A taxonomic revision of the Cymindis (Pinacodera) limbata species group (Coleoptera, Carabidae, Lebiini), including description of a new species from Florida, U.S.A.

Abstract The Cymindis (Pinacodera) limbata species group (Coleoptera, Carabidae, Lebiini) is a precinctive New World taxon with ranges extended from portions of temperate southeastern Canada and the U.S.A. through the montane regions of Mexico, south to the Isthmus of Tehuantepec. The group is distinguishable from all other members of the subgenus Pinacodera by males possessing a distinctive sclerite (endophallic plate) at the apex of the endophallus. In the past, a lack of material and misunderstandings of range of variation within species have contributed to confusion about how many species there really are. This revision of the limbata species group includes a classification, a key to groups within the subgenus Pinacodera and species within the limbata group, descriptions of species, re-rankings and new synonymies. In total 10 taxa are treated, with 6 new synonyms proposed, 1 new combination introduced and 1 new species described: Cymindis (Pinacodera) rufostigma (type locality: Archbold Biological Station, Highlands County, Florida, U.S.A.). Each taxon is characterized in terms of structural features of adults, habitat, geographical distribution, and chorological affinities. Available ecological information and treatments of variation are included.


Introduction
Genera of the subtribe Cymindidina (Lebiini) ( Table 1) include the southern Palaearctic-Oriental Trichis Klug, the southern Palaearctic-Afrotropical Hystrichopus Boheman, and Cymindis (s.l.) Latreille (Fig. 1). Cymindis is by far the most speciose and widespread, with a range that extends to all zoogeographic regions with the exception of Australia (Ball and Hilchie 1983). Species of Cymindis are arranged in four subgenera ( Fig. 2): Cymindis (s. str.) Latreille, Afrotarus Jeannel, Taridius Chaudoir, and Pinacodera Schaum. Both Afrotarus and Taridius are restricted to the Old World, Cymindis (s.str.) has a primarily Holarctic distribution with a few additional members in portions of the Oriental and northern Neotropical regions, and Pinacodera is a Western Hemisphere-restricted taxon with members ranging from southeastern Canada to Costa Rica in Middle America (Fig. 2).
Pinacodera includes more than 25 taxa arranged in two species groups, the limbata and latiuscula groups (Ball 2008, personal communication) that are not easily distinguished from one another. The less diverse limbata species group with 10 taxa, treated here, has members that are more northern geographically and males that are distinguishable from those of the latiuscula group by a sclerite (endophallic plate) located at the apex of the endophallus.
The taxonomic position of Pinacodera remains contested to some degree. Initially, Pinacodera was introduced and ranked as a genus by Schaum (1857). Schaum divided the genus Cymindis Latreille, 1806, after recognizing that specimens of species including C. limbata Dejean (the type species), C. fuscata Dejean, C. platicollis Say and C. complanata Dejean differed in having no setae (other than one pair of typical fixed setae located at the apex of each tarsomere) on the dorsal surface of the tarsi. As well, males of these species differed in having expanded meso-tarsi. The name Pinacodera (Greek, "flat+neck.") (Blatchley 1910), also indicates that Schaum recognized species of this group differed in having a pronotum that is flatter than those of Cymindis species. More than a century later, Pinacodera was ranked as a subgenus (Ball and Hilchie 1983) in the genus Cymindis (s. lat.) along with Afrotarus, Cymindis (s. str.), and Taridius. I accept here the ranking by Ball and Hilchie. At the time Schaum recognized Pinacodera, four other species of the limbata group (Table 1) had been described, including C. platicollis, C. complanata, C. chevrolati Dejean, and C. punctigera LeConte. In 1878, H. W. Bates described two additional species, C. amblygona and C. angulifera, but later realized (Bates 1883) that characters used to distinguish the species were variable and accordingly ranked them as varieties of "C. atrata" (a junior synonym of C. chevrolati). Two more varieties of "C. atrata", C. ruficornis, and C. laevior were also described by Bates (1891). During the same time period, Horn (1881) described two species of Pinacodera, and over a 7-year period in the early twentieth century, Casey (1913Casey ( , 1920 described 7 more. I became interested in this group after learning that due to both a lack of specimens available for description and misconceptions of variability in body size, proportions, and coloration, (Casey 1913(Casey , 1920 it was probable that several Pinacodera taxa were over ranked or incorrectly recognized as valid (Ball 2008, personal communication). A large amount of study material would provide the means to better partition variation within the group. Fortunately, a series of studies of Pinacodera are currently being undertaken at the University of Alberta and material had been borrowed from institutional collections from across the United States and Canada. Because of this, thousands of specimens have been available and a thorough examination of the taxa has been possible, using morphological features.
A cursory examination of the literature also revealed a distinct shortage of natural history information for the group despite the abundance of several of the species Table 1. Classification of supraspecific taxa of the subtribe Cymindidina, tribe Lebiini (Ball and Hilchie 1983 (Mahar 1978, Liebherr andMahar 1979). Because of the large numbers of individuals in certain areas, it can be assumed that Pinacodera taxa impact the ecosystems they inhabit (Mahar, 1978). This is another good reason why closer examination was warranted and fieldwork was desirable.
The following revision includes taxonomic treatments (descriptions and illustrations of structural features, notes about habitat, geographical variation and distribution, and postulated evolutionary and chorological affinities), and keys for identification of adults. In total 10 taxa are treated, with 6 new synonymies proposed, 1 new combination introduced and 1 new species described. Variation in wing length, body length, mental tooth form, and male genital characters receive detailed treatment.

Material
This revision is based on the study of more than 4000 adult specimens representing ten taxa belonging to the subgenus Pinacodera. Specimens of a few taxa were collected over the past two years or were already housed at the Strickland Museum, University of Alberta (UASM). Additional adult specimens were borrowed from the collections of various individuals and institutions listed below, along with a four-letter coden (Arnett et al. 1993) used to identify sources of specimens. Names in parentheses below, indicate curator or owner of collection.
Our first attempt (late June, 2007) at collecting specimens of species in the limbata group (Table 1) took George E. Ball, Norman Omoth, and myself, to several U.S. State Parks (Table 2) within the known range and habitat of several species in the limbata group. We were not able to collect C. complanata during this trip but we did collect other species in the limbata group including, C. limbata, C. platicollis platicollis, and C. platicollis atripennis. At localities in Pennsylvania, Virginia, and North Carolina (Ta-  ble 2), specimens of C. limbata and C. platicollis platicollis were found at night, in mixed forest, primarily on the trunks of aspen and oak trees more than 60 cm in diameter. At the time, the localities visited in Florida were very dry and specimens of C. platicollis atripennis were not active. We were fortunate to talk with P. E. Skelley (FSCA) who told us he had found C. platicollis atripennis while surveying squirrel nest insect fauna (Skelley 2007, personal communication). Almost all (16 of 18) individuals of C. platicollis atripennis collected during the trip were taken from squirrel nests.
A few months later Drew Hildebrandt (DAHC) caught two specimens at an ultraviolet (U.V.) light trap while on a collecting trip to George L. Smith State Park, Georgia, U.S.A. Based on this, in mid-February of 2008, Omoth and I collected at George L. Smith using sugar-bait on tree trunks (recipe in collecting methods). Less than an hour after our search began we encountered our first C. complanata specimen. Over the next several days we collected a total of 13 specimens of C. complanata in mixed forest of oak and pine, effectively increasing the known number of pinned adults by one third. All C. complanata were collected from slash pine (Pinus elliottii Engelm.). Previous to this the only known habitat data for this species was from one specimen labelled "under loose bark of live loblolly pine", (Pinus taeda L.) a tree species that is a close relative of slash pine (Petrides & Wehr 1998). With the ample number of hiding spots provided by the tree's flaky bark, along with the ferruginous surface of both the tree bark and the dorsal surface of C. complanata (Fig. 4), it seems possible that this species has come to rely on trees of these species for camouflage and perhaps uses them for other purposes as well.

Collecting methods
Collecting methods for this revision included: u.v. light trapping, beating vegetation, hand collecting from trees, and sugaring tree trunks. The most effective of these methods by far was the use of sugaring. Lindroth (1969) indicated that C. limbata had been recorded in numbers on sugar baits used for catching moths. I was able to confirm this after speaking with lepidopterist J. E. Rawlins (CMNH), who had observed similar activity while collecting with these baits. After some experimentation we found the following recipe to be effective. In a pot, mix two liters of red table wine, one can of beer, three packets of Fleischmanns instant-rise yeast, and four to six pounds of brown sugar. Stir well and bottle. Let stand with the cap off for several hours. Before use shake well to mix settled sugar and apply to trees at shoulder height with a wide paintbrush.

Preparation and examination of adults
Standard methods were used for mounting, dissecting, preparing genitalia, and other technical methods Hilchie 1983, Frania andBall 2007). Genitalia and other small structures were preserved in glycerine and stored in microvials that were pinned beneath the specimen from which they had been removed. Larger structures, including hind wings, were glued to cards and pinned beneath the specimens from which they had been removed.

Measurements
Measurements were made at 12×, 25×, and 50× with a Wild M5 stereoscopic microscope fitted with an ocular micrometer. Various measurements are expressed in the text by abbreviations previously used by Ball and Shpeley (2005):

HL
Length of head, measured on left side, from base of left mandible to posterior margin of compound eye. HW Width of head, maximum transverse distance across head, including eyes.

PL
Length of pronotum along midline. PWM Maximum width of pronotum.

EL
Length of elytra from basal ridge to apex.

EW
Maximum width of elytra. OBL Overall body length.
The shape of the head, pronotum, and metepisternum is shown by the ratio of the width over length (HW/HL; PWM/PL, ML/MW), and elytral shape is indicated by the ratio of the length to the width (EL/EW).
To indicate range in body size of each species, the overall body length (OBL) was measured (to the nearest 0.5 mm) from the apex of the extended mandibles to the apex of the elytra of both the largest and smallest individual of the species (Frania and Ball 2007).
Size of male genitalia was measured by drawing a straight line between the apical area and the basal lobe of the phallus.

Notes about synonymy
I rely on Lindroth (1955) and Lindroth and Freitag (1969) for information about type material for Say and Dejean names. Information about material of other authors was taken from notes by G. E. Ball (2008 personal communication).

Terms
Terms used for structural characters follow Ball and Hilchie (1983) and other authors (See also Figure 3 and Table 3). For some characters of the endophallus of males, no nomenclature has been developed, so in these instances I have used informal descriptive words or phrases.

Label data
All material has been databased and incorporated into the University of Alberta's Strickland Virtual Entomology Museum Database. This database includes UASM reference numbers, full locality data, date of collection, collectors, and codens.
Key to supraspecific taxa of subgenus Pinacodera, and to species of the C. (P.) limbata group, based on characters of adults. Dorsal surface of head between eyes with one shallow transverse line or smooth ( Fig 25A). Geographic range north and east of the Baja California Peninsula and south into mainland Mexico (Fig. 29) .Cymindis punctigera punctigera LeConte 11(9') Humeral macula of elytron with a testaceous patch extended from interval 6 (rarely 5) to lateral margin and extended posteriorly as far as one quarter (0.25) the length of the elytra (Fig. 8)  Head and pronotum rufo-testaceous and elytra bicolored, piceous with rufotestaceous macula medially (Fig. 18)  Color. Piceous to testaceous.
Microsculpture. Most specimens with microlines not visible on dorsum of head capsule; few with mesh pattern isodiametric to transverse between eyes. Elytra with mesh pattern isodiametric, microlines absent from apical half to moderately deep throughout.
Macrosculpture and pilosity. Head ventrally with evenly scattered setigerous punctures bearing setae or not. Pronotum with shallow to moderately deep scattered setigerous punctures, bearing pilose setae or not. Elytra with scattered setigerous punctures, pilose or not. Elytral epipleuron glabrous to moderately setose.
Fixed setae. Two pairs of supraorbital setae; clypeus with two lateral setae. Labrum with six setae along apical margin. Pronotum with two to four setae along each margin. Elytra with two setae in stria 3 and one posterior to end of stria 3; one seta at apex of interval 2; 14-18 lateral (umbilical) setae in the ninth interval; two setae on each of abdominal sterna III to VI (Fig. 3); four to six setae along apical margin of sternum VII (Fig. 3).
Luster. Head capsule and pronotum glossy, elytra glossy to moderately dull; ventral thoracic sterna and abdominal sterna glossy.
Female genitalia. Gonocoxite 2 (gc2) short and stout to long and narrow.
Habitat. Adults of this species group have been collected in temperate deciduous, coniferous and mixed forests, subtropical broadleaf forest, tropical montane forests, and acacia scrub environments. Adults are recorded from elevations ranging from sea level to ~3400 m. Geographical distribution. The range of this species group (Fig. 7,17,21,29,41 extends in eastern Canada from southern Quebec west to southern Ontario; in the United States from the Atlantic coast south to southern Florida and west to California. In Mexico, it is known throughout the montane regions, north of the Isthmus of Tehuantepec. Chorological affinities. Species of the limbata group are sympatric with members of the subgenus Cymindis (s.str) (Fig. 2) in portions of southeastern Canada, the U.S.A, and Mexico, and with the latiuscula group in portions of the southwestern United States and Mexico.
Taxonomic composition, and sequence of presentation. This group includes ten taxa in three species complexes (treated below). The sequence of presentation is based on geographic distribution of these three complexes, beginning in Eastern North America with the limbata complex; second, the monospecific primarily southwestern punctigera complex; and third, the northern Mexican chevrolati complex. Within each complex, the included species (or subspecies) treatments are in a sequence reflecting my preliminary thoughts about relationships, beginning with the most primitive and ending with the structurally most derived.

Figs 4-21
Diagnosis. Species in the limbata complex are distinguished by pale pronotal and elytral margins; several rows (two to four) of setigerous punctures extended the length of elytral interval 8. Most specimens of species in the limbata complex have contrasting punctation in alternate elytral intervals; intervals 1, 3 and 5 typically with one to two rows, interval 2, 4 and 6 typically with two to three rows. All species are macropterous.
Description. With character states of limbata species group, restricted as follows.
Macrosculpture and pilosity. Head capsule with shallow, evenly scattered setigerous punctures on dorsal surface from constriction of neck extended anteriorly toward clypeus. Elytra with striae shallowly impressed and punctulate throughout length; intervals almost flat to slightly convex (few with greater convexity in intervals 1, 3 and 5), one-two (mostly one) irregular rows of fine punctules extended the length of intervals 1, 3 and 5; two-three or three (mostly two) irregular rows of fine punctules extended the length of intervals 2, 4 and 6; interval 8 with two to four rows of fine punctules extended the length of the interval. Abdominal sterna with fine pilose punctures throughout.
Fixed setae. Elytra with two setae in stria 3 and one posterior to end of stria 3; one seta at apex of interval 2; 15-17 lateral umbilical setae.
Pronotum. Anterior and posterior transverse impression shallow; median longitudinal impression shallow; posteriolateral angles almost right-angled to rounded; posterior margin slightly lobate.
Geographical distribution. The range of the limbata complex extends in eastern Canada from southern Quebec west to southern Ontario; in the eastern United States from Maine south to southern Florida, west to eastern Colorado and Nebraska, and south to southern Texas. In Mexico it is known from Nuevo Leon, Mexico, in the northern portion of the Sierra Madre Oriental.
Chorological affinities. The geographical range of the limbata complex overlaps that of the punctigera complex and the chevrolati complex along the extreme southern and southwestern portion of its range.
Taxonomic composition. Five taxa are included in this complex: C. complanata Dejean; C. limbata Dejean; C. platicollis platicollis (Say); C. platicollis atripennis (Casey); and C. rufostigma sp. n.  Notes about synonymy. The above synonymy was established by Lindroth (1969Lindroth ( ), though previously (1955 he treated the names P. complanata Dejean and P. russata Newman as junior synonyms of P. limbata Dejean. Diagnosis. Adults of Cymindis complanata are distinguished from those of other species by ferruginous coloration throughout ( Fig. 4) (some slightly lighter in basal third of elytra), by the almost flat elytral intervals, setae extended almost to the constriction of the elytral epipleuron; a noticeable contrast in punctation extending the length of the intervals; scattered arrangements of one to two rows on intervals 3, 5 and 7 and scattered arrangements of two to four rows in intervals 2, 4, 6 and 8 (most specimens have one row of punctures on intervals 3, 5 and 7 and 3 three rows on 2, 4, 6 and 8). In males, the endophallic plate (ep) differs as it is almost flat, and apical endophallic lobe (ael) is enlarged (Fig. 6A). In females, the form of gonocoxite 2 (gc2) differs from all other species apices of ensiform setae extend to and often past gonocoxite 2 apex (Fig. 6B).
Color (Fig. 4). Dorsum of head entirely ferruginous to rufo-piceous; pronotum ferruginous with lateral margins ranging from translucent to lighter and creamy in appearance; elytra ranging from ferruginous to rufo-piceous, few specimens with basal third of elytra lighter in color, lateral margins translucent to somewhat translucent; antennae and other appendages ferruginous to brown.
Macrosculpture and pilosity. Dorsal punctures with short setae present, few to many visible on head, pronotum (Fig. 5), and at base of elytra (occasionally few setae extended toward apex of elytra). Elytral epipleuron with setae visible, extended to constriction. Elytral intervals with one to two rows of scattered punctures in intervals 1, 3 and 5 and two to three (rarely four) in intervals 2, 4, 6 and 8.
Fixed setae. Pronotum with two setae along each margin; 15-16 lateral (umbilical) setae. Two setae on each of abdominal sterna III to VI, four setae along apical margin of sternum VII (Fig 3).
Head. Fine scattered setigerous punctures from hind portion of eye to constriction of neck; also additional fine punctulae from base of neck extended laterally toward clypeus, otherwise glabrous.
Elytra. Elytral apices truncate, striae shallowly impressed and punctulate throughout length; intervals flat; a noticeable contrast of punctures extended interval lengths; scattered arrangements of one to two rows on intervals 3, 5 and 7 and scattered arrange-ments of two to four rows in intervals 2, 4, 6 and 8. Most specimens with a single row of punctures on 3, 5 and 7 and three rows on 2, 4, 6 and 8; epipleuron with setigerous punctures from base to constriction, in a few specimens beyond to apex.
Female genitalia. Gonocoxite 2 (gc2) (Fig. 6B) distinctly, obliquely truncate. Habitat, habits and seasonal occurrence. The known elevational range of C. complanata extends from sea level to 135 m on the eastern slopes of the Appalachian Mountains. All specimens that I have collected (13 in total) were taken from slash pine (Pinus elliotii Engelm.) though it has also been recorded from under bark of live loblolly pine (P. taeda L.). These species of pine are very similar to each other, sharing both reddishbrown bark coloration and an abundance of flaky bark to rest under. As well, they are common within the known range of C. complanata. Adults are crepuscular or nocturnal with most activity being observed on tree trunks. Most specimens have been collected in March and April. Methods of collecting include u.v. light, sugaring baits painted on tree trunks, and hand collecting at night.
Geographical distribution. The range of this species (Fig. 7) extends in the eastern United States east of the Appalachian Mountains from Maryland south to southern Florida, and westward on the Gulf Coast to western Alabama.
Chorological affinities. Cymindis complanata is sympatric in its entire range with C. limbata (Fig. 7), sympatric in the northern portion of its range with C. platicollis platicollis and in the southern portion of its range with C. platicollis atripennis.
Material  Lindroth (1969Lindroth ( : 1067 stated that the types of C. laevigata and C. fuscicollis are probably in the Zoological Museum of Moscow University (these types were listed at ZMMU by S. I. Keleinikova 1976). He had not seen them, and questions the synonymy of these names and with C. limbata Dejean. I note also that in 1829, some two years before the description of C. limbata Dejean, T. W. Harris had correspondence with fellow entomologist N. M. Hentz in which he referred to a Cymindis specimen from his collection that had an "ochreous elytral margin, with a humeral lunule of the same color" (Harris 1869). He refers to the specimen as C. comma. In 1835, the name C. comma appeared again in a list of the insects of Massachusetts that Harris compiled (Hitchcock 1835). I believe it likely that Harris was referring to what is now known as C. limbata but because the 1829 correspondence was not published until 1869, and was not a formal description, the name C. comma is not valid.
Diagnosis. Adults of Cymindis limbata are distinguished from those of other species by: a pale, testaceous humeral macula (Fig. 8) extended from interval 6 (rarely from interval 5) to the outer margin and posteriorly as far as one quarter (0.25) the length of the elytra; pronotum broadly rounded; antennomere 8, 3.0-3.9 × longer than wide. In males, genitalia with endophallus having a distinct sclerotized patch (Fig. 10A, 11A) medially, phallus apex with distinct curvature to the left when viewed from dorsal aspect (Fig. 11C); phallus apex broadly pointed and distinctly shaped (Fig. 10A, 11B). In females, gonocoxite 2 (gc2) long and narrow, sharply pointed at apex; apical ensiform setae curved outward and extended almost to gonocoxite apices (Fig. 10B).
Microsculpture. Microlines not visible on dorsum of head capsule and pronotum at 50× magnification. Elytra with mesh pattern isodiametric, microlines clearly defined throughout dorsal surface.
Macrosculpture and pilosity. Head capsule with very fine, randomly scattered setigerous punctures on dorsal surface (setae not visible or only barely so at 50× magnification) from constriction of neck extended anteriorly toward clypeus. Elytra with striae moderately impressed and punctulate throughout length; intervals slightly convex (few with greater convexity in intervals 1, 3 and 5); one or two (most specimens one) irregular rows of fine punctures extended length of intervals 1, 3 and 5; two or three (most specimens two) irregular rows of fine punctures extended the length of intervals 2, 4 and 6; interval 8 with two to four rows of fine punctules extended interval length. Abdominal sterna with fine pilose punctures throughout.
Habitat, habits and seasonal occurrence. The known elevational range of C. limbata extends from sea level to 1935 m. Specimens have been collected on and under bark, in leaf litter and from under stones in forests of oak, pine, tamarack, aspen, beechmagnolia and maple. Specimens have been found near creek and pond margins, among beach wash-up and have also been collected from squirrel nests in trees. Adults are crepuscular or nocturnal with most activity being observed on tree trunks. Adults are most commonly collected from May to August. Most teneral adults were found from late June to early July, suggesting emergence from pupal case also occurs around this time.
Methods of collecting include asafetida and molasses bait traps, sugaring baits painted on tree trunks, beating and sweeping vegetation, at incandescent light and u.v. light, Malaise traps, Lindgren funnels, Berlese traps, flight intercept traps (FIT's), pitfall traps, and hand collecting.
Geographical distribution. The range of this species (Fig. 7) extends in eastern Canada from southern Quebec west to southern Ontario, and in the eastern United States east of the Appalachian Mountains from Maine south to southern Florida, west to eastern Colorado and Nebraska, and south to southern Texas.
Chorological affinities. Cymindis limbata is sympatric in the northern and western portion of its range with C. platicollis platicollis, in the southeastern portion of its range with C. platicollis atripennis and C. rufostigma, and in the central portion of its range with C. complanata.
Fossil specimens. Fossil packrat (Neotoma sp.) middens have been used to determine late Quaternary insect assemblages in the Chihuahuan desert areas of northern Mexico and southwestern Texas (Elias 1992, Elias and Van Devender 1992, Elias et al. 1995. Particularily relevant to this treatment are middens examined from the Trans Pecos and Bolson de Mapimi areas where both C. limbata and C. platicollis subfossils had been reported (Elias 1992). These species are primarily warm-temperate deciduous forest-adapted eastern species and currently both have southwestern range limits that do not extend to the Trans Pecos and Bolsom de Mapimi areas. Unfortunately, the specimens were unavailable for reexamination but in their place Scott Elias graciously sent me two S.E.M. images (a pronotum and a single elytron) of the putative C. limbata fossils.
I was able to determine that neither fossil specimen was C. limbata, based on examination of these images. First, The fossil image of the pronotum has narrowly bordered pronotal margins and hind angles that are almost right-angled. This contrasts with the pronotum of C. limbata, which typically has a widely explanate margin and hind angles that are rounded, or at least widely obtuse. As well, I have not observed C. limbata with deep punctures either side of the median longitudinal impression of the pronotum, a feature characteristic of the fossil specimen. The S.E.M. specimen appeared to be a lebiomorph carabid and I thought it to be a member of the genus Calleida so I sent the information to carabidologist Achille Casale (University of Sassari, Italy) who has detailed knowledge of the Mexican Calleida fauna. He confirmed that the pronotum was "Calleida-like" and appeared not to belong to Pinacodera (Casale 2008, personal communication).
The S.E.M. of the single fossil elytron also differed from that of C. limbata. The width of the fossil elytron was more than 5 mm whereas C. limbata elytra range from 3.2 to 4.75 mm. Additionally, C. limbata typically have more than one row of shallow and fine punctures in the even elytral intervals and more than one row in interval 8. The fossil specimen has a single row in all intervals, which are obviously deeper and farther apart than observed in C. limbata. After considering these discrepancies I believe it more likely that the elytron belongs to C. punctigera punctigera LeConte as it is similar in size and morphological characteristics, and was found within the limits of the current geographical range of C. punctigera punctigera. Cymindis punctigera punctigera has been collected in the recent past from nests of packrats of the genus Neotoma Remarks. This is a polytypic species that includes two subspecies (C. platicollis platicollis (Say) and C. platicollis atripennis (Casey)).
Diagnosis. Adults of Cymindis platicollis are distinguished from those of other species in the limbata species group through the following unique combination of external and genitalic characters: translucently bordered pronotum and elytra (Fig. 12,  16, 19B); evenly brunneo-piceous to rufo-piceous elytral disc coloration and interval 8 with two to four rows of scattered setigerous punctures; apical area of phallus dimpled (at least at apex) on both dorsal and ventral surfaces (Fig. 14A, 15A-B); longitudinal striations absent.
Microsculpture. Individuals with microlines not visible (or hardly so) on dorsum of head capsule and pronotum at 50× magnification. Elytra with mesh pattern isodiametric, microlines clearly defined.
Macrosculpture and pilosity. Head capsule dorsally with very fine to coarse scattered setigerous punctures on dorsal surface from constriction of neck extended anteriorly toward clypeus, ventrally with scattered pilose punctures laterally, extended from constriction of neck to gula. Pronotum with shallow to somewhat deep and randomly spaced setigerous punctures, setal length very short to moderate at 50×; ventrally with randomly spaced setigerous punctures extended from margin of proepipleuron to apex of intercoxal process; setae visible on prosternum but not proepisterna. Elytra with striae shallowly to moderately deeply impressed, punctulate throughout length; intervals flat to slightly convex; few specimens with odd intervals somewhat raised and even intervals flat. Other details in "Variation" section below.
Fixed setae. Pronotum with two setae along each lateral margin. Elytra with 14-16 lateral (umbilical) setae; two setae on each of abdominal sterna III to VI; four setae along apical margin of sternum VII (Fig. 3).
Geographical distribution. The range of this species (Fig. 17) extends in eastern Canada from southern Quebec west to southern Ontario; in eastern United States from Maine south to southern Florida, west to eastern Colorado and Nebraska, and south to southern Texas. In Mexico it is known from Nuevo Leon in the northern portion of the Sierra Madre Oriental..
Macrosculpture and pilosity. Head capsule dorsally with fine, randomly scattered setigerous punctures (setae not visible or only barely so at 50× magnification) from constriction of neck extended anteriorly toward clypeus. Elytra with striae moderately impressed and punctulate throughout length; intervals slightly convex (few with greater convexity in intervals 1, 3 and 5); abdominal sterna with fine pilose punctures throughout.
Luster. Head capsule and pronotum glossyl elytra moderately glossy. Pronotum. Anterior transverse impression shallow (Fig. 13); posterior transverse impression moderately deep; median longitudinal impression moderately shallow; posteriolateral angles almost right angled to obtuse. Hind wings. Macropterous. Male genitalia. Phallus (Fig. 14A) length 1.70 -2.42 mm. Variation. Through the range of C. platicollis platicollis a three-phased cline is observed (Fig. 17, Table 4). Phase 1: northeastern specimens have an average overall body length of 8.83 mm, average phallus length of 1.76 mm (Fig. 15A) and are dorsally glabrous; (~75%) with a single row of ~50-80 (interval 2 with more than 70) punctures in interval 1-7, some (~25%) with one to two rows of punctures in intervals 2, 4 and 6, all others with one row of punctures (except interval 8 with two to four rows). Phase 2: more southern specimens have an average overall body length of 9.22 mm, average phallus length 1.82 mm and have dorsal setation on humeral area of elytra; some specimens with few setae visible on dorsum of head and disc of elytra; (~86 %) with odd intervals bearing one row of scattered setigerous puncture and even intervals bearing two or three rows of scattered setigerous punctures, others (~14%) with interval 2 having two or three rows of setigerous punctures (rarely one row), interval 8 with two or three rows, and all other intervals bearing one row of setigerous punctures. Phase 3: southwestern specimens (extreme south west Oklahoma to mid-western Texas south west to Nuevo Leon) have an average overall body length of 9.92 mm, an average phallus length of 2.08 mm (Fig. 15B) and are dorsally setose, most individuals having one or two rows (mostly two) of moderately deep, randomly spaced, pilose punctures in all intervals, interval 8 with two to three rows. Few with various combinations of above.
Habitat, habits and seasonal occurrence. The known elevational range of C. platicollis platicollis extends from 3 to 1935 m. Specimens were collected under stones, under and on bark of trees and associated mosses in forests of buckeye, beech-magnolia, elm, hackberry, hickory, juniper, mesquite, oak, oak-pine, and tamarack. It has been collected from shrub species Leucaena pulver (Schltdl.), from bromeliads associated with oak and also from the nest of woodrats, Neotoma micropus Baird. Adults are crepuscular, and most commonly collected from late February through July. I witnessed several pairs in copula over a three-week period of collecting in Georgia and Florida from late February to mid-March of 2008. Of these mated pairs I brought 9 females and 10 males back to the University of Alberta to attempt rearing larvae. I kept them all in a single plastic container with substrate from under the trees they were captured on. A wet ball of tissue provided moisture, and several 3rd to 5th instar larvae of cabbage looper moth species Trichoplusia ni (Hübner) were introduced each week for food. All individuals (with the exception of one female) survived for the first three months in captivity. By mid-June (three months after capture) males started to die, and within the next two weeks all had expired. All of the remaining eight females lived at least until mid-September (6 months after capture) with the last individual dying in early November (7.5 months after capture). Males lived an average of 82 days after capture and females lived more than twice as long with an average lifespan of 166.5 days after capture.
All beetles were removed from the container every week and the substrate was searched for eggs and larvae. No evidence of oviposition was found. Other attempts to rear larvae from mated Pinacodera adults (Mahar, 1978) were also unsuccessful. Many lebiines are known to have unusual ovipositional habits or needs; that may also be the case in Pinacodera and a reason why rearing is problematic.
Collecting methods include asafetida and molasses traps, sugar baits painted on tree trunks, beating and sweeping vegetation, at light and u.v. light, Lindgren funnel traps, Berlese traps, Malaise traps, flight intercept traps (FIT's), pitfall traps, hand collecting, and sticky traps.
Geographical distribution. The range of this subspecies (Fig. 17) extends in eastern Canada from southern Quebec west to southern Ontario; in the eastern United States from Maine south to mid-Georgia west to eastern Colorado and Nebraska south to southern Texas. In Mexico it is known from Nuevo Leon in the northern portion of the Sierra Madre Oriental.
Evolutionary affinities. This subspecies is, by definition the closest relative of C. platicollis atripennis.
Microsculpture. Microlines not visible on dorsum of head capsule and pronotum at 50× magnification. Elytra with mesh pattern isodiametric, microlines clearly defined throughout. Macrosculpture and pilosity. Head capsule, dorsal surface with shallow, randomly scattered setigerous punctures on dorsal surface from constriction of neck extended anteriorly toward clypeus. Elytra with striae moderately impressed and punctulate throughout length; intervals 2, 4, 6 and 8 typically with two to three rows of scattered punctures; all other intervals with one row of punctures.
Metepisternum. Distinctly longer than wide. Hind wings. Macropterous. Male genitalia. Phallus (Fig. 20A) length 1.70 -2.00 mm. Collection notes and habitat. The known elevational range of C. platicollis atripennis extends from sea level to 90 m. Specimens were collected on forest floor, under and on bark of trees and associated mosses in forests of cypress, juniper, magnolia, several oak species and both slash and loblolly pine. This subspecies has been collected from sand dunes in close proximity to water as well as from squirrel and caracara nests in trees. Methods of collecting include: dusk-dawn suction traps, medfly traps, u.v. and light traps, FIT's, sugaring baits painted on tree trunks, beating vegetation, hand collecting, Steiner traps, pitfall traps, and sticky traps. Geographical distribution. The range of this subspecies extends from southern Florida north to mid-Georgia and west to southern Mississippi (Fig. 17).
Morphological affinities. This subspecies is, by definition, the closest relative of C. platicollis platicollis.
Chorological affinities. C. platicollis atripennis is sympatric in portions of its range with C. limbata, C. complanata and C. rufostigma. It is allopatric with C. platicollis platicollis and all other species in the limbata group. Type locality. Archbold Biological Station, Highlands County, Florida, U.S.A. Specific epithet. This is a two part feminine Latin noun in apposition, nominative case, based on the adjective rufus (red) and the noun stigma (mark), referring to the rufo-testaceous central mark on the elytra of adult members of this species.
Macrosculpture and pilosity. Head capsule with shallow, randomly scattered setigerous punctures on dorsal surface from constriction of neck extended anteriorly toward clypeus. Elytra with striae moderately impressed and punctulate throughout length; intervals 2, 4, 6 and 8 typically with two to three rows of scattered punctures; all other intervals with one row of punctures; some specimens with intervals 1, 3 and 5 somewhat raised toward apex.
Habitat and seasonal occurrence. The known elevational range of C. rufostigma extends from sea level to 65 m. Some specimens were collected in stands of slash pine. Methods of collecting include: dusk-dawn suction traps, u.v. light traps, CO2-baited insect light traps.
Geographical distribution. The range of this species extends from southern Florida north to northern Florida (Fig. 21). Chorological affinities. Cymindis rufostigma is sympatric in portions of its range with C. platicollis atripennis and C. limbata. It is allopatric with C. platicollis platicollis and all other species in the limbata group.
Material examined. The type material. See above for details.

punctigera complex
The punctigera complex includes only a single species; see C. punctigera LeConte below for details.

Cymindis punctigera LeConte http://species-id.net/wiki/Cymindis_punctigera Figs 22-29
Remarks. This is a polytypic species that includes two subspecies (C. punctigera punctigera and C. punctigera sulcipennis). The basis for including these forms in a single species is as follows: first, similarity in body proportions; second, males exhibit similarity in details of the apical area of the phallus (Fig. 27B-C) and everted endophallus form (Fig. 27A); and third, allopatric but proximate geographical distribution. Diagnosis. Adults of Cymindis punctigera are distinguishable from those of other species of the limbata species group through the unique combination of: pronotum with scattered and distinct punctures (Fig. 24), one row of punctures in each each elytral interval, and dorsal color that ranges from testaceous to dark brown, some specimens approaching piceous, but if so, the elytral margins are translucently bordered; in males the apical area of the phallus is dimpled on both the dorsal and ventral surfaces, longitudinal striations are visible from mid-way of apical area and extended past constriction ( Fig. 27B-C), and the endophallus is distinctly bent to the right and dorsad when everted and viewed from the dorsal aspect (Fig. 27A).
Microsculpture. Most individuals with microlines not visible on dorsum of head capsule and pronotum at 50× magnification; few specimens with mesh pattern isodiametric to transverse between eyes and on disc of pronotum. Elytra with mesh pattern isodiametric, microlines clearly defined throughout dorsal surface.
Macrosculpture and pilosity. Head capsule with randomly scattered setigerous punctures on dorsal surface from constriction of neck extended anteriorly toward clypeus.
Pronotum dorsally with shallow and randomly spaced setigerous punctures with setae length ranging from very short to moderate; ventrally with setigerous punctures extended from margin of proepipleuron to apex of prosternal intercoxal process. Elytra with striae moderately impressed and punctulate throughout length; intervals slightly convex to markedly raised towards center of interval; single regular to moderately irregular row of ~50-65 setigerous punctures within each interval, setae length ranging from very short to moderate at 50×. Abdominal sterna with fine pilose punctures throughout.
Fixed setae. Pronotum with two setae along each margin. Elytra with two setae in stria 3 and one posteriad of stria 3; one seta at apex of interval 2; 15-17 lateral (umbilical) setae; two setae on each of abdominal sterna III to VI; four setae along apical margin of sternum VII (Fig. 3).
Luster. Head capsule and pronotum distinctly to slightly glossy; elytra glossy to rather dull; ventral thoracic sclerites and abdominal sterna glossy.
Pronotum. Anterior and posterior transverse impression shallow; median longitudinal impression moderately shallow; posteriolateral angles from almost right angled to slightly obtuse; posterior margin slightly lobate.
Geographical distribution. The range of this species extends (Fig. 29) in the southwestern United States from Lake Tahoe, California south through southern Utah and Nevada to western Texas; south in eastern Mexico to Nuevo Leon in the northern portion of the Sierra Madre Oriental; in western Mexico, on the slopes of the Sierra Madre Occidental to Jalisco, and in the Sierra Transvolcanica, to Michoacan. Further west this species ranges through most of the Baja California Peninsula.
Chorological affinities. Cymindis punctigera is sympatric in the southern portion of its range with C. chevrolati and C. platicollis platicollis. [handwritten]. PARALECTOTYPE; second specimen female, labeled same as lectotype, and "punctigera 2"; third specimen male, labeled same as lectotype, and "punctigera 3". (Two more specimens, each labeled "Ariz" and "punctigera 4" and "punctigera 5", respectively, were probably assigned to this species at a later date and are not recognized as types). Notes about types, homonymy, and synonymy. The reason for assigning C. punctigera to Apenes (Blackwelder 1944: 62) is not apparent. The type specimens of the named forms, taken in isolation, are distinctive and differ from one another as previous authors indicate. Examination of material geographically adjacent to these forms reveals that this species is quite variable throughout its range, and characters from Casey's descriptions are not diagnostic.

Cymindis punctigera punctigera
Diagnosis. Specimens of this subspecies (Fig. 22A-B) exhibit little to no rugosity on dorsal surface of head from between eyes toward clypeus.
Metepisternum. Individuals are at least 1.44× as long as wide. Hind wings. Macropterous or brachypterous. Male Genitalia. Phallus (Fig. 26A) length 1.70 -2.04 mm. Variation. Within this subspecies, dorsal pilosity, body coloration and wing state are, to some extent, variable geographically. The least understood is seta length on the dorsal surface of individuals. In some areas (mainly montane) populations of C. punctigera punctigera have uniformly long setae covering their dorsal surface while others from surrounding localities have setae so short that they are hardly visible at 50× magnification. Color of the dorsal surface ( Fig. 22A-B) varies from dark rufous over most of the species range (Fig. 22A) to predominantly testaceous to rufo-testaceous (Fig. 22B) in the Big Bend area of southwestern Texas.
Wing state and wing length vary in this subspecies. Examination of a few hundred individuals, from all known localities showed that C. punctigera punctigera exhibits a general trend toward brachyptery in the periphery of its range and macroptery in the central portion (Fig. 29). Population samples from two localities (Washington Co., Utah and Riverside Co., California) included both macropterous and brachypterous individuals. As well, average length of wing rudiments of brachypterous specimens differed between localities with the shortest being from Nuevo Leon, the same place where members of C. chevrolati exhibit the shortest average wing rudiments.
Collection notes and habitat. The known elevational ranges of C. punctigera punctigera extends from 670 to 2500 m. Most specimens were found at elevations higher than 1000 m. Specimens were collected under stones and bark of trees in forests of oak, pine, mesquite (Prosopis glandulosa Torrey), in stands of acacia and desert willow (Chilopsis linearis Cav.). As well they occupy riparian temperate forests, meadow, desert and pond margin habitats. This subspecies has been collected from nests of packrats of the genus Neotoma.
Geographical distribution. The range of this subspecies (Fig. 29) extends in the southwestern United States from Lake Tahoe, California south through southern Utah and Nevada to western Texas south to the interior of Mexico, both sides of the Sierra Madre Occidental, east as far east as Nuevo Leon, south to Michoacan and as far west as Nayarit.
Morphological affinities. This subspecies is by definition the closest relative of C. punctigera sulcipennis.
Chorological affinities. Cymindis punctigera punctigera is allopatric with C. punctigera sulcipennis and all other species of the limbata group except C. chevrolati and C. platicollis platicollis, which are sympatric with it toward the eastern limits of its range. Type locality. La Paz, Baja California Sur, Mexico, here designated. Notes about synonymy. Horn (1881:40) distinguished Pinacodera sulcipennis from Pinacodera semisulcata through the former having raised elytral intervals. Careful examination of more than 80 specimens revealed that some members of this subspecies (32%) have raised elytral intervals, especially in the basal third. This is the rationale for combining P. semisulcata with P. sulcipennis.
Diagnosis. Most specimens of this subspecies (94.2%) have two to several rugulose transverse lines on dorsal surface of head between eyes with rugosity in some extended to clypeus (Fig. 25B).
Geographical distribution. This subspecies is restricted to Baja California in Mexico (Fig. 29), ranging from the southern tip of the peninsula, to as far north as San Quentin, Baja California Norte.
Macrosculpture and pilosity. Head capsule with evenly scattered setigerous punctures on dorsal surface from constriction of neck extended anteriorly toward clypeus. Prothorax ventrally with fine setigerous punctures extended from lateral margin of coxal cavity to apex of intercoxal process. Elytra with striae moderately impressed and punctulate throughout length; intervals slightly convex, single irregular row of ~30-45 punctures within each interval. Abdominal sterna with pilose punctures throughout, setae increased slightly in length toward baso-lateral margins.
Fixed setae. Two pairs of supraorbital setae; clypeus with two lateral setae. Labrum with six setae along apical margin. Pronotum with two to five setae along each margin. Elytra with two seta in stria 3 and one beyond apex of stria 3; one setae at apex of interval two; 15-17 umbilical setae; two setae on each of abdominal sterna III to VI; 4-8 setae along apical margin of sternum VII (Fig. 3).
Luster. Head capsule and pronotum glossy; elytra glossy to slightly glossy, ventral thoracic sterna and abdominal sterna glossy.
Geographical distribution. The chevrolati complex is known from all the major mountain systems of Mexico north of the Isthmus of Tehuantepec (Fig. 41). It is also known from the Pacific Tres Marias islands.
Chorological affinities. The geographical range of the chevrolati complex overlaps the range of one other member of the limbata group (C. punctigera punctigera LeConte) and the ranges of several species of the latiuscula group (Hilchie and Ball, in preparation).
Overall body length. Twenty individuals (f=10, m=10) from state population samples of each species in the chevrolati complex were compared (Fig. 38). In each species males are shorter on average then females. Overall body length between C. chevrolati from Durango and C. laevior from Oaxaca, while statistically significant, was not useful taxonomically because of extensive overlap in both males and females.
Hind wing length. The three species in the chevrolati complex are brachypterous, showing metathoracic wing reduction, from slight ( Fig. 39B) to extensive (Fig. 39A). Population samples (Fig. 40) were measured to determine possible trends in wing length. Overall, wing length ( Fig. 40) of C. chevrolati is the most variable of the three species both between and within populations. Of individuals examined, C. laevior had the lowest variation throughout its range, though a slight decrease in length is observed from north to south. Specimens of Cymindis ruficornis had the shortest average wing length.
In the range of distributional overlap between C. chevrolati and C. laevior, wing length serves as a diagnostic character between the two species in Hidalgo (Fig. 40). In populations from Puebla, however, some overlap does occur.
Metepisternum reduction is associated with wing reduction in Carabidae (Darlington 1943). Cymindis chevrolati has a metepisternum that is 1.66 to 2.00× longer than wide, C. laevior is 1.40 to 1.67× longer than wide and C. ruficornis is 1.35 to 1.45× longer than wide. This is strongly correlated with the reduction of wing length observed in chevrolati complex members.
Several hypotheses have been proposed for the evolution of brachyptery in insects (Darlington 1943, Kavanaugh 1985, Liebherr and Hajek 1986. Cymindis laevior and C. ruficornis (the postulated sister group and closest relative to C. chevrolati) have reduced wings that can be explained as a result of high altitude occurrence and relatively stable montane habitat. Cymindis chevrolati is also found almost exclusively at similar elevations in montane habitats but has a much greater range in wing length.
It is unclear what selective pressures may be affecting wing length in C. chevrolati. No geographical trends are apparent. Specific epithet. The original spelling is "chevrolatii", but relatively recent catalogues (Csiki 1932, Blackwelder 1944, Lorenz 2005, have omitted the terminal "i", thus "chevrolati". Such a modification produces an incorrect subsequent spelling, but I accept the catalogue entries as "prevailing usage", and the latter name as a correct original spelling (International Code of Zoological Nomenclature, 1999: 43, art. 33.3.1).
As implicitly indicated by Dejean (1836: 9), the name C. atrata Chevrolat was a junior homonym of C. atrata Dejean (1831, a species that was subsequently transferred to the genus Inna Putzeys (Chaudoir 1872) and then to Eucheila Dejean (Shpeley and Ball 2001). As a replacement name, Dejean proposed Cymindis chevrolatii, using as a voucher specimen a male received from Louis Reiche, labeled "chevrolatii mihi/ atrata Chevrolat/ Mexico, d. Reiche " [green paper, handwritten]. This specimen is in the Chaudoir/ Oberthür collection, in front of the box label "nigrita Chaudoir" (MNHP). That specimen might have come originally from the Chevrolat collection to Reiche, and then to Dejean (ultimately to Chaudoir). Chaudoir (1837:8) recorded that he had received specimens from Chevrolat of C. atrata, which he renamed C. nigrita, evidently being unaware that Dejean had provided a new name in 1836. These four specimens (3 males, 1 female, each labeled "Ex Musaeo/ Chaudoir" [red print] ), are in front of the box label "nigrita / Chaud./ Mexique/ Cruz Blanca/ Sallé" (MNHP). As demonstrated by similarity in the male genitalia, the male lectotype is conspecific with the Dejean voucher for C. chevrolatii, and for C. atrata Chevrolat.
Authentic specimens of C. amblygona Bates and C. angulifera Bates were not located in neither The Natural History Museum (London) nor the Museum National d'Histoire Naturelle (Paris), though they were sought in both of those collections that are known to house the Bates material on which his New World work was based. Bates (1883: 187) noted that the features he had originally used to distinguish these two forms were exceedingly variable, even within single populations, citing as an example the series collected at Tehuacan, Mexico. Regarding C. amblygona, he stated that "even the definition of the form as a variety is impossible". Possibly, then, he removed the type labels from what were the type specimens of these forms. Bates described two additional varieties of "C. atrata": C. a. ruficornis and C. a. laevior. These are treated below, as separate species.
Diagnosis. Adults of Cymindis chevrolati (Fig. 30) are distinguishable from those of other species of the limbata species group through genitalic characters: in males a distinct microtrichial patch (mp) on the basal endophallic lobe (bel) of the endophallus (Figs 34A, 36). This patch can be seen in many males through the cleared phallus with endophallus inverted, located near the apex of the phallus. From the right lateral aspect, an everted sac has the microtrichial patch located on the dorsal surface of the basal lobe of the endophallus. Female genitalia differ from other species in the short, stout form of gonocoxite 2 (gc2) (Fig. 35, A2).
Microsculpture. Elytra with mesh pattern isodiametric, microlines shallow in basal half of elytra and shallow to absent from apical half.
Macrosculpture and pilosity. Dorsal punctures with setae present though short and almost not visible at 50× magnification. Ventral surface of head with evenly scattered setigerous punctures (bearing somewhat long setae) from behind eye laterally toward mentum. Pronotum normally with relatively evenly scattered setigerous punctures throughout (Fig. 31A), more densely so toward margins; few specimens with setigerous punctures along margin and few to no punctures on disc (Fig. 31B). Elytral epipleuron glabrous.
Fixed setae. Pronotum with two fixed setae along each margin. Four to six setae (typically four) along apical margin of sternum VII (Fig 3).
Luster. Elytra glossy in basal two thirds, in some specimens slightly less so in apical third.
Female genitalia. Gonocoxite 2 (gc2) (Fig. 35A) short and stout. Collection notes and habitat. The known elevational range of C. chevrolati extends from sea level to 3400 m, though it is important to note that specimens found at or near sea level were collected from the Tres Marias Islands, the immediately adjacent mainland and one other additional record from El Ebano, San Luis Potosi, that may be mislabeled or misinterpreted by me as Ebano, San Luis Potosi. Typically this species is found further inland and at higher elevation, mostly between 2000-3000 m. Specimens were collected under and around woody debris and stones in forests of oak, pine, fir, juniper and alder, in thorn scrub, and in stands of yucca. As well, the species occupies meadow, desert and grassland habitats.
Geographical distribution (Fig. 41) The range of this species is restricted to Mexico, extending in the Sierra Madre Occidental from Chihuahua south to Jalisco, in the Sierra Madre Oriental from central Nuevo Leon south to Hidalgo, and in the Transvolcanic Sierra of central Mexico as far south as central Puebla.
Morphological affinities. Based on genitalic characteristics and wing length states (Fig. 40), I postulate that C. chevrolati is the closest relative of C. laevior + C. ruficornis.
Macrosculpture and pilosity. Dorsal punctures with setae present, very short, hardly visible at 50×. Head ventrally with sparse, scattered setigerous punctures from behind eye laterally toward mentum. Pronotum normally with relatively evenly scattered setigerous punctures throughout, more densely so toward margins; few specimens with setigerous punctures along margin and few to no punctures on disc. Elytral epipleuron glabrous.
Fixed setae. Pronotum with two setae along each margin. Four to six setae (typically four) along apical margin of sternum VII (Fig. 3).
Luster. Elytra glossy throughout. Head (Fig. 37E-G). Mental tooth form varied. Hind wings. Markedly reduced, 1.06-1.57 mm in length, mean 1.32 mm. Male genitalia. Phallus (Fig. 34B) length 2.40-2.60 mm. Female genitalia. Gonocoxite 2 (gc2) (Fig. 35B) long and narrow. Collection notes and habitat. The known elevational range of C. laevior extends from 1524 to 3400 m. Specimens have been collected under bark, in woody debris and in leaf litter associated with forests of oak, pine, alder, juniper and stands of yucca. They have also been collected from bromeliads growing on standing trees. Because adults are incapable of flight, their presence above ground indicates the ability to climb trees.
Geographical distribution (Fig 41) The range of this species is restricted to Mexico, extending from northern Hidalgo in the eastern Transvolcanic Sierra east to western Veracruz and west to eastern Jalisco. The range extends in the Sierra Madre del Sur southward through the Sierra Madre de Oaxaca south to the Sierra de Miahuatlan from southern Oaxaca, and as far west as western Oaxaca.
Evolutionary affinities. Based on genitalic characteristics and wing length states, I postulate that C. laevior is the closest relative of C. ruficornis.
Microsculpture. Elytra with mesh pattern isodiametric, microlines shallow in most specimens, in few specimens more so in basal third.
Macrosculpture and pilosity. Dorsal punctures with setae present, erect and easily visible at low magnification. Ventral surface of head with evenly scattered setigerous punctures (bearing somewhat long pilose setae) from behind eye laterally toward mentum. Pronotum with evenly scattered punctures and pilose setae over entire dorsal surface. Elytral epipleuron with scattered punctures and pilose setae, setae longer and more regular in apical half.
Fixed setae. Pronotum with two to five fixed setae along lateral margin; elytra with two setae in stria 3 and 16 to 18 umbilical setae; two setae on each of abdominal sterna III to VI; four to eight setae along apical margin of sternum VII, typically six.
Geographical distribution. The known range of this species is restricted to the easternmost portion of the Sierra de Atoyac, in eastern Guerrero (Fig. 41).
Morphological affinities. Based on genitalic characteristics and wing length states (Fig. 40), I postulate that C. ruficornis is the closest relative of C. laevior.
Chorological affinities. Cymindis ruficornis is allopatric in relation to the other members of the chevrolati complex and all other members of the limbata species group.
Material examined. I have examined 116 specimens; 10 males and 10 females were dissected. For details see University of Alberta Strickland Virtual Entomology Museum Database (University of Alberta 2009).

Concluding remarks
It it not often that as much material is available (over 4000 specimens) as was for this revision. The large number of both borrowed and collected specimens allowed for examination of most species of the limbata group. throughout their ranges. Beyond determining how many species are in the limbata group and their geographical distributions, other interesting findings were made in regard to habitat preferences, geographical variation, macroptery/bachyptery, and possible hybridization.
Field collecting and integration of available label data has allowed insight into the special niche requirements of C. complanata. All specimens of C. complanata collected for this revision (13) were taken from slash pine (Pinus elliotii Engelm.) in mixed forest. Some borrowed specimens were also recorded from loblolly pine (P. taeda L.), which is a very similar species of tree. Having evolved a flattened, reddish body that blends well into the flaky bark of these trees, it would appear that C. complanata utilizes these trees specifically. This is very interesting as no other members of the limbata group seem to have habitat preferences that are so specific.
I was very fortunate to examine nearly 900 specimens of C. platicollis platicollis, from localities throughout its range. This allowed for an examination of possible patterns of geographical variation. Consequently, a survey of the external morphology exposed a northeast to southwest cline of increasing dorsal seta length that coincided with body size increase and elytral puncture change. With this pattern understood, 2 new synonomies were uncovered that may have otherwise gone unnoticed.
The availablilty of large number of specimens of C. punctigera punctigera also allowed for a thorough examination of the wing states of individuals throughout its range. An interesting pattern of macroptery/brachyptery was revealed, with brachypterous individuals typically being found near the outer limits of the range and macropterous individuals in the center. Most brachypterous individuals of C. punctigera punctigera live in the Mojave, Sonoran and Chihuahuan deserts, so desert associated conditions almost certainly play a role in this observed pattern. Wing reduction or loss has been tied to hot, dry, desert conditions in members of several insect taxa (Brues 1903;Peck 2008). This is partly because wing surfaces provide a major outlet for water loss and because of this, are more readily selected against in areas with these condition (Tinaut 1992). More work has to be done however, to determine what other mitigating factors may be influencing the observed pattern.
Analysis of genitalic characters, wing length, mental tooth form, and body size variation affirmed the presence of three chevrolati complex. As well, dissections of many males of both C. chevrolati and C. laevior from their narrow range of overlap revealed that some individuals had a highly reduced (few small microtrichia) microtrichial patch on the basal endophallic lobe. This appears to be an intermediate genitalic feature between the two species as C. laevior has no microtrichial patch and members of C. chevrolati have a patch with several microtrichia. Interestingly, the majority individuals with a reduced microtrichial patch were also observed to have a mental tooth form that was also "intermediate". This suggests the possibility that gene flow between C. chevrolati and C. laevior may be occurring in their area of geographic overlap.
Many of these observations would not have been possible without the substantial amount of specimens available for this revision. This illustrates the importance of having as much material as possible when doing revisionary work. In the future, the incorporation of molecular methods will likely help to further refine phylogenetic relationships within the limbata group. It may also aid in an examination of gene flow in the areas of sympatry between species.