Corresponding author: Santiago ZaragozaCaballero (
Academic editor: Hume Douglas
A taxonomic revision of the genus
VegaBadillo V, Morrone JJ, ZaragozaCaballero S (2021) Revision of the genus
The genus
Morphological structures in the male reproductive apparatus of
The aim of this study is to revise the species of
A total of 85 specimens for the analysis were borrowed from the following collections (acronyms follow the Insect and Spider Collections of the World website [
The material was determined by means of existing taxonomic keys (
The taxonomic treatment includes information on the type specimens and the material examined. In the Remarks section, we comment on the morphological similarities and dissimilarities between phylogenetically closed taxa (
Body 2.8–16.0 mm long; interantennal distance less than or equal to the scape length; antennae with 12 antennomeres, antennomeres 4 to 11 each with two symmetrical rami, 1.5 to 3 times longer than the respective antennomere; mandibles long, thin and crossed; maxillary palpi 4segmented, last segment securiform; labial palpi 2segmented; two separated tentorial pits and gula with two sutures; pronotum longer than wide, anterior edge rounded, sides almost straight; each elytron 2.8 to 5.4 times longer than wide, leaving the last 3 abdominal segments exposed; tarsomeres simple, without ventral combs; claws simple, without any teeth; wing with radial cell closed, RP reaching half or length less than half of MP1+2; and aedeagus with parameres slightly widened towards the midlength, apex toothed mesad.
Species of the genus
Species  Author  Country  Holotype 



Mexico 



Mexico 



USA 



Mexico 



USA 

This work  Mexico 




Mexico 


VegaBadillo et al. 2021  Mexico 



Mexico 



Mexico 


VegaBadillo et al. 2021  Mexico 



USA and Mexico 



Mexico  CUIC 


Mexico 



Mexico 


VegaBadillo et al. 2021  Mexico 


VegaBadillo et al. 2021  Mexico 



Mexico 



Costa Rica 



USA  NHMUK 


Mexico 



Mexico 

This work  Honduras 




Mexico 

This work  Belize 



VegaBadillo et al. 2021  Mexico 



Mexico 



Mexico 


VegaBadillo et al. 2021  Guatemala 

This work  Costa Rica 

Unknown.
1  Pronotum as long as wide; integument smooth 

–  Pronotum longer than wide; integument chagreend or smooth 

2  Interocular distance 1.5 times longer than eye width in dorsal view; posterior angles of pronutum rounded (Fig. 

–  Interocular distance twice or longer than eye width; posterior angles of pronutum acute (Fig. 

3  Body length not exceeding 3 mm; eyes circular in lateral view; posterior radial vein (RP) absent in hind wing 

–  Body longer than 3 mm; eyes oval in lateral view (Fig. 

4  Interocular distance 2.0–2.3 times longer than eye width; terminal maxillary palpomere shorter than preceding three combined 

–  Interocular distance 2.5 times eye width; terminal maxillary palpomere as long as preceding three combined 

5  Integument smooth 

–  Integument chagreened 

6  Body length not exceeding 5 mm 

–  Body length longer than 10 mm 

7  Body pale brown; pronotum monochrome; antennal rami as long as respective antennomere; branching of anterior cubital veins (CuA) absent in hind wing; aedeagus with three teeth at the inner apex of paramere 

–  Body darker brown; pronotum bicoloured; antennal rami 1.5 times as long as respective antennomere; anterior cubital veins (CuA) branched into CuA 1, CuA 2, CuA 3+4; aedeagus with one spine at the inner apex of paramere 

8  Each elytron 4.0 times as long as wide; r3 vein absent 

–  Each elytron 4.5 times as long as wide; r3 vein present 

9  Antennae short (less than twice the length of the pronotum) 

–  Antennae long (more than twice the length of pronotum) 

10  Body pale brown or yellow; eyes 3/4 as long as head in lateral view 

–  Body brown or dark brown with pronotum yelloworange; eyes 1/2 or 3/4 as long as head in lateral view 

11  Each elytron 3.0 to 3.5 times as long as wide 

–  Each elytron 4.0 times as long as wide 

12  Body yellow; interocular distance 1.5 times eye width; terminal maxillary palpomere as long as the preceding three combined 

–  Body pale brown; interocular distance 2.0 times eye width; terminal maxillary palpomere shorter than preceding three combined 

13  Pronotal disc without groove along midline 

–  Pronotal disc with groove along midline 

14  Body dark brown with pronotum yelloworange; eyes 3/4 as long as head in lateral view 

–  Body entirely dark brown; eyes 1/2 as long as head in lateral view 

15  Interocular distance twice as long as eye width (Fig. 

–  Interocular distance 3.5 times eye width (Fig. 

16  Interocular distance 2.5 times width of eye; 4^{th} (terminal) maxillary palpomere longer than preceding three combined 

–  Interocular distance 3.0 or more than 3 times width of eye; 4^{th} maxillary palpomere shorter or equal to preceding three combined 

17  Antennomere 1 is longer than antennomeres 2 and 3 combined; 4^{th} (terminal) maxillary palpomere shorter than preceding three combined 

–  Antennomere 1 as long as antennomeres 2 and 3 combined; 4^{th} (terminal) maxillary palpomere equal to preceding three combined 

18  Interocular distance 3.5 to 4.0 times eye width 

–  Interocular distance 3.0 times eye width 

19  Pronotal disc without longitudinal carina 

–  Pronotal disc with a longitudinal carina in posterior portion of pronotum strongly visible, with a length that does not reach the median length of the pronotum 

20  Body length not exceeding 6 mm 

–  Body longer than 9 mm 

21  Body brown, except antennae yellowbrown; interocular distance twice eye width; antennomere 1 longer than antennomeres 2 and 3 combined 

–  Body brown, yellowish mandibles with darker tips; interocular distance 3.0 times eye width; antennomere 1 shorter than antennomeres 2 and 3 combined 

22  Interocular distance at most twice eye width 

–  Interocular distance more than twice eye width 

23  Body orange, except antennae, abdomen, hind wings and legs dark brown; terminal maxillary palpomere half as long as preceding three combined 

–  Body yellow or brown, pronotum yellow orange or brown; terminal maxillary palpomere equal or 2/3 as long as preceding three combined 

24  Pronotum with uniform colouration; posterior radial vein (RP) length twice less than the length of MP1+2 (Fig. 

–  Pronotum mottled with darker brown spots; posterior radial vein (RP) length 1.5 times less than the length of MP1+2 (Fig. 

25  Body brown, with pronotum dark brown near midline; elytra each 3.5 times as long as wide; r3 vein present 

–  Body yellow; elytra each 4.6 times as long as wide; r3 vein absent 

26  Body black; terminal maxillary palpomere as long as preceding three combined 

–  Body dark brown or body dark brown and pronotum yelloworange; terminal maxillary palpomere shorter to longer than preceding three combined 

27  Body dark brown; pronotal disc with a longitudinal carina in posterior portion of pronotum strongly visible, with a length exceeding the median length of the pronotum; elytra 5.4 times as long as wide 

–  Body dark brown and pronotum yelloworange, pronotal disc without longitudinal carina; each elytron less than 5.0 times as long as wide 

28  Antennomere 1 as long as antennomeres 2 and 3 combined; terminal maxillary palpomere longer than preceding three combined 

–  Antennomere 1 is longer than antennomeres 2 and 3 combined; terminal maxillary palpomere shorter than or equal to preceding three combined 

29  Terminal maxillary palpomere shorter than preceding three combined; antennal rami 1.5 times respective antennomere 

–  Terminal maxillary palpomere as long as preceding three combined; antennal rami twice as long as respective antennomere 

California, USA (Fig.
Integument smooth and pronotum bicoloured (yelloworange and dark brown); antennal rami are 1.5 times as long as the respective antennomere; clypeus totally sclerotised; anterior cubital veins (CuA) on hind wing branched; aedeagus with one spine at the inner apex of paramere.
Unknown.
USA: California (Fig.
Map of southern North America showing specimen localities for
“USA CA: 10 mi. NE of /Trimmer/ 24.VI. 93. / Lot 2 BF&JL/ Carr.” “J. & B. Carr Coll. / Bequest to
Jalisco, Mexico.
Integument smooth, antennae less than twice the length of the pronotum, antennal rami as long as the respective antennomere, pronotum as long as wide, each elytron 2.7 times as long as wide and branching of the anterior cubital veins (CuA) absent in the hind wing; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: Jalisco and Colima (Fig.
“MEXICO. Jalisco, San Buenaventura/ 19° 47'.614" N 104° /03'.324" O. Alt. 720 m/ TL3 09021997/ Cols. F. A. Noguera, S. / Zaragoza, E. Ramírez y /E. González” (1) 
Veracruz, Mexico.
Body brown, integument chagreened, antennae less than twice the length of the pronotum, antennal rami 1.5 times the respective antennomere and each elytron 5.1 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: Veracruz (Fig.
USA.
Integument chagreened, antennae less than twice the length of the pronotum, antennal rami twice as long as the respective antennomere, pronotum longer than wide and each elytron 3.3 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
USA (Fig.
Oaxaca, Mexico.
Integument chagreened, antennae long more than twice the length of pronotum, antennal rami twice as long as the respective antennomere, pronotum longer than wide and each elytron 3.8 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: Oaxaca (Fig.
San Luis Potosí, Mexico.
This species can be distinguished by the chagreened integument, antennae long, more than twice the length of pronotum, antennal rami 3 times the respective antennomere and each elytron 4.4 times as long as wide, with two longitudinal costae and aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: San Luis Potosí (Fig.
Species dedicated by the first author to Edgar Uriel Garduño Montes de Oca, her beloved life partner.
Yucatan, Mexico.
Integument chagreened, antennae long, more than twice the length of pronotum, antennal rami twice as long as the respective antennomere and elytra barely reaching the middle of the metasternum, each elytron 3.1 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: Yucatán and Quintana Roo (Fig.
“MEXICO: Quintana Roo/ 19 km N Carrillo Puerto/ 18VI1990 blacklight trap/ coll. M.C. Thomas” “
Hidalgo, Mexico.
Integument smooth, long antennae more than twice the length of pronotum, antennal rami 1.5 times the respective antennomere and each elytron 4.7 times as long as wide with whitish colouration at the apex; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: Hidalgo (Fig.
Durango, Mexico.
Integument chagreened, antennae long, more than twice the length of pronotum, antennal rami twice as long as the respective antennomere, each elytron 2.6 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: Durango (Fig.
Oaxaca, Mexico.
Integument smooth, antennae less than twice the length of the pronotum, antennal rami lanceolate, twice as long as respective antennomere, pronotum as long as wide and each elytron 3.1 times as long as wide; aedeagus with one spine at the inner apex of paramere.
Unknown.
Mexico: Oaxaca (Fig.
Coahuila, Mexico.
Head almost as wide as pronotum, integument chagreened, antennae less than twice the length of the pronotum, antennal rami twice as long as respective antennomere, terminal maxillary palpomere as long as preceding three combined, each elytron 3.6 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: Coahuila (Fig.
Texas, USA.
Integument chagreened, antennae less than twice the length of the pronotum, antennal rami twice as long as the respective antennomere and each elytron 3.5 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
USA: Texas (Fig.
“U.S.A: Texas J. Davis/ Limpia Cyn. / July 26 197/ J. E. Wappes”
Mexico.
Integument smooth, antennae long, more than twice the length of pronotum antennal rami, 3 times the respective antennomere, scutellum almost quadrangular, with small notch on posterior margin and each elytron 4 times as long as wide, with one longitudinal costa; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: Nuevo León and Tamaulipas (Fig.
“Mexico: Tamaulipas/ Gómez Farías km 7 a Julilo/ 13IV2003 /I. Pacheco L. Cervantes” “
Nayarit, Mexico.
We synonymise
Integument smooth, antennae long, more than twice the length of pronotum, antennal rami lanceolate in lateral view, 3.1 times the respective antennomere and each elytron 4.5 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: Nayarit, Guerrero, and Hidalgo (Fig.
“MEXICO: Hidalgo: PN Los Mármoles/ Minas Viejas, Bosque de encino/ 1892 m. N 20° 55' W 99° 12' 41.1”/ Trampa de luz 18VIII2007/ J. Márquez, J. Asiain y S. Sierra cols.” 
Veracruz, Mexico.
Head almost as wide as the pronotum, integument chagreened, antennae long, more than twice the length of pronotum, antennal rami twice as long as the respective antennomere; pronotum mottled with darker brown spots; each elytron 4.0 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: Veracruz, Hidalgo, San Luis Potosí, and Querétaro (Fig.
“Mexico: Hidalgo/: km 14 Carr. Huejutla /Atlepexco 13051999/ H. Brailovsky y E. Barrera” (1) 
Hidalgo, Mexico (Fig.
Body black, integument chagreened, antennae long, more than twice the length of pronotum, antennal rami twice as long as the respective antennomere, terminal maxillary palpomere is as long as the preceding three combined and each elytron 4.3 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: Hidalgo (Fig.
Map of southern North America showing specimen localities for
San Luis Postosí, Mexico.
Body yellow, except for head brown, integument smooth, antennae long more than twice the length of pronotum, antennal rami 1.5 times the respective antennomere, pronotum as long as wide and each elytron 4.3 times as long as wide; aedeagus with one spine at the inner apex of paramere.
Unknown.
Mexico: San Luis Postosí and Tamaulipas (Fig.
“MEXICO: Tamps. Mun. / Gómez Farías, Al/ Cimas, 1000 m. 22 / III1987 P. Kovarik/ R. Jones; UV light” “From the Michael / Ivie Collection” (1) 
Hidalgo, Mexico.
Integument smooth, head almost as wide as the pronotum, antennae long, more than twice the length of pronotum, antennal rami twice as long as respective antennomere, each elytron 6 times as long as wide; aedeagus one with spine at the inner apex of paramere.
Unknown.
Mexico: Hidalgo (Fig.
“MEXICO: Hidalgo, Cuautepec /Tezoncualpan “El Caminero” / Bosque de encino. / 19° 56' 53.8" N; 98° 16' 27.9" W. /Trampa intercepción de vuelo, / 22 a 29 VIII2009, M. Torres col.” (1) 
Monteverde, Costa Rica.
Integument chagreened, head less wide than pronotum, antennae less than twice the length of the pronotum, antennal rami twice as long as the respective antennomere and each elytron 3.5 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Costa Rica: Monteverde, Heredia and Puntarenas (Fig.
“COSTA RICA: Heredia/ La Selva, 75 m /
Texas, USA.
Integument chagreened, head wider than the pronotum, antennae short, less than twice the length of the pronotum, antennal rami twice as long as the respective antennomere and each elytron 3.5 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
USA: Texas; Mexico: Nuevo León (Fig.
USA: “Texas, Cameron Co. / Sabal Palm Grove/ Audobon Reserve/ 2628 May 1979/ N. M. Downie” (1) 
Mexico City, Mexico.
Body dark brown with pronotum yelloworange, integument chagreened, antennae long, more than twice the length of pronotum, antennal rami twice as long as the respective antennomere, terminal maxillary palpomere is longer than the preceding three combined and each elytron 4.5 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: Mexico City (Fig.
“MEXICO: Ciudad de Mexico/Jardín Botánico, /19°19'10" N 99° 11'37.25"/ W, 2321 m.a.s.l. 19X2017/ V. VegaBadillo y S. ZaragozaCaballero” (2) 
San Luis Potosí, Mexico.
Body dark brown, integument chagreened, head less wide than the pronotum, antennae long, more than twice the length of pronotum, antennal rami twice as long as the respective antennomere and each elytron 5.4 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: San Luis Potosí (Fig.
Honduras.
Body pale yellow, integument chagreened, head as wide as the pronotum, antennae long, more than twice the length of pronotum, antennal rami twice as long as the respective antennomere and each elytron 4.6 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Honduras (Fig.
The term sáasil means glow in the Maya language, which is spoken in Honduras.
Sonora, Mexico.
Body pale brown, integument chagreened, head wider than the pronotum, antennae less than twice the length of the pronotum, antennal rami twice as long as the respective antennomere and each elytron 4.1 times as long as wide.
Unknown.
Mexico: Sonora (Fig.
Belize.
Integument chagreened, head almost as wide as the pronotum, antennae less than twice the length of the pronotum, antennal rami 1.5 times the respective and antennomere, each elytron 2.8 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Belize (Fig.
The term “
San Luis Potosí, Mexico.
Body brown except for antennae yellowbrown, integument chagreened, antennae long, more than twice the length of pronotum, antennal rami 3 times as long as the respective antennomere, head almost as wide as the pronotum and each elytron 4.1 times as long as wide; aedeagus trilobed with three teeth at the inner apex of paramere.
Unknown.
Mexico: San Luis Potosí (Fig.
Veracruz, Mexico.
Body dark brown, integument chagreened, antennae less than twice the length of the pronotum, antennal rami 1.5 times the respective antennomere and each elytron 4.3 times as long as wide.
Unknown.
Mexico: Veracruz (Fig.
Puebla, Mexico.
Body brown, except for middle part of pronotum, integument chagreened, head almost as wide as the pronotum, antennae long, more than twice the length of pronotum, antennal rami twice as long as the respective antennomere and each elytron 3.7 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Mexico: Puebla, Hidalgo and Veracruz (Fig.
“MEXICO: Hidalgo, Tlanchinol/TiV1, 1 a 22  X 2011/ Cols. J. Márquez y J. Asiain” (1) 
Puerta Parada, Guatemala.
Integument chagreened, antennae long, more than twice the length of pronotum, antennal rami twice as long as the respective antennomere, terminal maxillary palpomere as long as the preceding three combined and each elytron 4.1 times as long as wide; aedeagus with three teeth at the inner apex of paramere.
Unknown.
Guatemala: Puerta Parada (Fig.
Cartago, Costa Rica.
Head orangebrown, pronotum orange, integument chagreened, head a little wider than the pronotum, antennae long, more than twice the length of pronotum; antennal rami 1.5 times the respective antennomere, terminal maxillary palpomere shorter than the preceding three combined and each elytron 1.8 times as long as wide; aedeagus with one spine at the inner apex of paramere.
Unknown.
Costa Rica: Cartago (Fig.
Species dedicated to our dear friend and colleague Dr. Martín Leonel Zurita García, entomologist who dedicated his life to the study of beetles.
The genera
Some intraspecific variation in colouration has been observed in some species of
Taxonomy is essential for biodiversity knowledge and a crucial part of ecosystem conservation. It is necessary to implement an adequate sampling programme to explore the distributional patterns of
To Paulina Cifuentes Ruiz, Edgar Uriel Garduño Montes de Oca and Hume Douglas for their review and comments that enriched this work. To Susana Guzmán Gómez for her technical assistance in taking the photographs. To Caleb Martins for making the