=2) mm, CR 0.61 (n=2), pigmentation pattern as in figure 3B. Terminalia (Fig.3G) tergite 9 long,
tapering, with very short and blunt apicolateral processes, with distinctly posteromedial notch; sternite 9 straight on

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distal margin. Gonocoxite two times longer than greatest breadth, ventral and dorsal root slender, elongated;
gonostylus with ½ distal portion slightly curved. Parameres separated, each with a sclerotized basal knob; stem
long, curved near base and sinuous and swollen in median portion ending in a ventral lobe; apical portion tapered,
abruptly bent, without lateral fringe of spicules. Aedeagus Y-shaped, lateral arms strongly sclerotized, basal arch
triangular extending 2/3 of total length; distal portion rounded without median or distal processes.
Type data and depository. Holotype female, COLOMBIA, Valle Rio Raposo, II.1965, light trap, Lee col.,
(CCER). Allotype male, Garachine RP, PANAMA, 24.II.1953, FS Blanton col. (FSP). Paratypes, five as follows:
two females, same data except V.1965 (CCER; MLP); one female, same data as holotype except VIII.1965 (MLP);
one female, PANAMA, Mojinga Swamp, Ft Sherman, 28.VIII.1951, FS Blanton col. (FSP); one male, same data as
allotype (FSP).
Distribution and bionomics. This species is restricted to Colombia and Panama (Fig. 9). It is associated with
mangrove areas in these countries.
Etymology. The name of this species is based on the presence of two palpal pit on third palpal segment
(Greek—diplóos = two).

Culicoides fluminensis Santarém & Felippe-Bauer, sp. nov.

Figs. 4A–D

Culicoides reticulatus: Lutz, 1913: 49 (female; Brazil - Rio de Janeiro, São Paulo and Bahia; fig. wing, male, larva; bionomy,
larva in crab holes in mangrove edge; in part material from Rio de Janeiro); Forattini, 1957: 432 (in neotropical catalog;
figs.; in part specimens from Rio de Janeiro); Wirth & Blanton, 1973: 446 (distrib.in part material from Rio de Janeiro);
Borkent & Spinelli, 2000: 41 (in catalog; distrib.; in part material from Rio de Janeiro); Borkent & Spinelli, 2007: 74 (in
catalog; distrib. in part material from Rio de Janeiro).

Diagnosis. Female: This species is easily distinguished by the following combination of characters: large sized
species; 3rd segment of palpus with a small, shallow, rounded, sensory pit in middle portion; PR 2.6–3.2; long
proboscis; single pale spot over R-M; anal cell with two distal pale spots, one near mediocubital fork and other not
reaching wing margin; two unequal spermathecae.
Female. Head. Brown. Eyes bare (Fig.4B), narrowly separated, smaller than ½ ommatidium. Pedicel brown,
flagellum pale brown, first eight flagellomeres paler on proximal ½; AR 0.95–1.12 (1.03, n=6); sensilla
coeloconica on flagellomeres 1,6–8, two on all flagellomeres. Palpus (Fig. 4C) brown; 3rd segment slightly
swollen with a small, shallow, rounded sensory pit in middle portion; PR 2.6–3.2 (2.9, n=4). Proboscis long; P/H
ratio 1.05–1.18 (1.14, n=9); mandible with 20 teeth.
Thorax. Dark brown. Prominent pattern of yellowish patches well defined; pale humeral depressions present.
Scutum with two pale areas in the anterior submedian portion arranged as leaf clover and two pale areas in
posterior submedian portion; lateral portion with anterior and posterior pale areas; pale prescutellar depressions
present; scutellum yellowish laterally; postscutellum brown. Wing (Fig.4A) with contrasting pattern: second radial
cell including dark spot; one pale spot over R-M extending from M1 to the costal margin, slightly subdivided in
some specimens; r3 with four small and separated pale spots: first, oval, between the second radial cell and M1;
second, poststigmatic, extending behind second radial cell, abutting wing margin; third, rounded, in middle of cell,
smaller than others; fourth, distal pale spot slightly reaching wing margin; m1 with two pale spots, the distal one far
from wing margin and greater than proximal one; m2 with four pale spots: one proximal, near CuA, two between
the medial and mediocubital forks, a distal one, small, not reaching wing margin; cua1 with a rounded pale spot in
the middle of cell; anal cell with a faint, basal pale area and two distal pale spots, one near mediocubital fork and
other, faint, not reaching wing margin; wing base with a pale spot on M; M1, M2 and CuA1 with pale apex;
macrotrichia scarcely distributed on the distal third of wing; wing length 1.03–1.16 (1.08, n=10) mm; breadth
0.49–0.54 (0.51, n=10) mm; CR 0.63–0.66 (0.65, n=10). Halter knob brown in middle, stem pale. Legs (Fig.4D)
mostly brown; femora with subapical pale bands, tibiae with subbasal pale bands; hind tibia pale apically; hind
tibial comb with four spines, the one nearest the spur longest.
Abdomen. Brown. Two unequal ovoid spermathecae in bad condition in slide mounted specimens. Long,
swollen, rudimentary third spermatheca present (12.5 µm).

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FIGURE 4. Culicoides fluminensis sp. nov., female. A. Wing; B. Eyes separation, anterior view; C. Palpus; D. Femora and
tibiae of fore-, mid- and hind legs (left to right).

Male. Unknown.
Type data and depository. Holotype female, BRAZIL, Rio de Janeiro, Macaé, Barra de Macaé, R. Idelbrando
Alves Barbosa, 22°20'S 41°45'O, 29.III.1989, New Jersey trap, FEEMA col. (CCER). Paratypes nine females as
follows: six, same data as holotype (CCER); three, same data as holotype, except 25.IV.1989 (2 CCER; MNRJ).
Distribution and bionomics. This species is restricted to Brazilian state of Rio de Janeiro (Fig. 9). It was
collected near mangrove areas from the municipality of Macaé.
Etymology. This species is named after the originaries of the state of Rio de Janeiro, where the specimens
were collected.

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Culicoides irregularis Santarém, Felippe-Bauer & Castellón sp. nov.
Figs. 5A–F; Fig. 9

Diagnosis. Female: This species is easily distinguished by the following combination of characters: palpus 3rd
segment with a multiple, shallow, irregular sensory pit; PR 2.0–2.9; moderately long proboscis; mandible with 16
teeth; two unequal ovoid spermathecae.

FIGURE 5. Culicoides irregularis sp. nov., female. A. Wing; B. Eyes separation, anterior view; C. Antenna; D. Fore-, mid-
and hind legs (left to right); E. Spermathecae; F. Palpus.

NEW CULICOIDES AND C. RETICULATUS LUTZ Zootaxa 3795 (3) © 2014 Magnolia Press · 265
 Female. Head. Brown. Eyes bare (Fig. 5B), narrowly separated, smaller than ½ ommatidium. Pedicel brown,
flagellum pale brown, first eight flagellomeres paler on proximal ½; AR 1.04–1.25 (1.12, n= 24) (Fig. 5C); sensilla
coeloconica on flagellomeres 1,6–8; two on 1, three on 6–7 and three - four on 8. Palpus (Fig. 5F) brown, 3rd
segment swollen with a multiple, shallow, irregular sensory pit in ventral, dorsal and lateral side of segment; PR
2.0–2.9 (2.6, n=31). Proboscis moderately long; P/H ratio 0.80–0.94 (0.84, n=28); mandible with 16 teeth.
Thorax. Dark brown. Scutum without distinct pattern in slide mounted specimens. Wing (Fig. 5A) with
contrasting pattern: second radial cell including dark spot, pale spot over R-M extending from M1 to the costal
margin, subdivided; r3 with four small and separated pale spots: first, rounded, between the second radial cell and
M1; second, poststigmatic, extending behind second radial cell, abutting wing margin; third, rounded, in middle of
cell; fourth, distal pale spot slightly reaching wing margin; m1 with two pale spots, the distal one far from wing
margin; m2 with four pale spots: one proximal, two between the medial and mediocubital forks, a distal one not
reaching wing margin; cua1 with a rounded pale spot in the middle of cell; anal cell with a basal pale area and a pale
spot near mediocubital forks; wing base with a pale spot on M; M1,M2 and CuA1 with pale apex; macrotrichia
scarcely distributed on the distal third of wing; wing length 0.85–1.05 (0.98, n=31) mm; breadth 0.37–0.47 (0.44,
n=31) mm; CR 0.58–0.68 (0.65, n=31). Halter knob brown in middle, stem pale. Legs (Fig. 5D) mostly brown;
femora with subapical pale bands, tibiae with subbasal pale bands; hind tibia pale apically; hind tibial comb with
four spines, the one nearest the spur longest.
Abdomen. Brown. Two unequal ovoid spermathecae (Fig. 5E), measuring 48.2 µm (n=25) by 37.5 µm (n=25)
and 39.5 µm (n=23) by 33 µm (n=23), with sclerotized ducts measuring 7.5 µm (n=20); long, slender, rudimentary
third spermatheca present (17.2 µm); slightly sclerotized ring present (7.5 µm).
Male. Unknown.
Type data and depository. Holotype female, BRAZIL, Roraima, Maracá, 26.VI.1987, CDC light trap, "mata
1m", E. Castellon & S. Gomes cols. (CCER); paratypes 32 females, same data except: three (2 CCER; ILMD);
three, "mata 10 m" (2 CCER; MNRJ); three, 15.VIII.1987, "mata 5m" (2 ILMD; CCER); one, 30.VII. 1987 "mata
10m" (CCER); one, "mata 5m" (CCER); one, 24.VI.1987 (ILMD); two, 24.VII.1987 "mata 5m" (CCER; ILMD);
one, 24.V.1987 "mata 5m" (ILMD); one, 25.VI.1987 "mata 5m" (CCER); one, 25.VII.1987 (ILMD); one, 25.VI.
1987 "mata 10m" (ILMD); one, 21.V.1987 (CCER); one, 17.VI.1987 (ILMD); one, 16.VI.1987 (CCER); one,
23.VI.1987 "mata 5m" (CCER); one, 23.VII.1987 "mata 5m" (ILMD); one, 27.VII.1987 (CCER); one, 27.V.1987
"mata 5m" (ILMD); one, 27.V.1987 "mata 10m" (ILMD); one, 28.VII.1987 (CCER); two, 22.VI. 1987 "mata 10m"
(ILMD); one, 31.VIII.1987 (ILMD); one, 31.VIII.1987 "mata alagada" (ILMD); one, Rio Pardo, Amazonas, 21-
22.VI.2010, "peridomicilio", CDC light trap, F. Pessoa col. (ILMD).
Distribution and bionomics. This species is restricted to Brazilian states of Roraima and Amazonas (Fig.9). It
was associated with forest environments far from coastal regions.
Etymology. The name of this species is related with the irregular sensory pit present in third palpal segment
(Latim—irregulare = irregular).

Culicoides profundus Santarém, Felippe-Bauer & Trindade sp. nov.

Figs. 6A–G; Fig. 9

Diagnosis. Female: This species is easily distinguished by the following combination of characters: palpus 3rd
segment swollen, with large, deep sensory pit in middle portion, opening in a small, rounded pore; PR 1.8–2.4;
moderately long proboscis; mandible with 16 teeth.
Female. Head. Brown. Eyes bare (Fig. 6G), narrowly separated, smaller than ½ ommatidium. Pedicel brown,
flagellum pale brown, first eight flagellomeres paler on proximal ½; AR 1.02–1.18 (1.10, n=19) (Fig. 6C); sensilla
coeloconica on flagellomeres 1,(5) 6–8, two on 1, one or two on 5, three on 6 and 7 and four on 8. Palpus (Fig. 6E)
brown; 3rd segment swollen, with a large, deep sensory pit in middle portion, opening in a small, rounded pore; PR
1.8–2.4 (2.1, n=25). Proboscis moderately long; P/H ratio 0.86–0.95 (0.90, n=25); mandible with 16 teeth (n=2).

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FIGURE 6. Culicoides profundus sp. nov., female. A. Wing; B. Scutum; C. Antenna; D. Fore-, mid- and hind legs (left to
right); E. Palpus; F. Spermathecae; G. Eyes separation, anterior view.

Thorax. Dark brown. Prominent pattern of yellowish patches well defined (Fig. 6B); pale humeral depressions
present. Scutum with two pale areas in the anterior submedian portion arranged as leaf clover and two pale areas in
posterior submedian portion; lateral portion with anterior and posterior pale areas; pale prescutellar depressions
present; scutellum yellowish laterally; postscutellum brown. Wing (Fig. 6A) with contrasting pattern: second radial
cell including dark spot; pale spot over R-M extending from M1 to the costal margin, subdivided; r3 with four, small

NEW CULICOIDES AND C. RETICULATUS LUTZ Zootaxa 3795 (3) © 2014 Magnolia Press · 267
and separated pale spots: first, irregular, elongate, transverse, S shaped in some specimens, between the second
radial cell and M1; second, poststigmatic, extending behind second radial cell, abutting wing margin; third, nearly
rounded, in the middle of cell similar to the second one; fourth, distal pale spot lightly reaching wing margin,
sometimes connected with third one; m1 with two pale spots, the distal one far from wing margin; m2 with four pale
spots: one proximal, near CuA, faint and small, two between the medial and mediocubital forks, a distal one not
reaching wing margin; cua1 with a rounded and small pale spot in the middle of cell; anal cell with a faint, basal
pale area and one distal pale spot near mediocubital fork; wing base with a pale spot on M; M1, M2 and CuA1 with
pale apex; macrotrichia scarcely distributed on distal third of wing; wing length 0.82–0.97 (0.93, n=26) mm;
breadth 0.40–0.50 (0.46, n=26) mm; CR 0.60–0.67 (0.64, n=26). Halter knob brown in middle, in distal ½ in some
specimens, stem pale. Legs (Fig. 6D) mostly brown; femora with subapical pale bands, tibiae with subbasal pale
bands; hind tibia pale apically; hind tibial comb with four spines, the one nearest the spur longest.
Abdomen. Brown. Two slightly unequal ovoid spermathecae (Fig. 6F), measuring 42 µm by 35,7 µm (n=25)
and 36 µm by 30,5 µm (n=22), with sclerotized ducts with 6.25 µm (n=16). Long, slender, rudimentary third
spermatheca present (22,5 µm); short, cylindrical sclerotized ring present (5 µm).
Types data and depository. Holotype female, BRAZIL, Vila de Patauateua, Pará, Ourém, "Fazenda Gavião
Real", 01o29'44" S 47o13'07" W, 28-29.XI.2008, CDC light trap, forest, Trindade & Guimarães cols. (MPEG);
Paratypes, 25 females, same data, except: one, Juruti, " Fazenda Rezende, Comunidade do Café Torrado",
02o18'77"S 56o05'24"W, 17-18.XII.2007 (CCER); one, 26-27.VI.2008 (MPEG); one, Paragominas, "Fazenda do
Dr. Jorge Artur, próximo ao igarapé da casa do caseiro", 03-04.XII.2008, Trindade & Quaresma cols. (MPEG); six,
04-05.XII.2008 (4 MPEG; 1 CCER; 1 MNRJ); seven, 02-03.XII. 2008 (4 CCER; 3 MPEG); seven, Amazonas, Rio
Pardo, VI-VIII.2010, CDC light trap, "borda de floresta", F. Pessoa col. (ILMD); one, same data except 17-
18.VII.2010, "peridomicilio" (ILMD); one, same data except 21-22.VI.2010, "floresta" (ILMD).
Male. Unknown.
Distribution and bionomics. This species is restricted to Brazilian states of Pará and Amazonas (Fig. 9). It
was associated with farm environments in three municipalities of Pará State and with forest areas in Amazonas
State, both far from coastal regions.
Etymology. The name of this species is related with the presence of deep sensory pit on third palpal segment
(Latim—profundu = deep).

Culicoides pseudoreticulatus Santarém, Felippe-Bauer & Castellón sp. nov.

Figs. 7A–F; Fig. 9

Diagnosis. Female: This species is easily distinguished by the following combination of characters: palpus 3rd
segment with a shallow, round sensory pit in middle portion of segment, with no well-defined contours on distal ½;
PR 2.0–2.7; proboscis moderately long; mandible with 19 teeth.
Female. Head. Brown. Eyes bare, (Fig. 7B) narrowly separated, smaller than ½ ommatidium. Pedicel brown;
flagellum pale brown, first eight flagellomeres paler on proximal ½; AR 0.90–1.04 (0.98, n=4) (Fig. 7C); sensilla
coeloconica on flagellomeres 1, 6–8; two on 1, 6 and 7 and three on 8. Palpus (Fig. 7E) brown, 3rd segment slightly
swollen, with a shallow, rounded sensory pit in middle portion of segment, with no well-defined contours on distal
½; PR 2.0–2.7 (2.4, n=6). Proboscis moderately long; P/H ratio 1.0–1.06 (1.03, n=4); mandible with 19 teeth.
Thorax. Dark brown. Scutum without distinct pattern in slide mounted specimens. Wing (Fig. 7A) with
contrasting pattern: second radial cell including dark spot; pale spot over R-M extending from M1 to the costal
margin, subdivided; r3 with four small and separated pale spots; first, rounded, between the second radial cell and
M1; second, poststigmatic, extending behind second radial cell, abutting wing margin; third, rounded, in middle of
cell, similar to the second one; fourth, distal pale spot slightly reaching wing margin; m1 with two pale spots, the
distal one far from wing margin and greater than proximal one; m2 with four pale spots: one proximal, faint, near
CuA, two between the medial and mediocubital forks, a distal one not reaching wing margin; cua1 with a rounded
pale spot in the middle of cell; anal cell with a faint basal pale area and a distal pale spot near mediocubital fork;
wing base with a pale spot on M; M1, M2 and CuA1 with pale apex; macrotrichia scarcely distributed on the distal
third of wing; wing length 0.80–0.90 (0.86, n=7) mm; breadth 0.35–0.45 (0.42, n=7) mm; CR 0.61–0.66 (0.63,

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n=7). Halter knob brown in middle, stem pale. Legs (Fig.7D) mostly brown; femora with subapical pale bands,
tibiae with subbasal pale bands; hind tibia pale apically; hind tibial comb with four spines, the one nearest the spur
longest.
Abdomen. Brown. Two slightly unequal ovoid spermathecae (Fig. 7F), measuring 49.5 µm (n=5) by 35 µm
(n=5) and 43.2 µm (n=3) by 35 µm (n=3), with sclerotized ducts measuring 6.25 µm (n=3); long, slender,
rudimentary third spermatheca (15 µm) present in some specimens.

FIGURE 7. Culicoides pseudoreticulatus sp. nov., female. A. Wing; B. Eyes separation, anterior view; C. Antenna; D. Femora
and tibiae of fore-, mid- and hind legs (left to right); E. Palpus; F. Spermathecae.

NEW CULICOIDES AND C. RETICULATUS LUTZ Zootaxa 3795 (3) © 2014 Magnolia Press · 269
 Male. Unknown.
Type data and depository. Holotype female, BRAZIL, Roraima, Maracá, 28.VII.1987, CDC light trap, "mata
10m", E. Castellon & S. Gomes cols. (CCER); paratypes, six females, same data except: one, 23.VII.1987, "mata
5m" (CCER); two, 30.VII.1987 (MNRJ; ILMD); one, 24.VII.1987, "mata 5m" (ILMD); one, 31.VIII.1987, "mata
1m" (ILMD); one, Amazonas, Rio Pardo, 21-22.VI.2010, CDC light trap, "galinheiro", F. Pessoa col. (ILMD).
Distribution and bionomics. This species is restricted to Brazilian states of Roraima and Amazonas (Fig.9). It
was associated with forest environments far from coastal regions.
Etymology. The name of this species is related with its similar congener, C. reticulatus Lutz (Greek—pseudo
= false; Latim—reticulatus = reticulated).

Culicoides rhombus Santarém, Felippe-Bauer & Castellón sp. nov.

Figs. 8A–F; Fig. 9

Diagnosis. Female: This species is easily distinguished by the following combination of characters: palpus 3rd
segment rhomboidal, with a shallow, well - developed irregular sensory pit that reaches the extreme distal portion
of segment; PR 2.5–3.0; proboscis long.
Female. Head. Brown. Eyes bare (Fig. 8B), narrowly separated, smaller than ½ ommatidium. Pedicel brown,
flagellum pale brown, first eight flagellomeres paler on proximal ½; AR 1.05–1.10 (1.06, n=4) (Fig. 8C); sensilla
coeloconica on flagellomeres 1,6–8, two on 1, three on 6, two on 7 and three on 8. Palpus (Fig. 8F) brown, 3rd
segment swollen at middle length, rhomboid, with a shallow, well - developed irregular sensory pit that reaching
the distal portion of segment; PR 2.5–3.0 (2.6, n=7). Proboscis long; P/H ratio 1.08–1.28 (1.13, n=6); mandible
with 23 teeth.
Thorax. Dark brown. Scutum without distinct pattern in slide mounted specimens. Wing (Fig. 8A) with
contrasting pattern: second radial cell including dark spot; pale spot over R-M extending from M1 to the costal
margin, subdivided; r3 with four small and separated pale spots: first, rounded, between the second radial and M1;
second, poststigmatic, extending behind second radial cell, abutting wing margin; third, rounded, in middle of cell,
similar to the second one; fourth, distal pale spot slightly reaching wing margin; m1 with two pale spots, the distal
one far from the wing margin and greater to the proximal one; m2 with four pale spots: one proximal, near CuA,
two between medial and mediocubital forks, a distal one not reaching wing margin; cua1 with a rounded pale spot
in the middle of cell; anal cell with a faint basal pale area near base and a distal pale spot near mediocubital fork;
wing base with a pale spot on M; M1, M2 and CuA1 with pale apex; macrotrichia scarcely distributed on the distal
third of wing; wing length 0.85–0.97 (0.92, n=7) mm; breadth 0.40–0.45 (0.43, n=7) mm; CR 0.64–0.66 (0.65,
n=7). Halter knob browm, stem pale. Legs (Fig. 8D) mostly brown, femora with subapical pale bands, tibiae with
subbasal pale bands; hind tibia pale apically; hind tibial comb with four spines, the one nearest the spur longest.
Abdomen. Brown. Two slightly unequal ovoid spermathecae (Fig. 8E), measuring 45.5 µm (n=6) by 37 µm
(n=6) and 38.7 µm (n=5) by 32.5 µm (n=5), with sclerotized ducts measuring 4.3 µm (n=6); long, slender,
rudimentary third spermatheca present (17.5 µm).
Male. Unknown.
Type data and depository. Holotype female, BRAZIL, Roraima, Maracá, 28.VII.1987, CDC light trap,
"mata", E. Castellon & S. Gomes cols. (CCER). Paratypes six females, same data except: one, 28.VI.1987, "mata
10m" (CCER); one, 30.VII.1987, "mata 5m" (MNRJ); one, 22.VII.1987, "mata 5m” (ILMD); one, 27.VII.1987,
"mata 5m” (ILMD); one, Amazonas, Rio Pardo, 17-18.2010, CDC light trap, Capoeira, F. Pessoa col.; one same
data except VI-VIII.2010, "borda de floresta" (ILMD).
Distribution and bionomics. This species is restricted to Brazilian states of Roraima and Amazonas (Fig. 9).
It was associated with forest environment far from coastal regions.
Etymology. The name of this species is related with the rhomb shape aspect of the third palpal segment
(Greek—rhómbos= rhombus).

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FIGURE 8. Culicoides rhombus sp. nov., female. A. Wing; B. Eyes separation, anterior view; C. Antenna; D. Femora and
tibiae of fore-, mid- and hind legs (left to right); E. Spermathecae; F. Palpus.

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FIGURE 9. Geographical distribution of Culicoides reticulatus and the seven new related species.

TABLE 1. Table with numerical characters to C. reticulatus Lutz and the seven new related species.
Speciesa Wing length C.R. A.R. Pattern P.R. P/H ratio Mandible
sensilla teeth
C. amazonicus sp. nov. 0.92 mm 0.64 1.0 1,6–8 2.8 1.11 24
(0.87–0.95) (0.62–0.66) (0.98–1.04) (2.6–3.2) ( 1.03–1.16)
C. diplus sp. nov. 0.96 mm 0.66 1.08 1,6–8 2.7 1.02 21
(0.94–0.98) (0.66–0.67) (1.04–1.13) (2.5–2.8) (1.0–1.05)
C. fluminensis sp. nov. 1.08 mm 0.65 1.03 1,6–8 2.9 1.14 20
(1.03–1.16) (0.63–0.66) (0.95–1.12) (2.6–3.2) (1.05–1.18)
C. irregularis sp. nov. 0.98 mm 0.65 1.12 1,6–8 2.6 0.84 16
(0.85–1.05) (0.58–0.68) (1.04–1.25) (2.0–2.9) (0.80–0.94)
C. profundus sp. nov. 0.93 mm 0.64 1.10 1,(5) 2.1 0.90 16
(0.82–0.97) (0.60–0.67) (1.02–1.18) 6–8 (1.8–2.4) (0.86–0.95)
C. pseudoreticulatus 0.86 mm 0.63 0.98 1,6–8 2.4 1.03 19
sp. nov. (0.80–0.90) (0.61–0.66) (0.90–1.04) (2.0–2.7) (1.0–1.06)
C. reticulatus Lutz 1.3 mm 0.64 1.02 1,6–8 2.8 1.16 23–25
(1.0–1.04) (2.6–3.0) (1.14–1.17)
C. rhombus sp. nov. 0.92 mm 0.65 1.06 1,6–8 2.6 1.13 23
(0.85–0.97) (0.64–0.66) (1.05–1.1) (2.5–3.0) (1.08–1.28)

a: arranged alphabetically.

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Culicoides sp.

We examined a single specimen of Culicoides collected near a cave in a forest area from Amazonas State, which
has a similar wing pattern to C. reticulatus Lutz. It differ from this species and the other seven new species
described here by the aspect of the third palpal segment, which is cylindrical, without pits, with scattered sensilla
distributed on the segment, P.R. 3.0. These data led us to conclude that this specimen belong to a new species.
However, due to the bad montage condition, the specimen don't offer accurate data to make a description and we
think eventually describe it after collection of more specimens in this Amazon site.
Specimen examined: 1 female, BRAZIL, Amazonas, Balbina, 08.V.1985, CDC light trap, E Castellón & S
Gomes cols. (ILMD).

Taxonomic discussion

The new species described herein are similar to C. reticulatus by the number and distribution of the pale spots of
the wing, by the leaf clover pattern of yellowish patches on anterior submedian portion of scutum and by the
distribution pattern of sensilla coeloconica on flagellomeres. The notorious differences are the aspect of 3rd palpal
segment and the number, shape and position of the sensory pit, when present.
C. reticulatus is now restricted to Brazilian States of Bahia and Pernambuco, while the others seven new
species are described for Rio de Janeiro State (C. fluminensis), Brazilian Amazon Region (C. amazonicus, C.
irregularis, C. profundus, C. pseudoreticulatus and C. rhombus) and Colombia and Panama (C. diplus). Culicoides
amazonicus and C.profundus were found in sympatry in the municipalities of Juruti and Paragominas, Pará State.
Culicoides irregularis, C. profundus, C. pseudoreticulatus and C. rhombus occur simultaneously in Rio Pardo, in
the municipality of Presidente Figueiredo, Amazonas State. Also, C. amazonicus, C. pseudoreticulatus and C.
rhombus occur in Maraca, Roraima State. Sympatric species, with the similar pattern of wing spots and presenting
differences in the aspect of palpal pits, were observed for the first time in Neotropical Region, by Felippe-Bauer et
al. (2003) for species of the paraensis species group.
Culicoides diplus and C. fluminensis are collected in mangrove areas, the same biotope cited by Lutz (1913)
for C. reticulatus. Although, C. reticulatus has wing length (1.3 mm) bigger than C. fluminensis (1.08 mm) and C.
diplus (0.96 mm). Also, C. diplus has smaller P/H ratio (1.02) than C. reticulatus (1.16) and C. fluminensis (1.14).
The others five new species described from Amazon region are associated to farm and forest areas. This can
explain the difference in palpal pit, since this structure is known by the olfactory function in host seek (Blackwell,
2004).
Culicoides irregularis and C. profundus are the only species that has a moderately long proboscis (P/H ratio
<1.00) and a lower number of mandible teeth (16 teeth), which differs from C. reticulatus and the other five new
species that presents a long proboscis (P/H ratio ≥1.00) and a higher number of mandible teeth (20-25 teeth). Also,
the spermathecae are subequal in C. reticulatus and C. diplus, slightly unequal in C. profundus, C.
pseudoreticulatus and C. rhombus and unequal in C. amazonicus, C. fluminensis and C. irregularis. Culicoides
fluminensis is the only species that has a single pale spot over R-M, whereas in the other species this pale spot is
subdivided. All these species were included in reticulatus species group.

Acknowledgement

To Gustavo Spinelli for send us the data of specimens deposited in Museo de La Plata. To Maria Anice Mureb
Sallum (Faculdade Saúde Pública, USP), Felipe Pessôa (Instituto Leônidas e Maria Deane, FIOCRUZ) and
Orlando Tobias Silveira (Museu Paraense Emilio Goeldi) for the loan of part of material for study. To Coordenação
de Aperfeiçoamento de Pessoal de Ensino Superior (Capes) for financial support.

NEW CULICOIDES AND C. RETICULATUS LUTZ Zootaxa 3795 (3) © 2014 Magnolia Press · 273
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