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Research Article
The Brachiacantha Dejean, 1837 (Coleoptera, Coccinellidae) of Central America
expand article infoJorge Ismael Nestor-Arriola, Víctor Hugo Toledo-Hernández, Ángel Solís§, Guillermo González|, Jaroslav Větrovec
‡ Universidad Autónoma del Estado de Morelos, Cuernavaca, Mexico
§ Museo Nacional de Costa Rica, Santo Domingo de Heredia, Costa Rica
| Unaffiliated, Santiago, Chile
¶ Unaffiliated, Buzulucka, Czech Republic
Open Access

Abstract

A revision of the Central American species of the genus Brachiacantha was undertaken to update the knowledge of the Central American species of the genus. Material of several collections was reviewed, using original descriptions and keys, and comparing with the type material. Twenty-five species of the genus Brachiacantha were found in Central American material, including nine new species: B. nubes Nestor-Arriola, Toledo-Hernández and Solís, sp. nov., B. dentata Nestor-Arriola, Toledo-Hernández and Solís, sp. nov., B. isthmena Nestor-Arriola, Toledo-Hernández and Solís, sp. nov., B. aurantiapleura Nestor-Arriola, Solís and Toledo-Hernández, sp. nov., B. invertita Nestor-Arriola, Toledo-Hernández and Solís, sp. nov., B. papiliona Nestor-Arriola, Toledo-Hernández and Solís, sp. nov., B. tica Nestor-Arriola, Toledo-Hernández and Solís, sp. nov., B. hexaspina González, Větrovec and Nestor-Arriola, sp. nov., and B. mimica Nestor-Arriola and Toledo-Hernández, sp. nov. Nomenclatural changes include Brachiacantha gorhami (Weise), comb. nov., B. guatemalensis (Gorham), comb. nov., and Brachiacantha duodecimguttata Leng, syn. nov. for B. lepida Mulsant. The male genitalia of the species B. fenestrata Gorhan, B. octostigma Mulsant, B. aperta Weise, and B. cachensis Gorhan are described and illustrated for the first time. New records include B. indubitabilis Crotch and B. bipartita Mulsant (Costa Rica and Guatemala), B. gorhami (Weise) (El Salvador), and B. cachensis Gorham (Panamá). A key to the species is included.

Keywords

Cusps, holotypes, Hyperaspidini, key, synonyms, tooth

Introduction

The genus Brachiacantha Dejean, 1837, belongs to the family Coccinellidae, also known as ladybugs, lady beetles, ladybirds or ladybird beetles. The most evident character of the genus is the tooth on the leading edge of the protibia; however, this tooth is variable in size and is very small in some species (Gordon 1985; Gordon et al. 2014). Twelve species were listed for Central America, México, and the United States in the Biologia Centrali-Americana (Gorham 1894). Some species were added later by Weise (1903) and Leng (1911). Several of the above-mentioned studies do not include a description of the male genitalia of species, which is one of the structures most commonly used to recognize species today. The genus belongs to the tribe Brachiacanthini, but Seago et al. (2011) synonymized this tribe with Hyperaspidini.

Leng (1911) was the first to divide the genus into groups; he described six groups based on external characters; however, some of the groups were inaccurate (Nestor-Arriola and Toledo-Hernández 2016). Gordon (1985) revised the North American species and divided them into four groups based on external characters and male genitalia; some of his groups resemble those of Leng (1911). Gordon et al. (2014) revised the species of South America and divided them into twelve groups based on male genitalia, and in only a few groups on external characters and female genitalia. Some of the South American groups are equated with those of North America; the bistripustulata group (Gordon et al. 2014) have the same diagnostic characters of the dentipes group (Gordon 1985). The same applies to the North American ursina group and the South American debbie group; therefore, the known Brachiacantha species can be separated into fourteen valid groups.

Materials and methods

The material of the Brachiacantha species studied comes from several collections. The acronyms used in the text are as follows:

CEAM Colección Entomológica del Colegio de Posgraduados, Campus Montecillo. Texcoco, State, México

CNIN Colección Nacional de Insectos, Instituto de Biología, Universidad Nacional Autónoma de México, México City, México

FSCA Florida State Collection of Arthropods, Gainesville, Florida, USA

MNCR Museo Nacional de Costa Rica, Santo Domingo, Heredia, Costa Rica

MUCR Museo de Insectos de la Universidad de Costa Rica, San José, Costa Rica

MZCR Museo de Zoología de la Universidad de Costa Rica, San José, Costa Rica

NMP National Museum, Prague, Czech Republic

OUMNH Oxford University Museum of Natural History, Oxford, United Kingdom

USNMNH United States National Museum of Natural History, Washington D. C., USA.

Genitalia of both sexes were dissected by softening the specimens in hot water, removing the abdomen, placing it in a dilute solution of potassium hydroxide until the fat and muscle were dissolved, then rinsing the abdomen and genitalia in clean water and placing the structures in microvials filled with glycerin for later examination.

The anatomy terminology used is based on Gordon (1985) and Gordon et al. (2014); and updated according to Ślipiński (2007). However, the terms for male genitalia position (dorsal, ventral) have been changed to “concave side” or “convex side” because the male genitalia change position and the terms “dorsal side” and “ventral side” can induce error.

The species were assigned to known species groups according to the diagnostic characters described by their authors. The species groups considered as valid are those created by Gordon (1985) and Gordon et al. (2014); a new group was created and described only when the characters of a species or group of species did not match with any known group.

To abbreviate the information, only new species have a complete list of material reviewed, with assignment of holotypes and paratypes. Reviewed material for known species includes only the number of specimens, locations, and the collections where the material is located. In the cases were the type material of a known species was not reviewed, the method of identification of the species is explained.

Results

Twenty-five species were found, nine of which are new species. The species studied here belong to eight previously described groups and two new groups. Four species could not be identified as members of a group due to the lack of males.

Some species of the groups lepida and dentipes were previously recorded in North America (Gordon 1985). South American species were not found in Central America, but some species show some affinities with southern fauna. The dentipes group was the most abundant in Central America.

Species key

1 Elytra orange or yellow without spots B. bipartita Mulsant
Elytra with spots or some marks 2
2 Dorsal color mainly black or dark brown 3
Dorsal color mainly yellow, orange, or pale gray 17
3 Elytra with apex of elytral suture dentiform B. mimica sp. nov.
Apex of elytral suture not dentiform 4
4 Elytra black with lateral and apical borders yellow B. guatemalensis (Gorham)
Each elytron with one to five spots 5
5 Pronotum with two convergent discal spots, each elytron with five pale spots B. fenestrata Gorham
Pronotum without discal convergent spots, elytral maculation variable 6
6 Each elytron with five pale spots B. octostigma Mulsant
Elytra with different maculation 7
7 Elytra black with a large orange macula covering lateral margin B. aurantiapleura sp. nov.
Elytra with different maculation 8
8 Each elytron with three pale spots: one near the scutellar shield and two near the elytral apex B. tica sp. nov.
Elytra with different maculation 9
9 Males with setal tuft on center of first ventrite, male genitalia symmetrical B. isthmena sp. nov.
Males without setal tuft on center of first ventrite, male genitalia asymmetrical 10
10 Males without humeral spot and without cusps on third ventrite, male genitalia asymmetrical B. indubitabilis Crotch
Males with humeral spot. Third ventrite of male with two cusps, male genitalia asymmetrical. 11
11 Fourth ventrite with cusps B. barberi Gordon
Fourth ventrite without cusps 12
12 Penis guide of male genitalia with apical hook 13
Penis guide of male genitalia without apical hook 15
13 Apical hook of the penis guide curved and robust B. robustihamata Nestor-Arriola and Toledo-Hernández
Apical hook not curved or robust 14
14 Cusps on third ventrite very conspicuous parameres of male genitalia robust and with a tooth on superior margin B. dentipes (Fabricius)
Cusps moderately conspicuous, parameres without tooth B. bistripustulata (Fabricius)
15 Elytron black without spots, apical margin of elytron red or orange B. erythrura Mulsant
Elytron spotted 16
16 Cusps medium sized B. subfasciata Mulsant
Cusps large B. truncata Nestor-Arriola and Toledo-Hernández
17 Elytra yellow with pale brown lines B. aperta Weise
Elytra with black spots 18
18 Each elytron with a large basal macula B. papiliona sp. nov.
Elytra without a large basal macula 19
19 Protibia, mesotibia and metatibia toothed, two spots on each elytron B. hexaspina sp. nov.
Mesotibia and metatibia without tooth, maculation variable 20
20 Elytra pale with several elongated black spots, tibial tooth small B. gorhami (Weise)
Elytral spots rounded, not elongated 21
21 Pronotum mainly black with only lateral borders, anterior border and middle line until center of the disc pale colored. Each elytron with five black spots B. nubes sp. nov.
Pronotum variable, if mainly black with several pale colored projections into the basal black macula. Elytra variable 22
22 Elytra yellow with five black spots on each elytron, pronotum with several black small spots B. invertita sp. nov.
Each elytron with two black or brown spots, elytral suture black or with a large dark macula 23
23 Male abdomen with central part of ventrites depressed and with dense, long pubescence B. cachensis Gorham
Male abdomen without long pubescence 24
24 Dorsal color mainly yellow, orange or pink, sometimes dark with 12 pale spots; male abdomen without modifications B. lepida Mulsant
Dorsal color yellow. Male abdomen with acute tooth in middle of first and second ventrites B. dentata sp. nov.

Species accounts

dentipes group

Abdomen of male with the third ventrite bicuspid, penis guide of male genitalia asymmetrical (Gordon 1985; Gordon et al. 2014).

Brachiacantha barberi Gordon

Brachiacantha barberi Gordon, 1985: 572. Nestor-Arriola and Toledo-Hernández 2019: 548.

Material examined

Material listed in Nestor-Arriola and Toledo-Hernández (2019).

Diagnosis

Length 3.2–3.9 mm. Width 2.6–2.9 mm. Dorsally black with two to three red to orange spots; pronotum mostly black. Third and fourth ventrites bicuspid. Male genitalia asymmetrical, penis guide without apical hook.

Distribution

From the southeast of the USA to Costa Rica.

Discussion

This species is similar to B. bistripustulata, but it can be distinguished by its smaller size and the fourth bicuspid ventrite of B. barberi.

Brachiacantha bistripustulata (Fabricius)

Coccinella laevis Thunberg, 1781: 20.

Coccinella bistripustulata Fabricius, 1801: 383.

Brachyacantha bis-tripustulata: Mulsant 1850: 528.

Brachyacantha bistripustulata: Gorham 1894: 188. Casey 1899: 119. Leng 1911: 296. Korschefsky 1931: 203. Blackwelder 1945: 449.

Brachyacantha erythrocephala Gorham, 1894: 188 (non Coccinella erythrocephala Fabricius). Crotch 1874: 211.

Brachyacantha decora Casey, 1899: 119. Bowditch 1902: 206.

Brachyacantha bistripustulata var. guttata Weise, 1885: 231. Leng 1911: 297. Korschefsky 1931: 203.

Brachyacantha bistripustulata var. decora: Leng, 1911: 298. Leng 1920: 213.

Brachyacantha bistripustulata var. minor Leng, 1911: 298. Leng 1920: 213.

Brachyacantha bistripustulata var. obscura Leng, 1911: 299. Korschefsky 1931: 204.

Brachyacantha bistripustulata var. quichiana Leng, 1911: 298.

Brachyacantha bistripustulata ab. decora: Korschefsky 1931: 203.

Brachyacantha bistripustulata ab. minor: Korschefsky 1931: 203.

Brachiacantha decora: Gordon 1985: 561.

Brachiacantha laevis: Pope 1987: 58.

Brachiacantha bistripustulata: Milléo and Almeida 2007: 420. Gordon et al. 2014: 10. Nestor-Arriola and Toledo-Hernández 2019: 539.

Material examined

496 specimens from Costa Rica: Alajuela, Cartago, Heredia and Puntarenas; El Salvador: Ahuachapán, La Libertad and San Salvador; Guatemala: El Quiché, El Progreso, Frontera, Jutiapa, Chimaltenango, Baja Verapaz, Santa Rosa and Zacapa; Honduras: El Paraíso, Tegucigalpa, Atlántida, Colón, Cortés, Comayagua, Francisco Morazán, La Paz, Olancho and Yoro; Panamá: Oeste, Coclé, Chiquirí and Darien (CEAM, CNIN, FSCA, USNMNH, OUNMH, MNCR, MUCR, MZCR).

Diagnosis

Length 3.2–5.4 mm. Width 2.4–4 mm. Dorsally black with three to four orange, yellow or red spots on each elytron, spots sometimes fused. Males with pale-yellow head and pronotal anterior border. Third ventrite bicuspid in males, cusps short, wide and with divergent apex. Male genitalia asymmetrical, penis guide with apical hook.

Distribution

From the South of the USA to Brazil and Peru, including Jamaica, Cuba, Puerto Rico and Dominican Republic.

Discussion

This species is very similar to others in the group, but the shape of the male ventral cusps is different in being wider than long, apically divergent and not strongly pronounced, and is usually enough to differentiate it.

Brachiacantha dentipes (Fabricius)

Coccinella dentipes Fabricius, 1801: 381. Oliver 1808: 1051. Say 1835: 202.

Brachyacantha dentipes: Mulsant 1850: 525. Crotch 1873: 378. Gorham 1894: 196. Casey 1899: 120. Nunenmacher 1909: 162. Leng 1911: 300. Korschefsky 1931: 204. Wingo 1952: 27. Chapin 1974: 44.

Brachyacantha socialis Casey, 1899: 119. Wingo 1952: 27.

Brachyacantha dentipes socialis: Leng 1911: 301.

Brachyacantha dentipes americana Leng, 1911: 302.

Brachyacantha dentipes var. separata Leng, 1911: 301. Wingo 1952: 27.

Brachyacantha dentipes ab. socialis: Korschefsky 1931: 204.

Brachyacantha dentipes ab. separata: Korschefsky 1931: 204.

Brachiacantha dentipes: Gordon 1985: 564. Flores-Mejía and Salas-Araiza 2004: 14. Nestor-Arriola and Toledo-Hernández 2019: 542.

Material examined

340 specimens from Costa Rica: Alajuela, San José, Guanacaste, Heredia, Puntarenas, and San José; Honduras: El Paraíso and Olancho (FSCA, USNMNH, OUMNH, MNCR, MUCR, MZCR).

Diagnosis

Length 3–4.5 mm, width 2.1–3.2 mm. Three to four spots on each elytron; the spots may be fused. Third male ventrite with large, subtriangular divergent cusps. Male genitalia with wide parameres with strong tubercle on the concave margin; penis guide with elongated apical hook.

Distribution

From the south of the USA to Costa Rica, including the Bahamas.

Discussion

This species is very similar to B. bistripustulata, but the male third ventrite has larger cusps and the male genitalia are more robust in B. dentipes.

Brachiacantha erythrura Mulsant

Brachyacantha erythrura Mulsant, 1850: 530. Crotch 1874: 211. Gorham 1894: 187. Leng 1911: 302. Nestor-Arriola and Toledo-Hernández 2019: 545.

Material examined

Material listed in Nestor-Arriola and Toledo-Hernández (2019).

Diagnosis

Length 3.8–3.9 mm, width 2.8–2.9 mm. Dorsally black, head and posterior margin of the elytron red, orange or yellow. Ventral surface light brown to orange. Third ventrite of the male with small cusps. Penis guide of the male genitalia asymmetrical, without apical hook.

Distribution

México and Central America.

Discussion

This species can be confused with some specimens of B. quadrillum LeConte, but the third ventrite cusps of B. erythrura are much smaller and the male genitalia lack the apical hook.

Brachiacantha robustihamata Nestor-Arriola & Toledo-Hernández

Brachiacantha robustihamata Nestor-Arriola & Toledo-Hernández, 2017: 48.

Material examined

Material listed in Nestor-Arriola and Toledo-Hernández (2017).

Diagnosis

Length 4.5 mm, width 3.2 mm. Oval body. Three spots on each elytron, including an incomplete transversal band. Third ventrite with a small narrow cusp at each side of middle. Apical hook of the penis guide of the male genitalia large and rounded.

Distribution

Guatemala and Costa Rica.

Discussion

This species is very similar to B. bistripustulata and other species of the dentipes group. The principal difference is the male genitalia with a robust curved hook and male ventral cusps longer than wide. The cusps in the third ventrite are similar to those of B. blaisdelli Nunenmacher, but the male genitalia of B. robustihamata are distinguished by the curved hook.

Brachiacantha subfasciata Mulsant

Brachyacantha subfasciata Mulsant, 1850: 527. Crotch 1874: 211. Gorham 1894: 187. Leng 1911: 302. Leng 1920: 212. Korschefsky 1931: 207.

Brachiacantha subfasciata Gordon, 1985: 574. Nestor-Arriola and Toledo-Hernández 2019: 550.

Material examined

Material listed in Nestor-Arriola and Toledo-Hernández (2019).

Diagnosis

Length 3.8–5 mm, width 2.8–3.3 mm. Oval body. Dorsally black with one to four spots on each elytron. Third ventrite of male with separated cusps connected by a curved emargination. Male genitalia asymmetrical; penis guide almost truncate, without apical hook.

Distribution

From the center of the USA to Honduras.

Discussion

This species is very similar to others of the dentipes group, but the male genitalia are different as they lack the apical hook on the penis guide.

Brachiacantha truncata Nestor-Arriola & Toledo-Hernández

Brachiacantha truncata Nestor-Arriola & Toledo-Hernández, 2017: 45.

Material examined

Material listed in Nestor-Arriola and Toledo-Hernández (2017).

Diagnosis

Length 3.2–5.3 mm, width 2.5–4.2 mm. Male with four spots on the elytron, females with three spots on the elytron. Males with a triangular, dull apex, cusp at each side of the middle line. Male genitalia with penis guide without apical hook, shorter than the parameres, oblique at apex, almost truncate.

Distribution

México and Honduras.

Discussion

The male genitalia with oblique apex and the ventral cusps with blunt apex are enough to distinguish the species from others in the group.

lepida group

Male genitalia with penis guide asymmetrical, apex obliquely truncate, penis without alae (Gordon 1985). Brachiacantha lepida and B. indubitabilis were classified in two different monospecific groups by Gordon (1985), distinguished by differences in the male abdomen; however, after Gordon et al. (2014) the male genitalia are almost the only criterion used for group recognition purposes. There are no significant differences between the genitalia of the species, and they are, therefore, classified in the same group.

Brachiacantha lepida Mulsant

Figures 1, 2

Brachyacantha lepida Mulsant, 1850: 523. Crotch 1873: 378. Crotch 1874: 210. Gorham 1894: 185. Leng 1911: 324. Leng 1920: 212. Korschefsky 1931: 205.

Brachyacantha duodecimguttata Leng, 1911: 289. Syn. nov.

Brachyacantha 12-guttata: Leng 1911: 294. Syn. nov.

Brachiacantha lepida: Gordon 1985: 599.

Material examined

271 specimens from Belize: Orange Walk; Costa Rica: Alajuela, Cartago, Heredia, Guanacaste, Limón, San José and Puntarenas; Guatemala: Chimaltenango, El Quiche, Guatemala City, Huehuetenango, Suchitepequez and Totonicapan; Honduras: Atlantida; Panamá: Chiriqui (CEAM, CNIN, FSCA, USNMNH, OUMNH, MNCR, MUCR, MZCR).

Diagnosis

Length 2.3–3.7 mm, width 1.8–2.8 mm. Oval body. Dorsally pale yellow to pinkish orange except pronotum with a multilobed basal macula; each elytron with 4 black spots, one spot in a sub-humeral position, another in the second half of the elytron, a discal spot on the elytral suture, and a subapical spot on the elytral suture (Figs 1, 2). Ventral color dark brown to light brown. Males with the ventrites V and VI slightly emarginated, not depressed. Male genitalia asymmetrical. Protibia flanged.

Figures 1–4. 

Brachiacantha sp. 1 Brachiacantha lepida, dark habitus (image by Jorge Valdez Carrazco) 2 B. lepida, common habitus (image by Jorge Valdez Carrazco) 3 Brachiacantha fenestrata, male 4 Brachiacantha octostigma, male (image by Jorge Valdez Carrazco).

Distribution

From southern USA to Panamá.

Discussion

The small size and coloration are usually enough to distinguish this species from other species of the genus. Some specimens from Guatemala display the darker dorsal coloration that was previously described by Leng (1911) as the species B. duodecimguttata. The holotype of B. duodecimguttata is a female; the genitalia of male specimens with morphology similar to that described by Leng (1911) were dissected and no differences were found from that of B. lepida. Therefore, B. duodecimguttata should be considered a junior synonym of B. lepida.

Brachiacantha indubitabilis Crotch

Brachyacantha indubitabilis Crotch, 1873: 379. Casey 1899: 120. Leng 1911: 315. Korschefsky 1931: 205.

Hyperaspis triplicans Casey, 1924: 163. Korschefsky 1931: 198. Dobzhansky 1941: 11.

Hyperaspis triplicans microsticta Casey, 1924: 163. Korschefsky 1931: 192. Dobzhansky 1941: 11.

Brachiacantha indubitabilis Gordon, 1985: 596. Flores-Mejía and Salas-Araiza 2004: 14.

Material examined

Costa Rica • 2♀, 6♂; Guanacaste Prov., 20 km N Liberia; 15 Jul 1989; David G. Furth leg. (USNMNH). Guatemala • 1♂; Chicacao; 07 Jul 1949; T. H. Farr leg. (USNMNH).

Diagnosis

Length 3.2–3.7 mm, width 2.5–2.8 mm. Dorsally black, pronotum with lateral and anterior margins yellow, orange, or red; each elytron with three orange or red spots, males and females without humeral spots. Males without ventral cusps. Penis guide of male genitalia asymmetrical.

Distribution

United States of America and México. New record to Costa Rica and Guatemala.

Discussion

This species shows a color pattern that is very common in species of the dentipes group; however, it can be recognized because it lacks ventral cusps and lacks humeral spots in males.

buckleyi group

Penis guide of male genitalia long, slender, apically truncated. Parameres longer than penis guide, very wide at base, narrowed to rounded apex (Gordon et al. 2014).

Brachiacantha nubes Nestor-Arriola, Toledo-Hernández & Solís, sp. nov.

Figures 21–24

Holotype

Male, pinned, with genitalia in a separate microvial. Original labels: “Est. Santa Elena, Viejo, Santa Elena, Las Nubes, Prov. San José, COSTA RICA, 1210 m, 23–28 Oct. 1995, col. E. Alfaro, L_S_371750_507800, #6385” (MNCR).

Diagnosis

Dorsal color pale orange with five black spots on each elytron, pronotum orange with a basal black macula covering more than the half of the pronotum.

Description

Male holotype. Length 2.4 mm, width 2 mm; body rounded, convex. Dorsal color pale orange except pronotum with large black macula covering more than the half except the lateral and anterior margins and the middle line in the apical half until the center of the pronotal disk; scutellar shield black; each elytron with five black oval spots: sub-humeral spot large, scutellar spot over the elytral suture near the middle of the elytra, mid-lateral spot on the apical half of the elytra, discal spot on the apical half of the elytra, sub-apical spot over the elytral suture (Fig. 21). Ventral surface with head, prosternum, metaventrite and abdomen dark brown; legs, mouthparts and antennae yellow. Head punctures small, separated by 1½ their diameter, each puncture as large as an eye facet; pronotal punctures larger than head punctures, separated by 1½× their diameter; elytral punctures as large as head punctures, separated by 1½× their diameter; metaventral punctures larger than pronotal punctures, separated by their diameter. Clypeus emarginated. Prosternal carinae not examined. Protibia not flanged, basal tooth small. Basal abdominal ventrite without setal tuft. Abdomen with postcoxal line on basal abdominal ventrite slightly flattened along posterior ventrite margin; ventrite with sparse, long pubescence and small, dense punctures. Ventrites IV–VI depressed and emarginated (Fig. 22). Ventrites II–VI pubescent throughout, punctures fine, dense; third ventrite with a tubercle on each side of central part. Genitalia with penis guide slightly shorter than parameres, symmetrical, sides slightly convergent, slightly wider before apex, apex truncate (Fig. 23); parameres curved, wide at base, narrowed to apex, apex acute, setae arising from the apex (Fig. 24). Penis not examined.

Female. Unknown.

Variation

Unknown.

Etymology

The name refers to the type location.

Type locality

COSTA RICA, San José: Santa Elena de General Viejo, Las Nubes Biological Reserve Ecocampus (Faculty of Environmental Studies, York University), 9°23'08.05"N, 83°36'11.82"W.

Distribution

Costa Rica.

Discussion

This species is easily identifiable by its dorsal coloration. The only species it could be confused with is B. lepida; however, the number and arrangement of elytral spots are different in these species as nubes has more spots than lepida.

Brachiacantha fenestrata Gorham

Figures 3, 25–28

Brachyacantha fenestrata Gorham, 1894: 190. Leng 1911: 322.

Material examined

160 specimens from Costa Rica: Alajuela, Guanacaste, Heredia, and Puntarenas; Panamá: Chiquirí (MNCR, MUCR, MZCR).

Diagnosis

Dorsally black or dark brown; pronotum with the anterior angles, the anterior margin and two convergent oval spots on the center of the disc, pale yellow; each elytron has five pale spots (Fig. 3). Male abdomen with ventrites I-V depressed, emarginated, and abundantly pubescent. Male genitalia with the penis guide apically truncate and slightly longer than parameres, symmetrical, with convergent sides (Fig. 25); parameres slender, apex rounded, setae arising from the convex side margin (Fig. 26); penis curved in basal 1/2, apex with small alae, basal capsule without crest, inner arm of basal capsule long and slender (Figs 27, 28).

Variation

In some specimens the elytral spots are fused.

Distribution

Costa Rica and Panamá. The records of México must be confirmed.

Discussion

This species is easily recognizable by the pronotal pale spots.

juanita group

Male genitalia with penis guide short, evenly, ovately narrowed from base to apex, apex narrowly rounded, sometimes acute; penis with inner arm of basal capsule apically bifid. Female genitalia with spermathecal capsule long, slender, basal ¼ widened, cornu bulbous or narrowed to acute apex (Gordon et al. 2014).

Brachiacantha octostigma Mulsant

Figures 4, 29–32

Brachyacantha octostigma Mulsant, 1850: 539. Crotch 1874: 212. Gorham 1894: 188. Leng 1911: 311.

Material examined

Guatemala • 2♀; Totonicapan, V. Sta. María; 1850 m; 19 Jun 1973; Ginter Ekis leg. (USNMNH) Panamá • 1♀; V. Chiquiri; 2000–3000 ft; Champion leg. (USNMNH).

Diagnosis

Oval body. Elytra black, each elytron with five yellow spots. Male abdomen with several ventrites emarginated and depressed. Male genitalia with penis guide wide, shorter than the parameres, not truncate at apex, symmetrical, sides parallel but convergent at apical ⅓ (Fig. 30); parameres curved, wide at base, narrowed to apex, apex acute, setae arising from the apex and the convex side margin (Fig. 29); penis curved in basal ½, apex without alae, basal capsule crested, inner arm of basal capsule long and slender (Figs 31, 32).

Distribution

México and Central America.

Discussion

The coloration of this species is very similar to that of the ursina group, but the male genitalia are different, with a shorter penis guide with the apex not truncate. The material was identified by comparison with previously identified as B. octostigma by J. Chapin in 1956 (USNMNH), the diagnosis of the male and the male genitalia is based on Mexican specimens of the same series.

Brachiacantha bipartita Mulsant

Figures 5, 33–36

Brachyacantha bipartita Mulsant, 1850: 521. Crotch 1874: 211.

Brachyacantha westwoodii Gorham, 1894: 185 (in part). Leng 1911: 305 (in part).

Brachiacantha bipartita: Booth and Pope 1989: 369. Nestor-Arriola and Toledo-Hernández 2016: 826.

Material examined

Costa Rica • 1♂; San José; 1000–1200 m. a. s. l.; Ago 1980; N. L. H. Krauss leg. (USNMNH). Guatemala • 1♀; Antigua; Oct 1965; L. H. Krauss leg. (USNMNH).

Diagnosis

Body oval. Elytron orange. Head pale yellow in males, black in females. Males with pronotum black except lateral and anterior margins pale yellow to yellowish white (Fig. 5). Male genitalia with penis guide shorter than parameres, symmetrical, sides convergent at apical ⅓ (Fig. 34); parameres curved, wide at base, narrowed to apex, apex acute, setae arising from the apex and the convex side margin (Fig. 33); penis curved in basal ½, apex with long slender alae (Fig. 36), basal capsule crested, apex long and slender (Fig. 35).

Figures 5–8. 

Brachiacantha sp. 5 Brachiacantha bipartita, female 6 Brachiacantha aperta, female 7 Brachiacantha isthmena sp. nov. 8 Brachiacantha cachensis.

Natural history

This species has been collected on coffee plants (Coffea sp.).

Distribution

Mexico and Central America. New record to Costa Rica and Guatemala.

Discussion

This species is easily recognizable by the orange elytra without maculation. The only species similar to B. bipartita is B. westwoodii, but B. westwoodii is much larger and has the pronotum almost entirely orange with only a basal black macula (Nestor-Arriola and Toledo-Hernández 2016). The material was identified following the description of Mulsant (1850) and considering the information of Booth and Pope (1989).

Brachiacantha aperta Weise

Figures 6, 37–40

Brachyacantha aperta Weise, 1903: 208. Leng 1911: 323.

Material examined

123 specimens from Costa Rica: Limón, Cartago, and Puntarenas (FSCA, USNMNH, MNCR, MUCR, MZCR).

Diagnosis

Color pale yellow or yellowish white. Pronotum with three or four brown spots. Elytron with a brown-colored branched line pattern (Fig. 6). Ventrites I-VI emarginated and depressed at center, depression becoming wider from the basal ventrite to apical ventrite. Ventrites I-III densely punctate, pubescent within the depression more abundant and longer than at the sides. Male genitalia with parameres and penis guide strongly wide; penis guide as long as parameres, wider at middle, symmetrical, apex rounded (Fig. 40); parameres wide, apex rounded, slightly tuberculated in the concave side margin, with abundant setae at apex and the convex margin, weakly curved (Fig. 39); penis with lateral slender alae at apex (Fig. 38), basal capsule not crested, inner arm slender and long, apex spender and long (Fig. 37).

Distribution

Costa Rica.

Discussion

This species is easily recognizable due to its unique coloration.

Brachiacantha dentata Nestor-Arriola, Toledo-Hernández & Solís, sp. nov.

Figures 41–45

Holotype

Male, pinned, with genitalia in a separate microvial. Original labels: “San Lorenzo, R. F. Cord., Guanacaste, COSTA RICA, 1050 m, abril 1991, col. C. Alvarado, L-N 287800_427600” (MNCR).

Paratypes

Costa Rica • 1♂; Alajuela, P. N. V. Tenorio, V. Rio Roble, Palmital, “arriba la caliza”; 1000–1100 m; 21 June-16 Sep. 2006; J. A. Azofeifa leg.; malaise trap; L_N_296500_426300, #87178 (MNCR). 1♂; Alajuela, Palmarena, Rio San Lorencito, 5 km N of R. F. Sn Ramón; 900 m; Mar. 1990; Curso Carabidae exped.; 244500-470700 (MNCR). 1♂; Cartago, A. C. Amistad, P. N. Tapantí, Rio Dos Amigos; 1480 m; Mar. 1994; G. Mora, A. Solís and E. Ulate leg; L N 187600_560250, #2782 (MNCR). 1♂; Cartago, Capellades; 2–5 Apr. 1942; Biering leg., on Carica papaya L.; det. J. B. Chapin 1991 (MUCR). 1♂; Cartago, P. N. Tapantí; 1450 m; 13–15 May. 2005; Barries, Cate and Nagy leg.; L_N_190920_560811, #86647 (MNCR). 2♂; Cartago, P. N. Tapantí, Quebrada Segunda; 1250 m; Nov. 1992; G. Mora leg.; L-N 194000_560000 (MNCR). 1♂, same data; Dec. 1992 (MNCR). 3♂; same data; ago.1992 (MNCR). 1♂; same data, sep.1992; (MNCR). 2♂; same data; July 1992 (MNCR). 1♂; same data, Oct. 1992 (MNCR). 1♂; same data; July 1992; det. Elena Ulate INBio June 2003 as B. cachensis (MNCR). 1♂; Cartago, P. N. Tapantí, Quebrada Segunda; 1300 m; Oct. 1993; G. Mora leg.; L-N 194000_559800, #2286 (MNCR). 1♂; Cartago, Ref. Nac. Fauna Silv. Tapanti; 1250 m; Aug. 1991; F. A. Quesada leg.; L_N_194000_559800 (MNCR). 1♂; Cartago, Turrialba, M. N. Guayabo; 1100 m; 21 June 1994; F. J. Corrales leg.; L N 570000_217400, #3028” (MNCR). 1♂. Cartago, Turrialba, Tayutic, Grano de Oro; 1120 m; Oct. 1993; P. Campos leg.; L_N_200250_595900, #2438 (MNCR). 2♂; Guanacaste, R. F. Cord. Guanacaste, R. San Lorenzo, Tierras Morenas; 1050 m; Dec. 1991; C. Alvarado leg. L-N-287800_427600 (MNCR). 1♂; Guanacaste, Rio San Lorenzo, Tierras Morenas; 1050 m; 8 Mar.- 26 Apr. 1995; G. Rodríguez leg.; malaise trap; L.N. 287800_427600, #4488 (MNCR). 1♂; same data; dic.1992; #1764 (MNCR). 1♂; Puntarenas, A. C. Arenal, Buen Amigo, San Luis Monteverde; 1000–1350 m; May 1994; Z. Fuentes leg.; L N 250850_449250, #2926 (MNCR). 1♂; Puntarenas, R. B. Monteverde; 1520 m; Nov. 1993; N. Obando leg.; L N 253250_449700, #2478 (MNCR). 1♂; Puntarenas, R. B. Monteverde, est. La Casona; 1520 m; Mar. 1994; N. Obando leg.; det. Elena Ulate INBio June 2003 as B. cachensis; LN 253250_449700, #2819 (MNCR). 2♂; Puntarenas, R. B. Monteverde, est. Leonel Hernandez; 1600 m; June 1991; E. Bello leg.; L-N-249750_450075 (MNCR). 1♂. Puntarenas, R.B. Monteverde, San Luis; 1040 m; 24 Aug.-15 Sep. 1992; Z. Fuentes leg.; L-N-250850_449250 (MNCR). 1♂; same data; Sep. 1992 (MNCR). 1♂; San José, P. N. Braulio Carrillo; 1600 m; 26 Aug. 1998, C. W. and L. B. O´Brien leg. (FSCA).

Diagnosis

Dorsal color pale yellow with brown or black spots, ventrally brown. Males with a sharpened tubercle or tooth on second ventrite and a sharpened tubercle on each side of the center of the third ventrite. Male genitalia with penis guide wide, convergent at apex; parameres wide at base, rounded at apex; penis with a subtriangular projection near the alae.

Description

Male holotype. Length 3.1 mm, width 2.5 mm; body oval, convex. Dorsal color pale yellow except pronotum with a basal, short and wide brown macula, and a small spot on each side of the center of the disc; scutellar shield brown; elytron with three pale brown marks: sub-humeral and sub-apical spots closely oval, sutural vitta over the elytral suture from near the scutellar shield to the elytron apex (Fig. 41). Ventral surface with prosternum, metaventrite and abdomen brown; legs, mouthparts, and antennae yellow. Head punctures small, separated by their diameter, each puncture slightly larger than an eye facet; pronotal punctures as large as head punctures, separated by their diameter or less; elytral punctures larger than pronotal punctures, separated by their diameter or more; metaventral punctures as large as elytral punctures, separated by more than three diameters. Clypeus straight. Carinae on prosternal process straight, convergent but almost parallel, not joined, larger than half the prosternum. Protibia only slightly flanged with the border almost parallel to tibia, basal tooth small. Basal abdominal ventrite without setal tuft. Abdomen with postcoxal line on basal abdominal ventrite slightly flattened along posterior ventrite margin, ventrite with sparse, long pubescence and small, dense punctures. Ventrite II with a spur in the center at the posterior margin; ventrite III with a spur on each side of the middle line at the posterior margin. Ventrites IV-VI depressed and medially emarginated (Fig. 42). Ventrites II-VI pubescent throughout, punctures fine, dense. Genitalia with penis guide as long as parameres, symmetrical, parallel sided, convergent at apex (Fig. 44); parameres wide basally, narrowed to apex, apex rounded and wide, setae rising from the convex side border and apex (Fig. 43); penis curved in basal ½, apex with small alae and a pair of flattened subtriangular appendages before the alae, basal capsule without crest, inner arm of basal capsule long and slender (Fig. 45).

Female. Unknown. There were several females with the same data as males identified as B. dentata, but it was not possible to differentiate them from females of B. cachensis, a very similar species of the same region. Males of B. cachensis and B. dentata have different sexual characters.

Variation

Length 2.6–3.2 mm, width 2–2.5 mm. The pronotal spots can be fused. On some specimens there are two spurs on the third ventrite, but others have only one.

Etymology

From the Latin dentatus (= toothed). The name refers to the ventral spurs or teeth.

Type locality

COSTA RICA, Guanacaste: Reserva Forestal Cordillera de Guanacaste, Rio San Lorenzo, 10°36'39.999"N, 84°59'44.9988"W.

Distribution

Costa Rica.

Discussion

This species is externally similar to B. cachensis but can be differentiated from it by the dentate ventrites of the male and the male genitalia with the penis guide pointed at apex. It was not possible to distinguish between the females of these species.

tucumanensis group

Male genitalia with penis guide about as long as parameres, slightly “pinched” laterally at basal ⅓, widened in apical ½, sides rounded to acute apex (Gordon et al. 2014).

Brachiacantha aurantiapleura Nestor-Arriola, Solís & Toledo-Hernández, sp. nov.

Figures 46–51

Holotype

Male, pinned, with genitalia in a separate microvial. Original labels: “COSTA RICA, PUNTARENAS, Monteverde, San Luis, 1000–1400 m, Feb. 1994, col. Z. Fuentes, L N 449250_250850, #2615” (MNCR).

Paratype

Costa Rica • 1♀; same data as the holotype; #2771 (MNCR).

Diagnosis

Dorsal color black with the head and the angles of the pronotum yellow in males; both sexes with a large lateral orange macula on each elytron. Dorsal surface with a slightly golden sheen.

Description

Holotype. Male. Length 2.4 mm, width 1.9 mm; body oval-rounded, convex. Dorsal color black except head pale yellow with black clypeus; pronotum with lateral angles and anterior margin pale yellow; elytron with a large oval orange lateral macula, the macula touches the lateral margin and the humeral angle (Fig. 46), the black color shows a slightly golden sheen. Ventral surface with head, prosternum, metaventrite and abdomen dark brown; legs, mouthparts and antennae yellow. Head punctures small, separated by their diameter, each puncture as large as an eye facet; pronotal punctures larger than head punctures, separated by their diameter to 1½× their diameters; elytral punctures larger than pronotal punctures, separated by their diameter; metaventral punctures as large as elytral punctures, separated by 1½× their diameter. Clypeus straight. Prosternal carinae not examined. Protibia not flanged, basal tooth small, sponda extended beyond protibial margin (Fig. 47). Basal abdominal ventrite without setal tuft. Abdomen with postcoxal line on basal abdominal ventrite curved, touching the posterior ventrite margin, ventrite with sparse, long pubescence and small, dense punctures. Ventrites II-IV with a tubercle on each side of middle, each pair of tubercles more separated than the anterior ventrite; ventrites V and VI flattened and emarginated (Fig. 48). Ventrites II-VI pubescent throughout, punctures fine, dense. Genitalia with penis guide as long as parameres, symmetrical, sides slightly convergent, slightly wider before apex, apex acute (Fig. 50); parameres curved, narrowed to apex, apex acute, setae arising from convex side and the apex (Fig. 49); penis curved in basal ½, apex without alae, apex with a large apical ampulla, basal capsule without crest, inner arm of basal capsule small and subtriangular (Fig. 51).

Female. Length 2.5 mm, width 1.9 mm. Similar to male except head, pronotum and basal part of femurs black; abdomen not modified. Genitalia not examined.

Variation

The only known variation is shown by the female paratype.

Etymology

From the Latin word aurantius (= orange) and pleurón (= sides). The name refers to the dorsal color pattern.

Type locality

COSTA RICA, Puntarenas: Monteverde, San Luis, 10°17'11"N, 84°48'5"W.

Distribution

Costa Rica.

Discussion

This species is easily recognizable by its dorsal coloration and the gold sheen on the dorsal surface. The species is very similar to Hyperaspis panzosae Gorham, but the specimen described by Gorham is a female and the author described the specimen having black legs (however, the figure in Gorham’s table has orange legs instead), while the female described here has only the basal half of femurs black; the author did not describe any sheen for H. panzosae, while aurantiapleura has a gold sheen on the dorsal surface. There are several species of Hyperaspis with similar coloration to panzosae like H. excelsa Fall and H. cruenta LeConte, so that without examining the original material there is no evidence to assume panzosae as Brachiacantha instead of Hyperaspis; therefore, we decided to maintain aurantiapleura as a separate species.

Brachiacantha isthmena Nestor-Arriola, Toledo-Hernández & Solís, sp. nov.

Figures 7, 52–55

Holotype

Male, pinned, with genitalia in a separate microvial. Original labels: “PANAMÁ, Cerro Campana, 3000’. July 29 1970, H. and A. Howden” (USNMNH).

Paratypes

Costa Rica • 1♂; Limón, Res. Biol. Hitoy Cerere; 100 m; 6–16 May 1992; G. Carballo leg.; L-N 184200_643300 (MNCR). 1♂; Limón, Valle de la Estrella, R. B. Hitoy Cerere; 100 m; 04 Mar. 1994; G. Carballo leg.; L-N 643400_184600, #2889” (MNCR). Panamá • 1♂; same data as the holotype; 31 July 1970 (USNMNH). 1♂; Pr. Cerro Campana; 8°40'N, 79°56'W, 850 m; 14 July 1971; H. A. Hespenheide leg. (USNMNH). 1♂, same data, 10 Feb. 1973; W. Bivin leg. (USNMNH).

Diagnosis

Dorsal color black or dark brown; pronotum almost completely yellow; each elytron with three or four yellow spots. Male abdomen with a setal tuft on the first ventrite and several ventrites emarginated and depressed.

Description

Holotype. Male. Length 3 mm, width 2.2 mm, body oval, dorsally convex. Dorsal color black except head pale yellow; pronotum pale yellow with only a basal black macula; each elytron with four yellow spots in humeral, discal, mid-lateral and sub-apical positions (Fig. 7). Ventral surface with head, prosternum, metaventrite and abdomen dark brown; legs, mouthparts and antennae yellow. Head punctures small, separated by their diameter, each puncture as large as an eye facet; pronotal punctures larger than head punctures, separated by their diameter; elytral punctures as large as pronotum punctures, separated by their diameter; metaventral punctures as large as elytral punctures, separated by their diameter. Clypeus emarginate. Carinae on prosternal process straight, convergent, joined near the anterior prosternal border, larger than half prosternum. Protibia not flanged, basal tooth small. Basal abdominal ventrite with setal tuft at center. Abdomen with postcoxal line on basal abdominal ventrite slightly flattened along posterior ventrite margin, ventrite with sparse, long pubescence and small, dense punctures. Ventrites II-VI depressed and emarginated, punctures fine, dense. Genitalia with penis guide as long as parameres, symmetrical, slender, sides slightly pinched at basal half, widened in the apical third, apex acute (Fig. 53); parameres slender, basally wide, narrowed to apex, apex rounded, setae rising from the convex side margin and apex (Fig. 52); penis robust, strongly curved in basal half, widened in apical third with a narrowing before apex, alae short and slender, basal capsule without crest, inner arm long, parallel sided, rounded at apex (Figs 54, 55).

Female. Unknown.

Variation

Unknown.

Etymology

The name refers to the Panamá isthmus.

Type locality

PANAMÁ, Panamá Oeste: Parque Nacional Altos de Campana, Cerro de la Campana, 8°42'49.4"N, 79°57'10.24"W.

Distribution

Costa Rica and Panamá.

Discussion

The coloration of the species is similar to that of the dentipes group of species, but B. isthmena lacks the ventral cusps of the male, instead it shows a setal tuft on the first ventrite.

ursina group

Abdomen of male with the 5th ventrite modified; penis guide of male genitalia symmetrical, apically truncate (Gordon 1985).

Brachiacantha guatemalensis (Gorham), comb. nov.

Figures 9, 56–58

Hyperaspis guatemalensis Gorham, 1894: 200, tab. XI, fig. 10.

Material examined

Guatemala • 1♂; Yepocapa; 01 May 1948; H. T. Palmat leg. (USNMNH).

Diagnosis

Length 3 mm, width 2.4 mm; body rounded, convex. Dorsal color black, pronotum with lateral angles and anterior border yellow, each elytron with an irregular yellow macula covering from the humeral angle to the posterior border (Fig. 9). Ventral surface with head, prosternum and metaventrite black; abdomen yellow except the center of ventrites I-III black; legs, mouthparts, and antennae yellow. Carinae on prosternal process parallel, convergent near the anterior margin of prosternum, not joined. Protibia not flanged, basal tooth small. Ventrites I-IV truncate at apex; ventrites V and VI emarginated and depressed. Male abdomen with several ventrites truncate at center. Genitalia with penis guide as long as parameres, symmetrical, sides slightly convergent, apex truncate (Fig. 58); parameres parallel sided, apex rounded, setae rising from the apex border long and curved; penis curved in basal ½, apex with subtriangular alae, basal capsule not crested, inner arm of basal capsule long and slender (Figs 56, 57).

Figures 9–12. 

Brachiacantha sp. 9 Brachiacantha guatemalensis comb. nov., male 10 Brachiacantha papiliona sp. nov., female 11 Brachiacantha mimica sp. nov., female 12 Brachiacantha gorhami comb. nov., female.

Female. Unknown.

Variation

Unknown.

Distribution

Guatemala.

Discussion

This species is easily identifiable by the dorsal black coloration with elytral and pronotal margins yellow. The specimen examined was identified following the description and illustrations of Gorham (1894) as Hyperaspis guatemalensis. Nevertheless, this species has all the diagnostic characters of Brachiacantha, including the tibial tooth, lack of large abdominal pores and eye emarginated by the eye canthus; in addition, there are several species of Hyperaspis with coloration similar to H. limbalis Casey and H. caseyi Gordon, but the individual examined presents all the external characteristics, size and distribution of H. guatemalensis. Therefore, a new combination is proposed. Several species described by Gorham have been reassigned into other genera due taxonomic revisions (Gordon 1981); about the genus Hyperaspis there are several examples: H. pauperula was reassigned to Calloeneis, H. adelaia is considered now as a synonym of Cyrea tessulata (Mulsant) and H. terminata was reassigned to Serratitibia (Canepari et al. 2016; Gordon 1981; Gordon et al. 2013). According the present evidence H. guatemalensis should be reassigned to Brachiacantha, but an examination of the type material is needed to ensure the taxonomic identity with the material examined here.

invertita group, new group

Male genitalia with penis guide with sides slightly convergent, almost parallel; apex almost truncate with a triangular projection (Fig. 58). Parameres slender, apically narrowed (Fig. 59).

Brachiacantha invertita Nestor-Arriola, Toledo-Hernández & Solís, sp. nov.

Figures 59–64

Holotype

Male, pinned, with genitalia in a separate microvial. Original labels: “COSTA RICA, PUNTARENAS, A. C. Arenal, Buen Amigo, San Luis, 1000–1350 m, May 1994, col. Z. Fuentes, L N 250850_449250, #2926” (MNCR).

Paratypes

Costa Rica • 1♂; Cartago, Birrisito; 1400 m; 07 Aug. 1980; D. R. Whitehead leg. (USNMNH). 1♀; Cartago, Turrialba, Tayutic, P. N. Barbilla, Sector Cerro Tigre; 1500–1600 m; 22 June-22 July 2001; W. Arana leg.; malaise trap; L_N_211700_602500, #64679 (MNCR); 1♀; Cartago, P. N. Tapantí, Quebrada Segunda; 1250 m; Oct. 1992; G. Mora leg.; L-N 194000_560000 (MNCR). 1♀; Guanacaste, R. F. Cord. Guanacaste, Rio San Lorenzo, Tierras Morenas; 1050 m; Dec. 1991; C. Alvarado leg.; L-N 287800_427600 (MNCR). 2♂; same data; ene.1992 (MNCR). 1♀; Guanacaste, Est. Cacao, SW side Volcán Cacao; 1000–1400 m; Apr. 1988; GNP. Biodiv. Survey exped.; 323300_375700 (MNCR). 1♂; same data; malaise trap; 1988–1989 (MNCR). 1♀, 1♂; Guanacaste, A. C. Guanacaste, Sect. Las Pailas; 800 m; 6–26 June 1994; K. Taylor leg.; L N 309500_389500, #3063 (MNCR). 1♀; Guanacaste, Tierras Morenas; 650 m; 8–10 Feb 1994; Z. Fuentes leg.; L S 283950_424500, #2616 (MNCR). 1♀; Guanacaste, P. N. Rincon de la Vieja, Est. Las Pailas; 800 m; 10–21 Nov. 1993; M. A. Zumbado and D. G. García leg.; L N 306300_388600, #2559 (MNCR). 2♀, 6♂; Guanacaste, sect. Las Pailas, 4.5 km SW Rincón de la Vieja volcano; 800 m; 12 Apr.-4 May 1995; K. Taylor leg.; L_N_306300_388600, #6194 (MNCR). 1♂; Guanacaste, Z. P. Tenorio, Rio San Lorenzo, Tierras Morenas; 1050 m; Aug. 1992; G. Rodriguez leg.; L-N 287800_427600 (MNCR). 1♀; same data, Oct. 1992 (MNCR). 1♂; Guanacaste, Z. P. Tenorio, Rio San Lorenzo, Tierras Morenas; 1050 m; 28 Mar.-21 Apr. 1992; M. Segura leg.; L-N 287800_427600 (MNCR). 1♀; Puntarenas, Monteverde, San Luis; 1000–1350 m, Feb. 1994; Z. Fuentes leg.; L-N 449250_250850, #2615 (MNCR). 1♀, 1♂; Puntarenas, Monteverde, San Luis; 1040 m; 24 Aug.-15 Sep. 1992; F. A. Quesada leg.; L-N 250850_449250 (MNCR). 1♀; Puntarenas, R. B. Monteverde, est. La Casona; 1520 m; July 1992; N. Obando leg.; L N 253200_449700 (MNCR). 1♀; San José, Moravia, San Jeronimo; 1400 m; 05 Dec. 2008; A. S. Saenz leg.; L N 222050_535800, #95353 (MNCR).

Diagnosis

Dorsal color yellow with black spots. Male abdomen with several ventrites depressed and emarginate. Penis guide of male genitalia with an apical triangular projection, penis with long slender alae.

Description

Holotype. Male. Length 3.6 mm, width 2.0 mm; body oval, convex. Dorsal color yellow except pronotum, with a small basal black spot on each side of the disk center, a small black basal spot and a black subquadrate spot on each side at ½ from the lateral margin; scutellar shield black; each elytron with five black spots: scutellar spot slightly oval, sub-humeral spot large and oval, mid-lateral spot at the apical half of the elytron, discal and sub-apical spots rounded, the spots do not touch the elytral margin, the elytral suture or the anterior border (Fig. 59). Ventral surface with head, prosternum, metaventrite and abdomen black; legs, mouthparts, and antennae yellow. Head punctures small, separated by their diameter, each puncture slightly larger than an eye facet; pronotal punctures larger than head punctures, separated by their diameter; elytral punctures as large as pronotal punctures, separated by their diameter; metaventral punctures larger than elytral punctures, separated by more than three diameters at center, about one diameter on sides. Clypeus straight. Carinae on prosternal process curved, divergent, not joined, larger than half the prosternum. Protibia not flanged, basal tooth small. Basal abdominal ventrite without setal tuft. Abdomen with postcoxal line on basal abdominal ventrite slightly flattened along posterior ventrite margin; ventrite with sparse, long pubescence and small, dense punctures. Ventrites III-VI depressed and emarginated (Fig. 60). Ventrites II-VI pubescent throughout, punctures fine, dense; third ventrite with a tubercle on each side of middle. Genitalia with penis guide as long as parameres, symmetrical, sides slightly convergent, apex almost truncated but with a triangular appendage (Fig. 61); parameres narrowed to apex, apex acute, setae rising from the convex side border (Fig. 62); penis curved in basal ½, apex with long slender alae, basal capsule crested, inner arm of basal capsule long and slender (Figs 63, 64).

Female. Similar to male except abdominal characters. Genitalia not examined.

Variation

Males. Length 2.9–3.1 mm, width 2.2–2.7 mm. Females. Length 2.6–3.8 mm, width 2–2.9 mm.

Etymology

From the Latin inversos (= invert). The name refers to the dorsal color pattern, yellow with five black spots, the inverse of that of many Brachiacantha species (black with five yellow spots).

Type locality

Costa Rica, Puntarenas: Monteverde, San Luis, 10°17'11"N, 84°48'5"W.

Distribution

Costa Rica.

Discussion

This species is easily identifiable by the dorsal coloration, yellow with black spots.

cachensis group, new group

Male genitalia with penis guide “guitar” shaped, parallel sided at basal ⅓, widened at 2/3, apically emarginated; parameres subtriangular, rounded at apex.

Brachiacantha cachensis Gorham

Figures 8, 65–68

Brachyacantha cachensis Gorham, 1894: 190. Leng 1911: 322.

Material examined

71 specimens from Costa Rica and Panamá (FSCA, USNMNH, MNCR, MUCR, MZCR). Costa Rica: Alajuela, Cartago, Guanacaste and Puntarenas. Panamá • 1♀; Chiriqui Prv., cont’l Divide trail; 04 Jul 1997; Wappes and Morris leg.; Brachiacantha sp. R. Gordon 1998 (USNMNH). 1♀, 1♂; Chiriqui Prv., cont’l Divide trail; 04 Jul 1997; Wappes and Morris leg. (USNMNH). 1♀, 1♂; Chiriqui Prv., cont’l Divide trail; 08 Jul 1997; J. Huether leg. (USNMNH). 1♀; Chiriqui Prv., “Finca La Suiza”; 07 Jul 1997; J. Huether leg. (USNMNH).

Diagnosis

Dorsal color pale yellow to gray with brown or black spots. Ventrites I-VI depressed and emarginated, ventrites I-III with abundant and longer pubescence in the depressed part. Genitalia with penis guide slightly longer than parameres, symmetrical, basal ⅓ with almost straight sides, abruptly wide at apical 2/3 (Fig. 65); parameres wide basally, narrowed to apex, apex rounded (Fig. 66); penis curved in basal ½, widened at the apical ⅓, narrowed before apex, alae rounded, basal capsule with small crest, inner arm of basal capsule long and slender with parallel sides (Figs 67, 68).

Variation

The basal macula over pronotum can be disrupted, forming a macula over the posterior margin with a pair of small spots at the center of the pronotum. In some specimens the sub-humeral spot is half-moon shaped.

Distribution

Costa Rica and Panamá. New record to Panamá.

Discussion

This species is similar to B. dentata, but the males can be differentiated by the lack of a ventral tooth and having the central part of the ventrites depressed with abundant pubescence. Within the groups defined by Gordon et al. (2014) for the South American species based on the genitalia of the male, this species does not correspond to any of them, due to the emarginated apex of the penis guide. The material was identified following the descriptions of Gorham (1894) and the keys and illustrations of Leng (1911).

Brachiacantha hexaspina González, Větrovec & Nestor-Arriola, sp. nov.

Figures 13–20

Holotype

Male, pinned, with genitalia in a separate microvial. Original labels: “PANAMÁ: Veraguas, 12.IX.2017, Santa Fé env., Alto de Piedra-Cerro Tute, 08°30.8'N, 81°07.2'W, 850–1360 m, lower montane forest, individual collecting, Fikàček, Seidel and Sekerka, lgt.” (NMP) (1♂).

Paratypes

Costa Rica • 1♀; Alajuela, P. N. V. Tenorio, Rio Robles, Palmital, “arriba la Caliza”; 1000–1100 m; 21 June-16 Sep.; J. A. Azofeifa leg.; malaise trap; LN296500_426300, #87178 (MNCR). 1♀; Limón, Res. Biol. Hitoy Cerere, est. Hitoy Cerere; 100 m; Oct. 1992; G. Carballo leg.; L-N 184600_643300 (MNCR). 1♀; Limón, Valle de la Estrella, R. B. Hitoy Cerere, send. Espavel; 560 m; 18 Sep.-5 Oct. 2003; B. Gamboa, E. Rojas and W. Arana leg.; malaise trap; L-S 401200_569800, #75488 (MNCR). 1♀; Puntarenas, Golfito, Est. Agujas, Cerro Rincon; 645–745 m; 15 Apr.–15 May 2000; A. Azofeifa leg.; malaise trap; L_S_275500_521950, #56675, sp#106 (MNCR).

Diagnosis

Dorsal color yellow with two black spots in each elytron. Pro-, meso-, and metatibia with external tooth. Male with ventrites II-V emarginate and depressed, forming a longitudinal fossa on abdomen. Male genitalia with penis guide symmetrical, apically notched, penis with a strong curvature at apical 1/6 toward external side.

Figures 13–20. 

Brachiacantha hexaspina sp. nov. male 13 Dorsal view 14 Abdomen 15 Tegmen 16 Tegmen lateral view 17 Penis 18 Penis apex 19 Paramere 20 Penis guide. Scale bar: 1 mm, except 15–20 0.5 mm

Description

Holotype. Male. Length 4 mm, width 2.9 mm; body oval, slightly elongate, convex. Dorsal color yellow except pronotum with two basal, irregular light brown transparent spots, roughly round, diameter approximately 2/5 the length of pronotum; scutellar shield dark brown; elytron with two black spots: humeral spot irregular and slightly subquadrate, sub-apical spot large, twice the size of the humeral spot, subquadrate and with the basal side trilobed, closer to the lateral edge than to the suture; elytral suture and anterior margin black, lateral margin black until apical ⅓ (Fig. 13). Ventral surface yellow with head and prosternum brown; meso-, metasternum and abdomen dark brown; mouthparts and antennae yellow, mandibular apex black; legs yellow, trochanters light brown. Head punctures small, separated by less than half their diameter, each puncture larger than twice an eye facet; pronotal punctures as large as head punctures, separated by half to one diameter, more compact towards the edges; elytral punctures as large as pronotum punctures, separated by two diameters, irregular lines of large and apparently black dots on the anterior margin of elytra, the basal half of the lateral margin and each side of the elytral suture, the lines are separated from the elytral margins and the elytral suture by approximately the scutellar shield length; metaventral punctures as large as elytral punctures, separated by 1½× their diameters. Clypeus straight. Carinae on prosternal process straight, larger than half prosternum, convergent to prosternal anterior border without reaching it, not joined. Protibia flanged, basal tooth large, curved, as large as protibial width; mesotibia not flanged, basal tooth as large as protibial tooth; metatibia not flanged, basal tooth shorter than protibial tooth. Basal abdominal ventrite without setal tuft. Abdomen with postcoxal line on basal ventrite slightly flattened along posterior ventrite margin, slightly recurved towards the anterior margin at the external ⅓, without reaching the lateral or anterior borders; all ventrites with short and sparse pubescence at sides, more long and less sparse from I–V; ventrite punctures variable, fine and dense, larger towards the lateral and posterior edges. Ventrites II–V emarginated and depressed at middle, ventrite VI flattened, slightly depressed centrally, posterior margin emarginated. Ventrites III-IV with a tubercle at each side of the emargination, II and V only with a slight convexity. Ventrites I and II with long, bushy pubescence at middle, III–V glabrous at the middle (Fig. 14). Genitalia with penis guide longer than parameres, symmetrical, wide, basal ⅓ with almost straight sides, abruptly wide at apical ⅔, wider at apex, sides of the wide part slightly emarginated, apex slightly emarginated (Figs 15, 20), apex sinuous in lateral view (Fig. 16); parameres wide basally, narrowed to apex, apex rounded, setae rising from the apex border (Figs 16, 19); penis curved in basal ½, bended forming a right angle at the apical 1/6, narrowed before apex, alae narrowed apically (Fig. 18), basal capsule crested, heavily sclerotized, inner arm of basal capsule perpendicular to penis, long and slender with parallel sides, external arm in the direction of the tube, twice as long as wide, ending in a short projection (Fig. 17).

Female. Similar to male except the abdomen characters; ventral surface with head, prosternum, metasternum and abdomen orange; legs, mouthparts, and antennae yellowish orange. Mesotibial basal tooth smaller than protibial tooth; metatibial basal tooth shorter and wider than mesotibial tooth. Genitalia not examined.

Variation

Length 3.6 to 4.6 mm. Width 2.8–3 mm.

Etymology

The name derives from Greek hexa (= six) and Latin spine (= thorn), referring to the six teeth of the tibiae, long and curved like rose thorns.

Type locality

PANAMÁ, Veraguas: Santa Fé, Alto de Piedra, 08°30.8'N, 81°07.2'W.

Distribution

Costa Rica and Panamá.

Discussion

This species is easily identifiable by the pale dorsal coloration and the toothed pro-, meso- and metatibia. The presence of tibial spines on the middle and hind legs is a feature not previously observed in the genus. According to the groups defined by Leng (1911), this species belongs to group V, characterized by the presence of an abdominal canal that covers ventrites II-V, within which only B. cachensis Gorham has a similar design but whose male genitalia are different. Within the groups defined by Gordon et al (2014) for the South American species based on the genitalia of the male, this species does not correspond to any of them, due to the notch at the apex of the penis guide. Despite these peculiarities, the similarities in the abdominal ventrite modifications with other species from Central America, the lack of abdominal primary pores, and the presence of protibial tooth make it prudent to keep it in Brachiacantha.

Species not associated with males.

Brachiacantha papiliona Nestor-Arriola, Toledo-Hernández & Solís, sp. nov.

Figure 10

Holotype

Female, pinned, with genitalia in a separate microvial. Original labels: “PANAMÁ, Chiquiri. Prov., cont’l Divide trail, 04-VII-1997, col. Wappes and Morris” (USNMNH) (1♀).

Paratypes

Costa Rica • 1♀; Guanacaste, Est. Cacao, SW side Volcan Cacao, 1000–1400 m; 1988–1989; GNP. Biodiv. Survey exped.; malaise trap; 323300_375700 (MNCR). 1♀; Puntarenas, Monteverde, San Luis; 1000–1350 m; Feb. 1994; Z. Fuentes leg.; L N 449250_250850, #2615 (MNCR).

Diagnosis

Dorsal color pale yellow with black macula. Elytron with part of the elytral suture black, a sub-apical black spot and a large black spot on the anterior margin.

Description

Holotype. Female. Length 3 mm, width 3.6 mm; body oval-rounded; convex. Dorsal color pale yellow except pronotum with the central part black; scutellar shield black; each elytron with three black spots: a macula over the elytral suture, a large oval spot at the center of the second half of the elytron and an irregular macula over the anterior margin of the elytron (Fig. 10). Ventral surface with head, prosternum, meso- and metasternum and abdomen black; legs, mouth parts and antennae yellow. Head punctures small, separated by their diameter or less, each puncture as large as an eye facet; pronotal punctures larger than head punctures, separated by less than their diameter; elytral punctures as large as pronotal punctures, separated by their diameter; metaventral punctures larger than elytral punctures, separated by 1½× their diameter to two diameters. Elytral suture with apex dentiform. Clypeus straight. Carinae on prosternal process convergent, joined, longer than half prosternum. Protibia not flanged, basal tooth small. Basal abdominal ventrite without setal tuft. Abdomen with postcoxal line on basal abdominal ventrite slightly flattened along posterior ventrite margin, ventrite with sparse, long pubescence and small, dense punctures. Ventrites II-VI pubescent throughout, punctures fine, dense. Genitalia with two curved sclerotized arms at bursa copulatrix; apical strut long, flattened laterally, straight sided, slightly widened at apex; spermathecal capsule slender, widened and rounded at cornu.

Male. Unknown.

Variation

Unknown.

Etymology

The name refers to the basal elytral spot in the shape of butterfly wings.

Type locality

Panamá, Chiriqui: Hornito, 8°44'50"N, 82°13'84"W.

Distribution

Panamá and Costa Rica.

Discussion

This species resembles B. hazel Gordon et al. (2014) for the basal macula on the elytron, but the color pattern is different in lacking the sub-apical spot over the sutural margin. Gordon and collaborators did not describe the female of B. hazel, but the missing sub-apical elytral spot is not associated with sexual dimorphism in Brachiacantha species. Another remarkable character of B. papiliona is the presence of a tooth at the apex of the elytral suture, a rare character among Coccinellidae.

Brachiacantha tica Nestor-Arriola, Toledo-Hernández & Solís, sp. nov.

Figures 69–70

Holotype

Female, pinned. Original label: “COSTA RICA, PUNTARENAS, Península de Osa, Rancho Quemado, 200 m, Sep. 1992, col. A. Marín, L-S 292500_511000” (MNCR).

Paratypes

Costa Rica • 1♀; Puntarenas, same data than the holotype; 10 Sep.-10 Oct. 1993 (MNCR). 1♀; same data as holotype; Dec. 1992; F. A. Quesada leg.; (MNCR). 2♀; Puntarenas, P. N. Corcovado, send. Rio Claro; 1991; L-S 508300_270500 (MNCR). 1♀; Puntarenas, P. N. Corcovado, Est. Sirena; 0–100 m; Mar.-June 1991; L-S 270500_508300 (MNCR). 1♀; same data, 17 June-4 Sep. 1991; malaise trap; (MNCR).

Diagnosis

Dorsal color black; pronotum anterior angles pale yellow; each elytron with three pale yellow spots, one near the scutellar shield and two near the apex, no humeral or mid-lateral spots. Sponda large and conspicuous.

Description

Holotype. Female. Length 2.3 mm, width 1.7 mm; body oval, convex. Dorsal color black except head pale yellow; pronotum with lateral angles pale yellow; elytron with three pale yellow oval spots: scutellar shield spot small, small discal spot in the apical half of elytra, sub-apical spot large and oval (Fig. 69). Ventral surface with head, prosternum, metaventrite and abdomen dark brown; legs, mouthparts and antennae yellow. Head punctures small, separated by their diameter, each puncture as large as an eye facet; pronotal punctures larger than head punctures, separated by their diameter; elytral punctures as large as those on pronotum, separated by their diameter; metaventral punctures larger than elytral punctures, separated by their diameter to two diameters. Clypeus slightly emarginated. Carinae on prosternal process straight, larger than half prosternum, convergent, joined near the anterior margin of prosternum but not touching it. Protibia not flanged, basal tooth small, sponda obvious (Fig. 70). Basal abdominal ventrite without setal tuft. Abdomen with postcoxal line on basal abdominal ventrite curved, only touching the posterior ventrite margin, ventrite with sparse, long pubescence and small, dense punctures. Ventrites II-VI pubescent throughout, punctures fine, dense. Genitalia not examined.

Male. Unknown.

Variation

Length 2.1–2.6mm. Width 1.5–1.8 mm.

Etymology

The species name is a popular name for the inhabitants of Costa Rica.

Type locality

COSTA RICA, Puntarenas: Península de Osa, Rancho Quemado, 8°40'54.88"N, 83°33'33.81"W.

Distribution

Costa Rica.

Discussion

Although having three spots on each elytron is common in several Brachiacantha species, the particular pattern of B. tica, with one spot in the basal half of the elytron and two in the apical half, is not shared with any other species of the genus.

Brachiacantha mimica Nestor-Arriola & Toledo-Hernández, sp. nov.

Figure 11

Holotype

Female, pinned. Original labels: “GUATEMALA, ANTIGUA, 1500–1600 m, 01-VII-1980, col. N. L. H. Krauss” (USNMNH).

Diagnosis

Dorsal color black with three pale yellow spots. Protibial tooth small. Apical elytral suture dentiform.

Description

Holotype. Female. Length 2.8 mm, width 2.2 mm; body oval rounded, convex. Dorsal color black except head pale yellow; pronotum with lateral angles and sides yellow; each elytron with three yellow spots in discal, mid-lateral and sub-apical positions (Fig. 11). Ventral surface with head, prostermum, and metaventrite brown; abdomen yellowish brown; legs, mouthparts, and antennae yellow. Head punctures small, separated by their diameter or less, each puncture as large as an eye facet; pronotal punctures larger than head punctures, separated by their diameter or less; elytral punctures as large as pronotum punctures, separated by their diameter; metaventral punctures as large as head punctures, separated by their diameter to two diameters. Elytral suture with apex dentiform. Clypeus curved. Carinae on prosternal process convergent, joined near the anterior prosternal border, larger than half prosternum. Protibia not flanged, basal tooth small. Basal abdominal ventrite without setal tuft. Abdomen with postcoxal line on basal abdominal ventrite slightly flattened along posterior ventrite margin, ventrite with sparse, long pubescence and small, dense punctures. Genitalia not examined.

Male. Unknown.

Variation

Unknown.

Etiology

The name refers to its similarity to other Brachiacantha species.

Type locality

GUATEMALA, Sacatepequez: Antigua Guatemala, 14°33'N, 90°44'W.

Distribution

Guatemala.

Discussion

This species is very similar to B. bistripustulata and allies, but the pale-colored head and the yellowish ventral color of mimica marks the difference. The apical dentate elytral suture is shared only with B. papiliona.

Brachiacantha gorhami (Weise), comb. nov.

Figure 12

Coccinella pantherina Gorham, 1892: 161, tab. IX, fig. 9 (non Coccinella pantherina Linnaeus, 1758: 368) (non Coccinella pantherina DeGeer, 1773).

Coccinella gorhami Weise, 1904: 357.

Material examined

El Salvador • 1♀; Montecristo, 23 km N of Metapan; 2300 m; 10 May 1971; H. F. Howden leg. (USNMNH).

Diagnosis

Dorsal color pale yellow except head with brown clypeus and black above the eyes; pronotum with a black macula on the posterior margin and two large convergent subtriangular spots at center; scutellar shield black; each elytron with five black marks: a vitta on the elytral suture extending laterally in the second half of the elytron, an oval spot at the humeral callus, an elongated spot parallel to the sutural margin in the basal half of the elytron, an oval spot at middle and a small spot on the lateral margin in the apical half of the elytron (Fig. 12). Ventral surface brown. Protibial tooth small.

Variation

Unknown.

Distribution

El Salvador and Guatemala. New record to El Salvador.

Discussion

The material was identified using the descriptions and tables of Gorham. The species was described by Gorham as Coccinella pantherina, but the name was preoccupied by a junior synonym of Adalia bipunctata (Linnaeus, 1758) and a junior synonym of Adalia decempunctata (DeGeer, 1773). Weise (1903) noticed the homonymy and created the name Coccinella gorhami for the species.

Although Gorham placed the species as a member of the genus Coccinella, he compared it with the species B. lepida, thus indicating the small size of the species. In the last century, several species originally described as Coccinella have been revised and assigned to other genera such as Mulsantina and Cycloneda, but there are still several species which need revision. The specimen examined corresponded to the description and figures of Gorham (1892), but the presence of eye emarginated by the eye canthus, the lack of large abdominal primary pores between fourth and fifth ventrites and the presence of a small tibial tooth identified the species as a member of Brachiacantha. The dorsal coloration is unique among Brachiacantha species but is similar to those of other genera like Dilatitibialis, Serratitibia, and Cyrea. A revision of the original material may be needed to confirm the taxonomic changes.

Figures 21–24. 

Brachiacantha nubes sp. nov., male 21 Dorsal view 22 Abdomen 23 Tegmen, lateral view 24 Tegmen, concave side view.

Figures 25–28. 

Brachiacantha fenestrata, male genitalia 25 Tegmen 26 Tegmen lateral view 27 Penis 28 Penis apex.

Figures 29–32. 

Brachiacantha octostigma, male genitalia 29 Tegmen, lateral view 30 Tegmen 31 Penis 32 Penis apex.

Figures 33–36. 

Brachiacantha bipartita, male genitalia 33 Tegmen, lateral view 34 Tegmen 35 Penis 36 Penis apex.

Figures 37–40. 

Brachiacantha aperta, male genitalia 37 Penis 38 Penis apex 39 Tegmen, lateral view 40 Tegmen.

Figures 41–45. 

Brachiacantha dentata sp. nov., male 41 Dorsal view 42 Abdomen 43 Tegmen, lateral view 44 Tegmen 45 Penis.

Figures 46–51. 

Brachiacantha aurantiapleura sp. nov., male 46 Lateral view 47 Protibia 48 Abdomen 49 Tegmen, lateral view 50 Tegmen 51 Penis.

Figures 52–55. 

Brachiacantha isthmena sp. nov., male genitalia 52 Tegmen, lateral view 53 Tegmen 54 Penis 55 Penis apex.

Figures 56–58. 

Brachiacantha guatemalensis comb. nov., male 56 Penis 57 Penis apex 58 Tegmen.

Figures 59–64. 

Brachiacantha invertita sp. nov., male 59 Dorsal view 60 Abdomen 61 Tegmen 62 Tegmen lateral view 63 Penis 64 Penis apex.

Figures 65–68. 

Brachiacantha cachensis, male genitalia 65 Tegmen 66 Tegmen lateral view 67 Penis 68 Penis apex.

Figures 69, 70. 

Brachiacantha tica sp. nov., female 69 Lateral view 70 Protibia.

Discussion

Twenty-five species of the genus Brachiacantha are found in Central America, including nine new species. Some species such as B. fenestrata, B. cachensis, or B. aperta are easily identifiable without doubt; however, a revision of the original material is pending for the species B. guatemalensis (Gorham), comb. nov. and B. gorhami (Weise), comb. nov. Although it is not advisable to describe species based only on female specimens, the species B. papiliona sp. nov. and B. tica sp. nov. have enough external characters to ensure that they were not previously described. The species B. mimica sp. nov. has a very common dorsal coloration for Brachiacantha and Hyperaspis species, but the apical tooth of the elytral suture has not been described for Coccinellidae species with the exception of B. papiliona.

Conclusions

Our knowledge of the Coccinellidae species in both North America and South America, has changed in the last century; however, Central America is still poorly studied. In this study several new species were discovered by reviewing collections of the United States (USNMNH, FSCA), England (OUMNH), México (CEAM, CNIN), Costa Rica (MNCR, MUCR, MZCR), and Czech Republic (NMP), but there are poorly known collections in Central American countries and the biodiversity crisis makes the need for new studies urgent to record the biological diversity of the region. A detailed study is needed of the phylogenetic relationships in the genus Brachiacantha to elucidate the origin of the male genitalia diversity within the genus and if the species groups are monophyletic units.

Acknowledgements

This study was made possible by the financial support of the National Commission of Science and Technology (CONACYT). For allowing access to the collections and facilitating the loan of collection material, we thank Natalia Vandenberg and Elisabeth Roberts (USNM), Paul Skelley (FSCA), Darren Mann and Amoret Spooner (OUMNH), Jesús Romero-Nápoles and José Refugio-Lomelí (CEAM), Santiago Zaragoza-Caballero and Cristina Mayorga (CNIN), German Vega (MNCR), Paul Hanson (MZCR) and Humberto Lezama Ulate (MUCR). Thanks to Jorge Valdez Carrasco (CEAM) for taking some of the photographs in this article.

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