CLASSIFICATION, NATURAL HISTORY, AND EVOLUTION OF THE EPIPHLOEINAE (COLEOPTERA: CLERIDAE). PART II. THE GENERA CHAETOPHLOEUS OPITZ AND PLOCAMOCERA SPINOLA

Abstract This study deals with the sister genera Chaetophloeus, new genus and Plocamocera Spinola of the checkered beetle subfamily Epiphloeinae. Chaetophloeus is monotypic and is described on the basis of C. hispidus, new species. Plocamocera is revised to include 35 species as follows: P. castanea, new species; P. pupula, new species; P. jayhawkalis, new species; P. confrater Kuwert; P. procera, new species; P. aliguantula, new species; P. minima, new species; P. manausensis, new species; P. coactilis, new species; P. sericella Spinola; P. auratilis, new species; P. argentea, new species; P. lucis, new species; P. sesquipedalis, new species; P. salasis, new species; P. iota, new species; P. prolixa, new species; P. quadrula, new species; P. baria, new species; P. aura, new species; P. buenavista, new species; P. taruma, new species; P. aspera, new species; P. paris, new species; P. bispina, new species; P. onorei, new species; P. carnegei, new species; P. santa, new species; P. sericellopsis, new species; P. insula, new species; P. ambra, new species; P. similis, new species; P. bolivari, new species; and P. selva, new species. A neotype is selected for P. sericella Spinola. Lectotypes are designated for P. confrater Kuwert and P. byssinus Erichson (junior synonym of P. sericella Spinola). Five new synonymies are recognized, they are P. confrater var similis Kuwert (junior synonym of P. confrater Kuwert), P. confrater var. sericelloides Kuwert (junior synonym of P. confrater Kuwert), P. impressicollis Pic (junior synonym of P. confrater Kuwert), Epiphloeus byssinus Erichson (junior synonym of P. sericella Spinola), and P. latefasciata Pic (junior synonym of P. sericella Spinola). Plocamocerans are diurnal, but are considerably active at night, highly cryptic on bark, when disturbed conduct flylike escape behavior, and are predators of lignicolous insects. The morphology of integumentary sensilla on the antenna, pronotum, and elytra are described, and SEM photographs of them is provided. Described for the first time are four types of antennal sensilla (one type of sensilla trichodea, two types of sensilla chaetica, and one type of sensilla basiconica), four pronotal trichobothria, and three types of pronotal sensilla trichodea, and on the elytra of Plocamocera confrater Kuwert eight trichobothria and two types of sensilla trichodea. It is postulated that filamentous sensilla trichodea on the antenna serve to perceive volatile tree trunk chemicals and/or aggregate pheromones of prey bark beetles. The study includes a review of trichobothrial function and distribution among insects. Morphological analysis is extended to include structure of the internal reproductive organs of some plocamoceran species. Also included are a discourse on species-level discontinuities; discussion of the specimen-study methods; methods for phylogenetic analysis; a treatise of evolutionary relationships based on Hennigian principles implemented in a data base analysis involving Farris's Hennig 86 which produced a computer-based phylogeny involving genera, species groups, and species; a list of specimen repositories (with e-mail addresses); key to species groups and species; table of character-state analysis; four color photographs; 20 SEM photographs; a total of 219 illustrations; 12 distribution maps; and one diagram of phylogeny that depicts a hypothesis of inter-and intrageneric evolutionary relationships. The 35 plocamoceran species are categorized into six species groups whose combined distribution extends from southern Mexico to southern Brazil. The members of the confrater and coactilis species groups are superficially very similar; their aedeagal structure and shape of antennomeres must be examined in detail for credible identifications at the species level. The sister group relationship between Chaetophloeus and Plocamocera is supported by the following four synapomorphies, they are pedicel very large, aedeagus lanceolate, male accessory glands reduced to two pairs, and metabasitarsomere one longer than metabasitarsomere two. Profuse distribution of bristlelike setae (chaetosomes) on the dorsum, forebody, and elytral disc and presence of moderately elongated sensilla trichodea on the antenna are autapomorphies for Chaetophloeus, whereas filamentous sensilla trichoidea on the antenna, boldly transverse pronotum, very robust metacoxa, very robust metafemora, presence of trichobothria on the epipleural margin, and sex dimorphism of abdominal color are synapomorphic character states that establish the monophyly of Plocamocera and define its ancestral basic plan. It is postulated that South America was the ancestral environs of Chaetophloeus and Plocamocera and that there were three independent northward extensions of major plocamoceran lineages into Central America.


INTRODUCTION
In a very relevant way, this study began in the laboratory of Ayodyha P. Gupta, Emeritus Professor of Entomology, Rutgers University. His stern, but fair, tutelage as a graduate advisor inspired uncompromising scientific writing and critical thinking in functional morphology. During that time, I met William F. Barr, Gupta's graduate advisor and renowned cleridologist. The setting for a career in Systematic Entomology was complete.
When one commits to a taxon specialization there is an innate impulse to borrow, collect, and borrow some more, then, to investigate live specimens in the field-all meant to gain a thorough perspective of the ''gestalt'' variations of the organisms under study. Lee Herman, of the American Museum of Natural History, and the late Hank Dybas, of the Field Museum of Natural History, were very generous stewards of collections, and with their constant support and encouragement they fueled the fire in an incipient cleridologist. Among their collections I found a few seemingly uninteresting clerids that showed three unusual features. First, the antennae were somewhat plumose; second, the hind femora and hind coxae were partic-ularly robust (fig. 18); and third, the pronotum showed a curious set of discal and paralateral ''hairs'' (long filiform setae set in a spheroid depression) (figs. 7, 9). The filiform hairs, subsequently identified as the trichs of trichobothria and signature autapomorphy (uniquely derived character state) of the Epiphloeinae, I presumed to be sensory organs, whereas I interpreted the stout development of the hind legs as a specialization involving some form of saltation.
Jumping, however, was not in the venue of behavior that I experienced during my early exposure to live clerids. These beetles scurry (Perilypus), play dead upon disturbance (Pelonium), or abruptly take flight (Phyllobaenus), never having a suggestion of jumping despite the moderately thickened femora of some species. Then, in 1981, a sojourn to the Amazonian rain forests of Brazil provided the opportunity to observe and collect a variety of clerids. In a forest clearing (much like the one depicted in fig. 4), aside the Rio Negro, I noted an inordinate quantity of small insects catapulting from a felled hardwood of Manilkara sp. At first, I presumed that these insects were flies, but to my astonishment they proved to be plocamocerans, and to my great delight I realized that, indeed, ''jumping checkered beetles'' do exist. This is the second of a series of publications intended to more clearly describe and make more accessible the known species of Epiphloeinae, to place the species into a credible arrangement of genera, and to provide hypotheses of evolutionary relationships at the species, genera, and suprageneric levels. The first contribution (Opitz, 1997), a synopsis of the epiphloeine genera, intended to familiarize collectors and curators with the generic entities of the subfamily. With subsequent contributions I hope to generate interest toward collection, fluid preservation, indentification, and field study of additional specimens of epiphloeines and to gradually build up an inventory of evidence toward more complete and heuristic classifications and hypotheses of epiphloeine evolution, as well as of other taxa within Cleridae akin to Epiphloeinae. With few exceptions, there is a great need for larval specimens in our studies of Cleridae systematics.
Inherent in my contributions is the strong desire to promote a most important edict of our profession, that is, it is imperative that systematists present the results of their efforts in such a way that they stimulate and serve the broader community of biologists. In my view, this is done by providing a detailed table of contents, thorough and efficient descriptions of species, user-friendly diagnostic keys, an abundance of relevant illustrations, and refutable syntheses of evolutionary relationships. In this context, the recent work of my colleague Roland Gerstmeier (1998) is an outstanding example. In that work Gerstmeier presents, in a most eloquent manner, the clerid fauna of the Western Palearctic. Ultimately, our goal is to establish logically sound, stable, and heuristic classifications. In my view, such classifications are of greatest value when they are based on substantive estimates of phylogeny. No one has expressed this view of classification more succinctly than did Charles Darwin (1859: 486) when he prophetically wrote, ''Our classifications will come to be, as far as they can be so made, genealogies. . . ''. Moreover, proposals of phylogenies /classifications, particularly at the suprageneric levels, which often involve cumbersome nomenclatural changes, should include ''ge-stalt'' level analyses and a comprehensive understanding of species-level systematics. The eminent G. H. Horn (1887: 7) implies this very point as he noted, ''Progress in Natural History necessarily starts from a basis of species, and until these are accurately described so that others can arrive at a knowledge of them no great advance is possible' '. Anatomically, chaetophloeans and plocamocerans are very distinctive among epiphloeines. However, at infrageneric levels, plocamocerans are notably homogeneous to an extent that very few variations of character states are suitable for hypotheses of evolution. The lengthy filamentous-like setae on the antenna ( fig. 10), boldly transverse pronotum ( fig. 7), sexual dimorphism in color of the abdominal venter, and presence of only one pair of male accessory glands are uniquely derived features within the subfamily and establish the hypothesized basic body plan of ancestral Plocamocera.

MATERIALS AND METHODS
This study is based on approximately 900 adult specimens, some of which I studied in the field during an expedition to the Amazon Basin in 1981. Specimens were observed during daylight and evening hours, collected, and then preserved in Pampel's fluid (Ekis, 1977: 6) for study of internal organs. Not all species descriptions contain information about internal visceral or internal reproductive structure. Inclusion of such information was based on availability of fluid-preserved specimens. A few specimens were completely disarticulated to determine the range of anatomical differences among beetles of different species and to establish a foundation of structural variations for future assessments of evolutionary relationships at suprageneric levels.
Techniques of dissection, illustration, and procurement of meristic data were similar to those I described in my work on Perilypus (Ekis, 1977: 6). Also, the conventional integumental terminology used herein stems from that paper and from Nichols (1989). Names of antennal and elytral sensilla were adopted from Callahan (1975: 390). In the illustration of the alimentary canal, only two of four cryptonephridial malpighian tubules are drawn. The figures of the female and male internal reproductive organs show only one of two ovaries and one of two testes and accessory glands, respectively. I examined the primary type specimen of all nominal species, except that of P. sericella. Spinolas' type is presumed lost; a neotype of P. sericella Spinola has been selected during this project.
To some extent, color of the elytral surface and distribution of elytral setae are diagnostic of plocamoceran species. The elytral patterns, however, are sometimes obscured by intraspecific variations and by the varied physical condition of the elytral surface. Therefore, in my drawings of elytral color (see fig. 52), I have selected a specimen that best exemplifies the elytral surface pattern of a species. I have illustrated a left elytron to represent the color pattern of nearly all recognized species. However, the outline, or shape, of these elytra should not be taken to signify the accurate representation of elytral form of the species represented. Further, the abovementioned elytral outlines do not show the full complement of elytral trichobothria. For example, the non-SEM assessment of elytral trichobothria is depicted in figures (see fig. 53), whereas during SEM analysis it was discovered that more, minute trichobothria actually occur. Moreover, it was determined that the level of development of the elytral trichobothria ranges from highly developed (fig. 16) to moderately developed ( fig. 17). Only those trichobothria that could be observed with certainty at 750ϫ were illustrated by hand. A definitive assessment of the number of elytral trichobothria for each species will have to be determined with SEM magnifications, as was done for the elytral trichobothria of P. confrater Kuwert. An important component of applied systematic work is to assure the unquestionable indentity of primary types. I have experienced considerable difficulty with type specimens established by some of our pioneer workers in Cleridae. To minimize future problems in this area, my descriptions include the exact sequence of labels associated with each primary type. Also, locality label information of each primary type specimen is duplicated in exact sequence in the descriptions, whereas such information affiliated with secondary types is placed in the following sequence . . . country: state or province: specific locality; dates of specimen collection, natural history data, collector, and repository. In the bibliography, I have appended my current name to my published former name. The name change resulted from an inquiry and discovery of the identity of my biological father, a casualty of WW II.

ASSESSMENT OF SPECIES-LEVEL DISCONTINUITIES
Species status was postulated in accordance with the biological species concept as advocated by Mayr (1969). Conjectures about reproductive isolation are primarily based on indirect evidence of structure and to a lesser extent on the basis of spatial occurrence and presumed extant and/or historical geographical barriers. Differences in male genitalia, body size, shape of antennomeres and pygidia, and elytral pubescence patterns were very helpful in predictions of gene-flow discontinuites, as was, to a lesser extent, the predicted species limits of distribution due to refugial systems (Haffer, 1969;Brown, 1975;Ekis, 1977).
Among species of Cleridae, variations of male, and in some cases female, genitalia are very reliable criteria for supporting notions about reproductive isolation. However, there are several groups within the family in which such variations are minimal, as is the case in Plocamocera. Similarly, I found diversity of genital structure wanting in various other clerid groups, such as in the largely Neotropical Enoclerus, Paleotropical Tenerus, and Australian Phlogistus complexes. How-9 OPITZ: EPIPHLOEINAE Fig. 1. Speciation event in population Aa involving times X to Z (see Phyllogenetic Methods). ever, even in these complexes significant genital differences are observable at species group levels, although the differences are minimal. Frequently, such variations appear among assemblages of species whose specimens are strikingly similar externally. It would be a serious oversight not to consider such characteristics in studies of Cleridae systematics, irrespective of categorical level. The nominal species presented in this study represent my best estimate of what comprise the biological species within Plocamocera.

PHYLOGENETIC METHODS
The phylogenetic relationships proposed in this and subsequent revisions are based primarily on principles pioneered by Willi Hennig (1965Hennig ( , 1966. The dataset selected for predicting evolutionary relationships was analyzed with Farris's computer package, Hennig86 version 1.5 (Farris, 1988). The application of Hennig's principles involves two inherent assumptions. First, we assume that new species evolve when a population diverges genetically into two populations that are no longer mixing genes. This is not an unreasonable assumption, as the gene pool of both the ancestral population and that of the incipient species populations will change in time. Further, the first assumption incorporates the falsifiability factor into the relationship statement because inherent in the assumption is the notion that every species must have a sister species. Once the sister species relationship is proposed it is subject to falsification as new information becomes available. The importance of falsifiability in scientific work has been proposed by Popper (1968) and fully discussed and debated by many authors among which there are the works of Bock (1973), Kitts (1977), and Gaffney (1977, 1978).
The second assumption in Hennigian systematics is that none of the species under study is the immediate ancestor of any of its congeners. This is also a reasonable assumption because once an ancestral gene pool (fig. 1a, population Aa) evolves nuances of genome, or reshuffles and isolates genotype by selection factors (or factors such as genetic drift), two or more genetically incompatible populations may evolve (fig. 1c, population Aa has dichotomized into population A and population a). The ancestral population (Aa), at times X and Y, is no longer genetically identical to population A of time Z in that a portion of the ancestral genome, with potentially unique alleles, has been lost to population Aa during genome Aa divergence, or ''polyvergence''. Therefore, population Aa of time X (the ancestral population) and that of time Y (the ancestral population in the process of, let us say, ''genetic drifting'') are at least potentially not genetically identical to populations A or a of time Z, even if new alleles have not been generated in the threeor-so million years of speciation time. . . an unlikely event. Thus, population Aa is essentially extinct in the purest sense because it has lost a portion of its ancestral genome, that is, the ancestral gene pool (represented by Aa) has lost its original allelic identity in the process, and as a result of the speciation events. Attemps to relate phylogenetically an assemblage of recently evolved taxa is the bane of species-level phylogeny and is perhaps the principle cause of the enigmatic ''phylogenetic bush''. NO. 280 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY When we attempt to reconstruct phylogenies according to Hennig's methods, we implement the principle of synapomorphy. To establish synapomorphies we must first determine whether the various expressions of a character (the specific alleles of a gene) are derived (apomorphic) or primitive (plesiomorphic). All taxa that share a particular apomorphic expression of a character (i.e., a synapomorphy) are joined to form a monophyletic group, with potential difficulties associated with evolutionary reversals being dully noted. The methods for character-state analysis have been amply discussed by various authors, some of which are cited in my earlier work on Perilypus (Ekis, 1977). Therein, I formulated and discussed five criteria by which one can predict whether a particular character state is apomorphic or plesiomorphic. The most useful of these criteria has been the ''criterion of frequency of occurrence'' conceptually aligned with the cannons of ''out-group comparisons'' discussed by Ross (1974: 152), Erwin (1975: 3), and Watrous and Wheeler (1980: 9). My analyses involving assessments of apomorphy or plesiomorphy of character states has included information from all epiphloeine, enopliine, and korynetine species known to me.

REPOSITORIES OF SPECIMENS
The following abbreviations (letter codons) indicate the repositories of specimens from which material was obtained. Letter codons used to abbreviate the name and address of collections were taken from the comprehensive work of Arnett et al. (1993). Implementation of these codons on a worldwide basis would greatly facilitate communication among world systematists and promote consistency in our collection-related language.
I am indebted to collectors, curators, and collection managers for arranging loans of specimens. E-mail addresses of these colleagues are included to facilitate collectionoriented communications.  Kunze. These workers sampled, by various collecting methods, tree branches and boles of the aforementioned tree species and others coexisting with Araucaria. Among the materials they gathered were 115 specimens of Chaetophloeus hispidus, n. sp., obtained from November to February. One hundrednine of these specimens were reared from branches and six were collected by beating. Part of the findings of these investigators was that C. hispidus, although apparently a general predator of a variety of lignicolous beetles, seems to prefer species such as Corthylus rufopilosus, and Xylechinosomus minimus (Scolytidae). The authors further suggest that these clerids are also significant predators of other Xylechinosomus species. By association, that is, by coemergence from the same tree branches, C. hispidus is thought also to feed on Baudonisia villosiventris (Buprestidae), Strangalia dimidiata (Cerambycidae), Helipodus tuberculatus (Curculionidae), and Copidita sp. (Oedemeridae).
Plocamocerans seem diurnal, with large populations congregating on felled tree trunks during mid-morning, late afternoon, and at night. A tropical rainforest with freshly cut tree trunks (figs. [2][3][4][5] is prime collecting habitat for epiphloeines and clerids in general. In Central America, specimens were collected during July and April, some on dry wood of Inga and Guarea. I observed large populations of P. manausensis, new species, of the Amazon Basin, assembled on a recently felled tree leaning against its stump. The insects seemed to congregate on the underside, (i.e., shaded side) of the log, especially during the more sunintensive hours (figs. [3][4][5]. They were also easily captured at night on the surface of the aforementioned felled tree. Wilford J. Hanson collected a variety of plocamocerans in a Malaise trap draped over trunks of recently felled trees ( fig. 2).
On the basis of mouthpart structure and stomodeal contents, I conclude that plocamoceran beetles are carnivorous and are consumers of lignicolous insects, such as larvae of wood borers. The extent to which these beetles blend into their bark surroundings is astonishing. Their elytral variegations blend in perfectly with the thin bark of tree genera such as Manilkara and represent an outstanding example of adaptive coloration. Unfortunately, such adaptations tend to homogenize the outward appearance among congeners, which makes species-level identifications particularly difficult. Also noteworthy is the manner by which these clerids escape capture. Their strategy most often involves an initial scurry movement, then catapult to full flight. Their jumping abilities coincide well with the extraordinary development of their metafemur and metacoxa. Hespenheide (1973) associated such locomotory behavior with fly mimicry.

INTEGUMENTARY SENSORY ORGANS
SENSILLA ON THE ANTENNAE, PRONOTUM, AND ELYTRA Antennae: The plocamoceran antenna comprises 10 antennomeres ( fig. 86); the scape is unusually long and curvate, the pedicel large and spheroid, and the flagellum is greatly expanded and highly setose distally. The initial four flagellar antennomeres are short and narrow, whereas the terminal three constitute a substantially expanded club that forms the organs most conspicuous portion. To date, four types of sensilla have been found on the plocamoceran antennae: one type of sensilla trichodea, two types of sensilla chaetica, and one type of sensilla basiconica.
striking feature of the plocamoceran antennae ( fig. 10). Sensilla trichodea are most abundant on the club antennomeres (no. 13 in fig. 11) and take the form of long, longitudinally grooved, hairlike organs set on the antennomeres in well-formed sockets. Stein-brecht (1970: 124) suggested that the long sensilla trichodea of the male silk moth are instrumental in the detection of female sex pheromone. Shorter, but anatomically similar, sensilla trichodea have been studied in various other families of beetles in which these sensory organs were interpreted as mechanoreceptors and chemoreceptors (Merivee et al. 1998: 312, and references therein).
Elsewhere (see Natural History), I discussed my field observations involving specimens of P. manausensis, new species, on a recently felled tree infested with bark beetles within a tropical rainforest. On the basis of this predatory-prey association, and in conjunction with findings of antennal microsculpture and published biochemical information, I suggest that the filamentous sensilla trichodea of the plocamoceran antenna, which greatly increase the surface area of that organ, have an olfaction function and serve to detect volatile tree compounds and/ or aggregate pheromones released by bark beetles. Borden and Wood (1966: 259) suggested that in the bark beetle Ips confusus (LeConte), sensilla trichodea are predominantly involved in afference of sex pheromones. Through this afferent mechanism, plocamoceran predators can readily enhance their ability to quickly find concentrations of prey species in an environment where wood decomposition is accelerated by tropical climatic factors and where availability of suitable prey species is temporally short. Kaissling (1971: 357) stressed the importance of antennal surface in relation to that organ's afferent properties. Furthermore, the plumose-like construction of the plocamoceran antenna is somewhat reminiscent of the antennal development of the Bombyx moth, in which it has been established that flagellar branches capture and absorb odor molecules of pheromones . Schneider (1969. Schneider ( : 1034 suggested that highly plumose antennae, as those found in the silkworm moth Bombix mori, are effective molecular sieves. Several North American studies have shown that checkered beetle predation on bark beetles involves sensitivities to tree compounds and/or barkbeetle pheromones. For example, Pitman and Vite (1971: 404) and Whittaker and Foemy (1971: 759) demonstrated that Enoclerus lecontei (Say) respond to conifer terpenes and aggregate hormones, respectively. Moreover, Harwood and Rudinsky (1966: 297) established that Enoclerus sphegeus (Fabricius) and Thanasimus undatulus (Say) were attracted to oleoresins collected from living trees of Douglas fir (Pseudotsuga menziesii (Mirb.)), ponderosa pine (Pinus ponderosa), and grand fir (Abies grandis). Thanasimus dubius (Fabricius) is known to respond to pheromones produced by bark beetles (Ips spp.) and to tree volatiles (Vite and Williamson, 1970: 238;Dixon and Payne, 1979: 180, 1980: 381, Billings and Cameron, 1984: 1545Mizell et al., 1984: 180;Billings, 1986: 488).
Type I sensilla chaetica (no. 12 in fig. 11) have been found on antennomere nine and type II sensilla chaetica (no. 2 in fig. 12) on antennomere ten. Type I emerge from a wellformed socket, varies in length, is suberect and fluted, and tapers to a fine point. In the leaf beetle Psylliodes chrysocephala Linnaeus these kinds of antennal sensory organs (identified as sensilla trichodea I) (Bartlet et al., 1999: 293) are known to be enervated by one sensory neuron attached to the base of the bristle shaft, an indication of mechanosensory function according to Zacharuk (1985: 26). Type II sensilla chaetica have heretofore been established to occur only on the terminal antennomere of the antennal club ( fig. 12). They stem from a cuticular plate, vary in length and undulations, are externally smooth, and are relatively blunt at the apex. They most closely approximate the sensilla trichodea type II of P. chrysocephala L. (loc. cit.) and others reported in various species of weevils (Bartlet et al., 1999: 298). Mustaparta (1973Mustaparta ( : 570, 1975 discussed similar antennal sensilla in the weevil Hylobius abietis and reported their electrophysiologically responsiveness to odors. The plocamoceran sensilla basiconica (no. 14 in fig. 11) are short, externally smooth and thin-walled, straight or curved, peglike sensilla set in an elevated base. Elaterid, scolytid, curculionid, and coccinelid beetles are known to have similar antennal sensilla (Marivee et al., 1998: 312;Bartlet et al., 1999: 298) responsive to odors in electrophysiological experiments (Mustaparta, 1975: 283).
The remaining vestiture of the pronotal surface involves three types of sensilla trichodea (I, II, III). Pronotal sensilla trichodea types I (no. 5 in fig. 14 and no. 2 in fig. 24) and II (no. 6,fig. 14) are similar in shape; both are recumbent, short, fluted, linear, or slightly curved chaeta that narrow to a fine point. However, pronotal sensilla trichodea II differ as they arise from deeper punctiform depressions (see figs. 22 and 23). Further, some pronotal sensilla trichodea II are associated with a minute tubular peglike process (no. 7 in fig. 15).
Pronotal sensilla trichodea III (herein referred to as trichosomes; no. 5 in fig. 7) are very sparsely distributed on the pronotal disc, being most clearly evident along the anterior and posterior margins. Trichosomes are very robust, slightly curvate vertical bristles set in large punctiform depressions that are more abundantly found on the elytral disc and elytral epipleural (no. 1 in fig. 16) and sutural margins. An interesting additional feature of the pronotal surface is the presence of integumental pores (figs. 22, 23, and no. . Distal extremity of metatibia (1, croun of apical setae; 2, metatibial spur). 21. Pronotal lateral trichobothrium (3, trabeculae fig. 24) sparsely distributed among the discal sensilla.
Heretofore, trichobothria have not been recorded in the Cleridae, although several workers have referred to the pronotal trichobothria as a ''pair of long sensory setae'' (Crowson, 1964: 306), ''thorax with a pair of discal and a pair of internal sensory setae'' (Barr, 1950: 488), ''Halsschild mit je einem Paar disk aler und seitlicher Sinnesborsten.'' (Winkler, 1961: 30). Prothoracic trichobothria are variously developed within Epiphloeinae, reaching a high development in Plocamocera. In this genus the thrichobothrial gross structures resembles those that Schuh (1975: 5) classified as structural type four in which the trich is deeply sunk into the integument. In plocamocerans the base of the pronotal trich shows a domelike scalloped structure at its core ( fig. 9). It remains to be determined whether the variation of epiphloeine trichobothrial development and the distribution of other long integumental sensilla represent adaptation to a greater efficacy of afference. Schaefer (1975: 239) noted that the trich component of the trichobothrium was particularly long among ground-living heteropterans but was short among heteropterans that live on plants. Gorner and Andrews (1969: 301) discovered that, ''long trichobothria are more sensitive to airborne movements of low frequency (65 cycles/sec.) than shorter ones''.
The function of the pronotal trichobothria in clerids remains uncertain. However, their gross anatomical construction is very similar to the filiform setae described in the caterpillar of Barathra brassicae Linnaeus (Markl and Tautz, 1975: 82) in which the trichs serve to detect vibrations generated by oncoming predatory wasps or parasitic flies. It is conceivable that the prothoracic trichobothria of plocamocerans, and perhaps those of other epiphloeines, serve to warn these clerid predators of other predatory species such as wasps or treetrunk-foraging lizards. Camhi (1980: 147) discovered that puffs of air made by a toad's lunge is perhaps the primary means through which the cockroach detects an attacking toad.
Plocamocerans display slightly different trichobothria on the epipleural margin of the elytra (no. 2 in fig. 16). Possibly these hair sensilla also serve as an antipredatory device. The development of elytral trichobothria, and apparently their numbers, may vary among the plocamoceran species. A complete survey of trichobothrial structure, and of the occurrence and distribution of extraordinary integumental sensilla, involving genera of Epiphloeinae is in progress. It is hoped that results from such a study will generate additional synapomorphies for a more thorough resolution of Epiphloeinae phylogenetics.

DIAGNOSIS OF CHAETOPHLOEUS, NEW GENUS
Specimens of this monotypic genus are most conveniently identified by the profuse presence of stout bristles (chaetosomes) on the dorsum of their forebody and elytra (figs. 26, 37). Other outstanding characteristics include the substantial length of the antennal scape, very globose pedicel, and lengthened antennal sensilla trichodea. These trichodea are slightly longer than the width of the antennomeres of the antennal club. The antennal sensilla trichodea of the antennal club are considerably longer in the members of Plocamocera, than they are in the members of this genus, the presumed sister taxon of Chaetophloeus. Venter and legs copiously vested with setae that are short and decumbent or long and suberect; dorsum copiously vested with stout, bristlelike setae. Head: Cranium subrugosely punctated; eyes prominently bulging, finely faceted, deeply incised along frontal margin ( fig. 27), incision half width of eyes when eye viewed from front; antenna (fig. 29) inserted at lower angle of eye incision, comprised of 10 antennomeres, loosely clubbed, vested with few lengthy setae, setae not longer than length of club articles; scape as long as combined length of funicular antennomeres, pedicel particularly large, subglobose, antennomeres four and six slightly expanded, antennomeres five and seven cylindric, club-antennomeres eight and nine subtrigonal, equal in length, tenth antennomere ovoid, only slightly longer than antennomeres eight and nine; labrum deeply incised ( Thorax: Pronotum moderately transverse, anterior margin sinuous, prominently projecting medially, posterior margin broadly sinuous, subapical depression not very prominent, pronotal disc slightly depressed at sides where discal and paralateral trichobothria are prominent; disc with pair of paralateral swellings; epimeral prolongations only feebly extended mesad; procoxal cavities open; elytra distinctly tapered, surface subrugose, punctations small and shallow; epipleural margin feebly explanate; mesoscutellum quadrate-transverse (fig. 31); protibia with stout spines, protibial spur absent, protarsus with three pulvilli; mesotibia with one spur, mesotarsus with three pulvilli; metatibia with one spur, metatarsus with one pulvillus; tibial spurs not particularly elongated; metabasitarsus much longer than metatarsomere two; tarsal claws with small basal dens. Abdomen: Six visible sterna; pygidium broad-scutiform. Male Genitalia: Ae- NO. 280 XI.1999, Ex. Araucaria angustifolia, leg.: Roland Mecke (MCNZ). (Specimen card mounted, sex label affixed to paper card, white, machine printed; locality label, white, cursive, MCNZ repository label white, machine printed; number label (R 2488), white, machine printed; collection label with number (215.508), white, outlined, machine printed; holotype label red, machine printed.) The specimens identified as Plocamocera in Mecke et al. (2001: 113) involve members of this species.

DESCRIPTION OF CHAETOPHLOEUS
PARATYPES: Fifty-nine specimens. Fiftyeight specimens from the same locality as the holotype (MCNZ, 3 DIAGNOSIS: Members of this monotypic genus can be conveniently distinguished from other epiphloeines by the profuse distribution of bristlelike setae on the head pronotum, and elytra. Further, from its sister genus Plocamocera, Chaetophloeus specimens are conveniently distinguished by the considerably shorter filamentous sensilla trichodea on the antenna. In Plocamocera the gula is crescentic, whereas in Chaetophloeus the gula has a more trigonal shape (compare figs. 28 and 126).
DESCRIPTION: Body length, width, form, and integumental color as in generic description. In addition, lower frons castaneous, upper frons and epicranium piceous; pronotum narrowly castaneous along anterior and lower lateral margins; elytron light-castaneous at humerus and sporadically vested with short silvery decumbent setae that are most prominent as three feebly formed setal pencils at elytral basal third, silvery setae also form oblique broad line at elytral apical half. Head, thorax, and abdomen as in generic description. In addition, frons distinctly concave, elytral disc depressed behind humerus, elytral punctations prominent throughout disc and not arranged in longitudinal rows, and fifth abdominal sternum narrowed distally in females, evenly arcuate in males; aedeagus as in figure 35.
VARIATION: The shape of the fifth visible abdominal sternum is sexually dimorphic. In females it is narrowed distally, whereas in males it is not narrowed distally.
NATURAL HISTORY: Mecke et al. (2001) studied the insect fauna associated with Araucaria trees. Among the insects that were found to frequent these trees were specimens of C. hispidus, n. sp., which readily emerged from Araucaria angustifolia (Bert) O. Kuntze. The available specimens were collected during January, February, June, October, September, November, and December.
DISTRIBUTION (map 12): These beetles are known only from southern Brazil.
ETYMOLOGY: The specific epithet, hispidus, is a Latin adjectival meaning ''bristly''. I refer to the bristlelike setae on the dorsum of these beetles.

DIAGNOSIS OF PLOCAMOCERA SPINOLA
The most outstanding characteristic of the members of this genus is the long filiform hairs on the antenna ( fig. 86). As a group, the elytra of the members of Plocamocera tend to be variegated in color, ranging from stramineous to castaneous with many species characterized by a distinct flavotestaceous humeral macula that divides posteriorly. Moreover, I found the presence of setal aggregates on the elytra and number of conspicuous trichobothria on the epipleural mar- NO. 280 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY gin diagnostic for some species. Unfortunately, the elytra of many of the older specimens were severely depiliated, which rendered their identification difficult. When depiliated specimens do not show the elytral trichobothria ( fig. 55), one can determine the presence of some of these sensory organs by the more conspicuous black indentations with which these filiform setae are associated. Males with depiliated elytra may be diagnosed on the basis of the configuration of the last three antennomeres, shape of the pydigium, and characteristics of the aedeagus. Severely depiliated females are particularly difficult to assign to species, but in most cases the shape of the antennomeres of the antennal club leads one to the correct identification. At present, I recognize 35 species of the genus Plocamocera.
In addition to the aforementioned characteristics, members of Plocamocera are readily identified by other characteristics such as: antennal scape very long, as long as combined length of funicular antennomeres; antennal pedicel very globose, much larger than antennomere three; pronotum boldly transverse ( fig. 7), and with its anterior margin extensively projected at middle; pronotal trichobothria ( fig. 9) well developed; elytra oblongovate, about three times longer than wide, epipleural margin arcuate when viewed from above, adorned with rows of stout chaetosomes, and vested with several trichobothria (fig. 53) that tend to diminish in length from posterior of elytron to anterior of elytron; elytra with or without maculae and with dense clusters of light and dark setae; metacoxa and metafemora particularly robust; metatarsal pulvillus present on third tarsomere; color of abdomen sex dimorphic, males piceous, females flavotestaceous in basal half piceous in remainder; spermathecal gland attached to subapex of spermathecal capsule; and males with one pair of accessory glands.  Kuwert, 1893: 492. Lohde, 1900: 88. Schenkling, 1903: 86, 88. Gahan, 1910: 73. Chapin, 1927: 5. Blackwelder, 1945: 388. Corporaal, 1942: 255. Winkler, 1961: 59. Opitz, 1997 DESCRIPTION: Size: Length 4.0-8.0 mm; width 1.5-2.8 mm. Form: Elongate, elytra somewhat ovate, about three times longer than wide; pronotum (fig. 7) conspicuously transverse; epipleural margin feebly or strongly arcuate. Integument: Head, thorax, and abdomen concolorous or bicolorous, if bicolorous stramineous or castaneous, frons and vertex often infuscated; antenna bicolorous, scape flavotestaceous, remainder piceous; pronotum usually vested with pale setae at sides, disc usually infuscated; elytral surface usually variegated, rarely concolorous, when bicolorous stramineous or castaneous, or mixture of both. Vestiture: Integument copiously vested with decumbent and declinate setae; antenna (figs. 6, 40) with very long sensilla trichodea, discal and paralateral trichobothrial setae of pronotum (figs. 7, 9) particularly well developed, sides of pronotal disc matted with light setae whose apices extend toward middle; elytral disc abundantly vested with stout bristles, bristles are particularly notable along sutural and epipleural margins, disc also vested with pale or dark patches of setae, patches often transverse, sometimes oblique or angular in shape, epipleural margin minutely serrulate, adorned with row of chaetosomes and with three to eight trichobothria of various degree of development (    The members of this species group are foremost diminutive, most being about 3 mm in length; the cranium is usually black, the only exception being specimens of P. salasis, n. sp., in which the cranial margins are castaneous; and the first two antennomeres of the antennal club are usually ovoid ( fig. 149), being particularly slender in P. manausensis, n. sp.; only in P. salasis, n. sp., does the eighth antennomere approach subquadrate. In aggregate, this group of species ranges from Guatemala to the Amazon Basin.

sesquipedalis group
The basal segment of the metatarsus is extraordinarily long in these beetles ( fig. 109); the tripartite humeral macula is lacking; and the aedeagus ( fig. 120) is particularly long. In the more northern specimens of P. sesquipedalis, n. sp., and in all specimens of P. lucis, n.sp., the relatively light color of the elytra accentuates the abundance and robustness of dark elytral bristles. The combined range of the four species of this group extends from Central Costa Rica, across Guyana, south through Ecuador and Peru, and to the forests of Matto Grosso, in Brazil.

castanea group
This group contains one species whose members have a stricking development of the antennal club ( fig. 144); the elytral surface is somewhat undulated; and the sutural margin is bordered by particularly robust sensilla trichodea. The species ranges from the Napo rainforests of Ecuador to the more southern Brazilian lowlands of Goias.
confrater group As a group, the members of these species show minimal interspecific variation of external structures. One must examine and correlate variations of the aedeagus with those of the antennal club to fully appreciate the taxonomic significance of subtle integumental differences. In confrater group specimens the eighth antennomere is subquadrate ( This group of South American species has been recorded from Guyana, Ecuador, Peru, Bolivia, and Brazil. NO. 280 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY coactilis group In these plocamocerans the basal antennomere of the antennal club is most often narrow, rarely broad-oval ( fig. 153); the pronotum is predominantly piceous; the tripartite humeral macula is well developed; and the female pygidium is broad-scutiform. The composite range of the species group extends from Ecuador to Brazil.

sericella group
The narrow-ovoid shape of antennomere eight, absence of the tripartite humeral macula, and presence of matted aggregates of white setae on the elytral disc characterize the members of this species group. Included in this species group is P. sericella Spinola, the most widely distributed species of Plocamocera; specimens are recorded from Mexico to Brazil.

DISCUSSION OF SPECIES KEY COUPLETS
Members of closely related species of this genus are very similar externally, yet subtle differences of the antenna and elytra usually corroborate the more obvious differences of the male genitalia. In general, it can be stated that the abovementioned subtle differences can best be ascertained with high magnification. I have used magnification in the 750-1000 range. When dealing with specimens of the confrater and sericella groups, the antenna of all available male specimens should be removed from the cranium and examined submerged in a fluid (such as water or glycerin) at a magnification of at least 750ϫ. This level of magnification will provide clear resolution of the precise configurations of the antennomeres.
Interpretations of body size may occasionally become an obstacle when using this key. All known members of the manausensis group are uniformly diminutive, whereas specimens of other species groups are greater than 4 mm and, therefore, not diminutive as defined herein; all of these specimens will correctly filter through the first key couplet. However, small specimens of P. sericella Spinola approach the 4 mm body size of P. manausensis, n. sp., beetles. When confronting such specimens one must check the color of the cranium, which is black in P. manausensis, but predominantly castaneous in P. sericella. Also, one may precisely measure the body length of the beetles before one can proceed with confidence beyond the initial key couplet.
The next potential source of difficulty involves interpretation of ovality of the eighth antennomere, which is particularly relevant to couplet nine in which one has to distinguish between P. sesquipedalis, n. sp. and P. similis n. sp. The degree of ovality is more pronounced in P. similis n. sp., but one must observe this characteristic at a magnification approaching 1000ϫ.
In couplet 10 one has to distinguish between a subquadrate and ovoid shape of the eighth antennomere. In most cases the two shapes are easily separated, but in P. ambra, n. sp., the eighth antennomere is slightly angular and possibly interpreted as subquadrate. However, in such specimens the elytra does not show a tripartite posthumeral macula, as is also the case in all other specimens of the sericella species group.
Lastly, the degree of ''narrowing'' of the distal extremity of antennomere eight is used, with some difficulty, to separate some specimens among the coactilis group (couplet 24 and 24Ј). The narrowing may be quite notable or deceptively short. A magnification of at least 90ϫ must be used to recognize some of the more subtle differences among antennal characteristics.
VARIATION: Except for the slight tone variation of elytral color, the available specimens are quite homogenous.
NATURAL HISTORY: These beetles were collected from the environs of Manaus, Brazil, in January, on bark of Manilkara, during day and night.
DISTRIBUTION (map 6): Known only form the type locality.
ETYMOLOGY: The name manausensis constitutes a noun in apposition and refers to the type locality. PARATYPES: Twenty-four specimens from the same locality as the holotype (BMNH 1; FMNH 2; USNM 16; WOPC, 5).

Plocamocera iota, new species
DIAGNOSIS: The single tibial spine on the protibia serves to identify members of this species. The males are also identified by the configuration of the phallobasic apodeme of the tegmen. Unlike in specimens of P. manausensis and P. minima, in which the phallobasic apodeme is slender and elongated, in P. aliguantula specimens the apodeme is short and obtuse (figs. 113-115). VARIATION: The dark fascies of the dorsum is more pronounced in some specimens.
NATURAL HISTORY: Twenty-four specimens were collected on dry wood (an trockenem holtz, Inga) from species of Inga and Guarea, during July. One specimen was collected during April.
DISTRIBUTION (map 5): Known only from Central Costa Rica. ETYMOLOGY: From aliguantulus, a Latin adjective meaning ''little''. I refer to the small size of these beetles. DIAGNOSIS: The most reliable characteristic that distinguishes the members of this species from members of the closely related P. aliguantula and P. manausensis is the shape of the tegmen of the male genitalia. In P. minima the aedeagus ( fig. 114) is proportionally small and the phallobasic apodeme is about as long as the length of the phallobase. DESCRIPTION: Size: Length 3.0-3.6 mm; width 1.0-1.2 mm. Integument: Cranium black; pronotum predominantly piceous, periphery castaneous; elytra variegated, flavous humeral macula well developed, post medial flavous macula transverse; flavous portions of disc densely matted with white setae, piceous portion vested with dark setae; legs flavous. Head: Antenna as in figure 44. Thorax: Pronotal anterior margin moderately projected anteriorly, discal swellings shallow; elytra without distinct trigonal post-humeral macula ( fig. 60); elytral epipleural margin with four conspicuous trichobothria; protibial anterior margin with four spines. Abdomen: Male pygidium broad-scutiform, subtruncate, female pygidium broad-scutiform; aedeagus as in figure 114.

Plocamocera minima, new species
VARIATION: Except for tone variation involving elytral color the available specimens do not vary significantly.
NATURAL HISTORY: These beetles were collected from the type locality during April through July, form species of Inga.
DISTRIBUTION (map 5): Plocamocera minima specimens are known only from the type locality.

Plocamocera prolixa, new species
VARIATION: The infuscation on the frons, vertex, legs, and elytral disc vary in intensity.
NATURAL HISTORY: The available specimens were collected during October, November, and December; the series from Ikuribisi on Eschwellera sagotianum. The two specimens from Rondonia, Brazil were collected at an altitude ranging from 160-350 m.
DISTRIBUTION (map 5): This South American species is known from Guyana, French Guiana, Ecuador, Peru, and Brazil.
ETYMOLOGY: The specific epithet stems from the Latin sesquipes (having a measure of one and a half) and the Latin suffix -alis (condition of). I refer to the extraordinary length of the metafemur. (Specimen point mounted, antenna and sex label affixed to paper point, white, machine printed; support card, white; locality label, white; ma-chine printed; QCAZ repository label, white, machine printed; holotype label, red machine printed; plastic vial with abdomen and aedeagus.) PARATYPES: One specimen from the same locality as the holotype (WOPC, 1).

Plocamocera lucis, new species
DIAGNOSIS: These beetles are very similar to those of P. sesquipedalis, new species. The males of these two species are most reliably distinguished by the shape of the apex of the phallus which is in P. similis n. sp. and lobate in P. sesquipedalis, n. sp. Females, and males, have the eighth antennomere subrectangulate rather than subovate as is the case in most of the P. sesquipedalis specimens. Females of P. similis, from the more southern latitudes, may be undistinguishable from females of P. sesquipedalis.
NATURAL HISTORY: Specimens were collected from Bolivia during August at 400 m, from Ecuador during February (450 m) and April, and from Brazil in November. In Ecuador, at  DISTRIBUTION (map 1): This species is known from Bolivia and the Amazonian region of Ecuador, Peru, and Brazil. NO. 280 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY ETYMOLOGY: Latin, the adjectival castanea (chestnut). The trivial name refers to the color of the dorsal facies of these beetles.
NATURAL HISTORY: The available specimens were collected from Central Brazil, in October.
DISTRIBUTION (map 5): This species is known only from central Brazil.
ETYMOLOGY: Latin, the noun pupula (pupil of the eye). I refer to the piceous macula behind the basal tumescence of the elytra. (Specimen point mounted, antenna and machine printed sex label affixed to paper point, white; locality label, white, hand printed; AMNH repository label, white machine printed; holotype label, red, machine printed; plastic vial with abdomen.) PARATYPES: None. DIAGNOSIS: Along with P. pupula, P. quadrula is the only other known species of the confrater group whose specimens do not have the distal narrowing of antennomere eight. Specimens of P. quadrula do not have the distal margin of antennomere eight feebly concave ( fig. 146), which is the case in P. pupula specimens ( fig. 145). Also, in P. pupula specimens the tenth antennomere is nearly twice as long as antennomere nine.

Plocamocera quadrula, new species
DESCRIPTION: Size: Length 5.7 mm; width 2.0 mm. Integument: Predominantly castaneous, frons feebly infuscated; pronotum predominantly castaneous, disc infuscated; elytra variegated, with flavotestaceous humeral macula oblique, with three irregular aggregates of white setae, golden setae abundant along sutural margin and in piceous regions of disc; legs flavotestaceous, metafemora faintly infuscated on inner distal surface. Head: Antennal club as in figure 146. Thorax: Anterior margin of pronotum boldly projected at middle; discal swellings prominent; elytral epipleural margin with five conspicuous trichobothria; protibial anterior margin with five spines, the second and third PARATYPES: One specimen examined. DIAGNOSIS: These beetles are most closely allied to the members of P. pupula, n. sp. from which they may be distinguished by the shape of the eighth antennomere. In P. jayhawkalis, n.sp., the outer angle of the anterodistal margin of the eighth antennomere is blunt ( fig. 212). In P. pupula, n. sp., the outer angle of the anterodistal margin of the eighth antennomere is distinctly acute ( fig. 195). Also, in P. jayhawkalis, n. sp., the ninth antennomere is abruptly constricted at its posterior limits, whereas in P. pupula, n. sp., the apical constriction is gradual.
DESCRIPTION: Size: Length 7.0 mm; width 3.0 mm. Integument: Predominantly castaneous; anterior disc of pro-mesotibiae infuscated, posterior disc metatibiae infuscated near femoral apex; elytra variegated, with two dark transverse bands near apex. Head: Antennal club as in figure 212. Thorax: Pronotal anterior margin broadly projected at middle; elytral epipleural margin with five conspicuous trichobothria; protibial anterior margin with five spines. Abdomen: The abdomen of the holotype specimen is missing.
VARIATION: Not studied.
NATURAL HISTORY: The holotype specimen was collected from Surinam, during June, at 25 m, by fogging, a splintered tree trunk with pyrethrum.
DISTRIBUTION (map 6): This species is known only from Surinam.
ETYMOLOGY: The specific epithet is a compound noun in apposition. It stems from the mascot name of Kansas University (''jayhawks'') and the Latin possessive suffix-alis. The author, a Kansas State ''wildcat'', wishes to extend his appreciation, and a tribute, to Kansas University for providing Kansas State University with many years of collegial athletic competition.
VARIATION: Not observed. NATURAL HISTORY: The available specimens were collected from southern Venezuela during February, at 140 m.
ETYMOLOGY: The trivial name constitutes a noun in apposition and refers to the type locality.  tions, 1921-1922. (Specimens point mounted, antenna and sex label affixed to paper point, white, machine printed; support card, white; locality label, white, machine printed, expedition label, white, machine printed; USNM repository label, white, machine printed; holotype label, red, machine and hand printed.) PARATYPES: None. DIAGNOSIS: Specimens of this species may be distinguished from other members of the confrater group by the following combination of characteristics: interstitial spaces of elytral disc shiny, eighth antennomere only feebly narrowed distally, and tenth antennomere only slightly longer than antennomere nine.
VARIATION: One specimen examined. NATURAL HISTORY: The only available specimen was collected from Bolivia, in December.
DISTRIBUTION (map 7): Known only from the type locality.
ETYMOLOGY: The trivial name aura (glow) is a Latin adjective. I relate the name to the bright reflection from the interstitial spaces of the elytra. (Specimen point mounted, sex label affixed to paper point, white, cursive; support card, white; locality label, white, machine printed; FSCA repository label, white, machine printed, holotype label, red, machine printed; plastic vial with abdomen and aedeagus.) PARATYPES: Six specimens from the same locality as the holotype (FSCA, 2; WOPC, 4).

Plocamocera buenavista, new species
DIAGNOSIS: The members of this species may easily be confused with the superficially similar specimens of P. aura n. sp., also from Bolivia. However, the members of this species are distiguished from those of P. aura, n. sp. by the gritty property of the elytral interstitial spaces and by the subtle, but consistent, differences of antennomeres eight and nine (compare figs. 217-218). The anterodistal margin of the eighth antennomere is much less concave than it is in the holotype of P. aura n. sp. The distal, or apical, portion of antennomere nine is gradually less narrowed in P. buenavista, n. sp., than it is in specimens of P. aura, n. sp. (compare figs. 217-218).
VARIATION: The elytral disc is considerably more pale than it is in the other specimens. Otherwise, the available specimens are quite homogeneous.
NATURAL HISTORY: The available specimens were collected during November. In a tropical transition forest at 405 m.

DISTRIBUTION (map 8):
This species is known only from the type locality.
ETYMOLOGY: The specific epithet is a noun in apposition. It is used as a tribute to the fine Bolivian people of Buena Vista. Figures 162,173,195;map 7 HOLOTYPE: Male. Brazil: Amazonas: Manaus, 1 km W. Taruma Falls, 19-I-1981, Weston Opitz (MZSP). (Specimen point mounted; antenna, pygidium, and machine printed sex label affixed to paper point; support card, white; locality label, white, machine printed; natural history label, white, hand printed; MZSP repository label, white, machine printed; holotype label, red, machine printed; plastic vial with aedeagus.)
DIAGNOSIS: This species belongs to the subgroup of P. confrater in which specimens have the pronotum predominantly castaneous and only faintly infuscated. Within this subgroup, P. taruma specimens most closely approximate those of P. confrater, from which they are distinguished by the proportionally longer 10th antennal article (compare figs. 152, 162).
VARIATION: The intensity of the cranial and pronotal infuscation varies.
NATURAL HISTORY: The three specimens from Taruma Falls were collected in January on freshly felled trunks of Manilkara, a genus of hardwood prominent in riverside habitats of the Amazonian Basin.
DISTRIBUTION (map 7): Known only from the Amazonian environs near Manaus and Matto Grosso highlands of Brazil.
VARIATION: One specimen examined. NATURAL HISTORY: The available specimen was collected during April with a Malaise trap at 150 m. DISTRIBUTION (map 9): This species is known only from the type locality.
ETYMOLOGY: The specific epithet procera is a Latin adjectival and means slender. I refer to the extended length of the aedeagus when compared to the aedeagus of males of P. confrater Kuwert. COACTILIS GROUP Plocamocera coactilis, new species 12,46,64,65,95,104,116,136,163,198 (Specimen point mounted, antenna and sex label affixed to paper point, white, machine printed; support card, white; locality label, white, machine printed; MZSP repository label; holotype label, red machine printed.) PARATYPES: None. DIAGNOSIS: Within the coactilis group specimens of P. aspera most closely resemble those of P. paris. However, only in P. aspera specimens is the cranium distinctly rugose and antennomere eight more ovoid than angulate; also, antennomere nine is more narrowly prolonged distally (compare figs. 153, 164).
VARIATION: Not observed. NATURAL HISTORY: The available specimen was collected from the type locality during December.
DISTRIBUTION (map 7): Known only from the type locality.
ETYMOLOGY: The specific epithet is a compound name formulated from the Latin prefix -by (two) and the Latin noun spina (thorn). I refer to the two spines on the protibia. HOLOTYPE: Female. Ecuador: Napo: Yasuni Research Station, 19-30 Oct. 1998, 250 m, W. J. Hanson (QCAZ). (Specimen point mounted, sex label affixed to paper point, white, machine printed; locality label, white, machine printed; QCAZ repository label, white, machine printed; holotype label, red, machine printed.) PARATYPES: None. DIAGNOSIS: Within the coactilis species group, these beetles may be conveniently identified by the combination of antenno-mere eight ovate and protibial anterior margin with two spines.
VARIATION: Not observed. NATURAL HISTORY: The only available specimen was collected from the type-locality, during October, at 250 m.
DISTRIBUTION (map 7): Known only from the type locality.
ETYMOLOGY: The trivial name represents a noun in apposition and refers to Professor Giovanni Onore, an Ecuadorian Entomologist very devoted to the advancement of Ecuadorian insect systematics and to the growth of the insect collection at the Catholic Pontifical University of Quito, in Ecuador. Figures 160,200;map 6 HOLOTYPE: Female. Brazil: Santa Catarina: Nova Teutonia, 9-I-1957 (IZAN). (Specimen card mounted, antenna mounted on mount card; abdomen mounted on paper point; locality label, light green, machine and hand printed; IZAV repository label, white, machine printed; holotype label, red, machine printed.) PARATYPES: One specimen from the same locality as the holotype (WOPC, 1).
VARIATION: The two available specimens do not vary appreciably.
NATURAL HISTORY: The type specimens were collected from southern Brazil during January.
DISTRIBUTION (map 6): Known only from southern Brazil.

Plocamocera auratilis, new species
DESCRIPTION: Size: Length 4.5-5.0 mm; width 1.5-2.0 mm. Integument: Predominantly light-castaneous, pronotal disc infuscated; mesoscutellum piceous; elytral disc infuscated at sides along epipleural margin and at apex, profusely vested with gold-yellow setae which are particularly prominent along sutural margin, with four patches of silvery setae contiguous with sutural margin ( ETYMOLOGY: From the Latin adjective auratilis (gold colored). I refer to the golden sheen characteristic of the elytral surface so apparent in these beetles. Figures 47,61,82,96,108,117,133,135,156,211;map 4 Plocamocera There are no appreciable differences between the type specimen of this junior synonym and the type specimen of the senior synonym . Corporaal, 1950: 255. DIAGNOSIS: The pronotal disc is piceous and the elytra do not have conspicuous macula, but they do have three irregularly shaped setal fascia that range from white to grey. The elytral humeral macula are more pronounced in South American specimens. These clerids are readily distinguishable from superficially similar specimens of P. sericellopsis, by the presence of four spines on the protibia. In P. cericellopsis the protibia show three spines.
VARIATION: The dark areas of the elytra become more pronounced in specimens from South America, especially those from French Guiana and Brazil. In some of these beetles the legs are infuscated and the elytra are marked with piceous maculae.
NATURAL HISTORY: These beetles were collected throughout the year from altitudes that range from 150 to 1300 m. Specimens  PARATYPES: None. DIAGNOSIS: Specimens of this species resemble very closely members of P. sericella. However, P. sericellopsis specimens differ by having only three spines on the protibia, the male pygidium is emarginate, and the aedeagus is shorter and more stout.
DIAGNOSIS: This species is closely related to P. auratilis, n. sp. and P. ambra, n. sp. However, in P. bolivari specimens the body is shorter and more squat. Further, P. bolivari specimens differ from P. auratilis specimens by having the pygidium feebly emarginated, and the integument is generally more piceous. In P. auratilis beetles the pygidium is truncate and the integument is predominantly light castaneous. From P. ambra specimens, P. bolivari specimens differ by having a considerably narrower phallobasic apodeme.
VARIATION: The piceous regions of the elytra vary in intensity.
NATURAL HISTORY: The available specimens were collected from the eastern Andes of Venezuela, at 500 m. in a Malaise trap.
DISTRIBUTION (map 6): Known only from the type locality.
ETYMOLOGY: The specific epithet is a surname patronym and is used to honor General Simon Bolivar, a great liberator of the fine people of South America.  . 155) will identify the members of this species within the sericella species group.
VARIATION: The intensity of the infuscation on the frons varies.
NATURAL HISTORY: The specimen from Alajuela was collected in November; those from Puntarenas were collected at 300 m.
ETYMOLOGY: The specific epithet is a Latin noun that means amber. I refer to the amberlike luster emitted by the elytra. ETYMOLOGY: The specific epithet constitutes a noun in apposition and refers to the type locality.

EVOLUTIONARY CONSIDERATIONS
The available evidence suggests that Chaetophloeus and Plocamocera are more closely related to each other than either one is to any other genus taxon within Epiphloeinae. Moreover, although the monophyly of Plocamocera is readily hypothesized on the basis of uniquely derived characteristics (apomorphies), I found it difficult to resolve all evolutionary pathways among its species groups and species. The task was impossible among members of the confrater, coactilis, and sericella groups in which only a few structural variations could be credibly evaluated as being apomorphic or plesiomorphic.
There are various other groups within Cleridae in which integumental variations are minimal. Indeed, finding structural diversity among taxa is no guarantee that such diversity will be useful towards credible, refutable hypotheses of evolution. I have found this a problem among a substantial variety of generic and suprageneric taxa within the family; for example, in such genera as Enoclerus, Cymatodera, Priocera, Phlogistus, and Eleale, and among the other genera of Epiphloeinae. Species of such groups may present observable differences, but in most cases few of such differences prove useful for predictions of relationships. Systematic progress within these difficult groups will have to await new information from nonintegumental characteristics, for example, information from internal anatomy, larval structure, or perhaps from nuances in nucleic acid research. Promising taxonomic results involving structural variations of the internal organs were published by Ekis and Gupta (1971), Crowson (1972), and Ekis (1978). Recently, Gerstmeier (2000) has pioneered some very exciting research in the Cleridae, setting the stage for innovative studies involving Cleridae DNA. It is hoped that such information will augment our present inventory of characteristics and contribute to the establishment of stable, heuristic classifications within Cleridae and Cleroidea.

CHARACTERS SELECTED FOR PHYLOGENETIC ANALYSIS
Twenty-six characters of Chaetophloeus and Plocamocera were used in the analysis, 25 from morphology and one from geographic distribution. Outgroups included representatives of the remaining 12 genera of Epiphloeinae and those of more distant outgroups such as genera of the Enopliinae-Korynetinae stock. Character states designated as ''0'' are considered plesiomorphic whereas those given a value of ' '1'' (or ''2'' in the case of transformation series) are judged apomorphic.
Character 0-Body size: (0) not diminutive, 5 mm or longer; (1) diminutive, less than 4 mm PHYLOGENETIC ANALYSIS OF CHAETOPHLOEUS AND PLOCAMOCERA A character matrix was coded for the 13 taxa listed in table 1 (see p. 63) and encoded into Hennig86. The analysis yielded one phylogenetic tree ( fig. 219) with a length index of 32, consistency index of 84, and retention index of 85.

PHYLOGENETIC INTERPRETATIONS
During the latter stages of this study I found among my fluid-preserved Cleridae one specimen of Epiphloeinae whose external features seem to bridge the evolutionary gap between Plocamocera and other Epiphloeinae genera such as Epiphloeus and Madoniella. It soon it became clear that this specimen represented a new genus and species most closely related to Plocamocera. Also, the serrulate condition of the epipleural margin present in all plocamocerans and in members of Madoniella californica (Van Dyke), M. merkely (Horn), and M. nebulosa (Chevrolat) suggest possible close kinships among the abovementioned genera. Stout elytral bristles (herein defined as chaetosomes), prominent among plocamoceran beetles and those of the monotypic genus Chaetophloeus, are also found, albeit in fewer numbers, in members of Epiphloeus mucoreus (Klug) and E. duodecimmaculatus (Klug). It remains to be determined whether some of the abovementioned similarities represent homoplasy or signify clues of suprageneric relationships. It is hoped that results from a survey involving structural variations of pronotal and elytral trichobothria, as well as elytral trichosomes within Epiphloeinae will contribute to a better understanding of sister-group relationships within this interesting subfamily. Figure 219 illustrates the extent to which my current knowledge of Chaetophloeus and Plocamocera permits predictions of phylogeny at generic, species group, and species TABLE 1 Character Matrix of 25 Morphological Characters and One Geographic Charcter levels. Table 1 enumerates the data on which the phylogenetic diagram and narrative of relationships are based. The evidence for the discussion seems minimal, but it is offered as a beginning to build upon as more information becomes available.
The ancestral progenitor of the Chaetophloeus-Plocamocera stock (ancestral species A) (see fig. 219) evolved a very robust pedicel, the metabasitarsus became relatively lengthened, the aedeagus narrowed and lengthened, and the male internal reproductive organs were modified to the extent that only one pair of accessory glands remained. In time, this progenitor evolved the sister genera Chaetophloeus and Plocamocera. The indications are that this initial divergence took place in South America during the mid-Tertiary when the southern continent was disconnected from lower Central America (Malfait and Dinkelman, 1972), which functionally served as a small archipelago. Such an island setting would have been very conducive for speciation events. The monotypic Chaetophloeus and 80% of the extant plocamoceran species occur in South America, with most species being distributed within the Amazon Basin and central highlands of Brazil. The conjecture of a South American origin for these two genera assumes three independent northern dispersals involving taxa of three species groups. Five are known to occur in Central America, and only P. sericella extends from Brazil to southern Mexico. Such an extensive distribution of a species is not novel among clerid genera. Indeed, widespread distributions of clerid species may be more common than has been heretofore suspected, in view of the strong flying habits of these beetles. For example, the enopliine Apolopha reichei Spinola is known to occur from Argentina to Mexico (Opitz, 1998). Widespread distribution of a single species within a genus is not novel among the Cleridae.
During the evolutionary progression from ancestral species A towards Chaetophloeus, chaetosomes became profusely distributed on the dorsal integument, and antennal sensilla trichodea were moderately lengthened. Antennal trichodea reached their maximal length in ancestral Plocamocera (i.e., ancestral species B), which was also characterized by an extensively transverse pronotum, very robust metacoxa and metafemur, an epipleural margin that evolved long trichobothria, and an abdominal venter that became sexually dimorphic in color. One of the early offshoots of the plocamoceran basic stock (ancestral species B) was the stem species of the P. castanea-confrater-coactilis group (ancestral species C), which eventually evolved into 14 known species, 13 of which stem from ancestral species D being characterized OPITZ: EPIPHLOEINAE . P. similis. 198. P. coactilis. 199. P. aspera. 200. P. santa. 201. P. onorei. 202. P. bispina. 203. P. carnegei. 204. P. paris. 205. P. ambra. 206. P. bolivari. 207. P. auratilis. 208. P. insula. 209. P. selva. 210. P. cericellopsis. 211. P. sericella. 212. P. jayhawkalis. . P. procera. 215. P. confrater. 216. P. buenavista. Antenna. 217. P. buenavista. 218. P. aura. by a clearly discernible tripartite humeral macula and an abundance of gold-colored setae. Ancestor D eventually diverged to produce the P. confrater (six species) and P. coactilis (seven species) species groups. During the evolutionary progression from pro-genitor E, there evolved a stem species (ancestral species F) that passed to its descendants (P. lucis and P. sesquipedalis) a substantially lengthened metatibial spur. During the evolutionary divergence of ancestral species E, there became established in its de- Fig. 219. Hypothesized phylogeny of Chaetophloeus and Plocamocera based on the Henig86 computer program. Letters at nodes represent ancestral species and numbers refer to apomorphic character states listed on page 92. High-set marks on numbers indicate a transformation series of a particular character, with the double marks representing the most derived condition. scendants a trend towards a reduction of the width of the antennal club, and there was a continuance of the lengthening of the metabasitarsomere, which reached its highest degree of development in P. lucis, P. sesquipedalis, and P. prolixa. Subsequent evolution of ancestral species G led to P. prolixa and to the complementary stock that produced ancestral species H. The latter proceeded towards a secondary reduction of metabasitarsal length evident in P. minima, P. aliguantula, P. manausensis, and P. iota. From the aspect of paleogeography, one may conclude that part of the evolutionary history of ancestral species G was a northward expansion from southern ancestral grounds during which populations of incipient species must have localized on the thendisjointed Central American archipelago. During this time, ancestral species H produced two major lineages, the P. sericella (seven species) group and its complementary stock (ancestral species I), in which there developed a considerable reduction of body length, the cranium became predominantly piceous, the antennal club reached its most slender form, and the tripartite elytral humeral macula became faintly visible. One stock from ancestral species I became an exclusively Central American progenitor (ancestral species J) that yielded P. minima and P. aliguantula. The complementary stock evolved ancestral species K, which remained in South America and evolved three species, P. salasis, P. manausensis, and P. iota, whose sister-group relationships cannot be postulated at present.