The Brachiacantha Dejean, 1837 (Coleoptera, Coccinellidae) of Central America

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.


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: 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

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

Brachiacantha octostigma Mulsant
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. 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).

Brachiacantha bipartita Mulsant
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). Brachyacantha aperta Weise, 1903: 208. Leng 1911 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).
Discussion. This species is easily recognizable due to its unique coloration. 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.

Brachiacantha dentata
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.
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). 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.

Brachiacantha aurantiapleura
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, ba-sal 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.
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. 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.

Brachiacantha isthmena
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).
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 5 th 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. 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 ir-regular 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).

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). 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.

Brachiacantha invertita
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).
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).
Discussion. This species is easily identifiable by the dorsal coloration, yellow with black spots.
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). 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.

Brachiacantha hexaspina
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  (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   gin, 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.
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.   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, proster-num, 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.

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

Brachiacantha tica
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) 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.
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.
Distribution. Guatemala. Discussion. This species is very similar to B. bistripustulata and allies, but the palecolored 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  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.

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.