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Four new species of Cymatodera Gray from central and southern Mexico (Coleoptera, Cleridae, Tillinae)

Abstract Four new species of Cymatodera are described from Mexico: Cymatodera tortuosa Burke & Rifkind, sp. n. from Hidalgo and Tamaulipas; Cymatodera ortegae Burke, sp. n. from Colima, Jalisco and Michoacan; Cymatodera gerstmeieri Burke & Rifkind, sp. n. from Chiapas; and Cymatodera mixteca Burke & Rifkind, sp. n. from Puebla and Guerrero. Male genitalia and other characters of taxonomic value are illustrated.


Introduction
As part of an ongoing effort to catalog the diversity of Mexican Cleridae, the present work describes four new species of Cymatodera Gray from the central and southern states of Mexico. As previously discussed (Rifkind 2014(Rifkind , 2015, the diversity of the clerid genus Cymatodera in Mexico is extensive, but our knowledge of the group remains rudimentary. Many dozens of species await description and many more are likely to be discovered, particularly in areas distant from paved roads. Recent descriptions of species belonging to this genus include many endemics (Burke 2013;Burke and Zolnerowich 2014;Rifkind et al. 2010;Rifkind 2014Rifkind , 2015, and here again, it is quite likely that the tally will increase as collecting efforts reach further into habitats such as relictual cloud forest and isolated mountain ranges that are known centers of endemism. Much attention has been focused on the rapid destruction or degradation of natural habitat in Mexico and other parts of Latin America, and the insufficiency of current resources dedicated to cataloguing disappearing biodiversity (Blackman et al. 2014;Armesto et al. 2007;Santibañez and Santibañez 2007;Trejo and Dirzo 2000). One positive trend is a recent increase in the number of trained taxonomists from these countries. Several workers in Mexico, for example, have undertaken faunistic and systematic studies of Cleridae with the result that the pace of description there is now higher than it has been in nearly a century (Burke 2013;Burke and Zolnerowich 2014;Rifkind et al. 2010, Toledo-Hernández et al. 2015. Should this trend continue, there may be more hope than previously thought for the assessment and conservation of critically threatened habitats in Mexico.

Material and methods
Genitalia extraction and dissection procedures are similar to those outlined by Ekis (1977). Most of the morphological terminology used follows the works of Ekis (1977), Rifkind (1996) and Opitz (2010). Morphology of the male genitalia and pygidia are considered of primary importance as characters for the determination of new species in this descriptive work. Specimens were observed using 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.
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 = Maximum elytral width, EL = Elytral length.
Type material is deposited in the following collections: The new species can be separated from congeners by its unique combination of body form, antennal shape, size, color, and elytral fascia pattern. Cymatodera tortuosa superficially resembles a number of species that share a similar pattern of fuscous and testaceous elytral banding, such as C. balteata LeConte, C. sirpata Horn, C. undulata (Say), and C. wolcotti Barr. Cymatodera tortuosa, however, can be readily differentiated from those species based on clear differences in the male and female pygidium as well as discontinuity in geographic distribution. Specifically, the new species has the male pygidium distinctly modified (Fig. 11) compared to similar species; the female pygidium is moderately, V-shaped emarginate (Fig. 16), rather than broadly rounded posteriorly, as observed in similar species. As no specimens are known outside of central Mexico, it is possible that this new species is restricted to that region, whereas those listed previously are distributed for the most part in the south and southwestern United States, with only C. balteata ranging into the Mexican border states of Nuevo Leon and Tamaulipas.
Legs: Femora clothed with short, recumbent setae interspersed with a few erect and semierect setae; tibiae moderately vested with some short and long erect and semierect setae; femora and tibiae transversely rugose.
Elytra: EL = 8.8 mm, EW = 2.6 mm. Anterior margin arcuately emarginate; wider than widest portion of pronotum; humeri pronounced; sides subparallel; widest on posterior fourth; disc convex; apex moderately dehiscent; elytral sculpturing as follows: anterior third set with regular, rather coarse and deep striae that abruptly diminish after anterior third and disappear entirely on posterior half; punctations at elytral base coarse and deep; surface moderately clothed with short, very fine, pale, recumbent setae intermixed with very few fine, long, erect setae.
Female: The female paratype is somewhat longer than the male, having a total body length of 13.8 mm. The female also differs from the male holotype by having the sixth ventrite strongly, deeply, V-shaped emarginate, lacking the pair of elongate, posterolateral extensions observed in the male (Figs 11,20). Additionally, the female has a slightly paler integument than the male, with a less distinct elytral pattern anteriorly by comparison.

Distribution.
Presently known only from the northern portion of Hidalgo and the southeastern part of Tamaulipas, Mexico. The vegetation at the collecting localities is predominantly pine-oak forest (Fig. 29).
Etymology. The specific epithet tortuosa (from the Latin tortilis or tortuosus, meaning twisted or winding), is a reference to the intricate and elaborate structure of the male pygidium of this species. Cymatodera ortegae Burke, sp. n. http://zoobank.org/27646149-9E6B-4397-ACD2- DB7E55F37A7C Figs 2,7,12,17,21,22,24 Type material (n = 30). Holotype red labeled, male: Jalisco, road to microondas Los Mazos, Sierra Manantlán, 1425-1610 m, 19°42'N, 104°24'W, 12 km SSD Autlan, mixed hardwood forest 15-VII-1993, R. L. Westcott; holotype deposited in CASC. Paratypes yellow labeled: 1 male: same data as holotype (WFBM); 1 female: Mexico: Differential diagnosis. The undulate fascia pattern on the elytral ground, the testaceous to slightly greenish integumental color, general body shape, and geographic distribution of the new species will, in combination, serve to separate it from other species of Cymatodera. Cymatodera ortegae appears to be allied to several Mexican congeners that share similar body shape, integumental color, brachypterous condition, and a reduced anterior elytral margin. Of these, C. barri Rifkind, C. maculifera Barr, and C. monticola Rifkind are most similar. Unlike C. ortegae, however, the males of C. barri and C. maculifera possess a distinct pair of feebly to moderately developed tubercles on the median posterior portion of the metasternum. Both sexes of these species lack the irregular, infuscate elytral pattern of the new species. Cymatodera monticola possesses distinctly different terminalia from C. ortegae, as well as sinuate elytral apices.
Description. Holotype male. Medium sized, moderately slender anteriorly, rather robust posteriorly; brachypterous, TL = 13.1 mm. Color: Head, pronotum, thorax, femora and anterior portion of tibiae brunneous; posterior portion of femora and tarsomeres testaceous; antennae and mouthparts fuscous; abdomen fuscous, slightly darker than thorax, distal portion of abdominal segments with a depressed testaceous mark; elytral ground light testaceous, with a slight greenish tinge. Each elytron bearing a pair of irregular, sinuate, darkened fasciae: the first fascia located on anterior third, extending from the elytral suture to the epipleural fold, conspicuously slender proximal to elytral suture then abruptly widening before epipleural fold; the second fascia located at the elytral mid-length, moderately wide, extending from the elytral suture to before the epipleural fold. Punctation on elytral ground infuscate (Fig. 2).
Legs: Femora clothed with short, recumbent setae intermingled with less numerous erect and semierect setae; tibiae vested with short and long erect and semierect setae; femora and tibiae transversely rugose; metathoracic legs with tarsomeres longer than those of pro-and mesothoracic legs.
Females of the type series differ from males by having the sixth ventrite ( Fig. 17) with lateral margins moderately oblique, feebly arcuate, and posterior margin very feebly, shallowly, narrowly emarginate, rather than broadly, deeply, semicircularly emarginate, as observed in males (Fig. 12); additionally, females have the sixth tergite (Fig.  22) subquadrate, with the lateral margins oblique and the posterior margin very feebly, shallowly, narrowly emarginate. Females closely resemble males in other respects.
Variation. Length of males range from 7.9-13.1 mm, females from 7.6-13.3 mm. Specimens examined have considerable variation in body size, and also in the shape of the fasciate pattern on the elytral ground, which ranges from almost incomplete and very narrow, to conspicuously wide, covering most part of the elytral ground. The color of these fasciae is also somewhat variable, ranging from dark testaceous to dark greenish. The elytral ground color ranges from light testaceous to light greenish.
Distribution. Available specimens were collected in the central-occidental part of Mexico, in the states of Colima, Jalisco and Michoacan. Distribution appears limited to the southern tip of the Sierra Madre Occidental (Fig. 29).
Note. This new species appears to be confined to mid-to-high altitude mountainous environments in the central-west portion of Mexico: specimens were collected on the slopes of Volcan Nevado de Colima in the states of Colima and Jalisco, and Cerro Tancitaro, in the northwestern region of the Mexican state of Michoacan. These mid and high altitude areas are dominated by pine and pine-oak forest stands.
Etymology. The specific epithet is a patronymic honoring Cristina Ortega, a friend of the first author. Differential diagnosis. Cymatodera gerstmeieri is similar to a number of New World tilline species that share a testaceous to ferrugineous integument and a median, dark fascia on the elytral ground; those closest include Cymatodera mitae Burke, Bogcia disjuncta Barr, and Cymatodera insignis Schenkling. The new species can be separated from the former as follows: male specimens of C. gerstmeieri have the eleventh antennomere medially depressed, acuminate posteriorly, and approximately 2× longer than tenth antennomere (Fig. 8), while males of C. mitae have the eleventh antennomere cylindrical in shape, rounded posteriorly, and approximately 2.5-3× the length of tenth antennomere. Furthermore, males of C. gerstmeieri have a feebly developed longitudinal carina on the first ventrite, but lack a carina on the second ventrite (Fig. 27), whereas males of C. mitae have a well-developed longitudinal carina on the first ventrite, but also a somewhat less developed carina on the second (Fig. 28). The females of both species lack these carinae, but can be distinguished by the presence of a fuscous macula located on each elytral humeral angle in C. mitae, absent in C. gerstmeieri. The feebly to moderately serrate antennomeres 4-10 of C. gerstmeieri (Fig. 8) will easily separate it from Bogcia disjuncta, which has strongly serrate antennae. Cymatodera gerstmeieri somewhat recalls the Central American species C. insignis, with which it shares similar integumental color, antennal structure, and a median, slightly oblique, dark fascia. However, C. insignis bears a dark macula on the humeral angles, a longitudinal black macula at the posterolateral margin of the pronotum on either side, and has the posterolateral margins of the elytral ground narrowly darkened. These markings are absent in C. gerstmeieri.

Cymatodera gerstmeieri
Description. Holotype male. Moderately small, rather robust, metathoracic wings complete, TL = 10.2 mm. Color: head, pronotum, prosternum, mesosternum ferrugineous-brown, the rest of the body uniformly testaceous brown. Each elytron with a fuscous, irregular fascia near the mid-length, extending from the elytral suture to the eighth row of striae, but not reaching the epipleural fold (Fig. 3).
Females can be distinguished from males based on the structure of the pygidium. The sixth ventrite and the sixth tergite are broadly rounded posteriorly in females (Fig.  18). Other characters are constant in both sexes.
Variation. Length of males ranges from 8.7-10.2 mm; length of females from 9.5-13.1 mm. The elytral fascia is slightly variable in width, extending from the elytral suture to the epipleural fold in one male and one female, but incomplete in remaining individuals. Two males and one female in the type series have slightly paler elytra than the male holotype.
Distribution. All specimens in the type series were collected in the locality of El Aguacero, approximately 10 miles northwest of Ocozocuautla, Chiapas, Mexico. The elevation at the type locality is approximately 650 m and the vegetation is predominantly tropical deciduous forest (Fig. 29).
Females of the type series can be differentiated from males by the shape of the sixth ventrite. This segment is broadly rounded posteriorly (Fig. 19), rather than shallowly emarginate, as in males (Fig. 14). Remaining characters are constant in both sexes.
Variation. Length of males ranges from 6.3-8.4 mm; length of females from 7.1-7.9 mm. Individuals in the type series vary somewhat in integument color, ranging from pale testaceous to brown. Such color variation is observable in male and female members in the type series. Remaining characters in the type series remain consistent.
Distribution. The type series was collected from various localities in the Sierra Mixteca of Mexico, specifically in the south and southwestern portion of the state of Puebla, and in central Guerrero state (Fig. 29). This region is characterized by tropical deciduous to thorny forest habitats.
Etymology. The specific epithet makes allusion to the regional home of the Mixteca people, and of this new species.
National Museum of Natural History, and the Texas A&M Insect Collection. This article is Contribution No. 15-389-J from the Kansas Agricultural Experiment Station (KAES) and was supported in part by KAES Hatch Project No. 353,Insect Systematics. Publication of this article was funded in part by the Kansas State University Open Access Publishing Fund.