Six new species of Cymatodera from Mexico and Central America and the retention of Cymatodera obliquefasciata as a valid name (Cleridae, Tillinae)

Abstract Six new Cymatodera speciesfrom the Mexican states of Jalisco and Chiapas, and the Central American countries of El Salvador, Honduras, Costa Rica and Panamá are described: Cymatodera rosalinae sp. n., Cymatodera capax sp. n., Cymatodera sinuosa sp. n., Cymatodera vittata sp. n., Cymatodera rubida sp. n. and Cymatodera limatula sp. n. Justification for retaining Cymatodera obliquefasciata within Cymatodera instead of transferring it to Bogcia is provided. Male genitalia and other characters of taxonomic value are illustrated.

the new species, C. limatula, described below. Therefore, C. obliquefasciata should be retained as Cymatodera based on the ungual structure of the protarsus.
This work is intended to be a small contribution towards a better understanding of the Cymatodera fauna of Mexico and Central America, and to shed some light on the complex relationships found in this group.

Methods
The methods used for genitalia extraction and dissection are similar to those described by Ekis (1977). Terminology used partially follows the work of Ekis (1977) and Rifkind (1996). I considered differences in the aedeagi as the primary evidence for determination of biological species (sensu Mayr 1963).
Specimens were observed using a Leica MZ APO stereomicroscope. All measurements were made using a stereomicroscope ocular micrometer and the software Leica Application Suite V. 3.4.0. Optic images were taken using a Leica DFC 500 digital camera. Scanning electron photographs were taken using a Hitachi 3500N variable pressure scanning electron microscope.
The following abbreviations are used in the description of the holotypes: TL= Total body length, HW= Maximum head width, HL= Head length, PW= Maximum pronotal width, PL= Pronotal length, EW= Maxium elytral width, EL= Elytral length.
Variation. Female specimens differ from males in the following respects: antennal serration somewhat more evident (Fig. 10); elytral margins less parallel, rendering a somewhat more robust appearance; posterior margin of fifth visible ventrite broadly, shallowly, arcuately emarginate (Fig. 31); sixth visible ventrite subtriangular, lateral margins strongly oblique, almost confluent apically, posterior margin rather acuminate (Fig. 31); posterior margin of fifth tergite narrowly, moderately emarginate (Fig. 45); sixth tergite semicircular, lateral and posterior margins broadly rounded (Fig. 45); sixth tergite fully covering sixth visible ventrite. Pronotum and elytral ground color rather inconsistent in both sexes, ranging from testaceous to fuscous-brown. Midelytral fascia variably marked, almost imperceptible in some specimens. Some individuals possess an irregular, midelytral black band ranging in size from a complete fascia, to a pair of inconspicuous maculae on each side of elytra. Differential diagnosis. Distinguishable from congeners based on its size, antennal shape, midelytral marking, terminal abdominal segments and male genitalia. This species appears most similar to the allopatric species Cymatodera obliquefasciata. Color, form, elytral sculpturing, median pale fascia and serrate condition of antennae are characters shared by C. rosalinae (Fig. 1) and C. obliquefasciata (Fig. 7).
This new species can be separated from the latter based on the following respects: C. rosalinae has the antennomeres 2-4 slender, antennomeres 5-10 transversally robust in dorsal view and somewhat serrate (Fig. 9), due to sexual dimorphism, serrate condition of female specimens is somewhat more evident (Fig. 10), last antennomere internally sinuate, and humeral maculae absent. C. obliquefasciata has the antennomeres 1-3 slender, antennomeres 4-10 transversally slender in dorsal view and boldly serrate (Fig. 16), and humeral maculae variably marked to absent. In addition, the moderately incised and arcuately emarginate posterior margin of the sixth visible ventrite in male specimens of C. rosalinae (Fig. 24) is absent in males of C. obliquefasciata (Fig. 30). The female of C. rosalinae (Figs 31,45) can be distinguished from females of C. obliquefasciata (Fig. 37, 50) by the distinctive shape of its abdominal terminalia.
Distribution. Known from the vicinity of the Chamela Biological Station, situated in the Chamela-Cuixmala region, on the western portion of Jalisco, Mexico.
Etymology. I am very pleased to name this new species in honor of my mother, Rosalina Roco, a cornerstone in my life, and a person whose endless efforts have been of inspirational support during my professional career.
Color: head and pronotum dark brown; rest of the body uniformly brown. Each elytron with a post median, irregular, narrow, obliquely directed, black fascia that extends from epipleuron to elytral suture, becoming somewhat paler near suture; this fascia is bordered anteriorly by an inconspicuous pale marking; ventrites 1-5 with a pair of irregular, testaceous maculae near sides.
Legs: somewhat covered with short and long semirecumbent setae that become more numerous on second half of tibiae; femora finely punctate, longitudinally rugose; tibiae coarsely punctate, transversely rugose.
Differential diagnosis. Separable from other Cymatodera species based on its shape, elytral marking, unique terminal abdominal segments and male genitalia (Fig.  53). Due to the serrate condition of the antennae (Fig. 11), general size, form, color and midelytral fascia, Cymatodera capax ( Fig. 2) appears closest to the Mexican Bogcia oaxacae (Fig. 8). The new species can be distinguished from the latter based on ungual differences of the protarsus. C. capax presents the claw of the protarsus conspicuously separated (Fig. 23) from the denticle, rather than closely approximated, as observed in B. oaxacae (Fig. 22).

Distribution.
Known from two localities in the Guanacaste Province, Costa Rica: Playa Naranjo, adjacent to Santa Rosa, Guanacaste National Park; and Sector Las Pailas, Rincón de la Vieja National Park.
Etymology. The specific epithet comes from the Latin word capax (= wide), a noun that makes allusion to the overall robust appearance of this new species.
Color: head and pronotum fuscous; elytra, scutellum and legs brown, except posterior half of femora dark brown; antennae and mouthparts testaceous; abdomen, meso and metasternum light brown. Each elytron with three irregular, variably sinuate, pale fasciae; first on anterior fourth, slender, extending from elytral suture to epipleuron, surrounding scutellum; next fascia on second fourth, broader than preceding band, extending from elytral suture to epipleuron; third fascia on last fourth, slightly shorter and narrower than preceding fascia, extending from elytral suture to tenth stria, not reaching epipleuron.
Legs: vested with short, recumbent setae intermixed with occasional long, suberect and erect setae that become more densely arranged on tibiae; femora rugulose; tibiae transversely rugose.
Elytra: EL= 5.5 mm, EW= 2.5 mm; length to width ratio: males average 2.26, females average 2.19; anterior margin arcuately emarginate, as wide as pronotum; humeri feebly indicated; sides subovoid, widest at second third; disc moderately flattened apically; apex broadly, separately rounded, dehiscent, covering sixth tergite; surface smooth, clothed with short, recumbent setae combined with less numerous, long, erect setae; sculpturing consisting on rather numerous, coarse punctations arranged in striae that abruptly reduce in size and become less numerous on last fourth; interstices about 2.0 × the width of punctation.
Variation. Males have the fifth visible ventrite convex, rugose, moderately, coarsely punctate, lateral margins oblique, posterior margin broadly, deeply, arcuately emarginate (Fig. 26); sixth visible ventrite rectangular, surface convex, rugose, moderately, coarsely punctate, with a longitudinal, median depression that extends from middle to posterior margin, lateral margins parallel, becoming somewhat oblique on last third, posterior margin broadly, deeply, arcuately incised, strongly elevated before emargination, forming a conspicuous ridge bordering the area preceding the posterior margin, then abruptly descending toward emargination, hind angles robust, strongly arcuate at apex (Fig. 26); fifth tergite punctate, rugulose, posterior margin broadly, shallowly, arcuately emarginate; sixth tergite rectangular, longer than broad, rugulose, surface convex, moderately, finely punctate, lateral margins subparallel, posterior margin shallowly emarginate, finely crenulate (Fig. 40). Elytral ground color is slightly variable in both sexes, ranging from brown to fuscous. Leg color ranges from bicolored to uniformly brown. Fasciae color is also rather inconsistent, ranging from stramineous to testaceous. One paratype female displays the posterior margin of fifth tergite broadly, shallowly, arcuately emarginated. Differential diagnosis. This species recalls various Cymatodera members that are similar in color, fasciae pattern, body proportions and antennal gestalt. Among these, Cymatodera sinuosa is most similar to the Central American Cymatodera parallela Gorham, 1882 and the Mexican Cymatodera grossa Gorham, 1882. C. sinuosa can be separated from the former as follows: anterior margin of elytra as wide as pronotum (wider in C. parallela); elytral margins conspicuously wide behind second half (somewhat subparallel in C. parallela); elytral apices ( Fig. 3) broadly rounded and dehiscent (moderately rounded and feebly confluent in C. parallela); third antennomere ( Fig.  12) as long as preceding antennomere (1.5 × the length of preceding antennomere in C. parallela). Furthermore, this new species is distinguishable from C. grossa based on its unique pronotal and elytral sculpturing. C. sinuosa has the pronotum finely, moderately punctate (coarsely, densely punctate, with punctations nearly confluent in C. grossa); pronotal disc conspicuously impressed in front of middle (flat in C. grossa); antescutelar impression strongly indicated (inconspicuously indicated in C. grossa); anterior half of elytral ground moderately punctate (densely punctate in C. grossa); interstices 2.0 × the width of punctation (less than the width of punctation in C. grossa). Likewise, the male of C. sinuosa can be distinguished from males of other species sharing similar fasciae pattern and remaining congeners, by the unique combination of elongated, deeply incised, posteriorly elevated, and abruptly descended emargination on the sixth visible ventrite (Fig. 26).
Distribution. The species is known from two localities: La Muralla National Park, situated in the department of Olancho, Honduras and El Boquerón National Park, in the Department of San Salvador, El Salvador.
Etymology. The specific name comes from the Latin noun sinuosa (=sinuous), and refers to the winding character of the fasciae found on the elytral ground of this species. Description. Size: TL= 7.8 mm, length of males 7.5 to 9.2 mm, length of females 6.5-7.8 mm, n = 9 (Fig. 4).

Cymatodera vittata
Color: head fuscous-brown; pronotum, mouthparts, mesosternum, metasternum and abdomen testaceous; elytral ground predominantly testaceous except median re- gion pale-testaceous; legs mostly testaceous, except posterior half of femora brown. Each elytron with two pairs of black, irregularly marked maculae; the first adjacent to anterior margin, extending from second stria to humeral angle; the second located on first half of second fourth, more faintly marked than preceding pair, extending from second to eighth stria; a long, irregular, transversally marked, black fascia located on third fourth, in the form of a vitta, extending from suture to epipleuron, covering about one fourth of elytral length.
Variation. Females differ from male specimens as follows: antennomeres 5-10 weakly serrate, sixth visible ventrite subtriangular, lateral margins oblique, posterior margin broadly, very shallowly emarginate (Fig. 34); sixth tergite subtriangular, lateral margins oblique, posterior margin feebly notched, hind angles broadly rounded (Fig.  48). Additionally, female specimens lack the setiferous area found on the metasternal region of males (Fig. 58). Elytral ground color is rather variable in both sexes, ranging from flavous to ferrugineous. Maculae on anterior half of elytral ground irregularly impressed, ranging from strongly marked to absent. Posterior fascia color ranges from light brown to piceous. Leg color is also rather inconsistent, ranging from uniformly brown to bicolored. One male with anterolateral carinae of sixth visible ventrite more strongly pronounced. One male with surface of sixth ventrite feebly convex. Two females with posterior margin of sixth tergite truncate. Differential diagnosis. Its distinct elytral ground color, slender form, shape of terminal abdominal segments and male genitalia will readily separate this species from congeners. Cymatodera vittata appears especially similar to C. rubida, a sympatric species described below. Differences in antennal shape (Figs 13,14) serve to separate this new species from its closest congener. C. vittata has the fourth antennomere conspicuously longer than preceding antennomere (antennomeres 3-4 about the same length in C. rubida); antennomeres 5-10 are somewhat longer and extend beyond basal sixth of elytral ground (antennomeres 5-10 are slightly shorter in C. rubida and do not extend beyond basal sixth of elytral ground). Additionally, the distinctive metasternal sensory area found in the male of C. vittata is poorly developed in males of C. rubida (Figs 58,59). Differences in terminalia (Figs 27,28,41,42) and male genitalia (Figs 55, 56) also distinguish this new species from C. rubida.
Distribution. Known from three localities in Panamá: El Llano, Panamá Province; 4 km south of Gamboa, Colón Province; and 4 km south of El Valle, Coclé Province.
Variation. Female specimens have the sixth visible ventrite subquadrate, rugulose, surface strongly convex, lateral margins oblique, posterior margin truncate (Fig. 35); sixth tergite campanulate, rugulose, surface convex, lateral margins oblique, becoming parallel on last third, then abruptly expanding before apex, posterior margin broadly, deeply notched (Fig. 49); posterior margin of sixth tergite extending beyond apical margin of sixth visible ventrite, posterolateral margins of sixth visible ventrite projecting laterally beyond sixth tergite (Figs 35,49). Elytral marking on anterior half ranges from strongly impressed to absent. Posterior fasciae color is also rather variable, ranging from light brown to piceous. Leg color ranges from uniformly brown to bicolored. One male with posterior margin of sixth ventrite slightly less incised than remaining male specimens in the type series.
Differential diagnosis. Cymatodera rubida is separable from congeners based on its size, shape, elytral markings, terminal abdominal segments and male genitalia. Most similar to C. vittata, but differs from this by having the fourth antennomere about the same size as third antennomere (Fig. 14), rather than 1.5 × longer (Fig. 13). Unlike C. vittata, antennomeres 5-10 of C. rubida are rather short; hence, antennae do not extend beyond basal sixth of elytral ground. Furthermore, male specimens of C. rubida do not display the moderately dense vestiture found on the metasternal area of males of C. vittata (Figs 58-59). Conspicuous differences in terminal abdominal segments (Figs 27,28,41,42) and male genitalia (Fig. 55, 56) also distinguish this new species from C. vittata.
Distribution. Known from five localities in Panamá: the Canal Zone, 5 km south of Gamboa; Fuerte Kobbe; the mountainous region of Cerro Azul; the surroundings of Palenque; and El Llano-Carti road, 12 km north of El Llano.
Etymology. The specific epithet comes from the Latin noun rubida (=red), this adjective indicates the general reddish color of this new species. Description. TL= 9.8 mm, length of males 10.2-11 mm, length of female 9.4 mm, n = 4 (Fig. 6).

Cymatodera limatula
Color: head predominantly black (Fig. 60), except gula and submentum ferrugineous; pronotum and mesosternum light ferrugineous; elytra, metasternum, legs, mouthparts and antennae testaceous; abdomen testaceous, except sixth visible ventrite pale testaceous (Fig. 29), and median region of ventrites 2-3 irregularly fuscous (Fig.  62); vestiture uniformly pale; a transversally directed, black fascia on anterior margin of pronotum covering about one fifth of pronotal disc, and two irregular, small, black maculae located on lateral area of pronotum adjacent to posterior margin. Each elytron with two broad, somewhat irregular, transversally directed, black fasciae extending from suture to epipleuron; first located on humeral region, covering about one sixth of elytral ground; second behind middle, somewhat longer than preceding band, covering approximately one fifth of elytral ground.
Variation. The only female in the type series differs from the male by having the first visible ventrite moderately longer and the posterior margin of ventrites 1-2 truncate (Fig. 61). Moreover, abdominal differences are as follows: fifth visible ventrite rugose, lateral margins oblique, posterior margin truncate (Fig. 36); sixth visible ventrite rugulose, semicircular, surface feebly convex, broader than long, lateral and posterior margins broadly rounded (Fig. 36); fifth tergite rugulose, lateral margins oblique, posterior margin truncate; sixth tergite rugulose, broader than long, surface inconspicuously convex, lateral and posterior margins strongly oblique, slightly acuminate posteriorly, almost semicircular. Posterior margin of sixth tergite extending slightly beyond sixth visible ventrite. One male paratype does not possess the irregular, small, black maculae located on the posterolateral area of pronotum.
Differential diagnosis. The distinctive coloration and elytral markings of this species will serve to separate it from congeners. No other species in the Cymatodera group has the distinctive piceus fasciae on the pronotum and elytral ground, and the predominantly black color on the head (Figs 6, 60). In addition, shape of terminal abdominal segments (Figs 29, 36), strongly serrate condition of antennomeres 4-10, shape of last antennomere (Fig. 17), and male genitalia (Fig. 57) are characters that serve to separate C. limatula from remaining Cymatodera species.
Distribution. This species is known from three localities in the state of Chiapas, Mexico: El Aguacero, in the periphery of Tuxtla Gutierrez; El Chorreadero, 8 km northeast of Chiapa de Corzo; and on highway 190, 17 km west of Tuxtla Gutierrez.
Etymology. The specific epithet comes from the Latin noun limatula (=distinctive), and makes allusion to the characteristic color pattern of this new species.