Taxonomic revision of the New World genus Callotillus Wolcott (Cleridae, Tillinae), with the description of the new genus Neocallotillus, and an illustrated key of identification to species

Abstract The New World checkered beetle genus Callotillus Wolcott, 1911 is revised and the new genus Neocallotillus established. The subspecies Callotillus elegans vafer Wolcott is synonymized with the nominal subspecies, Callotillus elegans elegans (Erichson), which is transferred to, and designated as the type species of Neocallotillus gen. n. as Neocallotillus elegans (Erichson, 1847), comb. n. Two additional species are transferred from Callotillus to the new genus: Neocallotillus intricatus (Wolcott & Dybas, 1947), comb. n. and Neocallotillus crusoe (Wolcott, 1923), comb. n., the latter tentatively and based on Wolcott’s original description. Callotillus is now composed of two species: Callotillus eburneocinctus Wolcott, 1911 and Callotillus bahamensis Vaurie, 1952. All abovementioned species except Neocallotillus crusoe are diagnosed and redescribed. In the absence of reference material of Neocallotillus crusoe, Wolcott’s original description is transcribed. An illustrated key to species is provided. Characters of taxonomic relevance are illustrated and discussed. Updated distribution maps and locality data for all specimens examined are presented.


Introduction
Callotillus Wolcott is a genus of checkered beetles restricted to the Americas. The group has a wide distribution in the New World, extending throughout much of North and Central America and the West Indies (Fig. 8). Wolcott (1911) erected Callotillus for Tillus elegans Erichson and Callotillus eburneocinctus Wolcott, designating the latter as the type species. In later works, C. vafer Wolcott, C. crusoe Wolcott, and C. intricatus Wolcott & Dybas were described from North America, Puerto Rico and Costa Rica, respectively (Wolcott 1921(Wolcott , 1923Wolcott and Dybas 1947). Barr (1950) recognized C. vafer as a subspecies of C. elegans on the basis of apparent discontinuity in the geographic distribution of these species and subtle morphological variations observable throughout their geographic range. Finally, an expeditionary work in the West Indies by Vaurie (1952) retrieved a new species, Callotillus bahamensis. The genus Callotillus thus presently includes five species and two subspecies: Callotillus bahamensis Vaurie, C. crusoe Wolcott, C. eburneocinctus Wolcott, C. elegans Wolcott (including the subspecies C. e. elegans (Erichson) and C. e. vafer Wolcott), and C. intricatus Wolcott. In this paper we investigate the species of Callotillus within the context of a generic revision. All species comprising the two genera are diagnosed and redescribed, except N. crusoe, where the original description given by Wolcott (1923) is transcribed. An illustrated key and updated distribution maps are given for all the species treated here.

Material and methods
The taxonomic sampling consisted of approximately 300 specimens collected in North and Central America, and the West Indies. Male genitalia were dissected if more than one male per species was available. Genitalia extraction and dissection procedures are similar to those outlined by Ekis (1977). Most morphological terminology follows the work of Ekis (1977), Rifkind (1993), and Opitz (2010). Specimens were examined with a Leica MZ 7.5 stereomicroscope. Images were taken and measured using a Leica DFC 500 digital camera, and stacked using the software Zerene Stacker V. 1.04.
Information reflected in the distribution maps was obtained from the locality data of the material examined in this study, the original descriptions of all the species, and the works of Wolcott (1947), Corporaal (1950), Papp (1960), Barr (1975), Rifkind (1996), Leavengood (2008), and Burke et al. (2015).
Material used in this work was borrowed from the following collections: 1 Antennomeres 4-9 of males pectinate ( Elytral disc finely punctate, punctations irregularly arranged; median region of each elytron adorned with a transverse, light testaceous to almost whitish, slightly protruding fascia, and one protruding macula on the anterior half of the elytral disc, this macula can be absent in some specimens ( Fig. 1A-D Each elytron with a testaceous, broad and procurved fascia that initiates at the elytral suture and extends from the median region of the elytral disc to the elytral apex, and small, narrow, moderately oblique marking at the median region of the elytral disc; last third of elytral disc with a semicircular macula (Fig. 1F)  Each elytron adorned with a pair of elaborate, pale fasciae, and one macula arranged as follows: macula located on the anterior fourth, posterior to the humeral angle; one fascia located on anterior half of elytral disc, and strongly procurved, initiating on elytral suture and ending just before epipleural fold;  Pronotal disc dark testaceous to rufous; anterior half of elytral disc same color as pronotum, posterior half piceous, color transition interrupted by a median, transverse, pale fascia that extends from epipleural fold to elytral suture; median region of elytral disc depressed in lateral view (Fig. 1H) (Erichson, 1847), here designated. Differential diagnosis. Neocallotillus is most closely related to Callotillus. The new genus can be differentiated from Callotillus based on the following combination of characters: in Neocallotillus species the male has antennomeres 1-2 filiform; the third antennomere is moderately serrate; antennomeres 4-9 are strongly pectinate; and the tenth antennomere is ovoid in shape and laterally depressed ( Fig. 3A-C); the length of the tenth antennomere may vary by species. Females have antennomeres 1-3 filiform; the fourth antennomere is moderately serrate; and antennomeres 4-9 are robust, moderately, and gradually increase is size toward the distal end ( Fig. 3D-F); the tenth antennomere of females is similar to that of the males. Males of Callotillus have antennomeres 1-2 filiform; the third antennomere is moderately serrate; antennomeres 4-9 are strongly serrate and approximately equal in length; and the tenth antennomere is broadly ovoid and about the same length as antennomeres 8-9 combined (Fig. 3G, H). The antennal structure of females is similar to that of males, except antennomeres 4-9 are moderately serrate and the tenth antennomere is cylindrical to moderately ovoid (Fig. 3I, J). Additionally, Neocallotillus species are relatively slender and elongate (Figs 1A-E, 4A), while Callotillus species are conspicuously more robust (Figs 1F-G, 4B). Neocallotillus species lack an elytral swelling present on the anterior third of the elytral disc of Callotillus ( Fig. 4A-B). Neocallotillus also resembles Barrotillus Rifkind (Fig. 1I), however, the antenna of Neocallotillus is composed of 10 antennomeres ( Fig. 3A-F), while the antenna of Barrotillus has 11 antennomeres (Fig. 3K). The restricted distribution of Barrotillus, recorded only from a confined locality in Honduras, will also serve to separate it from the widely distributed Neocallotillus.
Elytra: Slender to feebly expanded posteriorly, elongate; median region of elytral disc feebly depressed in lateral view; sides subparallel to moderately expanded posteriorly in dorsal view; elytral declivity feebly to moderately gradual; elytral markings always present in various shapes, may be protruding or not.
Etymology: This generic name, which is preceded by the Latin prefix neo (new), refers to the superficial similarity to the genus Callotillus.
Remarks. Expressing tentative assignment of some of his species to Callotillus, Wolcott (1923) wrote: "Callotillus crusoe, as well as C. elegans and C. vafer, are placed in Callotillus provisionally only, as it differs from the other members of the genus in several important characters. No doubt, eventually, the creation of a new genus will be necessary for the reception of this new species and C. elegans and C. vafer. In C. eburneocinctus, the terminal segment of the maxillary palps is sub-cylindrical, the eyes are emarginate internally and the abdomen has but five segments. In C. elegans, C. vafer, and C. crusoe the maxillary palpi have the terminal segment conical, the eyes are deeply emarginate anteriorly, and the abdomen has six distinct well developed segments". The morphological differences listed by Wolcott, together with the presence of pectinate antennae on males of Neocallotillus versus serrate antennae on males of Callotillus ( Fig. 3A-C, G-H), and an elytral swelling present in Callotillus but absent in Neocallotillus (Fig. 4 A-B), sup-port the recognition of two genera within the group. The monotypic Barrotillus was also examined in this study in order to assess possible congenericity with Neocallotillus. The structure of the antennae and number of antennomeres serve as evidence to conclude that these closely related genera should be considered as separate taxa ( Fig. 3A Differential diagnosis. N. elegans can be differentiated from similar species based on the integument color, fascia pattern, and wide geographic distribution. The species is most similar to N. intricatus but can be easily differentiated from the latter based on the fasciae pattern on the elytral disc. Neocallotillus elegans has the elytra adorned with a light testaceous to almost whitish median, longitudinal, slightly protruding fascia, and a pair of protruding maculae on the anterior half near the humeral angles ( Fig. 1A-C), these maculae may be absent in some individuals (Fig. 1D). Neocallotillus intricatus has the elytral disc decorated with an intricate design of light testaceous fasciae and a pair maculae arranged in the following manner: each elytron with one macula situated posterior to the humeral angle; one strongly procurved fasciae located on the anterior half of the elytral disc, this fascia initiates on the elytral suture and do not reach the epipleural fold; and a second fascia situated immediately posterior to the first, this band is strongly oblique, initiating on the epipleural fold and not reaching the elytral suture (Fig. 1E). The geographic distribution of these species can also serve to separate them. Neocallotillus elegans is found from the United States to Costa Rica ( Fig. 8A), while N. intricatus is restricted to Costa Rica and Panama (Fig. 8B).
Redescription. Form: Small individuals, feebly to moderately slender ( Fig. 1A-D). Body: elongate, slender. Color: body integument variously colored, from piceous to ferruginous, with tones ranging from fuscous to testaceous; each elytron with one macula and one fascia, both markings ranging from almost albus to testaceous; thefascia is located on the median region of the elytral disc and can range from conspicuously wide to almost imperceptible; the macula is located on the median region of the first third of the elytral disc, initiating on the epipleural fold and almost reaching the elytral suture; these markings can be medially interconnected or not. The maculae may be absent in some specimens (Fig. 1D).
Thorax: Pronotum longer than broad, moderately to strongly punctate, punctations ranging from coarse and deep to moderately shallow and fine; sides subparallel in dorsal view, then abruptly constricted on posterior fourth; disc feebly convex. Prosternum smooth to feebly punctate; punctations coarse, finely to moderately vested with pale, recumbent setae. Mesoventrite smooth, feebly punctate; finely vested with some pale, semi-recumbent to recumbent setae. Metaventrite moderately punctate; strongly convex; surface smooth, vested with fine, recumbent and semi-recumbent setae; longitudinal depression present; metaventral process absent.
Elytra: Humeri indicated, slender, elongate; lateral margins subparallel, slightly to moderately broader on last third, then moderately to strongly depressed on second third, and conspicuously convex again on last third; sculpture consisting on shallow, irregularly arranged punctations; elytral apices subtriangular to almost rounded, feebly dehiscent; interstices at elytral base about 3× the width of punctuation; scutellum subquadrate, not depressed; epipleural fold complete, narrowing toward apex.
Female variation: Females can be distinguished from males based on the antennal structure and the shape of the last abdominal segment. The antennal shape of females is moderately to strongly serrate; antennomeres 2-3 are slender, filiform; antennomeres 4-9 are serrate, the serrations gradually increase in size toward distal end (Fig. 3D, E). The posterior margin of the sixth ventrite of females is strongly procurved, producing a semicircular pygidium (Fig. 5C-D).
Remarks. The species Callotillus occidentalis, described by Gorham (1882) for individuals collected in Guatemala and Nicaragua, was later synonymized with C. elegans by Schenkling (1903). We examined one female paratype of C. occidentallis ( Fig. 2A) and agree with Schenkling's synonomy. Barr (1950) proposed that Callotillus vafer be reclassified as a subspecies of C. elegans on the basis of integument color, geographic distribution discontinuity, and differences in the structure of the elytral punctation. Individuals inhabiting Arizona, California, Nevada, New Mexico, western Texas, Utah, and the Baja California peninsula were classified by Barr as C. e. vafer while specimens from western Louisiana, eastern Texas, Mexico and Central America were recognized as C. e. elegans. Barr also indicated the existence of intermediate forms of these subspecies in the United States and Baja California; however, we have found intermediate forms exist throughout much of the geographic range of the species. It is possible to find both color morphotypes ( Fig. 1A-B), including intermediate forms ( Fig. 1C-D), as well as conspicuously similar antennal forms (Fig. 3A, B), throughout North and Central America. Aside from integument variation, no other morphological evidence was found to differentiate these subspecies as separate taxa. As more material from these subspecies has been accumulated, we consider N. elegans a species with a wide spectrum of color variation throughout an extensive geographic distribution (Fig. 8A).   Distribution. Puerto Rico (Fig. 8C). Differential diagnosis. N. crusoe is similar to N. elegans but differs from the latter species by the absence of seriate elytral punctures, its larger size, its broader form, and the impunctate metaventrite and abdomen. The differently formed and arranged raised fasciae or maculae are also distinguishing characters. The head and pronotum in N. crusoe are densely pubescent, sparsely so in the N. elegans species; the antennae are differently formed, having a greater number of triangular segments; the color pattern is unique; the arrangement of the pubescence in basal half of elytra is distinctive; and the densely pilose elytral tubercles are present only in C. eburneocinctus.
The following is Wolcott's original description. N=1. Form: Moderately slender. Color: Black. Dorsal surface rather feebly shining; ventral surface very shining; front of head narrowly rugulose; antenna (apical two segments black) and labrum at sides testaceous; elytra black with the apical half in large part pale yellow; a large, ovate, ante-apical, black maculation; sides at middle with an oblique, elevated, white maculation; a similar minute, slightly transverse maculation at basal fourth at middle of width of each elytron.
Head: Including the not prominent eyes, equal in width to pronotum at apex; surface rather coarsely rugoso-punctate; pubescence dense, semi-recumbent, grayish white. Antennae slightly longer than head and prothorax, ten-segmented; basal segment short, very stout; second small, subtriangular; third to ninth triangular, their apices acute; ninth and tenth forming an elongate ovate mass; tenth narrower than eighth, nearly as long as seventh and eighth together; color testaceous; ninth and tenth segments black, the former narrowly testaceous at base.
Pronotum: Slightly longer than wide; apical margin truncate; sides parallel to slightly behind the middle, then rather strongly arcuately narrowing to about basal fourth, thence subparallel to base; base truncate, the extreme edge with a fine elevated margin; subapical constriction wanting; subapical transverse impression nearly obsolete, only faintly indicated in certain lights; surface with sculpture same as that of head; pubescence same as that of head but with long, sparse, erect, black hairs intermixed.
Elytra: Base nearly twice as wide as pronotum at base; length 1/3× times width at base; humeri obtusely rounded; sides from humeri to middle straight, nearly parallel, behind the middle gradually broadening to apical fourth, thence arcuately narrowing to the conjointly rounded apices; color black, apical half pale yellow, anterior margin of yellow portion convex; in apical third a large, elongate ovate, common, sutural maculation, extending very nearly to apical margin, black; sides slightly anterior to middle with a feebly arcuate, linear, elevated, white maculation, this extending obliquely and attenuate forward from lateral margin halfway to suture; at basal fourth a minute, slightly transverse, elevated, white maculation midway between the lateral margin and the suture; base with a broad triangular area, having one angle on suture, and an oblique fascia each side, extending from immediately behind the humeri to the suture at a point slightly before the middle, composed of dense, coarse, grayish-white pubescence; a large, feebly elevated, subbasal tubercle, midway between lateral margin and suture, densely clothed with a tuft of long, black hairs; black portions densely clothed with short, semi-recumbent, black pubescence, longer and erect in humeral region; the yellow portion densely clothed with pale yellowish pubescence, a few nearly erect, long, black hairs intermixed; surface finely and sparsely punctate at extreme base, becoming closer at about basal fourth, and a little coarser toward the apex; punctuation irregular throughout, showing no tendency to become seriate.
Remarks. Wolcott (1923) described Callotillus crusoe from a single male specimen collected near Camuy, Puerto Rico. Wolcott concluded that this species was allied to C. elegans and C. vafer but could be differentiated from the latter two based on the absence of elytral punctations, a relatively larger size and broader body shape, and the absences of punctations on the metaventrite and abdomen. Based on Wolcott's illustration and his descriptive work, the shape of the antennae appear to be serrate, and the species seems to be comparatively larger and broader than remaining Neocallotillus species. These characteristics may suggest a relationship to Callotillus. Due to the absence of material of N. crusoe, a redescription of this species is not presented in this study; however, in order to complement the revision of the newly erected genus Neocallotillus, the descriptive work given by Wolcott (1923) is transcribed above. Based on Wolcott's assessment, we tentatively place C. crusoe within Neocallotillus. Further examination of material from this species will serve to corroborate the relatedness of this species with those species composing Neocallotillus, or conversely, its reassignment to Callotillus. (Wolcott & Dybas, 1947), comb. n. Holotype depository. Naturalis Biodiversity Center, Leiden, The Netherlands (RMNH). Holotype locality: Farm La Caja, 8 km. west of San Jose, Costa Rica.

Neocallotillus intricatus
Distribution. Costa Rica, Panama (Fig. 8B). Differential diagnosis. N. intricatus is most similar to N. elegans. The two species can be differentiated based on the fasciae pattern on the elytra disc. Neocallotillus intricatus has the elytral disc decorated with an intricate design of light testaceous fasciae and a pair of maculae arranged in the following manner: Each elytron with one macula situated posterior to the humeral angle; one strongly procurved fasciae located on the anterior half of the elytral disc, this fascia initiates on the elytral suture and do not reach the epipleural fold; and a second fascia situated immediately posterior to the first, this band is strongly oblique, initiating on the epipleural fold and not reaching the elytral suture (Fig.  1E). Neocallotillus elegans has the elytra adorned with a light testaceous to almost whitish median, longitudinal, slightly protruding fascia, and a pair of protruding maculae on the anterior half near the humeral angles ( Fig. 1A-C), these maculae may be absent in some individuals (Fig. 1D). The geographic distribution of these species can also serve to separate them. Neocallotillus intricatus is limited to Costa Rica and Panama (Fig. 8B) while Neocallotillus elegans is found from the United States to Costa Rica (Fig. 8A).
Redescription. Form: Body elongate; head, pronotum and anterior half of elytra slender, feebly expanded behind second half of elytral margins. Color: Head, pronotum, thorax, abdominal segments 1-4 and femora griscent to fuscous; anterior margin of pronotum, antennae, mouthparts, tibiae, abdominal segments 5-6 and elytral apex light-ferruginous. Elytra adorned with an intricate array of two pale-testaceous fasciae and a pair of maculae of the same color, the position of these elytral markings is as follows: the first fascia is located on the anterior half of the elytral disc, this band is strongly procurved, initiating on the elytral suture and not reaching the epipleural fold; the second fascia is located immediately posterior to the first band and is moderately oblique, initiating on the epipleural fold and not reaching the elytral suture; the two macuale are located posterior to the humeral angles. Elytral pattern not elevated from elytral disc (Fig. 1E).
Thorax: Pronotum longer than broad; surface rugulose and strongly, finely punctate; punctations numerous, shallow; sides subparallel in dorsal view, then abruptly constricted on posterior fourth; disc convex. Prosternum feebly convex; surface smooth; conspicuously punctate, punctations shallow. Mesoventrite smooth; surface feebly punctate; finely vested. Metaventrite globate; surface smooth, strongly convex and finely punctate; longitudinal depression and metaventral process absent; metepisternum exposed but profusely covered with short, fine, pale setae observable in lateral view. Scutellum ovoid in shape. Elytra: Humeri indicated; slender on anterior half and then gradually expanding behind middle; surface convex on first third, then strongly depressed on second third, and then conspicuously convex on last third; elytral sinuosity observable in lateral view; sculpture on elytral disc consisting on abundant, very shallow, irregularly arranged punctations almost imperceptible in some individuals; elytral apices rounded, moderately dehiscent; interstices on elytral base about 2× the width of punctuation; epipleural fold complete, narrowing toward apex.
Female variation: The antennal shape of females is strongly serrate (Fig. 3F), rather than pectinate, as observed in males (Fig. 3C). Females have antennomeres 1-3 slender, antennomeres 4-5 are moderately serrate, and antennomeres 6-9 are strongly serrate, serrations gradually increase in size toward the distal end. Females also have the lateral and posterior margins of the sixth ventrite subquadrate (Fig. 5F), producing a somewhat semicircular pygidium. All females in the material examined were moderately larger than males.
Remarks. Wolcott and Dybas (1947) described Callotillus intricatus based on two specimens, one male and one female (Fig. 2B), collected from a single locality 8 km west of San Jose, Costa Rica. This species is here transferred to Neocallotillus based on the pectinate antennae of male individuals (Fig. 3C) and the feebly pectinate to almost serrate antennae in females (Fig. 3F), these antennal shapes are very similar to those observed on N. elegans (Fig. 3A-B, D-E). Other characters that are similar in these species are general body shape (Fig. 1A-E), elytral sculpturing, and the conical shape of the terminal segment of the maxillary palps.
Differential diagnosis. Callotillus is most similar to the new genus Neocallotillus. A list of useful characters to properly differentiate these genera is provided above in the differential diagnosis of Neocallotillus.
Redescription. Size: 6-10 mm. Color: Testaceous and ferrugineous to almost black; fasciae on elytral disc may be present or not, if present, ranging from light testaceous to brown. Body moderately robust, expanded posteriorly.
Thorax: Pronotum moderately to conspicuously globose, narrower than anterior margin of elytra; disc moderately to strongly convex, inconspicuously broader at middle, feebly sinuate behind middle, then feebly to moderately constricted on last fourth; anterior depression and antescutellar impression absent. Prosternum: smooth to feebly puncticulate; conspicuously wider than long. Mesoventrite smooth, feebly to moderately puncticulate; metepisternum partially visible in lateral view, not fully concealed by elytron. Metaventrite moderately to strongly convex; surface variously punctate.
Elytra: Moderately robust, elongate, expanded posteriorly; median region of elytral disc feebly to moderately depressed in lateral view; subbasal elytral swellings present; elytral apex declivity feebly to moderately steep; transverse fasciae may be present on elytral disc or not, if present elevated from elytral disc.
Remarks. In accordance to the taxonomic changes previously discussed, in this revision we treat Callotillus as a genus containing two species: C. bahamensis Vaurie (Figs1G, 2C), a species currently recorded from the Bahamas and the Cayman Islands (Fig. 8D), and C. eburneocinctus Wolcott (Figs 1H,4B), a species restricted to the southernmost tip of the Florida peninsula, including the Florida Keys (Fig. 8E). Vaurie, 1952 Holotype depository. American Museum of Natural History (AMNH). Holotype locality: South Bimini Island, Bahamas, British West Indies.

Callotillus bahamensis
Distribution. The Bahamas, Cayman Islands (Fig. 8D). Differential diagnosis. Callotillus bahamensis is most similar to C. eburneocinctus. The two species can be differentiated with ease based on the color pattern on the elytral disc. Callotillus bahamensis has the elytral disc predominantly piceous, except a light testaceous area surrounding the humeral angles, this testaceous area extends from the anterior fourth of the epipleural fold and may reach the scutellum or not (Fig. 1G). Callotillus eburneocinctus has the anterior half of the elytral disc rufous and the posterior portion fuscous, this coloration shift is interrupted by a transverse, moderately elevated, pale band which runs from the elytral suture to the epipleural fold (Fig. 1H).
Redescription. Form: Moderately robust; elytra gradually expanded toward apex, then abruptly narrowing behind distal fourth. Color: Anterior portion of femora, trochanters, coxae, and anterior fourth of the elytral disc light testaceous, this testaceous pattern on the elytral disc reaches the humeral region laterally and the scutellum internally; remaining body uniformly fuscous; the elytral disc is devoid of any bands or fasciae (Fig. 1G).
Aedeagus: Not available. Female variation: Females of C. bahamensis can be distinguished from male specimens based on the antennal structure and the shape of the last abdominal segment. The females have antennomeres 4-9 moderately serrate (Fig. 3I), rather than strongly serrate, as observed in males (Fig. 3G). Additionally, the last ventrite and the last tergite are subquadrate in shape (Fig. 5H), and not emarginate, as in males (Fig. 5G).
Remarks. In her original description, Vaurie (1952) indicated that C. bahamensis is most closely related to C. crusoe. After examination of specimens of C. bahamensis, this species is most similar to C. eburneocinctus. Based on Wolcott's description (1921), we place C. crusoe within Neocallotillus. Examination of material of C. crusoe will be essential to clarify the status of this rare species. Wolcott, 1911 Holotype depository. United States National Museum of Natural History (USNM). Holotype locality: Key West, Monroe Co., Florida.

Differential diagnosis.
Callotillus eburneocinctus is most similar to C. bahamensis. Characters to distinguish these species appear in the diagnosis section of C. bahamensis.
Redescription. Form: Body moderately elongate, robust; head and pronotum somewhat slender; elytra gradually expanded toward apex, then abruptly narrowing behind distal fourth. Color: head, antennae, mouthparts, pronotum, and anterior half of elytral disc testaceous to rufous; legs brunneous; distal end of mandibles, abdomen and posterior half of elytral disc fuscous; thorax bicolored, metaventrite anteriorly and internally ferrugineous, posteriorly and distally fuscous; each elytron with a transverse, median, pale fascia that runs from the epipleural fold to the elytral suture, this band may be protruded in most individuals (Fig. 1H).
Elytra: Robust; humeri indicated, gradually expanding toward elytral apex; surface convex on first third, then moderately depressed on second third, and conspicuously convex again on last third; a pair of long, stiff, erect tuft of dark setae located on the anterior fourth each elytron; surface of elytral disc rugulose; sculpturing absent; elytral apices rounded, moderately dehiscent; epipleural fold complete, narrowing toward apex.