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This Work Is Licensed Under the Creative Commons Attribution-Noncommercial-Share Alike 3.0 United States License This work is licensed under the Creative Commons Attribution-Noncommercial-Share Alike 3.0 United States License. To view a copy of this license, visit http://creativecommons.org/licenses/by-nc-sa/3.0/us/ or send a letter to Creative Commons, 171 Second Street, Suite 300, San Francisco, California, 94105, USA. Evarthrus sodalis sodalis LeConte Lexington, Kentucky Photograph by J. Scott A REVISION OF THE SPECIES OF THE GENUS EVARTHRUS LECONTE (COLEOPTERA: CARABID AE) RICHARD FREITAG Quaestiones entomologicae Departmen t of Biology 5 : 89-212 1969 Lakehead University Port Arthur, Ontario Within the genus Evarthrus, three subgenera, 43 species, and five subspecies are recog­ nized. The genus Evarthrus is described, and evidence is presented which removes Evarthrus as a unit from the Pterostichus complex to a position near the genus Molops in the tribe Pterostichini. A key to the species and subspecies is given. Each subgenus, species group, and species is described and synonymies are listed. The distribution of each species is presented by locality records and distribution maps. Structures which are used in identification are illustrated. The subgenus Fortax comprises six species of which one, iuvenis, is described as new. One genus group name and six species names are reduced to synonymy. The subgenus Cyclotrachelus includes 12 species, of which five, fucatus, macrovulum, texensis, parafaber, and levifaber are described as new. Six species names are listed as synonyms. The subgenus Evarthrus includes 25 species of which seven are new. The species sodalis LeConte and torvus LeConte are polytypic. Five genus group names and 26 species names are relegated to synonymy. A phytogeny is postulated for the subgenera, species groups and species. The geographical distribution of the genus is discussed. The endemic flightless Pterostichini of eastern North America are arrayed in a series of supraspecific taxa each of which is more or less easily defined; however, the relationships of these groups are at best uncertain. The most diverse of these groups is the genus Evarthrus, a complex of species included by some (Csiki, 1930; Lindroth, 1966) in the genus Pterostichus, and by others (Casey, 1918) treated as a group of related genera. (Ball, 1967 drew attention to this problem). I decided to attempt to solve these problems of relation­ ships and classification, but learned in the initial stages of the study that I would first have to undertake a revision of the species. The results of this investigation are presented in this paper. The revision is based on a study of adult specimens. The species were defined on the basis of evaluation of morphological and geographical evidence. Names were applied on the basis of study of the relevant type material. Sixteen species names were found to be junior synonyms, and 13 previously undescribed species were discovered and named. In addition to the formal taxonomic treatment of this group, I have presented my views on the phylogeny and geographical distribution of the extant species. 90 Freitag MATERIALS AND METHODS " Materials The material examined consisted of 7,600 adult specimens, which included the type specimens of Casey and LeConte. I have also briefly examined external structures and male genitalia of species of an additional 35 Nearctic and Palearctic subgenera of Pterostichns Bonelli, and of Abaris Dejean, Pseudabarys Chaudoir, Oxycrepis Reiche, Cratocerus Dejean, Catapiesis Brulle, Abax Bonelli, Molops Bonelli, Percus Bonelli, Lesticus Dejean, Piesmus LeConte, Stomis Clairville and Myas Dejean. Names of individuals and institutions from which material was borrowed are abbreviated in the text as follows: AMNH — American Museum of Natural History; ANSP — Academy of Natural Sciences; AU — Auburn University; BM — British Museum (Natural History); CAS — California Academy of Sciences; CM — Carnegie Museum; CNC — Canadian National Collection; CNHM — Chicago Natural History Museum; CU — Cornell University; DL — David Larson; DRW - D. R. Whitehead; FDPI - Florida Division of Plant Industry; GEB - G. E. Ball; INHS - Illinois Natural History Survey; ISU - Iowa State University; KLE - Kansas State University; MCZ — Museum of Comparative Zoology; MHNP — Museum d'Histoire Naturelle, Paris; MSU — Montana State University; NCSU — North Carolina State University; RCG - R. C. Graves; RF - R. Freitag; RTB - R. T. Bell; RU - Rutgers University; TAM — Texas A & M University; TCB — T. C. Barr; TE - T. Erwin; TH - T. Hlavac; UA — University of Arkansas; UASM — University of Alberta Strickland Museum; UK — University of Kansas; UL — University of Louisville; UMMZ — University of Michigan Museum of Zoology; UP - Purdue University; USNM - United States National Museum; UW — University of Wisconsin; VMK — V. M. Kirk. Methods General methods By making comparisons among their characteristics, specimens were sorted into demes, subspecies, species, and species groups according to degree of similarity and difference. The characteristics used were arbitrarily weighed, and the same characteristics were given different weights in different situations. The relationships revealed by these comparisons were interpreted and the evolution of the species and species groups was then inferred. Characters of adults Some structures which are used in the identification of the species of Evarthrus are discussed below to facilitate their use in the text. The lines of microsculpture of the integument of the dorsal surface are almost effaced, disoriented, and do not form meshes in specimens of some species. In specimens of other species the lines are close together and sinuate, but they do not form meshes. More frequently meshes are formed and are amorphic or isodiametric. The interspaces are flat or raised and bead-like. The lustre of the integument correlated with the microsculpture is as follows: shiny in the absence of meshes; dull with isodiametric meshes and flat interspaces; matte or velvet with isodiametric meshes and bead-like interspaces; iridescent with dense sinuate parallel lines. Females are always duller than males of the same species. Revision of Evarthrus 91 The frontal grooves of the head are usually of a particular shape, and are useful in recognizing specimens of a number of species. The grooves are straight, or crescent-shaped with the convexity directed medially or laterally (figs. 70-71). The number of setae on the penultimate article of the labial palpus is useful in delimiting species. Two dorsal "primary" setae are always present near the halfway point of the article. Three "secondary" setae are also present. One is apical, and arises from the ventral side of the article. Another is near the apical end of the article. It is dorsolateral and directed dorso­ lateral^. In a complementary position is the third seta. It is also near the apical end of the article on the dorsomedial side and is directed dorsomedially. The truly apical seta occurs more frequently than the other "secondary" setae but it is not always present. Other setae occasionally occur here and there near the "primary" setae (figs. 66-69). Horn (1881) noted that the number of setae on the labial palpus are not constant in Evarthrus and suggested that two groups may be recognized: a group with bisetose labial palpi and a group with plurisetose labial palpi. The mandibles of the type species, sigillatus Say, are illustrated (fig. 72). Details of form and structure of the pronotum are useful in recognizing subgenera and species. The general outline of the pronotum ranges in form from rectangular to cordiform (figs. 1-62). Another useful structure is the shape of the basal lateral fovea, which is puncti- form (figs. 1-7), monostriate or bistriate (figs. 8-20, 21-62). The position of the basal lateral seta is on (figs. 8-20), or beside the lateral bead (figs. 1-7 and 21-62). The lateral bead in specimens of a few species is broad posteriorly, but in most species it is narrow posteriorly as in fig. 25. The prosternal process which projects posteriorly between the front coxae has a longitudinal medial groove. This groove is deep or shallow. The apex of the prosternal process is or is not marginate. At least four setae are present on the anterior face of the middle femur and always in the same positions. A proximal pair of setae are located near the ventral side of the femur, and a distal pair near the dorsal side. In specimens of some species additional setae usually occur near either pair. The total number of setae ranges from four to eleven in the genus and seems to be a good character for grouping species (figs. 74-76). Setae are absent from the lateroventral margin of the claw bearing tarsal articles in specimens of four species of the subgenus Fortax. I have used the term "last abdominal sternum" in the text. This is morphological sternum VII, which is the apparent sternum VI in beetles (sternum 1 has disappeared). The male genitalia are very important structures in defining species and species groups. In fact, it would be exceedingly difficult to recognize and classify the species of Evarthrus without reference to these structures. Two parameres and a median lobe which contains an internal sac constitute the external male genitalia. The median lobe is a tube with a central bend. The portion of the lobe posterior to the bend is referred to here as the apical half, an<J that anterior to the bend the basal half. The lobe is always bent dorsoventrally with the convexity directed dorsally. In specimens of some species the apical half of the median lobe
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