PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM issued SMITHSONIAN INSTITUTION U. S. NATIONAL MUSEUM Vol. 104 Washington: 1955 No. 3342 BIOLOGY AND TAXONOMY OF NORTH AMERICAN BEETLES OF THE SUBFAMILY GEOTRUPINAE, WITH REVISIONS OF THE GENERA BOLBOCEROSOMA, EUCANTHUS, GEO- TRUPES, AND PELTOTRUPES (SCARABAEIDAE) By Henry F. Howden ^ Introduction A number of entomologists have been interested in the biology and taxonomy of the beetles in the subfamily Geotrupinae, family Scara- baeidae. While the adults have been collected and their habits recorded, little has been published on the biology of the larvae of these elusive beetles. Revisions of the genera Eucanthus, Bolbocerosoma, Geotrupes, and Peltotrwpes are included, with descriptions of seven new species and subspecies. Also, two previously unrecognized subspecies of Geotrupes are described. The key to the tribes and genera of North American Geotrupinae is on page 161, and in the review of the subfamily the genera and species are treated in the following order: Bolbocerosoma Schaeffer B. biplagiatum Dawson and McColloch B.farctum (Fabricius) B. pusillum Dawson and McColloch B. confusum Brown B. pusillum townesi, new subspecies B. tumefactum (Palisot de Beauvois) B. quadricornum Robinson B. bruneri Dawson and McCoUoch B. ritcheri, new species B. hamatum Brown B. cartwrighti, new species B. lepidissimum Brown B. elongatum, new species ' Department of Zoology and Entomology, University of Tennessee, Knoxville, Tenn., formerly of North Carolina State College, Raleigh, N. C. 327015—55- 151 152 PROCEEDINGS OF THE NATIONAL MUSEUM Bradycinetulus Cockerell B. Uebecki (Wallis) B. fossatus (Haldeman) B. filicornis (Say) B. ferrugineus (Palisot de Beauvois) B. floridensis (Wallis) B. rex Cartwright B. simi (Wallis) Bolborhombus Cartwright B. darlingloni (Wallis) B. carinatus (Schaeffer) B. alabamensis (Wallis) B. angulus (Robinson) Geotrupes Latreille B. parvulus Cartwright G. opacus Haldeman Bolbocerastes Cartwright G. ulkei Blanchard B. regalis Cartwright G. blackburnii (Fabricius) B. imperialis Cartwright G. blackburnii excrementi Say B. imperialis kansanus Cartwright G. egeriei Germar B. serratus (LeConte) G. hornii Blanchard B. peninsularis (Schaeflfer) G. balyi Jekel Bolbelasmus Boucomont G. stercorarius (Linnaeus) B. arcuatus (Bates) G. splendidus (Fabricius) B. minor (Linell) G. splendidus miarophagus Say B. hornii (Rivers) G. semiopacus Jekel Eucanthxis Westwood Peltotrupes Blanchard E. lazarus (Fabricius) P. profundus Howden E. lazarus subtropicus, new subspecies P. profundus dubius, new subspecies E. alutaceus Cartwright P. youngi, new species E. greeni Robinson Mycotrupes LeConte Bolboceras Kirby M. retusus LeConte B. obesus (LeConte) M. lethroides Westwood B. falli (Wallis) M. gaigei Olson and Hubbell B. thoracicornis (Wallis) M. cartwrighti Olson and Hubbell B. cornigerus Melsheimer M. pedester Howden In the discussion, an attempt is made to compare the habits of North American Geotrupinae with the habits of Geotrupinae from other continents, particularly Europe. The feasibility of using food and burrowing habits as aids in taxonomic and phylogenetic investigations are discussed, and one case is pointed out where they could be applied. ACKNOWLEDGMENTS Much of the work has been carried out at North Carolina State College under the able direction of Dr. P. O. Ritcher, Oregon State College, who aided me greatly in the field, taking many of the photo- graphs included here and making many helpful suggestions during the preparation of the manuscript. I am also extremely grateful to O. L. Cartwright of the U. S. National Museum for aid in checking identifications and for many helpful suggestions on the taxonomic aspects of the work. Special thanks are also due to Dr. Henry K. Townes, North Caro- lina State College, for his help both in the field and on the manuscript, and to Dr. T. H. Hubbell, University of Michigan, whose comments and manuscript enabled me to write the summary on Mycotrupes. NORTH AMERICAN GEOTRUPINAE—^HOWDEN 153 Without the generous loan of specimens from numerous institutions and individuals, the present work could not have been accomplished. In the following list of collections studied, the letters in parentheses represent the abbreviations used in the text when citing material studied; the name of the curator responsible for the loan of the speci- mens follows the abbreviation: Academy of Natural Sciences of Philadelphia (ANSP), J. A. G. Rehn; American Museum of Natural History (AMNH), Mont A. Cazier; British Museum (Natural His- tory) (BM), F. I. van Emden; California Academy of Sciences (CAS), H. B. Leech; Canadian Department of Agriculture (CC), W. J. Brown; Carnegie Museum (CM), G. Wallace; Chicago Natural History Museum (CNHM), Henry S. Dybas; Cornell University (CU), Henry Dietrich; Emory University (EU), P. W. Fattig; EmoryUni- versity Field Station (EUFS), S. Breeland; Illinois Natural History Survey (INHS), M. W. Sanderson; Iowa State College (laSt), J. L. Laffoon; Museum of Comparative Zoology (MCZ), P. J. Darlington; North Carolina Department of Agriculture (NCSM), D. L. Wray; North Carolina State College (NCSC), T. B. Mitchell; Ohio State University (OSU), J. N. KnuU; Oregon State College (OSC), P. O. Ritcher; Tulane University (TU), A. Miller; United States National Museum (USNM), O. L. Cartwright, E. A. Chapin and W H.. Anderson; University of Arizona (UA), George D. Butler; University of California, Davis (UCD), A. T. McClay; University of Idaho (Unid), W. F. Barr; University of Kansas (UnKa), R. H. Beamer and P. J. Spangler; University of Maryland (UnMd), E. N. Cory; University of Michigan (UnMich), T. H. Hubbell and A. L. Olson; University of Missouri (UnMo), G. W. Thomas; University of Wis- consin (UnWis), R. D. Shenefelt and E. W. King. In addition, I am indebted to the following individuals who kindly loaned material for study from their personal collections: O. L. Cartwright, T. Daggy, R. W. Dawson, B, K. Dozier, M. Robinson, P. J. Spangler, and F. N. Young. Thanks also are due to Drs. O. H. Fulkner, B. B. Fulton, D. L. Grosch, Z. P. Metcalf, T. B. Mitchell, and T. L. Quay of North Carolina State College, and to Dr. W. L. Engles of the University of North Carolina for suggestions and corrections in the manuscript. B. K. Dozier and W. H. Long were able field assistants on some occasions. Dr. F. I. van Emden, British Museum, and Dr. S. L. Tuxen, Zoologische Museum, Copenhagen, were extremely helpful in com- paring specimens with type material. My wife, Anne, was of tremendous help, both in the field and in the typing of the manuscript. Without her encouragement and help the work involved would have been much more difficult. 154 PROCEEDINGS OF THE NATIONAL MUSEUM vol. loi REVIEW OF LITERATURE Latreille (1796) was the first to propose the genus Geotrupes in the Scarabaeidae, not placing, at that time, any species in the genus. Two years later Fabricius (1798) again proposed the name Geotrupes with 65 included species, but used the name with an entirely different connotation than did Latreille. Latreille (1802, 1804) repudiated the Fabrician application of the name and listed some species that he believed should be included in the genus, and it has been his concept, and not that of Fabricius, that has been followed by subsequent authors. Recently, Potts (1948, 1951) has raised the question of the validity of the acceptance of G. stercorarius Linne as type of the genus, for it was not included in the Fabrician list of species in 1798. I believe, however, as did Latreille (1802), that the Geotrupes of Fabricius was not intended to be that of Latreille and is therefore a primary homonym, invalidating the Fabrician list for genotype selection in this case and leaving G. stercorarius Linne as type of the genus. This problem has been more fully discussed by Townes and Howden (1952) and has been referred to the International Commission of Zoological Nomenclature. It is still under consideration by the Commission. The erection of the subfamily (or family) Geotrupinae was credited to Latreille (1804). Following Latreille there were a number of more or less comprehensive taxonomic works on the subfamily which in- cluded the tribes Bolboceratini, Geotrupini, and Lethrini. Little of this work, however, dealt even sparingly with North American forms. The first comprehensive paper on North and Central American species was a monograph by Jekel (1865) that dealt solely with the tribe Geotrupini. This paper incited considerable interest and Horn (1868, pp. 313-322) published what is essentially a supplement to this work, commenting on synonymy and pointing out varieties within some of the species. He followed this in 1870 (pp. 48-50) with a key to the other North American genera, exclusive of Geotrupes, for which he subsequently published a key (1880, pp. 144-145). The information on the Central American Geotrupinae was assembled by Bates (1887), and one year later Blanchard (1888) published the most recent comprehensive work on the North American species in the genus Geotrupes. Bradley (1944) published a key to the species of Geotrupes as given by Blanchard, but without any consideration of synonymy or undescribed species. The information that had been rapidly accumulating before 1900 was finally brought together and greatly augmented by a series of extensive papers, worldwide in scope, by Boucomont. In 1902, in Wytsman's "Genera Insectorum," he set forth the genera and listed NORTH AMERICAN GEOTRUPENAE—HOWDEN 155 the described species under each. This was followed by a paper by Schaeffer (1906) in which was published a revision of the North American genera and species in the tribe Bolboceratini. In the same year Boucomont (1906) published his "Catalogue Provisoire des Geotrupidae." Then, in 1911, Boucomont published a revision of the genera of the Geotrupinae, dealing in large part with the Bolbocera- tini, in which he discussed Schaeffer's paper and proposed a number of new genera and subgenera. Perhaps the most useful part of Boucomont's work was his catalog of the literature of the subfamily published in 1912 in Schenkling's ''Coleopterorum Catalogus." He subsequently published a number of other papers, none of which dealt with the American fauna.