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Erwin 1970 Qev6n1 4 215 CC Released.Pdf 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. Brachinus janthinipennis (Dejean), male, Cornwall, Connecticut. A RECLASSIFICATION OF BOMBARDIER BEETLES AND A TAXONOMIC REVISION OF THE NORTH AND MIDDLE AMERICAN SPECIES (CARABIDAE: BRACHINIDA) TERRY LEE ERWIN Museum of Comparative Zoology Harvard University Cambridge, Massachusetts Quaestiones entomologicae U.S.A. 02138 6:4-215 1970 A taxonomic revision of the species of the North and middle American Brachinus Weber is presented. Sixty-two species are recognized as valid. Neobrachinus new subgenus, is erected to contain the New World species formerly included in Brachinus (sensu stricto). The following 14 taxa are described as new: Brachinus alexiguus, B. capnicus, B. chalchi- huitlicue, B. chirriador, B. cibolensis, B. javalinopsis, B. imporcitis, B. oaxacensis, B. galac- toderus, B. ichabodopsis, B. mobilis, B. explosus, B. aabaaba, B. sonorous. Twenty-five names are reduced to synonymy for the first time. A key to the species is given. Each species and species group is described and synonymies are listed. The distribution of each species is presented by locality records and distribution maps. Structures used in identification are illustrated. Two species of Pheropsophidius Hubenthal occur in Middle America and are included in the key to bombardier beetles of North and Middle America, one of these species is des­ cribed. Structures of both species are illustrated. A reclassification of the genera of the world bombardier beetles is presented. Seventeen genera are recognized as valid. These include 13 valid subgenera, one of which is new. Two tribes are included in the division Brachinida. One of these tribes includes four subtribes, two of which are new. Three generic names are reduced to synonymy for the first time. All taxa are described and keys for their identification are presented. A discussion of the general morphology of members of the genera is presented. Structures discussed are illustrated. A hypothetical phytogeny is presented for all genera of the division Brachinida and all species of Neobrachinus new subgenus. The geographical distribution of these taxa is also discussed. Figures and tables accompany the discussions. CONTENTS Materials and methods p 6 Comparative morphology p 11 Taxonomy p 27 Life histories and immature stages p 166 Phytogeny p 166 Zoogeography p 184 References p 209 6 Erwin The bombardier beetles are a complex and taxonomically difficult group belonging to the family Carabidae. In 1862, LeConte stated: "I must also add that I consider the species of this genus very decidedly opinionative, and that I am only impelled to the publication of this note by the necessity of giving names to a certain number of recognized forms, and of placing as synonyms some which I formerly considered as distinct, but which increased col­ lections have since shewn to be varieties." The situation had hardly changed 98 years later when Ball (1960: 164) wrote: "The taxonomy of the North American species of this group is very poorly understood and it is almost a waste of time at present to attempt to determine individuals to species." Until now, a taxonomic revision of all North and Middle American species of Brachinus has not been undertaken, probably because of the great morphological similarity among the species, a lack of traditional characteristics for separating the species, and the unavailability of much of the "type" material. The first of these obstacles was surmounted after the dis­ covery of external characteristics not used previously. The last problem was resolved with the help of G. E. Ball who compared my specimens with the type material of Dejean and Chaudoir in Paris. I was able to compare the LeConte and Blatchley types myself at the Museum of Comparative Zoology and Purdue University, respectively. This paper is an extension of my 1965 publication and deals with the taxonomy and dis­ tribution of Brachinus and Pheropsophidius in North and Middle America. I also propose here a reclassification of bombardier beetles of the world, including subgeneric components; discuss phylogenetic and zoogeographic hypotheses; and present a preliminary study of the comparative morphology of bombardier beetles. Bombardier beetles were first recognized formally when Linne described the species Carabus crepitans in 1758. Since that time, more than 600 trivial names have been proposed for members of the Brachinida. Dejean (1825, 1831), LeConte (1844, 1848, 1858, 1862), and Chaudoir (1868, 1876) have described most of the species in North and Middle America. Chaudoir's paper (1876) has been the only monographic treatment of the Brachinida as a whole. Basilewsky's 1959 revision of the Crepidogastrini must be considered monographic of that tribe. LeConte's 1862 paper and my revision of the California species (1965) contain the only new keys to North American species. (Blatchley's 1910 monumental "Coleoptera of Indiana" contains a key to the Brachinus species of Indiana, but it was based on LeConte's key.) Many faunal studies have included partial taxonomic treatment of some of the taxa of the Brachinida. Notable are publications by Antoine (1962), Basilewsky (1962), Darlington (1968), Habu (1967), Jeannel (1942, 1949), Liebke (1934), Peringuey (1885, 1888, 1896, 1898), and Reitter (1919). No complete taxonomic classification has been proposed for the group as a whole. Most workers still seem to rely more heavily on the catalogue classification established by Csiki (1933) than on any other reference source. MATERIALS AND METHODS Materials This study is the result of the examination of 28,633 specimens of North and Middle American Brachinus, and 2,172 specimens of other bombardier beetles. Most of these speci­ mens have been loaned to me by museums and private collectors in Canada, Europe, Mexico, and the United States. The others were collected by my wife and me in the United States. Were it not for the intensive collecting of carabids in Mexico by G. E. Ball and D. R. Whitehead, this area would have been omitted from this study due to insufficent material. Bombardier Beetles 7 Larval material was especially difficult to obtain. In addition to material from California (Erwin, 1967), I have seen one first instar larva of Brachinus mexicanus Dejean, and one unidentified last instar larva. This is still all that is known in North America, besides that which H. F. Wickham collected in the 1890's. The following abbreviations indicate the various museums and private collections from which specimens were borrowed: AMNH American Museum of Natural History, New York, New York 10024, P. Vaurie, L. Herman Jr. ANSP Academy of Natural Sciences, Philadelphia, Pennsylvania 19103, H. R. Roberts. AUAA Auburn University, Auburn, Alabama 36830, K. L. Hays. BMNH British Museum (Natural History), London, England, R. D. Pope. CArm C. Armin, 191 West Palm Avenue, Reedley, California 93654. CAS California Academy of Sciences, San Francisco, California 94118, H. B. Leech. CBak C. Baker, Boise State College, 1907 Campus Drive, Boise, Idaho 83707. CCha C. Chantal, 425 Saint Olivier, Quebec 4, Quebec. CEWh C. E. White, 2441 East Northview Avenue, Indianapolis, Indiana 46220. CMPP Carnegie Museum, Pittsburgh, Pennsylvania 15213, G. E. Wallace. CNC Canadian National Collection of Insects, Entomology Research Institute, Ottawa, Ontario, E. C. Becker, W. J. Brown. CNHM Chicago Natural History Museum, Chicago, Illinois 60605, H. Dybas. CPBo C. Bolivar y Pieltain, Instituto Politecnico Nacional, Mexico D. F. CUNY Cornell University, Ithaca, New York, 14850, L. L. Pechuman. CVMA Coachella Valley Mosquito Abatement District Collection, Thermal, California 92274. DDLa D. J. Larson, University of Calgary, Calgary, Alberta. DHKa D. H. Kavanaugh, 1121 Garfield Street, Denver, Colorado 80206. DRWh D. R. Whitehead, University of Alberta, Edmonton 7, Alberta. DTRT Division of Tropical Research, Tela Railroad Company, La Lima, Honduras, Central America, W. G. C. Forsyth. FDAG Florida Department of Agriculture, Gainesville, Florida 32601, R. E. Woodruff. GRNo G. R. Noonan, University of California, Riverside, California 92502 (including col­ lection of F. Andrews). HGou H. Goulet, University of Alberta, Edmonton 7, Alberta. HMO Hope Museum, Oxford, England, E. Taylor. ISNH Illinois State Natural History Survey, Urbana, Illinois 61803, L. K. Gloyd. ISUA Iowa State University, Ames, Iowa 50010, J. Laffoon. JHen J. Hendrichs, Apartado Posta, 11-774, Mexico 11, D. F. JSch J. Schuh, 4039 Shasta Way, Klamath Falls, Oregon 97601. KMTB Koninklijk Museum Voor Midden-Afrika-Musee Royal del' Afrique Centrale, Tervuren, Belgium, P. Basilewsky. KSU Kansas State University, Manhattan, Kansas 66502, H. D. Blocker. LACM Los Angeles County Museum, Exposition Park, Los Angeles, California 90007, C. L. Hogue. LBSC Long Beach State College, Long Beach, California 90804, E. L. Sleeper. LRus L. Russell, Oregon State College, Corvallis, Oregon, 97331. LSUB Louisiana State University, Baton Rouge, Louisiana 70803, J. B. Chapin. MSUM Montana State University, Missoula, Montana, 59801, N. Anderson. MCZ Museum of Comparative Zoology, Cambridge, Massachusetts 02138, P. J. Darlington, Jr. 8 Erwin MHNP Museum National d'Histoire Naturelle, Paris, A. Bons. MMM Moscow Museum, Moscow,
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